Ка 2 7 775 Annals of the Missouri Botanical Garden | С: | ар E м Flora of Panama. Part V. Fascicle 3. ВА second рагі) . Robert E. Woodson, Jr Robert M hA EM and Collaborators. 1—96 LISHED QUARTERLY АТ GALES SBU RG, ILLINOI 5, BY THE BOARD OF TRUSTEES OF 19, Mi oven BOTANICAL ASEM, ST. LOU Entered as IAEA pm at the Harp pes at fete шой, the Act of March. %; 1879 vant dois И! Wer оа oL Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation "with the Missouri Botanical Garden. | Information The ANNALS OF THE Missourr BOTANICAL GARDEN appears four times durirg the calendar year: February, May, September, and November. Four num е. i Subscription Price -$10.00 per volume Single Numbers 22. 2.50 each Contents of previous issues of the ANNALS OF THE Missouri BOTANICAL GARDEN are listed in the Agricultural Index, published by the H. W. Wilson pany. FLORA OF PANAMA Part V. Fascicle 3 LEGUMINOSAE’ Subfamily CaEsALPINOIDEAE (Caesalpinaceae of many authors) Trees, shrubs, or less frequently vines or herbs. Leaves usually compound, mostly pinnate, not infrequently bipinnate or occasionally unifoliate. Inflorescence mostly terminal or subterminal, usually racemose or paniculate of several racemes. Flowers mostly yellow or red; calyx usually with a short tube scarcely distinguish- able from the receptacular base, mostly polysepalous and 5-parted above, in Swartzia rupturing irregularly; petals 5, infrequently fewer or absent, imbricate, the uppermost within the others in bud; stamens usually 10, sometimes fewer, rarely more numerous; filaments usually not united; ovary free or adnate to the calyx-tube, sessile or stipitate. Legume of diverse types. A subfamily of attractive plants well represented in the tropics, of which many genera are of significant economic or ornamental interest. This subfamily received considerable attention from Bentham, whose masterly comprehension of the genera has to a great extent been accepted in later works. Britton and Rose have worked out the "Caesalpinaceae" for the ‘North American Flora’, but it is difficult to accept the multitude of segregate genera recognized by these authors, many of which are poorly delimited and nearly impossible to locate with confidence by use of the keys. a. Calyx entire, closed in bud, rupturing in anthesis; petals usually 1, i ) sometimes lacking; leaflets 1- to 5-foliolate (in Panama), odd-pinnate (SWARTZIEAE) 1. SWARTZIA profoundly 2-lobed) (ВлунгмтклЕ).........-...--.--..а. 1.1... а... а а... 2. BAUHINIA і іріппасе, or obviously 2-foliolate. c. Leaves pinnate or 2-foliolate. d. Anthers dorsifixed, versatile, longitudinally dehiscent. e. i ened base, synsepalous for at least a short distance above the receptacular portion. f. Leaflets many, small (2 cm. or less); perfect stamens 10; ovary short-stipitate, the stipe adnate to one side of the calyx; legume membranaceous, indehiscent (POEPPIGIEAE). To be expected in Panama . 3, PoEPPIGIA ff. Leaflets few, large (6-18 cm.) in Panamanian species; stamens 10, or 5 and staminodes 5; ovary sessile or nearly so; legume woody, 2-valved. g. Stamens 5, staminodes 5; long), complete (МокЕлЕ flowers larger (about 6 mm. ) . 4. Mora lIssued March 22, 1951. Continued from Ann. Mo. Bot. Gard. 37:314 (Fl. Panama 52:300). 950. (301) [Vor. ANNALS OF THE MISSOURI BOTANICAL GARDEN . Stamens 10; flowers smaller (about 2.5 mm. long), apetalous (apparently complete because of calyx- like а е. Calyx polysepalous above а thickened receptacular tube ог PM ase. lyx with a short or small receptacular base; ovary es- seatially free from the calyx or receptacle (CYNOMETREAE except Nos. 5 and 21). в. Petals 5 or 3 (see Priorta). h. Petals 3; filaments united неңді: legume winged from upper suture; leaflets a few pairs, several cm. lon hh. Petals 5; filaments free; урну; not winged; leaflets her pe smaller or of a sing . Leaves mostly 6- to 8- foliolates leaflets — strongly obovate; legume thin t (CAESALPINIEAE)............ esses 2-foliolate; эм larger, elliptic; legume gg. Petals lacking (sepals simulating petals in Priori). E tes ebracteate in anthesis; leaflets several, 30 - E Р. hh. RON 5-parted, conspicuously bracteate; leaflets 4, ff. TON. pi di a muore gross, tube-like receptacular ye to which the stipe of the ovary is variously adnate (A EAE). g. Leaflets 1 pair (in Panama); stipe of the ovary at least basally adnate to the stocky receptacular tube. amens 10 i. Stigma dilated; legume about 3 cm., 2-valved, ШЕ compressed or flattened; flowers small ii. геч. n egume large, indehiscent, terete; hh. Upper К ange the others rudimentary or lacking; rtile st gg. idus leaflets 2) o many pair; stipe of the ovary adnate the length of the elongate жү r tube. h. Leaflets large, relatively few; inflorescence condensed, үрөр sheathed by large pe fertile stamens г Flow ers wi ich dug rip calycine, ensheathing bract- P lets; stamens less than 12 (in Рапата)........................ ii. Flowers without de ensheathing bractlets; sta- mens 14—16 (in Panama) hh. — small кат, inflorescence expanded, race- ose; fertile stamen dd. Anthers basifixed, opening by d pores or slits (CASSIEAE). e. Petals 1-2 and minute, or lacking ee. cc. Leaves bipinnate (CAESALPINIEAE). d. Flowers very large, T petals about 6 cm. long; legume ligneous, elongate, frequently 4—6 dm. long. Introduced ornamental........ dd. Flowers smaller, the pde at most эсе 2 cm. long; legume scarcely ligneous, less than 1. on, e. Stigma pe tate; legume cb d on both margins; leaves and inflorescence densely ferruginous (їп Panama). Introduced tree ee. Stigma not peltate nor expanded; legume not winged; leaves and inflorescence rarely ferruginous (in Panama in Caesalpinia eriostachys only E igi d unilaterally adnate to calyx-tube; calyx-tube appear- ique or “lop-sided”; spatulate hum samaroid with a p С-ы inci seed; unarmed tree, the perds very large... (302) 8. PRIORIA 5. PHYLLOCARPUS 21. HAEMATOXYLON 6. CYNOMETRA 7. COPAIFERA 8. PRIORIA 9. PELTOGYNE 10. HYMENAEA 11. MACROLOBIUM 12. BRowNEA 13. BROWNEOPSIS 14. TAMARINDUS 15. DiALIUM 16. Cassia 17. DELONIx 18. PELTOPHORUM 19. ScHIZOLOBIUM 1951] FLORA OF PANAMA (Leguminosae ) 3 ff. Ovary free in calyx-tube from basal insertion; calyx-tu AM legume not spatulate and samaroid with a single cal seed; armed or unarmed tree, the vas mostly mo g Blants unarmed or E irregularly with recurved € leaves with a picuous primary rachis байн 8 outermost бу: рт cucullate or subcucullate a) 20. CAESALPINIA m n Pan gg. Plants find usually prominently e ed at the odes; leaves with a reduced or obsolete primary rachis; calyx-lobes, except in Haematoxylon, E a h. Outermost c lyx- lobes cuculla me a DES spicuous excentric lateral suture; ae id pod 21. HAEMATOXYLON usu ually obsolete hh. Outermost ns lobes about equal, not cucullate; legume with marginal sutures; primary leaf rachis mostly present doe gh often modified. pisos ius т йр from nodal spines; с? e; legume flattened, not torulose; to be expect "y in E 22. CERCIDIUM . Petiole and rachis of ч very short or almost ob- solete, consisting at least in part o prominent spine; rachis of pinna fat; legume torulose 23. PARKINSONIA 1. SWARTZIA Schreb. SwARTZIA Schreb. Gen. Pl. 2:518. 1791, nom. conserv. Tounatea Aubl. Pl. Guian. Franc. 1:549, pl. 218. 1775. Possira Aubl. loc. cit. 2:934, pl. 355. 1775 Rittera Schreb. loc. cit. 1:364. 1789; Sw. Fl. Ind. Oce, 935, f. 16. 1800. Hoelzelia Neck. Elem. 3:62. 1790 Riveria HBK. Nov. Gen. & Sp. 7: 266, A epi 1825. Dithyria Benth. in Hook. Jour. Bot. 2:8 Fairchildia Britt. & Rose, in N. Am. i p 51. igi other irish have been given for Swartzia (vide Dalla Torre & Harms, Index Small trees, usually glabrous and with 1- to 5-foliolate leaves (in Panama). Smaller branches usually conspicuously lenticellate. Leaves monofoliolate or pin- nate, stipulate, with petiole and rachis (if present) usually obsagittate-alate or at least angled; leaflets ovate to elliptic, membranaceous to coriaceous, prominently veined, with principal lateral veins anastomosing towards the margin; petiolules (in Panama) short and terete. Inflorescence few- to many-flowered, bracteate; buds diagnostic, clavate-pedicellate, globose, entire and (calyx) rupturing into 3—5 irregular sections at anthesis. Flower apetalous or 1-petalate, petal usually large and suborbicular, clawed; stamens usually many, generally of two sorts, fewer (less than 15) larger and longer ones ventrally, with many smaller and somewhat shorter ones above; anthers versatile, with conspicuous connective. Legume 1- to few-seeded, short or elongate, subterete to flattened. A distinctive genus among Panamanian Leguminosae, easily recognized by the entire buds and mono- or apetalous flowers. However, specific bounds within sec- tions of the genus have not been clear, and a number of "species" have had to be condensed here as S. simplex. (303) [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN e Fig. 105. Swartzia panamensis Predominantly neotropical from Mexico to southern South America; Africa. Common in Amazon basin, especially along the upper Rio Negro. a. Flowers apetalous; — igs terminal leaflet generally 15-30 cm. long; legume elongate, s 2. S. NUDA aa. Flowers monopetalate; viens smaller, terminal leaflet generally 6—20 cm. long; legume short, or large and broa b. Inflorescence elongate, Mode 20—40 cm. long, many-flowered; legume flat, broad . S. PANAMENSIS bb. Inflorescence short, usually less than 10 cm. long, few- eso iia legume subterete, turgid. c. Petal "i if any s than the calyx; stamens relatively е 18 . S. ARBORESCENS cc. Peral T longer than the calyx; stamens ma d. At least some leaves 3- foliolate; flowers orem less robust, (304) FLORA OF PANAMA (Leguminosae) 5 petal usually less than 2.5 cm. wide ‚4. S. SIMPLEX var. DARIENENSIS dd. Leaves 1-foliolate; flowers robust, petal often more than 3 cm. wide 5. S. SIMPLEX 1. SWARTZIA PANAMENSIS Benth. in Mart. Fl. Bras. 157:38. 1870. Swartzia pinnata Seem. Bot. Voy. Herald, 113. 1853, nec Willd., nec Willd. ex Vog., nec ittera pinnata Vahl. Tounatea panamensis (Benth.) Taub. in Bot. E 47:392. 1891 Fairchildia panamensis (Benth.) Britt. & Rose, in N. Am. Fl. 23:348. 1930. Trees to 20 m. tall, branchlets glabrous, ME lenticellate. Leaves large, 5-foliolate; stipules lanceolate, 1-2 mm. long; petiole subterete to angular, 1-6 cm. long; rachis not winged, 2.5—9 cm. long; leaflets glabrous to lightly pubescent below along chief nerves, ovate-lanceolate to elliptic- -lanceolate, long caudate-acuminate apically and broadly acute basally, 4-20 cm. long and 1.5-7 cm. broad, submembranaceous, prominently nerved beneath; petiolules gross, sul- cate above, 3-7 mm. long. Inflorescence many-flowered, up to 40 cm. long; peduncle lightly pubescent, somewhat angled; pedicels 1.3-2.5 cm. long; mature buds globose, verrucose, 7—9 mm. in diameter. Flowers 1-petalate; petal yellow, irregularly orbicular, dentate-fimbriate marginally, up to about 3.8 cm. long and broad, claw about 7 mm. long; calyx rupturing irregularly into 3—5 reflexed lobes; stamens many, apparently subequal, 8 or so lower filaments thicker than the rest, up to 1.5 cm. long; anthers bilocular, apically acute-subcaudate, up to 5 mm. long in larger anthers. Legume large, broad, flat, beaked, 2-2.5 dm. long and up to 1 dm. wide, splitting first along the ventral suture, the valves thick and somewhat elastic; seeds few, large, flattened, oval, 6-8 cm. in diameter, exarillate. Panama and Honduras. ANAL ZONE: Gatun River kasi Pittier 6511; between R. Pequeni and R. Indio Stik “a Allen 16786; Quebrada Ancha, Dodge & Steyermark 16786a; dr River, Pittier 2010. DARIEN: mouth of Rio Yapé, werd 324. SAN BLAS: Puerto Obaldia, Pittier 4324. 2. SWARTZIA NUDA Schery, in Ann. Mo. Bot. Gard. 30:92. 1943. Glabrous trees with terete branches. Leaves 3- to 5-foliolate; petiole (with rachis) 10—13 cm. long, flattened above and 2-3 sulcate, swollen at the nodes; leaflets elliptic, 14—32 cm. long, 6—13 cm. broad, acute or obtuse basally, acute and briefly attenuate apically, with about 12—20 confluent, arcuate, scarious-pubescent, lateral nerves; petiolules terete, canaliculate above, 4—8 mm. long. Inflorescence axillary from non-foliate nodes, spicate, many-flowered, 5-16 cm. long, short- pubescent; pedicels 1-1.5 cm. long; buds globular, about 6 mm. in diameter. Flowers apetalous; calyx rupturing into 3—5 irregular lobes; stamens many, gla- brous, in 2 series, shorter ones 1—1.5 cm. long, longer ones 1.9—2.1 cm. long; anthers bilocular, smaller ones 1.5—1.8 mm. long, larger ones 2—2.5 mm. long; ovary glabrous, linear-stipitate, including the style 2-3 cm. long; ovules about 13; style 6—7 mm. long; stigma truncate-capitate. Legume elongate, 12-20 cm. long, subterete, stipitate, apically attenuate; loculi 1—2, 1.3—2 cm. broad, interlocular constriction 0,2—1 cm. broad; seeds arillate, about 5 cm. long and 0.7 cm. broad. (305) [Vor. 38 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 106. Swartzia nuda Northwestern Panama. OCAS DEL TORO: Isla Colón, von Wedel 1073, 1107, 1224; Water Valley, von Wedel 909, 957, 172 3. SWARTZIA ARBORESCENS (Aubl.) Pittier, in Jour. Wash. Acad. 11:157. 1921. Possira arborescens Aubl. Pl. Guian. a 2:934, pl. 355. 1775. Possira triphylla Sw. Prodr. Veg. Ind. 1788. Rittera triphylla Sw. Fl. Ind. Occ. T i s Swartzia triphylla Willd. Sp. Pl. 2:1220. 1800. Swartzia parviflora DC. Mem. Leg. 403, pl. 60. 1825, fide Index Kew., Hemslev, Britton. Tounatea arborescens (Aubl.) Britton, in Bull. Torrey Bot. Club 16:325. 1889 Swartzia rariflora Hoehne, in Comm. Linh. е Estrat. Matto Grosso (Publ. 74] Annexo 5, Bot. pt. 12:16, pl. 188. 1922, fide Duc (306) 1951] FLORA OF PANAMA (Leguminosae ) A Tree to 10 m., branchlets glabrous to pubescent. Leaves glabrous, 1- to 3- foliolate; stipules setaceous; petiole with rachis 1—4 cm. long; rachis narrowly winged, auriculate at least at articulation of terminal leaflet; leaflets ovate to ovate- elliptic, rounded or cuneate at the base, obtusely short-acuminate, subcoriaceous, lustrous, the terminal leaflet 5—10 cm. long. Inflorescence axillary or terminal; the short slender, glabrous peduncles with 2—4 flowers; pedicels filiform, 1—1.5 cm. long; bracts small, setaceous; buds glabrous, subglobose, hardly 4 mm. in diameter. Flowers l-petalate; petal orbicular, unguiculate, a little longer than the calyx; stamens 18—20, almost all equal, twice longer than the calyx; anthers ovate; ovary stipitate, narrow, glabrous, 5- to 6-ovulate, attenuate to a short style; stipe a little shorter than the calyx. Legume short-stipitate, obliquely ovoid, long-acuminate, 4—5 cm. long, thick and carnose; seed obliquely ovoid, the aril lacerate. Panama (fide Seemann) ; northern South America from Colombia to Brazil. No Panamanian specimen of S. arborescens has come to our notice. Possibly the species was incorrectly reported from Panama by Seemann (Bot. Voy. Herald, 112. 1853), as S. triphylla, and does not really exist there. The above description is after Pittier (in Jour. Wash. Acad. 11:157. 1921). 4. SWARTZIA SIMPLEX (Sw.) Spreng. var. darienensis (Pittier) Schery, comb. nov. Swartzia darienensis Pittier, in Jour. Wash. Acad. 11:159. 1921. Swartzia myrtifolia Pittier, loc. cit. 158. 1921 (in part), not S. myrtifolia J. E. Smith, in es, Сус]. 34:no. 5. 1819, fide Wn. ?Swartzia trifolia Pittier, тос. E 158. 192 Tounatea subcoriacea Britt. in N. Am. Fl. 55 345. 1930. Tounatea cuneata Britt. loc. cit. 346. 1930. Trees 15 m. tall, upper branches glabrous, prominently lenticellate- dotted. Leaves glabrous, 1- to 5-foliolate; stipules linear, 1-5 mm. long; petiole narrowly obsagittate-winged, 0.5—3 cm. long, gross and terete near the axil, wing expanded and auriculate at insertion of petiolule(s); rachis, if present, similarly alate and more prominently so, wing at juncture of terminal leaflet 1.5-3 mm. wide, auricled; leaflets elliptic, abruptly acute or bluntly acuminate apically, cuneate to subobtuse basally, membranaceous or submembranaceous, prominently reticulatc-veined; lateral leaflets (if present) usually about 4-8 cm. long and 2-5 cm. broad; terminal leaflet 6—20 cm. long and 3-7 cm. broad; petiolules 2 mm. or less long, terete. In- florescence 3- to 8-flowered, bracteate, axillary or (less often) terminal on upper branchlets; bracts bidentate, dentae linear-lanceolate, about 1 mm. long; peduncles 1-6 cm. long, subterete, lightly pubescent; pedicels glabrous, 0.5-1.5 cm. long; buds globose, up to 8 mm. in diameter. Flowers 1-petalate, petal yellow, cordate- suborbicular, about 23 mm. wide; calyx splitting into 3—4 irregularly ovate sec- ions up to 1 cm. long, scurfy within, glabrous without; stamens many; anthers bilocular, basally versatile, larger ones about 2 mm. long, with conspicuous dark connective; filaments glabrous, about 10 of them grosser and longer than the others, up to 2 cm. long; ovary stipitate, stipe 4—7 mm. long, up to 13-ovulate; (307) [Vor. 38 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN style glabrous, about 6 mm. long; stigma obscurely bilobate. Legume glabrous, obliquely ovoid, caudate-beaked, apparently 2-valved, about 3.2 cm. long, 1.6—1.8 cm. broad and 1.2-1.6 cm. thick; seed 1, reniform, conspicuously arillate, about 2.5 cm. long. Endemic to Panama. ZONE: Mamei Hill, Pittier 3800 (TYPE). DARIEN: vicinity of La Palma, Pittier 16676; near mouth of Río Yapé, Allen 323. PANAMA: vicinity of Campana, Allen 2145; Taboga Island, Macbride 2801. PEARL IsLANDs: Trapicho Island, Allen 2621. The synonyms listed for S. simplex var. darienensis are mostly so interpreted from description analysis evaluated in the light of study of what specimens are available. In any group as polymorphous and inconstant as the "S. myrtifolia” group appears to be, it seems unjustifiable to base specific delimitation upon char- acters of rachis-wing venation, minor differences in leaf size and shape, distinction between subcoriaceous and membranaceous leaves, etc. Specimens seen show inter- gradation on all such characters, often on the same sheet. Possibly complete monographic study would show S. simplex var. darienensis to be synonymous with some other name, perhaps older than the Pittier publication of S. darienensis. However, for the present there seems no practical alternative but to accept the Pittier name, reduced to varietal status, as inclusive of several newer Britton species, as well as (doubtfully) Pittier’s S. trifolia. It is probable that S. ¢rifolia should be listed as a separate variety of S. simplex, but inasmuch as the type is without flowers no such step is here taken. A clear-cut delimitation between S. simplex var. darienensis and $. simplex is virtually impossible. 5. SWARTZIA SIMPLEX (Sw.) Spreng. Syst. Veg. 2:567. 1825. Possira simplex Sw. Prodr. Veg. Ind. 82. 1788. Rittera grandiflora Vahl, Eclog. 2:37. 1798 (see Excluded Species). Swartzia simplicifolia Willd. Sp. Pl. 2:1219. 18 Swartzia grandiflora (Vahl) Willd. Sp. РІ, Ө 1800 e UN Species), Tounatea simplex (Sw.) Taub., in Bot. Centralb. 47:391. 18 Tounatea penomenensis Britt. N. Am. Fl. 23:343. 1930. Tounatea gatunensis Britt. loc. cit. 344. 1930. Tounatea Williamsii Britt. loc. cit. 345. 1930. Tounatea Hayesii Britt. loc. cit. 345. 1930. Small tree to 10 m. tall, branchlets glabrous. Leaves 1-foliolate, stipulate, glabrous; petiole 2-20 mm. long, terete (especially basally) to narrowly alate- auriculate (especially apically); leaflet usually subcoriaceous, elliptic to ovate- lanceolate, bluntly acute to acuminate apically, rounded to cuneate basally, 4-20 cm. long, 2—8 cm. broad, with chief lateral nerves subparallel but confluent mar- ginally; petiolule about 1 mm. or less long, terete. Inflorescence 2- to 6-flowered, axillary or terminal, up to 10 cm. long, with axis lightly pubescent to glabrous; pedicels 5—20 mm. long, bearing globose buds about 8 mm. in diameter. Flower yellow, 1-petalate; petal irregularly orbicular, usually about 3 cm. tall and at least (308) 1951] Fig. 107. Swartzia simplex as broad, claw about 5 mm. long; stamens of 2 types, 8—12 larger longer ones at least 2 cm. long, and many shorter smaller ones less than 1.5 cm. long; anthers oblong, truncate, basally versatile, larger ones about 2 mm. long, smaller ones about 1 mm. long; ovary arcuate, long-stipitate, glabrous. Legume sigmoid-ovoid to asymmetrically oblongoid, subterete, up to 6 cm. long, attenuate-beaked, usually 1-seeded; seed prominently arillate, oblong-ovoid. Central America and West Indies. BOCAS DEL токо: Chiriqui Lagoon, von Wedel 1400. CANAL ZONE: Ancon, Piper 6024; Barro Colorado Island, Shattuck 808, Standley 41013, 40839; Fort Kobe Road, Allen 1888; Victoria Fill, Allen 1710. сосі.Е: Penonomé, Williams . COLON: Tumba Vieja, odge, Steyermark & Allen 16925. DARIÉN: Marraganti, Williams 005. PANAMÁ: Pacora, Bro. Paul 333; Chepo, Kluge 2. saN BLAs: Perme, Cooper 650. This species is listed as S. sim plex following more or less the concept of Pittier (in Jour. Wash. Acad. 11:157. 1921). Several of the Britton species are listed as synonyms following Pittier and from description analysis, without the types having been seen. Again S. simplex may not be the correct name, but serves as a con- venient catch-all until monographic study can determine more precisely the specific limits of certain ill-defined sections of the genus. The species as here considered (309) [Vor. 38 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN encompasses a number of intergrading and indefinite forms. It likewise grades into S. simplex var. darienensis, and some names and specimens that possibly could refer to the latter are included here. EXCLUDED OR DUBIOUS SPECIES SWARTZIA MYRTIFOLIA J. E. Smith in Rees, Cycl. 34:no. 5. 1819, was con- sidered by Pittier (Jour. Wash. Acad. 11:158. 1921) as occurring in Panama. Britton, however, believed plants referred by Pittier to this name to be a new species (Tounatea cuneata), which is here treated as a synonym of $. simplex var. darienensis. On the basis of plants seen and Britton's conclusion that the true S. myrtifolia does not occur in Panama, I hesitate to list S. sim plex var. darienensis as synonymous with the older West Indian S. myrtifolia. SWARTZIA GRANDIFLORA Willd. Sp. Pl. 2:1220. 1800, is referred to by Hemsley (Biol. Centr.-Am. Bot. 1:322. 1879-88) as occurring in Panama. However, neither Pittier (in Jour. Wash. Acad. 11:155—60. 1921) nor Britton and Rose (N. Am. Fl. 23:342-49. 1930) mention this name, even as a synonym, and Britton and Killip (in Ann. N. Y. Acad. Sci. 35:192. 1936) record it by name doubtfully in Colombia. “Index Kewensis’ (after? Benth. in Martius, Fl. Bras. 152:18. 1870) regards S. simplex as a synonym of S. grandiflora, and Vahl's description (as Rittera grandiflora) notes the great similarity of R. grandiflora and R. simplicifolia (= S. simplex). There seems little doubt but that S. grandiflora should be con- sidered a synonym of S. simplex. 2. BAUHINIA L. BauHInia [Plum.] L. Sp. Pl. 374. 1753 (originally in L. Gen. ed. 1:126. 1737). Pauletia Cav. Ic. 5:5, t. 400, 410. 1799. Amaria Mutis ex Caldas, in Seman. Nuov. Rein. Gran. 2:25. 1810. Schnella Raddi, in Mem. Soc. Ital. Modena 18:411. 1820. Lacara Spreng. Neue Entdeck. 3:56. 1822 Casparea HBK. Nov. Gen. & Sp. 6:317. 1823. Caulotretus Rich. ex Spreng. Syst. 4:Cur. Post. 406. 1827. Perlebia Mart. Reise 2:555 | 838. Alvesia Welw. Apont. (587:п. 47. 1858), fide Ind. Kew. Caspareopsis Britt. & Rose, in N. Am. Fl. 23:217. 1930. Other synonyms occur for Baubinia. Shrubs or small trees, or more generally vine-like and climbing, supported by other vegetation, unarmed or less frequently armed; trunk or stem often flattened, usually with hard wood and longitudinally striate bark. Branchlets with con- spicuous alternate nodes, often appearing somewhat zigzag-jointed, sometimes tendriled. Leaves diagnostic, inconspicuously caducous-stipulate, petiolate, simple but usually profoundly 2-lobed or sometimes 2-foliolate, rounded to cordate basally, conspicuously callused at insertion of leaf and petiole, bilobed apically, lobes more or less lanceolate. Inflorescence terminal or axillary near end of the branchlets, few- to many-flowered. Flowers usually whitish, conspicuous; calyx 5-parted, (310) 11 1951] FLORA OF PANAMA (Leguminosae) with a short or long conspicuous tube, the limb often spathaceous; petals 5 free, small or large, mostly unequal, clawed; fertile stamens 10 or 5 or in introduced species 1 or 3; anthers versatile or subversatile, usually sagittate basally; ovary usually stipitate or substipitate; legume compressed, elastically dehiscent. Old and New World tropics. 20—30 cm. long; inflorescence not known 13. B. MANCA lon . Leaves m. large, aa. Leaves small to large; usually 5—15 cm. lon Calyx үнер tubular, limb vubspathacesus or with elongate reflexed lobes nthesis. Creat la t 5 stamens Ir ur native plants. d. реді linear- lemen tou Plants armed; leaves dial. with рна ae calyx in bud ery narrow- -tubular, less than in the upper portion 1. B. PAULETIA E- ee. o unarmed; leaves larger, with acute lobes; calyx in stockier, at z t 4 mm. wide in upper рогїїоп...... e bud dd. Petals broader, epee to sp 2. B. UNGULATA tulat KE Dear із cleft, 7- to 9- ж. 3. B. EMARGINATA to e. Twigs armed; lea ee. Twigs ibd: ука larger, shallowly cleft, 11- nerved 4. B. LIGULATA cc. One or 3 stamens лил dea intro cultivated plants. st 5. B. PURPUREA r with оя ntherifero 6. B. MONANDRA d. Fl dd. Flower with 1 antheriferous stam o oblongoid, not | dodi but toothed or ncate = anthesis. mall, calyx never over 1 cm. preety to deeply cleft. . Calyx teeth prominent, erect, lanceolate to subulate. e. Leaves dull, often appressed- рота t above, the lobes blunt; erulen subtome long; leaves entire but B. STANDLEYI ee. Leaves shiny-glabrous above, the lobes acute; young twigs rufous-subhirsute or almost glabrous. н sate oe calyx about 8 mm. long or longer; oung twigs pubes o glabrate . B. CUMANENSIS Le Leaves E irs a ‘heir length; calyx about 6 mm. long; ung twigs rufous 9. В.5токкп dd. Cd. rest dns expan a ке ыг or minute. e. Calyx teeth rigidly reflexed, 5 . long, obovate-attenuate; br зс апа petiole dark red, сбн 10. B. REFLEXA ee. Calyx teeth not reflexed, vate or var Жы: branchlets and per ine oe, Ты Ыш! to subglabro Lh ida teeth manifest, obovate T. B. OBOVATA B. ЕХСІЅА ff. x teeth minute, calyx undulate to subtrunca сс. Flowers ioe calyx 1.5 cm. long or longer; deca suis. 2-folio d. onger than 1 cm.; antheriferous stamens 10...... IM ‘pedicels > ; antheriferous stamens 5...... 1 14. B. EUCOSMA dd. Mature pedicels shorter than 1 cm. 1805. 15. B. HYMENEAEFOLIA 1. Влонгміл Pautetia Pers. Syn. Pl. 1:455. Pauletia aculeata Cav. Ic. 5:6, pl. 410. 1799 Baubinia spinosa Poir. in Lam. Encycl. Supe. 1:599. Baubinia ite ip Spreng. Syst. Veg. 2:334. 25 к; parvifolia Seem. Bot. Voy rer vee 185277, non Hochst., fide Ind. Kew. Baubinia cblorantba Brandeg. in Zoe 5:200. Baubinia longiflora Rose, in Contr. U. S. Nis “Hach. 10:97. 1906. Shrub or small tree to 6 m. tall, branchlets pubescent, armed with stout prickles at the base of the petioles. Leaves ovate-orbicular, up to 6 cm. long and broad, pubescent to glabrous (especially above), rounded to truncate basally, cleft apically 1810. (311) [Vor. 38 I» ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 108. Bauhinia Pauletia up to 1⁄2 the length of the leaf, lobes rounded-obtuse; petioles short, pubescent, about 1 cm. long, callused at insertion of leaf; stipules linear. Inflorescence terminal, racemose, up to 25 cm. long. Flower large, greenish, up to 10 cm. long; calyx elongated spathaceous, tube about 1.5 cm. long; petals linear-elongated, up to 10? cm. long; fertile stamens 5, about 10 cm. long, with 5 smaller, narrower staminodes alternate with them; anthers linear, subbasally attached, sagittate basally, short-acuminate apically, about 2.5 cm. long or longer; ovary stipitate, pubescent; legume linear, compressed, long-stipitate, pubescent, up to 25 cm. long and 1.5 cm. wide. (312) 1951] FLORA OF PANAMA (Leguminosae) 13 Mexico to northern South America. CANAL ZONE: vicinity of Miraflores Lake, Pittier 2202, P. White 268. HERRERA vicinity of Chitré, Allen 1086. PANAMA: Juan Diaz, Standley 30408; Las Sabanas, Standley 25841, 31801; Matias Hernandez, Pittier 6801, Standley 28014. PROV. UN- KNOWN: without locality, Seemann 223. 2. BAUHINIA UNGULATA L. Sp. Pl. 374. 1753. Pauletia inermis Cav. Ic. 5:6, pl. 409. 1799 Bauhinia inermis (Сау.) Pers. Syn. Pl. 1:455. 1805. Baubinia Cavanillei Millsp. in Field Mus. Publ. Bot. 1:364. 1898. Unarmed shrub or small tree to 7 m. tall, branchlets brown-pubescent when young. Leaves ovate, glabrous above and pubescent below, conspicuously 9- to 11-nerved, up to 12 cm. long and almost as broad, cleft apically up to V5 their length, basally rounded to subcordate, lobes lanceolate, acute; petiole about 2 cm. long, calloused basally and at insertion of the blade. Inflorescence a terminal raceme up to 10 cm. long or longer, rufous-pubescent, with pedicels up to 2 cm. long. Flowers whitish, about 4.5 cm. long; calyx elongated, about 4.5 cm. long, subregular, tube about 1 cm. long; petals linear, elongated, about 3 cm. long; fertile stamens 10, up to 4 cm. long, in 2 series; anthers linear, basally sagittate, about 1 cm. long in bud; ovary stipitate, pubescent. Legume linear, lightly pubescent, stipitate, up to 20 cm. long and 1 cm. wide. Mexico to northern South America. снікюші: Gualaca, Allen 5061; San Felix, Pittier 5281. veracuas: headwaters Río Cafiazas, Allen 181. 3. BAUHINIA EMARGINATA Mill. Gard. Dict. ed. 8, no. 5. 1768. ?Baubinia rotundata Mill. Gard. Dict. ed. 8, no. 7. 1768, fide Britt. & Killip. Baubinia mollicella Blake, Contr. Gray Herb. 53:32. 1918, fide Britt. & Killip. Bauhinia mollifolia Pittier, Arbol. & Arbust. Venez. Dec. 6-8:88. 1927, fide Britt. & Killip. Prominently armed shrub or small tree, with young branchlets pubescent. Leaves small, orbicular-oblong, up to 6 cm. long and about as wide, more or less glabrous above, pubescent below, 7- to 9-nerved, truncate-subcordate basally, cleft apically about 15 their length, lobes ovate, blunt, rounded, more or less spreading, with a subulate apicule at the base of the cleft; petiole pubescent, up to 1.5 cm. long, callused only at insertion of the leaf; stipules linear, caducous. Inflorescence terminal or subterminal, pubescent, with stout buds on short pedicels. Flowers conspicuous, white; calyx-tube about 5 mm. long, limb spathaceous, up to 5 cm. long; petals obovate-spatulate, clawed, pinnate-nerved, 3—4.5 cm. long, up to 1.5 cm. broad; fertile stamens 10, about 3—4 cm. long, the alternating ones somewhat shorter; anthers linear, basally subsagittate, 6—7 mm. long; ovary stipitate, hirsute, with elongated style and truncate-capitate stigma. Legume not seen. (313) [Vor. 38 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama and northern South America. cocLÉ: La Venta, Muenscher 16310. PANAMA: Pacora, Woodson, Allen & Seibert 735 bis. Considerable uncertainty was experienced in selecting the name for the speci- mens cited. In general appearance they are much like B. albiflora Britt. & Rose of Salvador, but possess 10 fertile stamens whereas B. albiflora is described as with only 5. B. Schultzei Harms, of Colombia, is very similar but seems to have a more deeply cleft differing leaf. The older name, B. emarginata, was finally selected after comparison with some South American material, in the belief that this species in its broader sense would include the Panama specimens. Britton and Rose (М. Am. Fl. 23:203. 1930) do not list B. emarginata as occurring in Central America. 4. BAUHINIA LIGULATA Pittier, in Contr. U. S. Nat. Herb. 20:112. 1918. A large tree with unarmed, glabrate twigs. Leaves broadly ovate, 4-10 cm. long and 4—7.5 cm. broad, glabrous and somewhat shining above, pale and puberu- lent beneath, coriaceous, prominently 11- to 13-nerved, basally subcordate, apically cleft for only about 1% their length; petioles about 2 cm. long, sulcate; stipules minute, caducous. Inflorescence terminal or axillary-subterminal, racemose to somewhat paniculate, with ferruginous-pubescent buds. Flowers lilac, about 3 cm. long, with ferruginous-pubescent pedicels 2-6 mm. long; calyx-tube (and receptacular portion) obconical, about 7 mm. long, the limb splitting after flower- ing into 5 narrow, reflexed lobes about 14 mm. long, often more or less adnate; petals 5, ovate-elliptic, apically acute, basally attenuate, about 3 cm. long and 6 mm. broad, short-clawed, sinuate-margined; fertile stamens 10, 5 long and 5 short, free, glabrous; filaments incurved, up to 25 mm. long; anthers ovate-elliptic, about 5 mm. long; ovary essentially glabrous, stipitate, basally adnate to tube of recep- tacle and surrounded by 2 spathaceous ligules, 5- to 6-ovulate; style thick, the stigma papillose and somewhat 3- to 5-lobed. Panama. SAN BLAS: near Puerto Obaldia, Pittier 4334. Known only from the type (Pittier 4334) from the San Blas coast of Panama. Remarkable for its size, reported as a large tree up to 40 m. high and 80 cm. in trunk diameter. Named for the unusual ligules surrounding the base of the pistil. 5. BauHINIA PURPUREA L. Sp. Pl. 375. 1753. Baubinia retusa Poir. in Lam. Encycl. Suppl. 1:599. 1810, fide Spreng. Bauhinia triandra Roxb. Fl. Ind. 2:320. 1832, fide Ind. Kew. Bauhinia platyphylla Zipp. ex Span. іп Linnaea 15:201. 1841. Introduced, unarmed, ornamental shrub, branchlets glabrous to lightly pubes- cent. Leaves broadly orbicular, cordate to truncate basally, usually prominently 9-veined, glabrous to lightly pubescent below, up to 13 cm. long and 16 cm. broad, shallowly cleft apically, lobes obtuse, rounded; petiole angled, callous-swollen apically and basally, up to 4 cm. long. Inflorescence terminal or subterminal, (314) 1951] FLORA OF PANAMA (Leguminosae) 15 Fig. 109. Bauhinia Standleyi several- to many-flowered. Flowers conspicuous, 3-4 cm. long; calyx scarcely spathaceous, tube up to 1 cm. long, limb up to 2.5 cm. long; petals clawed, spatulate-obovate, about 3.5 cm. long; fertile stamens 3, glabrous; anthers linear- oblong, versatile, 7 mm. long in bud; ovary long-stipitate, densely pubescent, with a truncate stigma. Legume smooth, linear, up to about 30 cm. long. Introduced to New World tropics from Asia. CANAL ZONE: Balboa, Steyermark s.n. (Jan. 7, 1935); Barro Colorado Island, Shattuck 176. 6. BAUHINIA MONANDRA Kurz, in Jour. Asiat. Soc. Bengal 427:73. 1873. Bauhinia Kappleri Sagot, in Ann. Sci. Nat. Bot. VI, 15:317. 1882. Bauhinia Krugii Urban, in Ber. Deut. Bot. Ges. 3:83. 1885. Caspareopsis monandra (Kurz) Britt. & Rose, in N. Am. Fl. 23:217. 1930. Small cultivated tree with young branches lightly pubescent. Leaves ovate- orbicular, up to 20 cm. long and almost as wide, chartaceous to subcoriaceous, glabrous above, pubescent on veins below, basally cordate to truncate, apically cleft up to 14 the length of the leaf, lobes blunt, rounded; petiole lightly pubescent, up to 6 cm. long, with bilobate callus at insertion of the leaf. Inflorescence a terminal few-flowered raceme. Flowers large, showy; calyx about 3 cm. long, (315) [Vor. 38 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN spathaceous, pubescent without, tube (including pedicellar part) slender, 2-2.5 cm. long; petals obovate-oblanceolate, 4—5 cm. long, uppermost maculate; fertile stamen 1, arcuate, about 4 cm. long, other stamens rudimentary; anther linear, versatile, sagittate, about 5 mm. long; fruit reported linear, flat, up to 22 cm. long. Naturalized in the West Indies and northern South America; native to India. No specimens are recorded from Panama; the plant is reported in Colombia and likely may be cultivated in Panama as well. 7. BAUHINIA SrANDLEYI Rose, in Jour. Wash. Acad. 17:166. 1927. Schnella Standleyi (Rose) Britt. & Rose, in N. Am. Fl. 23:206. 1930. Large tendrilled vine, sometimes armed basally, with pubescent branchlets. Leaves broadly ovate, 3-9 cm. long and broad, prominently 9-nerved, lightly appressed-pubescent above and more heavily so below, subcordate basally, notched apically for about 44-2 their length, with a subulate apicule at the base of the notch, both inner and outer margins of the lobes rounded; petiole terete, pubescent, 2-4 cm. long, callused at insertion of leaf, caducous-stipulate. Inflorescence terminal or subterminal, up to 10 cm. long, pubescent, linear-bracteate. Flowers whitish, 15-17 cm. long; calyx pubescent, more or less bilabiate, subulate teeth (2 above, 3 below), about 2.5 mm. long, tube 4-6 mm. long; petals oblanceolate, up to 17 mm. long and 7 mm. wide, hirsute within basally, smaller petal maculate; fertile stamens 10, glabrous, alternate ones longer (6 mm. long) and with thicker filaments; anthers ovate, versatile; ovary setose-hirsute. Legume spatulate, 6—7 cm. long. Panama and Costa Rica. ZONE: Miraflores Lake, P. White 269; Palo Seco, Allen 2896; Ма: Fill near елімде со Allen 1711; without ары, Seemann 222. COCLÉ: nomé, Williams j4. PAN vicinity of Pacora, Allen 1125; near Panamá, Standley pum ; near Punta ЖАИ Standley 26247; Tabosa Island, Macbride 2800. This species apparently differs little from B. cumanensis HBK. (?B. glabra Jacq.). 8. BAUHINIA cumanensis HBK. Nov. Gen. & Sp. 6:521. 1824. Bauhinia columbiensis Vogel, in Linnaea 13:313. 1839, Ar DN Schnella brachystachya Benth. in Hook. Jou p? ria 2:9 Bauhinia brachystachya Walp. Rep. 1:852. Schnella columbiensis (Vogel) Benth. Bot. Voy. СЕ 89. 1844 ?Schnella heterophylla Benth., Griseb. Cat. Pl. Cub. 81. 1866, non B. beterophylla HBK. A scandent, usually tendrilled vine, the twigs pubescent to glabrate. Leaves ovate-orbicular, cordate, deeply lobed apically, 2-12 cm. long and almost as wide, glabrous and shining above, puberulent and pallid below, the lobes acutish, spread- ing; petioles up to 6 cm. long, glabrous. Inflorescence terminal, racemose, several- flowered, linear-bracteate. Flowers whitish; calyx pubescent, the tube about. 8 mm. long, 10-ridged, the teeth linear, 4-6 mm. long; petals oblanceolate, 16-25 mm. long, rounded at apex, pubescent without; stamens up to 12 mm. long in 2 (316) 1951] FLORA OF PANAMA (Leguminosae ) 17 series; ovary densely brown-lanate; style short, glabrous; legume broadly oblong, 6-10 cm. long, 2-2.5 cm. wide, glabrate, short-stipitate, few-seeded. Panama and northern South America to Venezuela and the Guianas; West Indies. Hemsley (Biol. Cent.-Am. Bot.) records the species (as B. columbiensis) from Coiba Island, Panama; Seemann (Bot. Voy. Herald) records it (as S. columbiensis) from “mouth of the Rio Grande de Panama”; Bentham (Bot. Voy. Sulphur) lists it by the same name from “Panama.” Neither authentic specimens of B. cuman- ensis nor B. columbiensis have been seen. Bentham’s judgment (Martius, Fl. Bras. 152:212. 1874) that B. columbiensis is synonymous with B. cumanensis 15 accepted for convenience, even though Britton and Killip (Ann. N. Y. Acad. Sci. 35:163. 1936) consider it distinct, although possibly the same as B. glabra Таса. В. cumanensis is included in this Flora only upon the basis of the references cited, and whether or not it really occurs in Panama as B. cumanensis is impossible to say at this time. The above description is taken from the original and that appearing in the ‘North American Flora.’ 9. Влоніміл Storkii (Rose) Schery, comb. nov. Schnella Storkii Rose, in М. Am. Fl. 23:206. 1930. Tendrilled vine, branchlets densely rufous-pubescent. Leaves ovate-orbicular, 4—9 cm. long and 4—8 cm. broad, glabrous above, pubescent beneath, especially along the nerves, prominently 11-nerved, deeply cordate basally, obcuneate-notched apically for not more than М the length, with a subulate apicule about 5 mm. long at base of the notch; lobes bounding notch cuneate-lanceolate to bluntly obtuse, rounded on outer margin, straighter on inner margin; petiole terete, rufous-tomen- tose, 2-3 cm. long, with a bilobate callus at insertion of the leaf; stipules linear, caducous. Inflorescence a terminal, several-flowered raceme, 2—6 cm. long, linear- bracteate, rufous-tomentose. Flowers pale pink, 12—15 mm. long; calyx pubescent without, more or less bilabiate, lower 3 and upper 2 dentae subulate, about 1.5 mm. long, tube 4—5 mm. long; petals linear-oblanceolate, 1.2-1.5 cm. long, up to 4 mm. wide, hirsute basally within, uppermost linear, smaller, maculate; stamens 10, all fertile alternate ones longer, 5 mm. vs. 4 mm. long; alternate filaments wider; anthers ovate-orbicular, versatile, bilocular; ovary setose-hirsute, short. Western Panama. BOCAS DEL TORO: Н. von Wedel 487. PROV. UNKNOWN: "western Panama", Stork 10. BauuiwrA reflexa Schery, sp. nov. Frutex scandens inermis ramulis rufo-hirsutis; foliis orbicularibus usque ad 12 cm. longis latisque, 11—13-nervatis supra glabris subtus appresso-pubescentibus, base cordatis apice lobatis ad %—М; longitudinem folium, lobis margine exteriore rotundatis margine interiore rectis apice brevi-acuminatis inflexis; petiolis sub- hirsutis 3—7 cm. longis; inflorescentiis racemosis terminalibus dense pubescentibus, rachibus conspicue bracteatis, bracteis linearibus acuminato-attenuatis ca. 1 cm. (317) [Vor. 38 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN longis; calyce campanulato exteriore dense pubescente 15-nervato coriaceo, ca. 8 mm. longo latoque, dentibus ligulato-oblanceolatis reflexis valde coriaceis, ca. 5 mm. longis, terminaliter attenuatis; petalis obovatis ca. 2 cm. longis, exteriore setoso-pubescentibus, base carnosis angustatis prominente biauriculatis, auriculis 1.5 mm. longis; staminibus fertilibus 10, 5—6 mm. longis, filamentis glabris, antheris 171.5 mm. longis; ovario sessili setoso-hirsuto pauci-ovulato, stigmate obliquo. 9, a GS vi aes qeu ч (7 Ж S S $, утә. i / f { Fig. 110. Baubinia reflexa Unarmed woody vine, with subhirsute (young) branchlets. Leaves orbicular, 5-12 cm. long and broad, prominently 11- to 13-nerved, glabrous above and con- spicuously reticulate, somewhat appressed-pubescent below (hirsute on principal veins), cordate basally, lobed apically for about V4—!4 their length, with a minute caudiform apicule 2—3 mm. long at base of the cleft, lobes rounded on the outer margin, straight on the interior, with acute to short-acuminate tips somewhat in- flexed; petiole subhirsute, 3-7 cm. long, inconspicuously callous-rugose apically and (318) 1951] FLORA OF PANAMA (Leguminosae ) 19 basally, with a typical bilobate callus at insertion of the leaf; stipules caducous. Inflorescence a terminal raceme, densely pubescent, rachis conspicuously bracteate, bracts subpersistent, linear, attenuate-acuminate, about 1 cm. long, lightly pubes- cent. Flowers pink, small, congested; calyx campanulate, densely pubescent with- out and very dark-hirsute-setose at insertion of pedicel and on main longitudinal ridges, glabrous within except at the base, thickly coriaceous, 15-ridged, about 8 mm. tall and broad, sinuately 5-lobed, each lobe about 1.5 mm. long and bearing a conspicuous reflexed fleshy tooth about 5 mm. long, the teeth pubescent, ligulate- oblanceolate, attenuate apically, about 2 mm. broad at insertion with lobe and bearing there on each margin a conspicuous fleshy horn about 0.5 mm. long; petals obovate, about 2 cm. long, densely setose without, especially laterally and near the middle, within only on the claw, claw thick-fleshy, linear, about 9 mm. long and 2 mm. wide, internally densely brown-setose on the margin and thus ridged down the middle, glabrous at the base, with a fleshy ridge towards either margin sur- mounted by a prominent brown-setose auricle about 1.5 mm. long; stamens 10, all antheriferous, those opposite the petals smaller, about 5 mm. long, those alter- nate with the petals larger, about 6 mm. long; filaments fleshy, glabrous; anthers small, bilocular, ovate-lanceolate, about 1.5 mm. long on larger stamens and 1 mm. long on smaller stamens; ovary sessile, setose-hirsute, few-ovulate, with an oblique, cleft, more or less sessile stigma. Legume unknown. This species is especially marked by the leaf characters; the condensed, promi- nently bracteate inflorescence; the unusual calyx teeth; the auricled petal-claw; the subsessile ovary and stigma; and the long red-brown general pubescence. It is close to B. Storkii and B. obovata. Panama. ANAL ZONE: Barro Colorado Island, Kenoyer 377» apo 994, Woodworth & Vestal x vicinity Salamanca Hydrographic Station, Woo Allen & Seibert 1623 epica Bot. Gard., TYPE); ?Rio Chagres between Río Indio & "Río Pequeni, Steyermark 9 AI 16762 (sterile). 11. BAUHINIA OBOVATA Blake, in Jour. Wash. Acad. 14:286. 1924. Schnella obovata (Blake) Britt. & Rose, in N. Am. Fl. 23:207. 1930. Unarmed vine, branchlets rufous-puberulous, soon glabrous. Leaves broadly ovate, up to 8 cm. long and as broad or broader, appressed-pubescent below, gla- brous above, subcoriaceous, prominently 9- to 11-nerved, subcordate, bilobed for about 14 their length, with a minute apicule at base of cleft, lobes lanceolate, shortly acuminate-tipped, tips incurved; petioles up to 4 cm. long, pubescent, somewhat rugose-calloused apically and basally, with a prominent callus at insertion of leaf. Inflorescence racemose, dense, many-flowered, rufid-puberulous, up to 8 cm. long; bracts obovate, about 4 mm. long. Flowers with bibracteate pedicels 6—9 mm. long; calyx campanulate, densely pubescent, up to 10 mm. long, teeth obovate, about 3.5 mm. long; petals obovate, clawed, rufous-pilose without, about (319) [ Vor. 38 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN 12 mm. long; antheriferous stamens 10, unequal; filaments glabrous; ovary sessile, densely rufous-pilose; stigma small, oblique. Panama. DARIEN: Sambi River, Pittier 5568. 12. BAUHINIA EXCISA (Griseb.) Hemsl. Biol. Centr.-Am. Bot. 1:337. 1880. Schnella excisa Griseb. Fl. Brit. W. Ind. 214. 1860. Baubinia Tbompsonii Johnston, in багш: 8:140. 1949. A glabrescent vine, the old stem often flattened and perforated, branchlets puberulous. Leaves subcoriaceous, ovate-orbicular, up to 20 cm. long and almost as wide, glabrous above, minutely pubescent below, basally shallowly cordate, apically cleft to V4 or more the length of the leaf, with a small apicule at base of the cleft, lobes bluntly acuminate-acute, the tips somewhat incurved; petiole up to 6 cm. long, basally and apically swollen and callous-rugose, with a bilobate callus at insertion of the leaf. Inflorescence racemose, many-flowered, with pedicels 1-4 mm. long. Flowers moderate; calyx pubescent, campanulate, ventricose in bud, subtruncate or broadly undulate, 5-6 mm. long and as broad, with 5 minute or shallow teeth; petals short, oblong, obtuse, 15 mm. long, externally villous; stamens 10, almost 1 cm. long; anthers about 1 mm. long; ovary sessile, villous; legume 6-7 cm. long, 2-2.5 cm. wide, apiculate, short-stipitate, compressed, arcuate. Panama; Trinidad. AS DEL TORO: ?Changuinola Valley, Dunlap 337 (mutilated: fruit only). САМАТ. ZONE: Balboa, Gillespie P-26. DARIÉN: La Palma, Pittier 5500. PEARL ISLANDS: San José, Anderson s. n. (1852). Exact application of this name (from Trinidad type) to Panamanian material awaits monographic study. Dr. I. M. Johnston is certain that the name B. excisa does not apply to the Panamanian citations, and indeed comparison with the orig- inal description of Grisebach would tend to support this view. Dr. Johnston has proposed the name B. Thompsonii to include the Panamanian entity. However, Broadway 2218, from Trinidad, is almost identical with the Panamanian material. Other specimens, likewise nearly identical, including some from Panama, have in the past been determined at various herbaria as B. excisa. Very similar also are B. platycalyx Benth. (to judge from herbarium material so determined), B. umbri- ana Britt. & Killip, B. breviloba Ducke (which is very similar to Panamanian material), and B. sericella Standl. In Johnston's several collections from San Jose Island of the Pearl Islands of Panama, considerable variation has been noted in the depth of the terminal notch or split of the leaf in what are obviously plants of the same species. Likewise leaves from the same plant may have differing numbers of prominent veins (1.е., either 9 or 11). Also variation in floral and leaf pubescence and in other floral characters can be noted. Perhaps Grisebach, in writing his original description, siezed upon a rather atypical form for the type. He did men- tion with the original description that the same species was known from Panama. (320) 1951] FLORA OF PANAMA (Leguminosae) 21 Until examination of the type is possible, along with abundant material from northern South America where the complex including the Panamanian entity seems to center, it is perhaps wisest to retain the name B. excisa. 13. BAUHINIA MANCA Standl. in Field Mus. Nat. Hist. Bot. 18:511. 1937. Unarmed, cirriferous, climbing vine, branchlets more or less angled and pubes- cent. Leaves very large, to 30 cm. long, prominently 11- to 13-nerved, glabrous or pubescent below, subcoriaceous, when dry greenish above and brownish below, basally rounded-subcordate, deeply bilobed or essentially 2-foliolate, lobes lanceo- late, up to 12 cm. wide, typically long-acuminate apically but often otherwise on same plant; petiole angled, about 12 cm. long, callous-thickened apically and basally. Inflorescence unknown. Costa Rica and Panama. Standley has described this species on vegetative characters alone, listing Frost 112 and Bailey 278 of Barro Colorado Island, Canal Zone, as probably conspecific with it. The species is distinct in the unusually large leaves, but exact specific delimitation and relationship is still in doubt. Fig. 111. Bauhinia eucosma 14. BauHINIA EUCOsMA Blake, in Jour. Wash. Acad. 14:286. 1924. Schnella eucosma (Blake) Britt. & Rose, in N. Am. Fl. 23:207. 1930. Large tendrilled vine, with terete, glabrous stems. Leaves so profoundly bi- lobed as to be 2-foliolate, subchartaceous, glabrous except occasionally strigose below especially on veins, lobes ovate-lanceolate, up to 10 cm. long and 4.5 cm. wide, prominently 5-nerved, rounded basally, bluntly acute or obtuse apically, basa! (321) LIB RARY MEDICAL SCHOOL [Vor. 38 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN sinus deep and narrow, apical sinus with a subulate apicule at base extending from the lower surface of the petiole and leaflets; petiole slender, glabrous, 3-6 cm. long, callous-rugose at base; stipules caducous. Inflorescence terminal, racemose, glabrous to lightly strigose, up to 10 cm. long or longer, bibracteate at base of pedicels. Flower white, fragrant; calyx large, campanulate, prominently longi- tudinally (15) ridged, 2-2.5 cm. long, with subulate teeth 3-4 mm. long; petals large, obovate, short-clawed, up to 5 cm. long, strigose without and on the claw within; antheriferous stamens 10, in 2 series, about 15 mm. long. Legume sessile, oblong to oblong-obovate, lightly strigose, up to 13 cm. long and 4 cm. wide. Panama. AL ZONE: vicinity Miraflores Lake, С. White 150. PANAMA: Casa Larga, Allen 5068; Matias Hernandez, Pittier 6782. 15. BAUHINIA HYMENEAEFOLIA Triana, Hemsl. Diag. Pl. Nov. 48. 1880. Schnella bymenaefolia (Triana) Britt. & Rose, N. Am. Fl. 23:208. 1930. An unarmed vine, young branchlets puberulent. Leaves bifoliolate, the leaflets ovate to oblong, oblique, coriaceous, 5-7.5 cm. long, obtuse, glabrate, 5-nerved; petiole slender, glabrous, about 6 cm. long. Inflorescence racemose, terminal or lateral, few-flowered; pedicels short. Flowers large, white; calyx broad, 15 mm. long, 15-ridged, puberulent, lobes small, ovate-oblong; petals 3.5 cm. long, short- clawed, densely pubescent without; antheriferous stamens 5, free, alternating with 5 sterile ones; filaments glabrous; anthers barbate; ovary sessile, densely hirsute. Panama and Colombia? This species was described at Kew from the S. Hayes 635 specimens from Panama, a photograph of which has been examined. No other Panamanian speci- mens representing this species have been seen. Thus its separation from the pre- ceding species is maintained on the basis of its description, in which the species is listed as having only 5 antheriferous stamens. SPECIES OF DOUBTFUL OCCURRENCE IN PANAMA BAUHINIA SPLENDENS HBK. Nov. Gen. & Sp. 6:321. 1824. BAUHINIA SUAVEOLENS HBK. loc. cit. 320. 1824. Both are South American species, listed by Hemsley (Biol. Centr.-Am. Bot. 1:340) as occurring in Panama from reference to specimens in the Kew Herbarium. Not listed in more modern works as occurring in Central America nor Colombia. Several sterile specimens, evidently Bauhinia, have been collected in Panama. On vegetative characters alone these appear different from any of the species here listed. Probably several additional species will eventually be known from Panama. 3. POEPPIGIA Presl Poepricia Presl, Symb. Bot. 1:15. 1830. This genus, as represented by P. procera Presl, loc. cit., has not yet been re- ported from Panama, but is to be expected there. It is known from Colombia, Venezuela, Peru, Brazil, Cuba, northern Central America and Mexico. (322) 1951] FLORA OF PANAMA (Leguminosae) 23 4. MORA Schomb. Mora Schomb. ex Benth. in Trans. Linn. Soc. 18:210, /. IÓ. 1839. Unarmed, gregarious trees of low swampy areas. Young stems lenticellate. Leaves (in Panamanian species) simply conjugate-pinnate, short-petiolate, incon- spicuously or not stipulate, with few, large, coriaceous leaflets. Inflorescence spicate, dense. Flowers small; the calyx campanulate, with a short tube an spreading 5-lobed limb; petals 5, small, subequal, imbricate; stamens 5, opposite the petals, free, equal, alternating with 5 apically dilated staminodes; anthers ob- long, longitudinally dehiscent; ovary subsessile, free, few-ovulate, with a short style and smal! terminal stigma. Legume coriaceous to ligneous, compressed, 2- valved, with a large suborbicular seed. This genus has been conservatively listed under Dimorphandra (Schott, in Spreng. Syst. 4, Cur. Post. 404. 1827). However, most modern treatments of the Leguminosae consider it sufficiently distinct (chiefly on the basis of simply pinnate leaves and large seed) to merit separate generic rank, based on Bentham’s well-described type Mora excelsa. A single species is known from Panama. 1. Mora oLrirERA (Triana) Ducke, in Arch. Jard. Bot. Rio de Janeiro 4:45. 1925 Dimorphandra oleifera Triana ex Hemsl. Bot. Voy. p 3: s 1885. Dimorpbandra megistosperma Pittier, in Jour. Wash. Acad. 5:472. 1915. Mora megistosperma (Pittier) Britt. & Rose, in М. Am. . Fl. 214 1930. Erect tree to 45 m. high, with brown, glabrous branches. Leaves simply ріп- nate, bijugate; petioles and petiolules rugose-verrucose, the former 2—5 cm. long, the latter about 3 mm. long; rachis more or less plane above, terete below, 2.5-9.5 cm. long, glabrous; leaflets opposite, asymmetrically ovate to oblong-acuminate, 6—18 cm. long, 4-7 cm. wide, apically obtuse or acute, basally somewhat inequi- lateral, coriaceous, glabrous, reticulate, dull below. Inflorescence terminal or sub- terminal, densely spicate, 8-10 cm. long. Flowers white, sessile; calyx glabrous, with a short tube, the lobes orbicular, ciliate, unequal, 3—4 mm. long; petals oblong, attenuate basally, rounded-emarginate apically, about 6 mm. long and 2.5-3 mm. wide, marginally scarious-ciliate; fertile stamens 5, 7—8 mm. long, with thick filaments, the anthers lanose-barbate apically; ovary subsessile, lanose, 2- to 3-ovulate; style glabrous. Legume monoseminate, up to 25 cm. long and 13 cm. broad, glabrous, woody-coriaceous, dehiscent; seed up to 18 cm. long and 12 cm. broad. Panama and Colombia. DARIEN: Sumacate, Pittier 6593. PANAMA: La Capitana, Pittier 4582. An interesting species producing one of the largest Dicotyledonous seeds known. The ripe pods are said to twist open on the tree, letting the enormous seeds fall to the boggy soil below where they germinate almost immediately. Many trees have (9233 [Vor. 38 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN huge, buttressed trunks, with flat primary roots extending outward on top of the ground for as much as 15 m. The species was originally described from a Pana- manian collection of S. Hayes. 5. PHYLLOCARPUS Riedel PHyLLocarpus Riedel, Endl. Gen. Suppl. 2:97. 1842. Large, unarmed trees. Leaves once-pinnate; petiole and rachis eglandular; leaflets a few pairs, moderately large; stipules conspicuous, caducous. Inflorescence of several to many condensed racemes clustered on older branches. Flowers con- spicuous, prominently pedicellate; calyx 4-parted, polysepalous above the short receptacular base, strongly imbricate; petals 3, imbricate, the uppermost smaller and obscured; stamens 10, diadelphous by basal fusion of (9) filaments, the fila- ments free above; anthers versatile, longitudinally dehiscent; ovary free from the calyx; style expanded or clavate apically. Legume flattened, winged along upper suture, indehiscent. Northern Central America and Amazonia. A single species is found in Panama, sometimes cultivated but according to Paul H. Allen definitely indigenous to the Madden Lake forests. 1. PHYLLOCARPUS SEPTENTRIONALIS Donn. Sm. in Bot. Gaz. 55:433. 1913. Tree, the branchlets puberulent and often verrucose when older. Leaves mod- erate; petiole usually 1—2 cm. long, callous basally, canaliculate above, puberulent; rachis several cm. long, similar to petiole; stipules narrowly lanceolate-falcate, about 7 mm. long; leaflets 4—6 pairs, elliptic, mostly 6—8 cm. long and almost 3 cm. wide, acute (and tip blunt) or obtuse apically, rounded and inequilateral basally, glossy above, puberulent below especially on veins and marginally, finely reticulate. Racemes few-flowered, 2—3 cm. long, clustered on older wood; pedicels about 1 cm. long, bibracteate basally. Flowers red; calyx with a very short recep- tacular tube less than 1 mm. long; calyx-lobes 4, elliptic, about 8 mm. long, glabrous except ciliate marginally, coriaceous; petals 3, about as long as the sepals; stamens 10, ng ee filaments as much as 2.3 cm. long, the lowermost united basally for about 14 their length; anthers oblong, about 1 mm. long, bilocular; ovary short-stipitate, free, glabrous; style (with ovary) about 2 cm. long, clavate apically; stigma terminal. Legume (reported) oblong, 12-17 cm. long and 4.0—4.5 cm. wide, thin, 1- to 2-seeded, the wing from the upper suture about 1 cm. wide. Guatemala and Honduras? to Panama; introduced into Florida, the Canal Zone and elsewhere. mit (cultivated), Allen 2053; Tumba Vieja, Steyermark & Allen Nar ibis y ci Higgins 259. Reported to be an attractive ornamental, the abundant scarlet flowers rivaling in showiness those of the Royal Poinciana (Delonix regia). Paul H. Allen has the following to say about the occurence of this species in Panama: “The collection cited from Tumba Vieja was from wild trees which are quite common there in (324) 1951] FLORA OF PANAMA (Leguminosae ) 25 areas of climax rain forest. They average about 90 feet in height, and are very conspicuous in December when they are in flower. The Allen and Higgins col- lections are from cultivated plants." 6. CYNOMETAA L. (Reviewed by John D. Dwyer, Union University) CYNOMETRA L. Sp. Pl. 382. 1753. Iripa Adans. Fam. 2:508 There „1763; here occur two additional synonyms not concerning the genus in Central America. Unarmed forest trees. Leaves evenly pinnate; leaflets a single pair (in Ameri- can species), oblique, subcoriaceous, eglandular. Inflorescence racemose to fascicu- late, axillary. Flowers small; calyx essentially polysepalous, the 3—5 lobes reflexed from a small, central, disc-like receptacle and often caducous, imbricate in bud; petals 5, subequal to somewhat unequal, inserted with the stamens upon the recep- tacle; stamens 10, free, the filaments glabrous; anthers small; ovary sessile to short- stipitate, mostly free, 1- to 2-ovulate; style filiform; stigma terminal, truncate or capitate. Legume rather small, usually oblique or curved, coriaceous, swollen; seeds arillate. Mexico through Central America; West Indies; South America; Africa; Asia and Pacific islands. Only a single species has been reported from Panama, unconfirmed by recent listings or specimens. 1. CYNOMETRA BAUHINIAEFOLIA Benth. in Hook. Jour. Bot. 2:99. 1840. Cynometra crassifolia Benth. loc. cit. 100. 1840. Tree with slender puberulent branches. Leaves 1-jugate, short-petiolate; leaf- lets 2, subovate, 2.5—4 cm. long, up to 2 cm. wide, glabrous, submembranaceous, obtuse, somewhat inequilateral, 2- to 3-nervate, pubescent on the nerves. In- florescence fasciculate, axillary, short-pedunculate; bracts about 1 mm. long. Flowers pedicellate; pedicels pubescent, about 8 mm. long, ovate-bracteate basally; sepals 4, about 3 mm. long, membranaceous, deciduous; petals 5, contracted basally, unequal, inserted upon the disc-like receptacle; ovary pubescent; style glabrous, inflexed; stigma capitate. Legume small, plane-convex, fleshy, verrucose and pubescent without. Panama?; Guianas to Colombia and Peru, Brazil and Argentina. This species from a British Guiana type is reported by Hemsley (Biol. Centr.- Am. Bot. 1:342) as occurring in Panama, based upon a specimen (5. Hayes 20) at the Kew Herbarium. The genus is to be expected in Panama, having been re- ported from both Colombia and Costa Rica, but whether the name listed by Hemsley is strictly accurate is doubtful. The more recent "North American Flora' gives no Cynometra from Panama, and C. baubiniaefolia is neither listed from Costa Rica nor Colombia. No specimens of Cynometra from Panama nor of C. bauhiniae- folia have been examined, and the above description is taken from the original. (325) [Vor. 38 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7. COPAIFERA L. (Reviewed by John D. Dwyer, Union University) СорАІЕЕКА L. Sp. Pl. ed. 2, 557. 1762, nom. conserv. Copaiva Jacq. Enum. Pl. Carib. 65. 1760. Copaiba Adans. Fam. Pl. 2:341. 1763. There also exist three African synonyms for Copaifera. Unarmed trees up to 30 т. high, with glabrescent, distinctly lenticellate branchlets. Leaves simply pinnate, the petiole and rachis usually glabrous; leaflets one to many pairs, usually alternate except terminally, small, coriaceous to charta- ceous, more or less inequilateral and falcate, commonly glabrous, reticulate, frequently punctate, short-petiolulate. Inflorescence a terminal or subterminal panicle of multiflowered spikes, the axis minutely caducous-bracteate. Flowers small, apetalous, reportedly yellow-white; calyx 4-parted, polysepalous, basally somewhat disc-like, subvalvate in bud, subtending bracts caducous except in very young bud; petals lacking; stamens usually 10, free, glabrous; anthers relatively large, bilocular; ovary free, substipitate, biovulate. Legume short, somewhat oblique, compressed to turgid, 2-valved, 1-seeded; seed large, arillate. Panama, West Indies, South America; Africa. Center of New World distribu- tion in Amazon valley of Brazil. a. Leaflets generally 3-5 cm. long, short-acuminate; legume usually less than 2 cm. long aa. Leaflets averaging larger (about 7 cm. long), long-acuminate; legume em. long or 1олдег.................................................................................... 1. C. PANAMENSIS 2. C. AROMATICA 1. COPAIFERA PANAMENSIS (Britt.) Standl. in Trop. Woods 34:41. 1933. Copaiva panamensis Britt. in М. Am. Fl. 23:222. 1930. Unarmed trees, with lenticellate, glabrous branchlets. Leaves essentially glabrous, 6- to 12 (generally 6- to 8) -foliolate; petiole short; leaflets obliquely ovate or ovate-elliptic, 3—5 cm. long, short-acuminate apically, obtuse or rounded basally, coriaceous, markedly reticulate, with a prominent midvein and many con- spicuous lateral veins, usually alternate except terminally on leaf; petiolules flat- tened, rugose when dry. Inflorescence of many small flowers, as described for the genus. Flowers apetalous; calyx-lobes 4, narrowly ovate to elliptic or lanceolate, about 3 mm. long, subhirsute within, glabrous without, spreading; stamens usually 10, 7—8 mm. long; filaments slender, glabrous; anthers versatile, oblong, almost 2 mm. long; ovary suborbicular, flattened, hirsute marginally; style arcuate, up to 5 mm. long, bearing a capitate stigma. Legume orbicular or broadly ellipsoid, about 15 mm. broad, glabrous; seed almost enclosed by the aril. Panama. CANAL ZONE: vicinity of Miraflores Lake, P. White 257; R. Chagres, Steyermark & Allen 16791. сосі.Е: Nata, Allen 816; El Valle de Anton, Allen 2005; Penonomé, Wil- liams 50; La Pintada, Allen 3605. The species constitutes a source of the oleoresin "copaiba balsam", used medic- (326) 1951] FLORA OF PANAMA (Leguminosae) 27 inally and in the manufacture of varnishes. It is readily confused with C. officinalis, but has narrow-elliptic leaflets їп 3—6 pairs, whereas the latter has wide-ovate leaflets in 2-4 pairs. 2. CoPAIFERA AROMATICA Dwyer, in Trop. Woods 83:15. 1945. Tree to 30 m. tall, with aromatic bark. Leaves 10—30 cm. long, with glabrous petioles; leaflets 8—12, alternate or opposite, distinctly inequilateral, narrowly ovate-oblong or broadly oblong, 2.5-9 cm. long, 1.3-3.5 cm. broad, falcate- acuminate, apically attenuate and minutely retuse, basally obtuse or cuneate, gla- brous, often punctate, principal veins somewhat pubescent below. Flowers unknown. Legume (immature) obovate-rotund, about 3 cm. long and broad, obtuse apically and basally; seed obovate-oblong, 1.5 cm. long and 1.1 cm. broad, invested by a dark-red aril 1.7 cm. long. Panama. осі: Horconcitos, Pittier 5118; San Felix, Pittier 5263. DARIEN CHIRI : Pinogana, Pittier 6080. PANAMA: Bella Vista, Macbride 2755. VERAGUAS: Karszenisz s. n. This species was described from fruiting (immature) material, and perhaps when flowering material becomes available specific delimitation will be more precise. Apparently it is intermediate between South American species (C. guianensis) and C. panamensis. NAMES OF UNCERTAIN APPLICATION Whether the name C. officinalis L., type for the genus, is at all applicable to Panamanian species of Copaifera cannot be stated certainly. Standley (Contr. U. S. Nat. Herb. 27:203. 1928) listed it (as Copaiva officinalis) as occurring in the Canal Zone, but other works tend to regard C. officinalis as restricted to the Antilles? and northern South America (although Britton & Killip, Ann. N. Y. Acad. 35:165. 1936 do not list it from Colombia). Dr. Dwyer believes that two Standley specimens (26185, 30608), both sterile, from Panama, are C. guianensis Desf. He states: "The leaflets of . . . . these col- lections . . . . are typically those of C. guianensis. From the labels, however, one learns that both specimens were taken from ‘vines’. As all of the Copaiferas range in size from small to very large trees, this notation is certainly of great interest." We must thus regard the occurrence of C. guianensis in Panama as yet highly uncertain. 8. PRIORIA Griseb. Prioria Griseb. Fl. Brit. W. Ind. 215. 1860. Large unarmed tree, with glabrate to lightly pubescent branchlets. Leaves simply pinnate, bijugate except occasionally monojugate, short-petiolate; leaflets pellucid-punctate, coriaceous to submembranaceous, arcuate-oblique and thus some- what asymmetrical, reticulate, glabrous; petiole and rachis lenticellate; petiolules short, callous-rugose. Inflorescence a terminal panicle of multitudinous spicate flowers. Flowers small, apetalous, apparently with petals (calyx-lobes) if sub- tending bracts are regarded as part of the perianth; calyx bibracteate (bracts some- times lobed and simulating a calyx), essentially polysepalous but basally more or (327) [Vor. 38 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN less disciferous, the sepals 5, orbicular, imbricated in bud; stamens 10, free, sub- equal; ovary subsessile, 1- to 2-ovulate. Legume flattened verrucose, ligneous, tardily 2-valved, monoseminate. The genus consists of a single species. Fig. 112. Prioria Copaifera 1. Priorta Copatrera Griseb. Fl. Brit. W. Ind. 215. 1860. Trees up to 40 m. high, with spreading crown and large trunk. Leaves 4-(2)- foliolate; petioles about 1—2 cm. long, basally callous-rffgose; leaflets elliptic- lanceolate, 6—16 cm. long, 4—8 cm. wide, rounded-obtuse and somewhat unequal basally, bluntly short-acuminate apically; rachis 2—5 cm. long; petiolules 2-6 mm. long. Inflorescence a panicle of many flower-bearing spikes each up to 10 cm. long. Flowers white-yellow, basally bibracteate; bracts broadly orbicular, about 1.5 mm. long, ensheathing the disciferous part of the calyx, often lobed and simulating a calyx; sepals scarious-margined, ciliate, about 2.5 mm. long; petals lacking; stamens 10, about 5 mm. long; filaments basally and internally lightly lanose; anthers or- (328) t951] FLORA OF PANAMA (Leguminosae) 29 bicular, bilocular, loculi markedly separated by the connective; ovary subsessile, lightly lanose marginally, with a short attenuate style and inconspicuous stigma. Legume suborbicular, up to 10 cm. long and almost as wide, prominently veined, verrucose, lepidote at least in youth; single seed large, flat, almost filling the pod. Panama to Nicaragua; Colombia; West Indies. BOCAS DEL TORO: Changuinola Valley, Cooper & Slater III. CANAL ZONE: Barro Colorado Island, Bangham 524, Salvoza 943; Chagres River, Allen 909; Trinidad River, Pittier 3071. DARIEN: vicinity of Pinogana, Allen 277, 932. The species is a source of "Copaiba balsam”, used medicinally and pharmaco- logically. With some specimens it is stretching the imagination to regard the flowers as apetalous, so much do the bracts simulate a calyx and the calyx a corolla. 9. PELTOGYNE Vogel PELTOGYNE Vogel, in Linnaea 11:410. 1837. Unarmed trees, the branchlets terete. Leaves simply pinnate, short-petiolate; leaflets one pair, coriaceous, punctate, short-petiolulate to subsessile, inequilateral. Inflorescence racemose, terminal or subterminal, few- to many-flowered. Flowers small, whitish; calyx receptacular and disc-like basally, the lobes 4, imbricate in bud; petals 5, sessile, punctate, subequal; stamens 10, free, glabrous; ovary few- ovulate, short-stipitate the stipe adnate to the receptacular portion of the calyx; style slender, with a peltate-capitate stigma. Legume 2-valved, flat. Primarily a South American genus centered in northern Brazil. A single species is known from Panama. PELTOGYNE PURPUREA Pittier, in Jour. Wash. Acad. 5:471. 1915. A tall forest tree with slender, glabrous branchlets and hard wood, usually found in localities with a dry season. Leaves deciduous, glabrous, reported with — caducous, membranaceous, acuminate stipules about 1 cm. long; petiole to 2 cm. long; leaflets 2, 5—7 cm. long and 2—3 cm. broad, markedly reticulate, subfalcate, inequilateral, acuminate, obliquely rounded or obtuse basally, short-petiolulate, the petiolules 3—4 mm. long. Inflorescence as described for genus. Flowers not de- scribed nor examined. Legume pedicellate (about 8 mm.), broadly obovate, about 3 cm. long and 1.6 cm. broad, flattened, glabrous, verrucose-reticulate with narrow sutures, apically mucronulate, slightly arcuate above, rounded below, 1-seeded; seed almost 2 cm. long, obliquely ovate, depressed, persistent on the dehisced fruit hanging by the funicle; funicle dilated into a narrow, cupuliform aril. Southern Darién, Panama; Costa Rica? DARIEN: La Palma, Piftier 6621 (sterile); Patino, Pittier 5708, 6610; Yaviza, Pittier 6586 (sterile). As is common for the "purple heart" genus, the wood is very hard, with white sapwood and beautiful purple heartwood, valuable in various uses. Until recently the species has not been known in flower, and its exact placement and affinities un- confirmed. Paul H. Allen states that he has collected abundant flowering material (329) [Vor. 38 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN of presumably the same species from Costa Rica, but I have not had opportunity to examine these collections. 10. HYMENAEA L. HYMENAEA L. Sp. Pl. 1192. 1753. Courbari Adans. Fam. 2:317. 1763. Tanroujou (Juss. Gen. 351. 1789) has also been given as a synonym for Нутепаса, referring to a Madagascar fruit. Unarmed resinous trees, with spreading crown. Leaves l-jugate, petiolate; leaflets 2, obliquely asymmetric, coriaceous, glandular-punctate, short-petiolulate to subsessile. Inflorescence terminal, short, subcorymbose. Flowers moderately large, with a short, gross, pedicel and thick receptacle; calyx 4-parted, the lobes imbri- cate; petals 5, slightly unequal, sessile, glandular; stamens 10, free, glabrous; anthers longitudinally dehiscent; ovary few-ovulate, short-stipitate, the stipe adnate to the receptacular portion of the calyx; style slender; stigma terminal. Legume ligneous, thick, indehiscent, few-seeded; seeds exarillate. A neotropical genus, best represented in the Amazon valley. A single species is known from Panama. 1. HvMENAEA Coursarit L. Sp. Pl. 1192. 1753. Hymenaea resinifera Salisb. Prodr. 327. 1796. H ymenaea animifera Stokes, Bot. Mat. Med. 2:449. 1812, fide Index Kew. Hymenaea Candolleana HBK. Nov. Gen. & Sp. 6:323, f. 566. 1824. Hymenaea retusa Willd. ex Hayne, Darst. u. Beschreib. Arzneigew. 11:sub, Ё. 12. 1856. Inga megacarpa (M. E. Jones, Contr. West. Bot. 15:140. 1929) is listed in the ‘Nort American Flora’ as a synonym of this species. Jones! description would preclude this pos- sibility, but if Britton had reference to the actual type perhaps the description was incor- rectly drawn by Jones. Description of the fruit would match H. Courbaril, but description of the leaves would not. Tree to 30 m. tall, with smooth bark, the trunk to 2 m. in diameter, the wood hard and reasonably durable, the branchlets glabrous. Leaves glabrous; petiole about 1-2 cm. long, moderately thick, rugose when dry; stipules caducous; leaflets 2, narrowly oblong to elliptic-lanceolate, 4-10 cm. long, 2—5 cm. wide, obliquely asymmetric, the outer portion broadly rounded basally and without, the inner por- tion narrow and only slightly rounded, apically short-acuminate, subsessile, coriaceous, punctate, dull below, shiny above, with midvein very prominent below. Inflorescence articulate, several-flowered, the pedicels puberulent, the bracts caducous. Flowers whitish, gross, the receptacular portion (of calyx) about 8 mm. long; calyx-lobes ovate to oblong, about 15 mm. long, densely puberulent, verrucose, coriaceous, easily caducous; petals elliptic, up to 2 cm. long, mem- branaceous; stamens about 3 cm. long; anthers elliptic, versatile, bilocular; ovary elliptic, oblong or obovate, compressed, glandular, dark; style up to 2.5 cm. long, glandular. Legume oblong, turgid-compressed, 5—15 cm. long, few-seeded. (330) 1951] 31 Fig. 113. Hymenaea Courbaril Mexico through Central America; West Indies; northeastern South America. CHIRIQUÍ: Caldera, Pittier 3351. cocrÉ: vicinity of El Valle, Allen 1766; Penonomé, Williams 193. VERAGUAS: EAR Rio Cafiazas, Allen 199. The species is of economic importance, both as timber and as a source of the resin "South American copal.” The pulp of the fruit is edible. The strong, heavy wood is employed in various kinds of construction. The resinous exudation from the trunks is of use medicinally, in varnish manufacture, and as an incense. Indians are reported eating the fruit pulp, and using the bark in making canoes. 11. MACROLOBIUM Schreb. MaAcROLOEB!UM Schreb. Gen. 1:30. 1789, nom. conserv. Vouapa Aubl. Hist. РІ. pen Тү 1:25, 4. 7. EH Outea Aubl. ES СС 128; 775: Kruegeria Scop. Introd. inc Rm 314. 12271 Antbonatba aun Fl. d'Oware 1:70, | 4 jim fide Dalla Torre & Harms. Utea J.. St. Ee Expos. ded 2: 1: Vuapa Ktze. Rev. Gen. 1:212. 189 Орзу ramet Britt. & Killip, 3 in et N. Y. Acad. 35:166. 1936. (331) [Vor. 38 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN Unarmed trees, of small or moderate size. Leaves pinnate, generally caducous- stipulate, with 2 to many leaflets (in Panamanian species a single pair); leaflets small or large, coriaceous or almost so, often unequal basally. Inflorescence usually racemose, terminal to axillary on older wood, bracteate but the bracts generally caducous. Flowers small to moderate, ensheathed to a greater or lesser extent by the (2) fused bracteoles, this bracteolar sheath cleft on one side; receptacular por- tion of calyx short; calyx-lobes 4 or rarely 5, slender, imbricate; visible petal 1, large, clawed to subsessile, folded-cucullate in bud, other petals squamiform or lacking; perfect stamens 3, elongate, free, in some species with up to 7 accompany- ing staminodia; ovary 2- to many-ovulate, subsessile to stipitate, the stipe or base of ovary adnate to the receptacle on one side; style slender, with a terminal small or capitate stigma. Legume flattened, 2-valved, small, orbicular to oblong; seeds 1 to few, large, ovate to orbicular. Panama; South America, most abundant in the Amazon section of Brazil; Africa. a. Petal about 1.3 cm. long 1. M. MODICOPETALUM aa. Petal about 4 cm. long 2. M TIERI Fig. 114. Macrolobium modicopetalum 1. MACROLOBIUM MODICOPETALUM Schery, in Ann. Mo. Bot. Gard. 30:88. 1943. Tree with subglabrous to glabrous branchlets. Leaves glabrous, 1-jugate; petiole short, terete, deeply canaliculate; leaflets subsessile, coriaceous, elliptic, basally unequal and subacute, apically short-attenuate and obtusely mucronate, (332) 1951] FLORA OF PANAMA (Leguminosae ) 33 12-23 cm. long and 4—9 cm. wide, the veins prominent beneath and laterally con- fluent. Inflorescence spicate, glabrous or very short-pubescent, bracteate, arising laterally or subterminally from the branchlets. Flowers narrow-pedicellate; the sheath (fused bracteoles) obovate, about 7—9 mm. long, bilobate, the lobes about 4 mm. broad; sepals 4, oblong, about 6-7 mm. long and 2.5-4 mm. wide, obtuse, glabrous; petal 1, white, ovate-lanceolate, 12-13 mm. long and 6—8 mm. wide, concave, undulate-margined, briefly unguiculate; stamens 3, the filaments linear, 11-19 mm. long, the anthers ovate, about 3 mm. long; ovary compressed, mar- ginally pubescent, substipitate, 4-ovulate; style glabrous, with the ovary up to 20 mm. long. Northwestern Panama. BOCAS DEL TORO: Fish Creek, von Wedel 2200, 2226, 2291, 2399. N Млскоговіом Pittieri (Rose) Schery, comb. nov. Vouapa Pittieri Rose, in N. Am. Fl. 23:226. 1930. Small tree. Leaves 2-foliolate, subsessile; leaflets broadly oblanceolate, 3—3.5 dm. long, sometimes 1 dm. broad, chartaceous, basally oblique, apically acute or acuminate. Inflorescence lateral, borne on older branches, 3—5 cm. long, short- pedunculate. Flowers pedicellate, the pedicels 4—6 mm. long, glabrous; sheath (fused bracteoles) 9—11 mm. long, glabrous, cleft for half its length; calyx gla- brous, the lobes imbricate in bud, linear-oblong to linear-obovate and about 2 cm. long in anthesis; petal 1, elliptic, about 4 cm. long and 13 mm. wide, thin, erose- undulate margined, glabrous, short-clawed; ovary lightly pubescent on margins, the style glabrous; legume oblong, 15 cm. long and 5 cm. wide, glabrous, the valves twisting in age; seed oblong, 2 cm. long, flattened, rugose. Northeastern Panama. SAN BLAS: Puerto Obaldia, Pittier 4355. 12. BROWNEA Jacq. Brownea Jacq. Enum. Pl. Carib. 6. 1760 (as Browneae) , nom. conserv. Hermesias Loefl. Iter Hisp. 278. 1758, hyponym. Shrubs or trees, branchlets pubescent or glabrous. Leaves large, simply pinnate, few- to many-foliolate; leaflets in pairs, opposite to subalternate, large, entire, short-petiolulate. Inflorescence capitate to short-racemose, terminal to cauliflorous, bracteate, the bracts often colored and caducous. Flowers large, showy, ensheathed basally by the coalesced bractlets; calyx 4- to 5-parted, submembranaceous, with an elongate, thickened receptacular base; petals 4—5, membranaceous, clawed; stamens 10—15, basally connate. Ovary stipitate, the stipe adnate the whole length of the receptacular tube. Legume flat, 2-valved. Panama to Brazil; southern West Indies; center of dispersion apparently interior Venezuela. (333) [Vor. 38 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN This striking genus is apparently in need of specific condensation. Plants from different localities (bearing different names) appear highly similar in the herbarium, and specific limits are poorly defined. The specimens are usually bulky and incom- plete оп herbarium sheets,—the sheets often being without reference to location of inflorescence or other so-called key characters, and with parts of the large in- florescence or flowers missing or caducous. The plants themselves are extremely variable in the number of leaflets on various leaves of the same plant. The specific names listed must temporarily serve until monographic study can determine their true reference and complete synonymy. a. Leaflets elliptic to ovate, usually rounded or obtuse basally, s glabrous below; floral sheath mostly rusty-tomentose or tomentulos 1 and scarcely 3 cm. long . B. Rosa-DE-MONTE аа. Leaflets narrowly e lliptic, — cuneate Mrs lly or narrowing below the middle, pubescent below, at least alon Pw midvein; floral — canescent-tomentose, usually fe 3.5 cm. “ina . B. MACROPHYLLA 1. BROWNEA Коѕл-ре-момте Berg. in Phil. Trans. Roy. Soc. London 63:174, pls. 8 9 0. 1773. Brownea Rosa Pers. Syn. Pl. 2:237. 1807. Brownea rosea Otto, in Otto & Dietr. Allg. Gartenzeit. 23:147. 1855, fide Index Kew. Brownea Princeps Linden ex Otto, in Otto & Dietr. loc. cit. 1855, equals B. Ariza, fide Index Kew PBrownea Ariza Benth. РІ. Hartw. 171. Brownea speciosa Rchb., Griseb. Fl. Brit. W. Ind. 212. 1860. (DC. da 2:477. 1825), Brownea coccinea Ідей. ex Griseb. loc. cit. 1860, поп В. coccinea Jac Hermesias Rosa O. Ktze. Rev. Gen. 191. 1891. Brownea rosa-montis Pittier, in Contr. U. S. Nat. Herb. 18:156. 1916. Small tree, with lenticellate, glabrous (in Panama?) branchlets. Leaves few- to several-jugate, glabrous, short-petiolate; the rachis stout, lenticellate, up to 40 cm. long or longer; leaflets elliptic (lowermost on leaf usually ovate or subovate), up to 20 cm. long or longer, 2—9 cm. wide, dull, apically attenuate-cuspidate, basally obtuse to rounded, with a gland below to one side on base of the midrib; petiolules stout, 3-6 mm. long. Inflorescence on older branchlets, condensed, bracteate, the bracts puberulent, broadly ovate to linear, several cm. long; com- mon peduncle 2—3 cm. long, tomentose; pedicels about 3-5 mm. long, pubescent. Flowers conspicuous, scarlet-red; bracteolar sheath relatively small, usually 2.5—3.0 cm. long, bilobed, tomentulose to tomentose, usually ferruginous; sepals (i.e. lobes) 4—52, oblong to linear, up to 25 mm. long and 13 mm. wide, subglabrous, free, often caducous; petals 5, usually oblong, about 4 cm. long and 1—1.5 cm. wide, glabrous, clawed, claw about 1 cm. long; stamens 10-11, about 8 cm. long, gla- brous, connate below for about 2 cm.; ovary tomentose, stipitate, with style about 8 cm. long. Costa Rica to Northern South America and Trinidad. CANAL ZONE: Ancón, Pittier 2722; R. Boqueron, Steyermark & Allen 17226; R. Indio Dodge & Allen 17309. pAmiÉN: El Real, Allen 045. saN BLAS: Puerto Obaldia, Pittier 4408. (334) 1951] FLORA OF PANAMA (Leguminosae) 59 Fig. 115. Brownea Rosa-de-monte Intergradation occurs between this and B. macrophylla. Intensive study of the genus in northern South America may show the differences to be of subspecific value only. B. Ariza (B. Princeps, fide Index Kew.) is tentatively listed as a synonym on what evidence can be ascertained from examination of Panamanian and South American specimens and its own brief description, but the type has not been seen. Standley (Contr. U. S. Nat. Herb. 27:202. 1928) and Pittier (Contr. U. S. Nat. Herb. 18:153? 1916) both list B. Ariza as occurring in Panama as a distinct species. 2. BROWNEA MACROPHYLLA Linden, Cat. no. 18:11. 1863; Mast. in Gard. Chron. И ОТ TAO LSZ 3 Brownea antioquensis Linden loc. cit. по. 23:3. 1869. [Vor. 38 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN Small tree, the branchlets generally tomentulose. Leaves few- to several-jugate; the petiole stout, to 10 cm. long; the rachis puberulent to glabrous, elongate; leaflets narrowly elliptic to elliptic-oblanceolate (sometimes more or less ovate in basal pair), up to 30 cm. long and 9 cm. wide, apically long attenuate-cuspidate, basally acute to cuneate (infrequently obtuse or rounded), pubescent at least along the midvein below, dull, with a basal gland; petiolules to 5 mm. long, callous and often tomentose. Inflorescence axillary on older stems to subterminal, bracteate; the bracts to 7 cm. long, glabrous to lightly pubescent; common peduncle about 3 cm. long, tomentose, many-flowered; pedicels 4—7 mm. long, pubescent. Flowers large, red; bracteolar sheath about 3.5 cm. long, spathaceous and bilobate, usually canescent-tomentose; calyx-lobes 5, linear-oblanceolate to narrowly elliptic, 2.5-3.5 cm. long, up to 8 mm. wide, glabrous; petals 5, rounded apically, up to 4 cm. long, the claw about 1 cm. long; stamens 10—11, glabrous, basally connate, usually 8—9 cm. long; ovary narrow, stipitate, tomentose, in- cluding style about 10 cm. long. Northern South America and Panama. L ZONE: Fort Randolph, -— m Fort Sherman, Standley 31005; France Field, Stevens 987; between France Field and Catival, Standley 30105. согом: Porto Bello, Peterson 6522. DARIEN: Boca ife PME Pittier 5591; Cerro Рігге, Pittier 6073; Pinogana, Pittier 6571. PANAMA: Juan Diaz, Hunter & Allen 931. The original “description” and ‘Gardeners’ Chronicle’ figure of B. macrophylla are nearly useless in determining correct application of this name. Inasmuch as the type specimen was not seen, I have followed generally Pittier’s interpretation of the species (Contr. U. S. Nat. Herb. 18:155. 1916), except for Peterson 6522 which seemed to have more affinities here than with B. Rosa-de-monte. 13. BROWNEOPSIS Huber BnowNEoPsis Huber, in Bol. Mus. Goeldi 4:565. 1906. Tree to 30 or more meters. Leaves even-pinnate, short-petiolate, the rachis extended, the leaflets subopposite in pairs. Inflorescence terminal to cauliflorous, congested, in more or less conspicuously bracteate heads. Flowers ebracteolate, sessile; calyx with a thick basal, obconic, angular, receptacular portion, and 4- parted limb, 3 sepals similar, the fourth different, frequently circumscissile; petals 3 or 4, rudimentary to partially petaloid-expanded; stamens 12-16, the filaments connate below for about half their length, somewhat unequal; ovary slender, stipitate, the stipe completely adnate to the receptacular tube; style elongate, with a capitate stigma. Legume large, flat, 2-valved. Panama to Brazil. A genus allied to Brownea, in a section of the Leguminosae where generic limits are often difficult to determine. Browneopsis is separated from Brownea primarily because of the lack of a bracteolar sheath in the flowers of the former. Macbride (Field Mus. Bot. Ser. 13%:135. 1943) has referred the species back to Brownea. A single species is known from Panama. (336) 1951] FLORA OF PANAMA (Leguminosae) 37 1. BROWNEOPSIS EXCELsA Pittier, in Contr. U.S. Nat. Herb. 18:157, pl. 63. 1916. Brownea excelsa (Pittier) Macbride, in Field Mus. Bot. Ser. 13:135. 1943. Large unarmed tree, with terete, glabrous, lenticellate branchlets. Leaves 4- to 6-foliolate; petiole short, gross basally, with rachis up to 10 cm. long; rachis flattened and usually 1-sulcate above, glabrous; leaflets ovate to lanceolate or ob- long, 5—11 cm. long, 2—5 cm. wide, apically usually very long-acuminate, basally truncate to obtuse to cuneate, glabrous, dull, chartaceous, minutely reticulate; petiolules about 6 mm. long. Inflorescence a condensed, short-peduncled, few- flowered, cone-like “head”, terminal or axillary and subterminal; bracts broadly ovate, to 3.5 cm. long, ciliate, somewhat puberulent, longitudinally rugose, over- lapping. Flowers cream to pinkish, showy; calyx with a thick basal receptacular portion about 8 mm. long, the 4 petaloid sepals (lobes) elliptic, one shorter than the other 3, to 2.5 cm. long, often caducous; petals 3 or 4, 2? of them narrowly elongate-obovate, about 3 cm. long, clawed, glabrous; stamens 14—16, mona- delphous, glabrous, up to 3.5 cm. long; anthers linear-oblong, up to 1 cm. long, versatile, bilocular; ovary narrow, stipitate, densely pubescent, with style up to 4 cm. long or longer; style glabrous; stigma terminal, rounded. Legume linear- oblong, 15—20 cm. long, 3—4 cm. wide, thin and flattened, densely puberulent, obliquely attenuate-mucronate apically, long-stipitate basally, few-seeded. Darién, Panama. RIÉN: Garachine, Pittier 5511; Marraganti, Williams ТОІІ; between Pinogana and Gen. Allen 261; El Real, Allen 946. 14. TAMARINDUS L. TAMARINDUS L. Sp. Pl. 34. 1753. Medium-sized trees of lower elevations. Leaves evenly pinnate, moderate-sized, short-petiolate; stipules minute, caducous; leaflets small, numerous, sessile, paired. Inflorescence short, usually terminal, few-flowered, racemose. Flowers moderately large, attractive; calyx 4-parted, the lobes reflexed above the receptacular base, imbricate in bud; petals very unequal, the 3 uppermost expanded and the 2 lower- most rudimentary; stamens 3, monadelphous; staminodia minute; anthers longi- tudinally dehiscent; ovary stipitate, the stipe adnate to the receptacular tube; style elongate, with a terminal subcapitate stigma. Legume linear-oblong, curved, thick, indehiscent, septate between the obovate-orbicular seeds. Native to the Far East but planted and naturalized throughout the world tropics. The genus is monotypic. 1. TAMARINDUS INDICA L. Sp. Pl. 34. 1753. Tamarindus occidentalis Gaertn. Fruct. 2:310, £. 146. 1791. Tamarindus umbrosa Salisb. Prodr. 323. 1796. Tamarindus officinalis Curt. Bot. Mag. pl. 4563. 1851. (337) [Vor. 38 > oe Ар» mu Pro, SEK Fig. 116. Tamarindus indica Unarmed, naturalized trees, with puberulent to glabrous branchlets, spreading crown, and rough, brown bark. Leaves 6—12 cm. long, glabrous or nearly so; leaflets 6—18 pairs, oblong, 1—2.5 cm. long, rounded to retuse apically, obliquely obtuse to subtruncate basally, chartaceous, reticulate. Inflorescence as described for the genus; pedicels slender, 6—10 mm. long. Flowers yellow striped with red; sepals 5, elliptic-lanceclate, 6-10 mm. long; larger petals 3, cbovate, 8-12 mm. long, subfimbriate; stamens arcuate, investing the ovary below, up to 14 mm. long; anthers oblong, about 2 mm. long; cvary somewhat pubescent basally. Legume 5—15 cm. long, about 2 cm. thick, lepidote, with an acid pulp surrounding the seeds; seeds about 1 cm. wide. World tropics. PANAMA: Taboga, Hayes 201. The species is commonly cultivated. The pulp of the fruit, said to be rich in formic and butyric acids, is used in preparation of a refreshing drink. The fruit is also reported to have medicinal properties, used as a laxative and in preparation of a gargle. 15. DIALIUM L. DrALIUM L. Mant. 1:3. 1767. Arouna Aubl. Pl. Guian. pog 1: ^ Bs 1775. Aruna Schreb. Gen. РІ. 1:26. 178 Cleyria Neck. Elem. 2:183. gr d Codarium Sol. ex Vahl, Enum. 1:302. 1805. (338) 1951] FLORA OF PANAMA (Leguminosae ) 39 Unarmed trees, usually large. Leaves once-pinnate; leaflets few, moderately large, mostly alternate, coriaceous to submembranacecus; stipules caducous. In- florescence paniculate, axillary or terminal. Flowers small, caducous-bracteate; calyx 5 (rarely 4) -lobed, the lobes imbricate, the receptacular portion short, thick; petals 1-2 and minute, or lacking; stamens 2 or rarely 3, distinct, with short fila- ments; anthers oblong, basifixed, erect, dehiscent by slits; ovary sessile or sub- sessile, generally 2-ovulate; style short, with terminal stigma. Legume subglobose to ellipsoid, sparingly fleshy, indehiscent; seed usually 1, somewhat compressed. Essentially a genus of the Old World tropics; with a single species in the New World, ranging from Guatemala to eastern Brazil. 1. DIALIUM GUIANENSE (Aubl.) Sandwith, in Lloydia 2:184. 1939. Arouna guianensis Aubl. Pl. Guian. Franc. 1:16, 7. 5. 1775. Aruna divaricata Willd. Sp. Pl. 1:156 ae Dialium dum Vahl, Enum. 1: 303. Dialium acuminatum Spruce ex Williams, in iud Mus. Publ. Bot. 15:201. 1936. Tall tree, with glabrous or puberulent branchlets. Leaves mostly 5- to 7- foliolate, caducous-stipulate, short-petiolate; rachis about 5-8 cm. long, often puberulent; leaflets ovate to elliptic-lanceolate, apically attenuate, basally rounded to cuneate, 3-12 cm. long, 1.5-4 cm. wide, glabrous, conspicuously reticulate, alternate or subopposite; petiolules about 3 mm. long, flattened or sulcate above, somewhat callous. Inflorescence an extended, branching panicle, terminal or sub- terminal, puberulent to glabrous; bracts caducous; pedicels very short. Flowers small, inconspicuous, apetalous; calyx pubescent without, densely so in bud, the receptacular portion short and stocky, the 5 lobes expanded, imbricate, obtuse, up to 3 mm. long; petals lacking; stamens 2, short; anthers bilocular, erect and basi- fixed, dehiscing by slits from the apex, the connective conspicuous; ovary sessile or subsessile, pubescent, 2-ovulate; style short, with a small terminal stigma. Legume subglobose to ellipsoid, up to 2.5 cm. long, very short-stipitate, with a fragile exocarp and somewhat fleshy endocarp. Central and South America, from Guatemala to Peru and Brazil. DARIEN: vicinity of La Palma, Pittier 5486. saN BLAS: Permé, Cooper 240. The species furnishes a very heavy, durable, brownish wood, used in construc- tion and for posts, but difficult to work because of its hardness. The fruit is edible, and is often consumed by various animals. At least one additional species of Dialium, presumably exotic (Steyermark s. n.), is in cultivation at the Summit Plant Introduction Garden, Canal Zone. 16. CASSIA (Tourn.) L. Cassia (Tourn.) L. Sp. Pl. 376. 1753. Senna (Tourn.) Mill. Gard. Dict. ed. 4. 1754. Chamaecrista Moench, Meth. 272. Cathartocar pus Pers. Syn. Pl. 1:459. 1805. Grimaldia Schrank, in Denkscht! Akad. München, 103. 1808. (339) [Vor. 38 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bactyrilobium Willd. Enum. Hort. Berol. 1:439. 1809. Cassiana Raf. in Am. Monthly Mag. 2:2 1818 Chamaecassia Link, Handb. 2:139. 1831 Chamaefistula С. Don, Gen. Hist. "He PI. 2:451. 1832. Xamacrista Raf. loc. cit. 127. 1838 Mac-Leayia Montr. in Mem. Acad. Lyon 10:198. 1860. Chamaesenna Raf.; Pittier, Arbol. & Arbust. Legum. 130. 1928. Cowellocassia Brite. à in N. Am. Fl. 23:251. 1930 Desmodiocassia Britt. & Rose, loc. cit. 244. 1930. Echinocassia Britt. & Rose, ag cit. 251, 1930. 30. Phragmocassia Britt. & Rose, loc. cit. 245. 1930. Pseudocassia Britt. & Rose, loc. cit. 230. 1930. Psilorbegma (Benth.) Britt. & Rose, E cit. 255. 1930. 1930 Xerocassia Britt. & Rose, loc. cit. 246. 1930. Trees, shrubs or herbs, or sometimes vine-like. Leaves pinnate, 2- to many- foliolate or occasionally aphyllous, often with petiolar or rachial glands; leaflets entire, often inequilateral. Inflorescence bracteate, axillary or terminal, racemose or paniculate or the flowers rarely subsolitary. Flowers usually yellow, conspicuous; calyx with a disc-like base and 5 usually imbricate sepals; petals 5, distinct, ex- panded, often unequal; stamens 10—4, the 3 uppermost usually rudimentary or lacking, the 4 median similar, usually intermediate in size, often conspicuously rostrate; the 3 lowermost similar or dissimilar, rarely lacking, variously modified but usually rostrate, usually larger than the median stamens; anthers dehiscent by terminal or basal pores or sometimes (in addition) lateral slits, basifixed and erect; ovary sessile or stipitate, several- to many-ovulate. Legume dehiscent or sometimes indehiscent, terete or more often compressed or flattened, the seeds often horizontal, the funicle filiform to very short. (340) 1951] FLORA OF PANAMA (Leguminosae) 41 A very large natural genus of world-wide distribution, common mostly to the tropics and subtropics; very abundant in the Americas a. Leaves without petiolar or rachial glands. eaves erp: multifoliolate, with 4 (very infrequently) or more hg о . Three joo stamens with slender filaments at least 5 times the length of the ovate anthers; anthers dehiscent basally as well as apically. d. Leaflets a or die и large, 8—20 cm. long; flowers about 4 cm. 1. C. FISTULA broad; in uced t dd. Leaflets р to 20 pairs, small, 6 cm. or less long; flowers about е. Calyx c canescent-tomentulose without; median stamens 5, sub- equal, the anthers lightly pilose; ovary tomentose; flowers pink or whitest. oun 2. C. GRANDIS ee. Calyx puberulent to subglabrous without; median stamens the anthers glabrous; ovary essentially glabrous; АКЕ yellow MM n m 3. C. MOSCHATA ес Юа stamens mostly with the oblong anthers longer than the filaments more than about twice in no case anther; anthers uri pm unusu n in Panama............ 29. C. MULTIJUGA 12—40 pairs, small nthers short-rostrate; йн normally 6—15 pairs, narrow or бике н, not en- adn the u m ermost lateral anthers about 7 mm. long, only lightly ek the шо anthers; legume ether turgid-quadrangular E if flattened, relatively few- dd. L larger 12 cm. long an ЖИ көө жылуы рии nther; ovary pubes ent; legume flattened; introduced tree... 4. C. SIAMEA ff: Leaflets “коо, acute apically, lightly pubescent b fila ments 2 m. the anthers; ovary glabrous; nee 1 5. C. SPECTABILIS vi pied тие как (mostly 1—2.5 cm. long) om the buds; lowermost Дыр т anthers ог longer, ied exceeding е y in m bt se) and m nate apically; stipules in reniform, about 3 cm. ine = ae pubescent; m eoe introduced 6. C. DIDYMOBOTRYA ts glabrous; ovary glabrou т. 7. C. NICARAGUENSIS в? p oblong to obovate, M 2— . wide, rounded and blunt apically, som ik pn mucrone; ded inequi- laterally lance olate, about 1 c Leaflets conspic E tomentuose below, especially on .. 8. C. RETICULATA LATA gg. Leaflets glabrous or e so; legume winged................... bb. Leaves normally 4- or 2-foliolate quic 5: leaves up to 10-foliolate in C. emarginata). c. Shrub or small tree; mature leaf larger, г leaflets 3—8 cm. long; ted in Panama... 10. C. EMARGINATA legume elongate, at least 15 cm. long; t expect cc. eigene ог ie bg pu ы pios 0.5—3 cm. ions: legume shorter, less than 8 cm. lon dr alle normally 4; d in Кб terminal racemes; stipules minut e. tae flowers larger (sepals mostly 8 mm. long or longer) ; M as essentially glabrous below 1. C. HISPIDULA (341) [Vor. 38 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN ee. Leaves and flowers smaller (sepals mostly 7 mm. г унар leaflets more or less setulose or pubescent с ЗАКИР lla. C. HisPIDULA var. KirLIPII dd. Leaflets normally 2; flowers 1-2 from the axils; stipules about ‘oles 12. C. ROTUNDIFOLIA do. RR E. oo ice ымы ралды ы ылы ы OA aa. At least most leaves with glands on — or rachis Glands borne on the i сар ea lowermost Pair of leaflet c. Stamens similar (although addi. jo» nthers erostrate and i uberulent мк э nal ridge bandi glands concave Wit а apically escis or sessile), borne on mid or upper petiole; leaf- lets mostly 0.5—2.5 cm. long; “Chamaecrista”. d. Leaflets 1—3 pairs; stipules cordate- мімен pair 13. C. DIPHYLLA а мач prostrate; petiolar gland үн! e small, rcel m. wide; legume small, abou lon ЖК ff. Plane hrub. со petiolar gland cae е larger, at ;1 3 cm. long 14. C. TAGERA 15. C. BREVIPES C. FLEXUOSA about 4 mm. lo 17. C. STENOCARPA gg. Flowers larger (normally 1 cm. long or longer); largest hers 7-12 mm long; ovary strigose- -pilose or tomen- h. Flowers scarcely more than 1 cm. long; longer anthers about 7 mm. long; petiolar gland usually short- dn с 7 o дынын ынена ыйын ыны дыы ыы: 17a. C. STENOCARDA V STE касынан hh. Flowers mostly 12-15 mm. lon ng; longer anthers usuall 11-12 mm. long; petiolar gland usually slen- 18. C. FLAVICOMA erly sti e ff. Petiolar glands sessile, patelliform; stem pubescence of mostly s i &. Leaflets pubescent, the surface (in dried mater ial) ap- earing verrucose with the мавту нац hair bases; stem bescence ascending... LLL 19. C. PATELLARIA glabr g 20. C. SIMPLEX cc. е dissimilar, the anthers short-rostrate and glabrous; glands x, usually bene close to petiolar insertion; leaflets mostly че 10 ст. goes d. Plan nt pubescent to hirsute; legume elongate and very narrow, wide асг. НАР ОКЕАННИНГ РЫ 21; C. LEPTOCARPA var, HIRSUTA dd. = igi or puberulent; legume shorter and broader, about e. > Рае ORT globose; leaflets т slightly larger (2—10 M шанды ныны дана А ae 22. C. OCCIDENTALIS (including C. sopHera) ee. menu әмер пере cylindric; leaflets averaging slightly dee СЕ Ош. on а coronae ыа даа 23. C. LIGUSTRINA - - б 5 а. a © д в о 'o 5 et S. % n1 £2 бо 5.8 a - 8 o 2 [e] [0] 5 e > % S 2 б la] 3 5 - et "a Б. " о ^ ч E. a o 24 leaflets or their scars, c. Leaflets 3 or more pai d. Leaflets yr 3 pairs, Е below; legume usually strongly arcuate or falcate-curved. e. Flowers larger, iioi about 20- ei mm. long; at least some rhai elongate, with anthers 8-10 mm. long; le egume NEM ОИС EE O АИ 25, C. LEIOPHYLLA ee. iii smaller, petals up to 12 mm. sai all stamens short, nthers 2—4 mm. long; legume зиЬсегетге.............................. 6. C. TORA (342) 1951] FLORA OF PANAMA (Leguminosae) dd. At least some leaves with more than 3 pairs of leaflets, а subglabro glaucous "v (C. Lic alan bera Dos d pairs p Suites has a broader legume usually 1 cm. o . laevigata, with either 3 or 4 КЕ pe leaflets, has me күөс, legume straight or inconstantly a e. Upper stem usually hirsute with hairs about 8 Rd slands Jong niis leaflets 30 or more um ee 1 wide) Д к nt with hairs about 1 mm. long; rachial glands а исне to subsessile; айе ts fewer (except pou in C. multijuga), mostly oblong or wider (at least 5 mm f. lc о а, А io rachis 10-25 cm. long; leg at, about 15 ЕЕ renee 10 or fewer кї E rachis 1-14 cm. Jong; legume if flat n ver 5 mm. wide, otherwise Ша ot ove g. Flowers borne in pairs on filiform pedicels from m а lowermost anthers very long- gone le or less wide, flat....... bm по lowermost 2 ia жине) 43 28. C, WILLIAMSII 29. C. MULTIJUGA 30. C. BIFLORA 88- rostrate; legum mm. or more wide, subte h. Upper stems and lower surface of leaflets айны lowermost eee anthers long-rostrate la. hh. Upper stems and lower surface 8 оши glabrous; lowermost lateral anthers short-rost i. Leaflets obovate, rounded to wi РИ apically, most С less long; larger anthers conspicu- usly rostrate 31. ii. Leaflets ovate, acute-a “сае, mostly 3—10 ст. Apu anche scarcely rostr 24. cc. Leaflets on d. Ty realy herbaceous legume very slender, only 3—4 mm. wide; ie nce of | airs 27. dd. Tre ees or shrubs legume 6— о тт. wide; pubescence, if any, of КО ait HAM leaflets . Large к bracts land stipules) persistent; small (not over 4 de), inequilaterally subfalcate and bcne nate. . Floral bracts about 2 mm. long i: Floral bracts about 8 mm. long or longer ee. Bracts (and stipules) iwi caducous (if occasionally per- sistent bs C. Maxonii, smaller and not foliaceous); leaflets with varicus but not all of the 3 characters listed above (e). P Leaflets only slightly тара асиїе, tomentulose be- low and prominent brownish vein g. Fertile stamens ia 4 gg. d stamens more than 4 ff. gp if acute markedly inequilateral, e ibo acumi- or caudate, puberulent below with very short or app а hairs or glabrous; fertile stamens usually g. Leaflets markedly inequilateral е bipes 3 anthers dehiscent a a na term gg. Leaflets or but slightly cda all anthers riens by paired terminal pores (although ie pores often somewhat fluent i er h aflets acuminate; petioles short, usually about 3 cm ng; 3 lowermost anthers shorter than 4 median ones; 1s we е lon e lower leaf surface puberulo hh. Leaflets caudate; жад i 10 cm. long; 3 lowe most anthers бдай the 4 median ones; iie C. BICAPSULARIS var. PUBESCENS C. BICAPSULARIS C. LAEVIGATA C. PILIFERA . C. UNICA C. UNDULATA . C. MAXONI C. OXYPHYLLA var. DARIENSIS . C. FRUTICOSA a. C. FRUTICOSA Var. GATUNENSIS 37. C. CAUDATA leaf surface glabro (343) [Vor. 38 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 117. Cassia Fistula 1. Cassia FisTULA L. Sp. Pl. 377. 1753. Cathartocarpus Fistula (L.) Pers. E М, :459 Bactyrilobium Fistula (L.) Willd. ‚Чо p] 1:440. 1809. Cassia rbombifolia Roxb. Fl. Ind. 2:334. 1832, fide Bent Cathartocarpus excelsus е . Don, Gen. Hist. Dic An 2:455. 1832. Cathartocarpus rbombifoliu s (Roxb.) G . Don, loc. cit. 18 Cathartocarpus fistuloides (Collad.) G. Dos, loc. cit. 454. ius Tree up to 20 m., usually glabrous. Leaves large, several-foliolate; petiole about 5 cm. long, glabrous, eglandular; rachis usually 2—4 dm. long, like the petiole; stipules small, caducous; leaflets normally 4—8 pairs, яд 8—20 cm. long and up to 8 cm. wide, ovate to lanceolate, acute apically, very obtuse basally, (344) 1951] FLORA OF PANAMA (Leguminosae) 45 puberulent to glabrous above, lightly pubescent below; petiolules up to 1 cm. long. Inflorescence a large, graceful, pendent, many-flowered raceme; pedicels slender, usually 3—4 cm. long. Flowers large, showy, yellow; sepals 5, comparatively small (usually about 6 mm. long), ovate or oblong, puberulent; petals 5, large, about 2 cm. long or longer, ovate-orbicular, short-clawed, venose; stamens 3-morphic; the 3 lowermost almost 3 cm. long, the anthers ovate-oblong, 4—5 mm. long, glabrous, dehiscent apically and basally; the 4 median stamens about 1 cm. long, the anther ovate-oblong, sagittate, about 4 mm. long, dehiscent from the basal lobes (and apical pores); 3 uppermost stamens shorter and smaller, somewhat unequal, the anthers similar to the median ones; ovary slender, lightly pubescent. Legume (reported) cylindric, about 50 cm. long, indehiscent, with horizontal seeds. Central America; West Indies; northern South America: native to Asia. CANAL ZONE: Ancón, Zefek 10; Balboa, Standley 30847 (sterile). The tree, indigenous to Asia, is planted extensively as an ornamental in tropical America. The pulp of the fruit is sweetish, and is said to be useful as a purgative or laxative. 2. Cassia GRANDIS L. f. Suppl. Pl. 230. 1781. Cassia brasiliana Lam. Encycl. 1:649. 1785. Cassia mollis Vahl, Symb. 3:57. 1794, fide Benth. Cathartocarpus grandis Pers. Syn. Pl. 1:459. 1805. Cathartocarpus brasilianus Jacq. Fragm. 58, t. 85, fig. 3. 1809. Bact yrilobium molle Schrad. in Gött. Gelehr. Anz. 713. 1821, fide Benth. Cassia regia Standl. in Contr. U. S. Nat. Herb. 18:103. 1916. Tree to 30 m., the branchlets pubescent towards the tip, later glabrous. Leaves large, many-foliolate; petiole short, eglandular, tomentose, canaliculate above; rachis up to 30 cm. long, like the petiole; stipules small, linear, caducous; leaflets up to 20 pair, oblong, 3-6 cm. long and up to 1.5 cm. wide, rounded or obtuse apically and basally, somewhat inequilateral basally, entire, tomentose below, more lightly pubescent and darker above, with about 20 pairs of lateral veins; petiolules tomentose, 1-2 mm. long. Inflorescence of several- to many-flowered racemes, axillary from older wood to subterminal; racemes to 20 cm. long, tomentose, the bracts caducous; pedicels slender, about 2 cm. long in lower flowers. Flowers showy, reported pink to white; sepals 5, unequal, oblong, the larger about 8 mm. long and 5 mm. wide, rounded apically, canescent-tomentulose without; petals ovate-orbic- ular, to 12 mm. long, short-clawed, glabrous; stamens 3-morphic, the anthers lightly pilose, the filaments glabrous; 3 lowermost stamens about 2 cm. long, the filaments gracefully arcuate and thickened towards the middle, the anthers basi- fixed, short-oblong, about 3 mm. long and 2 mm. wide, subsagittate below, de- hiscent both apically and basally; 5 median stamens about 1 cm. long, the filaments linear, the anthers ovate-orbicular, about 1.5 mm. long, more or less versatile and dehiscing basally; remaining 2 stamens rudimentary or obsolete; ovary linear, arcuate, tomentose. Legume very large, up to several dm. long and 5 cm. broad, subterete but margined, indehiscent, ligneous, transversely rugose. (345) Fig. 118. Cassia grandis (346) 1951] FLORA OF PANAMA (Leguminosae) 47 Mexico to northern South America; West Indies. BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 133. CANAL ZONE: Gatun- cillo, Piper 5127; Miraflores Lake, P. White 754; New Frijoles, Christopherson 131; Paraiso, Pittier 2532; near Vigia, Dodge, Steyermark & Allen 16576. сосіЁ: El Valle, Allen 4471. DARIEN: Marraganti, Williams 643. PANAMA: Chepo, Kluge 46; Chorrera, Allen 4262; Taboga Island, Macbride 2807. Cassia regia Standley is included as a synonym of С. grandis, although differing in degree of pubescence and certain other minor ways. Examination of the type number (Pittier 2532) indicates gradation towards C. moschata, and Standley’s type may have been a hybrid between С. grandis and С. moschata, a putative hybridization that seemingly has not again occurred (i.e. not represented by a second collection). 3. Cassia MOsCHATA HBK. Nov. Gen. & Sp. 6:338. 1824. S E mme [ex ==: 22 —_ 4 2 4” NT 49 À N £2» NS || |

4 Fig. 119. Cassia moscbata (347) [Vor. 38 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cathartocarpus moschatus С. Don, Gen. Hist. Dichl. Pl. 2:453. 1832. Tree to 12 m., the young branchlets puberulent, longitudinally ridged. Leaves large, many-foliolate; petiole short, eglandular, puberulent; rachis up to 30 cm. long, like petiole; stipules triangular-bilobed, caducous; leaflets up to 20 pairs, oblong, 4—5 cm. long and up to 1.5 cm. wide, obtuse apically, rounded and unequal basally, puberulent below and above (more lightly so above), dull below, with about 20 pairs of lateral veins. Inflorescence terminal or axillary from upper branches; bracts caducous; pedicels 1—2 cm. long, lightly puberulent. Flowers yellow; sepals ovate to orbicular, about 7 mm. long, puberulent to glabrate, re- flexed, dark; petals obovate to orbicular, up to 1.5 cm. long, short-clawed, some- what venose; stamens 4-morphic, glabrous; the 3 lowermost gross, their anthers ovate-orbicular, 3-4 mm. long, with terminal pores; the 4 median anthers oblong, sagittate, 2-3 mm. long, with basal pores; 2 smaller, orbicular anthers less than 1 mm. long; and 1 stamen rudimentary; ovary essentially glabrous. Legume terete, elongate, up to 5 dm. long and 2 cm. wide, septate, glabrous. Central America to British Guiana. CANAL ZONE: Chivi-chivi Trail, Piper 5137, Johansen 16; Mojinga Swamp, Allen 010. PANAMÁ: vicinity of Chorrera, Allen 1607; Río Tapia, Standley 28148. This species differs from C. grandis in the much more slender fruits, їп the lesser degree of pubescence in both flower and vegetative parts, and in the longer, pendulous racemes of differing colored flowers. 4. CASSIA SIAMEA Lam. Encycl. 1:648. 1785. Cassia florida Vahl, Symb. 3:57. 1794. Cassia sumatrana Roxb. Hort. Beng. 31. 1814, fide Benth. 825. Chamaefist ula B (DC.) G. Don, Piel Hist. Dichl. Pl. 2:452. 1832. Cassia arborea . Fl. Jam. 1:343. Sciacassia siamea bam) Britt. in N. PA “FL 23:252. 1930. A moderate-size tree, the branchlets puberulent. Leaves moderately large, with several to many pairs of leaflets; petiole short, eglandular, puberulent; rachis eglandular, flattened and sulcate above, somewhat nodose and cross-partitioned at insertion of the petiolules; stipules small, caducous; up to 30 leaflets, ovate-oblong to oblong, usually about 6 cm. long and 2 cm. wide when well-developed, obtuse to rounded-subtruncate and slightly emarginate apically, obtuse basally, minutely puberulent below, lustrous above; petiolules about 3 mm. long. Inflorescence usually a terminal panicle of short racemes. Flowers rather short-pedicellate; yellow; sepals suborbicular, up to 6 mm. long, puberulent; petals up to 15 mm. long, clawed, glabrous; stamens 10, only 7 fertile, dehiscent by terminal pores, the pores somewhat confluent; 3 lowermost stamens larger (the center one some- what smaller), the anthers up to 7 mm. long, short-rostrate apically, sagittate basally; 4 median stamens shorter, the anthers 5-6 mm. long, less noticably ros- trate; 3 uppermost stamens small, the anthers up to 3 mm. long, apparently non- (348) 1951] FLORA OF PANAMA (Leguminosae) 49 Legume linear, 2—3 dm. long and about 12 mm. wide, flattened, puberulent, undulate between the margins, dehiscent. Native to East Indies; planted in American tropics. functional; ovary pubescent. CANAL ZONE: Balboa, Allen 4555, Standley 30822; “Canal Zone,” Johansen 30. DARIEN: Cana, Williams 948. Fig. 120. Cassia spectabilis 5. CassiA SPECTABILIS DC. Cat. Hort. Monsp. 90. 1813. Cassia speciosa HBK. Nov. Gen. & Sp. 6:338. 1824; non Schrad., fide Benth. Cassi j Humboldtiana DC. Prodr. 2:489. 1825. Cathartocarpus Humboldtianus Loud. Hort. Brit. 167. enth. Pseudocassia spectabilis (DC.) Britt. & Rose, in N. Am. Fl. 23:230. 1930. Tree to 10 or more m. high, the branchlets usually tomentose when young. Leaves moderately large, an average leaf about 20-foliolate; petiole short, pubescent, eglandular; rachis usually about 2 dm. long, eglandular and otherwise like the (349) [Vor. 38 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN petiole; leaflets several to many pairs, lanceolate, 3-8 cm. long and usually about 2 cm. wide, acute apically, obtuse basally, pubescent below, especially along the veins, puberulent to subglabrous above and less dull than below, opposite on the rachis, with 10 or more pairs of prominent lateral veins; petiolules 2-3 mm. long, pubescent. Inflorescence of several terminal or subterminal several-flowered racemes; bracts lanceolate, a few mm. long, caducous. Flowers yellow; sepals 5, obovate-orbicular, markedly unequal, up to 1 cm. long and broad, glabrous to lightly puberulent; petals 5, mostly obovate, markedly unequal, up to 2.5 cm. long and 1.5 cm. broad, subglabrous, venose, short-clawed; stamens 10, 3-morphic; the 3 lowermost the largest, their anthers oblong, about 7 mm. long, short-rostrate apically and dehiscent by terminal pores, the loculi somewhat converging termi- nally; anthers of 4 median stamens 5-6 mm. long, similar to the 3 lowermost except the rostrum reflexed and the loculi divergent terminally; 3 uppermost stamens markedly dissimilar, more or less rudimentary, the anthers distinctly bilobed, each lobe reniform and dehiscent the length of its outer margin; ovary linear, glabrous. Legume linear, turgid-quadrangular, up to 2 dm. long and 1 cm. wide, transversely multiseptate, tardily dehiscent along one margin. Mexico to northern South America; West Indies. CHIRIQUÍ: San Felix, Pittier 5133. The tree is often planted in sections of the American tropics. d CASSIA DIDYMOBOTRYA Fresen. in Flora 221:53. 1839. Shrub or small tree, the branchlets puberulent. Leaves several- to many- foliolate, up to 3.5 cm. long; petiole about 3 cm. long, puberulent, eglandular; rachis similar to petiole; stipules ovate-rotund, acuminate, up to 12 mm. long; leaflets normally 8-16 pairs, oblong or elliptic, up to 5 cm. long and almost 2 cm. wide, basally inequilaterally obtuse, apically obtuse or acute and mucronate, sub- sessile, lightly pubescent, the hairs from a pustuliform base in dried material. In- florescence of a few subterminal racemes up to 3 dm. long, the buds approximate and ensheathed by ovate bracts about 12 mm. long. Flower yellow; sepals elliptic, imbricate, sessile, about 15 mm. long; petals obovate to elliptic, up to 2 cm. long, markedly venose, short-clawed; stamens 4-morphic; anther of lowermost center stamen about 7 mm. long, with terminal pores and lateral slits, prominently hastate basally, this anther bounded on either side by a stamen with a large anther about 12 mm. long, similarly dehiscent, sagittate basally, arcuate; anthers of 4 median stamens subequal, about 5 mm. long, long-rostrate, sagittate, dehiscent as the lowermost; uppermost 2—3 stamens rudimentary, distorted; ovary tomentose. Legume broadly linear, up to about 10 cm. long and 1.5 cm. wide, flat, dehiscent, multiseptate; seeds transverse. Native to northern Africa; introduced into American tropics, Hawaii, and elsewhere. CANAL ZONE: in cultivation at summit, Jobansen s. n. (350) 1951] FLORA OF PANAMA (Leguminosae) 51 If the Johansen specimen typifies the species, it is very little different from C. nicaraguensis. It differs chiefly in pubescence characters and in having smaller, different-shaped stipules. The plant is reported to emit an unpleasant odor when bruised. Fig. 121. Cassia nicaraguensis 7. CASSIA NICARAGUENSIS Benth. in Trans. Linn. Soc. 27:552. 1871. Cassia Seleriana Harms, in Bull. Herb. Boiss. 7:551. 1899, fide Britt. & Rose. Chamaesenna nicaraguensis (Benth.) Britt. & Rose, in N. Am. Fl. 23:250. 1930. Small, glabrous tree. Leaves many-foliolate, usually 20—30 cm. long; petiole elongate, basally rugose-callous; rachis flattened above, eglandular; stipules foli- aceous, about 3 cm. long, rounded apically, acuminate-attenuate basally, indented on inner side at insertion, deciduous from older leaves; leaflets 5—20 pairs, oblong or elliptic, usually about 6 cm. long and 2 cm. wide, basally obtuse, apically obtuse or acute and mucronate, dull below, markedly reticulate; petiolules about 1 mm. long. Inflorescence of several racemes terminal or axillary from the upper petioles; bracts large, caducous; pedicels 1-2 cm. long. Flowers showy, yellow; sepals 5, (351) [Vor. 38 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN obovate to orbicular, up to 2 cm. long, imbricate, sessile, venose; petals obovate to elliptic, up to 2.5 cm. long, very short-clawed, strikingly venose, the veins very ark; stamens with short filaments, 4-morphic; the anther of the lowermost cen- ter stamen about 5 mm. long, with terminal pores and lateral slits, prominently hastate basally, this anther bounded on either side by a stamen with a very large anther about 1.5 cm. long, similarly dehiscent, basally unequally sagittate, the larger lobe on the inner side; 4 median stamens moderate, the anthers dehiscent by terminal pores and lateral slits, apically long-rostrate, basally briefly sagittate; uppermost stamens 2—3, rudimentary, distorted, about 3 mm. long; ovary linear, glabrous. Legume flat, broadly linear, margined, septate, up to 12 cm. long and 1.5 cm. wide. Mexico and Central America. ZONE: vicinity of Е С. White 190, Р. 6 С. White 43; К. Azo САМА Caballo, D odge, Steyermark © Alle 23. PANAMÁ: ЕРІ, ажа. Pittier 2363; vicinity of Arraijan, Allen 1625. VERAGUAS: vicinity of Santiago, Allen 1 8. СазяА RETICULATA Willd. Enum. Hort. Berol. 1:443. 1809. Cassia strobilacea HBK. Nov. Gen. & Sp. 6:347. 1824, fide Benth. Cassia Tarantan HBK. loc. cit. 348. 1824, fide Britt. & Rose. Cassia dumetorum Bert.; DC. Prodr. 2:499. Chamaesenna reticulata Willd.) Pittier, in Trab. Mes. Com. Venez. 3:160. 1928. Small tree, the branchlets puberulent, terete, stout, with a large pith. Leaves large, up to 24-foliolate; petioles relatively short, similar to the rachis; rachis up to 3 dm. long or longer, puberulent, flattened and longitudinally margined above, somewhat nodose and subseptate between insertion of the leaflets, eglandular; stipules lanceolate, up to 1 cm. long, spreading at the base and subauriculate by oblique insertion, puberulent, subpersistent; leaflets 8-12 pairs, oblong to obovate (uppermost), up to 10 cm. long and 5 cm. wide, apically rounded to subtruncate and mucronulate; basally rounded or obtuse and somewhat inequilateral, tomentu- lose below, puberulent above, dull, reticulate; petiolule about 1 mm. long. In- florescence terminal or axillary from the upper nodes, puberulent or tomentulose, racemose, condensed above, the pedicels short; bracts persistent until anthesis, ovate-orbicular, up to 2 cm. long, usually cuspidate apically, finely reticulate, more or less imbricate and ensheathing the buds of the apical portion of the raceme. Flowers yellow, moderately large; sepals elliptic, up to 13 mm. long, delicately veined, the veins mostly longitudinal and subparallel; petals elliptic-orbicular, 16-17 mm. long, clawed, conspicuously dark-veined, glabrous; fertile stamens 6 (4 others present but rudimentary) ; the lowermost center one with a long filament (about 4 mm. long), its anther markedly sagittate, about 3 mm. long; anthers of the 2 lowermost lateral ones large, linear-elliptic, falcate, about 11 mm. long, dehiscent by a pair of terminal pores, apically short-rostrate and acute, basally unequally sagittate; 4 median anthers oblong, about 4 mm. long, prominently rostrate apically, dehiscent by a pair of terminal pores, basally sagittate; 3 upper- most stamens rudimentary, the anthers 2—3 mm. long; ovary linear, tomentulose. (352) 1951] FLORA OF PANAMA (Leguminosae ) 53 Legume linear-oblong, up to 15 cm. long and 2 cm. wide, thin, septate, glabrous, short-stipitate, margined but not alate; seeds transverse linear. Mexico to Brazil and Bolivia; Trinidad. BOCAS DEL TORO: Changuinola River, Dunlap 205; Old ees Island, von Wedel 1626 1862, 1988; Water Valley, H. von Wedel , 1757. CANAL ZO Ancón, Mason б; "ob pres Fendler 95; Gatün, ans 4791; Miraflores, P. © б. "White 44. COCLÉ: Penonomé, Williams 364. DARIÉN: Garachiné, Pittier 5509. ANAMA: Juan Diaz, Standley 30482; К. Tapia. Standley 28304; Las Sabanas, Standley 252. 9. Cassia ALATA L. Sp. Pl. 378. 1753. Cassia berpetica Jacq. Obs. Bot. 2:24, BRE. 20 1757. Cassia bracteata L. f. Suppl. 232. ізі, үр enth. Herpetica alata (L.) Raf. Sylva Tellar. 123. 1838. Shrub to 3 m. tall, the branchlets minutely puberulent to subglabrous, stout, terete. Leaves large, up to 28-foliolate; petioles relatively short, like the rachis; rachis elongate, up to 3 or more dm. long, flattened and margined above, somewhat nodose and septate between the insertions of the paired leaflets, minutely puber- ulent, eglandular; stipules lanceolate-subulate, usually about 1 cm. long, basally subauriculate by oblique insertion; leaflets 5-14 pairs, oblong to obovate or the lowermost sometimes ovate, up to 17 cm. long and 9 cm. wide, the uppermost largest, apically rounded and mucronulate (occasionally subretuse), basally rounded- subtruncate and obliquely inequilateral, subglabrous above and below, dull below, chartaceous, reticulate; petiolules 1-2 mm. long. Inflorescence terminal or sub- terminal, the many-flowered raceme appearing spike-like because of the short pedicels; bracts foliaceous, ovate or oblong, up to 2.5 cm. long and 2 cm. wide, persistent until anthesis and ensheathing the upper raceme, minutely puberulent, imbricate. Flowers yellow, conspicuous; sepals oblong, up to 15 mm. long and 8 mm. wide, delicately reticulate-venose, minutely puberulent or subglabrous; petals obovate to orbicular, up to 2 cm. long and 12 mm. wide, prominently dark-veined, clawed; stamens 10, 4-morphic; center stamen of the 3 lowermost with an elongate filament 6—7 mm. long, the anther small (about 4 mm. long), obliquely rostrate and dehiscent by 2 terminal pores, subsagittate basally; 2 lateral stamens of the lowermost group large, falcate, the filaments 2-3 mm. long, the anthers 11—12 mm. long, unequally bilocular, apically obliquely short-rostrate and dehiscent by a pair of terminal pores, basally each loculus prolonged as a subulate tip; 4 median stamens with filaments about 2 mm. long, the anthers 3-4 mm. long and similar to that of the lowermost center stamen; upper 3 stamens minute, non-functional, twisted, about 2 mm. long; ovary densely puberulent, arcuate. Legume broadly linear, about 15 cm. long and 1.5 cm. wide, longitudinally alate from the middle of each valve (the wing about 5 mm. wide), dehiscent, septate, straight, charta- ceous; seeds transverse, compressed, rhombic. Mexico to Paraguay; Old World tropics. (353) [Vor. 38 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONE: France ngs Standley 30438. сосіЕ: Ola, Pittier 5088. PANAMA: etween Panamá and Chepo, Dodge, Hunter, Steyermark © Allen 16646; Rio Tecümen, Б 26677; Taboga iden Macbride 2704. УЕКАСОА$: Sona, Allen 1075. Distinguished by the alate legume, foliaceous bracts, large lateral anthers, venose perianth, and the many subseptate, eglandular rachial (foliolar) nodes. 10. CASSIA EMARGINATA L. Sp. Pl. 376. 1753. Cassia atomaria L. Mant. 68. 1767, fide N. Am Cassia grisea A. Rich. in байга, Hist. Cubá Not. LI 1845, ex char., fide Benth. Cassia emarginata subunijuga Robin. & Bartl. Proc. Am. Acad. 43:53. 1907, fide N. Am. Fl. als emarginata (L.) Britt. & Rose; Britt. & Wilson, Scientif. Surv. Porto Rico & n Isls. 5:374. 1924. Shrub or small tree, the branchlets pubescent. Leaves small or moderate, usually 4-foliolate, petioles relatively long, up to 6 cm., more or less terete, tomentulose; rachis shorter than petiole (occasionally absent in 1-jugate leaves), up to 3 cm. long, similar to petiole, eglandular; stipules small, linear-lanceolate or subulate, 2-3 mm. long; leaflets ovate, obovate or elliptic, up to 10 cm. long and 5 cm. wide (but usually smaller), 1—5 pairs, apically rounded or obtuse and usually retuse or emarginate and mucronulate, basally rounded or obtuse, puberu- lent above, tomentulose beneath, secondary veins prominent and subparallel; petiolules 2—5 mm. long. Inflorescence racemose, axillary or subfasciculate from the upper nodes, short, few-flowered, pubescent; pedicels up to 3 cm. long; bracts lanceolate, about 3 mm. long, caducous. Flowers small, yellow; sepals ovate- orbicular, markedly unequal, the largest about 6—7 mm. long and 5 mm. broad; petals mostly elliptic or obovate, unequal, up to 15 mm. long and ! cm. wide, glabrous; fertile stamens 7, similar but somewhat unequal; anthers usually 4—5 mm. long, linear-oblong, erostrate and subtruncate apically or essentially so, de- hiscent by a pair of terminal pores; 3 uppermost stamens modified as staminodes or rudimentary, flat, about 2 mm. long, with lateral slits; ovary linear, subglabrous. Legume up to 3.5 dm. long and about 1 cm. wide, linear, straight or only slightly curved, glabrous, flattened, not impressed but the margins often undulate, in- dehiscent. Mexico; Central America; West Indies; Colombia and Venezuela. This species has not to my knowledge been collected in Panama, but certainly is to be expected there. It is found in the lowlands in both Costa Rica and Colombia. 11. Cassia HisPIDULA Vahl, Eclog. 3:10. 1807. Cassia procumbens Mill. Gard. Dict. ed. 8, no. 20. 1768, non L. Sp. Pl. 1753, fide N. Am. FI. (354) 1951] FLORA OF PANAMA (Leguminosae) 55 Cassia hispida Collad. Hist. Cass. 118. 1816, fide Benth. Cassia tetraphylla Collad. loc. cit. 130. 1816, non Desv. 1814, fide N. Am. Fl. Cassia pauciflora HBK. Nov. Gen. & Sp. 6:360. 1824, fide N. Am. Fl. Cassia lotoides HBK. loc. cit. 361. 1824, fide Benth. Cassia fagonioides Vogel, Syn. Gen. Cass. 50. 1837, fide Benth. Cassia leiantha Benth. in Hook. Jour. Bot. 2:78. 1840, fide Benth. Grimaldia hispidula (Vahl) Britt. & Rose, in N. Am. Fl. 23:299. 1930. Semiprostrate herb from a woody rootstock, the slender, frequently elongate branches viscid-pubescent, usually with both long and short hairs. Leaves small, typically 4-foliolate; petioles elongate, longer than the leaflets, 2-3 cm. long, pubescent like the stem but otherwise eglandular; rachis short, less than 1 cm. long, similar to the petiole although less pubescent; stipules linear-lanceolate, minute; leaflets ovate-orbicular to obovate-orbicular, up to 2 cm. long and almost as broad, obtuse or rounded both apically and basally, essentially glabrous, short- petiolulate. Inflorescence usually a few-flowered terminal raceme; the pedicels slender, pubescent like the stem, minutely bibracteate. Flowers yellow-orange, fairly conspicuous, about 2 cm. wide, borne in few-flowered terminal racemes; sepals ovate to ovate-lanceolate, about 1 cm. long, frequently setose without; petals obovate, 1.5—2 cm. long, rounded apically, subcuneate basally and narrowed into a claw-like base; stamens 10, all similar and subequal, the anthers linear, 5—7 mm. long, longitudinally pubescent laterally, dehiscent by paired terminal pores; ovary pilose, short. Legume linear or linear-oblong, 3—5 cm. long and about 7 mm. wide, flat, setose-hirsute, elastically dehiscent, the seeds obliquely transverse. Mexico to Brazil and northern South America; West Indies. The species is reported from Panama, and a specimen from Costa Rica has been examined. 11а. Cassia HisPIDULA Vahl var. Killipii (Rose) Schery, comb. nov. Cassia Killipii Rose, in Jour. Wash. Acad. 17:167. 1927. Grimaldia Killipii (Rose) Britt. & Rose, іп N. Am. Fl. 23:301. 1930. Generally of smaller proportions than the species, the flower scarcely 1.5 cm. broad and the leaflets mostly 1 cm. or less in length. The leaflets are more or less pubescent or setulose below, the key character used by Britton & Rose (N. Am. Fl. 23:299. 1930) in separating Grimaldia Killipii from G. hispidula. This char- acter alone seems to show some intergradation and lack of correlation with other characters, necessitating, in the author's opinion, reduction of Cassia Killipii to varietal status. Possibly C. fagonioides Vog., listed among synonyms for the species, is synonymous with this variety, being described as pubescent by Bentham. British Honduras, Panama, Colombia. cocLÉ: La Pintada, Hunter & Allen 521; Nata, Allen 813; Ola, Pittier 5014; Penonomé, Williams 104, PANAMA: Pacora, Paul 258; Río Tapia, Killip 3281, Standley 28186, 30656. (355) [Vor. 38 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN 12. CASSIA ROTUNDIFOLIA Pers. Syn. Pl. 1:456. 1805. руне rotundifolia (Pers. ) Greene, in Pittonia 4:31. nal synonyms are given by Bentham, especially for his South American varieties baubiniacfolia and grandiflora Semiprostrate herb, as of savanna habitat, the stems pubescent to sub- glabrous. Leaves bifoliolate, small; petiole short, not exceeding the stipules, egland- ular, pubescent like the stem; stipules lanceolate-cordate, ciliate or glabrous, up to about 1 cm. long; leaflets 2, asymmetrically subrotund to broadly obovate, rounded apically, 0.5-3 cm. long, sometimes ciliate, epetiolulate. Flowers 1-2 from the axils, small, yellow, the pedicel more or less filiform; sepals lanceolate, usually ciliate, up to 5 mm. long; petals obovate, about 6 mm. long, glabrous, sessile; fertile stamens apparently 5, all similar although somewhat unequal, the filaments very short; the anthers linear-oblong, up to 2 mm. long, essentially glabrous and erostrate, dehiscent by paired terminal pores; ovary pubescent. Legume linear, 1.5—4 cm. long and 3—5 mm. wide, flat, elastically dehiscent, the seeds obliquely transverse. Mexico; southern Central America; northern South America; West Indies. CHIRIQUI: Boquete, Davidson 700, Maxon 5131, Pittier 3311; Gualaca, Allen 5044. Possibly the Boquete specimens cited represent an undescribed variety of the species, all of them being more pilose, with smaller leaflets, and with shorter pods (fewer loculi) than is normal for South American, West Indian and Mexican specimens. Bentham, however, considered plants of larger proportions as varieties of the species, and it would perhaps be presumptuous to describe the Panamanian plants as a new variety without first comparing them with additional and more representative (European) material than is here available. Apparently the species, considered in its broadest sense, is quite variable. 13. CASSIA DIPHYLLA L. Sp. Pl. 376. 1753. Chamaecrista diphylla (L.) Greene, in Pittonia 4:28. 1899. Distinctive annual or perennial herb of open or seasonally dry areas such as savannas, the stems glabrous. Leaves small, bifoliolate, glabrous; petiole 5-6 mm. long, bearing 1—2 subcupuliform glands on the upper side towards the middle; stipules large, lanceolate-cordate, up to 15 or more mm. long, foliaceous and sub- ensheathing the stems, membranaceous; leaflets asymmetrically obovate, usually about 2 cm. long and 1 cm. broad, the conspicuous veins subparallel, epetiolulate, with a broad (about 2 mm.), callous insertion. Flowers yellow, borne singly from the axils on elongate petioles, the petioles up to 5 cm. long; sepals 5, gluma- ceous, unequal, up to 1 cm. long and 1-2 mm. wide, glabrous, apiculate, the prominent veins subparallel; petals obovate, about as long as the sepals, glabrous to puberulent, sessile; stamens 10, all similar, the anthers linear although somewhat (356) 1951] FLORA OF PANAMA (Leguminosae ) 27 Fig. 122. Cassia diphylla wider basally than apically, truncate, dehiscent by paired terminal pores, sub- glabrous; ovary pilose, short. Legume linear, up to 9 cm. long and about 6 mm. wide, flat, oblique apically and basally, lightly pilose, dehiscent. Mexico; Central America; West Indies; northern and eastern South America. ocLÉ: Nata, Allen 815. panamá: R. Azote Caballo, Dodge, Steyermark & Allen me El Vigia, Pittier 2301; Taboga Island, Allen 109. A species very different from the tree forms of Cassia, easily distinguished by characteristic leaves and stipules and glumaceous sepals. It is similar in habit and generally savanna habitat to C. brevipes, C. Tagera, and C. rotundifolia. 14. Cassta ТАСЕВА L. Sp. Pl. 376. 1753. Cassia ciliaris Collad. Hist. Cass. 98. Cassia Kunthiana Cham. & Schlecht. in titan 5:598. 1830, fide Benth. Tagera filiformis Raf. Sylva Tellur. 129. 1838. Chamaecrista Tagera Standl. in Contr. U. S. Nat. Herb. 18:104. 1916. Prostrate herb or subshrub, the branchlets somewhat pubescent, at least near the tips. Leaves very small, the leaflets 2-3 pairs; petiole 5 mm. or less long, bearing a stipitate gland above the middle, usually pubescent or ciliate above; rachis about 2 mm. long, mucronulate apically; stipules lanceolate-cordate, 4—10 mm. long, subensheathing the stem, glabrous to ciliate; leaflets very small, at most 10 mm. long and 5 mm. wide, asymmetrically obovate or obcuneate, subglabrous to ciliate, epetiolulate, with 2-3 prominent primary veins. Flowers yellow, solitary from the axils, the pedicels usually exceeding the leaf; sepals 5, narrowly ovate- (357) [Vor. 38 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN lanceolate, about 3 mm. long and 1 mm. wide, unequal, acuminate; petals 5, obovate, up to 5 mm. long, glabrous or lightly ciliate; stamens 7 or less (usually only 4 or only 4 larger and fertile), unequal but similar, the anthers linear, up to 2 mm. long, truncate apically and dehiscent by paired terminal pores; ovary pilose, short. Legume oblong, scarcely 1 cm. long and 4 mm. wide, oblique apically and basally, flat, strigose, pubescent, 1- to 4-seeded, evidently dehiscent. Mexico to northern South America. ZONE: Las Cruces trail, Hunter 9 Allen 754. cocré: Aguadulce, Pittier Pn Ж. Williams 105. PANAMA: Capira-Potrero, Dodge 9 Hunter 8606; sabanas near Chepo, Hunter & Allen 18; Hacienda La Joya, Dodge, Hunter, Steyermark & Allen 16011; Pacora, Killip 3243. veracuas: Sona, Allen 1040. Panama plants of this species seem to have a consistently smaller legume than is common for plants from northern Central America. 15. Cassia BREVIPES DC.; Collad. Hist. Cass. 119. 1816. Chamaecrista brevipes (DC.) Greene, in Pittonia 4:31. 1899. A shrub of open, seasonally dry places, the branchlets usually pilose-tomentose. Leaves small, bijugate; petiole about 5 mm. long, pubescent, bearing a sessile, cupuliform gland in the upper portion; rachis shorter than the petiole; stipules lanceolate-cordate, up to 1 cm. long, subensheathing the stem, venose with promi- nent subparallel veins; leaflets 4, asymmetrically obovate or obovate-elliptic, up to 2 cm. long and 5-10 mm. wide, subcoriaceous, glabrous to lightly pubescent, the prominent veins subparallel. Flowers arising singly from the axils, yellow, usually short-pedicellate; sepals 5, unequal, narrowly lanceolate, up to 1.5 cm. long and 5 mm. wide, scarious-coriaceous or almost glumaceous, the veins subparallel; petals 5, obovate, up to 23 mm. long and 13 mm. wide, glabrous, membranaceous; fertile stamens evidently 10 all similar, the anthers linear, 7-9 mm. long, puberulent laterally along the margins, dehiscent by paired terminal pores; ovary pilose, short. Legume oblong, up to 3.5 cm. long and 1.2 cm. wide, oblique apically and basally, flat, pilose, elastically dehiscent, the seeds transverse, linear. Central America and northern South America. cocLÉ: Olá, РИНет 5013; Penonomé, Williams 126. 16. Cassia FLEXUOsA L. Sp. Pl. 379. 1753. Cassia arenaria HBK. Nov. Gen. & Sp. 6:370. 1824, pos = bématerisia amlistipulata 1. Contr. 1). S. Nat. "id 12: 267. 1909, pl N. Am. Fl. Cassia picachensis Brandg. in Univ. Calif. Publ. Bot. 6:180. 1915, fide М. Fl. Small perennials of open slopes or moist savannas, erect from a woody root, the stems flexuous, glabrous or pubescent. Leaves moderate, multifoliolate; petioles short, scarcely 5 mm. long, usually pubescent above and bearing 1-2 glands, the glands sessile to stipitate, terete, expanded apically, concave at the apex; rachis up to 8 cm. long, flattened and somewhat sulcate above, more or less (358) 1951] FLORA OF PANAMA (Leguminosae) 59 cross-partitioned at insertion of the leaflets; stipules obliquely cordate, lanceolate, up to 1 cm. long and 5 mm. wide, erect, longitudinally venose; leaflets up to 50 pairs, linear, up to 1 cm. long and 1.5 mm. wide, inequilateral basally, obliquely mucronulate apically, with 2-3 prominent subparallel veins. Flowers mostly solitary from the axils, slender-pedicellate, yellow; sepals ovate-lanceolate, about 8 mm. long and 3 mm. wide, acute or acuminate apically; petals up to almost twice as long as the sepals, obovate, scarcely clawed; stamens 10, unequal but all apparently fertile; the anthers linear (although narrower apically), about 8 mm. long in the largest and scarcely 3 mm. long in the smallest, subsessile, dehiscent by paired terminal pores; ovary pubescent, linear. Legume linear, about 5 cm. long and 4—5 mm. wide, flat, scarcely stipitate, elastically dehiscent, transversely (obliquely so) several-seeded. Mexico to Brazil, Paraguay and Bolivia; West Indies. cocLÉ: Penonomé, Williams 114. 17. CassiA STENOCARPA Vogel, Syn. Cass. 68. 1837. Chamaecrista stenocarpa (Vogel) Standl. in Contr. U. S. Nat. Herb. 18:104. 1916. Cassia Brougbtonii Fawc. & Rend. in Jour. Bot. 55:37. 1917 C riparia DAS in Bull. Torrey Bot. Club. 44: 11, in part. 1917, not Cassia riparia HBK. 1824. A more or less erect annual or subperennial up to about 1 m. high, the branch- lets pilose or subpilose with long spreading hairs. Leaves moderately small, multi- foliolate; petiole short, usually 3-4 mm. long, pubescent, bearing on the upper side above the middle 1 (sometimes 2) gland, the glands stipitate, expanded or flaring and concave at the apex; rachis pubescent, sulcate above, more or less cross- partitioned at insertion of the petiolules; stipules lanceolate, up to 14 mm. long and scarcely 2 mm. wide, erect, acuminate apically, somewhat oblique канш): leaflets up to 25 pairs, linear-oblong, usually about 12 mm. long and 2-3 mm. wide, lightly pubescent or ciliate, the midvein excentric, mucronate e. in- equilateral basally, sessile. Peduncles short, 1- to 3-flowered, borne on the inter- node a short distance above the axils; pedicels about 15 mm. long, bibracteate. Flowers yellow, rather inconspicuous; sepals 5, ovate-lanceolate, about 7 mm. long, ciliate-pubescent, acuminate-attenuate; petals obovate, 6—7 mm. long, sessile; stamens evidently 10, similar although markedly unequal; the anthers linear, up to 4 mm. long, dehiscent by paired terminal pores; ovary linear, pilose. Legume linear, up to 6 cm. long and about 4 mm. wide, flat, lightly pubescent, elastically dehiscent, the seeds obliquely transverse. Mexico to northern South America; West Indies. ANAL ZONE: Balboa, Standley 25231, 27142, 20239; Cerro Gordo, Standley 26031; Culebra, Pittier 2104; Frijoles, Pittier 6332, Standley 27657; Gamboa, Standley 28337. : Chepo, Pittier 4749; Matías Hernandez, Pittier 6874; К. Tecümen, Standley 20475. The entity here considered as C. stenocarpa is scarcely distinct from many allied “species” of Central America, South America and the West Indies. For (359) [Vor. 38 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN example, no definite delimitation seems possible between specimens of C. stenocarpa compared with C. riparia (of HBK.), C. glandulosa, C. Chamaecrista and a num- ber of Britton & Rose segregates as they are found in the herbaria. To avoid further confusion, most of these various species are here regarded as distinct as they are found in Panama; however, it is recognized that reference to types and intergrading forms from all of tropical America would likely necessitate extensive condensation and reduction. The name C. glandulosa L. or C. Chamaecrista L. might well profitably apply to this whole complex of similar forms, with a few varietal names maintained for certain localized "species." "e Ap 422, FR 2% / Fig. 123. Cassia stenocarpa var. stenocarpoides 17a. Cassia STENOCARPA Уор. var. stenocarpoides (Britt.) Schery, comb. nov. Chamaecrista stenocarpoides Britt. in М. Am. Fl. 23:293. 1930. Cassia stenocarpoides (Britt.) Lundell, in Phytologia 1:215. 1937. Vegetative characters and inflorescence as described for C. s/emocarpa, except the petiolar glands usually shorter or thicker. Flowers larger than in the species; sepals ovate-lanceolate, about 8 mm. long and 2-4 mm. wide; petals obovate, the (360) 1951] FLORA OF PANAMA (Leguminosae) 61 larger about 1 cm. long; stamens apparently 10, the filaments almost obsolete, the anthers all similar but markedly unequal; largest anthers about 7 mm. long, de- hiscent by terminal pores and longitudinally pubescent-ridged laterally; ovary usually strigose-pilose. Legume as described for the species. Costa Rica and Panama; probably northern Central America and northern South America as well. снікюсі: R. Chiriqui Viejo, Peggy White 36. As stated in discussion of C. stenocarpa, correct application of the specific name cannot be certain without reference to types and additional collections not here available. While this variety seems fairly distinct in Costa Rica, both to the north and in South America various "species" appear doubtfully different. 18. Cassia FLAVICOMA HBK. Nov. Gen. & Sp. 6:366. 1824. Cassia stipulata G. Don, Gen. Hist. Dichl. Pl. 2:448. 1832, fide Benth. Chamaecrista chiriquensis Britt. & Rose, in N. Am. Fl. 23:287. 1930. Small suffrutescent shrub, the branchlets usually pilose with prominent yellow hairs. Leaves moderate, multifoliolate; petioles about 5 mm. long, heavily pubes- cent, bearing a solitary, stipitate, subpeltate gland above the middle; rachis usually 5—7 cm. long, pubescent like the petiole; stipules narrowly lanceolate, 1 cm. long or longer, acuminate, ciliate-margined; leaflets up to 30 pairs, linear, up to 18 mm. long and 2-3 mm. wide, membranous, glabrous or nearly so above, appressed- pubescent below, rounded and mucronate apically, inequilateral basally, the veins faint. Inflorescence borne from the internode, 1- to few-flowered, the peduncle very short; pedicels almost 1 cm. long, pubescent, bibracteolate above the middle. Flowers yellow, usually 12 or more mm. long; sepals ovate-lanceolate, 9—12 mm. long and mostly 1-2 mm. broad, strigose; petals obovate, unequal, up to 14 mm. long and 10 mm. broad; stamens 10, all similar but markedly unequal, the fila- ments essentially obsolete, the anthers dehiscent by paired terminal pores and with lateral pubescnt ridges; 3 longest anthers 11—12 mm. long, linear, strongly arcuate; remaining anthers 2-5 mm. long; ovary about 4 mm. long, subtomentose. Legume linear, about 5 cm. long and 6 mm. wide, dehiscent, oblique basally and apically, lightly pubescent to subglabrous on the valves; seeds obliquely transverse. Mexico, Panama, South America, West Indies. CHIRIQUÍ: San Felix, Pittier 5216. The single specimen known from Panama is the type for Chamaecrista chiri- quensis Britt. & Rose. There seems to be no definite, tangible difference betweén it and Mexican, West Indian and South American specimens listed in herbaria as Cassia flavicoma, C. glandulosa and C. riparia. In the absence of types, C. flavi- coma is chosen as the name most appropriate to the specimen in question; but, as mentioned in discussion of C. stenocarpa, monographic work would perhaps show this to be best treated as a variety of some earlier name such as C. glandulosa. (361) [Vor. 38 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN 19. CasstA PATELLARIA DC.; Collad. Hist. Cass. 125. 1816. Chamaecrista patellaria (DC.) Greene, Pittonia 4:32. 1899. An herbaceous perennial (or annual?), erect, to a few dm. tall, the branchlets densely pubescent with ascending hairs. Leaves moderately small, multifoliolate; petiole short, 2-7 mm. long, pubescent, bearing on the upper side 1—2 sessile, patelliform glands about 1 mm. long; rachis usually 4—5 cm. long, sulcate above, more or less cross-partitioned at insertion of the petiolules; stipules linear-lanceolate, up to 13 mm. long, oblique basally, attenuate-acuminate apically, ciliate; leaflets 25 or fewer pairs, linear-oblong, up to 16 mm. long and 3 mm. wide, inequilateral basally, mucronate apically by extension of the midvein, strigose-pubescent, the hairs from a pustuliform base (in dried material), the midvein excentric. Flowers 1—4 from the internodes, yellow, the common peduncle very short, the pedicels at most 1 cm. long, bracteate; sepals ovate-lanceolate, about 5 mm. long, pubescent; petals obovate, up to 11? mm. long (usually about 5 mm. in Panama), glabrous, sessile; stamens usually 8—10, all similar although markedly unequal, the anthers linear or linear-oblong, 1-3.5 mm. long, longitudinally pubescent laterally, de- hiscent by paired terminal pores; ovary densely pubescent. Legume linear, up to 5 cm. long and 5 mm. wide, flat, pubescent, elastically dehiscent, the seeds ob- liquely transverse. Mexico to northern South America; West Indies. NAL ZONE: Balboa, Standley 26437, 321543 Corozal, Standley 27300; Culebra, Pittier 4828; Summit, Standle ey 30053. PANAMA: Camino de las Sabanas, Bro. Heriberto 260; Corozal road, Standley 26798. The species is very similar to C. simplex, differing chiefly in the pronounced leaf pubescence. 20. CASSIA SIMPLEX (Standl.) Standl. in Contr. U. S. Nat. Herb. 27:199. 1928. Chamaecrista simplex Standl. in Contr. U. S. Nat. Herb. 18:103. 1916. Chamaecrista Browniana Britt. & Rose, in N. Ат, Fl. 23:293. 1930. An erect annual (or perennial?) to several dm. tall, the stems appressed- pubescent with short, incurved hairs. Leaves moderate, multifoliolate; petiole about 5 mm. long, pubescent like the stem, bearing on the upper side 1—2 sessile, patelliform glands about 1 mm. long; rachis similar to petiole, up to about 10 cm. long, more or less cross-partitioned at insertion of the petiolules; stipules linear- lanceolate, up to 15 mm. long, oblique basally, attenuate-acuminate apically, ciliate; leaflets up to 25 or more pairs, linear-oblong, usually about 8 mm. long and scarcely 2 mm. wide, glabrous, mucronate apically by extension of the excentric midvein, inequilateral basally. Flowers usually 2—3 from the internodes, yellow, the common peduncle very short, the pedicels up to 4 mm. long, bibracteate; sepals ovate-lanceolate, 5-9 mm. long, lightly pubescent; petals obovate, up to about 1 cm. long, sessile, glabrous; stamens 10 (sometimes less?), markedly unequal, all similar, the anthers linear, 2—5 mm. long, longitudinally puberulent laterally, de- (362) 1951] FLORA OF PANAMA (Leguminosae) 63 hiscent by paired terminal pores; ovary pilose. Legume linear, 3—4 cm. long and up to 5 mm. wide, flat, elastically dehiscent, pubescent, the seeds obliquely trans- verse. Guatemala, Panama and Colombia. CANAL ZONE: Ancón, Killip 3023; Standley 25104, 26342. PANAMA: Dormisolo, Pittier 4655; Hacienda La Joya, Dodge, Hunter, Steyermark & Allen 16805; Isla Taboga, Woodson, Allen 9 Seibert 1430; Standley 28014; between Pacora and Chepo, Woodson, Allen & Seibert 1659; Pacora, Killip 3324; R. Tecümen, Standley 22660. The species is very similar to C. patellaria, differing in having glabrous leaves and more appressed stem pubescence. Judging from material seen, these differ- ences are constant and without significant intergradation. The Dodge, Hunter, Steyermark & Allen 16805 specimen cited differs from most specimens in having larger, more abundant leaflets and the petiolar glands more or less oblong. It thus could be regarded as Chamaecrista (Cassia) Browniana Britt. & Rose, were that species to be considered valid. However, the only distinguishing feature of Chamaecrista Browniana seems to be the larger longitudinal proportions of the petiolar gland, a character scarcely of specific weight in a group where petiolar glands are notably variable. Unless further collections can prove the contrary, it would seem wise to include Chamaecrista Browniana as but a variation of Cassia simplex. 21. CASSIA LEPTOCARPA var. HIRSUTA Benth. in Trans. Linn. Soc. 27:531. 1871. Ditremexa leptocarpa (Benth.) Britt. & Rose, in N. Am. Fl. 23:256. 1930 (in part). Shrub to 2 m., with pubescent branchlets. Leaves pubescent (in Panama), several-foliolate; petiole up to 10 cm. long, angled, with a conspicuous basal gland about 6 mm. above the insertion; rachis lightly pubescent to hirsute; stipules linear, about 1 cm. long, caducous; leaflets usually about 5 pairs, ovate to ovate-lanceolate, acute and somewhat acuminate apically, obtuse to acute basally, 3—10 cm. long and up to 3.5 cm. wide. Inflorescence a terminal or axillary, several-flowered, condensed raceme; pedicels usually about 1.5 cm. long, pubescent. Flowers yellow; sepals elliptic to suborbicular, 6—8 mm. long, the outermost densely pubescent; petals obovate to suborbicular, up to 14 mm. long, venose, short-clawed; stamens 4-morphic, all dehiscent by apical pores, 7 fertile; the lowermost with a slender filament, with the anther oblong, about 3 mm. long, briefly sagittate basally, bounded on each side by a large stamen with a thick filament, with the anther 6—7 mm. long, very briefly sagittate basally, apically with an erect orbicular rostrum above the pores; 4 median stamens similar to large ones but their anthers 4—5 mm. long; 3 upper stamens non-functional, the anthers flat, orbicular-oblong, about 2 mm. long; ovary setose-hirsute; the style short. Legume linear, up to 30 cm. long and 3—4 mm. wide, thin, flattened, pubescent (in Panama), the margins ridged. Mexico; Central America; West Indies; South America to Paraguay. (363) Vor. 38 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 124. Cassia leptocarpa var. hirsuta AS TORO: ини ИР, von pre I3; vicinity of Chiriquí Lagoon, von Wedel i. "nut | Chorrens Porterfield Bentham’s variety /irsuta is not recognized by Britton & Rose in the ‘North American Flora.’ 22. Cassia OCCIDENTALIS L. Sp. Pl. 377. 1753.1 Cassia fal cata L. loc. cit. 1753. 30. Cassia caroliniana Walt. Fl. Car. 134. 1788, = Britt. & Rose. Cassia linearis Michx. Fl. Bor. Am. 1:261. 18 1Britton and Rose (N. Am. Fl.) regarded C. Sophera б distinct from С. occidentalis. ever, the spe E ns co Panama hs h the ey consi ide = to e C. S ү the typical ciden d, comparison of W ial C. bid n 4. Me 85, cited by зден ы diii ‘hat аа two species are (364) 1951] FLORA OF PANAMA (Leguminosae) 65 Cassia foetida Pers. Syn. Р]. 1:457 (sub E occidentalis). 1805, fide Benth. Cassia ciliata Raf. Fl. Ludovic. 100. 1817, fide Britt Cassia geminiflora Schrank, Hort. Monac. e 26. 1819, fide Benth. Cassia obliquifolia Schrank in Denkschr. Bot. Ges. Regensb. 2:40. 1822, ex char., fide Benth. Senna occidentalis Link, Handb. 2:140. 1831. Ditremexa que. bes 22, Britt. & Rose ex Britt. & Wilson, Scientif. Surv. Porto Rico & Virgin Isls Glabrous shrub or herb to 2 m. tall. Leaves moderate, generally 10-foliolate; petiole 2-5 cm. long, somewhat flattened and sulcate, bearing a dark, sessile, globose gland a few mm. from the base; rachis up to 15 cm. long, sulcate, eglandular; stipules ovate to narrowly lanceolate, caducous; leaflets normally 4—6 pairs, ovate or ovate-lanceolate, up to 10 cm. long and 3 cm. wide, apically acuminate, basally rounded or obtuse and somewhat oblique, glabrous, obscurely reticulate, dull, membranaceous; petiolules about 2 mm. long, with a few short, curved hairs above. not specifically distinct, and that C. зна (with its (us qn synonyms) probably should be included among the synonyms of C. occidentalis, or at merely as a variety of that species (into which Fendler $$ and Hayes 880 specimens wo Midi then seem to fall). As a variety, Sopbera would be expected to have proportionally shorter, more turgid iet less coarseness; smaller average leaflet; and possibly smaller, narrower bracts of the x a cence. There is, however, no good di а tie-in with this separation. It would thus seem best to regard C. Sopbera and C. occidentalis as cific until intensive subspecific research can indicate accurately if and how any further блг: eee may occur. Cassia Sophera carries with it a great number of t listed a Britt. & Wilson, Scientif. Surv. orto = and Vien va 5:372. 1924. Bentham, in "Trapa Linn 2:2 путп Cassia ruscifolia Taeq 1 c P0 70125178 96, ex icone. 2055 { С 5С 2 1 char Cassia aegyptiaca Willd. Enum. Hort. ae 442. 1809, ex char. j . . Cassia sopheroides C Colla. Ha Саз, ме Cassia Barclayana Sweet, Fl. Austral. 4 p 18 ане Sb. G. Don, Ge en. Hist. pii РІ. 2:452. 1832. Senna esculenta Roxb. Fl. Ind. 2:346. 183 Cassia ira ifo he DC. in Mem. Soc. Phys с f 299. 1836. Cassia atroviridis? Span. in Dunes 15: 201. 1841, char Cassia lanceolata Forsk. Fl. Aegypt.-arab., ex parte; Beck. in Bot. Zeit. 897, /. IO. 1850, non alio QE Шы, ы . in Herb. Mus. Brit. TORULO 9) а Jacq. " Collect. 1: б Cassia torosa Cav. Descr. 131. 18 char Cassia indica Poir. in Lam. En 2 Meth Suppl. 2:1272 1811. Cassia torulosa Poir. loc. cit. 126. Chamaefistula chinensis С. Don, Gen. ны. и pl^2:452. 1832. Chamaefistula torosa С. Don, loc. cit a 5 с. Кеў. ей. ap f 89. Cassia coromandeliana Jacq. Fragm. Bot. 67 1000. 1809, ex char. Стено о ии С. Don, Сеп. (зө Dichl. Pl. 21452: 1832] For var. PURP Cassia purpurea Roxb. Hon. us she 2. Senna purpurea Roxb. Fl. Ind. 2:342. Senna Sophera var. ee Roxb. E cit. 347. 1832. (365) [Vor. 38 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 125. Cassia occidentalis Inflorescence terminal to axillary in lower leaves, few- to several-flowered, race- mose or sometimes paniculate; bracts lanceolate, about 1 cm. long, caducous; pedicels rather short. Flowers yellow; sepals ovate to obovate, up to 9 mm. long, glabrous, imbricate; petals oblong or obovate, up to 15 mm. long, short-clawed, glabrous, venose, the veins scarcely reticulate; stamens 4-morphic, 6—7 of them fertile and dehiscent by a pair of approximate, oblique, apical pores; lowermost center stamen with a relatively long filament and small rostrate anther, the anther usually rudimentary; 2 lowermost lateral stamens with stout filaments, the anthers elliptic-subfalcate, about 7 mm. long, rostrate; 4 median anthers oblong, 4—5 mm. long, short-rostrate; uppermost 3 stamens rudimentary, represented by flattened, clawed, subspatulate staminodes; ovary linear, pubescent. Legume linear-oblong, up to 15 cm. long and 9 mm. wide, glabrous, flattened, margined, scarcely stipitate, septate, slightly curved; seeds transverse. Southern United States to Paraguay; Old World tropics. BOCAS DEL TORO: von Wedel 260; Isla Colón, von Wedel 478; “western Panama", Stork 74. CANAL ZONE: “Chagres”, Fendler 85; Matachin, Pittier 4055. CHIRIQUÍ: Boqueté, Davidson 704; Puerto Armuelles, Woodson t$ Schery 834. PANAMA: Chepo, Pittier 4456; Juan Franco race track, Standley 27746; Las Sabanas, Standley 25847; Taboga Island, Standley 27861; R. Tapia, Standley 28213. (366) 1951] FLORA OF PANAMA (Leguminosae ) 67 23. Cassia LIGUSTRINA L. Sp. Pl. 378. 1753. Cassia occidentalis var. glabra Stahl in Est. Puerto Rico 3:112. 1885, not DC., fide N. P эле ligustrina (L. ) p & Rose, in Britt. & Wilson, Scientif. Surv. Porto Rico and Virgin Isls. 5:372. 19 Herb or subshrub to 2 m. high, the branchlets sparsely puberulent to glabrous. Leaves moderate, 8- to 18-foliolate; petiole up to 5 cm. long, somewhat flattened and sulcate above, bearing near the middle or base a slender elongate-cylindric gland 1-3 mm. long; rachis up to 15 cm. long, eglandular, similar to the petiole; stipules ovate or lanceolate, caducous; leaflets mostly about 7-8 pairs, narrowly ovate or lanceolate, 1.5-6 cm. long, up to 1.5 cm. wide, apically acuminate, basally rounded or obtuse and somewhat oblique, glabrous or often ciliolate, obscurely reticulate, dull, membranaceous or submembranaceous; petiolules mostly about 1 mm. long, puberulent above. Inflorescence terminal or axillary in upper leaves, several-flowered, paniculate (of several racemes); bracts ovate-lanceolate, about 5 mm. long, caducous; pedicels in age as long as the flowers. Flowers yellow; sepals ovate or oblong, about 7 mm. long and 5 mm. wide, subglabrous; petals oblong- obovate, 12-16 mm. long and up to 1 cm. wide, scarcely clawed, subglabrous, venose; stamens 4-morphic, 6—7 of them fertile, the anthers dehiscent by a pair of approximate, oblique, terminal pores; lowermost center stamen with a relatively long filament and small, short-rostrate, rudimentary? anther; 2 lowermost lateral stamens with stout filaments about as long as the anther, the anther subfalcate, about 6 mm. long, short-rostrate; 4 median anthers oblong, 4—5 mm. long, short- rostrate; uppermost 3 stamens rudimentary, represented by flattened, clawed, sub- spatulate staminodes; ovary pubescent. Legume linear, about 10 cm. long and 6—7 mm. wide, glabrous, flattened, margined, short-stipitate, septate, slightly curved, the seeds obliquely transverse. Panama and West Indies. BOCAS DEL TORO: Isla Colón, von Wedel 517. The species is very similar to C. occidentalis (including C. Sopbera) and C. leptocarpa. Тс differs primarily in the shape of the petiolar gland, a character which in final analysis may not be of specific value. 24. CassiA LAEVIGATA Willd. Sp. Pl. 441. 1809. Adipera laevigata 94 кей Britt, & Rose, ex Britt. & Wilson, Scientif. Surv. Porto Rico & Additional synonyms gine by Bentham don Linn. Soc. 27:527. 1871) include: Cassia aurata Roxb., Cassia corymbosa Ort. (not Lam.), Cassia elegans HBK., Cassia flori- bunda Cav. ч part, not Collad., Cassia DIEI Desf., Cassia Herbertiena Lindl., Cassia hybrida Ten., Cassia DI ын Zolling, Cassia septentrionalis Zucc. ex Colla A Cassia tropica Vell., Cassia vernicosa Closs, ex char., Chamaefistula floribunda (Сау.) G Don, Chamaefistula SE (Lindl) С. Don, Chamaefistula laevigata (Willd.) G. Don, Senna aurata Rox (367) [Vor. 38 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN Small tree to subherbaceous, usually shrub-like, the branchlets terete, glabrous, unarmed. Leaves moderately large, once-pinnate; petioles mostly 3—6 cm. long, glabrous, eglandular, sulcate above, callous basally; rachis somewhat longer than the petiole, glabrous, bearing dark, conical-cylindric glands just above insertion of each pair of leaflets (occasionally lacking at terminal pair) ; opposite leaflets 6 or 8, ovate-lanceolate, 3—10 (mostly 5-6) cm. long and 1—3 cm. wide, pronouncedly acute-acuminate apically, cuneate to rounded basally, entire, glabrous, dull, short- petiolulate; stipules linear, caducous. Inflorescence of axillary and terminal short corymbose racemes; peduncle shorter than the leaves, usually about 5 cm. long, glabrous, the bracts caducous; pedicels about 2 cm. long in basal flowers, glabrous. Flowers yellow, conspicuous; sepals 5, free, ovate to obovate, very unequal, up to 1 cm. long, glabrous; petals 5, free, mostly obovate, up to 15 mm. long, con- spicuously reticulate-venose; stamens 7 (plus 3 small staminodes), trimorphic; lower center stamen moderately filamented, the anther moderate; 2 lower lateral stamens long-filamented, large-anthered, the filament gross, about 12 mm. long, the anther about 8 mm. long, basifixed, dehiscing by confluent terminal pores; 4 central stamens short, the filaments 1-2 mm. long, the anthers about 5 mm. long, dehiscing by paired terminal pores; all anthers scarcely rostrate; staminodes orbicu- lar, 1-2 mm. long; ovary slender, glabrous, the stigma obliquely terminal. Legume linear, 6—9 cm. long and at maturity about 1 cm. wide, turgid-terete, pulpy, glabrous, more or less bordered marginally, the seeds many, transverse. Mexico to South America; West Indies. Introduced throughout World tropics. PANAMA: "Island in Gulf of Panama, Wellesley Hills", Purdie s. n. (1887). As far as is known, the sole record for the species in Panama consists of the cited specimen, from uncertain location. The species exhibits moderate variability, but is readily distinguished from other species of Cassia in Panama, particularly by the glabrous, few-foliolate leaves, without petiolar glands. In many places the species is a common weed of disturbed or cultivated ground. Standley and Steyer- mark report the seeds used as a substitute for coffee in Guatemala. 25. CASSIA LEIOPHYLLA Vogel, Syn. Cass. 25. 1837. Cassia — Mart. & Gal. in Bull. Acad. Brux. 10?:307. 1843, non Collad. 1816, fide М. Am. FI. Cassia pu ond i Mart. & Gal. loc. cit., non DC., fide Bentham Vogelocassia Trina (Vogel) Britt., in N. Am. Fl. 23: 259. 1930. Herbs or herbaceous shrubs or small trees up to a few m. tall, the branchlets glabrate to pubescent. Leaves small, normally 6-foliolate; petioles about 2 cm. long, ridged, usually somewhat pubescent; rachis up to 4 cm. long, glanduliferous between lower 2 pairs of leaflets; stipules linear, about 1 cm. long; leaflets obovate, up to 7 cm. long and 3.5 cm. wide, but usually 3-4 cm. long, rounded or obtuse and sometimes mucronulate apically, basally acute to irregularly rounded, dull, usually pubescent; petiolules 1-2 mm. long. Inflorescence few-flowered, usually axillary from the terminal leaves or upper branchlets, the short peduncles often (368) L9 FLORA OF PANAMA (Leguminosae) 69 with only 1—2 flowers on slender, pubescent pedicels 1—3 cm. long. Flowers yellow; sepals unequal, ovate to obovate, up to about 1 cm. long; petals obovate, rounded, about 2-2.5 cm. long, short-clawed, venose; fertile stamens 7 (the others rudi- mentary), 2-morphic, glabrous, dehiscent by terminal pores; the 3 lowermost with linear-oblong anthers 8-10 mm. long, markedly rostrate; the 4 median stamens with short-beaked anthers about 6 mm. long; ovary linear, pubescent. Legume linear-falcate, up to 15 cm. long and 4 mm. wide, flattened, somewhat pubescent, the margins conspicuously ridged; the seeds oblique, prominent. Mexico; Central America; South America to Peru and Brazil. BOCAS DEL TORO: се 22. Carleton III. CANAL ZONE: Las Cruces Trail, Hunter & Allen 714. DARIEN: between Pinogana and Yavisa, Allen 275. PANAMA: between Capira and $us Dodge 4. оне [o Juan Díaz, Standley 30945; Matías Hernández, Standley 28765; Panama, Standley 26831. vERAGUAS: Sona, Allen 1058. 26. "CASSIA TORA 12 Sp. Р 376. 1755. Cassia obtusifolia L. loc. cit. 377. Cassia pentagonia Mill. Gard. on d '$, no. 18. 1768. 2. Lis а Fl. Aegypt.-arab. 86. 1775, fide Benth. ncycl. Meth. 1:643. 1783, fide Benth., поп L. Cassia Tala Desv. in t BOR, Bot. 3:73. Cassia toroides Roxb. Hort. Beng. 31. M fide Benth. Cassia bumilis Collad. Hist. Cass. 96. 181 Cassia gallinaria Collad. loc. cit. 1816, fide P th. Chamaefistula contorta G. Don, Gen. Hist. Dichl. Pl. 2:452. 1832, fide Benth. Senna toroides Roxb. Fl. Ind. 2:341. 1832, fide Benth. Senna Tor к Roxb. loc. cit. 340. Cassia co a (Don) Vogel, Syn. Cass. 20. 1837, fide Benth. Diallobus jin Raf. Sylva Puer 128. 1838. Diallobus uniflorus Raf. loc. cit. Cassia candenatensis pr in Ste id Nos Bot. ed. 2, 304. 1841, ex ic. cit., fide Benth. NE i i "Pe tt, & Rose in Britt & Wilson, oam Surv. Porto Rico & Virgin Isls. Annual herb or small shrub up to about 1 m. tall, the branches glabrous or nearly so. Leaves moderately small, normally 6-foliolate; petiole angled, sub- glabrous, flattened or sulcate above, 1—3 cm. long; rachis similar and about as long, bearing between the lowermost pair of leaflets a linear-subulate gland about 2 mm. long; stipules linear-subulate, about 1 cm. long, caducous in age; leaflets obovate, up to 4 cm. long and 2 cm. wide, apically obtuse to rounded or subtruncate and usually mucronulate, basally obtuse to subcuneate and unequal, glabrous above, glaucous and somewhat appressed-pubescent below, membranaceous, obscurely reticulate; petiolule about 1 mm. long. Inflorescence of 1—2 flowers from (each) upper nodes. Flowers yellow, moderate; sepals ovate to oblong, up to 8 mm. long and 4 mm. wide, unequal, scatteringly pubescent with few long hairs; petals ovate to obovate, up to 12 mm. long, venose, glabrous; fertile stamens 7, all dehiscent by a single terminal pore; anthers of the 3 lowermost rostrate, about 4 mm. long, with short filaments; anthers of the 4 median ones oblong, truncate, 2-3 mm. long, (369) [Vor. 38 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN with filaments only slightly shorter than in the lowermost stamens; ovary linear, pubescent. Legume linear, usually about 15 cm. long and 4 mm. wide, arcuate, subglabrous, dehiscent, margined-subterete; seeds obliquely longitudinal, shiny. Cosmopolitan in world tropics and warmer temperate sections; native to India. CAS DEL TORO: Isla Colón, von Wedel 51. CANAL ZONE: "Chagres", Fendler 92; Cule ven Pittier 2116, mte 26041; Summit, Standley 27310. PANAMA: Bella Vista, Standley 2315; between Panamá and Chepo, D Dodg e, Hunter, Steyermark 8 Ай» 16627; anamá, Bro. Paul 580; Rio Tapia, Standley 20076. VERAGUAS: Sona, Allen 1060. 27. CASSIA PILIFERA Vogel, Syn. Cass. 23. 1837. ?Cassia cubensis Hoffmansegg, Pfl. Verz. 1:209. 1824 Cassia maritima Willd. in Vogel, loc. cit. 1837, fide Benth. Emelista pilifera (Vog.) Pittier, in Jour. Wash. Acad. 19:176. 1929. Shrub or large herb, the branchlets pilose or setose. Leaves moderate, normally 4-foliolate; petioles up to 4 cm. long, loosely pilose; rachis much shorter than the petiole, up to 1 cm. long, normally bearing between both pairs of leaflets a linear- subulate gland 3—4 mm. long; stipules linear-filiform, usually about 1 cm. long, pilose like the rachis and branches; leaflets elliptic to obovate, up to 6 cm. long and 3.5 cm. wide, apically narrowly rounded to obtuse or subacute, basally unequal and oblique, pubescent above and below, dull above and somewhat glaucous below, obscurely reticulate, membranaceous; petiolules 1-2 mm. long, pubescent like the petiole. Inflorescence of few axillary, subterminal flowers, often umbellate (up to 4-flowered) from a common peduncle in the lower axils. Flowers yellow, large; sepals relatively small, ovate to lanceolate, up to 9 mm. long and 4 mm. wide, very unequal, sparingly long-pubescent; petals ovate or suborbicular, up to 2.3 cm. long and almost as wide, clawed, venose, lightly pubescent to glabrous in age; fertile stamens 7, dehiscent by a pair of approximate terminal pores, the 3 lowermost un- equal, linear-oblong, rostrate, arcuate, the anther of the largest about 1 cm. long with an elongate filament; 4 median stamens unequal, short-rostrate (the rostrum strongly reflexed) , the largest anther about 5—6 mm. long; ovary linear, pubescent. Legume up to 16 cm. long while only 3—4 mm. wide, subquadrate, margined, de- hiscent, arcuate, tomentulose to pilose; seeds longitudinal, rectangular, about 3 mm. long. Mexico to Argentina; Cuba. ANAL ZONE: Balboa, Standley 26057, 27118, 20303; Las Cruces Trail, ек 20010. PANAMÁ: Alhajuela, Pitlier 2367; between Capira and Potrero, Dodge & Hunter 8610; Chepo, Mell s.n.; Corozal road, Standley 26880; Matias Hernandez, Standley eo, 31017. 28. Cassia WiLLiAMsm (Britt. & Rose) Standl. Field Mus. Bot. Ser. [Fl. Costa Rica] 18:519. 1937. Peiranisia Williamsii Britt. & Rose, in N. Am. Fl. 23:265. 1930. Shrub or small tree to 3 m., the branchlets hirsute or pilose (the long hairs occasionally deciduous or lacking). Leaves elongate, multifoliolate; petioles short, glabrous below, puberulent and somewhat flattened above; rachis up to 20 or more (370) 1951] FLORA OF PANAMA (Leguminosae ) үл Д Pp S ANNIS W —— n8 257 > 2а аа 3 SSS = ~ TN SSS ХУМ $ 4 MMS OLY AT OSes % S ed TRAE Mn OA | = E ^ f 254 " < В N } DNY 5 PS ) қ Х Lr = 2 W NN ү € » \ К N У _ TAa S ANS 1 s \ B p or uit ES SS “ Ж” pr £ 7224 ча 22% 2 225 sss Fig. 126. Cassia Williamsii cm. long, subglabrous, bearing between the lowermost (and usually second) pair of leaflets a prominent, stalked, clavate gland 4—5 mm. long, and between all or many of the remaining pairs of leaflets thin acicular glands; stipules linear-falcate, about 8 mm. long, caducous; leaflets 30-60 pairs, small, narrowly oblong, up to 15 mm. long and 3 mm. wide, apically rounded, basally inequilateral, few-veined, glabrous; petiolules less than 1 mm. long. Inflorescence axillary from the terminal or subterminal axils, few-flowered; peduncle up to 4 cm. long, subglabrous; pedi- cels (usually 3—4) up to 3 cm. long, condensed or subumbellate from the upper peduncle; bracts lanceolate, about 3 mm. long. Flowers large, yellow; sepals very unequal, ovate to obovate, the largest about 15 mm. long, glabrous, membranaceous, somewhat maculate especially basally; petals oblong-obovate to orbicular, unequal, (371) [Vor. 38 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 3 cm. long and (the largest) almost 2.5 cm. wide, glabrous, short-clawed; fertile stamens 7, with short filaments; anthers of the 3 lowermost linear-oblong, falcate, about 7 mm. long, long-rostrate and dehiscent by a pair of approximate terminal pores; anther of 4 median stamens about 6 mm. long, subtruncate apically by oblique reflexion of a short rostrum, dehiscent by a pair of terminal pores; uppermost 3 staminodes flat, 2-3 mm. long, clawed; ovary linear, arcuate, pubes- cent. Legume linear, about 14 cm. long and 4-5 mm. wide, glabrate, stipitate, septate between the seeds; seeds transverse, rhombic, 2-3 mm. long. Panama and Costa Rica. CHIRIQUÍ: Boquete, Davidson 845; Hato del Jobo, Pittier 5406, 5407. сосі.Е: Bismark, Williams 567; El Valle, Allen 115, 1180, Hunter 9 Allen 301. Cassia Williamsii is closely related to C. Mutisiana of Colombia, which species, however, has the leaflets pubescent below. 29. CASSIA MULTIJUGA Rich. in Act. Soc. d’Hist. Nat. Paris 1:108. 1792. Cassia calliantha G.F.W. Mey. Prim. Fl. Esseq. 169. 1818, fide Benth. Cassia Richardiana Kunth, Mim. 139, ¢. 42. 1819, fide Benth. КА, gà Ep TN қ ES NI (9 & 51 ON \ Е POR 7) А Fig. 127. Cassia multijuga (372) 1951] FLORA OF PANAMA (Leguminosae ) 73 Cassia semifalcata Vell. Fl. Flum. Ic. 4:4. 68. 1827, fide Benth. Cassia Selloi G. Don, Gen. Hist. Dichl. Pl. 2:442. 1832, fide Benth. Cassia ampliflora Steud. in Flora 26:760. 1843, fide Benth. dditional synonyms are given by Bentham Small tree, the branchlets puberulent or subglabrous. Leaves moderately large, multijugate; petiole relatively short, 1-2 cm. long, subsulcate above except base- ward where more or less terete and callous; rachis 10—25 cm. long, sulcate above and cross-partitioned at insertion of the petiolules, usually puberulent marginally, bearing in most cases a prominent, elongate, cylindric-subconic gland between the lowermost pair of leaflets; stipules linear-acuminate, caducous; leaflets (on at least most leaves) 12—40 pairs, linear-oblong or oblong, 1—4 cm. long and up to 1 cm. wide, rounded and mucronulate apically, obtuse or rounded and slightly inequi- lateral basally, dull but conspicuously lighter below, glabrous or puberulent; petiolule about 2 mm. long. Inflorescence terminal or subterminal, paniculate of several racemes; bracts like the stipules, caducous; pedicels in age as long as 2.5 cm. Flowers yellow, showy; sepals ovate, up to 7 mm. long and 5 mm. wide, sub- glabrous; petals ovate to obovate or suborbicular, very unequal but the largest about 2 cm. long, clawed, glabrous or subglabrous; fertile stamens 7, of 2 sorts; lowermost up to 13 mm. long, the anther linear-oblong, 7-10 mm. long, falcate, long-rostrate, dehiscent by paired terminal pores; 4 median anthers oblong, 5—6 mm. long, short-reflexed-rostrate, subsessile; 3 uppermost stamens rudimentary or lacking; ovary glabrous or nearly so. Legume broadly linear, up to 20 cm. long and about 1.5 cm. wide, flat, septate, shiny, short-stipitate, the seeds transverse. Brazil, Peru, the Guianas; Colombia, Mexico and West Indies? fide Bentham, but not listed there in recent floras. CANAL ZONE: Balboa, Mell 12. This is apparently a new record for Panama. The specimen cited was likely from an introduced plant, having been collected in a populated area (Balboa). It had been determined incorrectly as C. moschata, and the label used had been de- signed originally for "Plants of Mexico." The species is rather widely planted as an ornamental. 30. CassrA BIFLORA 1. Sp. Pl. 378. 1753. Cassia tenuissima L. loc. cit. 1753, fide М. Am. FI. Cassia galegifolia L. Syst. Nat. ed. 10, nee "p fide Benth. Cassia Marimari Aubl. Pl. Guian. Franc. 382. 1775, ex ic. Plumieri cit., fide Benth. Cassia crista Jacq. Icon. Pl. Rar. 1:8, t. 74. 1781—86, fide Benth Cassia frondosa Ait. Hort. Kew. ed. 1, 2:53. 1789, fide Benth. Cassia pallida Vahl, Eclog. 3:12. 1807, fide N. Am. FI. Cassia semperflorens DC. Cat. Hort. Monsp. 90. 1813, fide N. Am. Fl. Cassia geminiflora Moc. & Sessé; эв Hist. Cass. 103, 7. 3. 1816, fide N. Am. FI. Cassia nemorosa HBK. Now. Ges & Sp. 6:353. 1824, fide N. Am. FI. Cassia acapulcensis HBK. loc. ae 1824, fide N. Am. Fi. C HBK. loc. c А Е Cassia biflora var. semperflorens DC. Prodr. 2:496. 1825, fide N. Am. ЕІ. Cassia Berteriana Balb. in DC. loc. cit. 1825, fide Benth. (373) [Vor. 38 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cassia oxyadena DC. loc. cit. 495. 1825, excl. syn. NE & Mill., fide Benth. Cassia fulgens Macfad. Fl. Jam. 1:342. 1837, fide Benth. ?Cassia xipbioidea Bertcl. Fl. Guatemal. 15. 1840, E WE * Ard Benth. Cassia biflora angustisiligua Griseb. Fl. Brit. W. Ind. Cassia pallidior Rose, in Jour. Wash. Acad. 17:167. Туы Peiranisia Бі оға (L.) Pittier, in Trab. Mus. Com. Venez. 158. 1928. Small shrub to about 2 m., the branchlets lightly pubescent to glabrous. Leaves moderate, up to about 20-foliolate; petiole relatively short, usually lightly pubescent, flattened above, somewhat margined; rachis elongate, usually about 10 cm. long, similar to the petiole, nodose at insertion of the leaflets, bearing a promi- nent obovoid or clavate gland 2—3 mm. long between lowermost pair of leaflets, and a small, linear, terminal process; stipules linear-subulate, almost 1 cm. long; leaflets 4-10 pairs, mostly elliptic or oblong, up to 3.5 cm. long and 14 mm. wide, apically rounded and mucronulate, basally subcuneate and somewhat unequal, sub- glabrous, glaucous below, dull above; petiolules lightly pubescent, 1-2 mm. long. Inflorescence mostly axillary, the slender peduncle commonly divided into 2 fili- form pedicels about 2 cm. long, frequently bearing 1—2 glands between the pedi- cels; bracts small, caducous. Flowers large, showy, yellow; sepals ovate to orbicular, up to 9 mm. long and broad, unequal, glabrous; petals ovate or obovate, 2-3 cm. long, short-clawed, glabrous, inconspicuously veined; fertile stamens 7, 2-morphic; anthers of the 3 lowermost oblong-falcate, 8—10 mm. long, very long- rostrate, the cylindric rostrum about 3 mm. long and dehiscent terminally by a pair of subconfluent pores; 4 median anthers oblong-subfalcate, 4—5 mm. long, apically obliquely truncate by strong reflexion and adnation of a short 2-pored rostrum; other stamens rudimentary or lacking; ovary linear, tomentose. Legume linear, 5-15 cm. long and 5 mm. wide, lightly margined, flat and very thin, im- pressed between the seeds, lightly pubescent to glabrous. Mexico; Central America; West Indies; northern South America. É: Aguadulce, Allen 4031. HERRERA: Chitré, Allen 1007. PANAMA: Alhajuela, Pittier. 3244. 3465. VERAGUAS: Santa Fe, Allen 4426. The species is quite variable and undoubtedly subspecific entities could be es- tablished. Such variation is especially noticeable in vegetative characters. The specimens cited, for example, differ markedly in leaflet pubescence, the Allen 1097 collection having quite pubescent leaflets tending towards the condition found in C. velutina Britt. & Killip, while the Pittier specimens have essentially glabrous leaflets (= C. pallidior Rose). 31. Cassia BICAPSULARIS L. Sp. Pl. 376. 1753. Cassia sennoides Jacq. Collect. 1:74. 1786. Cassia coluteoides Collad. Hist. Cass. 102, f. 12. 1816. Senna bicapsularis Roxb. Fl. Ind. 2:342. 1832. Cassia Augusti Harms, in Fedde Rep. Sp. Nov. 18:93. 1922, fide Macbride. Adipera bicapsularis (L.) Britt. & Rose, in Britt. & Wilson, Scientif. Surv. Porto Rico and Virgin Isls. 5:370. 1924 (374) FLORA OF PANAMA (Leguminosae) 75 n addition the following synonyms are apparently correctly given P aoe in his fee work, Revision of the genus Cassia, in Trans. Linn. Soc. 27:503. j 1 . 1783: Cassia Berterii Colla, Hort. Ripul. 30, £. 24. 1824. Cassia inflata Spreng. Syst. Veg. 2:336. 1825. Cassia dormiens Vell. Fl. Flum. Ic. 4:7. 6 Chamaefistula pendula G. Don, loc. cit. 1832. Cassia chrysoloma DeNot. Ind. Sem. Fort. Bot. R. Arch. Gen. 1840 (ex Linnaea 15:Litt. Ber. 92. 1841, ex char. Cassia UE Benth. in ET 22.5275 1849: Cassia glandulifera Reinw. in Blume, Cat. Hort. Bog. 68, ex Miq. Fl. Ind. Bat. 11:92. 1855. Cassia uiis um Hort. Bi (286. 1844), in Ann. Sci. Nat. Bot. II, 14:58. 1840, ex Miq. loc. cit. = Shrub or small tree, "m glabrous throughout. Leaves moderately small, 6- to 10-foliolate; petiole up to 3 cm. long; rachis up to 4 cm. long, somewhat flat- tened or sulcate above, bearing between the lowermost pair of leaflets a stocky clavate gland 1-2 mm. long; stipules caducous; leaflets obovate, up to 3 cm. long and 1.5 cm. wide, the terminal pair largest, apically rounded to subtruncate, basally cuneate (upper pairs) to obtuse (lower pairs) and somewhat unequal, subsessile, glaucous below, dull above, obscurely reticulate. Inflorescence terminal to axillary from the upper leaves, the individual racemes several-flowered; bracts small, Janceo- late, about 1 mm. long when caducous; pedicels about 1 cm. long at anthesis. Flowers yellow; sepals obovate to ovate or lanceolate, up to 12 mm. long, imbri- cate, unequal, greenish; petals oblong or obovate, up to 16 mm. long and 9 mm. wide, imbricate, glabrous, venose, sessile; fertile stamens 7, trimorphic; staminodes 3, flat, cuneate-deltoid, 4-5 mm. long, clawed, apically truncate; the lowermost- center anther linear, about 8 mm. long, apically rostrate and dehiscent by a single terminal pore, its filament about 4 mm. long; the two marginal anthers (of lower- most stamens) similar to the center one but more robust, with stout filaments up to 1 cm. long; 4 median anthers linear-oblong, about 5 mm. long, short-rostrate and dehiscent by 2 terminal pores, with short filaments; ovary linear, glabrous or pubescent, sessile. Legume linear-cylindric, about 12-15 cm. long and usually 1-1.3 cm. wide when mature, glabrous, dehiscent into 2 septate halves, stipitate, straight or only slightly curved. Mexico to Paraguay and introduced into Old World tropics. CANAL ZONE?: without locality, Hayes 746, Seemann 529. состЁ: Aguadulce, Pittier 31a. CASSIA BICAPSULARIS L. var. PUBESCENS Benth. in Trans. Linn. Soc. 27:525. 1871 Cassia indecora HBK. Nov. Gen. & Sp. 6:344. Chamaefistula indecora С. Don, Gen. Hise Dichl. Ұл 2:452. 1832. Cassía yen Willd. Herb. ex Vogel, Syn. Cass. 18. 1837. (375) [Vor. 38 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cassia ovalifolia Mart. © Gal. in Bull. Acad. Brux. 10:305. 1843. Cassia manzanilloana Rose in Contr. U. S. Nat. Herb. 1:325. 1895. Cassia bicapsularis var. indecora Urban, Symb. Ant. 2: 268. 1900. Adipera indecora (HBK.) Britt. & Rose, in N. Am. Fl. 23:239. 1930. The variety differs from the species in being more pubescent (upper twigs and lower leaflet surface usually puberulent) and in having the lowermost lateral sta- mens long-rostrate (beak 1-2 mm. long vs. 1 mm. or less in the species). While at their extremes the species and the variety are quite distinct in these character- istics, intermediacy on both the staminal and pubescence characters is not in- frequent in northern Central America and Mexico. Furthermore, correlation between these two characters is imperfect. Thus it would seem wise to follow Bentham and Urban in considering the pubescent forms as of varietal rank (C. bicapsularis L. var. pubescens Benth.), and not of specific rank as do Britton and Rose (C. indecora HBK.). Mexico to northern South America; West Indies. CANAL ZONE: Summit, Standley 30000. сосі.Е: El Valle, Hunter & Allen 337. 32. Cassta unica Schery, nom. nov. Cassia falcinella Standley, in Contr. U. S. Nat. Herb. 18:102. 1916, non Oliver. Chamaefistula falcinella (Sandl) "Brite. & Kors, іп М. Am. Fl. 23:238. 19 Shrub with terete or obscurely angled, cinereous-puberulent stems. Leaves 4- foliolate, small to moderate; petiole about 2.5 cm. long; rachis about 1 cm. long, usually subulate-tipped, bearing between each pair of leaflets а slender-cylindric, acute, black gland; stipules linear-falcate, about 1 cm. long and 1 mm. wide, cinereous-puberulent or subglabrous; leaflets oblong-obovate to elliptic-oblong, up to 9 cm. long and 4 cm. wide, abruptly long-acuminate apically, the tip up to 14 mm. long, acute, basally obtuse or rounded and unequal, chartaceous, glabrous, lustrous above. Inflorescence dense, many-flowered, paniculate, densely cinereous- puberulent; bracts small, about 2 mm. long, linear-lanceolate and not falcate. Flowers yellow; sepals oblong-ovate, up to 5 mm. long, obtuse, subequal, pubescent with short appressed, curved, yellow hairs; petals oblong, about 13 mm. long, obtuse, clawed, puberulent without; fertile stamens 7, anthers of the 3 lowermost slightly smaller, 5-7 mm. long, somewhat rostrate, dehiscent by a pair of sub- confluent terminal pores; 4 median anthers oblong, about 8 mm. long, obliquely very short-rostrate and subtruncate, dehiscent by a pair of terminal pores; upper- most 3 stamens minute and rudimentary; ovary linear, arcuate, tomentose; stigma and upper style as wide or wider than the ovary, oblique. Legume not known. Western Panama. CHIRIQUI: vicinity of San Felix, Pittier 5147. Unfortunately it has been necessary to provide a new name, C. unica, to C. falcinella Standley, in that a previous African species had been described as C. falcinella by Oliver (Fl. Trop. Afr. 2:281. 1871). Perhaps our species should not be distinct from Cassia undulata, although it is here listed separately because (376) 1951] FLORA OF PANAMA (Leguminosae) 77 C. undulata constantly has larger bracts. It would appear as though a minor variation were superimposed upon the "undulata complex", a variation that likely has not been able to perpetuate itself, in that no second collection of C. unica has appeared (type collected 1911). 33. Cassi UNDULATA Benth. in Hook. Jour. Bot. 2:76. 1840. Chamaefistula undulata (Benth.) Pittier, in Trab. Mus. Com. Venez. 5:151. 1928. Small shrub, often vine-like, the older branchlets glabrous. Leaves small to moderate, 4-foliolate; petioles elongate, up to 5 cm. long, glabrous or very lightly pubescent; rachis somewhat angled, up to 3 cm. long, bearing a subconic gland between each pair of leaflets; stipules linear-falcate to lanceolate-falcate, about 7 mm. long, subpersistent; leaflets lanceolate-subfalcate, up to 11 cm. long and 3.5 cm. wide, inequilateral, apically usually attenuate-acuminate, basally obtuse or rounded, the margins somewhat undulate, glabrous, shiny above, not conspicuously reticulate; petiolules short, callous. Inflorescence racemose-paniculate, terminal or subterminal, several-flowered, puberulent; pedicels up to 2 cm. long; bracts ovate- lanceolate, about 8 mm. long, mucronate, persistent. Flowers yellow; sepals elliptic to obovate, 7-8 mm. long, puberulent; petals elliptic to obovate, up to 15 mm. long, puberulent, venose; stamens 7, of 2 sizes, essentially similar; anthers oblong, somewhat curved, glabrous, basally rounded-subtruncate, apically truncate but with oblique rostrate or subrostrate pores to one side, larger (4) anthers about 8 mm. long, smaller (3) anthers 4-5 mm. long; ovary appressed-tomentose, style short. Legume 10—20 cm. long, about 1 cm. thick, turgid, glabrous, basally obtuse. Southern Mexico; Central America; Trinidad; northern South America. CANAL ZONE: Ancén Hill, Pittier 1724, Williams n Bellavista, Macbride 2734, Piper 5132; "Chagres", Fendler 86; 21. ded 2006; pire to Mandinga, Piper 51 138, 5141; Frijoles, Maxon 4704, Pittier 2688; Las Cruces pics Standley 20231. PANAMA: Bismark, Williams 506; Juan Diaz, 5, 30572; Panama City, Macbride 2734; Tumba Muerto road, Standley 20704. vERAGUAS: west of Sona, Allen 1043. 34. Cassia Maxonii (Britt. & Rose) Schery, comb. nov. Chamaefistula Maxonii Britt. & Rose, in N. Am. Fl. 23: D 1930. 3 19 Қ МК Из Williamsi Bart & Rose, loe: cit. 236. 1930. Small to moderate trees, the branchlets puberulent to glabrous. Leaves nor- mally 4-foliolate, moderately large; petiole 2-4 cm. long, stout, puberulent to glabrous, somewhat sulcate above; rachis similar, usually about 3 cm. long, with a prominent subconic gland between the lower pair of leaflets and generally a smaller gland or protuberance apically; stipules linear-lanceolate, curved, about 1 cm. long; leaflets ovate to elliptic, up to 15 cm. long and 7 cm. wide, acute to acuminate apically, obtuse or rounded and slightly inequilateral basally, glabrous above, puberulent-tomentulose and with raised prominent veins below, the veins brown against a dull-green background. Inflorescence terminal or subterminal, of (377) [Vor. 38 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1 to few usually flexuous racemes or panicles, puberulent; bracts linear-lanceolate, caducous; pedicels generally 2-3 cm. long. Flowers yellow, showy; sepals oblong to narrowly ovate, less than 8 mm. long, rounded apically, puberulent; petals larger, obovate, up to about 2 cm. long and 1.5 cm. broad, lightly puberulent; fertile stamens 4 (the median ones), all similar, the others more or less abortive or reduced or caducous; anthers oblong, about 8 mm. long, somewhat curved, apically short-rostrate, basally rounded-subtruncate, dehiscent by terminal pores; ovary linear, densely pubescent. Legume linear, angled-subterete, up to 25 cm. long and about 2 cm. wide, straight, smooth; seeds transverse. Mexico; Central America; northern South America. ANAL ZONE: Ancón, Bro. Heriberto 38, Killip 12055, Williams 9; Balboa, Standley Poe 26447, 28492; Bellavista, Macbride 2750, Piper 5357, p ndley 25346; near Culebra, Pittier 2312, Standley 25057; Empire to Mandinga, Piper 5142. "aen between Las Margaritas an s El Valle, Woodson, Allen & Seibert dg NE Williams 135. PANAMÁ: Matías Hernández, Standley 28060; between Panamá and Chepo, Dodge, Hunter, Steyermark " Allen 16647; near Panamá, Standley 26844, 27766; Río Tapia, Standley 28183; Río Tecümen, Standley 26568; Taboga win: , Standley bie 27086. SAN BLAS: Puerto Obaldia, Pittier 4405. UNKNOWN: “Pan , Seemann 22 Central American specimens with only 4 "e functional stamens are found distributed now and then as C. bacillaris (= С. fruticosa) and very frequently as C. oxyphylla. Drawing from specimens distributed under these two species (along with many specimens undetermined to species), markedly inequilateral leaves, usually larger buds and perianth parts, and appressed lower leaflet pubescence seem linked with a 7-stamen characteristic. Such characters would determine C. fruticosa, while C. Maxonii (often found in herbaria as C. oxyphylla) would then be distinguished by the reduced functional stamen number, the less inequi- lateral leaves, generally smaller petals and sepals, narrower buds, more tomentulose raised pubescence of lower surface of the leaflet, etc. C. Maxonii seems to differ from C. oxyphylla of South America only in pos- sessing 4 rather than 7 functional stamens. The writer was tempted to list C. Maxonii as a variety of C. oxyphylla, under the descriptive varietal name of quadristaminea, but refrained from so doing because: (1) the staminal difference would appear without intermediacy, and (2) the "tail would then wag the dog", the 4-stamen entity having been accumulated in the herbaria in a near absolute majority for Central America for almost a century (as C. oxyphylla following Bentham, or undetermined to species). Unfortunately, on priority grounds the name C. Maxonii must be given this species, although C. Maxonii was described (as Chamaefistula Maxonii) inade- quately and apparently without comprehension of the staminal difference separating it from C. oxypbylla. Yt thus may be more useful in analyzing the entity here resolved as C. Maxonii to refer to my description rather than to the original, and to regard most of the specimens cited as more "typical" of the entity than is the type (Maxon & Harvey 6611). At an opposite extremity of “untypicalness” from “Chamaefistula Maxonii”, but connected by intergradation within the entity, is (378) 1951] FLORA OF PANAMA (Leguminosae) 79 “Chamaefistula anconis Britt. & Rose". The latter was not selected as name- bringing synonym since its type is in fruit only. For the same reason, although more "typical" in foliage characters, “Chamaefistula Williamsii Britt." was not chosen. Another synonym, “Chamaefistula Hayesiana Britt. & Rose", was rejected because of poor condition of the type and "untypical" leaflet pubescence approach- ing that of C. fruticosa var. dariensis. Possibly monographic study will some day indicate an earlier name for this species, but inasmuch as most older descriptions, the types for which are not available to us, fail to stress the staminal character- istics, I am unable to list with certainty any name taking precedence over С. Maxonii. 35. CASSIA OXYPHYLLA Kunth. var. dariensis (Britt. & Rose) Schery, comb. nov. Chamaefistula dariensis Britt. & Rose, in М. Am. Fl. 23:238. 1930. Small or large trees, the branchlets usually puberulent. Leaves moderately large, 4-foliolate; petioles about 1—2 cm. long, puberulent, subterete; rachis similar, up to 4 cm. long, with a prominent subcylindric gland about 2 mm. long between lower pair of leaflets; stipules linear, caducous; leaflets ovate to elliptic, up to 15 cm. long and 7 cm. broad, acuminate apically, basally inequilaterally acute to obtuse, membranaceous, prominently reticulate, essentially glabrate above, pubes- cent and dull below; petiolules terete, about 3 mm. long, puberulent. Inflorescence a generally flexuous, terminal or subterminal, few- to several-flowered raceme or panicle; pedicels 1-2.5 cm. long, densely puberulent; bracts linear, small, caducous. Flower yellow; sepals relatively small, oblong to narrowly ovate, 5-8 mm. long, puberulent, reflexed in fruit; petals relatively large, oblong to ovate, up to 2.3 cm. long and 1 cm. wide, markedly venose with 3 prominent longitudinal veins, puberulent, very short-clawed; fertile stamens 7; anthers of the 3 lowermost smaller, oblong-falcate, about 6 mm. long, apically long-rostrate, dehiscent by 2 terminal pores, glabrous; anthers of the 4 median ones about 8 mm. long, ob- liquely rostrate apically but otherwise subtruncate and similar to the lowermost; ovary linear, tomentose, arcuate, the style thick and short. Legume (immature) tetragonal, rufous-puberulent, about 15? cm. long and 6? mm. wide, arcuate, not constricted, the seeds transverse. Panama. RIEN: Boca de Сире, Allen 010; Риман Pittier 6579; Rio Yape, Allen 327. PANAMA: Rio Tataré, Woodson & Schery The Britton and Rose species, here reduced to a variety of C. oxyphylla, may not be, in final analysis, distinct from C. oxyphylla. The type of C. oxyphylla is not available for examination, but the original description with plate (Kunth, Mim. 129, /. 39. 1819) is excellent. Comparison of C. oxyphylla var. dariensis with it shows the variety to differ principally in having longer, narrower petals and broader leaflets than the species—characters which may not be beyond the range of variability of the species. (379) [Vor. 38 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN 36. Cassia FRUTICOSA Mill. Gard. Dict. ed. 8, по. 10. 1768; Vogel, in Linnaea 15:67. 1841. ?Mimosa nodosa L. Sp. Pl. 516. 1753, fide Benth. Cassia bacillaris L. f. Suppl. 231. 1781, fide Benth. Catbartocarpus Bacillus Pers. Syn. Pl. 1:459. 1805. ?Inga nodosa Willd. Sp. Pl. 4:1016. 1806, fide Benth. Bactyrilobium bacillare Hornem. Hort. Bot. Hafn. 1:392. E Cassia puberula HBK. Nov. Gen. & Sp. 6:341. 1824, fide Ben Chamaefistula bacillaris (L. f£.) С. Don, Gen. Hist. ag Pl. m 451. 1832. Chamaefistula puberula (HBK.) С. Don, loc. cit. Cassia cartbaginensis Willd. Herb. ex Steud. Nom. E e 2, 304. 1841. Cassia Fockeana Miq. in Linnaea 18:579. 1844, fide Benth. Chamaefistula fruticosa (Mill.) Pittier in Trab. Mus. Com. Venez. 3:152. 1928. Chamaefistula Valerioi Britt. & Rose, in N. Am. Fl. 23:236. 1930, fide Standl. Shrub or small tree up to 10 m. tall, the branchlets usually glabrous. Leaves large, 4-foliolate; petiole longer than the rachis, 2—6 cm. long, terete to somewhat angled or flattened above; rachis 1-4 cm. long, usually glabrous, bearing between the lower leaflets an oblong-conic gland; stipules linear, early caducous; leaflets large, inequilateral (the apical ones pronouncedly so), ovate to elliptic-lanceolate, up to 18 cm. long and 9 cm. wide, usually glabrous above and lightly short- puberulent below, acute and somewhat acuminate apically, obliquely rounded to obtuse basally, the veins prominent above and below, more or less concolorous with the background; petiolules stout, 3—4 mm. long. Inflorescence terminal or sub- terminal, paniculate or of several racemes from the upper axils, puberulent; bracts caducous; pedicels up to 4 cm. long, puberulent. Flowers yellow; sepals large, more or less oblong, up to 13 mm. long and 7 mm. wide, appressed-pubescent, persistent; petals oblong to obovate-orbicular, up to 3 cm. long and 2 cm. wide, short-clawed, puberulent, lightly venose; functional stamens normally 7, bi- morphic; the 3 lowermost anthers conspicuously rostrate, falcate, 8—9 mm. long, the beak about 2 mm. long and dehiscent by a single upward-directed terminal pore; the 4 median anthers less curved, oblong, about 9 mm. long, subtruncate apically and basally, the short beak oblique and dehiscent by 2 terminal pores; other stamens rudimentary; ovary linear, sessile, tomentose. Legume elongate, linear, up to 30 cm. long and 1 cm. wide, terete or subterete, straight, glabrous, minutely verrucose; seeds transverse. Tropical America from Mexico to Brazil. CAS DEL ToRO: Changuinola Valley, Cooper & Slater 100, 130; Darkland, von Wedel 2626; Isla Colén, von Wedel 93; Shepherd Island, von Wedel 2730; Water Valley, von Wedel 613. CANAL ZONE: Barro Colorado Island, Shattuck 54; Bellavista, Piper 5131; “Chagres”, Fendler 87; Fort Randolph, Standley 28671, 28607; Gatun, Hayes 360, 360. CHIRIQUÍ: San pae Pittier 5146. сост: Bismarck, Williams 560. PANAMA: Alhajuela, Pittier 2319; between p and Potrero, Dodge & Hunter 8600; Juan Diaz, Standley 30546; Pacora, zb Y £10; О. Tranquilla, Hunter 9 Steyermark 17213; Rio a Hunter & Allen 236, Standley 26757, 20450. UNKNOWN: "western Panama”, Stork 2 (380) 1951] FLORA OF PANAMA (Leguminosae) 81 Fig. 128. Cassia fruticosa This species is more commonly located in the herbaria and literature under the name C. bacillaris, a name which it would be well to conserve were there any provision to do so in the international rules. 36a. Cassia FRUTICOSA Mill. var. $atunensis (Britt.) Schery, comb. nov. Chamaefistula gatunensis Britt. in N. Am. Fl. 23:234. 1930. Small viney tree, the branchlets puberulent. Leaves moderate to large, 4- foliolate; petioles about 3 cm. long, the rachis similar and only slightly longer; gland between lowermost pair of leaflets cylindric-subconic, about 3 mm. long; (381) [Vor. 38 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN leaflets elliptic, only slightly inequilateral and differing from the species in this respect, puberulent below, more so and with slightly longer hairs than in the species, glabrous above and otherwise much as in the species. Inflorescence several- to many-flowered, puberulent; flowers moderately large, yellow; sepals oblong to elliptic-ovate, somewhat unequal, up to 1 cm. long; petals elliptic to obovate, up to 2.5 cm. long, tomentulose; functional stamens 7, the three lowermost smaller, about 8 mm. long, and more markedly rostrate, dehiscent by paired terminal pores; the 4 uppermost subtruncate, about 1 cm. long, dehiscent by paired terminal pores from a short, sharply reflexed rostrum; ovary linear, tomentose; fruit pos- sibly somewhat broader than typical for the species. Panama and Colombia? ANAL ZONE: Barro Colorado Island, Bangham 603, Kenoyer 381; Gatun, Hayes 548. COLÓN: Río Indio de Fató, Pittier 4253. This variety approaches C. oxyphylla var. dariensis, from which it may be distinguished in having the "fruticosa" lower leaf pubescence (shorter, more ap- pressed hairs), while C. oxyphylla var. dariensis has the more tomentulose, raised- hair pubescence characteristic of C. Maxonii in Panama. 37. Cassta CAUDATA Standl. in Contr. U. S. Nat. Herb. 18:102. 1916. Chamaefistula caudata (Standl.) Britt. & Rose, in N. Am. Fl. 23:237. 1930. Shrub or small tree, the branchlets terete, glabrous. Leaves large, 4-foliolate; petioles elongate, about 10 cm. long, subterete, glabrous; rachis like petiole, about 7 cm. long, bearing a prominent subconic gland between the basal leaflets and a smaller apical gland or protuberance; stipules linear, caducous; leaflets ovate to elliptic, up to 25 cm. long and 7 cm. wide, obtuse or rounded basally, acuminate apically into a caudate tip about 3 cm. long, glabrous above and below; petiolules about 5 mm. long, dark, rugose. Inflorescence axillary or terminal, paniculate, several-flowered, puberulent; bracts linear, inconspicuous, about 2 mm. long; pedicels slender, up to 4 cm. long. Flowers yellow, large and showy; sepals un- equal, ovate to oblong-orbicular, up to 9 mm. long and 7 mm. wide, blunt or rounded, puberulent; petals mostly obovate, up to 3 cm. long and 17 mm. wide, venose, puberulent on the veins; fertile stamens 7; the 3 lowermost with large, oblong, markedly falcate anthers almost 1 cm. long, apically long-rostrate and dehiscent by a pair of terminal pores; 4 median anthers oblong, about 7 mm. long, only slightly falcate, short-rostrate, dehiscent by a pair of terminal pores; ovary linear, tomentose. Legume not known in maturity; seeds transverse. Panama and Costa Rica. PANAMA: upper Mamoni River, РИ ет 4401. NAMES OF UNCERTAIN APPLICATION TO CASSIA IN PANAMA C. confusa Rose. Apparently no description of this species ever appeared, although it is listed in the key in Standley's ‘Flora of Panama Canal Zone’ (Contr. (382) 1951] FLORA OF PANAMA (Leguminosae ) 83 U. S. Nat. Herb. 27:199. 1928). The position in Standley's key might indicate the plant in mind to have been C. leptocarpa var. hirsuta, although this variety has not to our knowledge yet been reported from the Canal Zone. In any event, the specific name "confusa" would be untenable due to prior use in Cassia. C. TRISTICULA HBK. (Nov. Gen. & Sp. 6:367. 1824). Hemsley reports this species from Panama, but no specimen so identified has come to my attention. Without reference to type material or without Hemsley's cited specimen, it is im- possible to be certain whether or not the name has any reference to Panamanian material, as a synonym or otherwise. Judging from the original description, the species would fall close to Cassia stenocarpa in the “Chamaecrista”? section of the genus. Kunth (HBK.) mentions its affinity to C. patellaria, but if the description be correct it differs from C. patellaria in having stipitate petiolar glands. 17. DELONIX Raf. Drroix Raf. Fl. Tellur. 2:92. 1836. Moderate, unarmed trees. Leaves twice-pinnate, the pinnae several pairs and opposite on the rachis, each pinna with numerous opposite leaflets; petiole mod- erately short, expanded basally, eglandular; rachis elongate, flattened above, egland- ular; stipules caducous; ultimate leaflets small, short-petiolulate. Inflorescence terminal or axillary, subcorymbose (of 1 to several racemes). Flowers showy; calyx-tube short; calyx-lobes subequal, much exceeding the tube, valvate; petals separate and spreading, long-clawed; stamens 10, subdeclinate, free, the anthers versatile and longitudinally dehiscent; ovary scarcely stipitate; style slender; stigma truncate. Legume thick-flattened, dehiscent, the valves ligneous; seeds transverse. Africa and Madagascar. 1. Drrowix REGIA (Bojer) Raf. Fl. Tellur. 2:92. 1836. Poinciana regia Bojer in Hook. Bot. Mag. pl. 2884. 1829. Spreading tree, the branchlets subglabrous, lenticellate, corky in age. Leaves large; petiole usually about 10 cm. long, flattened and somewhat sulcate above, swollen basally; rachis up to 5 dm. long, somewhat nodose and cross-partitioned at insertion of the pinnae, tomentulose or puberulent; stipules sagittate-bifurcate (the lobes frequently dissected), caducous; pinnae up to about 20 pairs, spreading, the rachis about 12 cm. long, tomentulose; ultimate leaflets up to 40 pairs, oblong, usually nearly 1 cm. long and 3—4 mm. wide, inequilateral basally, rounded or obtuse apically, usually tomentulose or puberulent above and below especially along the veins, lighter below. Inflorescence as described for the genus; bracts ovate- lanceolate, about 6 mm. long; pedicels up to 10 or more cm. long in lowermost flowers. Flower red, very ornamental; calyx-tube from pedicellar articulation about 7 mm. tall, shallow within; calyx-lobes spatulate-lanceolate, 2.5—3 cm. long, valvate in bud, reflexed in age; petals large, about 6 cm. long, spreading, the claw about 3 cm. long, expanding apically, tomentulose within, the blade suborbicular, (383) [Vor. 38 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN usually 3—3.5 cm. wide, glabrous; stamens about 4 cm. long, tomentose basally; anthers ovate, 4—5 mm. long, bilocular; ovary linear, 1 cm. long or longer, lightly hirsute; style about 3 cm. long, glabrous; stigma terminal, pubescent, unexpanded. Legume broadly linear, frequently 4—6 dm. long and 5—7 cm. wide, scarcely curved, compressed, 2-valved, the valves ligneous; seeds oblong, transverse. Native to Madagascar, introduced into world tropics and subtropics; commonly planted in Central America, West Indies and South America. CANAL ZONE: Balboa, Standley 30821; Bellavista, Macbride 2764. This colorful tree, the "Royal Poinciana” or “Flamboyant,” is one of the most frequently planted tropical ornamentals. Its brilliance during the relatively short owering season offers Temperate Zone visitors a taste of the expected but often lacking "floral splendor of the tropics.” 18. PELTOPHORUM Walp. PELTOPHORUM (Vogel) Walp. Rep. 1:811. 1842, nom. conserv. Baryxylum Lour. Fl. Cochinch. 266. 1790. Unarmed spreading trees. Leaves twice-pinnate; petiole and rachis eglandular; stipules small, caducous; pinnae several; ultimate leaflets numerous, relatively small. Inflorescence terminal, of few to several racemes. Flowers yellow; calyx-tube short, patelliform; calyx-lobes 5, imbricate, subequal; petals 5, spreading, subequal; stamens 10, inserted with petals on calyx-tube, similar; filaments free, pubescent basally; anthers versatile, longitudinally dehiscent, ovary free, usually stipitate; style elongate; stigma expanded, peltate. Legume indehiscent, winged on both margins, few-seeded. West Indies, South America, Africa, eastern Asia, East Indies, and Pacific islands. A Far Eastern species has been introduced into Panama. 1. PELTOPHORUM INERME (Roxb.) Naves, ex Villar in Blanco, Fl. Filip. Nov. App. 69, 2. 335. 1880. Caesalpinia inermis Roxb. Fl. Ind. 2:367. 1832. ое ferruginea Decne. in Nouv. Ann. Mus. d'Hist. Nat. Paris 2:462. 1833; Miq. » Bat, 1:111. 1855. Caesaipinis arborea Zoll. in Nat. en Geneesk. Archief 3:65. 1846; Miq. loc. cit. 112. , fide Villar in Blanco, loc. cit. 18 i ferrugineum Benth. Fl. Мини. `2:279. 1864. Tall tree, the branchlets puberulent to glabrous in age. Leaves large; petiole about 5 cm. long, ferruginous-tomentulose; rachis similar, up to 2 or more dm. long; pinnae several pairs, opposite, borne from flattened upper surface of rachis; ultimate leaflets several to many pairs to the pinna, oblong, about 15 mm. long and 6 mm. wide, markedly inequilateral basally, rounded-subtruncate apically, glabrous and darker above, ferruginous-puberulent to subglabrous below; stipules bifurcate, early caducous. Inflorescence densely ferruginous-tomentose, a terminal panicle of racemes up to a few dm. long, or the racemes axillary and subterminal; pedicels (384) 1951] FLORA OF PANAMA (Leguminosae) 85 less than 1 cm. long; buds globular. Flowers yellow; calyx-lobes elliptic, almost 1 cm. long, ferruginous-tomentulose without, glabrous within, imbricate; petals obovate, about 1.5 cm. long, contracted basally into a ferruginous-hirsute claw; stamens free, about 1 cm. long, the filaments densely pubescent basally, the anthers about 3 mm. long; ovary ferruginous-pubescent; stigma terminal, light in color, expanded and peltate. Legume narrowly elliptic, about 10 cm. long and 2 cm. wide, flat, indehiscent, longitudinally venose, with an undulate marginal wing 2-3 mm. wide; seeds longitudinal. Endemic to Malayan area; introduced into New World tropics. CANAL ZONE: Balboa, Allen 4468, Standley 30852; without locality, Johansen 27. This ornamental shade tree is reported fairly common in the Canal Zone, Ч чу 525 Э д in R ё Y 2 | Қы a ze = ЖА ; ЖЫ еб A TR Fig. 129. Peltophorum inerme (385) [Vor. 38 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN 19. SCHIZOLOBIUM Vogel SCHIZOLOBIUM Vogel, in Linnaea 11:399. 1837. Tall, unarmed trees, the leaves large, bipinnate, with numerous, small leaflets. Inflorescence axillary or terminal, racemose, or paniculate of several racemes; bracts small. Flowers yellow, perfect; calyx with a short, somewhat inequilateral tube, the lobes imbricate, longer than the tube, reflexed in anthesis; petals separate, im- bricate, narrowed baseward, subequal; stamens 10, free, subdeclinate, inserted with the petals on upper portion of calyx-tube; the anthers all similar, small, bilocular, versatile, dehiscent by longitudinal slits; ovary stocky, tomentose, short-stipitate, affixed basally on lower side to the calyx-tube; style about twice as long as the ovary; stigma terminal, minute. Legume flattened, coriaceous, tardily dehiscent, bearing a single seed in the expanded apical portion. The genus seemingly consists of a single species. DV Эллэ, au рг pum Ж MET P^ ер, p v ES b» ав 5 ТА ЖОК ат E. 7 i gg Л сырыы, СЕ / WAS > k ) aX 1% : 4 Vi 2 | 224220 2 Ж. 7 | Уу, Ж 4С 2 Уа 7% A YD Nee Eff Foeschner Біз. 130. Schizolobium parahybum (386) 1951] FLORA OF PANAMA (Leguminosae ) 87 1. SCHIZOLOBIUM PARAHYBUM (Vell.) Blake, in Contr. U. S. Nat. Herb. 20:240. ESI. Cassia parabyba Vell. Fl. Flum. 168. 1825; Ic. 4:f. СЯ 1827. Schizolobium excelsum Vogel, in Linnaea 11:399. 183 Schizolobium glulinosum Tul i in Archiv. Mus. oe ‘Nar Paris 4:157. 1844. Schizolobium Kellermannii Pittier, in Contr. U. S. ee ИЗД. 18:232. 1917. A tall, buttressed tree to 25 m. or more, the branchlets subglabrous. Leaves very large, twice-pinnate, multifoliolate; petiole stout, 1 dm. or more long, evi- dently eglandular, glabrous or somewhat viscid, slightly flattened or sulcate above; secondary petioles similar, about 1 cm. long, callous basally; rachis several dm. long, eglandular, flattened and margined above, the pinnae (several to many) arising opposite in pairs on the upper side; stipules apparently small, caducous; leaflets several to many pairs on each pinna, oblong or linear-oblong, 1.5-3 cm. long, 4-7 mm. wide, rounded apically and basally, dark and puberulent to glabrous above, lighter and appressed-pubescent below, subcoriaceous, the midvein very prominent, the lateral veins obscure; ultimate petiolules about 1 mm. long. In- florescence multiflorate, as described for the genus; bracts lanceolate, scarcely 2 mm. long; pedicels up to 1 cm. long in age, articulate above the middle. Flowers attractive, yellow; calyx-tube turbinate, 2-3 mm. long, dark and usually tomentu- lose without; calyx-lobes ovate-elliptic, about 6 mm. long and 3 mm. wide, puberu- lent; petals obovate-spatulate, almost 2 cm. long and usually 4—6 mm. wide, sub- glabrous, obscurely veined; stamens 10, as described for the genus, about as long as the petals, the filaments broader and scurfy basally, the anthers broadly elliptic, about 2 mm. long; ovary subfalcate, about 5 mm. long and 2 mm. broad, hispid- tomentose with dark hairs, few-ovulate. Legume obovate-spatulate, narrowed baseward, about 10 cm. long and 2.5-5 cm. wide, glabrous, bearing the solitary seed apically; seed flattened-ovoid, up to 2 cm. long. Mexico; Central America; South America. ANAL ZONE: along R. Chagres, Steyermark & Allen 16785. PANAMA: Chorrera, ie 1500. 20. CAESALPINIA L. CAESALPINIA L. Sp. Pl. 380. 1753. Poinciana L. loc. cit. 380. 1753. Guilandina L. loc. cit. 381. 1753. Poincianella Britt. & Rose, in N. Am. Fl. 23:327. 1930. Many other synonyms can be found for Caesalpinia considered in its broad sense. Some that 12. ол so included, in addition to the ones cited above, are: Adenocalyx Bert.; Balsamocarpon Clos; Bian Todaro; Brasilettia (DC.) өзгөр: Cam pec. d суза жайы tia Zoll.; шла; Vog.; Conzattia Rose; Coulteria HBK.; Ery Dunes Link; Guaymasia Britt. & Rose; Guilandia P. Br.; Larrea Ortega; Lebidibia ma. Melano- sticta DC.; Moparia Britt. & Rose; Nicarago Britt. & Rose; Pomaria Cav.; Pseudosantalum (387) [Vor. 38 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN е Russellodendron Britt. & Rose; Schrammia Britt. & Rose; Tara Molino; Ticanto кенеді to warrant inclusion іп опе genus, w while if considered as separate genera they cause in the herbarium considerable confusion and obscuring of the species, to sors nothing of — мө then in the almost impossible task of organizing a able generic r purposes of the M of Panama,’ Caesalpinia is Md in its broader sense, els following Bentha Trees, shrubs or sometimes vine-like or subherbaceous, armed or unarmed. Leaves twice-pinnate; petioles and rachis eglandular, glabrous or variously pubes- cent; pinnae usually several and opposite or subopposite on the rachis; ultimate leaflets few to many, opposite in pairs, or less frequently alternate, in the pinna. Inflorescence mostly racemose, variously pubescent; bracts normally caducous. Flowers yellowish or less commonly pinkish; calyx with a conspicuous tube and 5 imbricate lobes, the outermost lobe more or less cucullate and enclosing the bud; petals free, nearly equal, inserted upon the upper calyx-tube; stamens normally 10, free, inserted with the petals on the calyx-tube, usually subdeclinate; filaments mostly pubescent, often glandular; anthers small, bilocular, longitudinally de- hiscent, versatile; ovary free, sessile or short-stipitate, inserted at base of calyx- tube; stigma terminal, usually not dilated. Legume compressed or flattened, usually unarmed and dehiscent; seeds transverse. Widely distributed, in tropics and subtropics of both hemispheres. a. Shrubs or trees, lightly armed or unarmed; bracts of энин їп- conspicuous or missing; legume ME Suid or coiled. b. Inflorescence glabrous, large and showy; n естің шү 7- ‚ long; пеки Ке, exserted, about 5 cm. long; plant often Hitlers Min bb. — ан ber = to tomentose, smaller; pedicels less than 2 ‚ long; s short, less than 1.5 cm. long; plant unarmed. 6 feu sonas, red 10 or more cm. long, with a ferruginous- stellate tom owers larger, 2—3 cm. wide; legume fla (valves may it ge after dehiscence) сс; opem condensed, mostly less than 4 s long, ng vo коа rulent; flowers rs pone less than wide; leg coiled.. 3. C. CORTARIA aa. Vine- ‘like shrub of sea beaches, the stem an * v T heavily a arm 4 wit recurved thor PR of w Қамы about 1 cm. lon , persistent at least until нгаб legume armed, compressed, but not flat ИРИП 4. C. CRISTA . C. PULCHERRIMA C. ERIOSTACHYS 1. CAESALPINIA PULCHERRIMA (L.) Sw. Obs. 166. 1791. Poinciana 2225 ӛр. РІ. 380. 175 Poinciana bijuga ЕІ. ee 260. ug non L. Poinciana elata p p cit. 261. 1790. Lightly or scarcely armed shrub or small tree, the branchlets glabrous. Leaves moderately large, twice-pinnate; petiole 2-8 cm. long, terete, eglandular, glabrous; rachis like petiole, up to 2 or more dm. long; stipules lanceolate, minute, caducous; pinnae up to 8 or more pairs, opposite on the rachis; leaflets about 10 pairs to the pinna, subopposite, mostly oblong, 10-23 mm. long and 5—10 mm. wide, rounded apically, obtuse and somewhat inequilateral basally, glabrous, membranaceous, petiolulate. Inflorescence usually terminal, a corymbose raceme, several- to many- (388) 1951] FLORA OF PANAMA (Leguminosae) 89 flowered; bracts similar to stipules, caducous; lower pedicels elongate, not uncom- monly 7-8 cm. long, glabrous. Flowers ornamental, red to yellow; calyx-tube narrowly turbinate, 3—4 mm. long, the pedicellar stalk subarticulate a few mm. below the tube; mature calyx-lobes ovate or lanceolate to obovate, about 1 cm. long, glabrous, imbricate in bud, the outer calyx-lobe cucullate and somewhat larger; petals free, obovate, about 2 cm. long, glabrous, clawed; stamens 10, in- serted with petals on rim of calyx-tube, strongly exserted; filaments free, about 5 cm. long, glabrous except basally where viscid-pubescent; anthers ovate, 1-2 mm. long, versatile, bilocular, longitudinally dehiscent; ovary linear, glabrous, stipitate from base of calyx-tube. Legume linear-oblong, up to 12 cm. long, usually some- what wider apically than basally, obliquely acute both apically and basally, short- stipitate, flattened, glabrous, elastically dehiscent; seeds ovate, transverse. Widely cultivated or escaped throughout world tropics; probably native in northern Central America. CAS DEL TORO: Changuinola Valley, Dunlap 423; Old Bank Island, von Wedel 1860. ANAL ZONE: Balboa, Standley 30834; Paraiso, Standley 30009. PANAMA: Taboga Island, Bro. Celestine ОТ. An attractive ornamental frequently seen in cultivation in the Canal Zone and other tropical regions. The legume is reported to contain abundant tannin, but it is not a recognized article of commerce as is the "divi-divi" (legume of Caesal pinia coriaria). The leaves have been reported used for fish poison. 2. CAESALPINIA ERIOSTACHYS Benth. Bot. Voy. Sulphur, 88. 1844. Schizolobium Covilleanum Pittier, in Contr. U. S. Nat. Herb. 18:231. 1917. Poincianella eriostacbys (Benth.) Britt. & Rose, іп М. Am. Fl. 23:332. 1930. A small or moderate, unarmed tree of the Pacific ара the branchlets usually stellate-pubescent. Leaves moderately large, twice-pinnate; petioles somewhat flat- tened above, up to 2 cm. long, ferruginous-stellate-pubescent and also with some lighter, simple hairs, eglandular; rachis 10—20 or more cm. long, similar to the petiole, usually bearing 11—21 pinnae; stipules oblong, about 6 mm. long, rounded at the apex, caducous; pinnae usually subopposite on the rachis, up to 8 cm. long; ultimate leaflets 14—29 to the pinna, alternate, more or less oblong, 5-10 mm. long and 3—5 mm. broad, strongly inequilateral basally, rounded apically, sessile, darker, and usually subglabrous above, pubescent with simple hairs or subglabrous below. Inflorescence terminal or subterminal, consisting of a number of several-flowered racemes, the peduncle and pedicels densely ferruginous-stellate-pubescent; bracts similar to stipules; mature pedicels 1—2 cm. long. Flowers yellow, ornamental, often appearing before the leaves; calyx-tube cupulate, 5—6 mm. long, densely ferruginous-stellate-pubescent; calyx-lobes ovate-elliptic, about 8 mm. long and 3—5 mm. broad, imbricate and outer one subcucullate in bud, ferruginous-stellate- pubescent without, light-tomentose within; petals suborbicular, up to 14 mm. long and 12 mm. broad, clawed, dark-dotted towards the middle, pubescent base- ward on the claw; stamens 10, free, inserted with petals on the calyx-tube; fila- (389) [Vor. 38 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 131. Caesalpinia eriostachys ments about 12 mm. long, more or less glandular-pubescent, densely pubescent basally; anthers ovate, 1-2 mm. long, longitudinally dehiscent, bilocular; ovary sessile from base of calyx-tube, short, pubescent, few-ovulate; style elongate; stigma terminal, small. Legume linear-obovate, about 10 cm. long and 2—3 cm. broad, flattened, elastically dehiscent, obliquely beaked apically, short-stipitate, puberulent or tomentulose; seeds few, oval, about 12 mm. long, obliquely transverse. Northern Mexico to Panama; Cuba. сос: Aguadulce, Pittier 5105; R. Mata Ahogado, Allen 131. PANAMA: Matias Hernández, Pittier 6916; San José Island, Johnston 565. 3. CAESALPINIA CORIARIA ( Jacq.) Willd. Sp. Pl. 2:532. 1799. Poinciana coriaria Jacq. Select. Stirp. Am. 123. 1763 Caesalpinia thomaea Spreng. Syst. 2:343. 1825, fide М. Am. FI. Libidibia coriaria Schlecht. in Linnaea 5:193. 1830 An unarmed, low, crooked, much-branching tree, the branchlets glabrous. Leaves moderate, twice-pinnate; petiole rather stout, usually 1-2 cm. long, lightly pubescent, eglandular, terete but somewhat flattened above, callous and expanded basally; rachis similar, nodose at insertion of the pinnae, eglandular; stipules cadu- (390) 1951] FLORA OF PANAMA (Leguminosae) 91 cous; pinnae usually about 15, 3-5 cm. long, opposite or subopposite on the rachis except for the (odd) terminal one; ultimate leaflets as many as 35 pairs to the pinna, linear-oblong, 3-9 mm. long and up to 2 mm. wide, rounded apically, in- equilateral and subcordate basally, glabrous. Inflorescence a cluster or panicle of a few condensed racemes, the racemes several- to many-flowered, not exceeding the leaves; bracts caducous; pedicels scarcely 3 mm. long. Flowers small, yellow, fragrant; calyx-tube turbinate, about 2 mm. long, glabrous; calyx-lobes ovate, about 3 mm. long, imbricate and somewhat contorted in bud; petals suborbicular, 3-4 mm. long, short-clawed, glabrous; stamens 10, about 6 mm. long, inserted with petals on rim of calyx-tube; filaments expanded and pubescent basally; anthers ovate, scarcely 1 mm. long, bilocular, longitudinally dehiscent, versatile; ovary sublinear, glabrous, scarcely stipitate, inserted on base of calyx-tube; style glabrous, scarcely longer than the ovary; stigma truncate. Legume oblong, up to 6 cm. long and 2 cm. wide, flattened, thick, much coiled in age, apparently indehiscent. Mexico to Panama; West Indies; northern South America. NAL ZONE: Exp. Gardens, Lindsay 253; Summit, Mell 11. сосіЕ: Aguadulce, Pittier 4974. The legumes and bark of this tree are relatively rich in tannin, the pods report- edly having as much as 30 per cent tannin content. Besides being utilized locally in the preparation of dyes and inks, the pods are an item of considerable export interest from the tropical countries, especially the north coast of South America. The matured pods usually appear in the trade under the name “divi-divi,” a com- mercial source of tannin for many years. 4. CAESALPINIA CRISTA L. Sp. Pl. 380. 1753.2 Guilandina Bonducella L. D Pl. ed. 2, 545. d (in part). Guilandina semina Lour. Fl. Cochinch. 265. 1 ?Caesal pinia Bonducella da Fleming, Asiat. e 11:159. 1810 (= C. Вомрос, fide Guilandina Bonduc var. minus DC. Prodr. 2:480. 1825. Guilandina crista (L.) Small, EL Deer U. S. 591. 1903. Vine-like shrub of sea beaches, heavily armed with recurved thorns and forming impenetrable thickets above tide level, the branchlets pubescent. Leaves large, twice-pinnate; petiole usually 5-10 cm. long, subterete except basally where swollen and flattened above, tomentose, armed like the stems; rachis 2 or more dm. long, eglandular, tomentose and armed like the petiole; stipules foliaceous, usually of 2 “leaflets” a few cm. long; pinnae about 7 pairs, up to 15 cm. long, opposite andy and Exell (Jour. Bot. 76:175-180. 1938) show that the name C. crista as commonly E should refer to a different, smooth-fruited species. It should then be correctly replaced by C. Bonduc (L.) Roxb., and the species in recent times generally аеру as C. Bonduc should 1 become C. major (Medic.) Dandy & Exell. Thus our gray-seeded, armed-fruited species here listed as C. crista, and so regarded in most 20th-century literature and in быз. is probably in reality Bonduc, whil А C. E 7 E urere literature and most herbaria is C. j the names C. cris h be found applying to er of two different ja ista and С ith I have followed Е the EAT no fide Dandy & Exell), practice of listing the gray- seeded nickar-nut as C. crista. (391) [Vor. 38 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 132. Caesalpinia crista from the upper surface of the rachis, armed, tomentose; ultimate leaflets about 8 pairs to the pinna, ovate-elliptic, 2-6 cm. long and 1-2.5 cm. wide, obtuse and mucronulate apically, rounded basally, puberulent on veins and margins. In- florescence axillary or subaxillary, racemose, several- to many-flowered, the peduncle armed; bracts linear-acuminate, about 1 cm. long, subpersistent, reflexed, tomen- tose; pedicels about 4 mm. long, rufous-tomentose. Flowers moderately small, brownish-yellow; calyx-tube broadly turbinate or subcupulate, 2-3 mm. long, rufous-tomentose without; calyx-lobes ovate to obovate-elliptic, 5-8 mm. long, imbricate in bud, the outermost subcucullate, rufous-tomentose; petals narrowly (392) 1951] FLORA OF PANAMA (Leguminosae) 93 oblong, scarcely longer than the calyx-lobes, subglabrous, clawed; stamens 10, free, inserted with petals on rim of calyx-tube; filaments about 7 mm. long, pubescent; anthers ovate, about 1 mm. long, versatile, bilocular, longitudinally dehiscent; ovary ovate-oblong, stipitate from base of calyx-tube, pubescent, the style short, the stigma small. Legume oblong-orbicular, usually 5-6 cm. long and about 4 cm. wide, compressed, short-stipitate, densely covered with sharp prickles, tardily dehiscent; seeds usually 2, ovoid, about 2 cm. long, grayish. Widely distributed in world tropics, apparently native to eastern Asiatic region. CANAL ZONE: For ив. 22 di 213. cHIRIQUÍ: San Bartolomé, уа 6 Schery 943. p Santa Clara beach, Woodson, Allen & Seibert 1703. cor Fató, Pittier 3038. PANAMA: “Bella Vista, pd 12005, Standley 25310; San José Island of Pearl Islands, Johnston 722 A very distinctive е easily distinguished from all other species in Central America except C. Bonduc (— C. major) by its strand habitat, dense armament, and characteristic legume. It often forms impassible thickets on the Pacific shores of Panama, where it is studiously avoided by the passer-by. CAESALPINIA UROPHYLLA (Donn. Smith) Standl. (C. bonducella var. uro- phylla Donn. Sm.; Guilandina urophylla Britt. & Rose), armed much as is Caesal- pinia crista L. of the coastal areas, has been found at high elevations in Costa Rica. Possibly this unusual vine will be found in western Panama also. 21. HAEMATOXYLON L. HAEMATOXYLON L. Sp. Pl. 384. 1753. Haematoxyllum Scop. Introd. Hist. Nat. 225. 1777. Haematoxylon Brasiletto Karst. (Fl. Colomb. 2:27, pl. 114. 1862) is reported from both Costa Rica and Colombia, and is to be expected in Panama. It is a small armed tree, with leaves of about 3 pairs of obovate-cuneate, glabrous leaflets; small, yellow, long-pedicellate, short-racemose flowers; and a flat, thin, oblong legume. The wood is said to be employed for the same purposes as true logwood, H. campechianum. H. boreale S. Wats. is apparently a synonym of Н. Brasiletto. 22. CERCIDIUM Tul. The genus Сексілом Tul. (Arch. Mus. d'Hist. Nat. Paris 4:133. 1844) (Rhetinophloeum Karst.), as represented by C. praecox (К. & P.) Harms, may eventually be found in Panama. It has been collected in Colombia and other South American localities, and in Mexico. The plant is similar to PARKINSONIA, but bears the small bipinnate leaves distinct from the spines. The rachis of the pinna is terete and less than 4 cm. long. The inflorescence is a short corymb scarcely ex- ceeding 3 cm., and the calyx-lobes are valvate or induplicate-valvate. 23. PARKINSONIA L. ParKINSONIA L. Sp. Pl. 375. 1753. Small trees or shrubs, the branches prominently armed. Leaves twice-pinnate, (393) [Vor. 38, 1951] 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN although sometimes not obviously so because of (1) frequent fall of ultimate leaflets from flattened secondary rachis, and (2) extreme condensation of the primary rachis; pinnae few, approximate, almost "axillary" from the upper surface of a stout spine (spine— at least partially the modified petiole and primary rachis) ; stipules apparently modified as small lateral thorns from base of larger spine; leaf- lets of the pinna many, minute, often caducous; rachis of pinna flattened, green, eglandular. Inflorescence racemose, several-flowered, arising from the axils. Flowers moderate; calyx with a short, turbinate tube, the lobes imbricate, a few times longer than the tube; petals free, clawed, subequal; stamens 10, free, the anthers versatile and longitudinally dehiscent; ovary essentially free, straight, slender, subterete, scarcely stipitate. Legume tardily dehiscent, subterete and swollen at point of seed development, constricted and flattened between the seeds; seeds few, longitudinal. American tropics and subtropics; Africa. 1. PARKINSONIA ACULEATA L, Sp. Pl. 375. 1753. Parkinsonia Thornberi M. E. Jones, Contr. West. Bot. 12:12. 1908, fide I. M. Johnston. Xerophytic shrubs or small trees to several m. tall, the branchlets subglabrous, more or less flexuous and gnarled. Leaves multifoliolate, twice compound as de- scribed for the genus; petiole and rachis modified into a spine 5—15 mm. long, bearing from the upper side 1-3 pairs of pinnae; pinnae multifoliolate, the rachis linear, 2—3 dm. long and 1-2 mm. wide, flattened, green; ultimate leaflets about 25 pairs, linear-oblong, 2-8 mm. long and up to 2 mm. wide, often caducous. Racemes up to 2 dm. long, subglabrous; bracts lanceolate, about 1 mm. long; pedicels slender, 1-2 cm. long in age. Flowers yellow; calyx-tube turbinate, scarcely 2 mm. long; calyx-lobes ovate-lanceolate, 6-7 mm. long and about 3 mm. wide, subglabrous; petals 5, about 13 mm. long, the blade suborbicular. about 8 mm. long, claw pubescent at base, about 5 mm. long; stamens 8—9 mm. long, pubescent basally; ovary linear-oblong, about 4 mm. long, hispid; style linear, about equalling the stamens, glabrous; stigma terminal, minute. Legume linear, up to about 15 cm. long, nodose by constriction between the seeds, glabrous, striate-nerved; seeds about 1 cm. long. Found escaped or planted in the Americas from southern United States to Argentina and West Indies; from uncertain (American: Mexican?) origin. The species has been reported from Panama, although no specimens collected there have yet come to our attention. (Leguminosae to be continued in Part V, Fasc. 4) (394) JESSE MORE GREENMAN (1867—1951) Annals of the Missouri Botanical Garden Vol. 38 MAY, 1951 No. 2 JESSE MORE GREENMAN (%1867-11951) ROBERT E. WOODSON, JR. The many friends of Dr. J. M. Greenman were saddened by his death on January 20, 1951, at the age of eighty-three years. Jesse More Greenman was born on December 27, 1867, at North East, Erie County, Pennsylvania, the son of James William and Clarissa (More) Greenman. Entering the University of Pennsylvania in 1888, he served successively as assistant (1890—92) and as instructor (1893—94) in botany, graduating in 1893 with the degree of Bachelor of Science. In 1894 he transferred to Harvard University where he was assistant in the Gray Herbarium until 1899, when he was awarded the degree of Master of Science. It was during these years at the Gray Herbarium that he became associated with the late Benjamin Lincoln Robinson and was a fellow student with the late Merritt Lyndon Fernald. At that time botanists of the United States were only beginning to appreciate fully the rich opportunities in the floristics of tropical America, and particularly of Mexico. Following the example of his illustrious predecessors at the Gray Herbarium, Robinson plunged into the determination of the fascinating collections of Edward Palmer, C. J. Pringle, L. C. Smith, E. W. Nelson, and others. In this he was aided materially by Greenman and to a lesser extent by Fernald (whose real interests already were in the flora of the north- eastern seaboard). In these years Robinson evidently became a model for the somewhat younger Greenman, both professionally and personally, and the fancied emulation has been apparent to many of the friends and students of both men, amongst whom the writer of this notice is fortunate to be included. Both could be recognized in- stantly as "gentlemen of the Old School": courtly and immaculate, soft-spoken and deliberate in word and action; innately conservative and studied, perhaps, to most of their younger contemporaries. In the case of Robinson, lightened daily by a characteristic humane warmth and tolerance even in his declining yeers, the (95) [Vor. 38 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN total effect possibly was more original to those who knew them both. In Green- man, it has long appeared to the writer of this notice that the true man lay some- what deeper beneath the surface, and was only occasionally evident in delightful flecks of Barriesque humor which his friends will always cherish. The parallelism of Robinson and Greenman is striking in their professional careers: while Robinson elected the vast genus Eupatorium as a life work, Green- man chose the even more appalling genus Senecio. Neither lived to complete his task. While Robinson went to the University of Strasbourg to receive his doctor- ate of philosophy under De Bary, Greenman went as a Kirkland Fellow (1899— 1901) to study under Adolf Engler at the University of Berlin, which conferred upon him the Ph.D. in 1901. Returning to Harvard in 1902, he married Anne Louise Turner and resumed his duties as assistant at the Gray Herbarium, having also been appointed instructor in botany (1902—05) at the same time that Fernald received a similar promotion. Greenman’s teaching assignment, interestingly, was to conduct a class in plant geography; Fernald, with only his B.S. from Harvard College, was to conduct an elementary class. (Robinson, save for "research in- struction," never taught a formal class during his entire career at Harvard!) In my mind's eye I have long projected a fancied crisis in the little group of botanists at the Gray Herbarium in the year 1905. In that year there developed the possibility for a promotion to assistant professor at Harvard for one of the two younger men; there also came a request from the Field Columbian Museum of Chicago for an applicant for the post of assistant curator of botany under the late Charles F. Millspaugh. I have long taken it as a fitting tribute to the perception and moral strength of Robinson that he chose Fernald to remain with him at Har- vard rather than Greenman, with whom he had so much more in common. The decision may have reflected the needs of the forthcoming seventh edition of ‘Gray’s Manual’ (published in 1908), but I am inclined to believe that Robinson saw in Greenman too much of himself for the best interests of the Gray Herbarium. I am sure that the decision was a bitter disappointment to Greenman; but his ad- miration for Robinson was never dimmed. Both he and Mrs. Greenman long half- expected their eventual return to Cambridge, which they loved so well. Greenman took up his duties as assistant curator at Chicago in 1905. In 1908, perhaps to offset the defection-from-the-ranks of taxonomy by the late John Merle Coulter, he was appointed assistant professor at the University of Chicago, teaching taxonomy and plant geography. I suspect that working conditions at the Field Museum were none too pleasant for all concerned, and also that the emphasis placed upon morphology and Coulter’s later contempt for taxonomy (which seems to continue at that University to this day) were rather discouraging to a young botanist in the tradition of Engler, Robinson, Watson, and Gray. At any rate, when he was invited to join the staff of the Missouri Botanical Garden and the Shaw School of Botany of Washington University in 1913, Greenman was quick to accept. With him to St. Louis came the Senecios which were to be his com- panions until his death. 1951] WOODSON—JESSE MORE GREENMAN ЭУ. I believe that the Greenmans discovered іп St. Louis at least partial consolation for the loss of Cambridge. At the Missouri Botanical Garden the new director, George T. Moore (an old friend from Harvard days), had gathered together a small group of other former Harvard associates which could compare in brilliance and accomplishment with the botanical staff of their old alma mater herself. With its almost unexcelled botanical library, its large herbarium of unplumbed value ("unplumbed" because it was so disorganized!), its new and modern physiological laboratories, its greenhouses, its newly launched ANNALS, and perhaps above all its Lackland Fellowships sagaciously maneuvered by Dr. Moore from his Board of Trustees, the Missouri Botanical Garden and the Shaw School of Botany achieved national, if not international, significance almost over night. Dr. Greenman became curator of the herbarium of the Missouri Botanical Garden and associate professor of botany at Washington University in 1913; in 1917 he was promoted to full professor at the University. Аг the Garden, he found an Augean stable in the herbarium. Founded by the purchase of the per- sonal collection of Bernhardi in 1857 and the bequest of that of Engelmann in 1884, it had been augmented copiously but indiscriminatingly and presided over rather casually in the succeeding years. No detailed inventory of its contents existed. Greenman estimated the total number of specimens at 614,660, and began a painstaking system of accession records. Then began the almost hopeless task of orderly arrangement of this considerable museum, the laborious identification of the extremely valuable historical material from the scrappy bits of evidence ac- companying them (particularly in the Bernhardi herbarium), and of course the never-ending struggle to keep abreast of current accessions as well—all virtually single-handed. With his rather stern sense of duty as he saw it, it was inevitable that his own research should be thrust almost out of sight into the background. There can be little doubt that Greenman’s new role as a teacher in the Shaw School of Botany utilized his character and training most advantageously. He was a willing, thorough, and patient teacher. Where, in his day (or in ours?) could one find another prepared (or even interested) to offer courses of graduate level in bryophytes, pteridophytes, gymnosperms, angiosperms, and plant geography? Neither was he averse to undergraduate instruction, and for a number of years conducted the beginning course in botany in addition to his other duties. Although not an original lecturer, Greenman’s gentle courtliness everlastingly endeared him to all his students at Washington University. It was not so much what he said as the way that he said it; a rather pedantic reference to dichotomy could be accompanied by a glance that would warm his hearer with ill-defined pleasure. But at once the most cultivated and gracious of men, he could be the most punctilious and unyielding. Organizational activities were attractive to Greenman, and he was in frequent attendance at scientific conventions. He was a member of the Botanical Society of America, the Ecological Society of America, the American Society of Natural- ists, the New England Botanical Club, the Germanistic Society of St. Louis, the American Association of University Professors, the New York, the Illinois, and [Vor. 38 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN the St. Louis academies of science, and the honorary societies of Phi Beta Kappa, Sigma Xi, and Phi Sigma; he was a corresponding member of the Academy of Natural Sciences of Philadelphia, an honorary chairman of the University of Penn- sylvania Bicentennial Committee, a fellow of the American Association for the Advancement of Science (chairman of Section G in 1936-37), and president of the American Society of Plant Taxonomists for 1938. His social clubs included Sigma Alpha Epsilon fraternity, the University and the Harvard clubs of St. Louis, and the Faculty Club of Washington University. He was a Democrat and a nominal Presbyterian. Although socially gifted and inclined, Dr. Greenman's herbarium, his students, and his Senecios more and more circumscribed his life. Except for short vacations during the lifetime of Mrs. Greenman and for a collecting expedition of three months to Central America with his younger son, Milton T. Greenman, in 1922, all his waking hours seemed to be spent in the big brick building on Tower Grove Avenue. Apparently never a strong man, a series of severe illnesses beginning in 1925 took their toll of him. In 1936 Mrs. Greenman died, leaving him without the care with which she had attended him; but his steely determination carried him on. In 1945 Dr. Greenman was the victim of a cerebral hemorrhage. Upon his partial recovery he was appointed professor emeritus by the University, and in 1948 as curator emeritus by the Garden. Now that it was too late, he was given leave to return to his Senecios as he had planned in his youth. Walking daily from his home on Magnolia Avenue to the Garden, he pursued his remaining joy. For one who had suffered for so long, the end was a gentle withdrawa The fame of some men rests not so much upon what they have done of them- selves as upon what they have done through others. So it is with Jesse More Green- man. It may be said with little dispute that his impress upon American taxonomy, through his students, is second to none in America. The reverence and affection in which they will always hold him is still reflected in the salutation of the Festschrift! with which they greeted him upon the seventieth anniversary of his irt December 27, 1937. Dear Dr. We, dcs peu have gathered er to greet you and to celebrate your academic festival. To commemorate the day, s of us have brought to Ee feast table these con- tributions. тр and of themselves, e are not of great significance. They are impressive because of [: е evidence they offer of the degree to which your — с has con i uits of i ч i er of "noa ago, one of us, in talking with your own teacher, Pu jo referred то to himself as ап academic grandchild, much to En gler's interest. and amusement, Engler's generation is passed now, and you have become the dean. Your own academ қаға ан аге beginning to carry forward into male generation the essence 4. дез 4. ship which you have given us. We are deeply Vot 5 н ай з yours, for with it we have found n to And s 1t emulate you. ow, 2 the full fruition of your years labor, не іп ~ herbarium vh cho Souls бот and will have created. Мау these rice eak of our appreci our respect, and our cni for they Redi vividly to us that ЕСІ” псе and unassa ‘lable kindliness which guided u prwee Mo. Bot. Gard. 25:1—453. 1938. 1951] WOODSON—JESSE MORE GREENMAN 99 CHRONOLOGICAL LIST OF PUBLICATIONS OF JESSE MORE GREENMAN dicit B. L., and J. eenman. dew SW bes Vcn known plants ке їп Mexico by C. с. Ping i in es summer . Am. Acad. 29:382-394, 189 ———, ---. the Flora of the езе uiis as piis by the collection " Dr. G. Baur. Am. Jour. s Tai 135-149. 1895 -------:->--- and noteworthy plants chiefly from Oaxaca collected by Messrs. C. G. Pringle, ШС Smith E . W. Nelson. Ibid. 150—168. 18 ------; ————. ptic revision of the genus Lamourowsia Ibid. 169—174. 1895. ,——.. MEO us new species. Ibid. 76. -----,------ А new genus of Sterculiaceae, idi some other noteworthy plants. Bot. Gaz. 22:168-170. 1896. — ————, ------ Revision of the genus Tridax. Proc. Am. Acad. 32:3-10. 6. ——‚ ------. Synopsis of the Mexican and Central American species of the genus Mikania. Ibid. 10—13. 1896. --------, ————. А revision of the genus Zinnia. Ibid. 14-20. 1896. — — ——, ------ Revision of the Mexican and Central jac species of the genus Calea. Ibid. 20—30. 1896. ------ -----. A шерә key to the species of Porophyllum ranging north of the Isthmus of Panama. Ibid. 31-33. 18 ----- ————. iu E new or little known phanerogams, chiefly from Oaxaca. Ibid. 34—51. 1896. Greenman, J. M. Revision of the Mexican and Central American species of Houstonia. Ibid. 283-293. 1897 . Key to the c species E Liabum. lbid. 293-294. 18 Жанн of new and little known plants from Mexico ТЫЧ. 897. — vision the Мес and nt American species of Galum pé амы. Ibid. 33: 4555170. 189 Diagnoses d new and critical Mexican rn ыа A rm 0. 898 . Some new and ie pepe plants of the Northwest. . Gaz. 25: 261-269. 1898. ——.. Revision of the genera Montanoa, Perymenium, ee ae Ee Am. Acad. 34:507- 534. 1899. Synopsis of the genus Verbesina, with an analytical key to the species. Ibid. 534—566. 1899 Some new сы кү ranges, and newly noted identities among the Mexican БААС О Аз: Ibid. . Revision of "d e genus Gym ӨКҮН Proc. Boston Soc. Nat. Hist. 29:87—104. 1899. Supplementary notes upon Calea, Tridax, and "Mikania. Ibid. 105-108. 1899. New species and varieti es ge Mexican plants. Proc. Am. 2 35:307-315. 1900. The Re Senecio in New кас Rhodora 3:3—7. 190 Monographie rii nord- und кіна ы Arten a Gattung Senecio. I. Allgemeines у Morphologie. ышы Dissertation. 39 pp. Wilhelm Engelmann: Dp 19 А new western Camassia. Bot. Gaz. 34:307-30 Senecio Robinsonianus Greenm. In Sargent's т ind Shrubs 1:19—20, pl. ro. 1902. Кока Greenm. Bot 5:214. 190 -------. New and otherwise я Mic Eb. Mexico and Central America. Proc. Am. Acad. 39: 69-120. 19 -----. On the n me Urbinella and Altamirania. Contr. Gray Herb. Harv. Univ. N.S. 1. No. 25. Suppl. L 1 -------. Notes on Кеней Ке, Ma xican plants. Bot. Gaz. 37:219—222. 1904. T B. L., and J. M. Greenman. Revision of the genus Sabazia. Proc. Am. Acad. 40:3—6. ————, . Revision of the Mexican and Central American species of Trixis. Ibid. 6—14. 1904. c, Revision of the Mexican and Central American species of Hieracium. Ibid. 14—24 Сгеептап, J М ee and synonymy of Mexican and Central American Spermatophytes. Ibid. 28-5 19 -----. De асл 98 Se from the southwestern United States, Mexico, and Cen- tral America. Ibid. 41:235-270. А new ко "Bon Gaz pud 146- 147. 190 . Two new species from northwestern America. bid. 42:146-147. 1906. [Vor. 38 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sene In C. V. Piper’s Flora of the State of Washington. Contr. U. S. Nat. Herb. EET 1:595 2601. "1906. New species of Senecio and Schoenocaulon from Mexico. -------. Ргос. Ат. Асай. 43:19-21. 1907. ------. Studies іп the genus Citharexylum. Field Col. . Bot. Ser. 2:185-190. 19 ------. New or — i. rthy Spermatophytes from т Саша America and the West Indies. DIC 247-287. 19 Notes on E genus Senecio. Rhodora 10:68—69. 1908. n.] L. In G New Manual = шетті ed. 7, рр. 852-855. 1908. . The n name ga Bot. = ‚ 45:19 ay’s New Manual of Botany. [A r isn “Tid ge 153-154. 1909. к, ane undescribed psal from | Ibid. 48:146—148. 1909. Ottawa Nat. 25:114-118. rica. —————. Sir pecie ry Hooker (w =, с" sp of Cuban "scimus Field Col. Mus. Bot. Ser —328. 1912 p new spec E and notes on other Doxcastépbetón. ыу from Mexico and "mm са Ibid. 329—350. 1912. ——————. Taxonomy and 7 botany In the American Year Book 1913:685—687. 1914. Greenman, J. M., and C Thom ere rr chiefly from the south- estern United Stat e Mexico. Ann. Mo. Bot. = 1:405—418, pls. 24-26. 1914 бы J. M. Deina of North A merican жоу bid 263—290, pls. 10—14 1914. -- raph of the North and Central American species of the genus Senecio—Part II. Ibid. 2:573-626, pls. 17-20. 1915; 3:85- 194, pls. 3-5. 1916; 4:15-36, үп 4. 1917; 5:37- 108 E + 4-6. 1918. . Senecio. In L. H. Bailey's The а C Two exotic Composites in Nor HK = ес 6: УМ 3153. 1917. - rica. 289-292, pl. 10. 1917. Greenman, J к and Norma E. Pfeiffer. A new Selaginella from Mexico. lbid. 5:205—210, pls 11-12. temen; J. м. new arnt е the West Indies. ac 8:97—102, Без 1-2. 1921. St «cd y South Am n Senecios—I. Ibid. 10:73—110, pls. 4; -------. Opportunities for boten research in Ded America. n TII. State Acad. Sci. 16:76-81. ІІ. ge-and-Area hypothesis hp special reference to the Flora of Tropical America. FE Toar. ЕЎ 12:1 жыды 1 та ------- cio L. In Sen M e Standley’ s on and Shrubs of Mexico. Contrib. U. S. Nat. Herb. 231 1621- 1636. —. Botany. In the American Year Book 1927:651-655. Greenman, J. M., and E. M. F. Roush. New Agaves from нонии United States. Ann. Mo. t I] [^] ы - с. T о va ос o | - © N confusus, a novel house plant. Mo. Bot. Gard. Bull. 23: Я 1935 Noteworthy collections in the herbarium of the Missouri Botanical Garden. Mo. Acad. Sct, Proc, 3:65. 1937 Stu ies of South American m Ann. Mo. Bot. M 25:795—822. 1938. — xonomy as a field for research. Science, n.s. 87:265—266. 1938. —. Senecio schizotrichus, S. esp or neri and S. Тобы In Standley's Flora Costa Rica. Field Mu t. Hist. Bot. Ser. 18:1518—1519. 1 Genera from the чара of morpho logy. Bull. wey Bot. Club 67:371—374. 1940. the Missouri eres al Garden in relation to our knowledge & Succak ent Soc. America 13:125-1 27. 194 —. Studi pl ros an ntral снаа species е Bar yp a Ceiba 119—1 0. In addition to above public cations, num reviews abstracts of gus literature were published in Ad Botanical Gazette," vols. 47-61. 1909-1916, and in “Botanical Abst vols. 1-15. 1918-1926 tracts,” CHROMOSOME NUMBERS OF CALIFORNIAN DELPHINIUMS AND THEIR GEOGRAPHICAL OCCURRENCE! HARLAN LEWIS*, CARL EPLING*, GUSTAV А. L. MEHLQUISTt AND С. С. WYCKOFF} The genus Delphinium is represented іп California by about twenty-six species. Several of horticultural interest have been studied by Mehlquist; others have been studied extensively with respect to their interfertility and evolutionary relation- ships by Lewis and Epling. In the course of these investigations most of the California species of Del phinium have been examined cytologically and a summary of our combined observations on chromosome numbers (Table I) and cytological behavior is presented here. The materials used were either squash preparations of buds fixed in 1:3 acetic alcohol and stained in aceto-orcein or sectioned root tips fixed in CRAF and stained in safranin-gentian violet. These buds and roots were collected in the field or were taken from wild transplants grown in the garden. CYTOLOGICAL OBSERVATIONS The basic haploid chromosome number in the genus Delphinium is 8 (Darling- ton & Janaki-Ammal, 1945). We have found no deviations from this basic num- ber although three of the species examined, D. variegatum, D. banseni, end D. gypsopbilum, have tetraploid races (Table I). Тһе karyotypes of Delphinium species are morphologically very similar (Langlet, 1927; Tjebbes, 1927; Lewitsky, 1931; Lawrence, 1936; Gregory, 1941). Each genome has two long chromosomes, four of intermediate length and two quite short ones. The centromeres of the long chromosomes are submedian; those of the remainder, subterminal. The rela- tive lengths at mitotic metaphase of each chromosome and the positions of their centromeres are shown diagramatically in fig. 1A. This diagram is based upon measurements from root tips of D. banseni, D. parishii, D. besperium, D. varie- gatum, D. parryi, D. gypsophilum, and D. recurvatum. Chromosomes a and b are readily distinguishable during mitosis or meiosis by the relative lengths of their two arms. Chromosomes c, d, and f are difficult to distinguish from each other in mitotic divisions and are indistinguishable in meiosis. Chromosome e can usually be recognized in somatic divisions by the presence of a small satellite at the end nearest the centromere but usually can not be told from c, d, and f during meiosis. The two short chromosomes can usually be recognized by their small size but can- not always be distinguished from each other. * University of 2-5 Los Angeles. Do Botanical Garden. Formerly University of California, Los Angeles. s paper represents a report of two largely separate studies, one i поа ар 5 d other n pos juis d Wycko for Mehlquist alone or Mehlquist and others. Lewis and Epling are responsible for the remainder of the cytological data and all systematic determinations. (101) 102 [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN The tetraploid races of D. variegatum, D. hanseni and D. gypsophilum have indistinguishable karyotypes and differ from the diploids only in the double number. Meiosis has been studied in pollen mother cells of 45 diploid transplants and 36 tetraploids as follows: Diploids pm (1) S ррррррру > » > variegatum Tetraploids D. gypsophilum (4) D. hanseni (22) D. variegatum (10) TABLE I Number | Number chromosome of о number populations plants D. andersoni Gray 16 3 8 D. californicum T. & С 16 2 10 * D. cardinale Hook. 16 6 26 D. decorum Ее & Mey. ssp. Tracyi Ewan 16 2 8 D. glaucu 16 1 6 D. grac не жайы. tum Greene 16 1 1 D. gypsophilum ins 16 16 { 156 32 19 | 222 D. banseni Greene 16 66 437 32 116 1143 D. Резфетінт Gra 16 11 30 D. hesperium f. della cens Ew 16 14 71 D. hesperium var. cuyamacae T ТИНИ Jeps. 16 1 9 D. inopinum 16 3 6 TD. nudicaule 'Т 16 7 26 D. nuttallianum Pritz 16 1 5 D. parisbii же. 16 25 D. parryi Gra 16 32 104 D. parryi ssp. A editiosum (Jeps.) Ewan 16 11 21 D. parryi var. blochmanae (Greene) Jeps. 16 4 24 D. patens Benth. 16 11 29 D. polycladon Fastw. 16 1 1 D. purpusii Brandg. 16 3 9 "a rvatu e 16 3 17 D. trolliifolium Gray 16 5 10 D. uliginosum Curran 16 1 10 D. umbraculoru 16 8 22 D. variegatum T. & G. 16 (22 (99 32 16 | 128 Tot. 301 "Тос. 1603 * Previously reported by Mehlquist et al. (1943). 1). {Previously reported by Gregory (194 1951] EWIS, EPLING, MEHLQUIST, WYCKOFF——DELPHINIUMS e. © 2, % 4 ы: С ---. == Fo “ уз К ” M A \ NS \ M —? Бы - --- ` 27 айры \ б Ры , / , М , NS ГА 2 122 br ATE. Fig. 12 ‚ diagram of the relative lengths of mitotic metapha ase chromosomes representative of a species com plex ferred to in the text. В, represe meiotic metaphase I of a diploid (D. parisbii). C, anaphase I of a diploid showing a bridge and fragment (f). The bridge involves 1 ] i tetraploid showing delayed separation of (D. gypsobbilum). se I of a tetraploid showing 1 E, apha and 1 foli edes chain (D. variegatum). chromosomes a or b bivalents, 1 quadriv Psi ring, [Vor. 88 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chiasma frequencies observed at diplotene are nearly constant for all the species examined and agree with those previously reported for D. cardinale (Mehlquist et al, 1943). The longest chromosome, a, has a chiasma frequency close to 1.0 for each arm. However, a chiasma in only one arm is sometimes found and a second chiasma may occasionally occur in one or both arms. Моге than two chiasmata per arm have not been observed. Chromosome b has a chiasma fre- quency close to 1.0 in the longer arm; one is usually present in the shorter arm but its precise frequency has not been determined. Chromosomes c, d, e, f, g and b generally have but one chiasma per bivalent, which is in the long arm, although an additional chiasma may rarely occur in the short arm. Eight bivalents occur regularly in the diploids at metaphase I, but quadrivalents are frequent in the tetraploids. In 418 tetraploid cells, 183 (44 per cent) had 1 quadrivalent, 144 (34 per cent) had two quadrivalents, and no cells had more than two. The quadrivalents were confined to the two long chromosomes a and b, the former being quadrivalent in 70 per cent of the cells examined and the latter in 44.5 per cent. Of the possible quadrivalent types (Darlington, 1937) only a simple ring of four or a chain of four has been found (fig. 1E). Disjunction of the quadrivalents at anaphase may result in either adjacent or alternate chromosome separation (fig. 1E). However, configurations indicating alternate separation are about twice as numerous as those indicating adjacent sep- aration. Either type of separation produces nearly 100 per cent visibly good pollen and the plants concerned are fully fertile. Trivalents have rarely been observed between any of the chromosomes. The shortest chromosomes frequently separate precociously and can often be seen as univalents at diakinesis, indicating either that no chiasmata were formed or that terminalization and separation occurred during prophase. Segregation of these chromosomes is usually equal, although it is occasionally 3 and 1 in the tetra- ploids. Diploid gametes carrying an extra short chromosome can function, for at least one plant grown in the greenhouse had an extra b chromosome. Unequal segregation of this pair in diploids apparently results in non-functional gametes. Terminalization of chiasmata is usually complete in the six shorter pairs at metaphase I but is often incomplete in one or both arms of chromosomes a and b (fig. 1B). Sometimes separation of these chromosomes is greatly delayed at ana- phase I in both diploids and tetraploids (fig. 1D). In this case the unseparated ends of the chromosomes remain on the equator until after the centromeres have reached their respective poles. The chromosomes are stretched thin and are ob- viously under tension, but there is no evidence in subsequent stages that breakage or abnormal separation occurs. It has been shown on a number of occasions that terminalization of chiasmata is arrested by changes in homology (Sansome, 1932; Catcheside, 1932; Darlington, 1937) and, further, that such an arrest need not lead to breakage of chromatids (Darlington, 1931). Stretched chromosomes showing delayed separation have recently been found by Marta Walters (1950) in the hybrid Bromus trinii X B. maritimus where she has suggested that the delayed 1951] LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS 105 separations may be due to changes in homology of the chromosomes. In the present instance the anaphase stretching may result from the formation of chias- mata in inversion heterozygotes, but proximal to the inversions. This is further suggested by the fact that the delayed separations are often observed in the same plants that show inversion bridges. Pachytene has been difficult to study in our preparations. Our observations on the occurrence of inversions is accordingly based upon the anaphase. Bearing this in mind, it would seem that most of the inversion bridges apparently involve the longer chromosomes, but it has not been possible to determine either the num- ber of inversions or the particular chromosomes in which they occur. Anaphase bridges characteristic of those produced by crossing over within an inversion were found in 10 (22 per cent) of the diploids examined and 25 (70 per cent) of the tetraploids. In many instances the accompanying fragment was clearly visible (fig. 1С); in others it may have been obscured by other chromo- somes. It is probable that inversions were present in more plants than is indicated inasmuch as bridges in plants where they did occur were seen in only 0.8-1.7 per cent of the anaphase and telophase cells examined. The possibility of inversions passing unnoticed if a limited number of cells can be examined is, therefore, very great. In view of the high frequency of inversion heterozygotes found in them the tetraploids may carry still other inversions than those of their corresponding diploids. However, even if the diploids and tetraploids do not differ in the par- ticular inversions which they carry, the chances of a tetraploid with 4 homologous chromosomes being heterozygous for any given inversion is materially increased. More than one bridge has not been observed in the same cell, which might sug- gest that but a single inversion is involved in all of the plants. However, the frequency of bridges is so low that the chances of observing two bridges in a single cell from crossing over in two inversions would be very small indeed. All of the species in which inversion bridges have been found are interfertile to some extent, and several of them grow in close proximity to one another or even together. The inversion heterozygotes referred to may therefore be the result of interspecific hybridization. GEOGRAPHICAL DISTRIBUTION The geographical distribution of the colonies in which chromosome counts were made, including their precise locations, are indicated in the following list, together with label data which will identify the vouchers on file in the herbarium of the University of California, Los Angeles. The number of plants at each locality of which the chromosome number was determined is shown in parentheses. In some cases two collections from the same colony are combined. The species concerned are listed alphabetically. [Vor. 38 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN DELPHINIUM ANDERSONI Gray (2n — 16) LASSEN CO.: ы id ipt n. of Ravendale, Meblquist & Jenkins 380 (3). MODOC со. inder-slope of Sconchin Butte, Lava Beds Nat'l. Mon., Meblquist 83 мемы pi Ts n. е of Caldwell Butte, Lava Beds Nat'l. Mon., Aoc rhe еј жұты (2). DELPHINIUM CALIFORNICUM Т. & С. (2n = 16) CONTRA COSTA CO.: near Juniper Camp, Mt. Diablo, Meblquist & Compton 305 (5). MONTEREY CO.: near Aromas, Meblquist 300 (5). DELPHINIUM CARDINALE Hook. (2n — 16) LOS ANGELES CO.: Sepulveda Canyon, Santa Monica Mts., Mehlquist & Compton 301 (6); San Antonio Wash near Claremont, Mehlquist & Compton 206 (5 SAN BERNARDINO CO.: 2 mi. e. of Upland on Highland Ave., жет 306 (11). SAN DIEGO CO.: near — Е & Geissman 404 (4). SANTA BARBARA CO.: c, Meblquist 20 (5); 3 ті. s. of Lompoc оп the Santa Rosa Rd., Mehlquist 205 y os янв ae) (5). DELPHINIUM DECORUM Fisch. & Mey. subsp. rRAcYr Ewan (2n = 16) BOLDT CO.: 2 mi. s. of Kneeland on the rd. to Bridgeville, Meblquist & Jenkins 368 | (4); South Fork of Van Duzen R., Meblquist & Jenkins 374 (4). DELPHINIUM GLAUCUM Wats. (2n = 16) SAN BERNARDINO CO.: Dobb's Cabin, San Bernardino Mts., Meblquist & Compton 302 (6) DELPHINIUM GRACILENTUM Greene (2n — 16) BUTTE CO.: 2.5 mi. n. of Little Butte Creek bridge on the Paradise to De Sabla Rd., May 25, 1941, Lewis & Epling (1). DELPHINIUM GYPSOPHILUM Ewan, Diploid (2n = 16) (Fig. 2) RN CO.: 1 mi. s. w. of Poso e on the Bakersfield to Woody Rd., April 14, 1944, Lewis & Epling (15); т. S. Hwy. 466, 4.2 mi. n. of the w. rd. to. Caliente, April 22, 1943, Lewis & Epling (15); 1.5 mi. n. of McKittrick on Hwy. 33, Meblquist 411, April 23, 1943, Lewis & Epling (19); 5.0 mi. s. w. of Maricopa on U. S. Hwy. 399, Lewis & Epling 531, Epling & Miles 545 (47). MONTEREY CO.: 2.4 mi. w. of the jtn. at Pleyto, Lewis & Epling 697 (1); 3.0 mi. w. of the jtn. at сом I 9 Ерін E 699 (1); 4 mi. w. of the jtn. at Pleyto, Lewis © Epling 698 (8); . е. of King City on the rd. to Hollister, ten & Meblquist 431 (1); Bradley to Jalon M. at jtn. arad rd. to Pleyto, Lewis & Epling 694, 782 (9). SAN BENITO CO.: 0.8 mi. w. ds Bitterwater, Lewis & Epling 820 (1). SAN LUIS OBISPO C hr ев of U. S. Hwy. 101 on rd. to Cambria WS снр Lewis & Epling 064 (4); of Klau, Lewis & oet pes ч (13 )› mi. w. of Hwy. 41 on the Hillman а to сені, rd., Epling 735 (6); of U фта . 466 оп the rd. to Simmler, Lewis 9 Epling 715 (3); near Adelaida, Lk e oral 4% (9). DELPHINIUM GYPSOPHILUM Ewan, T'efraploid (2n = 32) (Fig: 2) SNO С 5.5 mi. n. of Coalinga on Hwy. 33, Lewis & Meblquist 451 (3); 7.2 mi. n. of Кайны є on Hwy. 33, Lewis & Epling pis Lewis & Mehlquist 455 (9); 13.6 mi. n. of Coalinga on ат 33, Mehlquist 414, Lewis 9 Meblquist 454 (5); 14.9 mi. n. of Coalinga on Hw . Lewis & Epling 530 (38); 5.2 mi. w. of Coalinga оп Hwy. 198, Lewis & Meblquist 2 (48). 1951] LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS 107 po TETRAPLQ Eth Univenerry се caciroamia Pi MM CALIFORNIA— NEVADA Fig. 2. Distribution of D. gypsophilum showing stations from which plants have been Phe cytologically. KERN co.: about 2 mi. e. of McKittrick, Meblquist 8 Wyckoff 332 (4); 8.7 mi. w. of McKittrick, Apr. 23, 1943, Lewis 9 Epling (25); Cottonwood Pass on Hwy. 41, 0.6 mi. e. of the San Luis Obispo Co. line, Apr. 24, 1943, Lewis & Epling (13); 0.4 mi. e. of Caliente R. R. Station, Lewis 9 Epling 403 (5); 1 mi. w. of Caliente R. К. Station, Lewis 8 Epling 404, Epling 585 (20); 4.2 mi. w. of Caliente К. К. Station, Apr. 22, 1943, Lewis & Epling (6); U. S. Hwy. 466, 0.4 mi. n. of the w. rd. to Caliente, Apr. 22, 1943, Lewis & Epling (6); U. S. Hwy. 466, 0.9 mi. n. of the w. rd. to Caliente, Apr. 22, 1943, Lewis 6 Epling (4); U. 5 Hwy. 466, 1.0 mi. n. of the w. rd. to Ca pu Lewis 9 Epling 529 (10); U. S. Hwy. 466, 2.9 mi. n. of the w. rd. to сане Арг. 22, 1943, Lewis © [Vor. 38 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN Epling (3); U. S. Hwy. 466, 5.0 mi. n. of the w. rd. to Caliente, Apr. 22, 1943, Lewis & Epling (12); U. S. Hwy. 466, 4.2 mi. n. of the w. rd. to Caliente, Lewis, Epling 9 Mehl- quist 458 (9). KINGS CO.: hills e. of Avenal, Meblquist & Blodgett 331 (6). SAN JOAQUIN CO.: 1 mi. e. of Midway, Lewis 9 Meblquist 452, Epling & Miles 543 (4). SAN LUIS OBISPO CO.: 2.8 mi. e. of the jtn. of U. S. Hwy. 466 and Hwy. 41 near Cholame, Apr. 24, 1943, Lewis & Epling (31); U. S. Hwy. 466, 1.4 mi. e. of jtn. of Hwy. 41 near Cholame, Lewis & Epling 713 (9); 6.1 mi. s. of U. S. Hwy. 466 on the rd. to Simmler, Lewis & Epling 715 (1); 5.1 mi. n. е. of Creston on the rd. to Shandon, Lewis 816 (1); 5.2 mi. s. w. of U. S. Hwy. 41 on the Shandon to Creston rd., Lewis & Epling 21 (1). DELPHINIUM HANSENI Greene, Diploid (2n = 16) (Fig. 3) AMADOR CO.: 2.6 mi. n. of Jackson, May 20, 1945, Epling (1); 3.5 mi. s. of Jackson on Hwy. 49, Epling 584 (3); 3.7 mi. s. of Jackson on Hwy. 49, May 20, 1945, Epling (6). BUTTE СО.: 2.5 mi. e. of Oroville city limits, Epling 666 (1). CONTRA COSTA СО.: Marsh Creek Rd., 5.8 mi. e. of the Curry Creek Park Rd., Lewis & Epling 703 (3). ELDORADO CO.: 0.5 mi. w. of Garden Valley, May 26, 1941, Lewis & Epling (1). KERN CO.: mi. e. of Hwy. 65 on the rd. to Woody, Epling 560 (1); 1.9 mi. sw. of Woody, Meblquist 338 (1); 3.1 mi. s. w. of Woody, Мау 18, 1945, Epling (9); 3.6 mi. s.w. of Woody, Lewis 9 Epling 528 (9); 5.3 mi. s. w. of Woody, May 18, 1945, Epling (7); 5.5 mi. s. w. of Woody, May 18, 1945, Epling (5); 3.7 mi. s. of White River, Epling 561 (6); Glennville Church, Meblquist & Wyckoff 327 (3); 1.2 mi. e. of Glennville, Lewis, Epling 8 Meblquist 460 (5); 2 mi. e. of Onyx, Lewis & Epling 484 (16); 3.7 mi. e. of mouth of Kern River Canyon, Meblquist 9 Wyckoff 323 (3); about 5 mi. e. of the mouth 30, 1946, Lewis & Epling (1); 3.6 mi. s. of Bodfish, Lewis & Epling 485 (16); 10.1 mi. s. of Bodfish, Lewis & Epling 487 (16); 12.5 mi. s. of Bodfish, May 24, 1943, Lewis, Epling 9 Mehlquist (13) ; 1.3 mi. e. of the cemetery, Walker Basin, Lewis & Epling 488 (30); Oiler Canyon Rd., 3.5 mi. s. of the jtn. at the s. w. margin of Walker Basin, Lewis 9 Epling 480 (14); Oiler Canyon Rd., 6.5 mi. s. of the jtn. at the s. w. margin of Walker Basin, Lewis & Epling 400 (3); Oiler Canyon Rd., 8.5 mi. s. of the jtn. at the s. w. margin of Walker Basin, Lewis & Epling 402 (20); 1 mi. w. of Caliente R. R. Station, Apr. 22, 1943, Lewis & Epling (1); 3.5 mi. w. of Caliente R. R. Station, Lewis & Epling 405 (19); 1.8 mi. e. 5. Hwy. 466 on the w. rd. to Caliente, Meblquist ё Lewis 437, Lewis, Epling & Meblquist 457 (8); U. S. Hwy. 466, 0.4 mi. n. of the w. rd. to Caliente, Apr. 22, 1943, Lewis & Epling (1); U. S. Hwy. 466, 0.6 mi. n. of the w. rd. to Caliente, Lewis 9 Epling 406 (7); U. S. Hwy. 466, 1.5 mi. n. of the w. rd. to Caliente, Epling 550, Apr. i. n. of t i Apr. 22, 1943, Lewis & Epling (4); U. S. Hwy. 466, 3.8 mi. n. of the w. rd. to Caliente, Apr. 22, 1945, Lewis & Epling (4); U. S. Hwy. 466, 4.2 mi. n. of the w. rd. to Caliente, Lewis, Epling 9 Meblquist 450, 461 (21); U. S. Hwy. 466, 4.1 mi. e. of the w. rd. to Caliente, Lewis 6 Epling 500 (13). DERA CO.: Madera to Raymond Rd., 1.1 mi. e. of the jtn. with the Chowchilla Rd., Epling 564 (4) ; Raymond to Coarsegold Rd., 4.9 mi. e. of the jtn. to Knowles, Epling 568 (1); 1.0 mi. s. of Raymond, Epling 567 (2); 1.4 mi. s. of Raymond, Epling 566 (3); 1.6 mi. s. of Raymond, Epling 565 (2) MARIPOSA CO.: 6.8 mi. s. of Mariposa, Meblquist 346 (4); Hwy. 140, 15 mi. e. of the rd. to LeGrand, Epling 570 (1); 8.1 mi. s. of Coulterville, Meblquist 347 (3); Merced River, 1.3 mi. s. of El Portal, Lewis 761 (2). 1951] LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS 109 Pies tee ads us arora "des озын 7 TETRAPLOID о | UnivinerT or CALIFORNIA ранае CALIFORNIA—NEVADA BERKELEY AMD LOS ANGELES Distribution of D. hanseni showing stations from which plants have been examined m E LARE co.: Kern River near Durrwood, Lewis, Epling & Meblquist 509 (5); Kern River at Welch (near Fairview), May 24, 1943, Lewis, Epling 9 Meblquist (2); 1.9 mi. e. of Lemon Cove, May 18, 1945, Epling (2) ; 4.2 mi. e. of e Cove, Epling 563 (5); 0.3 mi. n. of White River, Lewis, s 9 Meblquist 470 (1); 0.7 mi. n. of White River, Lewis, Ерін 9 Meblquist 472 (2); 1.0 mi. n. of White River, Meblquist © Wyckoff 329 (2); about 1.5 mi. n. of White River, Meblquist 8 Wyckoff 330 (3); 1.9 mi. n. of W River, May 18, 1945, Epling (4); 2.6 mi. п. of White River, Lewis, Epling © Койын 473 (1). [Vor. 38 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN TUOLUMNE CO.: 4.8 mi. e. of Mather Ranger Station, Epling 770 (6); 5.9 mi. e. of Mather Ranger Station, Epling 771 (4); Jacksonville, 0.6 mi. w. of the jtn. of Hwy. 49 and Hwy. 120, Epling 579 YOLO CO.: 3.8 mi. w. of Winters, Meblquist & Baker 314, Lewis & Epling боб (5). YUBA CO.: 3.4 mi. e. of Oregon House P. O., Lewis 764 (3). DELPHINIUM HANSENI Greene, Tetraploid (2n = 32) (Fig. 3) AMADOR CO.: 2.6 mi. n. of Jackson, May 20, 1945, Epling (4). CALAVERAS CO.: 1.6 mi. n. of Copperopolis Ranger Station on ый 4, Epling 582 (4); 4.5 mi. n. of Hwy. 120 on the rd. to Copperopolis, С 581 KERN CO.: U. S. Hwy. 466, 4.2 mi. n. of the w. rd. to Caliente, Meblquist & Lewis 425, 426, 427, 460 (81). MADERA CO.: e ping to Madera Rd., 0.6 mi. s. of Raymond Rd., Epling 569 9; ; Madera to Raymond Rd., 1.1 mi. e. of jtn. n. with Chowchilla Rd., Epling 564 (2); n. of the viaduct over the 5 River at Madera, Epling 9 Mile es 544 (1); 4.7 mi. n. pr Madera, Mehlquist 345A (5); 5 n. of Madera, Lewis & Meblquist 438 (3); 10 mi. n. of Madera, Lewis & Meblquist jon 423 (4); 13 mi. n. of Madera, Meblquist 345B (3). TULARE CO.: 1.6 mi. n.e. of Richgrove, Lewis, Epling & Meblquist 475, Epling & Miles 540 (1); 0.7 mi. n. of White River, Meblquist © Wyckoff 328, Lewis, Epling 9 Meblquist 471 (20); 2.6 mi. n. of White River, Lewis, ст, © ipei 474 (4). TUOLUMNE со.: Jacksonville, 0.6 mi. w. of the of Hwy. 49 and Hwy. 120, Epling 578 (4); 0.5 mi. n. w. of Jamestown on the сек қ лай Rd., Ad 723 (1). DELPHINIUM HESPERIUM Gray (2n = 16) (Fig. 4) o.: 5 mi. e. of the Lake Co. line on the Lakeport to Hopland Rd., Lewis, Epling 5 Stebbins 246 (4). NDOCINO СО.: 9.6 mi. s. of Rock Creek, Mehlquist & Jenkins 366 (2). co.: 6 mi. s. of Monticello on the rd. to Napa, Meblquist 355 ҚУА 6.8 mi. s. of “Жылы on the xk to Napa, Meblquist & Paddock 308, Meblquist 356 (4); 7 mi. s. of Monticello on the rd. to Napa, Mehlquist 357 (2); 8.4 mi. s. of Monticello on the rd. to Napa, Meblquist 358 (2); Wooden Valley, Meblquist 400 (1). SAN MATEO CO.: Page Mill Rd., near Stanford Univ., Lewis 9 Epling 436 (3). SANTA CLARA CO.: Uvas Creek, Lewis © Epling 435 (2). SONOMA CO.: 1 mi. e. of Bodega Bay, Lewis & Epling 515 (2); Mark West Springs, Meblquist & Jenkins 362 (5). DELPHINIUM HESPERIUM Gray forma PALLESCENS Ewan (2n = 16) (Fig. 4) This entity, described by Ewan (1945) as D. hesperinm sen pallescens, is most con- spicuously distinguished from the typical form by its white or pinkish flowers and is a readily recognized geographical race of the drier more etes, foothills of the Coast Ranges (fig. 4). In our opinion it я warrants subspecific status. The com- bination required will be made in another publica CONTRA COSTA CO.: Marsh Creek Rd., 1 mi. w. eee Curry Creek Park Rd., Lewis 9 Epling 702 (1); rd. to Livermore, 0.1 mi. s. of the Marsh Creek Rd., Lewis % Meblquist 44 (3); Beck Rd. (to Livermore), 1.0 mi. s. of the Marsh Creek Rd., Lewis & Epling 580 (22); Beck Rd. (to "uu E Ж s. of the Marsh Creek Rd., Мау 6, 1944, Lewis & Epling (5); rd. to Antioch, n. of the Marsh Creek Rd., Lewis 5 мен: quist 446 Ou Antioch Golf Course, Pede © ры ГЫ 445 (5). LAKE mi. n. of Clear Lake on rd. to Stubbs, Meblguist & Jenkins 364 (1). NAPA CO.: 2 mi. n. of Knoxville on the rd. to e Meblquist 9 Paddock 300, Mehl- p & Baker 319 (5); 4.6 mi. s. of Monticello on Napa Rd. ‚ Meblquist 354 (3). N BE iw CO.: 0.5 mi. s. Lo Paicines, Lewis & Meblquist 450 (4); 2.8 mi. s. of the en he Pinnacles, Lewis & Meblquist 449 (3); 4.1 mi. s. of Pinnacles P. 25 Meblquist 346 (6); 9.9 mi. s. of Paicines, Lewis 9 Epling 700 (4). 1951] WYCKOFF—DELPHINIUMS 111 LEWIS, EPLING, MEHLQUIST, аа Жек E sera prs m | D. HESPERIUM | UM | | | | Е ШЕ: С (UYAMACAE © | | | | | | | | | | | | | "EI NIE CALIFORNIA — NEVADA Fig. 4. Distribution of D. besperium showing stations from which plants have been examined cytologically. yolo co.: Putah Creek Canyon, 8 mi. w. of Winters, Meblquist © Baker 315 (4). DELPHINIUM HESPERIUM Gray var. CUYAMACAE (Abrams) Jeps. (2n — 16) (Fig. 4) SAN DIEGO CO.: Cuyamaca Lake, Mehlquist & Geissman 403, Ensign & McClintock 554 (9). [Vor. 38 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN DELPHINIUM INOPINUM (2n — 16) have evidence to indicate that D. parisbii Ж inopinum Jeps. represents a good species. The new combination required will be made elsewhere VENTURA СО.: 1.0 mi. w. of С Sa E Station, Ball 807, Lewis 630 (3); 13.2 mi. s. of Ozena оп U. S. Hwy. 399, Lewis 9 Epling 755 (1); 13.6 mi. s. of Ozena on U. S. Hwy. 399, Lewis & Epling 756 (2) DELPHINIUM NUDICAULE T. & С. (2n = 16) HUMBOLDT со.: S. Fork Van Duzen Rd., Meblquist & Jenkins 375 (3). MODOC CO.: s. slope of Caldwell Butte, Lave Beds Nat'l. Mon., Meblquist & Jenkins 379 (5). NAPA CO.: 12 mi. n. of Monticello on rd. to Knoxville, Meh! Ys 6 Baker 318 (3); 11 mi. n. of Monticello on rd. to Knoxville, Meblquist 350 (3); 16.5 mi. s. of Monticello on rd. to Napa, Mehlquist 359 (4); Nuns Canyon, Mehlquist 9 ‘Beker 321, Mehl quist & Jenkins 361 (5). SANTA CRUZ CO.: 0.9 mi. n. of Brookdale Resort, Santa Cruz Mts., Meblquist 333 (3). DELPHINIUM NUTTALLIANUM Pritz. (2n = 16) NEVADA CO.: Soda Springs Hotel, Meblquist & Jenkins 381 (5). DELPHINIUM PARISHII Gray (2n = 16) (Fig. 5) INYO CO.: Westgaard Pass, Bohart 410 (1); Canyon near Hells Gate, Death Valley Nat'l. Mon., Lewis 9 Lowe 682 (3); Emigrant Canyon, Death Valley Nat’l. Mon., Lewis KERN CO.: rad to Cuddy Valley, 0.1 mi. n. of the у Valley Rd., Lewis & Dunn 478 (6); 1.0 mi. w. of Chuchupate Ranger Station, Lewis 740 (1); 1 mi. e. of the summit о er Pass, Lewis, Epling & Meblquist 467 (2); ET 27% alker Pass, Lewis, Epling 5 Меш 466 (2); 7.1 mi. w. of the summit of Walker Pass, Lewis, Epling 9 Meblquist 468 (1 » GELES CO.: Palmdale, Epling 745 (2); 23.5 mi. e. of Palmdale, Apr. 17, 1941, Lewis 8 "Ebling (1); near Lancaster, Meblquist (5). RIVERSIDE CO.: Palm Springs, mouth of Tahquitz Canyon, Lewis STI (1). SAN BERNARDINO CO.: 4.9 mi. n. of the jtn. of the Mojave River and Deep Creek, Apr. 25, 1941, Lewis & Epling (1); Morongo Valley, Apr. 27, 1941, Lewis & Epling (1); grade between Morongo Valley and Yucca Valley, Meblquist. 333, Lewis © Epling 508 (15); 4.8 mi. s. of Barstow on the rd. to Stoddard Well, Apr. 27, 1941, Lewis & Epling (1); 8.2 mi. s. of Barstow on the rd. to Stoddard Well, Apr. 27, 1941, Lewis & Epling (1); the Mojave River Narrows at Hy Mae Apr. 27, 1941, Lewis & Epling (1); = Baldwin Lake, Meh/quist 421, 422, 424 (6); Holcomb Valley, Meblquist 420 (1); watz Mts., C. kid ( Meblquist mi я ). SAN DIEGO С 1.2 mi. w. of bridge in Sentenac Canyon, ra apr © н 407 (2); 2.4 mi. w. РЄ; bridge, Meblquist & Geissman 408 (2); mouth of Coyote Canyon, Meblquist & Geissman 400 (2); mouth of Borego Palm Canyon, Mehlquist 3 Geissman DELPHINIUM PARRYI Gray (2n = 16) (Fig. 5) INYO CO.: Sherman Hills above Paradise Camp, n. of Bishop, Bohart 410 (2). LOS ANGELES CO.: Santa Catalina Island, west end of Whitley Ave., Avalon, Lewis 631 (1); Mulholland Drive e. of Topanga Canyon Rd., Meblquist 9 Wyckoff 301, 302 (5); Lopez Canyon, Mehlquist 312, 40016 (9). MONTEREY CO.: 1.0 mi. w. of the jtn. at Pleyto, Lewis & Epling 605 (4); 1.5 mi. w. of the jtn. at Pleyto, Lewis & Epling боб (2); Bradley to Jolon Rd., at jtn. with Pleyto Rd., vaa Aida (1). RIVE CO.: бап ummit, Lewis & Epling 482 (1); 3 mi. s. of Banning on the rd. to > Idyllwild, 5. ks (3). 1951] LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS 113 <А M ж ex \ дуй" onera ebrii CALIFORNIA— NEVADA кактау AMD LOS Anar Fig. 5. Distribution of D. parryi and D. parisbii showing stations from which plants have been examined cytologically. SAN BERNARDINO CO.: Cedar Springs, Lewis 9 Epling 534 (1); 5 mi. = of Verdemont, Apr. 1944, Lewis (1); near the Zanja, Mentone, Apr. 20, 1946, Lew SAN DIEGO CO.: Lakeside Park, Froelich 313 (4); апо bor 307 (4); near Fallbrook, Like ed 9 Eggers 393 (2 OBISPO CO.: 1.9 mi. w. Dr Pozo, May 24, 1941, Lewis 9 D (1); 6.1 mi. of U: wy. 466 on rd. to Simmler, Es 9 Ebling 714 (2); 7.1 mi. w. of Simmler, Lewis С “Ebling 501 (7); 4.1 mi. s. and 1.1 mi. e. t Atascadero, De & Brown 2383 mi. s. of Klau, Lewis 6 Epling 662 (3); 8.5 mi. n.w. of Adelaida Cemetery, eae " Epling 650 (2); 9.3 mi. n. ж. of Adelaida Hei diam Lewis © Epling 658 (3); 9.9 [Vor. 38 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN mi. w. of U. S. Hwy. 101 on the n. rd. to Adelaida, Lewis & Epling 687 (2); 3.5 mi. e. of Santa Margarita on the rd. to 5 Lewis 9 Epling 653 (2); Santa pon coms Mebl- quist 380 (5); rd to Creston, 4.2 т of Santa M to Pozo Rd., Ball 802 (3). SANTA BARBARA СО.: Davy Amby Саш Figueroa Mt., Epling 477 (12); 0.8 mi. s. of Davy Brown Camp, Figueroa Mt., Lewis 720 (7); 1.1 mi. n. of Red Rock Mine on the "X us wierd to Davy Brown Rd., Lewis 722 (3); ар хэр. Meblquist 390, 481 (3). co.: Ojai, 1942, Мей (1); U. S. Hwy. 399, 9 mi. s. of jtn. with Mt. Pinos "Rd, (ames © Ebling 754 (6). DELPHINIUM PARRYI ssp. SEDITIOSUM (Jeps.) Ewan (2n — E (Fig. 5) FRESN 6 mi. w. of Coalinga, Lewis & Meblquist 434 (2); 7.6 mi. w. of jtn. of the Los E Creek Rd. and the rd. to Coalinga, Lewis & Epling 628 RT 10.9 mi. w. of jtn. of the Los Gatos Creek Rd. jy E rd. to Coalinga, Lewis © Epling 624 (1). ; МеА- pe 385 (2); 6.7 mi. n. of Pinnacles P. O., Meblquist бс (3); 5.1 mi. s. of Pinnscles О.; Meblquist 367 (2). SANTA CLA 2“ со.: Livermore-Mt. Hamilton Rd., 0.9 mi. n. of Del Puerto Rd. t Patterson, Lewis T MAE 610 (2); depu e ной Rd., 1.8 mi. n. x Del Puerto d. to Patterson, Lewis & Epling боо (1); Livermore-Mt. Hanita Rd., 1.3 mi. s. of Del Puerto Rd. to nct гучы m иы; 615 (1). w \o B 3 т о шад] к 5 „В i) p. о ж- "2 ‚© - 5-6 A ОЕ. ч жЕ" бо ‚л, + w N \о 3, 5 © A vo 5 5 к ” г) La - и”, ‚О DELPHINIUM PARRYI Gray var. BLOCHMANAE (Greene) Jeps. (2n = 16) (Fig. 5) SAN LUIS OBISPO CO.: 5.5 mi. n. of Guadalupe, Meblquist 513 (3); dunes to the w. of the Nipomo to Guadalupe Rd., Lewis & Epling 504 (9); 5.3 mi. n. of Veterans Memorial in Guadalupe, Lewis & Epling 685 (4); U. S. Hwy. 101, 11.1 mi. n. of Los Alamos, Lewis 6 Epling 505 (8). DELPHINIUM PATENS Benth. (2n = 16) O.: 6.5 mi. s. of Walker ren on hen Oiler Canyon Rd., Lewis & Epling 491 (1); Kern River — about 5 m of mouth, Meblquist & Wyckoff 326 (3 LOS ANGELES CO.: Pacoim vy cingi, San Gabriel Mts., Meblquist & Froelich 306 (5); near Mt. Wilson Toll Rd., Meblquist 398 (2 MONTEREY СО.: near Priest Valley, Meblquist 208 (2); near Aromas, Meblquist 200 (2). NAPA CO.: 11 mi. n. of Monticello, on the rd. to Knoxville, Meblquist 316A (1). RIVERSIDE CO.: about 15 mi. s. of Banning on the Idyllwild Rd., NE 399 (3). SAN DIEGO CO.: Banner Grade, Mehlquist & Geissman 405 (3); 406 (2). SANTA BARBARA CO.: San Marcos Pass Rd., 5 mi. n. of San Marcos Ranch, Mehlquist 390 (3). VENTURA CO.: Mt. Pinos Public Camp, Lewis & Dunn 480 (1). DELPHINIUM POLYCLADON Eastw. (2n = 16) MARIPOSA CO.: e. end of Lake Teneya, Aug. 22, 1941, Lewis (1). DELPHINIUM PuURPUSII Brandg. (2n = 16) KERN CO.: about 2: of the mouth of the Kern River Canyon, Meblquist * Wyckoff 324 (3); 5.0 mi. e. en the mouth, Lewis, Epling & Meblquist 464 (1); 3.7 mi. of the mouth, Mus oor (59; DELPHINIUM RECURVATUM Greene (2n = 16) CONTRA COSTA CO.: 0.5 mi. w. of Hwy. 120 on the Byron Hot Springs Rd., Epling & Miles 542 (3). FRESNO CO.: 7.2 . of Coalinga on Hwy. 33, Meblquist 412 (3); 13.6 mi. n. of 1). n Coalinga on Hwy. 33, pue 413, Lewis & Meblquist 453 (1 1951] LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS 115 DELPHINIUM TROLLIIFOLIUM Gray (2n — 16) UMBOLDT CO.: 1.5 mi. n. of Kneeland, Meblquist & Jenkins 367 (3); Yager Creek, Meblquist 6 Jenkins 370 (3); Yager Creek bridge, 8 mi. n. Bi Bridgeville, он аг % Jenkins 371 (2); 16.2 mi. п. of Bridgeville, Meblquist & Jenkins 372 (1); 1.1 mi. w. of South Fork Van Duzen Rd. on rd. to Dinsmore, Meblquist & Jenkins 373 "n DELPHINIUM ULIGINOSUM Curran (2n = 16) LAKE СО.: 2.7 mi. e. of Middletown on rd. to Lower Lake, Meblquist 9 Baker 320, Meblquist & Jenkins 363 (10). DELPHINIUM UMBRACULORUM (2n — 16) Delphinium rici id ili is an undescribed species. А formal taxonomic description will be presented elsewher SAN LUIS OBISPO CO.: 1324. 0 mi. s. of Klau, Lewis & Epling 663, бот (3); King City to Jolon Rd., 1.2 mi. n. of Avila, Lewis & Epling 701, 781 (1); Santa Margarita—Pozo Rd., 10.6 mi. e. of rd. to Creston, Lewis 813 (1). SANTA BARBARA CO.: San Rafael Mts., Davy Brown Camp pec Lewis 717 SE 0.8 mi. n. of Davy Brown Camp Ground, Lewis & Mathias 701 (1); mi. n. of D Brown Camp Ground, Lewis 718, Lewis 6 Mathias 792 (3); about 5 mi. w. of Las Cruces- Lompoc Rd. on the rd. to Jalama, Meblquist 512 (2); 0.3 mi. w. of Las Cruces-Lompoc Rd. on rd. to Jalama, Lewis 767 (8). DELPHINIUM VARIEGATUM T. & G., Diploid (2n = 16) (Fig. 6) ALAMEDA CO.: 10.7 mi. s. of Livermore P. O. on the Tesla-Corral Hollow Rd., Lewis & Epling 608 (15). CALAVERAS CO.: 7.8 mi. f Mokelumne Hill, Epling 583 (4); 4.5 mi. n. of Hwy. 120 on the rd. to CS s "Еріне 580 D 2) CONTRA COSTA СО.: rd. to Livermore, 0. . of the Marsh Creek Rd., Lewis © ie 443 (6); 1.0 mi. s. of the Marsh pec Rd. on the Beck Rd. to елан; Lewis & Epling 589 (5). E Lake-Napa County line, rd. Knoxville to Reiff, Mehlquist & Paddock 3II (5). MARIPOSA CO.: 3.8 mi. n. of Bagby, Epling 572 (3); Hwy. 140, 16 mi. e. of the rd. to Le san Ерін. pa (5): M 5.2 mi. s. of ыы. оп U. S. Hwy. 101, Meblquist 9 Jenkins 365 (2). NAPA CO.: 2144 n. of Monticello, Meblquist © Baker 318, Meblquist 351 Cre i. n. of Knoxville, Meblquist " "Paddock 309 (5); 6.8 mi. n. of Monticello on коле Meblauist 9 Baker 316 (3); 0.7 mi. s. of Nuns Canyon, Mehlquist 9 as $ 60 (3). SAN LUIS OBISPO CO.: 2.9 mi. w. of Pozo, Lewis & Epling 502 (12); 7.8 mi. w. of Pozo, Apr. 24, 1943, Lewis & Epling (10); near Santa Margarita, Lewis & Epling 503 (6). SANTA CLARA CO.: Mt. Hamilton, Paddock 402 (3); уе to Mt. Hamilton Rd., 0.2 mi. w. of Mines Rd., Lewis & Epling 611 (1); Livermore to Mt. Hamilton Rd. at jtn 3 (2). TULARE CO.: 2552 Valley (near So Lewis € Epling 526 (1). TUOLUMNE CO.: 5.5 mi. e. of Hangman's Tree, Bret Harte, Epling 730 (1); Hwy. 108 at jtn. with rd. to a areas Epling pe (3). PLACER CO.: Weimar, Mehlquist & Jenkins 382 (5). DELPHINIUM VARIEGATUM T. & G., Tetraploid (2n = 32) (Fig. 6) ELDORADO CO.: 0.5 mi. w. of Garden Valley, Lewis & Epling 442 (2). [Vor. 38 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN e үе be z г = Т RENE. ы ae таз | | D. VARIEGATUM | | | | DIPLOID . | з. Жс) | (PE E ЕНЕ o! | \ iia | | | | | | | | | | T-——L cj | | | T id — UnivanatTy OF Cai iP OR ala Pena CALIFORNIA — NEVADA 6. Distribution of D. variegatum showing stations from which plants have been ex- dud cytologically. A CO.: 0.5 mi. n. of the n. end of the bridge over the Fresno R. at Madera, Epling & Miles 541 (2); 10 mi. n. of Madera, Lewis & Meblquist 433 (3); 21.8 mi. s. of Merced, Lewis & 4. 447 (7). MERCED со.: Pacheco E Meblquist 416, 417 (9). TUOLUMNE CO.: Hwy. 108, 0.7 mi. e. of the rd. to Soulsbyville, Epling 575 (5). 517 19511 LEWIS, EPLING, MEHLQUIST, WYCKOFF—DELPHINIUMS LITERATURE CITED Cytologia 4:68-1 The chromosomes of a new haploid Oenothera. i Jour. саш Catcheside, D. С. (1932). Darlington, C. D. (1931). The cytological theory of inheritance in Oeno thera. 0 4. 112 Recent advances in R 2nd ed. 671 pp. Philadelphia-London E. K. Janaki-Ammal (1945). Chromosome atlas of cultivated ho. 397 pp. A synopsis of the North American species of Delpbinium. Univ. Colo. Stud., 27 Series D, 2:55-24 Am. Phil. oy baa a and cytological studies in the Ranunculaceae Juss. 3- Padi zur Zytologie der Ranunculazeen. Svensk Bot. нра 21:1-17. Lawrence, №. A C. (1936). The origin of new forms in Delphinium. Genetica 18:10 А u systematics. (English summary) pull. yes Bot. Lewitsky, G (1931). The “karyotype RAE Mehlquist, G. A. О. Blodgett, and L. Bruscia (1943). Colchicine induced tetraploidy in Del- phinium ups Jour. d 7-1 Sansome, E. R. (1932). Segmental interchange n Pisum sativum. Cytologia 3:200—219. Tjebbes, K. (1927). vo apr aid E thre Walters, Marta S. (1950). Spontaneous brea Bromus trinii X B. бака. Бене i а ee De finm 522 Hereditas 10:160-16 Кав ved reunion of meiotic AE i in the hybrid STUDIES IN THE APOCYNACEAE. УШ! AN INTERIM REVISION OF THE GENUS ASPIDOSPERMA МАКТ. & ZU CC. ROBERT E. WOODSON, JR. Contemporary systematic botany in South America has attained to such a high degree of proficiency and eminence that a word of apology, at least, becomes the outland botanist who undertakes a taxonomic revision of an almost wholly South American genus. I am able to make excuse with a whole heart for attempting a revision of the apocynaceous genus Aspidosperma, for I had truly hoped never to feel called upon to do so. There are numerous South American botanists, par- ticularly in Brazil, who could present a much more detailed picture of the variation and system of the genus than I. The species of Aspidosperma are very widespread and frequent trees in tropical America; and since I am regarded in certain quarters as something of an authority on the Apocynaceae, many specimens of the genus have been submitted to me for naming over a period of years. From the very first I was vexed by the lack of decisive criteria in the literature and baffled by discordant specimens filed under the same name in our herbaria. I soon became aware that I, myself, was contrib- uting to the confusion in both respects. The need for a revision was all too apparent. The publication of this study of the genus, and at this time, has a rather fortuitous history. One day in the year 1938, I was writing to Dr. K. H. Rechin- ger, of Vienna, in connection with a loan of Apocynaceae which was being returned to him. After telling him how greatly I had profited from his kindness, I re- marked wistfully that "someday" I hoped to request the loan of his Aspidospermas which I knew would be rich in the older types required by a monographic study. Out of the goodness of his heart, Dr. Rechinger must have misunderstood my in- tentions, for, to my consternation, all of the Aspidospermas in the Naturhistorisches Museum descended upon me by return mail! Since I was already engrossed in another major project at the time and needed encouragement for an additional task, I wrote for counsel from my old friend, Dr. Friedrich Markgraf. 'То my relief, he replied that Dr. J. G. Kuhlmann, of Rio de Janeiro, was already engaged in a study of Aspidosperma: professional ethics appeared to have furnished me an alibi, and I could return the Aspidospermas to Vienna with a clear conscience! I there- upon relinquished the loan to my herbarium superior for the required "official transmittal." Within three months central Europe was in turmoil; but for me, fresh horror was added when I learned that my loan had not been forwarded, as I had trusted, and could not then be sent with safety. Unstudied, yet, the Viennese Aspido- spermas reposed first in neutral and later in "enemy" internment for nearly ten years! lTssued June 18, 1951. (119) [Vor. 38 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN АП this time unnamed Aspidospermas continued to arrive, and I continued to make my haphazard way amongst them with increasing misgivings. Finally I wrote to Dr. Kuhlmann in desperation for news of the long-awaited revision, but without reply. After a second and a third unanswered inquiry, my impulse toward revising Aspidosperma, chiefly for my own use, could be repressed no longer, and the Vienna loan with its precious types was unpacked again and put to tardy use; at length being returned to Dr. Rechinger in 1949. Later, Dr. Kuhlmann re- sponded by forwarding to me generous portions from the types of several of his species of Aspidosperma. These specimens, not represented in any North American or European herbarium, have been invaluable to me. Aspidosperma has been in need of a revision for many years. As a matter of fact, not even a complete presentation of the literature alone has been prepared since the publication of A. de Candolle's treatment for the ‘Prodromus in 1844. Probably well over 100 species have been described since that time. Jean Mueller's account for ‘Flora Brasiliensis’ in 1860 is roughly in synoptic form and deals with 46 names applied to the Brazilian species; but today species of Aspidosperma are known from every country of continental Latin America except El Salvador and Chile, as well as from Hispaniola and Trinidad. Since Mueller's time, studies of three small groups of species have been pub- lished by Markgraf (in Notizbl. Bot. Gart. Berlin 12:553. 1935; 13:464. 1937). Pichon's perspective of the genus (in Bull. Mus. Nat. d'Hist. Nat. ser. 2, 19:362. 1947) provides a detailed analysis of a dozen more or less natural sections with enumerations of representative species for each, but without keys to the species and without synonymy; however, it is such a major contribution toward alleviation of the chaos that the need for my own study is greatly reduced. Finally, even with this present study which has occupied so much of my attention for several months, tropical American dendrology still requires the field observations of Dr. Kuhl- mann, and the appearance of his investigations of Aspidosperma will be awaited by no one more eagerly than me. GENERAL MORPHOLOGY Habit.—The species of Aspidosperma are trees from 2 to 60 m. in height, growing in a variety of habitats from the dry campos of south-central Brazil, Paraguay, and Argentina, to the inundated river margins of the Amazon valley, and at elevations from a few meters above sea level to approximately 2000 m. in eastern Peru and Bolivia. The largest and most distinctive trees are found in my series NrTIDA, where heights of 40 m. are common, frequently with boles over 1 mm. in diameter b. h. The boles of this series, also, are characterized within the genus by very striking buttresses or lamellations which are frequently noted by collectors of botanical specimens. Thickness and texture of the bark vary considerably and apparently without phylogenetic significance except in the series NoniILES, where, with the exception of Aspidosperma decussatum, obvious lenticels in the leafy twigs are absent. Probably similar environmental conditions have 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 121 resulted in very thick, corky periderm and disproportionally shortened internodes in A. verbascifolium and A. tomentosum of the series MACROCARPA and PYRICOLLA, respectively. Both are species of the Brazilian and Paraguayan campos. Trees of Aspidosperma may be deciduous, as in series MACROCARPA and Рүкт- COLLA, or apparently evergreen, as in the remaining series. It is obviously difficult to ascertain vernation habits of the species without extensive field experience; my own observations are based only upon a limited number of herbarium specimens. Nevertheless, it is obvious that species of Macrocarpa bloom frequently upon leafless twigs or together with early leaves only. Twigs of these species have more or less obvious seasonal articulations; although the rounded terminal buds are ap- parently naked, the terminal internodes are conspicuously foreshortened—protec- tion possibly being furnished in this way in lieu of bud scales. In series PyRICOLLA, however, not only are the terminal internodes shortened, but the terminal buds are protected by strongly imbricate scales. In this series, consequently, the young twigs bear conspicuous seasonal articulations. Terminal buds of series RAMIFLORA and PoLyNEuRA are essentially similar to those of Macrocarpa (although those of the latter apparently produce scales at times): in all four series, the bud is obviously terminal, with the subtending internode conspicuously thicker than the terminal leaf petiole. In series NITIDA, STEGOMERIA, and Nonirzs, however, the buds are naked and consist of a few leaf initials as the terminal internodes are not sensibly foreshort- ened. Since in these series the internode subtending the terminal bud is only of about the same thickness as that of the terminal leaf petiole, the terminal bud appears pushed to one side in a pseudo-lateral position recalling that of many species of fig, and this aspect is of aid in the casual assignment of specimens to their series. Twigs of the QuEBRACHINES are notable within the genus in producing a char- acteristic bushy growth through the use of abundant axillary buds with closely imbricate scales. The twigs, however, are not conspicuously articulated. Latex bears interesting correlation with floral and fruit characters in Aspido- sperma: in series NOBILES the latex apparently usually is reddish, although in some parts of the plant, as in the stem, it may be white although red in the inflorescence and fruit for example. Traces of the red latex may frequently be found upon herbarium specimens. In species of PoLvNEURA the latex appears generally to be colorless or opalescent and more scanty than in the other series, to judge from the notes of collectors. In the remaining series the latex is milky. Тһе latex of no species seems to have suggested economic use, and I am not aware of any special study of it. The wood of very many Aspidospermas appears to be tough and strong, and an important local source of structural timber. Other woods or bark are used as sources of tannin, or even as a substitute for quinine, due to the astringent crystal- line inclusions. A special study of the wood anatomy of 29 species of Aspidosperma was published by Milanez (in Physis 15:428. 1939) and will be discussed more fully in succeeding paragraphs. It may be noted here, nevertheless, that Milanez [Vor. 38 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN generally found "annual rings" in the wood of species with scaley buds, or in those with naked buds but with congested upper internodes and blooming before or with production of young leaves: "annual rings" were absent or inconspicuous in other species. А general account of the woods of Aspidosperma, including their economic uses, will be found in Record and Mell's "Timbers of Tropical America', рр. 507—515 (Yale University Press, 1924). Leaves.—The leaves of Aspidosperma frequently aid in the assignment of speci- mens to series, although less often to species. Phyllotaxy is alternate or approxi- mate typically, except іп A. decussatum of the Nosies, which has broad decussate foliage, and in the two species of series QUEBRACHINES in which the small spine- tipped leaves are opposite or ternate apparently rather indiscriminately. Extremely useful in the separation of many species of NrrIDA are the peculiar leaves with margins more or less conspicuously auriculate-revolute at the base: many species of the same series are without this character, however. Leaves of Nirma, QuE- BRACHINES, and Мовп.вв habitually are of coriaceous texture and those of the other series membranaceous. Leaf venation always is pennate: In NoniLEs the secondary veins are nearly perpendicular to the midrib and appear to continue without dichotomy to the margin. In PoLvNEUmA and Мітірл the secondary veins are more arcuate and dichotomize to form a rather definite marginal vein close to the leaf margin; in the other series, particularly in PyRICoLLA, the broadly arcuate secondaries dicho- tomize and anastomose to form the marginal vein farther from the leaf margin. My series PoLvNEURA coincides essentially with Pichon’s section RETICULATA, which bears an appropriate name recalling the very dense and conspicuous tertiary reticulation of the habitually rather thin, pale foliage. Inflorescence.—Inflorescence structure іп Aspidosperma would furnish a sep- arate study of considerable morphological interest because of its wide range of modification and close correlation with other morphological criteria used in the delimitation of series and species. Although the structure is basically cymose throughout, it shows definite tendencies toward indeterminate composition, par- ticularly the thyrse. In series МАСКОСАКРА and 5ТЕСОМЕЮІА the inflorescence probably shows its most simple, if not primitive structure. Here it is a terminal or, more usually, a subterminal cyme of relatively few flowers, without obvious bracts and lacking the determinate flower, at least in the lower dichotomies. The primary peduncles very frequently are attached to the stem some distance above the "subtending" leaf which bears an axillary, vegetative bud: this prompts the interpretation of the inflorescence as morphologically a determinate axis as is shown more clearly in the interpetiolar cymes of Asclepiadaceae. Although in Macrocarpa and ЅТЕСОМЕВІА, which are without well-defined seasonal growth, the lateral inflorescences are spaced at moderate distances, in species of PYRICOLLA, where flowering occurs at or near the greatly condensed terminal nodes at the beginning of the season, the inflorescences (also dichotomous 1951] WOODSON—STUDIES IN THE APOCYNACEAE. УШ 123 cymes) become congested into a corymbiform mass. In one species of the latter series, namely A. pyrifolium, it is helpful to observe that the determinate flowers occur at every dichotomy to the base; this quite possibly is the original design for the "empty" dichotomies produced by other species of the genus. In the closely related A. multiflorum, determinate flowers may occur at the upper dichotomies although absent at the lower. Although inflorescences of the genus habitually are terminal ог subterminal upon the younger branches, in the series RAMIFLORA and STEGOMERIA the lateral inflorescences are produced much farther below the apical meristem, very fre- quently upon completely suberized stems, and are described as "cauliflorous" in this study. In the series Nitipa, the rather numerous species may have either of two types of inflorescence: laterally subterminal and dichotomously cymbiform, as іп PYRI- COLLA, but with more or less obvious and irregularly placed but persistent bracts; or terminal and thyrsiform, also with similar bracts. The latter type, although appearing thyrsiform, actually is of the type of cyme which has been called the “aggregate dichasium” (cf. Woodson, in Ann. Missouri Bot. Gard, 22:1. 1935). Since the ultimate terminal and lateral branches of the thyrsiform type cor- respond closely to the cymiform structure of the first type, the two appear to be correlated. However, whether the thyrsiform type represents a condensation of a shoot apex bearing several cymiform branches, with resultant reduction of the foliage, or whether the more simple type is a simplification of the thyrsiform, is difficult to decide. I am somewhat of the opinion that the first view is more plausible. A rather similar situation exists in the four species of series POLYNEURA, with A. dispermum and A. cylindrocarpon representing the cymiform subterminal type, A. cuspa the thyrsiform terminal type, and A. polyneuron occupying an intermediate position. In series Noses the case in favor of inflorescence condensation may be fol- lowed somewhat more clearly. In what I believe to be the most "primitive" type of inflorescence, shown by that of A. verruculosum, the flowering axis is terminal and bears few or several alternate, rather distantly and uniformly spaced flowering branches with congested determinate divisions. In the remaining species of the series, however, the inflorescence is of a very characteristic type: terminal or sub- terminal with a more or less elongate naked primary peduncle, at the tip of which are borne the flowering peduncles in a congested more or less umbelliform false whorl. 'The degree of compounding and congestion of these varies even upon a single individual, and it is very easy to visualize the primitive condition as being that similar to A. verruculosum. Неге in NoBILES one may observe the more or less obvious and irregularly placed bracts, such as those of NITIDA, very conspicuous in some species and quite obscure in others. A type of inflorescence which appears to have been reduced from that of NoniLEs is found in the QUEBRACHINES. Неге the inflorescences are produced very abundantly in an axillary position as a rule. The axes are reduced to rela- | Vor. 38 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN tively few flowers each so that the whole appears racemiform in fact; but it is interesting to observe that the branches are opposite or ternate, as are the foliage leaves. Неге, although bracts are produced, they are small and caducous. Having already written more fully concerning the inflorescence of Aspido- sperma than my comprehension of it warrants, perhaps, I can end only in stating that while I believe there to be evidence to interpret the thyrsiform aggregate dichasium as derived from the bracteate, cymiform structures in such series as NITIDA, POLYNEURA, and QuEBRACHINES, the phylogenetic relationship of these to the lateral, ebracteate cymes of PyricoLLa, МАСКОСАКРА, and STEGOMERIA certainly is less than clear. Flowers.—The calyx of Aspidos perma typically consists of five imbricate, equal or subequal segments which are free to the receptacle. They always are destitute of internal glandular squamellae. In the more advanced series, however, there is a pronounced tendency for the outermost pair of calyx lobes to be larger than the inner three. This tendency is particularly strong in certain species of series NITIDA, as in A. discolor, in which the innermost segment may be entirely suppressed. The climax is reached in the species of SrEGOMERIA, in which the two outer lobes become connivent for about half their length (except in A. illustre) and com- pletely enclose the two inner lobes which are much smaller and are free. This very unusual calyx prompted me some years ago to erect a distinct genus, Cufodontia, for the single species then known (C. stegomeris Woods.). Discovery of the fruit of Cufodontia and appreciation of the tendency to zygomorphy of the calyx in many Aspidospermas renders the genus untenable in my opinion. The corolla of Aspidosperma is of two general types: salverform with reflexed limb and tube constricted at the orifice as in series MACROCARPA, RAMIFLORA, and PynicoLLa, and tubular or tubular-salverform with erect or laxly ascending limb respectively, and unconstricted tube, as in the remaining series save опе. The most specialized of the latter type are found in series Noses. Here, with the excep- tion of A. Fendleri, the lobes are abruptly caudate-acuminate and nearly filiform from near the base; they are very tightly contorted spirally in the bud, spreading horizontally as they unroll, and become erect at full anthesis. An intermediate type is found in A. rigidum, which comprises the series Ricipa, in which the corolla lobes are reflexed, although the throat is not constricted. A peculiar characteristic of the corolla which has been ignored generally is the series of longitudinal clefts or fissures which occur behind the stamens. These clefts originate as external invaginations of the corolla tube at the place of attach- ment of the epipetalous staminal filaments, eventually passing around the base of the filaments, as is shown diagrammatically in fig. 1; they may be reclosed almost immediately, or may continue some distance toward the corolla throat. Photo- micrographs to illustrate the clefts in representative series are provided in pl. 1; these were prepared from microtome sections obtained from herbarium specimens, and tissue distortion caused by desiccation is apparent. Although I have found no verbal reference to the clefts, their use in dissections is shown by the very neat 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. Vti 125 drawings prepared both by Dr. Kuhlmann and by Sr. David de Azambuja, both of the Jardim Botanico of Rio de Janeiro. The corolline clefts, in fact, seem to provide a rather precise morphological criterion to separate Aspidosperma from other closely related genera, for I have found them in all species of that genus, and quite absent in such relatives as Microplumeria, Geissospermum, and Diplorbyncbus. A photomicrograph illus- trating staminal insertion in Diplorhynchus is included in pl. 1. What the function of the corolline clefts of Aspidosperma may be is quite obscure to me: if the corollas were only larger, they might possibly be construed as "short cuts" for a pollinating proboscis, or even to drain excess water; but their very small size would appear to preclude these. Fig. l. Anatomy of the corolla in Aspidosperma: diagrams from serial microtome sections showing origin of the fissures. Explanation in the text. Instead of functional innovations, it seems more likely that the epistaminal slits of Aspidosperma actually may be an atavism. In his study of the development of the perianth in Vinca [Lochnera] rosea, Boke’ found that the upper corolla tube (i.e. above the insertion of the stamens) is formed by the ontogenetic union of the bases of the "petals", with the united epidermal layers remaining distinct as late as the organization of the staminal archesporium although lost at maturity. Subtle differences of texture of the corolla of Aspidosperma have been quite noticeable to me, but I have found them difficult to convey to others and have omitted them from my keys. In the species of MAcrocarpa and RAMIFLORA the texture appears rather thickly membranaceous, and more delicately so in PyRICOLLA and STEGOMERIA; dried corollas of these series imbibe water very freely and section in paraffin very readily, as the photomicrographs of pl. 1 show. In Nonis, NITIDA, QUEBRACHINES, and most POLYNEURA, on the other hand, the texture appears to be subcoriaceous or at least pergamentaceous; corollas of NrrIDA contain many cells packed with tannin and crystalline inclusions, and those of NortLrs are heavily suberized: this renders them very much more difficult to section, as the photomicrographs also testify. 7 тВоке, N. H. Amer. Jour. Bot. 35:413. 1948. [Vor. 38 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN The stamens of Aspidosperma consist of two bilocular thecae which are wholly fertile to the base and free from the stigmatic clavuncle, as in most other Plumeroideae. The outer locule of either theca is slightly longer than the inner, under the base of which it tends to incurve. This tendency varies amongst the species and I have tried to use it as a supporting character for the series, but with- out success. The pollen is granular. The stamens of the genus may be uniform amongst themselves or more or less dissimilar in size of anther and length and relative height of insertion within the corolla tube. This tendency is rather more marked and variable in the NiTwa than in the other series. Here, too, the tendency does not appear to be of sufficient uniformity to use as a "key character" even for the NirIDA. Botanists who follow the literature of Apocynaceae will notice that my descrip- tion of the thecae of Aspidosperma coincides with that of Pichon (in Mem. Mus. Nat. Hist. Nat. n. s. 27:183, 196. 1948) for the African genus Diplorbyncbus: Pichon, in fact, distinguishes Aspidosperma and Diplorbyncbus upon the characters > of "anthéres à sacs polliniques égaux . . .” and "anthéres à sacs polliniques extérieurs ‚ in part, respectively. However, it will become ap- ээ depassant les interieurs . . . parent to any one carefully examining a series of dissections that the pollen sacs of Aspidosperma are far from equal, and accord with Pichon's description for Diplorbyncbus. Although I have specimens of only one species of Diplorhynchus available for study, namely D. condylocarpon (Muell Arg.) M. Pichon, I have found the staminal filaments to bear a small laminate appendage at their insertion to the corolla tube, these resembling the filament appendages of my subgenus Pferantbera of the South American genus Forsteronia which are borne higher on the filament. Pichon does not mention these appendages, if indeed they occur in the other species of Diplorhynchus, but perhaps his description of the filaments as keeled (“carénés” may be intended to refer to them somewhat obliquely. The pistil always is bicarpellary and the ovaries are free essentially from the receptacle to the common style, or somewhat united at the base. The ovules are peltate and borne upon a binate ventral placenta, the number of rows upon each placenta-half varying from опе to six. In PoLtyNeuRA the number of ovules is characteristically smaller than in the other series, being reduced to two in A. dis permum. The stigmatic clavuncle is relatively simple and small, and from capitate to fusiform in shape (occasionally somewhat umbraculiform) and bluntly or sharply apiculate. It has not appeared sufficiently useful to be employed systematically. It has become habitual to describe the pistil of Aspidosperma as possessing an annular nectary or "discus", and amongst such authors I must include myself (in N. Amer. Flora 29?:119. 1938). It is true that in A. macrocar pon the outer wall of the ovary is irregularly tumid and glandular toward the base, and that this region certainly represents a strongly adnate nectary; but in the remaining species 1951] WOODSON—STUDIES IN THE APOCYNACEAE. УШ 127 even such evidence of a nectary is wholly lacking. It surely is indicative of the prevalent ignorance of the genus that so many authors, including myself, have actually distinguished Aspidosperma from related genera by invoking this essen- tially imaginary structure whose rather variable appearance in one or two species certainly is of phylogenetic rather than of diagnostic importance. Fruit.—The fruit of Aspidosperma is a dry, dehiscent, more or less woody follicle containing few or numerous, large, papery-winged seeds from which the generic name is derived. The follicles are strongly compressed, as a rule, but may be nearly cylindric normally, as in А. cylindrocarpon, or as apparently individual or racial anomalies in such species as A. quebracbo-blanco. The size varies greatly, the smallest (274 cm. long) in A. dispermum and A. cuspa of the POLYNEURA, and the largest (9-15 cm. long) in A. macrocarpon and A. verbascifolium of the MACROCARPA. The shape of the follicle generally is broadly rounded and may be seen to con- sist of several types as expressed roughly in degrees of curvature of the placental margin, which always occupies the greater arc. The shape is simplest in the NoniLEs, where the margins are more or less equally biconvex with resultant ellip- tical or circular configuration: the placenta may be described in this series as rotating about 120°-180°. In the series МітірА and STEGOMERIA the follicle is nearly circular for quite another reason, for the placental margin may rotate as much as 300? and the shape is more accurately defined as very unequally convex- concave or convex-plane and very broadly dolabriform. In all three of these series the seeds are nearly circular with the embryo more or less central. In Macrocarpa and particularly in PyrIcoLLA the follicle is pyriform and unequally biconvex with the placental margin the greater (180^—225^), and grad- ually attenuated to а basal stipe. Although the seeds of Macrocarpa are nearly circular, those of PYRICOLLA are ovate and acute at the base. In POLYMERA (except A. dispermum) the follicles are falciform to clavate-subcylindric and the seeds are very distinctive, bearing a much reduced basal wing quite excentric to the embryo; this tendency is climaxed in A. cuspa, in which the cotyledons are asymmetric as well, with one cordate lobe shorter than the other. In A. cuspa, also, the wings of the seed appear as alternately basal and apical as seen in the opened follicle, although all are chalazal, and therefore basal as determined by the position of the embryo: these fruits correspond almost exactly to those of the African genus Diplorhynchus. Additional useful characters of the fruits of Aspidosperma are found in the relative development or suppression of a basal stipe, the presence or absence of obvious lenticels, quality or absence of indument, etc. Particularly useful in the recognition of species of the NirIDA are the warty or spiny excrescences of the pericarp. [Vor. 38 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN PHYLOGENY OF THE SERIES Pichon’s synopsis of Aspidosperma (in Bull. Mus. Nat. d’Hist. Nat. ser. 2, 19:362. 1947) is based almost entirely on relative dimensions of the corolla tube and lobes, and results in the erection of 12 major divisions which he called "series," but the names of which are constructed as recommended for sections by the Inter- national Rules and which furthermore are preceded by the sign ($) ordinarily reserved to designate sections. I am referring to these categories as sections in this discussion. Two of Pichon's sections consist of species which I am excluding from Aspidosperma; otherwise, the enumeration of his sections are as provided below, each preceded by an alphabetical index for use in our further discussion: (a) "TETRAsTICHA (f) LAEVIFoLIA (b) GLABRIFLORA (д) Міскогова (с) Рилғіока (h) RETICULATA (4) MacroLoBA (i) CRASSITUBA (e) MACRANTHA (j) PUNGENTIA My own major divisions coincide with the sections of Pichon essentially only where Pichon’s are not based upon relative dimensions of the corolla tube and lobe (1. е. in his sections CRASSITUBA, PUNGENTIA, and in part RETICULATA). This is my major reason for adopting an entirely different nomenclature as being less liable to confusion with his. Whether an intrageneric group is to be designated as a section or as a series, of course, is not entirely a matter of convenience how- ever weighty that argument may be; in addition, my groups, although I believe them to be quite natural, lack the distinction which I should prefer sections to have. The tendencies themselves may well lead in time to the definition of fully formed genera in fact, but at the present time they are too fluid. At any rate, my series are as follows. For sake of contrast with Pichon’s sections, I have appended to the name of my series a sequence of letters to stand for Pichon’s placement of species with reference to the enumeration of his series in the pre- ceding paragraph. The actual names of the species would make the list too cumbersome. (1) Macrocarpa—c, с; (6) Мітша--с, e, #,, (2) RAMIFLORA—e (7) STEGOMERIA—e (3) PYRICOLLA—a, b, су, dy, е, f, g,» (86) QUEBRACHINES—j, (4) PorvNruRA—a,, b, d, h,, (9) NomniLEs—i,, (5) Ricma—a, b, с Without attempting a mathematical analysis, the degree of correlation. may be envisaged by means of a scatter diagram (text-fig. 2). For many years I have looked forward to the opportunity of comparing two independent taxonomic studies, such as this. The opportunity is greater yet, since there exists a recent "monograph" of Aspidosperma by Milanez (in Physis 15:429. 1939) which is based upon wood anatomy instead of organography. Milanez was able to examine material of 29 species of Aspidosperma (a thirtieth, A. quadri- 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 129 Pichon jo. B „ 122 фе o ө е " "T? 1 ә о ° Je A e 9 Ф P e 9 o 9 5 e ө 9 1 1 | І И i І І 1 8 5 4 Б 6 " 8 9 Woodson Fig. 2. Correlation of Pichon's sections and Woodson’s series of Aspidosperma. Explanation in the text. ovulatum Pittier, actually being a species of Rauwolfia and is excluded from this discussion). As a result of his investigations Milanez was able to distinguish four "sections" based upon the position of the wood parenchyma: whether terminal, diffuse, metatracheal, or paratracheal, representing what he believed to be a phylo- genetic sequence following Jeffrey. An interesting contrast of Pichon's system and my own is obtained by regression of either upon that of Milanez, whose sections are indicated by Roman numerals consecutively in the appended scatter diagrams (text-fig. 3). MILANEZ’ SYSTEM / PICHON’S MILANEZ’ SysTEM / WoopsoN's Т ites Gy 8) 21 L 1,44 П. ce @ lu I. 25.4, HP fh NL 44.4] IV. es, ig IV. lud [Vor. 38 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN & b c d e f$ ес h 1 j Pichon Milanez 1 12. 1 2 3 4 5 6 7 8 9 Woodson Fig. 3. Comparison of Күнес нн of Aspidosperma with those of Pichon and Woodson, respectively. Explanation in the It will be noticed that Pichon's system and mine agree in one very important detail, and that is lack of agreement with Milanez in the disposition of two of his species in Section IV. These species are cited by him as "А. LeCointei" and A. Duckei. The first is cited in ‘Index Kewensis’ as a typographical error for the second! It appears almost certain that the material which Milanez studied was misdetermined. Since it apparently was a large-flowered species, upon the basis of our very close agreement, I suspect that his trees may actually have been of one of the large-flowered species of my series Nrripa. (VI), which bear a superficial 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 131 resemblance to those of my Macrocarpa (I). I have tried to locate herbarium specimens to accompany Milanez' wood samples, both at the Yale School of Forestry and at the Jardim Botanico of Rio de Janeiro, but without success. In view of their dubious identification, I have drawn a line to separate their scores upon the scatter diagrams. The reticulate nature of phylogeny and the barriers to adequate serial presenta- tion are so familiar to every evolutionary biologist that it scarcely is necessary to point out that perfect correlation of two or more independent systems of any group of organisms is almost impossible of fulfillment. SPECIATION The persistent doubt in the minds of some well-informed persons, even amongst biologists, whether, upon describing a new species, a systematist is describing only a single organism (his type specimen) or a population, never ceases to amaze me. One would suppose it beyond the limits of profitable discussion to question whether, in erecting a species even upon the basis of a single specimen, a systematist visual- izes himself as expressing at least the approximate mode of a population frequency distribution. This should be evident in the invariable expression "species nova”, since a species is a concept of natural populations. Undeniably it happens ali too frequently that a type specimen of an author may be found later not to represent the mode of a distinct natural population but actually, say, a certain quartile or decile (or whatever) of a frequency distribution previously envisaged by another author: his species name thereupon becomes a synonym of the aggregate. This, essentially, is the function of monographic study, i.e. to establish the frequency distributions of natural populations and also to assign the relative position of previous population concepts within those distribu- tions. In 1936 R. A. Fisher (in Ann. Eugenics 7:179. 1936) published his process of "discriminate functions" by which it is possible to demonstrate mathematically the reality of two or more natural populations. There can be little doubt that in defining his systematic categories a systematist operates in a closely analogous man- ner more or less subconsciously, by seeking the ratio of the difference between the means of a compound measurement to its standard error. This, of course, is what we mean by the need of a systematist for experience and judgment: experi- ence in the degrees of variance of the organisms which he studies, and judgment of their values relative to others. I am convinced that the contours of the normal distribution are quite as deeply impressed in the minds of the best of our system- atists as they are amongst statisticians in general. The lot of the systematist, how- ever, deserves the sympathetic commiseration of all professional statisticians because of his usual lack of linear series and the general paucity of his samples. [Vor. 38 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN Adequate samples are as desirable to a systematist as to a mathematical statis- tician. But whilst a mathematician would seldom attempt to find a frequency distribution from a single case, a systematist is called upon to do so quite regularly: he seeks to do this approximately by extrapolation from his experience with other related organisms: systematic biology would as surely be thrown into confusion if he did not do so, as mathematics would be if mathematicians were to follow suit. I have found Du Rietz's definition of taxonomic categories (in Svensk Bot. Tidskr. 24, H. 3:333. 1930) very stimulating and helpful, although certainly idealistic and occasionally difficult to satisfy in many problems. As we have noticed already, an experienced taxonomist usually will be able to distinguish a species from a single specimen. In the case of a subspecies this usually will be impossible, if we are to keep the subspecies in its geographic or ecologic connotation which I feel to be a sine qua non. I personally can see little value іп the concept of variety as a non-geographic or non-ecologic variant: perhaps it might be used as a provisional status where the habitat factor is yet to be demonstrated fully. I have not considered my present data sufficient to warrant the establishment of infra-specific categories in Aspidosperma. Nowadays the imaginations of plant systematists may be somewhat over- stimulated in the detection of hybrids; but in Aspidosperma it appears to me that there are two very obvious instances of it: perhaps the simpler case occurs in northeastern South America. There are three species of the series NopiLes: Aspido- sperma album, A. Spruceanum, and A. Fendleri. The first is the most widespread, ranging from the Magdalena River in Colombia to the lower Amazon in Brazil. It has very distinctive inflorescence structure of a peculiar scorpioid aspect, and nar- rowly stipitate follicles which are almost exactly circular in outline and with a relatively thin, minutely yellowish-pulverulent pericarp; the leaves of specimens to the extreme north and south of its distribution are wholly or essentially glabrous. Both A. Spruceanum and A. Fendleri have corymbiform inflorescences, leaves which are white-tomentulose beneath, and much more woody follicles which are sessile or very shortly stipitate; the former is found in the upper Amazon and Rio Negro basins, and the latter in Venezuela. The two species have several differentia, but the chief is found in the corolla lobes: whether narrowly caudate-acuminate and tightly spiral in the bud (А. Spruceanum), or rather broadly ovate and acute, and scarcely spiral in the bud (A. Fendleri). It so happens that in much of the Guianas, Venezuela, and the middle Amazon, many specimens with inflorescences and pods characteristic of A. album are found with leaves more or less white-tomentulose beneath (such, in fact, are the type specimens not only of A. album but of all its synonyms—the populations which I consider as genetically most pure are without separate nomenclatural designation), or again, with inflorescences midway between scorpioid and corymbiform, or, yet again, with woody, sessile follicles with the indument as for A. album. It seems more than likely that such variation, in the light of the specific characters of the geographically neighboring species, may be taken as evidence, in herbario (!), of 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 133 hybridization. That album is involved is very apparent; whether both Spruceanum and Fendleri are involved as well, or only the former, is not so clear—but all o the putative hybrids collected in flower show the attenuate and tightly rolled corolla lobes, so that I am inclined to think the former the more likely comple- mentary parent if but one is to be selected. No less than ten species names have been proposed in the past to describe indi- vidual variants of the populations of which I speak, and each one could be repre- sented in a key which might be adequate to distinguish the specimens presently at hand but which would cause increasing difficulty as each additional specimen accrued to our collections. This would be natural to expect as the result of random segregation of multiple factors in two or more interbreeding populations. Since these species of Aspidosperma occupy rather distinct geographical ranges, the genetic intergrades may be found to compose a geographical gradient or genocline. In а well-known floristic region, such as the eastern United States, it might be possible, with hundreds or thousands of herbarium specimens, extensive field ob- servation, and by dint of several years of exhaustive study, to establish the contours of such a genocline if such it is. Contemporary systematic botany cannot wait for such minutiae for many genera and species, quite obviously. Unless utter chaos is to result, the major emphasis of plant systematists must continue to be spent on the "exploratory" phase, guided by more occasional excursions into the "experi- mental" phases; until, in fact, there is nothing left to "explore", and that will not be for some time to come. If I knew as much about each of the 52 species of Aspidosperma as I do about two of the subspecies of Asclepias tuberosa after six precious years of study (cf. Ann. Missouri Bot. Gard. 34:353. 1947), or as Professor Babcock knows about Crepis after almost a lifetime of study, I might map the various clines with con- siderable precision. Then I might be able to designate a certain specimen, for instance, as “Aspidosperma album X Spruceanum (F;)”, ог go into detail if a backcross. As it is, some complicated backcrosses are apt to appear pretty homozygous to me; what am I to do? I believe that, as a systematic "explorer", what should be done is to con- struct my keys to species elastically enough to include genetic variants with the species most cognate to them, even at the expense of verbal contrast, giving warn- ing in a footnote if hybridization is suspect. I should consider myself insincere to attempt unreserved specific segregations if the slightest evidence of hybridization (or areal or ecological clines) were manifest. That has been my guide in this study of Aspidosperma, which probably recognizes fewer species than some readers will expect. My chief fear is that there still may be too many. At the other end of South America, in the campos of southern Brazil and adjacent Paraguay, Argentina, and Bolivia, a fascinating pattern of variation, pos- sibly the result of multiple-factor recombination following hybridization, is found to connect no less than four species of series PyricorLa: A. tomentosum, A. australe, A. subincanum, and A. parvifolium. These species all inhabit the same [Vor. 38 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN general range, and all are represented in our herbaria by copious specimens in which the various characters of habit, type of bark, size and shape of leaf and flower, and particularly peculiar indument are strongly and positively correlated. These are the most frequently collected of Aspidospermas, and amongst the several scores of specimens in our museums there are sufficient to testify to the absence of absolute pollination and genetic barriers between them. The species, ber se, are so distinctive that a tremendous range of recombination patterns is produced, and these are well represented in the specimens cited in the following taxonomic section. I have attempted to arrange the specimens into eight groups as a very rough estimate of this putative hybridization: GROUP I. ("Pure" tomentosum): Thick, heavily corky stems; large sessile leaves; long corolla lobes; inflorescence very condensed; follicles without a definite midrib—all very densely yellow-tomentose. GROUP II. ("Pure" australe) : Slender stems with thin bark at least on upper branches; smaller, long-petiolate, glabrous leaves; short corolla lobes; inflorescence diffuse; follicles without a definite midrib or indefinitely striate. GROUP III. ("Pure" subincanum): Slender stems with thin bark; rather large, sessile leaves finely white-sericeous beneath and glabrous above; inter- mediate corolla lobes; inflorescence diffuse; follicles with a prominent mid- rib. GROUP IV. ("Pure" parvifolium): Slender stems with thin bark; small, moderately petiolate leaves; short corolla lobes; inflorescence few-flowered; follicles with a very prominent midrib—all densely reddish-tomentellous. GROUP V. (Gomezianum, etc.) : Rather slender, thinly rimose stems; rather large, sessile or short-petiolate leaves nearly or quite glabrous; intermediate corolla lobes; inflorescence much branched from the base, finely puberulent; glabrous follicles without definite midrib. (tomentosum X australe?). GROUP VI. (Chodatii): More slender, thinly rimose stems; large, long- petiolate leaves finely canescent to glabrate; long corolla lobes; inflorescence fairly long-pedunculate, finely puberulent to glabrate; follicles with a prom- inently elevated midrib. (tomentosum X subincanum 2). GROUP УП. (camporum, Warmingii, etc.): More slender, thinly rimose stems; small or medium-sized, short-petiolate leaves; shorter corolla lobes; shortly pedunculate inflorescences—all very densely orange-tomentellous; follicles with a prominent midrib. (tomentosum X parvifolium ?). It seems significant that this group is found in the southern coastal Brazilian states. GROUP ҮШ. (various backcrosses to fomentosum ?): Much as in Group I, but stems somewhat more slender and with less corky bark. The enumeration of such groups could be subdivided almost indefinitely to indicate accessory hybridizations particularly of australe with closely neighboring 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 135 species (vide infra). The collections which my friend Senhor Amaro Macedo has sent me for study are helpful in this connection, since he has found trees of four of these groups, I, VI, VII, and УШ, within a single day's collecting in the vicinity of Ituiutaba, Minas Gerais. I hope that this problem will not escape the attention of Brazilian botanists who are interested in natural hybridization in their rich flora. Another type of difficulty is represented in series PyrIcoLLa by the group of species comprising A. australe, A. pyricollum, A. Ulei, and A. Vargasii, enumerated in their approximate geographic sequence from south to north in eastern South America. As reference to the key to species will show, the morphological criteria characterizing these species are of a somewhat different character, and are generally more difficult to analyze than is true in the genus as a whole. They are capable of recognition, however, and the rather unsatisfactory morph- ological differentia appear to be reinforced by distinctive geographical ranges. Hybridization also may appear to be the agency in producing morphological blur- ring at the boundaries of the ranges. Such being the case, I suspect that the units which I am designating as closely knit species may, upon more adequate materials for study, prove to be geographic subspecies. I hesitate to change their presently accepted status because my rather paradoxical feeling is that the recognition of subspecies requires more detailed study than for species! But I believe that this viewpoint is easily defensible. A knotty problem of nomenclature which will have to be tackled sooner or later concerns type specimens which are shown to be heterozygous. Ап instance in point is the type specimen of Aspidosperma album (Richard s.n. in Herb. Haun.), which has the peculiar inflorescence and leaf outline of supposedly homozygous A. album as defined in this paper, but with the leaves white-tomentu- lose beneath as in A. Spruceamum. Should a "neotype" be selected from the homozygous populations and the historical holotype disqualified as heterozygous? Opinion may sanction that at length, but I believe that few contemporary botanists would allow a policy so gravid with confusion; for type specimens of incalculable number would be subject to change from the time of Linnaeus. The inference which I draw is that type specimens will be but of casual interest, once the “explor- atory” period of plant systematics is ended; until then, our “higher criticism” of them must be held in check. Economic UsEs AND VERNACULAR NAMES Notes on the economic uses of Aspidosperma as reported by plant collectors are scattered throughout the following text, under the appropriate species. Detailed information on the timbers of various species will be found in Record & Hess, ‘Timbers of the New World’, pp. 107-115 (Yale University Press, 1943). The trees are so familiar to the local inhabitants that vernacular names are legion. These are noted for each species, when reported by collectors of the speci- mens cited, and are gathered together in a special index preceding the taxonomic index. [Vor. 38 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN ӛторү MATERIAL I have been privileged to study herbarium specimens from the following insti- tutions, to the curators of which I am greatly indebted. I have used the symbols advocated by Lanjouw (in Chron. Bot. 5:143. 1939) in citing the specimens: C—Botanisk Museum, Copenh F—Chicago Natural History ds Field] Museum, Chicago. G —Conservatoire Botanique, Geneva GH—Gray Herbarium, Cambridge, Mass. P—Muséum National d’Histoire Naturelle, Paris. 5 Mafurbüdielika: erdt tockh = US—U. E ұлын Herbarium, Washin U—Botanisch Museum en Herba rium, na SABE OU on andas Museum, Vienna. COL—Instituto de Ciencias Naturales, Bogota. R—Museu Nacional, Rio de Janeiro. TAXONOMY AsPIDOSPERMA Mart. & Zucc. in Mart. Nov. Gen. & Sp. 1:57. 1824; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4:141. 1895; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:362. 1947, nom. conserv. (Т.: A. tomentosum Mart.). Macaglia Rich. ex Vahl, іп Skrivt. Nat. Selsk. Kjoebenh. 6:107. 1810. (T.: M. alba Vahl). Peltospermum DC. in Bibl. Univ. Genéve 17:133. UN т m m" DC.). Ostreocarpus Rich. ex Endl. Gen. Pl. 1396. 1840, Thyroma Miers, Apoc. So. Am. 22. 1878. (T.: Th. 5 (Muell. Arg.) n Cufodontia ro in Archivio Bot. Sist. Fitogeogr. & Genet. 10:39. 19 Сте: stegomeris Woodson). Coutinia Vell. бле Portug. 166, pl. 10. 1799. (T.: C. illustris Vell.). Trees, rarely shrubby, usually with milky or reddish latex. Leaves usually alternate or approximate, rarely decussate or ternate, estipulate, penninerved. In- florescence terminal or axillary, occasionally extra-axillary, determinate, dichasially cymose (frequently with the determinate flower suppressed) to aggregate-dichasial and thyrsiform, the secondary peduncles frequently fasciculate in umbelliform manner, bracts persistent, caducous, or suppressed; calyx usually of 5 free, equal to strongly unequal lobes, rarely 4 with the 2 outermost larger and strongly con- nate with the 2 inner and free lobes wholly included, always eglandular within; corolla salverform to tubular-salverform or tubular, the tube cylindrical to ampuli- form and constricted or not constricted at the orifice, fissured behind the anthers, the lobes sinistrorsely contorted in aestivation, reflexed to erect in anthesis; stamens inserted midway or higher within the corolla tube, filamented, the anthers minutely mucronulate, of 2 uniformly fertile thecae, each 2-loculate, the outer loculus some- what longer than the inner and usually somewhat inflexed at the base, the pollen 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 137 granular; pistil superior, 2-carpellate, the carpels essentially free from the style to the receptacle, sometimes apparently glandular toward the base but without a distinct disc, bearing 2-32 peltate 2—6-ѕегіате ovules upon a thin ventral placenta, the stigma variously clavate to subcapitate upon a more or less elongate common style. Fruit follicular, usually strongly compressed and more or less woody, more or less assymetrical from nearly circular to falciform or dolabriform, the seeds peltate, exalbuminous, greatly compressed, with a flat, papery, concentric or strongly excentric wing. KEY TO THE SERIES a. Corolla typically т the lobes reflexed or horizontally spread- ing, e con ted at the orifice; bracts wholly suppressed (ex- EI: b. Corolla 1.2-2.5 cm. long, of relatively thick textu c. Inflorescence terminal or subterminal, eem bie: follicles 254 dolabriform, subplano- 452 the placenta rota ting about 225? gradually narrowed to a conspicuous 45,9 ely papillae t te essentially glabrous, we obvious lenticele, pow ur larg seed i n wing I. Macrocarra (p. 138) o O 4 3 = © m бе БЕ O о 3 e о ө ы c тг е © mn o £ on л Lo] о т a [e] et EHI RES =: 6 LE e [77] Ч = >” оо 95 с. Es $ x =o. © 2 ы с is ^ [җа] © Ж pla nv the p papillat te to ау о de gelo vp эдетаты moderately large; seeds with a nearly circular concentric wing........ bb. Corolla 0.4—0.8 cm. long, or if Кед (А. pyrifolium) of delicate texture; follicles pyriform, unequally bi- -convex, the placenta rotat- ute II. RAMIFLORA (p. 142) A. tomentosum), conspicuously lentic ege seeds with балш. ovate acute at the base ПІ. Pvnicorra (p. 144) aa. Corolla tubular or tubular- iatis! An lobes erect or зе 054 spread- ing at anthesis (salverform with xed lobes in A. rigidum), the tube not constricted at the orifices ae usually manifest and per- sistent, occasionally wanting (caducous in QUE HIN b. Corolla b essentially tercte, c: lobes not Wess Ка -асит г loosely spiral in the bud; inflorescence ecce odd nor Е c. Leaves Шо or irregularly approximate, not spine-tipped. d. Calyx lobes 5, abnormally i distinct or barely united at the base, equal to more or m unequal. e. Corolla glabrous w DUM t “(occasionally velutinous-papillate toward the tip in рот f. Corolla tubular or ae salverform, the lobes erect or placenta rotatin ut 45 — sessile, essentially ee caapi е а еф with a arrow, strongly eccentric basal wing (except in sree um) IV. PorvNEURA (р. 157) . Corolla salverform, the lobes reflexed at anthesis; ovary polygonally angulate; follicles very br sa dolabriform, concave-convex, the place ed "^ with or without a midrib, sessile; seeds ich a concent wi V. КїсрА (p. 164) E ee. Corolla densely sericeous without; ovary died x or m 21 s very broadly dolabriform, conc E ual bi-convex, uu placenta rotatin ibd t о" adt or spiny, sessile or very a abru d tly id short- ing VI. Nitwa (p. 166) [Vor. 38 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN dd. Calyx lobes 4, abnormally 5 the outer pair much larger and "nd connate (except in А. illustre), completely Ж ч the r pair which are separate and much smaller; нев - е plac 27 sessile o ortly stipicate, ay loose, wrinkled, more or les ы ы periderm; seeds wih a concentric circular VII. STEGOMERIA (p. 176) cc. Leaves opposite or К. ‘pine жүред; follicles broadly oval to ee. v equally bi-c x, the placenta rotating about utely papillate to esse мез? м glabrous, obviously tat pei or very shortly stipitate (А. borco- prosara eeds with a nearly circular concentric wing VIII. QuEBRACHINES (р. 180) bb. Corolla tube strongly Шотана, the lobes Hr ду caudate- Fig oar and very tightly spiral-contorted i d (except in А. leri); primary bran d ge the iced bas whorled ог deci follicles somewhat pyriform to nearly circular or sub- cy in ric, | — equally bi-convex, the placenta rotating about 120°-180°, stipitate to sessile, densely tomen A пог obviously lenticellate; а with a concentric, nearly circular зуїпр................... IX. NoniLEs (p. 183) SERIES I. Macrocarpa Woodson, n. ser. 6 Macrantha M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947, in part. Trees with milky latex, apparently deciduous, with thick corky, or thin rimose bark, not conspicuously lenticellate. Branches with naked, rounded buds, the sub- tending internode definitely thicker than the terminal petiole, without definite seasonal articulations, but with internodes greatly shortened toward the tip of the branch. Leaves alternate, rather broad and subcoriaceous, with widely arcuate secondary veins reticulating distally and with apparent reticulate tertiary venation on the lower surface. Inflorescence terminal or subterminal, cymiform, without obvious bracts; corolla typically salverform, the lobes reflexed or horizontally spreading, the tube constricted at the orifice, relatively large and of thick texture, densely tomentose or tomentellous without. Follicles unusually large, broadly dolabriform, subplano-convex, the placenta rotating about 225°, gradually nar- rowed to a conspicuous stipe, minutely papillate to essentially glabrous, without obvious lenticels; seeds with a nearly circular, concentric wing. Species 1—2 Figure 4. KEY TO THE SPECIES Flowering gie сн е the internodes not greatly con- ете rata and rimose; leaves of moderate "ah abruptly ced rca der po etioles, iran pubescent to «epist indie aber. -sericeous without, the tube — fissured . А. MACROCARPON . Flowering ipte very stout, the internodes greatly condensed, de periderm very thick and suberose; leaves very large, decurrent to shor stout неми densely t tomentose to lne in age; corolla densely tom- the entose, the tube inconspicuously fi 2. А. VERBASCIFOLIUM (A. m Mc jan and A. se npe cam appear to hybridize freely) P ы = - M ASPIDOSPERMA MACROCARPON Mart. Nov. Gen. & Sp. 1:59. 1824; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947. (T.: Martius s. n.) eve codo We Muell. Arg. in Mart. Fl. Bras. 61:46. 1860; M. Pichon, loc. : St. Hilaire 8. Жы. м. (sic!) Mart. a, normale Muell. Arg. loc. cit. 47. 1860. (A. macrocarpon Mart., var. typ. 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 139 Fig. 4. Aspidosperma macrocarpon Mart.: Flowering branch, bud, dissected flower, fruit, and seed. Aspidosperma macrocarpum Mart. B. glabratum Muell. Arg. loc. cit. 1860. (T.: Riedel 540). Aspidosperma macrocarpum Mart. y. macrothyrsum Muell. Arg. loc. cit. 1860. (Т. Riedel 2770). Aspidosperma macrocarpum Mart. 8. lanatum Muell. Arg. loc. cit. 1860. %2 Riedel EN: Aspidosperma Gardneri Muell. Arg. loc. cit 1860; M. Pichon, loc. cit. 1947. (T ner 2000! 2.1. Gardneri Muell. Arg. a. ovatum Muell. Arg. loc. cit. 48. 1860. (A. Gard- р.). Aspidosperma Gardneri Muell. Arg. В. ellipticum Muell. Arg. loc. cit. 1860. (T.: Weddell мааа platyphylla pas Arg.) O. Ktze. n бев, 2:416. 1891. Macaglia macrocarpa (М p O. Ktze. loc. cit. Macaglia Gardneri (Muell g.) O. Ktze. loc. т i. Aspidosperma Duckei E in Archiv. Jard. Bot. Rio. Jan. 3:244. 1922. (T.: Ducke ! du Snethlagei Mgf. in Notizbl. 10:118. 1927. (Т.: Snethlage 676!). Aspidosperma LeCointei auct. ex Record, apud Milanez, in Physis 15:479. 1939 [Vor. 38 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN Trees 3-25 m. tall, the branches relatively slender, with thin, rimose bark. Leaves elliptic, obtuse at base and apex, about 10—17 cm. long and 5-8 cm. broad, firmly membranaceous, wholly glabrous to variously yellow-pubescent; petioles about 1-4 cm. long. Inflorescences clustered subterminally at the uppermost nodes, subumbelliform, several-flowered, densely and minutely ferruginous-tomentellous to yellow-tomentose; primary peduncle about 1—3 cm. long, the secondary pedun- cles obsolete or nearly so; pedicels 1-2 mm. long. Calyx lobes ovate, acuminate, 2-4 mm. long, nearly equal, appressed-tomentellous to tomentose. Corolla white, densely appressed-sericeous to yellow-tomentose without, the tube 6—7 mm. long, about 2 mm. in diameter at the insertion of the stamens, somewhat constricted at the base and at the orifice, deeply fissured, nearly glabrous within, the lobes oblong- elliptic, 7—9 mm. long. Stamens inserted at about midway within the corolla tube, the anthers about 1.5 mm. long. Ovary ovoid, about 1.5 mm. long, wholly gla- brous or tomentellous; stigma narrowly ovoid to subfusiform. Follicles semi- circular or dolabriform, 9—14 cm. long, 8—12 cm. broad, the stipe about 3 cm. long, very stout, pericarp dark brown to black, irregularly striate, minutely tomentellous; seeds circular, about 8 cm. in diameter. Central Brazil and adjacent Venezuela, Bolivia, and Paraguay; savannas and gallery forests; flowering chiefly from August until September. Vernacular names: Platanote (Venezuela—Ll. Williams); Muirajussára (Brazil —A. Ducke). VENEZUELA: AMAZONAS: El Ratón, alt. 100 m., Williams 13223 (F ). BRAZIL: ACRE: near mouth of Rio Маска (сынасу of Rio Yaco), Krukoff 5682 (К, МО, U, US). amazonas: Parintins, Ducke 24574 (К, US). coraz: Riedel s. m G . MINAS GERAIS: St.-Hilaire 1770 (P). PARA: Obidos, Ducke 11040 (G, P, US), 11402 (К, U); Rio Tapajóz, Bella Vista, Ducke 16489 (С, US); haut Ariramba (Trombetas) , Ducke 14889 (G); bords du Rio Toca qs Weddell 2470 (Р). PIAUHY: S. Filomena, Snethlage 676 (F); Gardner 2666 (P, U ). PARAGUAY: — y ago Hassler 10575 Tn P, W), 10805 (М). OLIVIA: LA tolomé (near Calisaya), Krukoff 10108 (MO, U, US), тотоёа (МО); Tuiri чы ат Krukoff 10802 (МО, О); Tumupasa, Cárdenas 1085 , NY). Aspidosperma macrocar pon is a rather uniform population with glabrous, long- petioled leaves and appressed ferruginous-tomentellous inflorescences borne upon leafless twigs in the extreme northern and western portions of its range. In the central range and in the south, however, it becomes more variable, the leaves tending to be increasingly shorter-petioled and the inflorescences borne with the new foliage, both being more and more yellow-tomentose. Since these tendencies approach the characters of A. verbascifolium, which is indigenous to southern Brazil, I assume that the two species hybridize. The most complete field notes on A. macrocarpon, unfortunately, are those by Ll. Williams accompanying the single, completely sterile specimen from Venezuela which is cited above: “N. v.: Platanote—A rbol de 20-22 m. de altura, lactifero; 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 141 copa angosta de pocas ramas erectas; el tronco es redondo, derecho, sin estribos, 25 cm. de diámetro, sin ramas por 9—10 m.; la corteza es áspera, gris claro y el liber rojo rosado exuda en abundancia un látex pegajoso; la madera es de color claro; las flores son terminales, de pétalos blancos; el fruto es dehiscente; en la márgen entre la sabana y la selva de galeria." In Bolivia, Cárdenas notes (sub 1085 in hb. N. Y.): "Big tree with very hard wood." 2. ASPIDOSPERMA VERBAsCIFOLIUM Muell. Arg. in Mart. Fl. Bras. 61:46. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947. (T.: St. Hilaire 825!). Macaglia verbascifolia (Muell. Ar 8.) О. Ktze. Rev. Gen. 2:416. 1891. Macaglia lanata О. Ktze. loc. cit. 37:197. 1898. (T.: Kuntze s. n.!). Aspidosperma lanatum (О. Ktze.) Malme, in АВ Bot. 21A, no. 6:20. 1927. Apparently rather small trees, the branches disproportionally stout, with greatly condensed internodes and very thick periderm, yellow-tomentose to glabrate. Leaves broadly elliptic to elliptic-obovate, apex broadly obtuse to rounded, broadly decurrent to the petiole, about 10—30 cm. long and 7-16 cm. broad, firmly mem- branaceous, densely yellow-tomentose on both faces to glabrate or glabrous above; petioles about 1—2 cm. long. Inflorescences clustered subterminally at the upper- ost nodes, corymbiform to subumbelliform, densely yellow-tomentose, 5-8 cm. long, the dichotomously branching secondary peduncles about equalling the primary, to more or less suppressed, the pedicels 3-5 mm. long. Calyx lobes ovate to ovate-lanceolate, acute, 3—4 mm. long, nearly equal, densely yellow-tomentose. Corolla white, densely yellow-tomentose without, the tube 6—7 mm. long, about 2.5 mm. in diameter at the insertion of the stamens, somewhat constricted at the base and at the orifice, slightly fissured behind the anthers, essentially glabrous within, the lobes oblong-elliptic, 9710 mm. long. Stamens inserted about midway within the corolla tube, the anthers about 1.7 mm. long. Ovary ovoid, about 1.5 mm. long, glabrous or papillate; stigma subglobose. Follicles semicircular or broadly dolabriform, about 15 cm. long and 10 cm. broad, rather abruptly pro- duced to a stipe about 6 cm. long, deep brown, irregularly striate and minutely velutinous-papillate; seeds circular, about 8 cm. in diameter South-central Brazil, in "cerrados"; flowering from August to October. Vernacular names: Panaceia; Peroba do campo (Oliveira) ; Moela de emei ( A. Macedo) ; Peroba amarga (Duarte). BRAZIL: GolAz: St.-Hilaire 825 Ou entre re et Ponte Laurada, Glaziou 21741 (С, F, P). Maro Grosso: Kuntze s. n. (W), .n. (R); Cuiabá, Kublmann 1227 (R). MINAS GERAIS: Uberava Resnell 8 71 (F, US); Мын "Macedo 542 (MO); Patos de Minas, carmo do Paranaiba, Oliveira 1330 (MO); Serra de Catiara, Duarte 2038 (MO). Very little is known about this species, which appears to hybridize with the preceding (vide ante). Glaziou (sub n. 21741 in Hb. Par.) describes the plant as “arbuste torteux"; Macedo (sub n. 542 in Hb. Mo. Bot. Gard.) describes it merely as "tree". [Vor. 38 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN SERIES П. RAMIFLORA Woodson, n. ser. Ser. Macrolobii K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?:141. poi ы in part; $ Macrantba M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947, in par Trees with milky latex, apparently evergreen, with rather thin, somewhat loose bark, rather inconspicuously lenticellate. Branches with naked, acuminate buds, the subtending internode definitely thicker than the terminal petiole, without definite seasonal articulations, but with internodes greatly shortened toward the tip of the branch, the two uppermost leaves appearing opposite as a rule. Leaves alternate, membranaceous, large, with widely arcuate secondary veins reticulating distally and with very apparent reticulate tertiary venation upon both surfaces. Inflorescence cauliflorous some distance from the tip of the branch, sessile, few- flowered, with persistent, evident bracts; corolla typically salverform, the lobes reflexed or horizontally spreading, the tube constricted at the orifice, relatively large and of thick texture, densely tomentellous without. Follicles rather large, broadly dolabriform, plano-convex, the placenta rotating about 270°, indefinitely papillate to essentially glabrous, very conspicuously lenticellate; seeds with a nearly circular, concentric wing. Species 3 Figure 5. 3. ASPIDOSPERMA RAMIFLORUM Muell. Arg. in Mart. Fl. Bras. 61:55. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. П, 19:364. 1947. (T.: Sellow 1651!). Geissospermum ? жүн Mart. ex Muell. Arg. loc. cit. 1860, лот. nud. in synon.; Miers, Apoc. So. Am. 1878. Trees 12-30 m. tall, with close, relatively thin bark. Leaves oblong to oblong- elliptic, acute at base and apex, about 9—20 cm. long and 4-8 cm. broad, mem- branaceous, glabrous; petioles 1—2 cm. long. Inflorescences sessile, few-flowered, cauliflorous on the younger branches several nodes below the tip. Calyx lobes ovate, acute, nearly equal, about 2 mm. long, densely ferruginous-tomentellous without. Corolla densely ferruginous-tomentellous without, the tube 7-10 mm. long, about 2 mm. in diameter at the insertion of the stamens, constricted at the thickened orifice and near the base, the lobes broadly oval to oblong-elliptic, obtuse to rounded, 6-12 mm. long. Stamens inserted about midway or slightly higher within the corolla tube, the anthers 1.0-1.5 mm. long. Ovary oblong-ovoid, about 1.5 mm. long, glabrous or somewhat ferruginous-pilose; stigma fusiform. Follicles obliquely oval, sessile or very shortly stipitate, 6—9 cm. long and 5-7 cm. broad essentially glabrous, nearly covered with whitish lenticels; seeds broadly oval, about 6 cm. long and 4.5-5.0 cm. broad. South-eastern Brazil and eastern Bolivia; dense forests; flowering from May until October. Vernacular names: Peroba amarella (Brazil—Servico Florestal do Brasil) ; Tambá (Brazil—Mexia). 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 143 Flowering branch, dissected flower, fruit, Aspidosperma ramiflorum Muell. Arg Fig. 5. and seed. Ввл?п.: Sellow 1651 (W); Schuch s. п. Uu СН, ХУ). MINAS GERAIS: Fazenda de Aguada, Е 5054 (С, СН, К, МО, NY, S, О, US). RIO DE JANEIRO: a e Cinqué, о 9 (C, P); S. Christorao, Morrada Telegrapho, Glaziou 6636 (C, F, P); Corco- vado a T de Freitos, Glaziou 17132 (C ad urbem in silvis montis Сы ёте, Kubl- mann 16366 (U, US); Matta da Fabrica Catipos, Serviço Florestal 105 ( BoLivia: LA PAZ: Guanai, Rusby 2649 (GH, NY, US); Huachi, White 1018 (NY). P); White (sub по. 1018) describes the wood as hard and white; Mrs. Mexia (sub no. 5054) notes that the trunks are straight and slender, with hard wood, and An attempt was made to distinguish the Brazilian popu- producing good lumber. lation from the Bolivian upon morphological grounds, but without success. [Vor. 38 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN Series ІП, PyricoLtta Woodson, n. ser. Ser. е” К. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 47:141. 1895, in part; e rolobii K. Sch. loc. cit. 142. 1895, in part; $ Tetrasticha M. Pichon, in Bull. = Hist. a II, 19:363. 1947, in part; $ Glabriflora M. Pichon, loc. cit. Т : p» part; 5. Piliflora M. Pichon, loc. cit. 364. 1947, in part; 5 — ba M. Pichon, loc. cit. 1947; $ Microloba K. Sch. ex M. Pichon, loc. cit. Trees, occasionally shrubs, with milky latex, 2. with moderately thin rimose bark, conspicuously lenticellate. Branches with scaly buds, with definite seasonal articulations and with the leaves rather crowded toward the tips. Leaves alternate or irregularly approximate, membranaceous to subcoriaceous, moderately large to rather small, with widely arcuate secondary veins reticulating distally Inflorescence terminal or subterminal, pedunculate, many- or few-flowered, with- out evident bracts; corolla typically salverform, relatively small, or if larger of delicately membranaceous texture. Follicles pyriform, unequally bi-convex, the placenta rotating about 180°, gradually narrowed to a conspicuous stipe, indefi- nitely papillate to essentially glabrous (densely yellow-tomentose in A. foment- osum), conspicuously lenticellate; seeds with a concentric, ovate wing acute at the base. Species 4-14 Figure 6. KEY TO THE SPECIES a. Corolla lobes twice as long as us tube or somewhat longer — —— small, about 7 mm. long, glabrous; inflorescence ят, ас -flowered, Kia with the terminal flowers absent below И "e relatively large, about 1.5 cm. appressed-sericeous without; inflorescen — al, the termina ki flower alw nspic эе ұла mtg? hine or dt «vet an in "me t midrib........ . А. PYRIFOLIUM aa. Corolla lobes much shor the tube, or about as long. b. — he үтеп as etree as the tube. mall-le sad "ine glabrous osos RR flowers solitary in the 4. А. MULTIFLORUM o a long, glabrous or somewh pe r lea 6. А. OLIGANTHUM cc. Large- loved. trees, кей pex I inflorescence densely yellow- tomentose; inflor ered; follicles densely yellow- tomentose leaves ane pe in follicles essentially glabrous ybrids) . А. TOMENTOSUM bb. бее eis ы а$ аз as the prk or shorter. c. Inflorescence eral-flow d. Corolla ең Гео acuminate, Ко. half as long as the tube; еи — about twice as long as ada — pa борене elevated midrib o n both f ai i entic 8. A. SUBINCANUM ne cu dd. Corolla pesa ovate to ren irt obtuse to rounded, one-third o one-quarter as long as the tube. e. af olla 1 c ree about one-quarter as long as as (the tube; inflorescence ve densely - elliptical, өл Эм» three times as long as broad, w E a conspicuously elevated midrib on both fac кай, reddish-brown, — red- undi Jn to тма 9 ee. Corolla lobes ovate, about one-third as long as the tube. f. Corolla tube relatively narrow, di ut one-fourth or less s wide as long; inflorescence white-, yellow-, or yellowish . A. PARVIFOLIUM g. Inflorescence somewhat shorter than the subtending peti- oles, white- өле еріні corolla 6.5-8.0 mm. long; follicles 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 145 sa circular, grayish when dry, without a prominent idrib 10. A. AUSTRALE g. "ra orescence somewhat longer than the subtendi ing Қ ел x dg nd Laie whee less than about half a oad s long, bro when dry, PAS ie wi e Ani seid cachet on either h. Inflorescence rather loose, am axes yellow-pubescent to glabrate; corollas 6.5—7.0 mm. long; leaves oblanceolate to obovate, rounded to buna: 11. A. PYRICOLLUM hh. Inflorescence quite dense, Ae axes ier eae elas Te piti corollas —5.5 опр; os ellip O acumina 12. A. VARGASII ff. Corolla E RNC broad, Kus half as puc as long; inflorescence yellow т; Viper about . long; follicles ot age vial abou s long as broad, te а ver reddish-brown 13.%А Ота cc. Inflorescence of solitary flowers in the upper leaf ane ora . 5 mm. long, the lobes about half as long as the еа 4. А. REDUCTUM 4. ASPIDOSPERMA MULTIFLORUM A.DC. in DC. Prodr. 8:397. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:363. 1947. (T.: Blanchet 2806!). Macaglia multiflora (A.DC.) О. Ktze. Rev. Gen. 2:416. E ma Monteroi Standl. ex Record & Hess, Timbers of the New World, 61. 1943; ecord, in Trop. Woods 80:1. 1944, nom. nud. Small to medium-sized trees with rather thin, reddish bark, glabrous through- out. Leaves elliptic, apex shortly and obtusely acuminate, base broadly obtuse to rounded, 5—11 cm. long, 3-6 cm. broad, membranaceous; petioles 1.5—2.5 cm long. Inflorescences clustered subterminally at the uppermost nodes, 3—5 cm. long, the peduncles slender, 3—4 times dichotomous with the terminal flowers ab- sent at the lower dichotomies, many-flowered. Calyx lobes ovate, acuminate, essentially equal, about 1.5 mm. long, scatteringly ciliolate. Corolla greenish white, glabrous without, the tube about 2 mm. long, about 0.7 mm. in diameter at the insertion of the stamens, somewhat constricted at the orifice and at the base, the lobes about 5 mm. long, narrowly oblong-ellipic. Stamens inserted at about midway within the corolla tube, the anthers 0.5—1.0 mm. long. Ovary ovoid, about 0.5 mm. long, glabrous, the stigma ovoid. Follicles compressed-pyriform, about 5 cm. long and 2.5 cm. broad, rather gradually narrowed to a stipe about 1 cm. long, with many conspicuous lenticels of varying size, with an indistinct midrib. Eastern and southern Brazil; in savannas; flowering in October. BRAZIL: ВАЇА: in certam fluvii S. Francisci, Blanchet 2806 (С, Е, P, W). MATO Grosso: Salto Belo, Rio Sacre, Baldwin 3115 (MO, US). para: Campina do Jupiry prope lacum Faro, Ducke 21801 (MO, US). The description of the fruit is drawn from Ducke 21801, which has been named A. multiflorum by my friend Dr. Markgraf. I am a bit doubtful of its real identity but do not know of a more sure disposition of it. I believe that A. multiflorum may hybridize with the A. tomentosum complex in Minas Gerais; this possibility will be discussed following the account of the latter species. [Vor. 38 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5. ASPIDOSPERMA PYRIFOLIUM Mart. Nov. Gen. & Sp. 1:60. 1824; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947. (T.: Martius s.n.!). pee refractum Mart. loc. cit. 1824; M. Pichon, loc. cit. 1947. (T.: Martius a s bicolor Mart. loc. cit. 1824. (T.: Martius s. n.!) еи populifolium A.DC. in DC. Prodr. 8:397. 1844; M. Pichon, loc. cit. 1947. : Gardner 2604! Aspidosperma Martii Мб ex Muell. Arg. іп Mart. Fl. Bras. 61:48. 1860; M. Pichon, loc 7. (T.: Martius 485!). уе. byrifolium Mart. В. molle Muell. Arg. loc. cit. 54. 1860. (T.: Martius s.n.!). i 78. cive Mg guaranticum Malme, in К. Sv. Vet. Akad. Handl. Bihang, 24, afd. 3, no. . I, fig. I. 1899; М. Bichon, loc. cit. 1947. (T.: Malme 1006!). sat to medium-sized trees 2—14 m. tall, usually with brittle, jointed, rimose branches, finely appressed-puberulent to glabrous when young. Leaves elliptic to ovate, frequently somewhat pandurate, apex acuminate to obtuse, base obtuse to rounded, 2—12 cm. long, 2—6 cm. broad, membranaceous, finely puberulent to glabrous; petioles 1-3 cm. long. Inflorescences borne with the young leaves, 2—5 cm. long, few- to several-flowered, regularly dichasial, the terminal flowers always present; pedicels 2-5 mm. long, minutely puberulent to glabrous. Calyx lobes ovate to ovate-lanceolate, acuminate, 2—3 mm. long, sparsely pilosulose. Corolla white, very fragrant, finely appressed-puberulent to essentially glabrous without, the tube 4—6 mm. long, about 1.5 mm. in diameter at the insertion of the stamens, the orifice constricted and thickened, the lobes lanceolate, narrowly acuminate, 1—2 cm. long. Stamens inserted in the upper third of the corolla tube, the anthers about 1 mm. long. Ovary ovoid, about 1 mm. long, glabrous; stigma subglobose, minutely pilosulose. Follicles nearly circular, shortly stipitate, the body 4—5 cm. long and 3—5 cm. broad, yellowish brown, conspicuously lenticellate, with or with- out an indistinct midrib, the stipe 0.5—1.0 cm. long; seeds broadly ovate, 5—6 cm. long and 3—4 cm. broad Widely distributed in eastern and southern Brazil and adjacent Paraguay; in dry scrub, thickets, and woodland (caatinga and varzea); flowering chiefly from September to January. Vernacular names: Pereiro (Brazil—Luetzelburg); Pereiro preto (Brazil— Lisboa); Peroba paulista (Brazil—Macedo); Ivahay, Palo de rosa (Paraguay— Malme). BRAZIL: ALAGOAS: Be = "P Falls, Chase 7821 (F, GH, MO, US). Baia: Caiete et praedium Maracas, . (W); in silvis catingas, Martius s. n. (W); Geremoabo, Schery 487 (MO); Calder "Ul; 7054. ds CEARÁ: Sao Estevam, Löfgren 770 (R); Sobral, Dahlgren 028 (F, MO); Rio осу at Fortaleza-Recife road, Drouet 2710 (Е, H, MO), 2724 (F, GH, ao, rert ekki Allemáo 967 (МО, P, R); Gardner 1753 (СН, P, US, W). Maro crosso: Cujaba, Martius 585 en. MINAS GERAIS: Ituiutaba, D951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 147 Macedo 2020 (MO), 2023 (MO). Pana: Vigosa, Lisbóa 2435 (US). PARAHYBA: tes Gonçalo, Luetzelburg 26800 (Е, MO); Serra Borborema, Luetzelburg 12356 (F, US). PERNAMBUCO: locality lacking, Pickel 3497 (GH, Е). prauny: locality гаа Gardner 2664 (P, US, W). RIO DE JANIERO (?): locality lacking, Glaziou 11184 (С PARAGUAY: Concepción, Hasler 7287 (MO, P, W), 7287a (MO); Colonia Risso pr. Rio Apa, Malme ee (R, S, US). The type specimen of E Martii may represent a hybrid between A. pyrifolium and A. macrocarpon. It consists of both fruits and flowers with foliage: the leaves and fruits can be referred to A. pyrifolium without difficulty, but the flowers are rather densely tomentellous and the lobes are slightly shorter than the tube and are somewhat too broad for true A. pyrifolium. In the original description Mueller suggested an affinity with A. macrocar pon. Flowers and foliage of A. pyrifolium appear to be somewhat more pubescent in its southern range than in the north, but I see no indications to suggest that this variation may be due to hybridization, although Sr. Macedo's specimens from Minas somewhat suggest A. subincanum, which occurs in the immediate vicinity. 6. AspIDOsPERMA Oliganthum Woodson, spec. nov. Frutex parvus ut dicitur ramulis fragilibus geniculatis tenue rimosis glabris. Folia obovata apice rotundata basi cuneata ca. 1 cm. longa 5-8 mm. lata mem- branacea glabra opaca; petiolis ca. 2-3 mm. longis. Flores in axillis folium superiorum solitarii; pedicellis 2-5 mm. longis tenuissimis glabris. Calycis laciniae ovato-lanceolatae anguste acuminatae ca. 1.5 mm. longae glabrae. Corollae extus glabrae tubo ca. 2.5-3.0 mm. longo ca. 1 mm. diam. lobis anguste lanceolatis acuminatis 3.0—3.5 mm. longis. Stamina in tubo corollae medio inserta antheris ca. 0.5 mm. longis. Ovarii carpella depresse ovoidea truncata ca. 0.25 mm. alta glabra ovulis 2 (vel 42); stigmate sessili fusiformi ca. 0.3 mm. longo. BRAZIL: Bafa: in der Catinga bei Tambury, October, 1906, Ule 7200 (K, түре). A most peculiar reduced form recalling A. reductum of Paraguay, but ap- parently most closely related to A. pyrifolium. Beside the tiny, solitary flowers and small foliage, the form of the ovary is remarkable. 7. ASPIDOSPERMA TOMENTOSUM Mart. Nov. Gen. & Sp. 1:58, 2. 34. 1824; М. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947. (T.: Martius s. m.!). dia ai du dasycarpon A.DC. in DC. Prodr. 8:396. 1844; M. Pichon, loc. cit. 1947. T.: Claussen 34 acie аш Советни A.DC. loc. cit. 397. 1844; M. Pichon, loc. cit. 1947. (T.: Gardner 835! Aspidosperma eld lu Fisch. ex Muell. Arg. in Mart. Fl. Bras. 61:45. 1860, nom nud. PENAS tomentosum Mart. B. velutinum Muell. Arg. loc. cit. 1860. (T.: Riedel 2, Aspidosperma tomentosum Mart. y. angustifolium Muell. Arg. loc. cit. 1860. (T.: Riedel 671!). [Vor. 38 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aspidosperma obscurum Muell. Arg. loc. cit. 48. 1860; M. Pichon, loc. cit. 1947. (T.: Pobl 4314!). m camporum Muell. Arg. loc. cit. 49. 1860; M. Pichon, loc. cit. 1947. (T.: el 586). Aspidosperma Hilarianum Muell. Arg. loc. cit. 50. 1860; M. Pichon, loc. cit. 1947. (T.: St.-Hilair ; Aspiesperma Warmingii Muell. Arg. in 2 Vidensk. Meddel. 101. 1869; M. Pichon, с. cit. 366. 1947. (T.: Warming s. n.!). уона a Muell. Arg. loc. cit. 1869; M. Pichon, loc. cit. 364. 1947. (Ts arming s. Macaglia tomentosa me O. Ktze. Rev. Gen. 2: E 1891. Маса Па dasycarpa (A.DC.) О. Ktze. loc. cit. 189 a арени Rojasii Hassl. in Fedde, Repert. 12: 257, 1913; M. Pichon, loc. cit. 363. (T.: Hassler [Rojas] 10629! Aspidosperma Quirandy Hassl. var. ыды ium Hassl. loc. cit. 260. 1913. (T.: Fiebrig Aspidosperma Chodatii Hassl. ех Mgf. in Notizbl. 8:427. 1923; M. Pichon, loc. cit. 366. . (T.: Hassler [Rojas] 10647!). Trees 4—25 m. tall, with disproportionally thick, heavily suberized branches with greatly condensed internodes (more slender and thinly rimose in putative hybrids). Leaves condensed at the tips of the flowering branches, sessile, ob- lanceolate to obovate, apex acute to rounded, base narrowly cuneate, 10—30 cm. long, 4—12 cm. broad, membranaceous, densely yellow-tomentose on both faces or glabrate above (smaller, petiolate, and wholly glabrous in putative hybrids). Inflorescences clustered subterminally in the axils of the upper leaves, 4—10 cm. long, densely yellow-tomentose, the flowers sessile or subsessile at the tips of the dichotomous peduncles. Calyx lobes narrowly ovate to lanceolate, acuminate to acute, 3—6 mm. long, densely yellow-tomentose. Corolla white to yellow, yellow- tomentellous without, the tube 3—5 mm. long, about 1 mm. in diameter at the insertion of the stamens, the lobes lanceolate, acuminate, 4—5 mm. long. Stamens inserted at about the upper third of the corolla tube, the anthers about 1 mm. long or somewhat less. Ovary ovoid, about 0.5 mm. long, glabrous; stigma fusiform, about 0.5 mm. long. Follicles broadly obovate, 5-6 cm. long and 3.0-3.5 cm. broad, shortly stipitate, densely yellow-tomentose, without a distinct midrib (gla- brate and with or without a midrib in putative hybrids); seeds ovate, about 4 cm. long and 3 cm. broad. Southern Brazil and adjacent Paraguay and Bolivia; in savannas and dry wood- lands (campos, cerrados); flowering from July to November. Vernacular names (Brazil): Pereiro do campo (Martius); Paroba, Pequía (Gardner); Pao pereira do campo (Warming); Tambá (Mexia); Guatambi, Guatambi do cerrado (Macedo) ; Pequía de pedra (Servico Florestal). BRaziL: Baia: Caeté, Riedel 671 (W). Mato crosso: Fazenda das Moças, Campo Grande, Archer & Gebrt 135 (US); locality lacking, Kuntze s. n. (NY); Cuiabá, Kuhl- mann 1230 (К). coraz: Anapolis, Oliveira 1342 (MO); Alexandre, Glaziou 21740 (P) 1951] WOODSON—STUDIES IN THE APOCYNACEAE. УШ 149 MINAS GERAIS: Ituiutaba, Macedo 520 (MO), 536 (MO), 711 (MO), 712 (MO), 737 (MO); Serra da Caraga, Glaziou 15211 (F, P); Serra do Lenheiro, prés S. Joao d'El Rei, ing s.n. (C, NY), Engle s. n. (C); Bello Horizonte, Mello Barreto 300 (F), 9222 (F, МО); Lagóa Santa, Mello Barreto 9213 (F, МО); Uberava, Regnell 871 bis (US); т Duarte 3204 (MO). PARANÁ: Jaguariahyva, Dusén 13116 (Е, GH, MO, US), 18022 (US); Itararé, Dusén 16514 (GH, MO), s. ». (GH, MO, nte US); Patrimonio, Dusén 16780 (US); Cachoeirinha, Whitford 4 А 126 (Е, СН, US). RIO DE JANIERO: Serra dos Orgãos, Gardner 835 (GH, NY, P, US); Ipanemil, Harsbberger 842 (US); locality lacking, Wi lkes Exped. s.n. E STA. CATHARINA: locality lacking, Mueller 170 (К). $АО PAULO: Araruguara, ае 036 (С). DATA LACKING: Sello 40 (0); Reichardt 13 (W), 32 (W); Pohl s. n. (ХУ); St.-Hilaire s. n. (F, P, US); Claussen s. n. (С), 326 (P), 327 (F, P), 346 (G Paracuay: Sierra de Amambay, E [Rojas] 10804 (МО, W), 10647 (MO, W), 106474 (МО, ХУ), 10620 (MO, W), 106294 (MO, W), тобот (MO, W); Concepción, Hassler 7100 (MO, W), 71994 (MO, W); Concepción, Balansa 1346 (K); zwischen Río pa und Río Aquidabán, Fiebrig 4202 (GH). BOLIvIA: LA PAZ: Sta. Ana, d’Orbigny 745 (P, W). This is the most frequently collected as well as the most variable species of Aspidosperma. strongly suspect the hybridization of it with three other species of the same area: A. subincanum, A. australe, and A. parvifolium. The four species are so distinctive in the characteristics of their stems, leaves, flowers, and fruits, as well as in their peculiar induments, that a tremendous range of recom- bination patterns is available, and these are well represented in the specimens cited above. I have attempted to arrange the specimens into five groups as a very rough estimate of this putative hybridization. I hope that this problem will not escape the attention of some Brazilian botanist who is interested in natural hybridization. GROUP 1. ("Pure" tomentosum): Thick, heavily corky stems; large sessile leaves; long corolla lobes; inflorescence very condensed; follicles without a definite midrib—all densely yellow-tomentose. GROUP II. (Gomezianum, etc.): More slender, thinly rimose stems; smaller, sessile or shortly petiolate, nearly or quite glabrous leaves; shorter corolla lobes; inflorescence much branched from near the base, finely puberulent; glabrous follicles without a definite midrib. (tomentosum X australe 2). GROUP Ш. (Chodatii): More slender, thinly rimose stems; large, long- petiolate leaves finely canescent or glabrate; long corolla lobes; inflorescence fairly long-pedunculate, finely puberulent to glabrate; follicles with a prom- inently elevated midrib. (tomentosum X subincanum ?). GROUP IV. (camporum, Warmingii, etc.): More slender, thinly rimose stems; small or medium-sized, shortly petiolate leaves; shorter corolla lobes; shortly pedunculate inflorescences—all very densely orange-tomentellous; follicles with a prominent midrib. (tomentosum X parvifolium 2). It seems significant that this group is found in the southern coastal Brazilian states. GROUP V. (БасЕсговвев to tomentosum ?): Much as in Group I, but stems somewhat more slender and with less corky bark. [Vor. 38 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN The collections which my friend Sr. Macedo has sent me for study are helpful in this connection, since he has found trees of three of these groups, I, II, and V, within a single day's collecting, in the vicinity of Ituiutaba, Minas Gerais. 8. ASPIDOSPERMA SUBINCANUM Mart. ех A.DC. in DC. Prodr. 8:397. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Martius 262!). bap de: subincanum Mart. B. tomentosum Muell. Arg. in Mart. Fl. Bras. 61:50. Claussen s. n.!). мезі я (Mart.) О. Ktze. Rev. Сеп. 2:416. 1891. Tree as much as 25 m. tall, with moderately stout, knotty, thinly rimose branches. Leaves broadly elliptic, abruptly acuminate, broadly decurrent to the petiole, 9-18 cm. long, 4-11 cm. broad, thinly membranaceous, finely and densely cinereous-puberulent to glabrous beneath, glabrous above. Inflorescences in the uppermost leaf axils, 3—8 cm. long, many-flowered, conspicuously pedunculate, densely cinereous-puberulent; pedicels 1-3 mm. long. Calyx lobes ovate-lanceolate, acuminate, about 2 mm. long, densely cinereous-pilosulose. Corolla yellow, finely pilosulose without, the tube about 4 mm. long and 1 mm. in diameter at the in- sertion of the stamens, constricted at the orifice and above the base, the lobes elliptic-lanceolate, obtusely acuminate, 2.0-2.5 mm. long. Stamens inserted in the upper third of the corolla tube, the anthers about 1 mm. long. Ovary ovoid, about 1 mm. long, minutely puberulent; stigma fusiform, about 0.5 mm. long. Follicles obovate, shortly stipitate, 3-4 cm. long and 2—3 cm. broad, with a prom- inent midrib, conspicuously lenticellate, essentially glabrous; seeds ovate, about 3.5 cm. long and 2.5 cm. broad. Southern Brazil; forests and woodlands (capoes, capeiras, cerradas) ; flowering from September until November. Vernacular names: Pão pereira do mato (Warming); Guatambí (Macedo); Carrasco (Krukoff). BRA GOIAZ: near Coco, upper Rio Tocantins, Krukoff 2067 (K, NY, U, US); Goiaz, Burchell 6501 (К), 7305 (К). MINAs GERAIS: Ituiutaba, Macedo 2021 (MO), 2025 (MO); Uberava, Regnell 868 (C, F, R, US); Lagóa Santa, Warming s. n. (C, NY) Bello Horizonte, Magalhães 652 (MO); locality lacking, Claussen s.n. (G), 328 (C, F, GH, NY, P, US); St.-Hilaire s. n. (P); Sebastianopolis, Pohl 2185 (W). MATO GROSSO: Cuyabá, Martius 262 (MO, P, W), Malme 1640 (R). são PAULO: Araruguara, Löfgren 972 (C). The possibility of hybridization of this species with A. tomentosa has been discussed under the latter; the unusually large leaves and longer pedunculate in- florescences of a specimen of A. pyrifolium collected at Ituiutaba by Sr. Macedo suggest possible hybridization with that species as well. 9. ASPIDOSPERMA PARVIFOLIUM A.DC. in DC. Prodr. 8:398. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Guillemin 604!). Thyroma parvifolia (А. x Miers, Apoc. So. Am. Aspidosperma — 4 Sch. in Engl. Bot. ты am Sai 67:30. 1901; M. Pichon, loc. cit. 1947. (T.: Glaziou 17697!). 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 151 Small or medium-sized trees, the branches somewhat stout, densely reddish- tomentellous when young, becoming glabrate. Leaves obovate-elliptic, acute to rounded, 4-10 cm. long and 1—4 cm. broad, firmly membranaceous, densely reddish-tomentellous to glabrate or glabrous beneath, glabrous above; petioles 1—2 cm. long. Inflorescences subterminal at the upper nodes, densely reddish-tomentel- lous, 2-4 cm. long, loosely or densely flowered; pedicels 1-2 mm. long. Calyx lobes ovate, acute to obtusish, 1.5-2.0 mm. long, densely reddish-tomentellous. Corolla greenish, densely reddish-tomentellous without, the tube 4—5 mm. long, 1.0-1.5 mm. in diameter at the insertion of the stamens, the lobes ovate-reniform, 1.0-1.5 mm. long. Stamens inserted at the upper third within the corolla tube, the anthers about 1 mm. long. Ovary ovoid, about 1 mm. long, densely and minutely reddish-puberulent, the stigma ovoid, about 0.5 mm. long. Follicles sub- elliptical, about 5-6 cm. long and 2.5—4.0 cm. broad, gradually narrowed to a stipe 1.5-2.0 cm. long, reddish brown, with a prominently elevated midrib; seeds oval, about 5 cm. long and 3 cm. broad. Eastern Brazil; in woodlands; flowering from January to July. Vernacular names: Piquía (Guillemin); Tambá café (Mello Barreto). ВкА7П.: Bafa: Machado Portello, Rose & Russell 10041 (NY, US). MINAS GERAIS: Fazenda da Cachoeira, Tombos, Mello Barreto 1393 (F), Oliveira 325 (MO). RIO DE JANEIRO: Sumaré, Kublmann 4373 (U, US); Alto Macahe, Glaziou 12075 (C, P), 17131 (P), 17607 (P), 18366 (C, P). são PAULO: Ubatuba, Guillemin 604 (G); locality lack- ing, Riedel s. n. (G, GH, U, W), Pohl s. n. (W Like the other species of the series PYRICOLLA, A. parvifolium appears to hybri- dize with others growing within pollinating distance. I interpret in this way the somewhat variable color and abundance of the characteristic indument as well as variability in leaf size and shape. The tendencies of this variation rather suggest A. pyricollum as a hybridizing agent. 10. AsPIDOSPERMA AUSTRALE Muell. Arg. in Mart. Fl. Bras. 61:58. 1860; М. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Sellow s. n.!). сои ы ыы Muell. Arg. in Pow Vidensk. Meddel. 104. 1869; M. Pichon, 63 : Warming s. n.!). Macaslia austral is PA Arg.) O. Ktze ee Gen. 2:416. 1891. ин етта о Hassl. ia Fedde, Repert. 12:259. 1913; M. Pichon, loc. cit. 366. 47. (T.: r 2329! ds osperma Qu abd v asl. var. campestre Hassl. loc. cit. 1913. (T.: Hassler 7199!). Aspidosperma Quirandy Hassl. э silvaticum Hassl. loc. cit. 1913. (T.: Hassler 10601!). Aspid Vin d australe Muell. Arg. var. estrellense Hassl. loc. cit. 263. 1913. (T.: Hassler fa) ! 51 ЕЕ australe Muell. Arg. var. erytbroxylum Hassl. Addenda РІ. Hassl. 12. 1917, d. nu Ge australe Muell. Arg. var. longepetiolatum Hassl. loc. cit. 1913, nom. nud. Aspidosperma occidentale Malme, Arkiv Bot. 21A%:10. 1927; M. Pichon, loc. cit. 366. 1947, non Mgf. (T.: Malme 2732!). Aspidosperma missionum Speg. in Physis 3:336. 1917. (T.: Spegazzini 1268 5—photo US!). Trees 3—20 m. tall, the trunk with suberized bark, the branches rather slender and thinly rimose. Leaves ovate- to narrowly oblong-elliptic, apex obtuse to [Vor. 38 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN acuminate, base obtusely cuneate to rounded, 5-12 cm. long, 1.5-5.0 cm. broad, firmly membranaceous, glabrous or inconspicuously pilosulose beneath when young; petioles 1.5—5.0 cm. long. Inflorescence densely clustered subterminally at the upper nodes, divaricately branched from near the base, 2-4 cm. long, densely appressed-puberulent; pedicels 2-3 mm. long. Calyx lobes ovate, acute, 1.5—2.0 mm. long, appressed-puberulent. Corolla greenish or yellowish, very densely white- sericeous without, the tube 5—6 mm. long, about 1.5 mm. in diameter at the insertion of the stamens, the lobes ovate, 1,5-2.0 mm. long. Stamens inserted at the upper third within the corolla tube, the anthers about 1 mm. long or somewhat less. Ovary broadly ovoid, about 1 mm. long, densely sericeous, the stigma ob- longoid, about 1 mm. long. Follicles nearly circular, 3-4 cm. long and 2.5-3.0 cm. broad, without a midrib or indefinitely striate, essentially glabrous, usually grayish, the stipe 0.5-1.5 cm. long; seeds ovate, about 3.5 cm. long and 2.5 cm. broad. Southern Brazil and adjacent Paraguay, Argentina, and Bolivia; in light woods; flowering in September and October. Vernacular names: Tamb verde (Brazil—Magalháes) ; Tambú canudo (Brazil —Magalhies, Oliveira). BRAZIL: MATO GROSSO: Corumbao, Malme 2732 (S). MINAS GERAIS: Belo Horizonte, СУЯ hi (MO), 689 (MO); Oliveira, pa iie 4447 (MO); Sta. Luzia, Magalháes 4446 (MO); Betim, Magalbáes 612 (M О); A raxa, 22! 1345 (МО); Arcos, Oliveira 252 at E арба Santa, Warming s. n. (C), Engel 10 6 (C). RIO GRANDE DO SUL: São Leopoldo, Leite x (СН); São Leopoldo, Dutra 772 oR). SAO PAULO: locality lacking, Mosen 2532 (C, P, К). DATA LACKING hus meridionalis", Sellow s. n. kp Villa Rica, Jorgensen 3600 (C, F, GH, МО, US); Gran Chaco, Hassler 2329 (K, MO, W); Estrella, Hassler [Rojas] 10651 (MO, W); Sierra de Amambay, Hassler [Rojas] 10755 © W). BOLIVIA: COCHABAM between Vila-Vila and Misque, Cárdenas E (MO, US). STA. CRUZ: Cer -— Steinbach 6394 (К); si id lacking, Herzog 1682 ARGENTINA: n Hort. Agron. La Plata (ex Misiones), BLU NM tipicos cultivados por el рг. Заа Наитап 3535 (MO The Hauman sheet was received without a name, and may represent А. mis- siomum Speg., since it corresponds well with the photograph of the type specimen. Aspidosperma australe is a very well-marked species, but appears to hybridize with both A. tomentosum and A. pyricollum. 11. AsPIDOSPERMA PYRICOLLUM Muell. Arg. in Mart. Fl. Bras. 61:58. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Weddell 445!). Aspidosperma Riedelii Muell. Arg. loc. cit. 56. 1860; M. Pichon, loc. cit. 1947. (T: Riedel 27711). е Sellowii Muell. Arg. loc. cit. 1860; M. Pichon, loc. cit. 1947. (T.: Sellow К olivaceum Muell. Arg. loc. cit. 57. 1860; M. Pichon, loc. cit. 1947. (T.: ellow s. n. siste — Muell. Arg. В. obtusifolium Muell. Arg. loc. cit. 58. 1860. (T.: ы Боен Muell. Arg. y. obovatum Muell. Arg. loc. cit. 1860. (T.: Sellow S. m.l Thyroma Riedelii (Muell. Arg.) Miers, Apoc. So. Am. 26. 1878. 1951] WOODSON—STUDIES IN THE APOCYNACEAE. УШ 153 Thyroma Sellowii (Muell. Arg.) Miers, loc. cit. 24. 1878. Macaglia olivacea (Muell. Arg.) O. Ktze. Rev. Gen. 2:416. 1891. j 1 ДІР Aspidosperma Riedelii Muell. Arg. var. genuinum Hassl. in Fedde, Repert. 12:262. 1913. Aspidosperma Riedelii Muell. Arg. var. genuinum Hassl. forma microphyllum Hassl. loc. (T.: Balansa 1344). Aspidosperma Sellowii Muell. Arg. var. genuinum Hassl. loc. cit. 263. 1913. Aspidosperma Sellowii Muell. Arg. var. collinum Hassl. loc. cit. 1913. (T.: Hassler 4044). pos n bello-borizontinum A. Silv. in Arch. Mus. Nac. Rio Jan. 23:159, pl. 1. 921; M. Pichon, loc. cit. 1947. (T.: Silveira 570). Кс з longipetiolatum Kuhlm. in Anais Prim. Reun. Sul-Am. Bot. 3:86, /. 15. 1940. (T.: Kublmann 29793). Trees 3—10 m. tall, the branches relatively slender, thinly rimose. Leaves ob- lanceolate to broadly oval-obovate, acute to rounded at the tip, base acutely cuneate, rarely broadly obtuse, 3-10 cm. long, 1—6 cm. broad, firmly membra- naceous, glabrous; petioles 1-2 cm. long. Inflorescence subterminal at one or more of the upper nodes, 2-5 cm. long, distinctly pedunculate, very inconspicuously yellow-puberulent to essentially glabrous; pedicels 1-2 mm. long. Calyx lobes ovate to ovate-lanceolate, acute to obtuse, 1-2 mm. long, inconspicuously pilosulose to essentially glabrous. Corolla greenish or yellowish, finely pilosulose or papillate without, the tube 5.0—5.5 mm. long and about 1.5 mm. in diameter at the insertion of the stamens, the lobes ovate, acute to obtuse, 1.5—2.0 mm. long. Stamens in- serted at about the upper third within the corolla tube, the anthers 1-2 mm. long. Ovary ovoid, about 1 mm. long, sparsely pilosulose, the stigma fusiform, about 1 mm. long. Follicles elliptic-pyriform, the body 4-6 cm. long and 2.5-3.0 cm. broad, with a more or less definite midrib, brown, gradually narrowed to a stipe 1.0-1.5 cm. long; seeds oval, 3-4 cm. long and about 2.5 cm. broad. Southern Brazil; in woodlands (restingas); flowering from October to December. Vernacular names (Brazil): Guatambu (Serviço Florestal, Magalhães, Malme) ; Pequía da restinga (Serviço Florestal); Amarellao (Ducke); Pequía (Glaziou). AZIL: ACRE: Rio Acre, Ducke 207 (US). MiNAs GERAIS: Caldas, Nova Ponte, | : P R, U, US); Serra do Cipó, Duarte үй (М PARAÍBA: Areia, Vasconcellos 243 (МО). RIO DE JANEIRO: Тејисо, Pohl 4315 en Corcovado, Gardner 5542 (К), Ducke & Kuhlmann 15387 (MO); Serra u orgaos, Barbosa s. (MO); Praia do Pinto, Con- stantino 2170 (U); horto о cultum, ао 7825 (0, US); Recreo dos Bandeirantes, Lutz 547 (Е, R, US); Mundo Novo, Kuhlmann 15346 (U, US); restinga da Copacabana, Glaziou pH (C), Raben s.n. (C); montem Dois Crimios, Warming s. n. (C); Ipanema, Riedel 2771 (W), Serviço Florestal 100 (MO); environs de Rio-Janeiro, Weddel 445 (P, W); Horto Florestal 108 (MO); St. Louis, Glaziou 4079 (C, F, P); Alto Macahé de N. Friburgo, Glaziou 18364 (C, P), 19630 (C, P); Carahy, ngle s. n. (С); Corcovado, Ducke & Kuhlmann 15387 „ aM oo Itajai Muller 121 (R). s&o PAULO: Jardim Botanico, Hoehne 28544 (F, GH, MO); locality lacking, Lund s. n. (C). LACKING: Glaziou 636 (C), 637 tis 1586 (C), 2023 (C), 5933 (C), 18365 (С); Mn n. (U, W); Riedel s. n. (GH W). This species is a bit difficult to distinguish from A. ih: at times, but usually may be told by the nearly glabrous inflorescence which appears to be rather flat-topped, while it seems to be almost spherical in A. australe. The fruits, of [Vor. 38 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN course, are quite different. I think it possible that A. pyricollum may hybridize with A. australe to the south and with A. Ulei to the north, while the plants usually referred to A. Sellowii show some indications of A. parvifolium. 12. AsPIDOSPERMA Vancasu A.DC. in DC. Prodr. 8:399. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (Т.: Vargas s. n.!). Маса а Vargasii (A.DC.) О. Ktze. Rev. Gen. 2:416. 1891. Trees 3-20 m. tall, the branches relatively slender, with close thin bark. Leaves elliptic to obovate, shortly acuminate, obtusely to acutely cuneate, 5-12 cm. long, 3-6 cm. broad, firmly membranaceous, glabrous; petioles 1.0—1.5 cm. long. Inflorescences clustered subterminally at the uppermost nodes, densely flow- ered, densely puberulent, 2-5 cm. long; pedicels about 1 mm. long. Calyx lobes ovate, acute, about 2 mm. long, minutely pilosulose. Corolla white, densely pilosulose without, the tube 3.5—4.0 mm. long, about 1 mm. in diameter at the insertion of the stamens, the lobes ovate, about 1.5 mm. long. Stamens inserted in the upper third of the corolla tube, the anthers about 1 mm. long. Ovary ovoid, about 1 mm. long, densely puberulent; stigma ovoid, about 0.5 mm. long. Follicles pyriform, about 4—5 cm. long and 2.5—3.0 cm. broad, gradually narrowed to a stipe 0.5-1.0 cm. long, with a definite midrib; seeds ovate, about 4 cm. long and 2.5 cm. broad. Venezuela and adjacent Colombia and Guiana; rocky arid slopes and transition forest; flowering from June to September. Vernacular names: Amarillo (Venezuela—Curran & Haman); Yema de huevo (Venezuela—Steyermark, Ll. Williams); Walababadan (Surinam—Boschwezen) ; Quillo bordón (Peru—Woytkowski). COLOMBIA: data — уне 5216 (US). VENEZUELA: ARAGU Parque Nacional, LI. баш 10125 (F). BOLIVAR: La Prision, Medio Caura, ШІ. "Williams II712 (F, K, MO, US); Guayapo, Bajo rie? E Williams 11820 (F, MO, US). DISTRITO FEDERAL: Antímano, РИ Шек 13381 (F, МО, 51 (GH, US), 8605 (GH, US), 10380 (GH, NY, US), 11888 (F, G, MO, NY, US), se 194 (F), Rose 21912 (GH, US), Fendler 1200 (GH—in part, К); Caracas, SUCRE: Cerro Imposible, Steyermark 62023 (MO, US); Río Guagua, Steyermark 62786 (МО). тасніка: San Cristóbal, Archer 3201 (US). a м: Sectie О, Wood Herbarium, Surinam 323 (K, U). U: HUANUCO: Shapajilla, Woytkowski 26 (F, MO). SAN MARTIN: Tarapoto, Li. on 6231 (F, MO, US). The timber is said to be rather hard, yellow, and with a bitter taste. Possible hybridization of A. Vargasii and A. Ulei is discussed briefly under the latter species. 13. AsPrmbosPERMA ULEI Mgf. in Notizblatt 9:78. 1924. (Т. Ule 8453, photo. O!). Aspidosperma occidentale Mgf. loc. cit. 15:133. 1940, поп Malme. (T.: Ule 0700!). Trees 3—40 т. tall, with relatively slender, thinly rimose branches. Leaves obovate to obovate-elliptic, obtuse or rounded to acute, base narrowly or broadly 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 155 Fig. 6. Aspidosperma Ulei Mgf.: Flowering branch, entire and dissected flowers, entire and split fruit showing seed. cuneate, 4-10 cm. long, 2—5 cm. broad, membranaceous, glabrous above, densely yellow-puberulent to glabrate beneath; petioles 1-2 cm. long. Inflorescences one to several at the upper nodes, densely yellow-puberulent, 3—5 cm. long, relatively lax and long-pedunculate; pedicels about 1 mm. long. Calyx lobes ovate, acute, 1-2 mm. long, minutely yellow-puberulent. Corolla greenish white to yellow, densely yellow-puberulent without, the tube about 3 mm. long and 1.5 mm. in diameter, the lobes ovate, about 1 mm. long. Stamens inserted at the upper third within the corolla tube, the anthers somewhat less than 1 mm. long. Ovary ovoid, about 1 mm. long, densely and minutely puberulent, the stigma ovoid, less than 0.5 mm. long. Follicles pyriform, about 4 cm. long and 3—4 cm. broad, rather abruptly narrowed to a stipe about 1 cm. long, with a prominently elevated midrib; seeds ovate, about 4 cm. long and 3 cm. broad. Northern Brazil and adjacent Venezuela; in mixed low bush and transitional forest; flowering from October to June. [Vor. 38 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN NEZUELA: GUARICO: between Ortiz and Guarico Bridge, Pittier 12225 (NY, US); near sein Para, Archer 3022 (US). British Guiana: Kumuparu, Demerara River, Forest Dept. 2536 (К); Kanuku Mountains, Takutu River, A. С. Smith 3151 (MO, U, US), 3389 (MO, U, US). IL: ACRE: Rio Acre, Ule 9700 T; Poser ONAS: Surum a Branco, Ule 8451 (С, U); Serra Grande, Rio Branco, Ducke 7 (U, 4 Bafa: Bonfim, Curran 148 (GH, MO, US). PERNAMBUCO: Tapera, Pickel 5. (F, GH). Markgraf cited both Ule 8452 and 8453; the latter was photographed by Mac- bride and prints are distributed in many herbaria. As far as I can learn, both sheets were lost in the destruction of the herbarium at Berlin-Dahlem, and I have been unable to find duplicates. However, Ule 8451 was collected at the same locality and on the same date as the type specimens and corresponds very closely with the photograph of Ule 8453 Aspidosperma Ulei appears to hybridize with both A. Vargasii and A. pyri- collum, to judge from our meagre herbarium representation: the specimens from British Guiana tend to glabrate foliage, and the fruits from Venezuela have in- definite midribs; the small, obovate-oblong leaves of the specimen from Pernambuco are very suggestive of those of pyricollum, although the fragments of inflorescence show the characteristically dense, yellow pubescence of A. Ulei I feel myself fortunate in having discovered the isotype of A. occidentale Mgf. non Malme (Ule 9700) amongst the undetermined specimens from Kew. It is flowering, and almost unquestionably referable to A. Ule Mgf. Before finding this specimen, the only one which I had to represent A. occidentale was Krukoff 5470, cited by Markgraf as a paratype and seen by me amongst the specimens at Geneva, Stockholm, and Washington. The difficulty lay in the fact that the specimen was in fruit, and that Markgraf did not describe fruit for A. occidentale. Furthermore, the fruits are of a type which is unfamiliar to me otherwise: it is strongly unequal-biconvex, about 5 cm. long and 4 cm. broad, apparently sessile, quite woody, dark brown with prominent lenticels. The whole suggests the fruits of A. ramiflorum to me, although it is not that species. It most definitely is not the fruit of A. Ulei, and may represent a new species, perhaps a second for the RAMIFLORA. However, this study has impressed upon me the dangers of describing new species of Aspidosperma from fruit alone, and I shall forbear in this case. 14. АвршоврЕкмл reductum (Hassl.) Woods., stat. nov. Aspidosperma Riedelii Muell. Arg. ssp. reductum Hassl. in Fedde, Керегі. 12:262. 1913. (T.: Fiebrig 3371). Small tree with relatively slender, gnarled, loosely rimose branches. Leaves obovate-spatulate, apex broadly rounded, narrowly cuneate to a subpetiolar base, 1.5-2.5 cm. long, 4-9 mm. broad, wholly glabrous. Flowers borne singly in the axils of the uppermost leaves; pedicels 2-3 mm. long, glabrous. Calyx lobes oblong-spatulate, obtuse or rounded, 2.5-3.0 mm. long, glabrous. Corolla appar- 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 157 ently greenish-white, glabrous or indefinitely papillate without, the tube about 3 mm. long, about 1 mm. in diameter at the insertion of the stamens, the lobes obovate-oblong, about 1.5 mm. long. Stamens inserted at about the upper third within the corolla tube, the anthers somewhat less than 1 mm. long. Ovary ovoid, about 0.5 mm. long, minutely sericeous, the stigma about 0.25 mm. long. PARAGUAY: Cordillera de Altos, Fiebrig 337 (F, G). Like many other small-leaved specimens of the series PyRICOLLA, particularly if sterile, this plant was associated with A. Riedelii by Dr. Hassler. The peculiar calyx lobes of this plant, however, are quite unlike those of any other of the genus which I have examined. A close parallel is found in the uniflorous A. oligantbum of northern Brazil, but the flowers of the two species are quite different. Series ТҮ. PoLyNEURA Woodson, n. ser. Ser. Macrolobii K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?:141. 1895, in part; Ser. E olobii К. Sch. loc. cit. 142. 1895, in part; $ Tetrasticha M. Pichon, in Bull. Mus. t. Hist. ser. 2, 19:363. 1947, in part; $ Reticulata M. Pichon, loc. cit. 366. 1947. Trees or shrubs with rather scanty colorless or milky latex, apparently ever- green, with rather close, thin, conspicuously lenticellate bark. Branches with naked or scaly (A. polyneuron), acuminate buds, the subtending internode defi- nitely thicker (or scarcely thicker in A. cuspa) than the terminal petiole, with (A. polyneuron) or without definite seasonal articulations. Leaves alternate, firmly membranaceous, rather small to moderately large, with prominent reticulate venation upon both surfaces, the secondary veins widely arcuate. Inflorescence terminal or subterminal, thyrsiform to cymiform, with persistent but irregularly disposed and frequently obscure bracts; corolla tubular or tubular-salverform, rather coriaceous, glabrous without (but more or less puberulent-papillate toward the upper tube and lobes in A. polyneuron and A. dispermum) ; ovary essentially terete. Follicles falciform to subcylindric, somewhat concave- to somewhat bi- convex, the placenta rotating about 45°-135°, sessile, essentially glabrous, con- spicuously lenticellate; seeds with a narrow, strongly excentric basal wing (con- centric in А. dispermum). Species 15-18 Figure 7. KEY TO THE SPECIES a. Inflorescences clustered subterminally in the uppermost leaf axils, dichasial, the br ich an ence or caducous, the flowers definitely s, pe el seed an excentric or concentric wing, the embryo ела е b. Coro m ore or r les puberulent Те the aes about half as long as th Ам inflorescence more or less densely erulent. s orescence Chev lax, dure pube ns pros ре has the r Sardis tertiary venation sed; folli icles broly рне, not dU lenticellate, de "dms ре л а wing ; 2. Ек Әкені dense, gray- Қара: эл. eii go to oblong-elliptic, the reticulate tertiary vena nent on both ee follicles clavate-oblong, ED n lenti- cellate, the seeds with an elongate, obtuse, basal w Гея 5. А. DISPERMUM о о 16. А. POLYNEURON [Vor. 38 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN bb. Corolla glabrous without, the lobes about twice as long as the tube; inflorescence very lax, glabrous or rarely — puberulent; leaves ovate-elliptic, the her. me tertiary venation emersed; follicles stout- ly c аулада тый» егу conspicuously Ша. the seeds with a short, acute, basal wing 17. A. CYLINDROCARPON aa. Inflorescence aos, usually terminal, monochasial, the bra per- sistent, relatively conspicuous, the flowers sessile; follicles Pion al obviou sly ndi the seeds with an excentric, elongate, obtuse, basal wing, the embryo bilaterally asymmetric 18. A. CUSPA 15. ASPIDOSPERMA DISPERMUM Muell. Arg. in Mart. Fl. Bras. 6':60. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Riedel rogr!). Macaglia disperma (Muell. Arg.) O. Ktze. Rev. Gen. 2:416. 1891. Trees about 6 m. tall, the branches relatively slender, minutely puberulent when young, glabrate and thinly rimose at maturity. Leaves broadly elliptic to ovate-elliptic, the apex abruptly and shortly acuminate to obtuse, the base broadly obtuse, 5-12 cm. long, 2-7 cm. broad, firmly membranaceous, the reticulate tertiary venation emersed, glabrous; petioles 2-3 cm. long. Inflorescences clustered subterminally in the uppermost leaf axils, dichotomously dichasial, 2-5 cm. long, the peduncles and pedicels densely white-puberulent, the bracts very minute; pedicels about 1 mm. long. Calyx lobes broadly ovate-trigonal, acute to obtuse, about 0.5 mm. long, densely white-puberulent. Corolla greenish, the tube about 2.5 mm. long and 1 mm. in diameter, glabrous without, the lobes oblong, rounded, about 1.5 mm. long, sparsely appressed-pilosulose without. Stamens inserted some- what above midway within the corolla tube, the anthers about 0.7 mm. long. Ovary ovoid, glabrous, about 0.5 mm. long, the stigma subcapitate. Follicles broadly elliptic, acuminate, 2-3 cm. long, 1-2 cm. broad, sessile, with a prominent midrib, not obviously lenticellate; seeds 2, oval, about 2 cm. long and 1.0-1.5 cm. broad, with a narrow concentric wing and a bilaterally symmetric embryo. Southeastern Brazil on rocky hillsides and carrascos. Vernacular name: Pereiro da serra (Mello Barreto). RAZIL: MINAS GERAIS: Serra da Lappa, Riedel 1001 (G, NY, U, W); Diamantina, ibi Barreto 0861 (F). Although transitional to the species of series PyrIcoLLA іп the concentrically winged seeds, the reduction of the seed number to two is the most advanced of that tendency in the series PoLYNEURA. It is odd that this species has not been collected more frequently, since Mello Barreto reports that it is very abundant in the municipality of Diamantina. Nevertheless, A. dispermum must be of relatively limited distribution; specimens from eastern Peru which have been referred to it clearly represent A. polyneuron. 16. AsPIDOSPERMA POLYNEURON Muell. Arg. in Mart. Fl. Bras. 6!:57. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.:Riedel 2332!). pua Peroba All. ex Sald. Config. & Descr. Madeiras Rio Jan. 9, 104. 1865. (T.: Beaurepaire s. n.! Aspidosperma venosum Muell. Arg. in Kjoeb. Vidensk. Meddel. 103. 1869. (T.: Warming $. %.1). 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 159 Thyroma polyneura (Muell. e ite Apoc. So. Am. 24. 1878. E polyneuron Muell. Arg. var. genuinum Hassl. in Fedde, Кереге. 12:260. Aspidosperma polyneuron Muell. Arg. var. longifolium Hassl. loc. cit. 1913. (T.: Hassler 104064!). Aspidosperma Dugandii Standl. in Trop. Woods 36:15. 1933. (T.: Dugand 355!). Trees 8—20 m. tall, the trunk as much as 80 cm. thick, the branches relatively slender, glabrous, thinly rimose. Leaves oblong- to obovate-elliptic, shortly and abruptly acuminate to obtuse or rounded at the apex, base acutely cuneate to obtuse, 4—12 cm. long, 1—4 cm. broad, firmly membranaceous, the reticulate tertiary venation very prominent on both surfaces, glabrous; petioles 1.0—1.5 cm. long or somewhat less. Inflorescences clustered subterminally in the uppermost leaf axils, dichasial, relatively dense, gray-puberulent, 1—3 cm. long, the pedicels about 1 mm. long, the bracts minute. Calyx lobes broadly ovate, acute to rounded, 0.5-1.0 mm. long, appressed-pilosulose without. Corolla whitish or yellowish, densely appressed-puberulent to nearly glabrous without, the tube 2.5-3.0 mm. long, about 1 mm. broad at the insertion of the stamens, the lobes ovate, 0.5-1.5 mm. long. Stamens inserted somewhat above midway within the corolla tube, the anthers about 0.5 mm. long. Ovary ovoid, about 0.5 mm. long, densely puberu- lent. Follicles clavate-oblong, 3-6 cm. long and 1.0-1.5 cm. broad, very con- spicuously lenticellate, the seeds 2—3.5 cm. long, the obtuse, basal wing about as long as the radially symmetric cotyledons. Northern Colombia to Paraguay and northern Argentina and eastern Peru, in forests and thickets from approximately 25 to 1000 m. elev.; flowering from February to June in the north and from November to March in the south. Vernacular names: Carreto (Colombia—Dugand); Comuld or Сити (Colombia—Killip et al.) ; Paroba and Paroba mirim (Brazil—Warming) ; Peroba rosa (Brazil—Whitford & Silveira); Perobinba (Brazil—Whitford & Silveira); Peroba and Palo rosa (Argentina—Denis). COLOMBIA: ATLANTICO: Luruaco, Dugand 582 lat pe I 1507 (Е, va near Barranquilla, Dugand 428 (F); Arroyo de Сайа, Dugand 9 Jaramillo 2700 (COL), Dugand 3 255 (F, US), 208 (F); alrededores de Galapa y i jane nd & Jeremillo 3442 BOLIVAR: Rio Sinu, Verken © нені 5,7. . CUNDINAMARCA: (C Tocaima, Spo 4689 (COL, US); Chucunda-Tocaima, Garcia 3083 (COL, US); east of Apulo, along trail to Anapoima, Killip, Dugand & rmt 36131 (COL, US) ; Hacienda El Cucharo, between Tocaima and Pubenza, Killip, Dugand & Jaramillo 38280 (COL, MO, US), Jaramillo 9 Mejia 233 (COL). MAGDALENA: Tocaima, Triana s.n. (COL); near Fonseca, Haught 4007 (COL, US); Procedencia ттн, Nunez Bossio 541 (COL); Río Tucurinca, Dugand 1024 (COL, F); Don Jaca, Santa Marta 325 Record бб (F). BRAZIL: ESPIRITO SANTO: dro. W hit ford B ‘Silveira 62 (Е, GH, US). MINAS GERAIS: Lagoa Santa, Warming s (С, МОУ age lacking, Aces 870 (F, NY). PARANA: curis W hitford 4 Silvela 125 (F, СН, US); Patrimonio, Dusén 16781 (GH, мо NY, US); Jaguariahyva, Dusén s.n. (F, MO); Volta Grande, Dusén s.m. (СН). RIO DE JANEIRO: Morro Azul, Riedel 2332 (F, GH, NY, P, U, W); Floresta da Upica, Glaz іои 11180 (C, P); Botsilical Gardens, Whitford 12 (GH, US); Mundo Novo, Kuhlmann 15344 (U, US); Matta da Fabrica Alianca, Servico Florestal 103 (MO) ;Avelar, Мийез 0 (R). SAO PAULO: Jardim Botanico, Hoehne 28669 (NY); Rio Claro, Löfgren 673 (C); Sao Paulo, Beaurepaire s. n. (R); locality lacking, Löfgren 32 (C), Lund s.m. [Vor. 38 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN PARAGUAY: Sierra de Amambay, Hassler 10408 7 Me ends (W). ARGENTINA: MISIONES: Puerto Bertoni, Denis 357 Peru: CAJAMARCA: Prov. Jaen, near Las nbi SM 7115 (СН, US). Professor Dugand, who bas been particularly interested in this species, reports that it produces excellent structural timber, the heartwood being handsome pink or orange in color and the sapwood dull gray-white. From the reports of collectors, the trees appear to be evergreen, with rather scanty, watery sap and not milky. The leaves of specimens currently assigned to A. Dugandii appear to be somewhat larger than strictly typical A. polyneuron, and the corolla lobes may average a bit longer, but I do not consider these slight differences to be of specific importance. 17. ASPIDOSPERMA CYLINDROCARPON Muell. Arg. in Mart. Fl. Bras. 61:54. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:366. 1947. (T.: Sello s. n.!). ы-і еден i lagoense Muell. Arg. in Kjoeb. Vidensk. Meddel. 102. 1869. (T.: Warming Арса brevifolia Rusby, in Bull. М. Y. Bot. Gard. 8:113. 1912. (Т.: К. S. Williams Aspiospera cylindrocarpon Muell. Arg. var. genuinum Hassl. in Fedde, Кереге. 12:260. Aspidosperma емее Muell. Arg. var. longepetiolatum Hassl. loc. cit. 261. 1913. ы cylindrocarpon Muell. Arg. var. macrophyllum Hassl. loc. cit. 1913. (T.: Hassler 11 Trees 4—12 m. tall, the branches relatively slender, glabrous or irregularly puberulent when very young, thinly rimose at maturity. Leaves ovate- to lance- elliptic, acutely acuminate to obtuse, base obtuse to acute, 5-12 cm. long, 1.5-6.0 cm. broad, firmly membranaceous, lustrous, the reticulate tertiary venation emersed, glabrous; petioles 2.0—2.5 cm. long. Inflorescences clustered subterminally in the uppermost leaf axils, dichotomously dichasial, very lax, 2—7 cm. long, glabrous or rarely irregularly puberulent, the pedicels 1-3 mm. long, the bracts minutely lanceolate, caducous. Calyx lobes ovate, acute, 1 mm. long, glabrous or minutely ciliolate. Corolla white, glabrous or rarely indefinitely puberulent-papillate with- out, the tube 2-3 mm. long, about 1 mm. wide at the insertion of the stamens, the lobes narrowly oblong, 5—8 mm. long. Stamens inserted about midway within the corolla tube, the anthers about 0.5 mm. long. Ovary ovoid, about 0.5 mm. long, glabrous, the stigma subglobose. Follicles stoutly clavate-oblong, 6-8 cm. long and 2.0—2.5 cm. broad, very conspicuously lenticellate, the seeds 3—5 cm. long, the acute basal wing somewhat shorter than the bilaterally symmetric coty- ledons or about as long. Southern Brazil and adjacent Paraguay and Bolivia, in woodlands and balcony forests of plains; flowering from September to November. Vernacular name: Carapanbuba (Brazil—Ducke). BRAZIL: MATO GROSSO: Caceres, Hoehne 4444 (MO), 5034 (R). MINAS GERA entre Sitio & Barbacena, Glaziou 12048 (C, P); Arcos, Oliveira 211 (MO); Bello Hori. 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 161 Fig. 7. Coe D Muell. Arg. (upper figures): Flowering branch, entire and diss ssected flowers; Aspidosperma cuspa (HBK.) S. F. Blake (lower figures): Flowering branch, entire and split fruit showing see onte, Magalhães 620 (MO); Lagóa Santa, Warming s. n. (C, Е, W); Santa Luzia, Mello Bare 3189 (F); Caldas, Regnell 870 bis (C, F, GH, K, NY, R, U, US); locality lacking, Regne ell s.n. (F), Sello s. n. (P, ХУ). PARANA: Jaguariahyva, ‘ad Ton. i Men ds usén 16071 (F, GH, MO, NY, US), ibid. in campo rupestre, Dusén s. n. (F, O). E JANIERO: Aldeia de São Pedro, Glaziou 12052 (C, K, MO, P, R); m сае аа Aldeia, Ule s.n. (R). são PAULO: Loreto, Vecchi 264 (R). Pataca Sierra de Amambay, Hassler [Rojas] 10500 (MO, W). VIA: SANTA CRUZ: Tarochito, Steinbach 8144 (К, U); Río San Juan, R. S. M 9 255 (NY). Although the inflorescence of this species typically is glabrous, occasional pubescence, combined with somewhat shortening of the corolla lobes, might be [Vor. 38 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN interpreted as evidence of occasional hybridization with A. polyneuron. This is particularly noticeable in the Bolivian specimen cited above. 18. ASPIDOSPERMA cUsPA (НВК.) S. F. Blake, ex Pittier, Man. Pl. Us. Venez. 110. 1926. Conoria ? Cuspa HBK. Nov. Gen. & Sp. 7:242. 1825. (T.: Humboldt & Bonpland s. n., hoto. GH!). Alsodeia Cuspa (HBK.) Spreng. n 4:Cur. Post. 99. 1827. €— ны Lbotzkianum Muell. Arg. in Mart. Fl. Bras. 61:60. 1860; M. Js in l. Mus. Nat. Hist. Nat. II, 19:366. 1947. (Т.: Lhotzky s.n., photo. MO! Tos scia румен Muell. Arg. in Linnaea 30:398. 1860; M. Pichon, loc. cit. 1947. (T.: Spruce 3617!). Aspidosperma sessiliflorum Muell. Arg. loc. cit. 399. 1860. E nin Fl. Trin. 53!). T byroma sessiliflorum (Muell. Arg.) Miers, Apoc. So. Am. 23. Thyroma decipiens (Muell. Arg.) Miers, loc. cit. 24. teli Thyroma Lhotzkiana (Muell. Arg.) Miers, loc. cit. 25. 1878. Macaglia decipiens (Muell. Arg.) O. Ktze. Rev. Gen. 2:416. 1891. Aspidosperma domingense Urb. Symb. Ant. 5:460. 1908. (T.: Eggers 2340). Aspidosperma lucentinervium S. F. Blake, in Contr. Gray Herb. n. s. 53:46. 1918. (T.: Curran aman 970!). ко elliptica Rusby, Descr. So. Am. Pl. 82. 1920; M. Pichon, loc. cit. 1947. 1). Aspidosperma Lbotzkianum Muell. Arg. var. hypoplasium Malme, in Arkiv Bot. 21A°:11. (T.: Malme 2745!). Жж абын Woronovii Standl. in Field Mus. Publ. Bot. 8:34. 1930. (T.: Woronov ! Shrubs or trees 3—8 m. tall, the branches relatively slender, glabrous, or in- frequently densely puberulent, thinly rimose at maturity. Leaves narrowly oblong- elliptic to broadly oval or ovate, apex obtuse or rounded, base broadly obtuse or rounded, 2-13 cm. long, 0.8-8.0 cm. broad, firmly membranaceous, yellowish- green, above glabrous, opaque or lustrous, beneath more or less glaucous, glabrous or infrequently more or less densely puberulent, the tertiary reticulate venation very prominent on both surfaces; petioles 2-9 mm. long. Inflorescences solitary, monochasial and rather thyrsiform, densely papillate or puberulent, 1-3 cm. long, terminal or less frequently apparently lateral and axillary or extra-axillary, the flowers sessile, the bracts 1-3 mm. long or less, persistent. Calyx lobes broadly ovate, obtuse, about 1 mm. long, papillate or minutely puberulent. Corolla greenish yellow or white, glabrous without, the tube 2.5—3.0 mm. long, about 1.5 mm. in diameter at the insertion of the stamens, the lobes broadly ovate, obtuse, about 1.0-1.5 mm. long. Stamens inserted somewhat above midway within the corolla tube, the anthers about 1 mm. long. Ovary ovoid, about 0.5 mm. long, glabrous, the stigma fusiform, about 0.3 mm. long. Follicles broadly or narrowly subreni- form, acute or rounded at the tip, with a prominent midrib, 2—4 cm. long and 1-2 cm. broad, obviously lenticellate, glabrous or minutely papillate; seeds 2.0-3.5 cm. long and 1.0—1.5 cm. broad, the obtuse basal wing about as long as the bilat- erally asymmetric cotyledons. 1951] WOODSON-—STUDIES IN THE АРОСҮМАСЕАЕ. УШ 163 Colombia to southern Brazil; eastern Ecuador; Hispaniola and Trinidad. In arid thickets, light woodlands, and savannas; flowering intermittently throughout the year. Vernacular names: Carreto, Amargo (Colombia—Dugand); Vara de piedra (Colombia—Bro. Elias); Cuspa (Venezuela—Humboldt, Steyermark); Cuspa negra (Venezuela—Steyermark) ; Amargoso (Venezuela—Curran & Haman). Harri: vicinity of Jean Rabel, Leonard & Leonard 12646 (MO, US); vicinity of Mole St. Nicolas, Leonard & Leonard 13351 (GH, NY, US); Massif des Mathieux, Ekman 6643 (US), 088 (US). INIDAD: Pointe Gourde, Britton & Broadway 2648 (GH, NY, US); Gasparee Island, Broadway 9426 (0); Camaronaro, Danouse 6070 (MO, NY, US); locality lacking, Sieber 53 COLOMBIA: ATLANTICO: Barranquilla, Elias E dd (F, MO), Dugand 37 (F), 3138 (COL); Puerto Colombia Dugand 632 (F, NY), Dugand & Jaramillo 3229 (COL, US). CUNDINAMA Apulo, trail to Ж Killip, Dugand 9 Jaramillo 38169 7075 (F). MAGDALENA: Santa Marta, Н. Н. Smith 836 (F, GH, MO, NY, U, US), Espina 28 (F, MO), Record 66 bis (GH, m 81 (F); Bonda, Castañeda 135 (COL, МО); Municipio Baraya, Caicedo 12434 (COL). VENEZUELA: AMAZONAS: Maypures, Spruce 3617 (NY, W). ANZOATEGUI: Guanta, Curran © Haman 1211 (GH, NY, US). ARAGUA: Maracay, Pittier 11362 (G, GH, NY, US). сакавово: Valencia, Pittier 0042 (GH, NY, US), 8708 (GH, NY, US). DISTRITO FEDERAL: between La Guaira and Río Grande, Curran & Haman 970 (GH, NY); La api Curran & Haman 840 (US), 038 (US); со. Pittier 10214 (СН, NY, US). STATE UNKNOWN: Cabo Blanco, Curran & Hoven 907 (GH, US), 938 (GH), 040 (GH, NY, US), 954 (GH), 955 (GH, NY); Camburi Chiquito, Curran 9 Haman 840 (G NY, US), 923 (GH, US); Rio Caribe, Curran & Иа 1260 (СН, NY, US); Cristobal Colon Pa ash 62 (GH, NY, US), 062 (NY, BAÍA: locality AA E 09 (NY). craRÁ: locality lacking, Lan s $. n. ЧС), Allemáo 968 puget P,R). : Goiaz, Burchell 7348 (К). Mato к mba, Malme 2745 (G, S US), 522 "S. US); Cuyabá, Malme 2567 (NY, US). PARAHYBA: S. Gongalo, Leutzelburg 26050 (NY). ко n. JANEIRO: Cabo Frio, A RUN 11185 (C, P). Ecuapor: EL ORO: between Portovelo and Río Cabra, Steyermark 54085 (F, MO, Y. This is an extremely distinctive species which shows no indications of inter- gradation with its congeners, At the same time, it is extremely variable, par- ticularly in leaf size, color, and indument or lack of it. The most outstanding of these variants are A. Woronowii (densely puberulent), A. domingense (unusually small leaves of rather heavy texture), and A. Lbofzkiama (inflorescences axillary or extra-axillary some distance from the apical meristem). These variants inter- grade, however, and I do not believe that our present knowledge is sufficient even to recognize them as varieties. An interesting aspect of A. cusa is shown by the Luetzelburg specimen from Baía and the Burchell specimen from Goiaz, which show the juniper-like deforma- [Vor. 38 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN tion of the inflorescence so common in species of Series NiripA. Burchell explains this by the remark “‘insectis deformat”. 1 am inclined to interpret this abnormality as a physiological character supporting the morphological affinity of Series Porv- NEURA and Nita which I first assumed from morphological grounds. Series V. Кісіра Woodson, n. ser. $ Tetrasticha M. Pichon, in Bull. Mus. Nat. Hist. Nat. 1947, in part; $ Glabriflora M. Pichon, loc. cit. 1947, in part; $ on м. Же, loc. cit. 364. 1947, in part Tall trees, apparently evergreen, with rather close, thin, conspicuously lenti- cellate bark. Branches with naked, obtuse buds, the subtending internode some- what thicker than the terminal petiole, without definite seasonal articulations. Leaves alternate, firmly membranaceous, moderately large, with rather indistinct, widely arcuate secondary venation. Inflorescence subterminal, dichotomously cymose, with persistent but irregularly disposed and frequently obscure bracts; corolla salverform, the lobes reflexed but the tube not definitely constricted at the orifice, somewhat coriaceous, glabrous without; ovary polygonally angulate. Fol- licles very broadly dolabriform, nearly circular, the placenta rotating about 300^, sessile, smooth, with or without a prominent midrib, glabrous, inconspicuously lenticellate; seeds with a concentric, nearly circular wing. Species 19 Figure 8. 19. ASPIDOSPERMA RIGIDUM Rusby, in Mem. N. Y. Bot. Gard. 7:323. 1927 (as rigida). (Т. Rusby 503!). Aspidosperma ал fa tapa Mef. in Notizbl. 12:300. 1935. (T.: dita 8120!). Aspidosperma acreanum Mgf. loc. cit. 15:133. 1940. (T.: Ule 0701 енн laxiflorum Kuhlm. in Anais Prim. Reun. Sul-Am. lg 3: 88, £. 17. 1940. : Ducke 22438!). CBS ap жим ИВ Kuhlm. loc. cit. 1940. (Т.: Ducke 24577!). Trees 8—40 m. tall, the branches relatively slender, glabrous. Leaves ovate- to oblong-elliptic, apex subcaudate-acuminate, base broadly decurrent to the petiole, 6—15 cm. long, 2—6 cm. broad, firmly membranaceous, opaque, glabrous; petioles 0.5—1.5 cm. long. Inflorescence either terminal or lateral to the leafy branches, or both, 3-4 cm. long, dichotomously cymose, more or less appressed-pilosulose, in- conspicuously bracteate, the pedicels 2-3 mm. long. Calyx lobes ovate, acute to obtuse, 1.0-1.5 mm. long, minutely tomentellous without. Corolla greenish-white, wholly glabrous without, the tube 2—3 mm. long, the lobes reflexed, oblong-elliptic, 5-7 mm. long. Stamens inserted about midway within the corolla tube, the anthers somewhat less than 1 mm. long. Ovary oblongoid, glabrous, about 1 mm. long, the stigma minutely capitate. Follicles nearly circular, 3.0—4.5 cm. in diameter, sessile or very shortly stipitate, smooth, glabrous, with a very excentric midrib; seeds circular or broadly oval, about 4 cm. in diameter. 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 165 Fig. 8. Aspidosperma rigidum Rusby: Flowering branch, bud, dissected flower, and fruit. Eastern Bolivia and neighboring Brazil, in forests (varzea and ferra firma); flowering from July to October. Vernacular name: Carapanahuba (Brazil—Krukoff, Ducke). BRAZIL: AMAZONAS: near mouth of Rio Embira, Krukoff 5172 (G, MO, S, U, US); б Humayta, КтибоЙ 6224 (МО, U, US), 63 98 ы U, US); ме к» em frente als. Paulo de Olivença, Ducke 24577 (МО). E: near mou Macauhan, Krukoff 5642 (MO); Seringal Oriente, Каден 498 (US): Pier Monte Mo, Ule 0701 (К). pará: Rio Tapajos, Ducke 22438 (MO). Borivia: LA Paz: Bopi River valley, Rusby 503 (NY); Province of S. Yungas, basin of Río Bopi, Krukoff 10148 (MO, U, US), 10170 (MO, U, US). sANTA CRUZ: Jorochito, Steinbach 8129 (F, ОН, К, S). Krukoff reports that the timber of this species is of excellent quality. [Vor. 38 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN Series VI. Nta Woodson, n. ser. Ser. Macrolobii K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?:141. 1895, in an Microlobii К. Sch. loc. cit. 142. 1895, in part; $ Piliflora M. Pichon, in Bull. M Nat . Hist. Nat. II, 19:364. 1947, in part; $ Macrantha M. Pichon, loc. cit. 1947, i in part; $ Laevifolia M. Pichon, loc. cit. 365. 1947. Trees with milky latex and conspicuously sulcate or lamellate boles, apparently evergreen, with tight dense bark but usually conspicuously lenticellate. Branches with naked, acuminate buds, the subtending internode scarcely thicker than the terminal petiole and the bud thus appearing pseudo-lateral, without definite seasonal articulations and with the uppermost internodes not definitely shortened. Leaves alternate, usually more or less coriaceous, frequently revolute-auriculate at the base. Inflorescence terminal or subterminal, thyrsiform or cymiform, with per- sistent but irregularly disposed and more or less evident bracts; corolla tubular or vubular-salverform, rather coriaceous, densely sericeous without, the lobes erect or ascending at anthesis, the tube not constricted at the orifice; ovary glabrous or sericeous, polygonally angulate or sulcate; calyx lobes 5, abnormally 4, distinct or barely united at the base, equally or very strongly unequal. Follicles very broadly dolabriform, concave-convex to very unequally bi-convex, the placenta rotating about 2257-3007, broadly warty to spiny, sessile or very abruptly and shortly stipitate; seeds with a concentric circular wing. Species 20—33 Figure 9. KEY TO THE SPECIES a. Corolla about 1.0—2.5 cm. lon b. Corolla lobes 2— E times as long as the tube. c. Corolla about 2.5 . long; inflorescence conspi icuously у. densely brown- ы ы moderately coriaceous, the sec ec venation obvious 0. A. INUNDATUM cc. Corolla about 1.3 cm. long; inflorescence inconspicuously bracteate, finely sulphur- — leaves very edd coriaceous, the secondary venation almost completely immerse bb. Corolla lobes about as long as ming кз то — i as long or рни c. Corolla lobes oblong, about as lon tube; leaves oblon obovate-oblong, broadly ro unded ғы the Ped gray-papillate ani d. Leaves heavily келісе се ghly lustrous above; inflorescence ter бөлеуі согоПа 1.5-1. long dd. ендің ди naceous or г атаса opaque above; inflores- ipn: orolla about 1.2 cm. long ec. Candi lobes ovate, about half as pls as the tube or somewhat » » аза elliptic, acute to obtuse at the base, yellow- isnt мз N — . A. SCHULTESII N N . А. MEGAPHYLLUM N w . А. MYRISTICIFOLIUM 24. A. CARAPANAUBA аа. TN э 3 ra b. Inflorscenc eiut to the leafy shoo ves not conspicuously calles с at the base. : ag adn usually broadest at about the middle or below, obtuse or ded at the base, the secondary veins of about 14—20 t e. Leaves ovate- oblong-e ep teed acute to obtuse at the tip, дан ый ог slightly сыба above, microscopically рарШаге to essentially glabrous beneath; flower — 4 subtended by brscteole "follicles abruptly constricted to a short e HN 25. A. MARCGRAVIANUM ee. Leaves broadly oval or ov = rounded to emarginat = hi shly 4. н nsely velutinous-papillate ai flowers distinctly ik tp follicles sessile 26. A. EXCELSUM dd. Leaves usually "me dos the middle, narrowly cuneate at 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 167 ni па highly lustrous above, the secondary veins almost in- rable and closely crowded; flowers priet pedicellate......27. A. EBURNEUM Cc: Leaves conspicuously аети auriculate at the d. piu dichotomously cymose, the pedu du shorter than e subtending peti dole or scarcely longer; gea rather narrowly Е. elliptic; indument gray dd. Inflorescence prom the peduncles much longer than the btendin 28. A. OBLONGUM su e. Leaves usn elliptic to oval, 4—8 cm. long, broadly obtuse to rounded at the tip; calyx lobes very unequal; indument gray 29. A. DISCOLOR ee. Leaves кн ovate to lanceolate, 6—12 cm. long, acute to acuminat in tip; calyx lobes a to somewhat unequal; ment brow 30. A. SALGADENSE bb. ГЕ lateral to oe leafy shoo c. Leaves ine to е! elliptic, n tip obtuse to abruptly and shortly ac d. tetas not revo КЕ auriculate at the base; inflorescence Кт condensed, the primary peduncle much shorter than the tending petiole 31. A. NITIDUM dd. gm Е ант at the base; inflorescence rather lax, ig pu е about as long as the subtending petiolés x what 32. А. AURICULATUM cc. Leaves СТОР elliptic to Td np d gradually and acute- ly subcaudate-acuminate, revolute into a narrowly cuneate base...... 33. A. PICHONIANUM 20. AsPIDOSPERMA INUNDATUM Ducke, in Archiv. Jard. Bot. Rio Janeiro 3:245. 1922; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:364. 1947. (T.: Ducke I7195!). Aspidosperma acanthocarpum Mgf. in Notizbl. 14:128. 1938. (T.: Ducke 24569!). Trees of moderate height, the branches rather stout, densely ferruginous- tomentellous when young, becoming glabrate. Leaves ovate to broadly oblong- elliptic, apex broadly acute to shortly acuminate, base obtuse, 7-16 cm. long, -7 cm. broad, glabrous and somewhat lustrous above, densely and minutely brown-tomentellous beneath; petioles 1.5-2.0 cm. long. Inflorescence both termi- nal and lateral to the leafy branches, stout, corymbose-thyrsiform, 4—9 cm. long, conspicuously bracteate, densely brown-tomentellous, the pedicels about 2 mm. long. Calyx lobes broadly ovate, acute, 2.5—3.0 mm. long, densely and minutely brown-tomentellous without. Corolla white, densely tomentellous without, the tube about 9 mm. long and 2.5 mm. thick at the insertion of the stamens, the lobes narrowly lanceolate-elliptic, 1.5-1.6 cm. long. Stamens inserted somewhat above midway within the corolla tube, the anthers about 2 mm. long. Ovary ovoid, about 2 mm. long, densely tomentellous, the stigma narrowly cylindric. Follicles nearly circular, 3.5—4.0 cm. in diameter, with a short stout stipe about 0.5 cm. long, densely and stoutly spinose; seeds circular, 3—4 cm. in diameter. Amazon valley, in forests; flowering during August and September. Vernacular names: Carapanatiba and Maparand (Ducke). [Vor. 38 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN BRAZIL: PARA: Gurupa, silva ab Amazonum fluvio JM Ducke 17195 (P, U, US); ad orientem lacus Salgado prope flumen Trombetas, Ducke 21808 (P, US); — 2 xt 15814 (MO). AMazoNas: São Paulo de price Rio а "Ducke 24569 (U, US). It is 52. doubtful whether А. inundatum and А. acanthocarpum actually are conspecific. Ducke 24569, which bears stoutly spiny fruits, also bears stout inflorescences which clearly indicate the plant to be of this affinity; however, they are much too young to show floral structure. In describing A. inundatum, Ducke describes the fruit in detail, but does not mention them as being spiny; a fruit which accompanies Ducke 17105 in the United States National Herbarium is too old and badly decomposed for accurate analysis. 21. АѕрІроѕрРЕКМА Schultesii Woodson, spec. nov. Arbor alta; ramulis crassiusculis nigris juventate minute pilosulis tandem gla- bratis. Folia alternata longissime petiolata lamina elliptica vel oblongo-elliptica apice obtusiuscula basi et in petiolo late decurrente 12-15 cm. longa 5-6 cm. lata rigidissime coriacea supra illustri venis secundariis emersis subtus pallida et minute denseque sulfureo- vel griseo-tomentella; petiolis crassiusculis nigris ca. 4 cm. longis. Inflorescentiae terminales vel subterminales dichotome cymosae ca. 6 cm. longae minute sulfureo-puberulae; pedunculis graciliusculis; bracteis vix bene visis; pedi- cellis ca. 1 mm. longis. Calycis laciniae subaequales late ovatae acutae 1.5-2.0 mm. longae extus minute sulfureo-tomentellae. Corolla lutea extus plus minusve sul- fureo-pilosa; tubo ca. 3 mm. longo ca. 1.5 mm. diam.; lobis oblongo-lanceolatis ca. 10 mm. longi. Antherae ca. 0.6 mm. longae. Ovarium depresse sphaericum longitudinaliter sulcatum ca. 0.5 mm. longum dense tomentellum; stigmate ovoideo apiculato. Fructus ignoti. AZIL: AMAZONAS: path between headwaters of lra-Igarapé and headwaters of Кы Abit, pow of Rio Taraíra, July 4-6, 1948, R. E. Schultes 9 F. López 10178 (MO, HOLOTYPE). This is such an unusual species that it is indeed unfortunate that the fruit is not known. The flowering specimen somewhat suggests the series NoBILEs in the black, obscurely lenticellate branches, heavily coriaceous leaves, and somewhat fastigiate inflorescences. To judge from stains upon the leaves, I suspect that the latex may be red, as well; although Dr. Schultes states that it is white. 22. АвршовреЕкмл megaphyllum Woodson, spec. nov. Arbor ca. 15 m. alta; ramulis crassiusculis juventate dense ferrugineo-papillatis mox glabratis cortice saturate fusco firmo striato haud evidenter lenticellato. Folia magna rigide coriacea lamina late oblonga apice latissime obtusa vel rotundata basi late rotundata 15-25 cm. longa 9-15 cm. lata supra valde illustri subtus pallida et dense griseo-papillata; petiolo nigro glabro 1.5-2.0 cm. longo. Inflorescentia terminalis dichotome cymosa repetite ramosa ca. 7-8 cm. longa dense ferrugineo- papillata; pedunculis crassiusculis; bracteis inconspicuis; pedicellis ca. 2 mm. longis. Calycis laciniae 4 (vel 5?), valde imbricatae inaequales exteriores late subreni- formes rotundatae 4.0—4.5 mm. longae 5—6 mm. latae, interiores suborbiculares 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 169 ca. 4 mm. diam., omnes dense ferrugineo-papillatae. Corolla extus dense ferrugineo- tomentella; tubo ca. 7-8 mm. longo 3.5-4.0 mm. lato; lobis late oblongo-ellipticis ca. 8 mm. longis. Antherae ca. 2 mm. longae. Ovarium depresse subsphaericum ca. 1.5 mm. altum dense ferrugineo-hirtellum. Fructus ignoti. ACRE: near mouth of Rio Macauhan, tributary of Rio Yaco, on terra firma, No 24, 1933, Krukoff 5639 (K, MO, ноготүрЕ, NY, U). Although referred to Geissospermum excelsum. Kuhlm. (Aspidosperma cara- panauba M. Pichon) by Markgraf, and undoubtedly closely related to it, I believe that the two species are sufficiently distinct. 23. AsPIDOsPERMA myristicifolium (Mgf.) Woodson, comb. nov. UTE (2) myristicifolium Mgf. in Notizbl. 11:787. 1933. (T.: Ruiz 6 Pavon, д, elatum E. L. Little, in Jour. Wash. Acad. Sci. 38:105, fig. 10. 1948. (Т.: Little 6517!) Trees up to 35 m. tall, with sulcate trunks as much as 1 m. in diameter, the branches relatively slender, very minutely appressed brown-pilosulose when young, soon becoming glabrate and rather conspicuously lenticellate. Leaves obovate- oblong, apex obtuse to shortly acute, base rounded to subtruncate, 20—25 cm. long and 7-10 cm. broad, firmly membranaceous, essentially glabrous, opaque above, beneath paler and with the relatively distant secondary veins very prominent, the tertiary venation immersed; petioles rather stout, 1.0-1.5 cm. long. Inflorescence axillary, dichasial, many-flowered, brown-tomentellous. Calyx lobes orbicular, about 2 mm. long, brown-tomentellous. Corolla densely brown-tomentellous with- out, the tube about 6 mm. long and 2 mm. broad, the lobes oblong, about 6 mm. long and 2 mm. broad. Stamens inserted about midway within the corolla tube, the anthers somewhat more than 0.5 mm. long. Ovary globose, about 1 mm. long, densely white-pilosulose. Follicles very broadly dolabriform, subplano-convex, sessile, about 11-12 cm. long and broad, very densely brown-velutinous; seeds with a circular concentric wing about 10 cm. in diameter. Western Ecuador, in forests. Vernacular names: Naranjo (Ruiz & Pavon); Naranjo de monte (E. L. Little). Ecuapor: GUAYAS: Guayaquil, Ruiz & Pavon s.m. (MO). Los Rios: Pichilingue, E. L. Little 6517 (MO, US). The discovery of the fruits and seeds of this species by Dr. Little verifies Pichon's opinion of its systematic position (in Bull. Mus. Nat. Hist. Nat. П, 19:368. 1947). 24. ASPIDOSPERMA CARAPANAUBA M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1948. (Based on the succeeding:) коо excelsum Kuhlm. in Archiv. Inst. Biol. Veg. Rio Jan. 2:89, pl. 7. 1935, on Aspidosperma excelsum Benth. (T.: Ducke 24471!). [Vor. 38 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN Large trees, the branches relatively stout, dark brown to black, minutely ferruginous-papillate when young, becoming glabrate with a close bark without apparent lenticels. Leaves broadly elliptic to oblong-elliptic, obtuse to abruptly and shortly acuminate, base acute to obtuse, 10—20 cm. long, 4—8 cm. broad, coriaceous, above minutely yellow-papillate to glabrate and lustrous, densely and minutely yellow-tomentellous beneath; petioles 1.5-2.0 cm. long. Inflorescence lateral to the young leafy shoots, dichotomously cymose, 5—6 cm. long, densely yellow-tomentellous, the peduncles rather stout, the bracts inconspicuous, the pedi- cels about 1 mm. long. Calyx lobes 5, broadly ovate, broadly acute to obtuse, 2.5-3.0 mm. long, densely and minutely yellow-tomentellous. Corolla white, densely yellow-tomentellous without, the tube 7.5—8.0 mm. long, about 3 mm. broad, the lobes ovate, about 3 mm. long. Stamens inserted within the upper third of the corolla tube, the anthers about 1.3 mm. long. Ovary broadly oblong- ovoid, densely yellow-tomentellous, about 2 mm. long. Fruits unknown. Central Amazon valley, in forests of high land; flowering in September. Vernacular name: Carapanauba (Ducke). BRAZIL: AMAZONAS: circa lacum Uaicurapá, prope Parintins, Ducke 24471 (P, U). Ducke cites the type specimen as mo. 24491, apparently a typographical error. 25. AspiposPpeRMA Marcgravianum Woodson, spec. nov. Arbor magna 20-60 m. alta; ramulis graciliusculis glabris cortice fusco valde lenticellato. Folia ovato- vel oblongo-elliptica apice acuta vel obtusa basi late obtusa vel rotundata 6—14 cm. longa 2—5 cm. lata crasse membranacea supra opaca glabra subtus minutissime papillata vel glabra; petiolis 0.8—2.0 cm. longis. In- florescentiae terminales thyrsiformes 3—6 cm. longae minute puberulae; pedunculis graciliusculis; bracteis minimis; floribus sessilibus. Calycis laciniae inaequales ovatae vel late oblongo-ovatae obtusae 2-3 mm. longae extus minute tomentellae bracteolis minimis 2—3 subtendentibus. Corolla virescens extus minute appresso- tomentella; tubo ca. 5 mm. longo ca. 1.5 mm. lato; lobis ovatis acutis ca. 1.5 mm. longis. Antherae prope ostio corollae tubae insertae ca. 0.6 mm. longae. Ovarium ovoideum longitudinaliter sulcatum ca. 1 mm. longum glabrum; stigmate fusiformi ca. 1 mm. longo. Folliculi fere orbiculares breviter stipitati 4—6 cm. diam. valde verrucosi, stipite ca. 0.5—1.5 cm. longo; seminibus 4—6 cm. diam. Surinam and northern Brazil to eastern Bolivia; forests in terra firma; flowering from August to October. Vernacular names: Parelbout and Witte parelbout, Apukuita (Surinam—Bosch- wezen; Carapanauba (Brazil—Ducke, Krukoff) ; Cipoal (Brazil —K rukoff) . SURINAM: Kaboerie, Boschwezen 5060 (NY, U, US); Brownsberg, Boschwezen I75I (U); Sanderij F, van Niel 6224 (U), Boschwezen 1424 (U), Archer 2775 (US); Sectie О, Boschwezen 1354 (U), 6395 (U, US). BRAZIL: AMAZONAs: Municip. Humayta, near Livramento, Krukoff 6843 (MO, US), 7207 (К, MO); Municip. Sao Paulo de Olivenga, basin of Rio Solimoes, Krukoff 12150 (NY, US); Manáos, Ducke 1163 (МО, NY, US). Pará: Belém, Bosque Municipa! 49 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. ҮШ 171 Ducke 1263 (MO, ноготүрЕ, US); Haut Ariramba (Trombetas), Ducke 14900 (US); Gurupo, Ducke 17224 (US). BOLIVIA: LA PAZ: Province of Larecaja, Tuiri, e Mapiri, Krukoff 10833 (U, US), 10872 (MO, U, US); Tumupasa, Cárdenas 1980 (K, NY). The majority of these specimens have been referred to previously as A. nitidum. Not only the position, but also the relative size of the inflorescences, as well as the shape, size, and general aspect of the foliage, however, demonstrate that A. Marc- gravianum is abundantly distinct from A. nitidum The inflorescences of many of the specimens cited above have been deformed, perhaps by insects, into an enlarged witches-broom affair in which all the floral parts have become calyx-like: the whole resembling a small branch of juniper. This disease also was noted іп two specimens of А. сира. 26. ASPIDOSPERMA EXCELSUM Benth. in Hook. Jour. Bot. 3:245. 1841; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1947. (T.: Schomburgk 4661). Macaglia excelsa (Benth.) O. Ktze. Rev. Gen. 2:416. 1891. Trees 15—35 m. tall, the branches moderately stout, essentially glabrous, at maturity with dark, rather indefinitely lenticellate bark. Leaves broadly oval to ovate, the tip broadly rounded to emarginate, base broadly obtuse to rounded, 7—12 cm. long, 3.5-7 cm. broad, heavily coriaceous, glabrous and highly lustrous above, pale and densely velutinous-papillate beneath; petioles 1-2 cm. long. Inflorescence terminal, corymbi-thyrsiform, about 5 cm. long, inconspicuously puberulent- papillate, the peduncles rather stout, branching repeatedly, the bracts rather incon- spicuous, the pedicels 2-3 mm. long. Calyx lobes subequal, broadly ovate, obtuse to broadly acute, about 2 mm. long, densely tomentellous-papillate. Corolla greenish-white, tomentellous-papillate without, the tube about 3 mm. long and 1.5 mm. broad, the lobes ovate, acute, about 1.5 mm. long. Anthers about 0.6 mm. long. Ovary ovoid, sulcate, about 1 mm. long, glabrous. Follicles broadly oval strongly spinose-verrucose, 5.5-9.0 cm. long and 5—6 cm. broad, sessile. British Guiana and Surinam, in "various types of forest [Sandwith]"; flowering from June to September. Vernacular names: Paddlewood (British Guiana—Schomburgk, Archer) ; Yaruru (British Guiana—Tutin) ; Wit parelhout (Surinam—Boschwezen). Вкітізн GUIANA: Moraballi Creek near Bartica, Sandwith 321 (NY, U); Tuicama Suri, Laekie 2037 (F); upper Mazaruni River, Leng 270 (NY); Bartica-Potaro Road, Tutin 233 (US); Macouri Creek, Essequibo River, Archer 2483 (US); locality lacking, M 468 (W RINAM: fluv. РЕТ Stahel © Gonggrijp 944 (0). Schomburgk noted that the narrow buttresses of this species prevented use of the timber for most purposes except the making of paddles; Tutin and Sandwith note that this practice is continued to the present day. [Vor. 38 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN 27. ASPIDOSPERMA EBURNEUM F. Allem. ex Sald. in Ann. Sci. Nat. V, 19:213. 1874; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1947. (T.: Glaziou 6046!) . Aspidosperma pruinosum Mgf. in Notizbl. 15:133. 1940. (T.: Glaziou 12076!). Aspidosperma compactinervium Kuhlm. in Anais Prim. Reun. Sul-Am. Bot. 3:87, £. IÓ. 1940. (T.: Machado Nuñes 26072). Tall trees about 20 m. in height, the branches relatively slender, essentially glabrous, with reddish-brown, clearly lenticellate bark. Leaves narrowly elliptic to elliptic-obovate, apex shortly subcaudate-acuminate to acute or rounded, base rather narrowly cuneate, 6—8 cm. long, 1.5—3.5 cm. broad, moderately coriaceous, olive- green and highly lustrous above with almost innumerable, ascending, closely crowded secondary veins, paler and minutely papillate beneath; petioles about 1 mm. long. Inflorescence terminal, thyrsiform, 2—3 cm. long, inconspicuously appressed-pilosulose, the peduncles rather slender, the bracts inconspicuous, the pedicels about 1 mm. long. Calyx lobes more or less unequal, ovate, acute, about 2 mm. long, tomentellous-papillate. Corolla greenish-white, minutely tomentellous without, the tube about 4 mm. long and 1.5 mm. broad, the lobes oblong-ovate, about 3 mm. long. Anthers about 1 mm. long, inserted at about midway within the corolla tube. Ovary ovoid, sulcate, about 0.7 mm. long, glabrous. Follicles nearly circular, about 3.0—3.5 cm. in diameter, with a stipe about 0.5 cm. long, with rather low, widely spaced warts. Southeastern Brazil in forests; flowering from August to October. Vernacular names: Pequía-marfím (Glaziou, Saldanha); Pequía amarella (Saldanha) ; Peroba de gomo (Mello Barreto). BRAZIL: DISTRITO FEDERAL: Rio de Janeiro, Glaziou 638 (C, P). RIO DE JANEIRO: floresta de la Tijuca, Glaziou 6046 (C, P), 12076 (C, P). MiNAs GERAIS: Belo Horizonte, Mello Barreto 10005 (F); Avellar, Kublmann 37745 (MO). Saldanha cited no specimens in describing A. eburneum, but gave the locality as "montagne de Tijuco". The original label of Glaziou 6046 bears the pencilled note “Tijuco” and the date 1870, and for this reason I have chosen that specimen as the provisional lectotype. 28. AsPIDOSPERMA OBLONGUM A.DC. in DC. Prodr. 8:399. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1947. (T.: Martin s. n.!). Macaglia oblonga (A.DC.) O. Ktze. Rev. Gen. 2:416. 1891. Aspidosperma Kublmannii Mgf. in Notizbl. 12:555. 1935. (T.: Kublmann 22453). Trees as much as 35 m. tall, with rather slender branches closely yellowish-gray puberulent when young, soon becoming glabrate with a close black, conspicuously lenticellate bark. Leaves narrowly oblong to rather broadly elliptic-oblong, apex acute to rather shortly and obtusely acuminate, base subtruncate and conspicuously revolute-auriculate, 7-14 cm. long and 2—4 cm. broad, firmly membranaceous, above opaque to somewhat lustrous, glabrous or glabrate at maturity, beneath pale 1951] WOODSON—STUDIES IN THE APOCYNACEAE. УШ 173 and minutely gray-papillate; petioles 0.5-2.0 cm. long. Inflorescence terminal, dichotomously cymose, 2-3 cm. long, densely gray-puberulent, conspicuously bracteate, the pedicels about 1 mm. long. Calyx lobes subequal, ovate, obtuse, about 1 mm. long, grayish tomentellous-papillate. Corolla greenish-white, densely and finely gray-tomentellous without, the tube about 3 mm. long, the lobes oblong- ovate, about 1.5 mm. long. Anthers about 1 mm. long, inserted about midway within the corolla tube. Ovary ovoid, tomentellous, about 1 mm. long. Follicles broadly oval to nearly circular, 4-6 cm. long, 3.5-5.0 cm. broad, verrucose, with a stipe about 0.5 cm. long. Guiana (and western Brazil?), in mora and mixed forest; flowering in Septem- ber and October. Vernacular names: Zwart parelbout and Jaroro kharemeroe (Surinam—Bosch- wezen). BnrrisH Guiana: Kamuni Creek, Groete Creek, Essequibo River, Maguire 9 Fanshawe 22832 (F, GH, MO, NY, U, US). SuRINAM: Sectie O, Boicbwezen 4830 (NY, О, US), Stabel 70 (U). GuiANE FRANCAISE: Maroni, Wackenbeim 133 (P); locality lacking, Poiteau s.m. (G, GH, W), Martin s. n. (P), Benoist 963 (P). I have not seen the type specimen of А. Kublmannii, but the description of the fruiting branches contains nothing which does not apply to A. oblongum. Kublmann 22453 was collected in western Mato Grosso near the Bolivian border, however, and would thus enlarge the known distribution of the collective species greatly. Without inflorescence final disposition of А. Kublmannii is impossible. 29. ASPIDOSPERMA DISCOLOR A.DC. in DC. Prodr. 8:398. 1844; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1947. (T.:Blanchet 3388!). Aspidosperma Francisii A.DC. loc. cit. 1844. (T.: Blanchet 28501). аст discolor A.DC. В. parvifolium Muell. Arg. in Mart. Fl. Bras. 61:60. 1860. Blanchet 2859!). Whose discolor (A.DC.) O. Ktze. Rev. Gen. 2:416. 1891. Trees as much as 20 m. tall, the branches moderately slender, minutely gray- papillate when young, soon becoming glabrate and with dark brown, irregularly lenticellate bark. Leaves broadly elliptic to oval, apex obtuse to rounded, base broadly rounded or abruptly cuneate and revolute-auriculate, 4—8 cm. long and 2—3 cm. broad, firmly membranaceous to moderately coriaceous, glabrous and olive- green above, pale and densely gray-papillate beneath; petioles 0.7-1.0 cm. long. Inflorescence terminal, corymbi-thyrsiform, 5—7 cm. long, the peduncles minutely puberulent-papillate, branching repeatedly, the bracts rather conspicuous, the pedi- cels about 1 mm. long. Calyx lobes very strongly unequal, 2.0—3.5 mm. long, tomentellous-papillate. Corolla greenish-white, densely tomentellous without, the tube about 4—5 mm. long and 2 mm. broad, the lobes ovate, about 3 mm. long. Anthers about 1.5 mm. long, inserted within the upper third of the corolla. Ovary ovoid, about 1.5 mm. long, hirtellous. Follicles broadly oval, about 6 cm. long and 4 cm. broad, spiny-verrucose. [Vor. 38 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN Eastern Brazil, in forests. Vernacular names: Quina de rego (Froes); Cabo de machado (Ducke). Bnazi.: Baía: Igreja Velha, Blanchet 3388 (NY, W); Serra Acurua, Blanchet e (NY, W); Jacobina, Blanchet 3761 (P); Ibyguara, Serra Sincora, Froes 4 (NY). NAMBUCO: Recife, Ducke 2230 aM O). Froes reports that the bark is very bitter and is used in treatment of malaria. 30. ASPIDOSPERMA SALGADENSE Mgf. in Notizbl. 12:554. 1935. (Т. Ducke 22456!). Trees as much as 30 m. tall, the branches relatively slender, densely grayish yellow-papillate when young, becoming glabrate and with dark brown, conspicu- ously lenticellate bark at maturity. Leaves ovate- to lanceolate-oblong, apex ob- tuse to abruptly acute, base broadly obtuse and revolute-auriculate, 6-12 cm. long and 2—5 cm. broad, moderately coriaceous, glabrate and rather lustrous above, densely yellowish gray-papillate beneath; petioles 0.5-1.5 cm. long. Inflorescence thyrsiform, 5—6 cm. long, densely yellowish gray-papillate, the peduncles rather stout, branching repeatedly and monochasially, the bracts conspicuous, the pedicels about 1 mm. long. Calyx lobes more or less unequal, broadly ovate, obtuse, 1.5—2.0 mm. long, grayish tomentellous-papillate without. Corolla white, densely tomen- tellous without, the tube about 4 mm. long and 1.5 mm. broad, the lobes ovate, about 1.5 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Ovary ovoid, sulcate, glabrous, about 1.5 mm. long. Lower Amazon valley, in forests on terra firma; flowering from August to October. Vernacular name: Сағарапайра (Capucho). IL: PARA: Lago Salgado, Rio Trombetas infer., Ducke 22456 (К, U, US); Boa узга, Tapajos region, Capucho 457 (F). 31. AsPIDOSPERMA NITIDUM Benth. ex Muell. Arg. in Mart. Fl. Bras. 61:59. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. П, 19:365. 1947. (T.: Spruce 1651!). Thyroma nitida (Benth.) Miers, Apoc. So. Am. 24 Md aquaticum Ducke, in Archiv. Inst. Biol Ver. Rio Jan. 4:59, pl. 3, fig. g. 38. (T.: Ducke 24570!). Small trees with slender glabrous branches with dark brown, conspicuously lenticellate bark. Leaves broadly elliptic to oval, apex broadly obtuse or rounded, base obtuse, not revolute-auriculate, 5-15 cm. long and 2.5—7.0 cm. broad, rather thinly coriaceous, glabrous and rather lustrous above, beneath glabrous or indefi- nitely papillate; petioles 1.0-1.5 cm. long. Inflorescence dichotomously cymose, lateral toward the tip of the leafy branches, 1.5—3.0 cm. long, rather few-flowered and indefinitely congested, puberulent-papillate, the bracts inconspicuous, the pedicels about 1 mm. long. Calyx lobes somewhat unequal, ovate, obtuse, 1.0—1.5 mm. long, grayish tomentellous-papillate. Corolla greenish, minutely tomentellous- papillate without, the tube about 3 mm. long, the lobes ovate, about 0.5 mm. long. Anthers inserted within the upper third of the corolla tube, about 0.5 mm. long. 1951] APOCYNACEAE. VIII 175 WOODSON—STUDIES IN THE 77229 РА ЖУ? 7 % h FRUH fy УЙ. Ji Муу“ САМА. е \ Aspidosperma nitidum Ben Flowering ip. Ie flower, and fruit; ох oblongum А. ОС. (lower [em lower face of Ovary ovoid, sulcate, glabrous, about 0.5 mm. long. Follicles broadly obovate cr oval, about 4 cm. long and 2.5—3.0 cm. broad, prominently verrucose. Upper Amazon valley, in inundated forests; flowering intermittently Carapanaiba (Ducke). NY, U, US), 660 (F, MO, NY, 5, Vernacular name: ZIL: AMAZONAS: Manáos, Ducke 24570 (G, K, MO Us), p» (F, MO, NY, US), 309 (F); Barro do Rio Negro, Spruce 1651 (GH, NY, P, W). 32. ASPIDOSPERMA AURICULATUM Mgf. in Notizbl. 12:299. 1935; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:365. 1947. (T.: Ducke 22450!). Large trees; branches relatively slender, inconspicuously puberulent-papillate when young, soon glabrate and with dark brown, inconspicuously lenticellate bark [Vor. 38 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN Leaves broadly oval, apex broadly rounded, base broadly obtuse and rather incon- spicuously revolute-auriculate, 8—12 cm. long and 4.0—6.5 cm. broad, moderately coriaceous, glabrous and opaque above, paler and very minutely papillate beneath; petioles 1.0-1.5 cm. long. Inflorescences lateral toward the tip of the leaf branches, about 4—6 cm. long, relatively lax, the peduncle inconspicuously puberulent-papil- late and several times dichotomous, the bracts inconspicuous, the pedicels about 1 mm. long. Calyx lobes nearly equal, ovate, acute, about 2 mm. long, pale yellowish tomentellous-papillate without. Corolla minutely yellowish tomentellous-papillate without, greenish white, the tube about 3 mm. long, the lobes ovate, about 1 mm. long. Anthers inserted within the upper third of the corolla tube, about 1 mm. long. Ovary ovoid, sulcate, glabrous, about 0.5 mm. long. Fruits unknown. Lower Amazon valley in forests on ferra firma; flowering in January. Vernacular name: Сағарапайба (Ducke). BRAZIL: PARÁ: Belém, inter locos Catu et Providencia, Ducke 22450 (MO, P, U, US). 33. АѕрІроѕРЕКМА Pichonianum Woodson, spec. nov. Arbores ca. 15—30 m. altae; ramulis graciliusculis juventate minute denseque luteo-papillatis maturitate fuscis et vix lenticellatis. Folia anguste elliptica vel elliptico-lanceolata apice gradatim acuteque acuminata basi anguste cuneata ibique margine revoluta 5-11 cm. longa 1.5—4.0 cm. lata moderate coriacea supra glabra illustria subtus pallidiora et minute griseo-papillata; petiolis ca. 1 cm. longis. In- florescentiae cincinnate laterales cymosae multiflorae 2—3 cm. longae hemisphaericae dense cinereo-puberulae; pedunculis repetite ramosis; bracteis vix manifestis; pedi- cellis ca. 1 mm. longis. Calycis laciniae plus minusve inaequales ovatae late obtusae 1.5—2.0 mm. longae extus dense griseo-luteae tomentello-papillatae. Corolla virescens extus luteo-tomentella; tubo ca. 3 mm. longo; lobis late ovatis ca. 1 mm. longis. Antherae ca. 1 mm. longae. Ovarium ovoideum ca. 1 mm. longum dense tomentello-papillatum. Fructus ignoti. Eastern Venezuela, in forested slopes; flowering in November. Vernacular names: Poretay-yek, Cabo de hacha (Steyermark). VENEZUELA: BOLIVAR: wooded slopes of Quebrada O-paru-má between Sta. авсны de Kavanayén and Río Pacairo (tributary of Río Mouak), alt. 1065-1220 m., Novembe 20-21, 1944, J. A. Steyermark 60378 (Е, MO, HOLOTYPE). A very unusual plant particularly because of the numerous, cincinnately arranged inflorescences. Dr. Steyermark reports that the sap in the leaves and stems is milky, and that the wood is very strong, being used for ax-handles and other implements. The species commemorates my friend M. Marcel Pichon, of the Muséum National d'Histoire Naturelle. Serres VII. SrEGOMERIA Woodson, n. ser. Trees with milky latex, apparently evergreen, with moderately thin rimose bark, conspicuously lenticellate. Branches with naked acuminate buds, the sub- tending internode scarcely thicker than the terminal petiole and the bud thus ap- pearing pseudo-lateral, without definite seasonal articulations. Inflorescence sub- 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ УЛ terminal and usually extra-axillary, pedunculate or cauliflorous, without evident bracts; calyx lobes 4, abnormally 5, the outer pair much larger and strongly con- nate (except in A. illustre) , completely enclosing the inner pair which are separate and much smaller; corolla tubular-salverform, submembranaceous, the lobes as- cending, the tube not constricted at the orifice, sericeous without; ovary essentially terete. Follicles very broadly dolabriform, plano- to concave-convex, the placenta rotating about 2752-3007, sessile or shortly stipitate, with a loose, wrinkled, and more or less lenticellate periderm; seeds with a concentric circular wing. Species 34—36 Figure 10. KEY TO THE SPECIES a. Corolla 2.6—2.8 cm. long; outer calyx lobes about 1.5 cm. long, free to the base or nd 50; а revolute-auriculate at the base................ 34. A. ILLUSTRE аа. аг about 1.2 cm. long; outer calyx lobes 2—6 mm. long, con- about half. m length; deve not i m 7 sit Bs Inflorescence wit e-dichot us peduncle lon ee fire eared ate to кекті gla -— т {ЕЛАТУ . А. STEGOMERIS nflor sessile, the obscure peduncle about long; follicles densely velutinous-papillate, not obviously E 36. A. CURRANII c — Eh oO a [e] © "5 о о < - 5 a с ы pat Е 34. ASPIDOSPERMA ILLUSTRE (Vell.) Kuhlm. & Pirajá, іп Arch. Jard. Bot. Rio Jan. 4:375, pl. 35. 1925. Coutinia illustris Vell. Quinogr. Portug. 166, pl. ТО. 1799. Tall trees with sulcate trunks 2-3 dm. in diameter, the branches rather stout, essentially glabrous, conspicuously lenticellate. Leaves oblong-elliptic, apex obtuse to broadly acute, base rounded and revolute-auriculate, 5-12 cm. long and 2—5 cm. broad, subcoriaceous, lustrous above, paler and submicroscopically papillate beneath; petioles 1.5—2.0 cm. long. Inflorescence terminal or subterminal, thyrsi- form, few- to several-flowered, somewhat shorter than the subtending leaves, con- spicuously bracteate, the pedicels about 1.5 cm. long. Calyx lobes very unequal, essentially free, the outer pair much the larger, broadly oval, rounded, about 1.5 cm. long, the inner much smaller, ovate, acute to acuminate, 2-5 mm. long. Corolla wholly glabrous without, pilosulose within, the tube 1.2-1.3 cm. long, the lobes broadly elliptic, 1.4—1.5 cm. long. Stamens inserted above midway within the corolla tube, the anthers about 3 mm. long. Ovary oblongoid, about 1.5 mm. long, glabrous. Follicles very broadly dolabriform, abruptly narrowed to a short stout stipe, about 10 cm. long and 7 cm. broad, smooth, glabrous; seeds with a nearly circular wing about 7 cm. in diameter. Eastern Brazil, in forests. Vernacular names: Tambú peroba, Quina de сататй (Serviço Florestal; Limeira). BRAZIL: BAÍA: s do Gronogogy, Limeira 17935 (MO), Curran 41 (GH, US). MINAS GERAIS: төлес dim Serviço Florestal 107 (MO). Before having seen the flowers of this species, generously given me by Dr. Kuhl- mann, I had placed this species in the series МїтїрА because of the revolute-auricu- [Vor. 38 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN late leaf bases. And that character of A. illustre, together with the separate outer calyx lobes, certainly must be very significant in providing a link between series Nitipa and StEGOMERIA. The follicles of А. illustre, however, are very typical of STEGOMERIA; furthermore, although there is a strong tendency toward inequality in the calyx lobes of Nitiwwa (cf. A. discolor), it is nowhere as marked as іп А. illustre. 35. AsPIDOSPERMA Ste£omeris (Woodson) Woodson, comb. nov. Ж” stegomeris Woodson, in Archivio Bot. 10:39. 1934; М. Am. Fl. 297:123. 938. (T.: Cufodontis 220!). Cufodontia Lundelliena Woodson, in Archivio Bot. 10:40. 1934; N. Am. Fl. 29°:124, (T.: Lundell 3408!). Backen arborea Woodson, in Ann. Missouri ot. Gard. 21:617. 1934; N. Am. Fl. 29?:123. 1938. (T.: Morton & Makrinius 2692! Aspidosperma Lundellianum Woodson, in Am ‚ Jour. Bot. 22:684. 1935; N. Am. FI. :120. 1938. (T.: Lundell 1284!). Cufodontia escuintlensis Matuda, in Madroño 10:174. 1950. (T.: Matuda 16978!). Trees as much as 33 m. in height, the branches rather slender, indefinitely puberulent-papillate when very young, soon becoming glabrate and with a thin, striate, conspicuously lenticellate, yellowish-gray bark. Leaves alternate, rather narrowly elliptic-oblong to broadly oval, apex shortly acuminate to broadly rounded, base broadly obtuse to rounded, 6—16 cm. long and 2.5-7.0 cm. broad, firmly membranaceous to subcoriaceous, above dark green, glabrous, and somewhat lustrous, beneath paler and indefinitely papillate to essentially glabrous; petioles 0.7-1.5 cm. long. Inflorescences lateral and extra-axillary near the tips of the leafy branches, cymose, several-flowered, more or less densely yellow-papillate, the peduncle once- to thrice-dichotomous and about 1-2 cm. long; bracts extremely inconspicuous or absent; pedicels 1-5 mm. long. Outer calyx lobes 3-6 mm. long, more or less densely yellow-papillate. Corolla yellowish-white, the tube 4—6 mm. long, about 1.5 mm. broad, densely and appressed velutinous-papillate with- out, the lobes oblong-elliptic, 5—6 mm. long, essentially glabrous without. Stamens inserted about midway within the corolla tube, the anthers about 1.2 mm. long. Ovary ovoid, glabrous, about 1 mm. long. Follicles broadly subreniform to nearly circular, 7-11 cm. long and 6.5—8.0 cm. broad, sessile, indefinitely papillate to essentially glabrous, covered with small lenticels. Southern Mexico to Costa Rica, in various types of forest; flowering from February to July. Vernacular names: Chichi blanco (Chiapas—Matuda); Pechmax, Pechnox (Yucatan—Flores); Malady blanco, Peechmaax (Lundell—British Honduras) ; Chichica (Standley—Guatemala). О: CAMPECHE: Tuxpena, Lundell 1284 (Е, GH, MO, NY, US). снілраѕ: Erg кү» 16078 (F); Esperanza, Matuda 17538 (F British Honpuras: El а District, Lundell 6220 (MO), 6 MO). ours A: PETEN: Libertad, Lundell 3408 (MO); салы Egler 42-320 (MO); Vaxactun, Bartlett =» (F, MO, NY, S, US). RETALHULEU: vicinity of Retal- huleu, Parr 88803 (F, MO). 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 179 55 T 7, мү SS & \ Lf Ж Wis \ " Қыз \ ag МУМ TU y Flowering branch, entire and Fig. 10. Aspidosperma stegomeris Sige Woodson: Hiei flower, dissected calyx, and fru Costa RICA: PUNTARENAS: Jiménez, Cufodontis 220 (W). The embarrassing synonymy of this species is largely due to my failure, on the one hand, to associate the peculiar flowering specimen of Cufodontia stegomeris with the fruit of Aspidosperma Lundellianum, and to appreciate the variability of the species, upon the other. The circumstances have been discussed by Mr. Matuda (loc. cit. 1950) Standley reports that in Guatemala the species is a common and important lumber tree; Matuda reports that in Chiapas it has no use as timber, but can be used for firewood. Matuda also says that the bark is used to make a bitter con- coction for the treatment of malaria. 36. AsPIDOSPERMA CURRANI Standl. in Trop. Woods 36:14. 1933. (Т.: Curran 294!) Trees as much as 30 m. tall, the branches relatively slender, glabrous, with rather thin, striate, yellowish brown, inconspicuously lenticellate bark at maturity. Leaves narrowly oblong-elliptic, apex acutely acuminate, base rather narrowly [Vor. 38 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN attenuate to the petiole, 7-15 cm. long and 2.5-3.5 cm. broad, firmly membra- naceous, glabrous; petioles 0.5—1.0 cm. long. Inflorescences lateral and extra- axillary somewhat below the tip of the leafy stems, virtually sessile, the obscure peduncle about 1—3 mm. long, bearing 2—5 sessile, greenish-yellow flowers. Outer calyx lobes about 2 mm. long, densely yellow-papillate. Corollas not seen (color from Castañeda). Follicles very broadly subreniform to nearly circular, about 6 cm. in diameter, densely velutinous-papillate, not obviously lenticellate. Northern Colombia, in lowland forests; flowering in April. Vernacular names: Carreto (Whitford & Pinzon) ; Caretillo (Castafieda). COLOMBIA: BOLIVAR: vicinity of Estrella, и 204 (GH). caLpas: Buenavista, Whitford & Pinzon 2 (Е, СН, К, NY, US). MAGDALENA: (MO); Pinijay, Castaneda 1105 (MO). Series VIII. QuEBRACHINES Woodson, Ser. Macrolobii К. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4? M 5 Tucurinca, Castaneda 636 n. ser. *:141. 1895; § Pungentia Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:368. 1947 Trees with thin, yellowish-gray, rimose bark, apparently evergreen. Branches with small scaly buds, but without definite seasonal articulations, the nodes knobby. Leaves opposite or ternate, small and narrow, spine-tipped, coriaceous (firmly mem- branaceous in А. horco-kebracho), the veins more or less immersed but crowded and sharply ascending. Inflorescences chiefly axillary at the upper nodes, thyrsi- form, usually rather few-flowered, with small caducous bracts; corolla tubular- salverform, glabrous or indefinitely papillate, somewhat coriaceous, the lobes nar- row and ascending, the tube not constricted at the orifice. to subcylindric, almost equally bi-convex, the placenta ro Follicles broadly oval tating about 45^-135^, indistinctly papillate to essentially glabrous, more or less obviously lenticellate, sessile but with the pedicel conspicuously accrescent in A. quebracbo-blanco— shortly stipitate and the pedicel not accrescent in A. horco-kebracho; seeds with a nearly circular concentric wing. Species 37-38 KEY TO THE SPECIES E L inflorescences several- to many-flowered, usually densely canes Figure 11. eaves coriaceous, with 20—30 pairs of closely crowded secondar гу veins; -flo cent; follicles woody, sessile but the fruiting pedicel accrescent to fo orm a 37. A. QUEBRACHO-BLANCC m m pe в ы < m N B: th ч E = pescar di to subcoriaceous, with about 10 pairs of ondary veins; inflorescences few-flowered, б ате istant ре coriaceous, ans cates icted t short, narrow stipe, fruiting pedicel not greatly accrescent the 38. А. HORCO-KEBRACHO 37. ASPIDOSPERMA QUEBRACHO-BLANCO Schlecht. in Bot. Zeit. 19:137. 1861; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:368. 5. ft.) Macaglia Quebracbo O. Ktze. Rev. Gen. 2:416. 1891. (Based 1947. (T.: Burmeister upon A. quebracho-blanco chlecht.). Macaglia quebracho-blanco (Schlecht.) Lyons, Pl. Names Sci. & Pop. ed. 2, 286. 1907. 1951] WOODSON—STUDIES IN THE APOCYNACEAE. VIII 181 Lo iios quebracbo-blanco Schlecht. ssp. brevifolium Hassl. in Fedde, Repert. 12:261. : Hassler 12003). үа са Speg. in Physis 3:333. 1917. (Т.: Spegazzini s. n.). Aspidosperma crotalorum Speg. loc. cit. 334. 1917. (Т.: Spegazzini s. n., photo. US!). Aspidosperma quebracbo-blanco Schlecht. var. ellipticum ме in Notizbl. 12: m 1955. Rojas 142 ов quebracbo-blanco uocis forma Schlechtendaliana Mgf. loc. cit. 13:467. A. quebracbo-blanco form. Aun quebracbo-blanco SEI NE Malmeana Mgf. loc. cit. 1937. (Т. Malme s. n. ші Gl quebracho- blanco Schlecht. forma Spegazziniana Mgf. loc. cit. 1937. (Based otalorum Speg.) Trees 5-20 m. tall, the branches slender, minutely papillate when young, soon becoming glabrate and with thin, orange-brown bark. Leaves opposite or ternate, narrowly oblong-elliptic to obovate or oblanceolate, apex gradually narrowed to a sharp spine, base gradually attenuate to a subpetiolar base, 3-5 cm. long and 0.5-1.5 cm. broad, coriaceous, light, frequently yellowish-green, glabrous or minutely papillate when young, the sharply ascending secondary veins of about 20—30 pairs closely crowded and more or less immersed within the thick mesophyll. Inflor- escences opposite or ternate in the upper leaf axils, thyrsiform, densely cinereous- papillate to nearly glabrous, several- to many-flowered, 1—3 cm. long, the bracts minute, caducous, the pedicels 2—3 mm. long. Calyx lobes ovate, obtuse, nearly equal, 1-2 mm. long, indefinitely papillate. Corolla white, yellow, or yellowish green, glabrous to indefinitely papillate without, the tube 3—5 mm. long, about 1.5 mm. broad, the lobes narrowly oblong-lanceolate, 6—7 mm. long. Stamens inserted about midway within the corolla tube, the anthers about 1 mm. long. Ovary ovoid, glabrous, about 1 mm. long. Follicles very variable, subcylindric to broadly oval, 4—10 cm. long and 1—7 cm. broad, very woody, minutely verrucose, with or without a midrib, indefinitely papillate, sessile, but the fruiting pedicel conspicuously accrescent. Southeastern Bolivia and roughly the northern half of Argentina and adjacent Paraguay and Uruguay, from 15 to 1800 m. elev., in xerophilous woodlands; flowering intermittently, but chiefly from October to February. Vernacular names: Quebracho blanco (Schlectendal et al.); Ubird-ro-puiita (Argentina—Spegazzini) ; Kacha-kacka (Bolivia—Cardenas). Paracuay: zwischen Río Apa und Río Aquidabán, Centurión, Fiebrig 4250 (GH US); Pilcomayo River, Morong 900 ( E sc MO, NY, US); locality lacking, donas Hk Sta. Solfia, orillas del Río Uruguay, Rosengurtt B-3240 (MO, US). LIVIA: COCHABAMBA: Misque, Cárdenas 2144 (GH). sTA. CRUZ: Mairana, Stein- Ур, 8278 (F, GH). ARGENTINA: CATAMARCA: Capital, Castillon 1560 (СН). снасо: Taco Pozo, Cab- rera 3168 (NY); Phu Meyer 10 (NY); Barranqueras, г 12 (05), 39 (US), 43 (GH, is US); locality uod Curran 653 (F, NY). corposa: Estancia Germania, Lorentz s. (GH); Cordoba, Hieronymus s.n. (F, US); Qu ilino, Curran 129 (US); Serrezuela, "Bartlett 19236 (US); Las Tapas, Bartlett 20640 (GH, US); Capillo del Monte, Kurtz 6631 (NY). CORRIENTES: Goya, Curran 242 (US), 235 (US); San Cosme, Parodi [Vor. 38 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN 11. Aspidosperma quebracho-blanco Schlecht.: Flowering branch, entire and dis- к ns fruit, and seed. 12085 (GH). ENTRE R10s: Gualeguaychú, Serie 9 Migoya 673 (MO). FORMOSA: Ро? del e. Parodi 8504 (GH); VER Morel 982 (F); locality illegible, NUR ros (GH, MO, US). Jujuy: San Pedro de Jujuy, Орен aa & Beetle 22511 (GH). La RIOJA: мез Varela 777 (US); Capital, ее s.n. (МО); Gral. Roca, Gómez бча (МО). МЕ Alto Pencoso, Jörgensen 103 (С). sALTA: Sierra Sta. Barbara, Schnel 47 (MO); Gis de la Candelaria, Venturi 0701 (GH, MO, NY); Quebrachal, Castellanos 1748 ( Is: Sierra Varela, Castellanos 2008. (MO); Quines, Castellanos 882 MO); Larca, Castellanos 2634 (MO); Sierras de La Quijadas, Castellanos 223 STA. FE: Lant ob (NY); covi, Venturi 7 (MO pto. Gral Obligado, Ragonese 3111 (MO); Dpto. San Justo, Ragonese 2734 SANTI T San Sebastian, Gramajo 10 (US); El Charco, Schreiter s. n. (GH). Tucuman: Vipos Venturi 1373 (F, GH, MO, US); Burroyaco, Venturi 7479 (F, US); Tapia, Rodriguez 105 (MO). 1951] WOODSON-—STUDIES IN THE АРОСҮМАСЕАЕ. УШ 183 Admittedly, A. quebracho-blanco is a variable species, but no more so, probably, than most other species of this genus or many others. Most variable, in this, is the type of fruit, which forms the basis for Markgraf's definition of forms. With- out personal experience in the field, I am hesitant to adopt these, for I think it very probable that the fruit types are not actually of three discontinuous sorts but rather form a continuous series which would prove embarrassing to one attempting infra-specific classification. This surely has been the case in the systematics of the North American forest flora. A. quebracho-blanco is a well-known timber tree of Argentina and Paraguay. Spegazzini reports that its bitter bark is used also as a treatment for malaria and other fevers. 38. ASPIDOSPERMA HORCO-KEBRACHO Speg. in Speg. & Girola, Descr. Maderas, 380. 1910; in Physis 3:335. 1917. (Т.: Spegazzini 1602, photo. US!). Trees as much as 25 m. tall, the branches relatively slender, glabrous, with rather thin, orange-brown bark at maturity. Leaves opposite or ternate, elliptic- to oblanceolate-oblong, apex acute to obtuse with a rather short spine at the tip, base acutely to obtusely cuneate to the subpetiolar base, 2-4 cm. long and 0.5-1.0 cm. broad, glabrous, firmly membranaceous to subcoriaceous, the secondary veins of about 10 pairs, relatively distant and plainly evident upon either surface. In- florescences opposite or ternate in the upper leaf axils, reduced to an umbelliform cluster of 2—4 flowers, occasionally 1-flowered, glabrous, the peduncles 2-4 mm. long, the bracts minute, apparently persistent, the pedicels 1-2 mm. long. Calyx lobes ovate, acute, about 1 mm. long, glabrous. Corolla reddish (Spegazzini), glabrous without, the tube about 4 mm. long and 1 mm. broad, the lobes narrowly oblong-lanceolate, about 3 mm. long. Anthers somewhat less than 1 mm. long, inserted about midway within the corolla tube. Ovary ovoid, about 1 mm. long, glabrous. Follicles broadly obovate to nearly circular, 3—5 cm. in diameter, cori- aceous, glabrous, not obviously lenticellate, with a more or less conspicuous mid- rib, abruptly constricted to a narrow stipe about 0.5 cm. long. Northern Argentina, in dry mountain forests; flowering from November to February. Vernacular names: Horco-kebracbo (Spegazzini) ; Quebracho-blanco (Curran). ARGENTINA: CHACO: Barranqueras, Curran 86 (F, GH, NY, US); Fontana, Meyer 7 (NY). sra. FE: Depto. Gral Obligado, Ragonese 3645 (MO). Spegazzini reports that the timber of this species is as desirable as that of A. quebracbo-blanco. Serres ІХ, NoniLES Woodson, n. ser. Ser. Macrolobii K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?:141. 1895, in part; $ Crassituba M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947. Trees with red or milky latex, apparently evergreen, with close hard bark, not obviously lenticellate (except А. decussatum). Branches with naked acuminate [Vor. 38 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN buds, the subtending internode definitely thicker than the terminal petiole, without definite seasonal articulations and with the leaves not definitely crowded toward the tips. Leaves alternate (opposite in A. decussatum), usually rather rigidly coriaceous, the secondary veins usually sharply ascending or horizontal and not forking toward the margin. Inflorescence variously thyrsiform, the primary branches usually whorled or fastigiate, terminal or subterminal, usually many- flowered, with persistent but irregularly disposed bracts; corolla tubular-salver- form, glabrous or essentially so, subcoriaceous, the lobes usually lance-linear and abruptly caudate-acuminate from near the base (except in A. Fendleri), tightly spiral-convolute in aestivation (except A. Fendleri), erect at anthesis, the tube not constricted at the orifice, longitudinally callose-angulate. Follicles somewhat pyriform to nearly circular or subcylindric, almost equally bi-convex, the placenta rotating about 1202-1807, stipitate to sessile, densely tomentellous, not obviously lenticellate; seeds with a concentric nearly circular wing. Species 30—52 Figure 12, KEY TO THE SPECIES а. ee агу of р се or corymbiform aspect, the oe nches D auge ie her ascending lateral i Tre the flower 2 at ab unded or Ime — b. Corolla weh pt онр as a ану as the ла: longer. ©. Corolla lobes ovate to ovate- "Арле acute, scarcely contorted aestivation; inflorescence blac x or dark gray-tomentellous; leaves oblong-obovate, 6—11 cm white-tomentellous — озеш E т eed sessile or LE black- to dark bro entellou Lm . long cc. Corolla las 5255 linear, abruptly dtl -acuminate from the эн Д I erii in aestivati 9. A. FENDLERI d. Pedic out s long as the calyx bis at anthesis; in- orescence white or "light. gray" -tomentellous; leaves p obovate, 10—20 cm. long, white- or light gray-tomentellous be- neath; lies elipsoid, broadly анын, dark brown-tomen- tellous, 10—1 long 40. A. SPRUCEANUM dd. Pedicels ies as n ng as the calyx lobes to somewhat shorter. e. Calyx lobes foliaceous, about 5 mm. long, as long as the d a tube or n good longer; inflorescence grayish — tomentellous; leaves oval-obovate, rigidly coriaceous, 8— long, yellow-t toment a é beneath P A. STEYERMARKII ee. Calyx lobes Bit pss "-— E : mm. long, about half as long as the corolla tube or som t les f. Stems with a thick corky d dn the inflorescence; и obovate to — poor rounded or very broadly obtus у ti long, persistently golden- omen ellou b iu. m scence brown- -tomentellous ШАН spi, sito m black-tomentel 42. A. NOBILE ff. Stems without a corky perider g- гм Филант umbelli form or * corymbiform, aggregated terminally and subterminally. h. ur hb А many, gray- Абе vii sak calyx lobes about half as long as the corolla ; leaves pw ceous, narrowly i wog to расво 8—17 long, acute to obtuse, prominently veined on Rn sur- faces, essentially g мын 5; folli чы Meroe circular, 6—9 in E narrowly stipitate, black-tomen- cm. tellous 43. A. STEINBACHH 1951] 185 WOODSON—STUDIES IN THE APOCYNACEAE. УШ hh. Inflorescences few, brown-tomentellous; calyx lobes less than half as long as the corolla tube; leaves cori- aceous, oblong-elliptic prios to obo 6—15 cm. long, glabrous, the venation obscure on both surfaces; fol- licles nearly ci ireular, rather ұйы stipitate, 7-8 cm. 44. А. OBSCURINERVIUM -tomenrel lous in diameter, whit gg. Inflorescences dud and Pme the uppermost leaf axils, paniculiform, the d prin A distant and alternate, white опека leaves elliptic- oblong, subcaudate-acuminate, 8—16 ong, mi -= white- tomentellous beneath, the a prominent u oth surfaces follicles oval, 4—7 cm. long, narrowly қы tellous 45. А. VERRUCULOSUM bb. Corolla lobes pk as pan as the corolla tube or les c. Leaves кош зарды obovate, 2—5 cm. long, cus yellow- puberulen cc. Leaves d ate or irregularly approxima d. Leaves persistently to ee papillate beneath. e. Inflorescence black- or atellouss corolla glabrous; leaves элле 315 lpi acute z subcaudate-acuminate, 4-13 cm. ellow tellou ura becoming gray in age; follicles nb 8- 20. apod psa stoutly anl 46. А. DECUSSATUM А. MELANOCALYX black-tomentellous ee. Inflorescence yellowish gray- төй кедү olla lobes in- сч у E te at the ; leaves br Srp ovate- Bis Fen unded or broadly obtuse at j^ and tip, 7-20 cm =; beneath А. ETEANUM ein tomente dd. Qe sentially glabrous beneath, or inconspicuously ETE P when very you e. Inflorescence «wu нар ате gon pedunculate; iie Mex о то ido adly m. long; follicles pengi val, 8—15 . long, Ems E pow ad sordid 1 49. А. MEGALOCARPON range-tomentellous, relatively vi pedunc- 5-15 С E : idee pale o acutely acuminate, ae leaves oblong-elliptic, о о 50. A. LEUCOCYMOSUM aa. Inflorescence ot distinctly cymose aspect, the determinate branches sub- e and much smell than the div атн. frequently scorpioid iiec vels. ers thus borne p . lo 8, урау glabrous beneath and nt with cro tomentellous beneath nd dn АҢ etr Шы Es 6—9 cm. in шы Мы, light brown- 1 51. A. ALBUM pulv rulent bb. ee elliptic to oblong- elliptic, 12725 cm. long, light p tom- entellous beneath and with rather distant secondary venation.............. 52. A. SANDWITHIANUM 39. Авгшроврекмл Fendleri Woodson, spec. nov. Arbores usque ca. 25 m. altae; ramulis sat crassiusculis juventate minute albo- puberulis tandem glabratis cortice atro-fusco vix perspicue lenticellato. Folia alternata vel approximata lamina late obovato-oblonga apice rotundata saepe emar- ginata basi obtuse cuneata vel rotundata 6-11 cm. longa 3—6 cm. lata coriacea supra nitida venis secundariis vix visis subtus minute denseque albo velutina venis secundariis subprominulis utroque latere ca. 12—16 subarcuatis; petiolo ca 1-2 cm. longo. Inflorescentia terminalis thyrsiformis valde plano-corymbiformis multiflora ca. 4—9 cm. longa, bracteis vix bene visis; pedicellis 1-4 mm. longis atro- griseo-tomentellis (pilis albis et nigris interspersis). Calycis laciniae ovato-lanceo- latae acutae 3-4 mm. longae atro-griseo-tomentellae. Corolla extus glabra; tubo valde angulato ca. 3—5 mm. longo et 1.5 mm. lato; lobis anguste ovatis vel ovato- [Vor. 38 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN lanceolatis acutis nec caudato-acuminatis nec alabastro valde spiraliter contorto more specierum propinquorum. Antherae in medio tubi corollae insertae ca. 1 mm. longae. Ovarium globosum са. 1 mm. altum glabrum. Folliculi plus minusve compresse ellipsoidei sessiles vel subsessiles 7-12 cm. longi 5—7 cm. lati plus minusve lignosi atro- vel griseo-tomentelli; seminibus late ovalibus ca. 6 cm. longis 5 cm. latis. Venezuela and British Guiana, in forests. Vernacular name: Karuad-pali (British Guiana—A. C. Smith). UELA: ARAGUA: La Mesa, Parque iesus Pittier 15231 (US), Pittier 9 Nakschennic 15340 (US), 15307 зү: DISTRITO FEDERAL: Macarao Valley, Delgado 207 (F, US); Caracas, Karsten s. ХУ). Merma: Tovar, Fendler 781 (С, GH, К, MO, 222. Us). 316 (GH, K). Вкітізн GUIANA: Isherton, basin of Rupununi River, A. C. Smith 2433 (F, С, NY, MO, S, U, US This species is quite unique amongst NoniLEs because of the broad, loosely contorted corolla lobes. The general relationship of the species appears to be most close to A. Spruceanum, which has considerably larger, obovate leaves and longer-pedunculate inflorescences tending to appear in the upper leaf axils, in addi- tion to the common caudate-acuminate, tightly spiral corolla lobes. The likelihood of hybridization of A. Fendleri and A. album will be discussed briefly under the latter species. 40. ASPIDOSPERMA SPRUCEANUM Benth. ex Muell. Arg. in Mart. Fl. Bras. 61:52. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947 (as A. album). (T.: Spruce 2265!). Macaglia Spruceana (Benth.) О. Ktze. Rev. Gen. 2:416. 1891. Aspidosperma igapoanum Mgf. in Notizbl. 12:298. 1935. (T.: Ducke 245681). Rather small trees (“рагуа” et "vix mediocris", fide Ducke), with rather stout, sulcate branches densely white-tomentellous when young, developing a tight, dark brown bark without obvious lenticels at maturity. Leaves alternate or approxi- mate, broadly obovate to oblong-obovate, rounded at the tip, rounded or broadly cuneate at the base, 10-20 cm. long, 4-11 cm. broad, coriaceous and slightly revo- lute, above lustrous and glabrous with the secondary and tertiary veins rather apparent, beneath densely white-velutinous with 25—30 subhorizontal secondary veins quite apparent; petioles about 1.5 cm. long. Inflorescences terminal and solitary in the uppermost leaf axils, 12-18 cm. long, corymbosely thyrsiform, the long primary peduncles with several secondary peduncles umbellately fasciculate at the tip, the bracts obvious; pedicels about 4 mm. long, all densely and minutely white-velutinous with few interspersed black hairs. Calyx lobes ovate, acuminate, about 2 mm. long, white- or pale grey-velutinous. Corolla pale yellow or white, glabrous without, the tube about 3 mm. long, the lobes ovate-linear, tightly spiral in the bud, about as long as the tube. Anthers inserted above midway within the corolla tube, about 0.7 mm. long. Ovary glabrous, about 0.5 mm. long. Follicles broadly ellipsoid, 10—11 cm. long and 5-6 cm. broad, woody, densely brown- tomentellous, with a short broad stipe 1.0—1.5 cm. long 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 187 Northern Brazil, in periodically inundated forests; flowering in June. BRAZIL: AMAZONAS: Santa Izabel, Rio Negro, Ducke 511 (F, MO, NY, E 24568 (U, US); San Gabriel do Cachoeiras ad Río Negro, Spruce 2265 (C, Е, GH, NY, W). The flat-topped, mostly axillary inflorescences and large obovate leaves of this species help in its identification, since the characteristic pedicels are not fully elongate until anthesis. As a rule, the species of М№тірл growing on inundated lands appear to be smaller trees than those growing on ferra firma. 41. AsPIDOSPERMA Steyermarkii Woodson, spec. nov. Arbores ca. 8—10 m. altae laticem sanguineum exudentes; ramulis crassiusculis juventate minute sulfureo-velutinis mox glabratis cortice atro-fusco valde com- presso. Folia alternata vel approximata late obovata vel elliptico-obovata apice latissime rotundata basi obtuse cuneata ca. 8—13 cm. longa 4.5-8 cm. lata rigide coriacea margine valde revoluta supra illustria venis leviter ascendentibus 25—30 numquam manifestis subtus dense fusco-tomentella venis satius visis; petiolo crasso ca. 2 cm. longo. Inflorescentiae dense corymboso-thyrsiformes multiflorae floribus valde congestis dense fusco-sulfureo-tomentellae ad apicem ramulorum fasciculatim aggregatae ca. 6—10 cm. longae, bracteis vix bene visis; pedicellis ca. 1 mm. longis. Calycis laciniae ovato-lanceolatae acuminatae ca. 5 mm. longae aliquando foliaceae extus fusco-sulfureo-tomentellae (pilis sulfureis et nigris interspersis). Corollae gilvae extus glaberrimae tubo valde calloso-angulato ca. 3.5 mm. longo; lobis ovato- linearibus valde caudato-acuminatis ca. 4 mm. longis in alabastro valde spiraliter contortis. Antherae in medio corollae tubo insertae ca. 1 mm. longae. Folliculi ignoti. VENEZUELA: BOLIVAR: Ptari- e 5% steep forested slopes at base of first line of sand- stone bluffs, o n south-facing part, east of "Cave Rock", alt. 2130 m., November 4, 1944, na. Seeman 59658 (F, MO, HO OI, NY). This species is so distinct that its relationships are conjectural wholly. Its habit suggests an affinity with A. nobile 42. ASPIDOSPERMA NOBILE Muell. Arg. in Mart. Fl. Bras. 61:44. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947. (T.: St.-Hilaire 760!). Aspidosperma Poblianum Muell. Arg. loc. cit. 1860. (T.: pes inea !). acaglia nobilis (Muell. Arg.) O. Ktze. Rev. Gen. 2:416 Macaglia Pobliana (Muell. Arg.) О. Ktze. loc. cit. 1891. Trees about 4 m. tall, the branches disproportionally stout, densely golden- tomentellous when young, covered with a thick corky bark at maturity. Leaves closely crowded, broadly oblong to obovate, rounded or broadly obtuse at the tip, broadly cuneate to rounded at the base, 8—30 cm. long and 3-11 cm. broad, heavily coriaceous, densely golden-tomentellous upon both surfaces when young, becoming glabrate and lustrous above at maturity, pale olive-green, the secondary veins about 40 pairs, broadly ascending, rather indistinct upon both surfaces; petioles 1.5—3.0 cm. long, stout. Inflorescences terminal and axillary at the uppermost nodes, [Vor. 38 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN densely golden-tomentellous, 6—12 cm. long, the primary peduncle relatively short with several secondary peduncles corymbosely or umbellately crowded at the tip, the bracts evident; pedicels about 1 mm. long. Calyx lobes ovate-elliptic, acute, about 3 mm. long, golden-velutinous. Corolla yellow, glabrous, the tube callose- angulate, about 4—5 mm. long, the lobes ovate-linear, caudate-acuminate and tightly spiral in the bud, 5-6 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Ovary globose, glabrous. Young follicles rather narrowly ellipsoid, with a conspicuous stipe, densely black- or dark gray- tomentellous, not known at maturity. Southern Brazil, in sandy and rocky campos and cerrados; flowering from May to e : GOIAZ: entre Bernardo Lobo et Jaragua, жеи к (C, P); Salinas, уйй р (Е, P); locality lacking, St.-Hilaire 760 (F, P, US); ad viam a Brejão Rio Bonaventura et ad Lavrinhas, Pobl 1220 (W); S lacking, Pobl 1835 (K, W). MATO GROSSO: Cuyabá, Malme s. n. (R, US); locality lacking, Kuntze s. n. (NY); Burchell 7505 (K). This utterly distinctive species forms an interesting parallel with A. verbasci- folium, a geographical relative, because of its thick corky bark. 43. ASPIDOSPERMA STEINBACHII Mgf. in Notizbl. 9:1158. 1927. (T.: Steinbach 7261!). Trees as much as 20 m. tall, the branches somewhat thick, essentially glabrous, developing a tight, black bark without evident lenticels at maturity. Leaves broadly ovate-oblong to narrowly oblong-elliptic, apex obtuse or rounded to shortly and obtusely acuminate, base rounded to narrowly cuneate, 8—17 cm. long and 2-7 cm. broad, firmly membranaceous or pergamentaceous, wholly glabrous or in- definitely papillate beneath when very young, lustrous above and somewhat paler beneath, the crowded, almost innumerable secondary and tertiary veins evident upon both surfaces; petioles 2-3 cm. long. Inflorescences crowded subterminally, densely grayish brown-tomentellous, broadly corymbose, 12—15 cm. long, the sec- ondary peduncles umbellated congested, the bracts fairly conspicuous; pedicels about 1 mm. long. Calyx lobes ovate-elliptic, acute, somewhat unequal, about 3 mm. long, densely grayish brown-tomentellous without. Corolla glabrous with- out, the tube callose-angulate, about 4.0—4.5 mm. long, the lobes ovate-linear, caudate-acuminate, tightly spiral in the bud, about 4 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Ovary about 1 mm. long, glabrous. Follicles broadly elliptic to nearly circular, 7-10 cm. long and 6—9 cm. broad, abruptly narrowed to a slender stipe 4—5 cm. long, densely black- tomentellous; seeds broadly oval or circular, 6—9 cm. in diameter. Eastern Peru and Bolivia, in forests; flowering from July to November. Vernacular names: Pinchi-caspi (Peru—Ll. Williams;) Gavetillo (Bolivia— Steinbach). 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 189 PERU: SAN MARTIN: Tarapoto, Ll. Williams 5816 (F, MO, US) BOLIVIA: STA. CRUZ: Buenavista, Prov. Sara, Steinbach 7261 E GH, MO, NY, U), 7356 (F, GH, MO, NY, U). La Paz: San Bartolomé, near Calisaya, Prov. S. Yungas, Krukoff 10290 (F, MO, U, US). Although not conveyed by the key, I believe that A. S£einbacbii is closely related to A. melanocalyx of eastern Brazil, largely because of the similar shape and in- dument of the fruits, although the leaves and indument of the inflorescence may ally it more directly to A. megalocarpon of Colombia and Central America. 44. ASPIDOSPERMA OBSCURINERVIUM Azambuja, in Rodriguesia 9:51, pl. Т. 1947. (T.: Ducke 931!). Tall trees with relatively slender branches inconspicuously gray-pilosulose when young, soon becoming glabrate and with a tight, blackish brown bark. Leaves broadly oblong-elliptic, obtuse to broadly acute at the tip, the base broadly obtuse, 6—15 cm. long, 2.5—5.0 cm. broad, coriaceous, the venation obscure on both sur- faces, glabrous and lustrous above, paler and submicroscopically papillate beneath; petioles 1-2 cm. long. Inflorescence terminal and few in the uppermost leaf axils, corymbose-thyrsiform, about 2—8 cm. long, the secondary peduncles more or less umbellately fasciculate at the tip of the primary peduncle, brown- tomentellous, the bracts inconspicuous; pedicels about 1 mm. long. Calyx lobes rather unequal, ovate, acute, somewhat less than 2 mm. long, gray-tomentellous without. Corolla yellow, glabrous without, the tube callose-angulate, about 3 mm. long, the lobes ovate-linear, caudate-acuminate, tightly spiral in the bud, about 3 mm. long. Anthers inserted about midway within the corolla tube, about 0.7 mm. long. Follicles nearly circular, 7-8 cm. in diameter, woody, densely white-tomentellous, with a rather broad stipe about 1—2 cm. long; seeds circular, about 7 cm. in diameter. Upper Amazon valley, in highland forests; flowering in May. BRAZIL: AMAZONAS: Manáos, ultra Flores, Ducke 931 (F, MO, NY, US), 1156 (MO, NY, US). Outstanding, particularly by reason of the white-tomentellous fruits. 45. ASPIDOSPERMA VERRUCULOSUM Muell. Arg. in Mart. Fl. Bras. 61:51. 1860. (T.: Spruce 3328!). Macaglia verruculosa (Muell. Arg.) O. Ktze. Rev. Gen. 2:416. 189 Aspidosperma paniculatum Azambuja, in Rodriguesis 10: 117, pl. í 71948. (Т.: Ducke 1168!). Rather small trees (fide Ducke), the branches rather slender, inconspicuously white-papillate when young, glabrate and developing a tight blackish brown bark at maturity. Leaves alternate, rather distant, narrowly elliptic-oblong to oblong- obovate, apex abruptly and shortly obtuse-acuminate, base obtuse to acutely cuneate, 8-16 cm. long and 2.5-6.0 cm. broad, subcoriaceous, the secondary veins almost innumerable, subhorizontal, crowded and apparent on both surfaces, above glabrous, olive-green and lustrous, beneath paler and microscopically white- (Vor. 38 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN papillate; petioles 1.0-2.5 cm. long. Inflorescence solitary and frequently axillary at the upper nodes, paniculiform, the branches alternate and relatively distant, 5—17 cm. long, the branches progressively shorter to the subsessile flowers, densely white-papillate, the bracts obscure; pedicels about 1 mm. long or somewhat less. Calyx lobes rather strongly unequal, ovate, acute, about 1 mm. long, densely and minutely white-tomentellous. Corolla white, glabrous without, the tube callose- angulate, about 3 mm. long, the lobes ovate-linear, caudate-acuminate and tightly spiral in the bud, 2.5-3.0 mm. long. Anthers inserted at about midway within the corolla tube, about 0.7 mm. long. Ovary glabrous, about 0.5 mm. long. Follicles elliptic or oval, 4-7 mm. long and 4—5 mm. broad, narrowed to a stipe 0.7-1.5 cm. long, finely brown-tomentellous. Northern Brazil, in sandy savannas; flowering from March to June. BRAZIL: AMAZONAS: Мапіоѕ, Rio Tarumá- -miry, Ducke 018 (F, МО, NY, US), 1164 (MO, NY, US), 1410 (US); ad fl. Casiquari, Vasiva et Pacimoni, Spruce 3328 (С, GH, NY, P, W); between headwaters of Ira-Igarapé and Igarapé-Abiu, affluent of Rio Taraira, Schultes & López 10190 (US). Azambuja's name for this species is so appropriate that it is rather a pity to have to discard it in favor of Mueller's. However, it appears quite unavoidable, since the foliage of Spruce's specimen corresponds so closely to that of Ducke's specimens. Spruce's fruits are somewhat smaller and have somewhat more slender stipes than those of Ducke, but I believe that this is due to their relative im- maturity. 46. AsPIDOSPERMA decussatum Woodson, spec. nov. Arbusculae vel frutices ca. 3 m. alti laticem lacteum exudentes; ramulis sat crassiusculis juventate minutissime gilvo-pilosulis numquam ancipitis maturitate glabratis et cortice fusco-brunneo paulo lenticellato acutis. Folia opposita decus- sata brevissime petiolata lamina late obovata apice rotundata basi obtuse cuneata 2-5 cm. longa 1.5-3.0 cm. lata rigide coriacea venis secundariis numerosis recte ascendentibus congestis utrimque prominentibus supra olivacea nitida subtus sub- concolore et obscure minuteque luteo-pilosula; petiolo ca. 0.5 cm. longo. Inflores- centia terminalis corymboso-thyrsiformis pauciflora breviter pedunculata ca. 2—4 cm. longa omnino fulvide gilvo-tomentella, bracteis vix bene visis; pedicellis ca. 1 mm. longis. Calycis laciniae subaequales ovali-ovatae acutae ca. 3.0-3.5 mm. longae dense gilvo-tomentellae. Corolla albida extus glabra odora, tubo calloso- angulato ca. 3.5-4.0 mm. longo, lobis ovato-linearibus in alabastro valde spiraliter contortis ca. 2.0-2.5 mm. longis. Antherae in medio tubi corollae insertae ca. 0.7 mm. longae. Ovarium late ovoideum ca. 1 mm. altum glabrum. Folliculi ignoti. EZUELA: BOLIVAR: Ptari-tepuí, on cee) south-facing slopes overlying sand- stone, on “Cave Rock” above “Cave Camp”, alt. 1810 m., October 29, 1944, J. A. Steyer- mark 50506 (Е, MO, ноготүрЕ, NY). Distinct, except for the QuEBRACHINES with which it is not likely to be con- fused, because of the decussate leaves of most unusual form for the genus. 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 191 47. ASPIDOSPERMA MELANOCALYX Muell. Arg. in Mart. Fl. Bras. 61:52. 1860; M. Pichon, in Bull Mus. Nat. Hist. Nat. II, 19:367. 1947. (Т.: St.-Hilaire пл) Aspidosperma leucomelanum Muell. Arg. B Kjoeb. Vidensk. Meddel. 105. 1869; M. ichon, loc. cit. 1947. (T.: Warming s. n.! МЕЕ ра Ж (Muell. Arg.) О. кш. Rev. Gen. 2:416. 1891. Large or small trees (8 m. tall, fide Ducke), the branches relatively slender, compressed-angulate, densely golden-papillate when young, glabrate and developing a tight gray-black bark without evident lenticels at maturity. Leaves narrowly oblong-elliptic, acute to subcaudate-acuminate, the base acutely cuneate to rounded, 4-13 cm. long and 1-5 cm. broad, subcoriaceous, the numerous broadly ascending secondary veins obvious upon both faces, olive-green and lustrous above, densely golden-tomentellous beneath when young, becoming gray-papillate in age; petioles 1.0-2.5 cm. long. Inflorescences terminal and axillary at the uppermost nodes, very densely corymbose-thyrsiform, very many-flowered, shortly pedunculate, 3—6 cm. long, black- or dark brown-tomentellous throughout, the bracts inconspicuous; pedicels about 1 mm. long or less. Calyx lobes 2-3 mm. long, ovate, blackish brown-tomentellous. Corolla yellow, glabrous without, the tube about 4.5 mm. long, callose-angulate, the lobes ovate-linear, caudate-acuminate, tightly spiral- contorted in the bud, 2.0—2.5 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Ovary about 0.5 mm. long, glabrous. Follicles oval, 8-10 cm. long and 6—7 cm. broad, woody, densely black-tomentellous, abruptly constricted to a stipe 2.0-2.5 cm. long. Eastern Brazil, in forests; flowering from February to April. BRAZIL: DISTRITO FEDERAL: Rio de Janeiro, Glaziou 20414 (C, F, МО, n Ру ШӘ). ad urbem loco Sumaré, Kuhlmann 19397 (Р, U, US). MINAS GERAIS: goa Santa, Warming s. n. (C, NY); Estagao Experimental, Mello Barreto 8704 (F, Mos 8705 (F, MO); locality lacking, St.-Hilaire s. n. (P). Well characterized by the very dense, shortly pedunculate, black-tomentellous inflorescences and small, narrow leaves. The leaves of the type of A. leucomelanum are somewhat larger and the inflorescence indument somewhat lighter in color than those of the type of A. melanocalyx, but I can observe no more important dif- ferences. Mello Barreto's flowering specimen deserves special mention: the inflorescence is rather loose and the corolla lobes somewhat longer than is typical of the other specimens cited. After dissection of several flowers, I have found the number of stamens to vary from 3 to the normal 5 and their position within the same corolla to vary, as well as their size. There is indeed a slight tendency for the anthers to vary in size within single flowers of both the Nostes and the МіТІрА, but it is so marked in the case of this specimen that I must interpret it as an abnormality. [Vor. 38 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN 48. ASPIDOSPERMA ETEANUM Mgf. in Notizbl. 12:297. 1935; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947. (T.: Ducke 22445!). Trees as much as 20 m. tall, the branches rather stout, densely yellowish gray- tomentellous when young, glabrate and developing a rather rimose brown bark without evident lenticels at maturity. Leaves alternate or approximate, broadly ovate-oblong, broadly rounded at the tip and at the base, 6—20 cm. long and 3—8 cm. broad, rigidly coriaceous, the numerous and dense secondary veins rather ob- scure, olive-green and highly lustrous above, pale gray-papillate beneath; petioles 1-2 cm. long. Inflorescences densely crowded terminally and subterminally, very broadly corymbose-thyrsiform, very many-flowered, shortly pedunculate, 9—15 cm. long and about as broad, densely pale yellowish gray-tomentellous, the bracts in- conspicuous, the pedicels about 1 mm. long. Calyx lobes ovate-reniform, rounded, about 2 mm. long, pale yellowish gray-tomentellous. Corolla yellowish, the tube glabrous, callose-angulate, 4-5 mm. long, the lobes ovate-linear, caudate-acuminate and tightly spiral in the bud, inconspicuously barbate, about 2 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Follicles un- nown. Northeastern Brazil, on ferra firma and hillside forests; flowering from August to October. Vernacular name: Muirapyranga eee BRAZIL: AMAZONAs: Parintins, Ducke 138 PARÁ: Oriximiná, Rio Trombeta infer., Ducke 22443 (US); Serra de Almeirim, JP 22445 (P, U, US); Bóa Vista, region de Tapajos, ELS 432 (F). 49. ASPIDOSPERMA MEGALOCARPON Muell. Arg. in Linnaea 30:400. 1860; M. Pichon, in Bull. Mus. Nat. Hist. Nat. IL, 19:367. 1947. (T.: Karwinsky 1348). Маса а megalocarpa (Muell. ANO O. Ktze. Rev. Gen. 2:416. 189 Aspidosperma cruentum Woods. in Amer. Jour. Bot. 22:684. Du fue cruenta). (Т.: Aspidosperma du NS H. H. Bartlett, in Publ. Carnegie Inst. Wash. 461:16. 1935, As Erara sanguinale H. H. Bartlett, loc. cit. 22. 1935, nom. nud. Aspidosperma Matudai Lundell, in Phytologia 1:339. 1939. (T.: Matuda 2030!). Aspidosperma chiapense Matuda, in Madroño 10:172. 1950. (T.: Matuda 16361!). қР Ж” chiapense Matuda forma tenax Matuda, loc. cit. 173. 1950. (Т.: Matuda 73 Trees 7-30 m. tall, the trunk 2-8 dm. in diameter and with whitish, rough bark; branches minutely gray-pilosulose when young, soon glabrate and developing a blackish bark without apparent lenticels. Leaves alternate, elliptic-obovate to narrowly oblong, apex acute to broadly obtuse, base acutely or obtusely cuneate, 5-25 cm. long, 2-9 cm. broad, firmly membranaceous to subcoriaceous, wholly glabrous, highly lustrous above, somewhat paler beneath, the secondary veins broadly ascending to subhorizontal, very numerous and crowded; petioles 1—3 cm. long. Inflorescences terminal and axillary at the uppermost nodes, broadly corym- bose-thyrsiform, shortly pedunculate, many-flowered, sordid brown-tomentellous, 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. VIII 193 the bracts suppressed; pedicels about 1 mm. long or shorter. Calyx lobes somewhat unequal, ovate, acute to obtusish, about 2 mm. long, densely sordid brown- tomentellous without. Corolla yellowish white, glabrous without, the tube callose- angulate, 3—4 mm. long, the lobes ovate-linear, caudate-acuminate, strongly spiral in the bud, about 1.5-2.0 mm. long. Anthers inserted about midway within the corola tube, about 1 mm. long. Ovary globose, glabrous, about 0.7 mm. long. Follicles broadly oval to nearly circular, 8-15 cm. long and 6—11 cm. broad, woody, brown-tomentellous, with a rather broad stipe 1—3 cm. long. Southeastern Mexico to Colombia and British Guiana, chiefly in dense, rather dry forest, but in forested river bottoms as well; flowering from February to August. Vernacular names: Chichi, Chichi colorado, Chichi prieto, Chichi amarillo (Mexico—Matuda); Colorado (Mexico—Ll. Williams); Volador (Mexico-Kar- winsky); My lady (British Honduras—Lundell, Burns, and Record); Red and White malady (British Honduras—Gentle) ; Chichique (Guatemala—J. D. Smith) ; Chaperno (Honduras—Standley); Caney, Mamey (Colombia—Castafieda) ; Sbi- badan (British Guiana—Forest Dept.). Mexico: cHIAPAsS: Escuintla, Matuda 2030 (F, GH, NY, US), 16361 (US). OAXACA: Ubero, Ll. Williams 0504 (Е, MO). veracruz: Fortuño, Coatzacoalcos River, Ll. Williams 8729 (F, MO). Britis Honpuras: Stann Creek, Gentle 2800 (MO, NY, US), 2801 (F), 2855 (F, K, MO, NY), 3271 (F, MO); El Cayo, Lundell 6326 (GH, MO, NY). GUATEMALA: ALTA VERAPAZ: Cubilgiiitz, TOM 44520 (F, MO); between Trece pn and yer Cook & Doyle 47 (US). EscuINTLA: Escuintla, J. D. Smith 2475 (F, GH, US). TEN: Уахассип, Bartlett 12570 (MO, NY, US), 12663 (MO, US); La Libertad, Lundell 3398 (F); locality lacking, Walker 1076 (F, GH, US). QuEzar- TENANGO: Río Ocosito, J. D. Буш 2477 (US). RETALHULEU Río Samalá, Sbannon 209 (F, US); Mazatenango, Bernouilli & Cario 1836 (K, S). SUCHITEPEQUEZ: Rio Sis, J. D. Smith 2476 (K); е Steyermark 47690 (F). HONDURAS: ATLANTIDA: Lancetilla Valley, Standley 53992 (F, US). I (US). RR. MAGDALENA: Caracolicito, region de Sta. Marta, Castaneda 96 (C e Tucurinca y Fundacion, Castañeda & Gonzalez R 365 (OOL, MO); E 4 Casteiteda 620 (MO); Petén, Dugand 821 VENEZUELA: AMAZONAS: San Carlos de fio Negro, Ll. Williams "Rd (F, US). British Guiana: sandhills, Demerara River, Fanshawe F1226 (K, M Xx. This species is obviously one of the most widespread as well as most frequently collected of the genus. Although segregates have been proposed, they are based upon minor variants which appear more or less throughout the entire range and are of little biological significance. Llewellyn Williams reports that the sapwood of the timber is almost white to light brown, and that the heartwood is vermillion or deep reddish brown and constitutes about two-thirds of the bole. Не also reports that although the wood frequently is attacked by white ants, it is tough and difficult to split and is used frequently for house construction, beams, and railroad ties in Mexico. Matuda states that the timber is esteemed by lumbermen of Chiápas, and "is so hard when cured that it is impossible to drive a nail into (Madrofio 10:173. 1950). [Vor. 38 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN 50. ASPIDOSPERMA LEUCOCYMOSUM Kuhlm. in Archiv. Inst. Biol. Veg. Rio Jan. 2:88, pl. б. 1935; M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947. (T.: Ducke 24472!). Aspidosperma leucostachys Kuhlm. ex Mgf. in Notizbl. 12:558. 1935. (T.:Ducke 24472!). Moderate-sized tree (fide Ducke) with rather slender branches inconspicuously white-papillate when young but soon developing a dark brown bark without evi- dent lenticels. Leaves alternate or approximate, oblong-elliptic, acutely acuminate, the base obtusely cuneate, 7-15 cm. long and 2.5—5.0 cm. broad, subcoriaceous, the veins rather inconspicuous upon both surfaces, rather pale olive-green and wholly glabrous, lustrous above; petioles 1.5—2.5 cm. long. Inflorescences corym- bose-thyrsiform, 4—9 cm. long, rather few-flowered individually but rather densely clustered terminally and at the congested upper nodes, pale orange-tomentellous, the bracts rather inconspicuous; pedicels about 2 mm. long. Calyx lobes ovate, broadly obtuse to rounded, about 2 mm. long, densely white-tomentellous without. Corolla creamy white, glabrous without, the tube callose-angulate, about 5 mm. long, the lobes ovate-linear, caudate-acuminate and tightly spiral-contorted in the bud, about 2.5-3.0 mm. long. Anthers inserted about midway within the corolla tube, about 1 mm. long. Follicles unknown. Northern Brazil; flowering in December. AZIL: AMAZONAS: ad ripas altas flum. Curicuriary (affl. Rio Negro super.), Ducke 8. (К, МО, Р, О, 05). 51. AsPIDOSPERMA ALBUM (Vahl) К. Ben. ex M. Pichon, in Bull. Mus. Nat. Hist. Nat. II, 19:367. 1947 (as to basinym). Macaglia alba Vahl, Skrivt. Nat. Selsk. Kjoebenh. 6:107. 1810. (T.: in Hb. Vahl., comm. Richard! ). Bignonia latisiliqua Poir. Encycl. Suppl. 1:632. 1810. (T.: Patris Aspidosperma latisiliquum (Poir.) A.DC. in DC. Prodr. 8:676. 1844 еф (шайды), Peltospermum Patrisii DC. ex A.DC. loc. cit. 1844, лот. nud. іп synon. Miara bacbypterum Muell. Arg. in Mart. Fl. Bras. 61:51. 1860. (T.: Spruce 334 Aspidosperine desmantbum Benth. ex Muell. Arg. loc. cit. 52. өзді. M. Pichon, in Bull. us. Nat. Hist. Nat. II, 19:367. 1947. (T.: Spruce 2618! Maca bacbyptera (Muell. Arg.) O. Ktze. Rev. oo 2:416. 1891. Macaglia desmantha (Benth.) О. Ktze. loc. cit. 1 Aspidosperma Woodsonianum Mgf. in Notizbl. 12: nr 1935. (T.: Boschwezen 2330!). Aspidosperma centrale Mgf. loc. cit. 560. 1935. (T.: Ducke 21593!). Trees 10—30 m. tall, with trunks 3-6 dm. in diameter; branches slender to rather stout, indistinctly papillate when young, soon developing a light brown or blackish bark without evident lenticels. Leaves alternate or approximate, obovate to oblong-elliptic, rounded to shortly acute at the tip, the base obtusely cuneate, 7-15 cm. long and 3-7 cm. broad, coriaceous or subcoriaceous, the secondary typically rather numerous and crowded, typically evident on both surfaces together with the reticulate tertiary venation, occasionally more or less immersed, glabrous 1951] WOODSON-—STUDIES IN THE АРОСҮМАСЕАЕ. УШ 195 14 ^ AP ы БМ / м SOHC ғы (P (QU Px 2, МИЧ? абр ) оха i 2 Й E Y rume a I 4 асрау. РУЛОН СУСЫЛ ОИУ Flowering bran entire opening bud, Inflorescence. Aspidosperma album (Vahl) К. Ben.: NUR ‘fully opened flower, fruit, and seed; Aspidosperma — "Benth. : on both surfaces typically, or more or less pruinose beneath, rather lustrous; petioles 2-3 cm. long. Inflorescences typically somewhat congested terminally and in the uppermost leaf axils, 3-12 cm. long, rather long-pedunculate, more or less white- or yellowish-tomentellous, of distinctly cymose aspect, the determinate branches subsessile and much shorter than the divaricate, frequently scorpioid lateral branches, the flowers nearly sessile. Calyx lobes more or less unequal, ovate, 1.5—2.0 mm. long, white- or yellowish-tomentellous. Corolla yellowish green, glabrous without, the tube callose-angulate, 2-3 mm. long, the lobes ovate-linear, caudate- acuminate, tightly spiral in the bud, about 3.5 mm. long. Anthers inserted about midway within the corolla tube, 0.5-0.7 mm. long. Ovary about 0.5 mm. high, [Vor. 38 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN glabrous. Follicles nearly circular, less woody than in other species, 6—9 cm. long and 5-8 cm. broad, rather thinly yellowish brown-puberulent, contracted to a rather slender stipe 1—4 cm. long. Northeastern Colombia to the Amazon valley, in highland forest (Mora and Wallaba forest in British Guiana) ; flowering from April to November. Vernacular names: Tun-yek (Venezuela—Steyermark) ; Guabadaro (Venezu- ela—Ll. Williams) ; Siba-danni and Shibadan (British Guiana—Forest Department) ; Siferoe adda, Mantjotjo, and Bitterbark (Surinam—Boschwezen) ; Piquía-marfin, Páo amarello, and Araraíba (Brazil—Ducke); Araracanga (Brazil—Huber) : Ararabuba (Brazil—Capucho). COLOMBIA: MAGDALENA: Salotal, Castañeda 1131 (MO). VENEZUELA: AMAZONAS: San Carlos, Río Negro, Ll. Williams 14510 (F, MO, US). BOLIVAR: Ptari-tepui, Steyermark 60644 (F, MO BRITISH GUIANA: Christianburg, Anderson I 125 (К); Essequibo, Tapakuma Lake, Anderson 337 (K); Simuni Creek, Rupununi River, Davis 2109 (K); Mazaruni Station, ау ead 5275 (К, M: Brownsbers, Boschwezen 1704 (U), 1708 (U); Sectie O, Stabel 121 (U), Маа 163 (U), 2330 (U, US), 2726 (U); Zanderij I, LS І 77 (U), Boschwezen "T Ried жік (О), 2525 (K, О), 2636 (NY, О), 2674 (МҮ, US). NA: Cayenne, Patris s. n. (С); locality dr Richard s.n. (C), rina п. E К, BRAZIL: AMAZONAS: prope Panuré, ad Rio Vaupés, Spruce 2618 (C, GH, NY, P); ad fl. КҮ Vasiva, ес Pacimoni, Spruce 3345 (С, С, GH, Р); Manáos, Ducke 7a (F), 536 (F, MO, NY, US); Serra de Parintins, Ducke 21593 (U, US); Manáos, Ducke 24576 (U, 15); São Paulo de Olivença, Ducke 22415 (MO). pará: Mosqueiro, Ducke 786 (F, MO, NY, US); Belém, Huber 3848 (P, US); Collares, Ducke 12632 (US); Bóa Vista, Tapajos, Capucho 573 (F). Aspidosperma album is a very variable species. In the lower Amazon valley the leaves are more thin in texture and are nearly glabrous beneath, with promi- nently reticulate tertiary and secondary venation upon both surfaces, and these characters also are shown by the one specimen to the extreme north of its range in Colombia, and by most specimens from British Guiana. On the other hand, most specimens from the central part of the specific range, in the upper Amazon and in the Guianas and Venezuela, show more coriaceous leaves with less prominent venation and with more or less of a white-tomentellous or pruinose indument on the lower surface; the peduncles also appear to be somewhat stouter, more yellowish in color, and less strongly cymose. I suspect that this variability may be due to hybridization with either A. Spruceanum or A. Fendleri; Y am rather inclined to interpret the immature but sessile follicles of the type specimen of A. pachypterum (Spruce 3345) as sug- gesting the latter species. Whatever may be responsible for the variability, I am not prepared to attempt resolving it by the labored recognition of several intergrading entities. Unfor- tunately, the type specimens of all the published species names enumerated at the head of this discussion are of “intermediate” aspect, while the more “pure” popu- lations of Pará and Magdalena are without a segregate name. 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ 197 The combination A. album (Vahl) R. Ben. ex M. Pichon was made to super- cede M. Spruceanum: however, examination of the type specimen of Macaglia alba in Vahl's herbarium at Copenhagen discloses the divaricate inflorescences which support Malme's observation "Secundum specimen in Mus. Haun. asservatum Macaglia alba Vahl est Aspidospermatis species, A. desmantbo Benth. peraffinis." (in Arkiv Bot. 21А, по. 6:7. 52. ASPIDOSPERMA SANDWITHIANUM МА}. in Notizbl. 12:561. 1935. (Т.: Forest Dept. 2108!). Trees as much as 30 m. tall, the branches relatively slender, densely yellowish brown-tomentellous when young. Leaves evidently alternate or approximate, ellip- tic to oblong-elliptic, apex rather broadly acute to obtuse, base obtusely cuneate, 12-25 cm. long and 4.5-7.5 cm. broad, coriaceous, the approximately 20 pairs of secondary veins impressed within the essentially glabrous, lustrous upper-surface and prominently elevated upon the light brown-tomentellous lower surface; petioles 2.5-4.0 cm. long. Inflorescence clustered terminally and subterminally, rather long-pedunculate, 4-14 cm. long, densely yellowish brown-tomentellous, of dis- tinctly cymose aspect, the determinate branches subsessile and much shorter than the divaricate, more or less scorpioid lateral branches, the flowers nearly sessile. Calyx lobes rather unequal, oblong-ovate, obtusish, about 2 mm. long, densely yellowish white-tomentellous without. Corolla pale yellowish green, glabrous with- out, the tube callose-angulate, about 3 mm. long, the lobes ovate-linear, caudate- acuminate and tightly spiral in the bud, about 7 mm. long. Stamens inserted above midway within the corolla tube, the anthers about 0.5 mm. long. Ovary globose, about 0.5 mm. long, glabrous. Follicles unknown. British Guiana, in high "miscellaneous forest"; flowering in August. BRITISH GUIANA: Simuni Creek, Rupununi River, Forest Dept. 2108 (К). Perhaps too closely allied to A. album, but with strikingly different leaves and indument, and with unusually long corolla lobes. EXCLUDED SPECIES Aspidosperma anomalum Muell. Arg. in Mart. Fl. Bras. 61:61. 1860 = Micro- PLUMERIA ANOMALA (Muell. Arg.) Mgf. in Notizbl. 13:458. 1937. Aspidosperma clerceanum Iljin & Kraschen, in Index Kew. Suppl. 7:21. 1929, sphalm. == AsTRAGALUS CLERCEANUS Iljin & Kraschen, іп Acta Hort. Bot. Acad. Sci. (ante Petrop.) 43:591. 1931. Aspidosperma condylocarpon Muell. Arg. in Mart. Fl. Bras. 61:55. 1860 = DieLORHYNCHUS CONDYLOCARPON (Muell. Arg.) M. Pichon, in Bull. Mus. Nat. Hist. II, 19:368. 1947 Aspidosperma macrophyllum Muell. Arg. in Linnaea 30:397. 1860 = PARALYXIA MACROPHYLLA (Muell. Arg.) Mgf. in Notizbl. 13:458. 1937. Aspidosperma quadriovulatum Pittier, in Bol. Cient. & Tecn. Mus. Com. Venez. 1:66. 1925 — RAUWOLFIA sp. 198 [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aspidosperma quebracho-colorado Schlecht. in Bot. Zeit. 19:137. 1861 = ScuiNop- SIS QUEBRACHO-COLORADO (Schlecht.) Bark. & Meyer, in Bol. Soc. Arg. Bot. 3:156. 1950. Aspidosperma sessilis Huber, in Bull. Soc. Bot. Genève, II, 6:200, fig. 13. — MICROPLUMERIA ANOMALA (Muell. Arg.) Mgf. in Notizbl. 13:458. 1915 1937. Aspidosperma tuberculatum (Vahl) R. Ben. ex Pichon, in Bull. Mus. Nat. Hist. Nat. П, 19:369. cit. 1947. 1947 — OcHROSIA TUBERCULATA (Vahl) M. Pichon, loc. INDEX TO EXSICCATAE talicized numerals refer to collectors’ number. s, s.n. (sine numero) to unnumbered 5 parenthetical numerals refer to the numerals of the species conserved in this vision. Allemáo, Е. 067 (5); 968 (18). Anderson, C. W. 125, 337 (51). Arbeláez, E. P. 4680 (16 Archer, W. A. 2483 (26); 2775 (25); 3022 (13); чан (12 Archer, A. Ge het. 135 (7): Balansa, B 0 OQ 4644 (37). Baldwin, A. A 5 (4). Barbosa, A. s.n. Ж? қ j 12570 (49); 12660 (35); 12663 (49); 10236, 20640 (37 eaurepaire, —. s.n. (16). Bénoist, R (28). Bernouilli, & R. Cario 6 (49). Blanchet, J S н (5); 2 (4); 2850, 3388, 3761 = . (51); 1354, 1424 (25); 1704, Esth 1751 (25); 2251,2330, 2389, 2525, 2636, 7074. 2726 (51) ; 4830 (28); 5060, б 395 (25 Britton, N. L. & W. E. Broadway. 2648 18 62, 062, 9426 (18). Bircadway, W.E 7305 (8); 7348 Burchell, W. % 6501, 18). 432 (48); 457 (30); 573 Cárdenas, M. T (25); 1085 (1); 2144 Castañeda, R. R Ó (49); 135 (18); 620 (49); 24 поў (36); 1131 (51). Castafieda, — Gonzales К. 365 (49). Castellanos, А. 492, 1748, 2238, 2634, 290 i Castillon, L. 1569 (37). Chase, А. 7821 (5). Claussen, P. s.n. (7, 8); 326, 327 (7); 346 (7). DE D. 8 Cook, O. F. & C. B. Doyle. Cufodontis, G. 4. 335, Curran, H. M. 9 (37); 41 (34); 43 (37); 86 TU A (37); 148 (13); 235, 242 07); 294 (36); 653 (37). Curran, H. M. & M. Haman. 586, 840, 907, 923 (18); 938, 940, 054, 055, 970 (18); 073 (12); 1211, 1260 (18). Dahlgren, B. E. 928 (5). Danouse, I. i 51). 104 (12); 297 (39). Denis, F. 3578 (16). Drouet, F. 2710 (5); 2724 (5) 2938 (2), 3204 (7); 22 Duarte, A. P. (11). Ducke, A. 7a 138 (48); soy (11) 51); 309 (31); 511 (40); 536 (5 (31); 786 (51); 018 (45); 031, 1156 (44); 1163 (25) ; 1168 (45); 1263 (25); 1410 (45); 2230 (29); 11040, 11402 (1); 12632 (51); 14880 (1); 14900 (25); 15814 (20); 16480 (1); 17105 К. (51); 22436 34947 (13). cke, А. & J. G. Kuhlmann. 15387 (11). Dugand, A. 37 (18); 208, 355, 428, 582 Sl МАСЫ Po 821 (49); 1024 (16); 3138 (18 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. УШ Dugand, A. & R. Jaramillo. 2790 (16); 3229 (18); 3442 (16). Dusén, P. s.n. (7, 16, 17); 13116 (7); 16071 (17); 16514, 16780 (7); 16781 8022 (7). Elias, Bro at^ E 1507 (16). Engel, 1056 (10 Engle, —. s.n. (7, 11) Espina, R. 28 (18). Eyerdam, W. J. & A. A. Beetle. 22511 (37). Fanshawe, D. Е.1226 (49); 5275 (51). Fendler, А. 781 (39); p in part. (12); 1299 in part (18); 131 aed) P K. 337 (14); 4250 (37); 420? ( Forest Demim шок Guiana). 2108 (52); 2 Froes, J. García-B., H CT 6). Gardner, С. 835 (7); 1753, 2664 (5); 2666 (1); 5542 (11). Gentle, P. 2800, с 2855, 3271 (49). Giocomelli, —. s. 37 Glaziou, A. 636, 27 (11); 638 (27); 1586, 2023, 4079, 5000, 5033 (11); 6046 (27); 6636, 0500 (3) ; 11184 (5); 11185 (18); 11180 (16); 12075 (9); 12076 (27); 12048 (17); 12049 (7); 12052 (17); 19211 (7); 17131 (9): 2713243); 17133 (2? 17607 (9); 18364, 18365 (11); 66 (9); 19630 (11); 20414 (47); 21740 (7); 21741 (2); 21742 (42 Gomes, К. 771 (37) Gramajo, —. JO (37) Harshberger, J. W. D 2320 (10); 7100, 71094 (7); Hassler, E. 7207, T2078 (5); 10408, 10408a (16); 10575 (1); 10590 (17); тобот, 10620, 10620a, 10647, тоб4ўа(7); 10651, 10755 (10); 10804 (7); 10895 (1). Haught, O. 4007 (16); 4212 (18). 0). Herzog, T. Hieronymus, G. . (37). Hoehne, 23 e. m (17); 28544 (11); 28660 Horto is 5 Jarmillo Mejia, R. 233 Job, M. M. 1180 (37). 199 Jórgensen, P. 193, 1958 (37); 3690 (10). $. 7%, arsten, H (39); Kenoyer, L. Н. бг (49) Killip, E. . Dugan Jaramillo. 38131 (16; 38169 qus қ й (16). Krukoff, ; 5172 (19); (43); 10833, 10872 (25); 10892 (1); 12150 (25). Ein L4 8 (19); 4373 (9); 1230 (7); 1227 (2); 15334 (16); 15346 m 16366 (3); 19307 (47): 37745 Kuntze; Ос Sa (1.200642) Kurtz, F. 6631 (37). Laekie, J. 2037 (26) Lankester, C. Н. з. м. (49) Leite, О, 2327 (10). Leng, H. 279 (26). Leonard, E. C. & С. M. Leonard. 12646, 13351 2 Limeira, —. 5 (328). Lisbóa, A 24 (1); 2435 (5). Little, E. 1. 23 Lófgren, > - p (16); 770 (5); 036 2 (8). Lorentz, P Sun. (32). iones Р. 200 (18); 12356, 26800 (5); 20059 (18). Lund, P. d ая B 16, 18). Lundell, C. 5); 3308 (49); L. 1284 (3 3408 (35); 6220 (35); 6326 (49); 6360 (35). Lutz, B. PEU. Macedo, A. 520, 536 542 (2); 711, 729; 2 o олко), 2023 (5); 2025 (8). ir. M. 203 (11); 378 uo 460 (11); 072 (10); 620 (17); 652 (8); 689, » 447 ir 0). Maguire, B. & F. Fanshawe. 22832 (28). Malme, G. A. . (42); 1006 (5); 1640 (9; 152567 (10) ; 2732 (10); 2745 (18); M T Martius, P. von. ( s. n. (28), (51). s.n. (5); 262 (8); 485 2030, 16361 (49); 16978, 35). Mello Barreto, Н. L. de. 300 (7); 1303 (9); 3189 (17); 8704. 5705 (47); 9215. 9222 (7); 0501 (15); 5 (27). Mexía, Y. 5054 (3); 0 yt 200 Меуег, Т. Lo IQ (37). Morel, I. 982 (37). Mosén, H. 2532 (10). Mueller, —. 170 (7). Muller, F. 121 (11). Mutis, J. E. 5216 (12). Niel, J. van. 6224 (25) Nuhes, M 6). Núñez Bossio, —. 541 (16). Oliveira, J. E. 211 (17); 252 (10); 325 (3); 1330 (2); v, E 1345 (10). d'Orbigny, A. - (7). Palmer, E. &. n. (3 Pis asm A. S504, 12085 (37). Patri A Pickel, D. B. po (13); 3497 7 (5). Pittier, H. 51, 7773, 8605 (12); 8708, 9042 (18); 0851 (12); 10214 (18); 10380 (12); 11362 (18); 11888 (12); 12225 (13); 133ó1 (12); 15231 ке Pittier, Н. & — Nakischenovich. 1534 КА, =, (39). Pohl, J. E. ‚ (7, 9); 1229, 1835 (42); 218$ (8); diis a 1). tim 1). Ragonese A. E. 2734, 3111 (37); 3645 (38). Record, S. ШЫ 66 (16); d bis, 81 (18). Regnell, A $. n. ; 868 (8); 860 (11); бо, (16); 870 f (17); 877 (2); $71 bis ; Reichardt, —. 3t, 32 тэ Richard, A. s 51 Riedel, L. 671 (7); 1091 5. 1, 9, 11) (15); 2332 (16); ү (1); 2771 (11). Rodriguez, —. 19 Rojas, T. 10590 a (ss also Hassler]. Rose, J. М. 21012 (12 Rose, ‚ М. & Р. Russell. 10041 (9). 240 ( £^ ‚ ravon. 5.7. Rusby, H. i 203 (19); 2649 (3). Rusby, . W. Pennell. 335 (18). St. lise E es s.n. (7, 8, 47); 760 (42); 825 (2); tiy (1% Sandwith, N I (26) chery, R 2 Schnel, —. 47 Schomburgk, К. 468 (26) Econ К. в (37) uch, Y {3 ). алы, К. E. & Е. López. 10190 (45). 10178 (21); [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sellow, F. s.n. (10, 11, 17); 49 (7); 1651 Ў Serie, — & — Migoya. 673 (37). бегуісо, Florestal. 103 (16); 105 (3); 107 (34); 109 (11). Shannon, L. W. 209 (49) Sieber, F. W. 53 (18). € A. C. 2433 (39); 3151 (13); 3380 13). Smith, H. Smith, Н. Н. 436 T 8). Smith, J. D. 2475, 2476, 2477 (49). Snethlage, Е.Н. 676 (1). pruce, R. 1651 (31); 2265 (40); 2618 (51); 3328 (45); 3345 (51); or Stahel, G. ^ (27); 77, Iar Stahel, G. & H. Gonggrijp. 4 (26). Pool P. C. 53002 (49); % (35). Steinbach, J. 6394 (10); f 7356 (43); 20 (19); 8144 (17); 8 (37). КЕС u^ EU о (49); 54085 (18); 59500 (46); 50858 (41); 60378 (33); 60644 (51); — (12); 62819, 62849 ( 18); 62923 (12 16). riana, J. J Tutin, T. G. Uh E ^» (72 4 (5); 7200 (6); 8451 9700 (3), 9701 (19). Varela, —. 777 (37). Vargas, J. M. s.n. (12). BE J. 243 (11). Venturi, S. {2 e Pie 9701 iat Verken, M & P. de Cambour s. n. (16). Wackenheim, —. 133 (28). Walker, —. 1076 (49) Warming, E. 5.п. (7, 8, 10, 11, 16, 17, 47 Weberbauer, А. 7115 (16). - 445 (11); 2001 (42); Weddell, 2470 (1). hite, O. Е. 1018 (3) Whitford, H x 12 (16). Whitford, Н. N. & J. Pinzon. 36). Whitford, H. x & F. Silveira. 62, 125 . sm. (7). Williams, LI. 5816 (43) ; 6231 (12); ane 0504 (49); IOI25, II7I2, 11829 (12 13223 (1); 14510 (51); 14519a (49). Williams, R. S. 255 (1 ood Ри 323 (12). Woronow, С. 7075 (18). Woytkowski, F. 26 (12). 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. INDEX TO VERNACULAR NAMES Guatambu Guatambü do Cid Horco-kebrac Ivaha Jaroro kharemeroe " acka y SUELE с ТЫСЫ ААКЫНЫН ЧАА RUE RE Mantjotjo Maparaná Mosla deemed uoo E Lh ee cene 2. ЫНЫ ЖЕСІН REM Рато гоа ue ааа еа лла ҮШ 201 Panaceia 141 Pão amarello 196 Pao pereira do campo 148 Рао pereira do mato. 150 Parelhout 170 P 148, 159 Paroba mirim 159 Pechmax 178 Pechnox 178 Peechmaax 178 Pequía 148, 153 Pequía amarella 172 Pequía da restinga 153 Pequía de pedra 148 E marfim 172 Pereir 146 Prid da serra 158 Pereiro do campo 148 Pereiro preto 146 Peroba 159 Peroba amarella .. 142 Peroba amar 141 Peroba de gomo -...... 172 Peroba do campo 141 Peroba paulista 146 roba rosa 159 Perobinha ----—-- 159 Pinchi-caspi ------- 188 i co 157 Piquía-marfin 196 Platanote ---—- 140 oretay-ye 176 Quebracho blanco 1815183 Gullo bordón + LL ———— 154 uina de camamü 177 Quina de rego -------- 174 Shibadan 193, 196 Siba-danni 196 Siferoe adda 196 Tambü 142, 148 Honbioifé. (c е. 5.0. EE 151 Tambü canudo 152 TENDO peroba Ss ole trit n — 177 Tambu verde s- 152 dli Gs a DESEE TM MOS Hm MTS 196 Ubira-ro-puütá 181 IE С га ыы 163 КОН ы ш e c E PS 193 ФАК "55 perite nin 154 wb ea СЕТИВА УНЕ oc 171 Witte parelhout ------- 170 NEU" ——— 171 Yema de huevo 154 Peart ра Боп. .—— 1. ee 175 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN SYSTEMATIC INDEX [Vor. 38 type indicates accepted, 2” names; italics indicates synonyms; bold man P emi new species or combination Alsodeia Cuspa жері Spreng ыбы: 162 Aspidosperma Mart. bas 126 acantbocarpum ” даь С >. acreanum Met, ———— 164 album (Vahl) R. Ben 194 anomalum Muell. Arg. —-—---- 197 aquaticum Ducke ---.-------------------------- 174 argenteum Mauell. Arg. -------------------- 151 БЕСЕ Ма a5 cen. 175 australe Muell. 151 var. erytbvosylates Hassl. . 151 estrellense Hassl. 2... 151 longipetiolatum Hassl. 2 151 bello-horizontinum A. Silv. 153 bicolor Mart. 146 brevifolia Rusby 160 camporum Muell Arg. |... 148 carapanauba M. Pichon |... 169 centrele МЕЁ So 194 chakenis м 181 chiapense Matuda 2. 192 forma tenax Matuda -------------------- 192 Chodatii Hassl . 148 clerceanum Піп & Kraschen —у........- 197 com pactinervium Kuhlm. |... 172 сопауіосатроп Muell. Arg. |... 197 crotaloram Speg. -aama 181 cruentum Wo x e ЖАН еН RR 192 Curranii Standl. 179 cuspa (HBK.) S. Е. Blake |... 162 Leonid ган Muell. Arg. .......... 160 ar. genuinum На. 2 160 longepetiolatum На]. 2 160 е НЗ. ес 160 тусетін ADCO. .................. 147 decipiens Muell. I NIE 162 ecussatum Woods. |... 190 desmantbum Benth. 194 MA Жалы Leod E 173 ar. B. ane сов Arg... 173 ы. Muell. Arg. ыы: 158 domingense Urb. 2-2. 162 Duckei Ны. 139 Dusendl Standl .. — 159 eburneum F. Allem. |... 172 elatum E. L. Little — 169 elliptica Rusby 162 eteanum 192 excelsum Benth, ое ——— 171 Fendleri Woods, 2-02 185 Francisii А КЕТЕ y t Gardneri Muell. Arg. Жа асе а. È B. ellipticum Muell. Arg. |... а, ovatum Muell. Arg. 2... orco-kebracho Speg. е СО igapoanum ү PIIR ыча ingratum lagoense Muell. DUE Бе ЕС. lanatum (О. Ktze.) т” Seaton latisiliquum (Poir.) A anum Woo megalocarpon Muell. Arg. |... nitidum Benth. |. ____ occidentale me occidentale Mgf. ... i hum Woods olivaceum Muell. Ase. ЕЕ nu А090 obtusifolium Muell. Arg. bechbyblerum Muell. Arg. __ pallidiflorum Muell. Arg. M 2. Sc 4257, illustre ( Vell.) Kuhim, & х Pirajá A 1951] WOODSON-—STUDIES IN THE APOCYNACEAE. paniculatum т E cer 189 parvifolium AU. Бе гык ш шы 150 РИСО АЛИШ о s 158 Pichonianum Woods. В-и УА platyphyllum r Arg. - DI CERT ЖЕ 138 Poblianum Muell Arg. ------------------- 187 polyneuron Muell. Arg. ——5 ——— 158 var. genuinum На]. -------------------- 159 longifolium На]. -------- 159 populifolium A.DC, ------ — 146 pruinosum Mat. o oe ж 172 py D ls Mu ell. ND NEIN 152 y. obovatum Mn um Е ад. [у н MER LE ues 146 B. molle Muell. Arg. ------- 146 quadri-ovulatum Pittier —----- 197 quebracho-blanco E uo s ФАП orma Malmeana Mgf. ------------------ 18 Mgf. . 1 ы e T зт a gazziniana Mgt. ----------- 181 var. ellipticum Mgf. ---------------------- 181 ssp. brevifolium Hassl. ------------------ 181 quebracho- oH Schlecht. ....... 198 Quirand y -- 148 var. pem ттт Mash ——— 148 m pestre ши = 157 silvaticum MHassl. ------------------ 151 ramiflorum Muell. Arg. -------------------- 142 rauwolfioides Mgf 164 reductum (Hassl.) Woods. -------------- 156 refractum rt 146 Riedelii Muell Arg Seat МИНЕН. ЕЕН 152 хаг. сепшиит FLASS. oooi 153 forma micropbyllum Hass]. — 153 ssp. reductu iM с NS 156 rigidum Rusby ------- 164 jasii Hass] 148 salgadense Mgf. 174 sanguinale Н. H. Bartl. —-—_______ 192 sanguineum Н. H. Bartl. ------------------ 192 Sandwithianum Mgf. ------------------------ 197 Schultesii Woods. |... 168 Sellowii Muell. Arg. -------------------------- 152 ar dade Fasal -——— ы + 153 unum Hass]. —._____- 153 То Muell. Arg. -----.----------- 162 sessilis Huber --- 198 Snethlagei Mgf. 139 Spruceanum Benth 186 stegomeris (Woods.) Woods. -..----- 178 Steinbachii Mgf. 188 Steyermarkii Woods ccc o e fpe 187 pu Mar 150 5 Muell. Arg — 150 mei ien pr ы чер 164 fomentosum: Mart. зеза 147 VII 203 y. angustifolium Muell. Arg. -------- 147 B. velutinum Muell. Arg. ---------—- 147 tuberculatum (Vahl) R. Ben. ---------- 198 lei Mgf. = 154 Vargas ALR. 2252.2... 154 velutinum Fisch 147 venosum Muell. Arg. ------------------------ 158 verbascifolium Muell. ЕС ы. 141 erruculosum Muell. Arg. ---------------- 189 Warmingii Muell. Arg. —------------------ 148 Woodsonianum МЕЁ. -~---------------------- 194 Woronovit Sand, „Е. 162 Astragalus Clerceanus Шіп & Kraschen 197 Ай latisiliqua: VOR ...------- 194 Conoria ? C Bgm 162 Continia. Vel. „Ааа 136 Miusiris ValL ке Sane ae re 177 Cufodontia Woods. ....----- 136 orea Woods e RE —— 178 escuintlensis Matuda -------------------------- 178 queis na Woods, 1a 178 stegomeris Woods. .— 178 biperhyachus condylocarpon КА, Ата) M Pichon ..————————— Geissospermum asus eS re 169 myristicifolium М. -------- 169 ramiflorum Mart. ---.------------------------- 142 Macaglia 2 NES АДА ызы с-қа 136 EINEN SL Ls л sae 194 EON (Muell. Arg.) О. Ktze. .. 151 bicolor (Mart.) О. Ktze. ----------------- 146 camporum vies Atg.) О. Ktze. _. 148 dasycarpa (A.DC.) с ОРЕ 148 decipiens (Muell. Arg.) y сау OM 162 desmantha (Benth.) О. Ktze. ---------- 194 discolor (A.DC.) О. Ktze.—— — — 173 disperma (Muell. Arg.) O. Ktze 158 excelsa (Benth.) О. Ktze.. 171 Gardneri (Muell. Arg.) О. Ktze.- 139 Gomeziana (A.DC.) О. Ktze.....— 48 Hilariana p Arg.) О. Ktze. __. 148 ENERO... eer nnne 141 macrocarpa (Mart.) O. aii ee 139 Martii (Manso) О. Etze. 146 о) Ktz megalocar pa (Muell. ер О. Ktze. 192 melanocalyx (Muell. Arg.) O. Ktze. 191 O 1 multiflora (A. ) a o V 45 nobilis (Muell. Arg.) O. Ktze.---------- 187 oblonga 9% ae PERA 172 obscura (Muell. Arg.) О. Ktze._-..... 148 olivacea (Muell. Arg.) O. Ktze 153 pachyptera (Muell. Arg.) O. Ktze.. 194 platyphylla (Muell. Arg.) О. Ktze... 139 Pobliana (Muell. Arg.) O. Ktz 187 populifolia (A.DC.) О. Ktze......-- [Vor. 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN byricolla (Muell. Ен a Ktze.... 153 Peltospermum P (Mart) O. Kte. — — 146 latisiliquum үгел DC. racho О. Ktze 18 Patrisii DC. Сас. blanco (Schlecht. Lyons 180 Schinopsis Mg oom colorado refracta (Mart.) O. Ktze. 146 (Schlecht.) Bark. & Meyer Spruceana (Benth.) 5 [m TOTIS 186 рутон Mera s-i ias subincana (Mart.) О. Ktze. 0 1 bicolor (Mart.) Міегѕз neni Кеч E т — 148 decipiens (Muell. Arg.) Miers Vargasi C.) O. Ktz 154 Lbotzkiana (Muell. Arg.) Miers 2. Аы pow frs о. Кое. 141 nitida (Benth.) Miers |... verruculosa (Muell. Arg.) О. Ktze.. 189 parvifolia (A.DC.) Miers |... Microplumeria anomala (Muell. Arg.) polyneura (Muell. Arg.) Miers Mgf. 1 Riedelii (Muell. Arg.) Miers Sichrocia — (Vahl) M. Pichon 198 ellowii (Muell. Arg.) Miers- Ostreocarpus sessiliflora (Muell. Arg.) Miers... Paralyxia са (Muell. Arg.) Mgf. 197 [Vor. 38, 1951] 206 ANNALS OF THE MISSOURI BOTANICAL GARDE PLATE 1 Anatomy of the corolla in Aspidosperma (all figures X 115) Fig. 1 macrocarpon Mart.: Origin of fissure at the insertion of the filament. Fig. 2 macrocarpon Mart.: Fissure at level of anther. Fig. 3 Steinbachii Mgf. Fig pyrifolium Mart. . stegomeris (Woodson) Woodson. . Schultesii Woodson. >> >> > > > mi Rod d d M моу polyneuron Muell. Arg. Diplorbyncbus cond ylocarpon (Muell. Arg.) M. Pichon: Absence of fissures. ж iu oo PLATE 38, 1951 Bor. GARD., Vor. ANN. Mo. 2..5 Й" ө Y. t. = tee = 424 е ее Ёз: + қ E e. 97 Жы. үз шашы А», Nae _ ОО v» dat / 4 с р 2. f VHI APOCYNACEAE. WOODSON-—STUDIES IN THE Annals of the Missouri Botanical Garden Vol. 38 SEPTEMBER, 1951 No. 3 A REVISION OF STILLINGIA IN THE NEW WORLD* DAVID JAMES ROGERS** Three previous treatments of the genus Sfillingia (ВаШоп!, Mueller’, Pax & Hoffman?) have done much to clarify the systematics of this group. However, these studies are based upon few specimens, little distribution data, and practically no habitat information. As a result, the species have remained conglomerate or split into ill-defined groups. An examination of annotations borne by herbarium specimens demonstrates the confusion concerning the identity of any one species. Furthermore, a clear differentiation has never been made between S/illingia and the closely neighboring genus Sapium, so that specimens collected without mature fruit can be assigned to neither with certainty. Pax and Hoffmann? have described twenty-six species of Sfillingia; thirteen in North and Central America, nine in South America, three in Madagascar, and one in the Fiji Islands of the southern Pacific. The present revision undertakes to re- examine the American species with the purpose of effecting a natural organization. The standard methods of the herbarium systematist have been employed for in- terpretation of the species, amplified by special field studies for two of the species of the southeastern United States. A conservative concept of species is adopted, and no infra-specific entities are recognized where there are insufficient data for delimitation and description of the variants. Since none of the species of Stillin gia are pantropic in distribution, no artificial separation is made by the omission of the species of the Eastern Hemisphere. The great reduction of the floral organs not only in S/illingia, but in most genera of the Euphorbiaceae, limits the amount of emphasis which may be placed on the structures of the pistillate and staminate flowers for delimitation of the i ane Сёп. pest 510. 1858, ex parte. . Arg. in DC. Prodr. 15?:1155. 186 Ж д А ий, in Engler, Pflanzenr. IV. Fam. 147. V:180. 1912 n investigation carried out in the graduate resi: atory of the Henry Shaw School of Botany of Washington Univer and PORS as a thesis in partial fulfillment of the requirements for the sity degree of Doctor of Philoso ** Allegheny College, yo Pennsylvania. Issued October 5, 1951 (207) [Vor. 38 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN species. This condition necessitates what may seem an over-emphasis on vegetative characters and differences of habitat and distribution. However, correlation of all factors demonstrates that those characters other than of the flower are of sufficient stability to justify their use in separation of species. History Stillingia was named by Garden‘ in honor of Benjamin Stillingfleet, an English botanist who lived from 1702 to 1771. Endlicher? first placed S/illingia in the Euphorbiaceae as a member of the tribe Euphorbieae. J. F. Klotzsch® placed the genus in the tribe Hippomaneae. His work included a study of South American representatives of several genera of Hippomaneae for which he gave a short description, usually followed by a list of species belonging to the genus. Klotzsch followed Endlicher in uniting Sapium and Stillingia, with Sapium as а section, maintaining S/illingia as the generic epithet. Не gave a short description of Sfillingia, followed by diagnoses of the sections EUSTILLINGIA and SAPIUM, but hc listed no species for the genus. Baillon's? systematic studies on the Euphorbiaceae employed such a broad in- terpretation of genera allied to Sfillingia that most of those genera of the presently recognized tribe Hippomaneae appeared as sections of S/illingia. In this work, he actually made no formal transfers, but merely listed the genera under the heading Stillingia. He apparently realized later? that the names in his ‘Etude Général des Euphorbiacées’ were not effectively transferred to Stillingia and tried to correct this situation in several volumes of 'Adansonia', in which he made the formal trans- fers, listed the basinyms, and cited specimens for each of his entities. Mueller? reseparated Stillingia and Sapium, and established Gymnostillingia as a separate genus, based on S. acutifolia. Later!? he reduced Sapium to the status of section іп Excoecaria, but maintained both Sfillingia and Gymnostillingia as separate genera. Bentham" placed Gymnostillingia without rank within the genus S/illingia, recognizing for the first time that the gynobase is the most important generic character. Mueller’ had previously noted the occurrence of this structure, but failed to appreciate its importance as a linking character between Stillingia and Gymnostillingia, 5. Watson!? placed the herbaceous species of the southwestern United States and northern Mexico within the genus S/illingia, allied to S. acuti- folia, but he did not definitely state whether these species should be treated as a section or as a subgenus, Рах and Hoffman! established a new section, LEPTO- x L. Mant. 1767. UA Gen. РІ. ty ae 1836-1840, Б parte. 9Klotzsch, in Wiegm. (Erichs.) Arch. 7:1 1841. 7 Baillon, Ét ud. Сеп. Euphorb. 510. ји ех parte 5 Baillon. in Adatwonla 1:350. 1861; 2:27. 1861; 3:162. 1862; 5:320. 1865. Mue rg. in РЬ 13Pax an Hoffmann, i in Engle; М! Iv. Tu ge} V:194. 1912. 1951] ROGERS—REVISION OF STILLINGIA 209 STACHYAE, for these four species. Pax and Hoffman" were the first to divide the tribe Hippomaneae; Stillingia appears as а member of the subtribe Stillingiinae. These authors have given the limits of the genus as it is known today, dividing it into six sections based largely on vegetative characters. I have not altered the generic limits of S/illingia since there is no confusion with the allied genera. Within the genus, however, new subgeneric categories are employed to emphasize the relationships and differences between the speciés groups. The subgenus is used in order to demonstrate a wide morphological diver- gence of two large species groups, and the series is used in an effort to show that boundaries between the groups within the subgenus are not distinct, and are at best tendencies toward morphological divergences. This seems to be a more logical division than to place all of the species groups on one level (as sections), indicating nothing of the stages of differentiation on a scale above the species. GENERIC RELATIONSHIPS Within the tribe Hippomaneae, Sebastiania, Excoecaria, Maprounea, and Sapium are the genera which have been most frequently confused with, and seem to be the most closely related to S/illingia. I am not prepared to comment upon the correct phylogenetic sequence of these genera, nor upon the position of Stillingia with reference to them. The genus Sapium has the closest morphological re- semblance to Stillingia, and there has been some confusion in the past concerning the characters which separate these two genera. In Sfillingia, ап abscission layer forms above the base of each of the cocci so that after dehiscence, a three- or two- rayed hardened portion of the pericarp remains attached to the pedicel. The hard- ened portion of the pericarp, called the gynobase in this study, is the most promi- nent distinguishing feature of Stillingia. None of the other genera have this structure. It is slightly more difficult to separate Stillingia from Sapium when flowering specimens without mature fruits are available. The following list of characters will serve to differentiate the two genera with ease when all of the characters are considered together (cf. fig. 1). Any one of these may fail if taken individually: Stillingia Sapium Pistillate Flower Sepals separate (except in S. saxatilis), Sepals united. or absent. Seed Testa hardened, smooth to rugulose, not Testa arillate in American species. arillate. Vegetative Characters Herbs, shrubs, or occasionally small trees. arge trees, infrequently subshrubs. соси, ас Баѕе leaf present or absent, Glands at base of leaf mostly present, mostly rar elongate, ritu or scutelli- elongate, tu pesci or cL. the o неу for of the glands wide, constricted, usu pposite, more a part o ые, ұй aes spac the petiole thee м de leaf blade. 14Pax and Hoffmann, loc. cit. 180. 1912. [Vor. 38 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN The three remaining genera are easily distinguished from S/illingia: Sebastiania and Excoecaria by the three to several stamens of the staminate flower, pistillate flower long-pedicellate, sepals united, gynobase absent; Maprounea by its compact inflorescence which appears almost as a capitulum, pistillate flower long-pedicellate, without gynobase. The following is a synopsis of Stillingia in the Western TNR as pro- jected in this study. oppositifolia beers ex n Arg. Il е.) D. J. Rogers peru D. у. p^s gers silica id Muell „А microsperma Hoffm. diphtherina D. И "ha ogers bicarpellaris S. Wats. aquatica Chapm. Series OPPOSITIFOLIAE эчи уыт Pp SHOR а Pax & Hoffm. Subg. STILLINGIA Series DICHOTOMAE й | . saxatilis Muell. PP Series SYLVATICAE . salpingadenia € Arg) Huber : 2. ета JR Dusenii P 13. 9 14. S 15. S ax & 16. S. zelayensis ты мыр Arg. 17. S. £ 18. S ы С нен ех "4 $ $ S S 5. Series 19. S. acutifolia (Benth.) Benth. ex Hemsl. |UTIFOLIAE b m 20. S. Treculiana ves a I. M. Johnst. Subg. GYMNOSTILLINGIA S 21. S. денесін 5. d 22. S. spinulos TRECULIANAE | 23 pss 5. “Wats. SYSTEMATIC CRITERIA The genus Séillingia is a diverse group of perennial herbs, subshrubs, shrubs, S. acutifolia, of southern Mexico and Guatemala, and S. oppositifolia, of southeastern Brazil, are the only species which have sufficient height and diameter of trunk to be called trees. and small trees. The tree habit is rare in the American species: Stems of the species of all series except svLvATICAE arise directly from a tap root. Although seldom found on herbarium specimens, the woody root ‘crown or The ad- ventitious roots of S. sylvatica and S. aquatica are distinctive, those of the former being stoutly elongate-fusiform, arising at wide intervals along the rhizome, and those of the latter filiform and very closely set on the primary root. The branching systems are opposite, alternate, approximate, or fascicled, the latter being a result of shortened internodes subtending the inflorescence and thus presenting a somewhat whorled appearance. rhizomes of species of svLvATICAE are, nevertheless, of diagnostic value. The fascicled branching pattern gives the upper portions of the plant a corymbiform appearance. The bark of the 1951] ROGERS—REVISION OF STILLINGIA т 211 е f g h i Fig. 1. Illustrations of taxonomic criteria for S/illingia and Sapiu Seed: тараны scutellifera, testa hardened; b—Sapium hae APER. JE testa. arillat Pistillate flower: c—Sapium marginatum, the sepals united; d—Stillingia salpingadenia, uj шені. separate. Pedicel after dehiscence of fruit: e—Séillingia sylvatica ssp. sylvatica, the gynobase present; f—Sapium HA M without gynobase. eaf: debes. scutellifera, with 2 basal scutelliform or cyathiform glands; h—S/illingia aquatica, E glands at base of blade; i—Sapium pedicellatum, with 2 basal, tubular glands. shrubby species is lenticellate in varying degrees, with the exception of those of series DICHOTOMAE, whose succulent species have no lenticels. S. sanguinolenta has an unusually rough bark due to the presence of a large number of lenticels. Succulence of stem and leaf is a definitive character of the species of series DICHOTOMAE. The leaves are widely spaced on the stem, except in S. Bodenbenderi and $. peruviana. In these species the leaves may be widely spaced but frequently they may arise in groups on peg-like short shoots. The outline, margin, venation, and texture of the leaves are important key characters. The leaves of the Mexican, Central and South American species of the subgenus SriL.LINGIA, with few excep- tions, are provided with two to several cup- or boat-shaped glands at the base of the blade. This is not a constant feature of each leaf, but generally more than one leaf of a standard herbarium specimen will have these modified serrations. The types of leaf texture occurring in Sfillingia are: thin to firmly membranaceous, definitely coriaceous, and sücculent or fleshy. "These conditions are usually char- acteristic of whole groups of species, but S. diphtherina is the only known species with coriaceous leaves. [Vor. 38 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN The organization of the inflorescence, a spiciform thyrse, is essentially the same throughout the genus. The flowers are monoecious, with the pistillate flowers always solitary in the cymule, but the number of staminate flowers in a cymule varies. The pistillate flowers are attached toward the base of the peduncle and either immediately subjacent to the lowest staminate cymule or with a distinct space on the peduncle separating the pistillate and staminate cymules. The in- florescence may be pedunculate below the point of insertion of the lowest pistillate flower, or sessile. Both the pistillate and the staminate cymules are subtended by a bract the shape of which is of some importance in species differentiation. The bracts are squa- maceous in all species except S. Dusenii, in which the bract of the pistillate cymule seems to be somewhat foliaceous. The bracts are subtended by two patelliform, cyathiform, tubiform, or urceolate marginal glands. Generally, these glands are simple, but in S. oppositifolia they may be twice or three times branched. These glands are considered as homologues of the cup-shaped serrations at the base of the leaves of several species. The staminate flowers are borne in groups of 3 to 13, or singly in the axil of a bract toward the apex of the inflorescence. This inflorescence structure is assumed to be derived from a more primitive type by reduction. The inflorescence structure of Hevea, for example, with several pistillate flowers on the primary and secondary axes, might represent a more primi- tive condition than that observed in Stillingia where the pistillate flowers are single in the cymules. A further reduction series is evident within the genus Stillingia in the staminate cymule structure. The species of subgenus STILLINGIA are char- acterized by three to many staminate flowers in each cymule, but the subgenus GYMNOSTILLINGIA has only one staminate flower for each cymule. A strengthening feature for the distinction between the two subgenera is found in the sepals of the pistillate flower. All of the species of subgenus STILLINGIA have three (or two in S. bicarpellaris) sepals which are distinct, or fused into a truncate annulus in S. saxatilis, and the remnants of these are occasionally apparent at the base of the gynobase even at maturity of the fruit. The name GYMNO- STILLINGIA is derived from the absence of a calyx, and all species of the subgenus except S. acutifolia are completely lacking it. S. acutifolia is a transitional species in this respect, with very small and fugacious sepals. No primary systematic importance has been attached to the staminate flowers since they are rather uniform and present no readily observed differences. Оп the other hand, relative congestion of the staminate cymules and the total number of staminate flowers of an individual cymule are used in the diagnostic treatment. The pollen grains are spheroid or nearly so, with three pores, except in species of series DICHOTOMAE in which they are ellipsoid with one lateral pore. The mature fruit of Stillingia affords the most striking feature of generic dis- tinction, but presents few characters for species diagnosis. The gynobase (described 1951] ROGERS—REVISION OF STILLINGIA 219 in the section оп Generic Relationships) is a constant feature of all species of Stillingia. The lobe length of the mature gynobase is consistently given in the descriptions, being measured from the central column to the apex of the lobe. The variability of this character within a species prevents inclusion as a key character. Seed size, contour, and surface are constant within the species, for the most part. Again, variation occurs notably in the extreme southeastern part of the range of S. sylvatica where introgression with S. aquatica is reflected in the vari- ability of the seed. The presence or absence of the caruncle is one of the best supporting characters for the division of the genus Stillingia into two subgenera. Subgenus STILLINGIA is characterized by the presence of a well-defined caruncle in all species. Subgenus GYMNOSTILLINGIA is not completely without this body, how- ever, and a very definite caruncle appears in S. Treculiana. Stillingia acutifolia, S. spinulosa, and S. linearifolia are ecarunculate; S. paucidentata is a transitional species in this respect, with a very minute and fugacious, but nevertheless definite caruncle. Attempts to count the somatic chromosomes of S. sylvatica using Perry's? techniques were not particularly successful, but were sufficiently good to show that a number in excess of that reported by Perry (2n — 36) were present in the specimens examined. Since this might indicate a polyploid race within the species S. sylvatica, an effort to correlate pollen grain size with polyploidy was made. Although there was some variability in pollen grain dimensions, there was no indi- cation of two different races, the range of variation being continuous. SUBGENERIC CATEGORIES The subgenus Srinata includes those species with 3 to 13 staminate flowers in a cymule, 3 to 2 definite sepals in the pistillate flowers, and a well-defined car- uncle on the seed. Species with a single staminate flower in a cymule, sepals and caruncle absent, are referred to subgenus GYMNOSTILLINGIA. There is a sufficient overlap of characters between the two subgenera, however, to prevent the estab- lishment of distinct genera. The series within both subgenera are divided largely on vegetative characters with only a few correlating floral characters. In subgenus STILLINGIA the first and most primitive series, OPPOSITIFOLIAE, is a group of woody shrubs or small trees with membranaceous or coriaceous leaves and spheroid pollen grains. The indi- vidual species of this series are quite distinct morphologically. Series DICHOTOMAE, a group of four shrubby species, is characterized by mostly succulent stems and leaves, and by ellipsoid pollen grains with one pore. This interesting group is poorly known and infrequently collected. Stillingia Uleana seems to be the connecting species between the species of series DICHOTOMAE and those of opposiTIFOLIAE. Further collections from eastern Brazil, to which region this group is confined, may throw more light on the inter-relations of these species. 15 Perry, in Am. Jour. Bot. 30:527. 1943. [Vor. 38 214 ^ ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 3. Inflorescence and flower structure of Stillingia paucidentata, typical of subgenus GYMNOSTILLINGIA, The third series of subgenus STILLINGIA, SYLVATICAE, is composed of the best known and most extensively collected group of species within the genus Sfillingia. This series represents a different type of specialization from that of series picHo- TOMAE. All of the species are suffruticose, but their floral structure is very similar to that of the less specialized species of ОРРОВІТІБОГЛАЕ, from which this group seems to be derived. Although the species are very closely related, they seém to have diverged and stabilized themselves sufficiently to be called species rather than subspecies of one large "cenospecies". Evidence of their close relation is seen not only in morphological characters, but also in the fact that they have apparently erected none or few genetic barriers to prevent rather free hybridization where their geographic margins overlap. Because of these putative hybrids, many names 1951] ROGERS—REVISION OF STILLINGIA 422 һауе been applied previously without any thought as to the explanation of the phenomenon, nor of the ensuing confusion. The three South American representatives of svLvATICAE, S. salpingadenia, 8. scutellifera, and S. Dusenii, are widely separated geographically from the three North American species. Although easily distinguished morphologically from 8. sylvatica of the southern United States, S. scutellifera resembles the Texan repre- sentative of the former in many particulars, both ecologically and morphologically. In only one case have I seen fit to designate a subspecies in this series, that of S. sylvatica ssp. tenuis, which occurs only in extreme southeastern Florida. Suf- ficient material, together with personal knowledge of the habitat through field studies, makes this possible. There seems little doubt, however, that at least one species in South America, S. өси/е! (ста, has a closely related form, exemplified by Hassler 5612 and called saxatilis var. salicifolia by Chodat and Hassler!", which possibly could be designated as a subspecies. Lack of sufficient knowledge makes such a step premature. There seems to be little or no genetic barrier between S. aquatica of Series OPPOSITIFOLIAE and S. sylvatica of Series sYLvATICAE. In the spring of 1950 local population samples were made at several points in Florida to determine the possi- bility of hybridization between such widely differentiated species. "The assembled specimens were studied and indexed according to techniques used by Anderson". Although larger samples would have been desirable, those actually made give rather positive evidence that free hybridization does occur. The subgenus GYMNOSTILLINGIA is composed of two series which are widely differentiated. Series ACUTIFOLIAE has one shrubby species, S. acutifolia, with broad, thinly membranaceous leaves. The second series, TRECULIANAE, is a group of perennial herbs with a compact growth habit and for the most part very small, narrow leaves. There seems little doubt that the four species of this series form a natural group of plants which have become adapted to the extreme environmental 2а conditions under which they exist. GEOGRAPHY In the accompanying maps (figs. 4 and 5) are shown the known areas of distri- bution of the species of Sillingia in the New World. In general, the North Ameri- can material has been sufficiently ample to plot distributions with some accuracy, but there are still many gaps and disjunctions which may be filled with further col- lecting, particularly in Central and South America. For the most part, the species of S#illingia are limited to areas of temperate or subtropical climate, those occurring closest to the equator being at higher altitudes. In South America, S. Bodenbenderi reaches farthest south, in the Sierra de Cordoba in Argentina; S. sylvatica is the most northern species in North America, reaching the 38th parallel in southern Kansas. 16Chodat & Hassler, in Bull. Herb. Boiss. II, 5:676. 1905. 17 Anderson, Edgar. Introgressive Hybridization. New York. 1949. [Vor. 38 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN (ЧЕГЕТ v Lat acm SERIES 1. OPPOSITIFOLIAE и | Ioni E S 2. DICHOTOMAE | SERIE Fig. 4. Ranges in S/illingia. Although the series OPPOSITIFOLIAE is very widely distributed in both conti- nents, the individual species normally occupy small ranges. With the exception of S. aquatica the species of this series are found in mountainous regions in meso- phytic habitats, mostly at altitudes above 500 meters. They generally occupy land masses of both continents which have been the longest exposed and the longest continuously available for plant growth (Weeks!5, Schuchert!?). S/illingia aquatica, on the other hand, is the only species of the genus found in marshy or swampy habitats only a few meters above sea level in a region of relatively recent origin. ~ A3 Weeks, in Bull. Geol. Soc. Am. 59:249. І 19 Schuchert, Historical Geology of the Antillean-Caribbean Region. New York. 1935. 1951] ROGERS—REVISION OF STILLINGIA Lit Fig. 5. Ranges in Stillingia. The species of Series DICHOTOMAE are found only in the dry, upland regions of eastern Brazil. Their areas of distribution are the least understood of any in Stillingia. At least one species, S. trapezoidea, occurs in the caatinga, or open scrub forest of southern Piauhy, Brazil. The exact type of habitat is not known for the other three species of this series, but their adaptation to a dry habitat is shown in succulence of stem and leaf; this, together with their locality, marks them as a derived group. With the exception of S. zelayensis of the Mexican and Central American high- lands, species of the series sYLvATICAE are found in regions of lower elevation than [Vor. 38 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN those of series OPPOSITIFOLIAE, generally from near sea level to altitudes of 300—500 m. Sfillingia zelayensis, however, is confined to upland regions, from 1200 to 2800 m., mostly in open pine forests. The greatest distribution areas of the species of this series are in regions which were inundated by Cretaceous seas, and which today are mostly covered with open forests or savannas in North and South America. Stillingia acutifolia, the only species of series ACUTIFOLIAR, occupies a small area in southern Mexico and Guatemala. It occurs in the mountainous regions, within a definite mesophytic habitat. The four species of series TRECULIANAE are rather narrow endemics in the arid desert or semi-desert regions of the south- western United States and northern Mexico. STUDY MATERIAL The herbaria where specimens have been obtained for study, together with the symbols?" employed in their citation, are as follows: A—Arnold Arboretum of Harvard University, Jamaica Plain, Mass. AN—Colegio Anchieta, Porto Alegre, Brazil. BR—Jardin Botanique de l'État, Bruxelles. C—Universitetes Botanisk Museum, Copenhagen. F—Chicago Natural History Museum (Field Museum). FLAS— Agricultural Experiment Station Herbarium, University of Florida, Gainesville. G—Institut de Botanique Systematique de l'Université, Genéve. GH—Gray Herbarium of Harvard University, Cambridge, Mass. K—Royal Botanic Gardens, Kew. LIL—Instituto Miguel Lillo, Tucumán, Argentina. MICH—University of Michigan, Апа Arbor. MO—Missouri Botanical Garden, St. Louis. NY—New York Botanical Garden. OKL—Bebb Herbarium, University of Oklahoma, Norman. P—Muséum National d'Histoire Naturelle, Paris. S—Naturhistoriska Riksmuseet, Stockholm. TEX—University of Texas, Austin. UC—University of California, Berkeley. US—United States National Herbarium, Smithsonian Institution, Wash- ington, D. С, I wish to acknowledge my indebtedness to the curators of these institutions, and to the Director and Staff of the Missouri Botanical Garden where this study was made. 29 The symbols used are as suggested by Lanjouw in Chron. Bot. 5:143. 1939, 1951] ROGERS—REVISION ОҒ STILLINGIA | 2%? TAXONOMY STILLINGIA Garden, ex L. Mant. 19. 1767; Baillon, Étud. Сеп. Euphorb.. 510. 1858, ex parte; Muell. Arg. in DC. Prodr. 15?:1155. 1866; Benth. in Benth. & Hook. f. Gen. Pl. 3:334. 1880; Pax, in Engl. & Prantl, Nat. Pflanzenf. ІП, 5:96. 1890; Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147. V:180. 1912. Seborium Raf. Sylva A 63. 1838. Ditrisynia Raf. loc. cit 183 Glabrous perennial herbs, shrubs, or small trees. Leaves simple, alternate, opposite, or verticillate, usually with 2 to several filiform, glandular stipules, en- tire to serrate, with or without 2—3 cyathiform or scutelliform glands at base of blade, the serrations glandular or occasionally the glands in the sinuses of the serra- tions. Inflorescence a terminal, spiciform thyrse, the cymules bracteate, the bracts with 2 marginal glands; flowers monoecious, monochlamydeous or naked. Pistillate flowers solitary, borne toward the base of the peduncle; sepals 3, rarely 2, mostly separate, infrequently united into a truncate annulus, imbricate, or the sepals ab- sent; ovary superior, 3- to 2-celled, a single pendulous, anatropous ovule in each cell; stigmas 3, rarely 2, recurved, more or less coalescent at the base into a single style. Staminate flowers solitary ог 3—13 in a cymule, occupying the upper por- tion of peduncle, the 2-lobed calyx always present; stamens 2, exserted, the fila- ments coalescent at the base, the 2 thecae adnate, extrorse, longitudinally dehiscent; pollen spheroid to ellipsoid, with 1—3 pores, the exine reticulate, granular, or punc- tate. Fruit a 3- to 2-celled, dry, septicidally dehiscent capsule, the accrescent gynobase 3- to 2-lobed, hardened and persistent after dehiscence of carpels; seeds 1 in each carpel, with or without a micropylar caruncle, the embryo central, the cotyledons broad and flattened, the endosperm mealy. Standard species: Stillingia sylvatica Garden, ех L. Mant. 19. 1767. KEY TO THE SUBGENERA AND SERIES A. Staminate flowers in clusters of 3 or more; sepals of pistillate flowers pr Sub esent, persistent; caruncle present Subgen. I. STILLINGIA (p. 220) ru r EN or Foe from a tap root; stems perennial, woody or su rts gray- Pes wil and frequently some- what blacke eg ге к Шо. obviou C. Stems woody; leaves membranaceous or coriaceous; pollen spheroid, with ores; tropical and subtropical Americas, exclusive al eastern and northeastern Brazil er. l. OPPOSITIFOLIAE (p. 220) GG Pd mostly succulent; leaves succulent; pom “рен wit 1 lateral pore; eastern eus ortheastern Bra 2. DICHOTOMAE (p. 230) BB. Sub ігі arising from an enlarged woody base; stems most Db nual or biennial, he rd ceous or subherbaceous, green to reddish s usually without lenticels 3. SYLVATICAE (p. 233) AA. Staminate flowers single; sepals of pistillate flowers absent, or re and fugacious; caruncle absent (except in $. Treculiana and S. pencidentata) ubgen. П. GvMNosrILLINGIA (p. 243) Shrubs or small trees, ee ME petiolate, the pin- nate venation prominent; sophytic habitats, Central i Ser. 4. ACUTIFOLIAE (p. 243) Americ DD. Tni herbs; cin sessile to fai on the venation not prominent (except S. spinulosa with palmate venation); xero- ie habitats, Sous eae United po and northwestern exico Ser. 5. TRECULIANAE (p. 245) [Vor. 38 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN Subgenus I. SriLLINGIA $ Eustillingia Kl. in Wiegm. (Erichs.) Arch. 7:187. 1841; emend. Muell Arg. in Linnaea 32967. 1869; Series 1. OPPOSITIFOLIAE D. J. Rogers, ser. nov $ Fruticosae Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, V:186. 1912. Caules lignosi; folia membranacea vel coriacea; pollen sphaeroideum, foramini- bus 3. Americae tropicae et subtropicae Brasilia orientali excepta. KEY TO THE SPECIES A. — usually broadest at the middle or above; secondary roots widely on the p grum not thickly ped seeds ellipsoid to ovoid, 3—6 pe чу мар» ^ ooth; tropical and subtropical America, ex- б a on well-drained soils. B. Pistils 3 hrak leaves narrowly to db elliptic; seeds rela- lowest жеге» cymule, the cymules decussate; glands of pistillate Lo ts frequently trifurcate; leaf margins finely to coarsely serrate; ; brancher ү dichotomous, never fascicled; keren stern Bra unculate below the 22 ien Mere the of pisti Шаке bracts sim margins venly serrulate; branches no dan ds dicho- tomous, sometimes fascicled. D. Leaves arisi sing in жиз on peg-like short shoots, or alternate ipeo outh Amer ves 1. S. oppositifolia асн petiolate to subsessile; bracts of staminate without m ; Argentina and Brazil . $. Bodenbenderi EE. Leaves “atively Tong- саа bracts of staminate cymules ate; 3 ‚ S. peruviana DD. te ‘solitary and pri че arising on main axis; Mexico and entra pak densely кенен branches opposit approximate, never fascicle di hogy aid 3-lobed; Mexico.. 4. FF, Bark sparsely enc bra requently fascicled; и іс shallo wly 3-lobed; Central e or occasionally S. sanguinolenta . long, 2—4 42. 5. S. microsperma GG. Leaves acute, coriaceous, 1.8— i үзе ст. long, 0.5 broad petioles short, 0. e cy .0 cm. ong; inflorescence open, the cymules distant, relatively few-flowered........ 6. S. diphtherina BB. Pistils 2-, rarely Nisi sen a linear- ед udi larger, mm. 7. S. bicarpellaris АА. Leaves usually br бекінді құлы the base; secondary roots — set n the primary, forming a mat; seeds subspherical, 2-3 mm. in diameter, rugose; Georgia ur "Florida, in swampy places or А intermittent ponds 8. S. aquatica STILLINGIA OPPOSITIFOLIA Baill. ex Muell. Arg. in DC. Prodr. 152:1160. 1866 (T.: Sellow 4985!). к. . Shrubs or small trees 1—5 т. tall; stems woody, frequently branched, the branches opposite, terete, slender, sparsely lenticellate, gray-brown, frequently somewhat blackened on older parts. Leaves decussate, widely spaced, petiolate, membranaceous, yellowish-green, elliptic or spathulate to narrowly rhombic, 3—9 1951] ROGERS—REVISION OF STILLINGIA ЖЕТ cm. long, 1-5 cm. broad, apex rounded to acute ог acuminate, base acute, cyathi- form glands infrequent, finely to coarsely serrate, the midrib prominent, secondary venation not obvious; petiole 0.3—1.2 cm. long, sulcate above. Inflorescence 1—2 cm. long, usually sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules decussate, crowded, distinctly separate upon the peduncle; bracts of pistillate cymule caudate-acuminate, the staminate broadly rhombic, with 3 mucros, the glands urceolate, the pistillate frequently trifurcate. Sepals of pistillate flower 3, linear-lanceolate, 2 mm. long or less; ovary sessile, 3-carpellate; styles 2-3 mm. long. Staminate cymules 3- to 5-flowered, the flowers subsessile, about 1-2 mm. long; calyx shallowly 2-lobed, the lobes entire; pollen spheroid, with 3 pores, the exine coarsely reticulate. Fruit deeply 3-lobed, about 5-6 mm. wide, the lobes of the gynobase 2-3 mm. long, seed not seen. A common shrub in Araucaria woods, between 600 and 1000 m. alt., with average rainfall of 1750 to 2500 mm., 15—17° C. average temperature, with oc- casional snowfall and —5? C. during winter. Flowers and fruits from October to March. BRAZIL: MINAS GERAIS: ad Lagoa Santa in silvis super rupes calcar., Warming 1525 (С, б, GH). RIO GRANDE DO SUL: Montenegro, S. Salvador, Friedrichs 32938 (LIL, S); San Francisco de Paula, Rambo 2278 (LIL), Rambo 44828, 46236 (АМ); Kappesberg, pr. Montenegro, S. Salvador, in silva, Rambo 2278 (AN, LIL), Rambo 43828 (AN); Passo do Inferno, pr. San Francisco de Paula, Rambo 4818, 4824 (AN); Nova Petropolis, pr. vut Rambo 6575 (AN) ; Nonoae ad fl. Uruguay superius, ad araucarietum, Rambo 28353 (АМ, МО); Caracol, pr. Canela, Rambo 28809 (AN); San Francisco de Paula, Vila Oliva, in abs Rambo 31144 (LIL); Bom Jesus, Facenda Bernardo Velho, in silva campestri, Rambo 34775 (AN, LIL, S); Vila Oliva pr. Caxias, Rambo 4313 36 (AN); Gramado, pr. Canela, Rambo 44085 (AN). WITHOUT LOCALITY: "Brasilia meridionali", Pi (Sello) 4985 (G, K, P I am indebted to Fr. Rambo for his data on the habitat and distribution of this species. Baillon?! published Stillingia oppositifolia Kl. as a nomen nudum, apparently taking this name from a specimen in the Berlin Herbarium annotated Sapium oppositifolium by Klotzsch. In the Atlas published with his Étud. Gén. Euphorb., Baillon designates the illustrations of Plate V, figs. 24 and 25 as Stillingia (Sapium) oppositifolia Kl. However, in the text (p. 513) he lists S. oppositifolium Kl. to- gether with ten other species in which the “5” clearly stands for Sapium, since the endings of the specific epithets agree with the endings of the neuter noun Sapium and not with the feminine S/illingia. Of the ten other species listed on page 513 of the text, two are designated in the Atlas as Sapium. Baillon’s taxonomy in- cludes all of these species under S/illingia, but he apparently did not make the necessary transfers in nomenclature. The name Stillingia oppositifolia was validated by Muell. Arg. in DC. Prodr. 157:1160. 1866. 21 Baillon, Étud. Сеп. Euphorb. 513. 1858. [Vor. 38 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. $тиллмсл Bodenbenderi (О. Ktze.) D. J. Rogers, comb. nov. Sapium Bodenbenderi O. Ktze. Rev. Gen. 32:292. 1898. (T.: Bodenbender 69021). Excoecaria Bodenbenderi (О. Ktze.) К. Schum. in Just’s Bot. Jahresb. 26':349. 1898. Sapium subsessile Hemsl. in Hook. Icon. Pl. t. 2684. 1901. (T.: Weir 315). Shrubs or small trees; stems woody, frequently branched, the branches alter- nate, opposite or approximate, sometimes fascicled, terete, slender, moderately lenticellate, gray or slightly reddish-brown. Leaves clustered on short side branches, alternate if single, crowded, subsessile, stiffly membranaceous, elliptic to obovate, 0.8—3.0 cm. long, 0.5—1.3 cm. broad, apex acute to rounded, base acute, usually with 2 cyathiform glands, callose-serrulate, the midrib prominent, the secondary venation obscure. Inflorescence 1-2 cm. long, slightly flexuose, the peduncle elongate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, widely spaced, not distinctly separate upon the peduncle; bracts of the pistillate cymule elliptic, mucronate, the staminate elliptic to ovate, rounded to acute, the glands patelliform. Sepals of pistillate flower 3, elliptic, mucronate; ovary sessile, 3-carpellate; styles unknown. Staminate cymules 5- to 9-flowered, the flowers subsessile; calyx shallowly 2-lobed, the lobes serrulate; pollen irregu- larly ovoid, with 3 pores, the exine reticulate to granular. Lobes of the gynobase 2-3 mm. long. Fruit and seed not seen. ARGENTINA: CORDOBA: Sierra de Córdoba, Sept., Bodenbender 6902 (NY, photo in F); Dept. de las Minas, Cuesta de las Chacras, Jan. 14, Hieronymus 817 (G, NY, US). Hemsley’s failure to recognize the affinities of his Sapium subsessile with Sapium Bodenbenderi O. Ktze. probably is due to the fragmentary nature of Bodenbender’s specimens. The plate of Sapium subsessile (in Hook. Icon. Pl. t. 2684. 1901), however, indicates that Hemsley's species is synonymous with Sapium Bodenbenderi and, furthermore, that the specimen is Stillingia, not Sapium. These drawings show a calyx of three separate sepals and a leaf base with cyathiform glands, both of which are characteristic of Stillingia rather than Sapium. n addition, the col- lection of Hieronymus (no. 877), referred by Hemsley to Sapium subsessile, pos- sesses a gynobase, one of the best morphological distinctions of Stillingia. Huber, in an appendaged note in Bull. Herb. Boiss. II, 5:452. 1906, first noted the affini- ties of Hemsley’s species со Stillingia, but he did not make a formal transfer. It is difficult to understand why Pax and Hoffmann failed to make the transfer to Stillingia, but maintained both Sapium Bodenbenderi O. Ktze. and Sapium subsessile msl I have not seen Weir 315 (the specimen at Kew having been temporarily mis- placed), and the wide geographic separation of this specimen (collected in the state of São Paulo, Brazil) from those of Bodenbender and Hieronymus (State of Córdoba, Argentina) is hard to explain. This may be a relict species on the old land masses of southern Brazil and in the Sierra de Córdoba, Argentina. 1951] ROGERS—REVISION OF STILLINGIA 223 Fig. 6. Stillingia peruviana. 3. STILLINGIA peruviana D. J. Rogers, spec. nov. (Т.: Stork & Horton 10014!). Frutices 2 m. alti, caulibus lignosis saepe ramosis; rami approximati vel fascicu- lati teretes graciles, cortice parce lenticellato cineraceo-brunneo, succo lacteo. Folia in ramulis perbrevibus congesta aut alternata si singularia petiolata mem- branacea anguste elliptica vel lanceolata apice baseque acutis, 2.0-3.5 cm. longa, 0.5—0.8 cm. lata, subtiliter serrulata vel crenulata, basi glandulis 2 minutis cyathi- formibus vel nullis, nervo medio prominenti venis secundariis in superficie inferiore manifestis, petiolis gracilibus 2-4 mm. longis supra sulcatis. Inflorescentia ca. 2.5 cm. longa constanter fusco-rubra sub cymula pistillata ima pedunculata, regione superiore staminali ab inferiore pistillato in pedunculo non distincte separato, cymulis pistillatis remotis staminalibus congestis, bracteae cymularum pistillatarum caudato-acuminatae marginibus saepe involutis, bracteae cymularum staminalium [Vor. 38, 1951] 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN late rhombeae mucronulatae, glandibus disciformibus sessilibus simplicibus. Sepala floris pistillati tria 1 mm. longa minusve. Ovarium sessile 3-carpellatum, stylis 3 mm. longis. Cymulae staminales 5- vel 7-florae; floribus subsessilibus ca. 1—2 mm. longis, calyce tenuiter 2-lobato, lobis serrulatis; pollen sphaeroideum foramini- bus 3 exosporiis granularibus. Fructus profunde 3-lobatus ca. 5 mm. latus, lobis gynobasis ca. 2 mm. longis; seminibus ellipsoideis vel ovoideis ca. 3 mm. longis 2.5 mm. latis laevibus pallide brunneis, caruncula parva sub micropylo affixo. Fairly common in shrubland along rivers at altitudes from 2500 to 2900 m. Fruit said to be edible. PERU: HUANCAVELICA: Prov. Tayacaja, Mantaro Valley, near La Mejorada, rainy green shrubwood, Mar. 21, Weberbauer 7605 (GH); 4 km. south of Mejorada, Mar. 14, Stork & Horton 10918 (F, HOLOTYPE). The nearest relatives of S. peruviana occur in the Sierra de Córdoba, Argentina, and in southern Mexico and Central America. In evolutionary sequence, it prob- ably is a connecting link between the more primitive shrubby species of the moun- tains of southern Brazil and the more advanced shrubs of the mountains of Central America and Mexico. The distinctive features of this species are its narrowly elliptic to lanceolate leaves, frequently grouped on short side branches, and its deep red, slender inflorescence. Vernacular name: Cabra-cabra (Peru). 4. SrILLINGIA SANGUINOLENTA Muell. Arg. in Linnaea 32:88. 1863. (Т.: Ebren- erg s. n.) Stillingia sanguinolenta a. lanceolata Muell. Arg. loc. cit. 1863. (T.: ibid.). Stillingia sanguinolenta B. angustifolia Muell. Arg. loc. cit. 1863. (T.: Schiede? [Ebren- berg] 12 Shrubs 1—3 m. tall; stems woody, frequently branched, the branches opposite to approximate, terete, slender, the lenticellate bark roughened, the sap milky. Leaves opposite, widely spaced, petiolate, membranaceous, narrowly to broadly elliptic, 1.7-8.6 cm. long, 1.0-2.6 cm. wide, apex acuminate to acute, base acute, usually with two cyathiform glands, serrulate, the midrib and secondary venation prominent; petiole 0.2-1.0 cm. long. Inflorescence 3.0-6.3 cm. long, shortly pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, widely spaced, not distinctly separate upon the peduncle; bracts of the pistillate and staminate cymules cuspidate, about 1.5-2.0 mm. long, the cyathiform glands sessile. Sepals of the pistillate flower 3, cuneate, serrulate, about 2 mm. long; ovary sessile, 3-carpellate; styles 2-4 mm. long. Staminate cymules 7- to 9-flowered, the flowers subsessile, about 2 mm. long; calyx shallowly 2-lobed, the lobes serrulate; pollen ellipsoid, circular in cross-section, with 3 pores, the exine reticulate to coarsely punctate. Fruit deeply 3-lobed, 6-8 mm. wide, the lobes of the gynobase 3—4 mm. long; seeds ellipsoid, about 4.5—5.0 mm. long, 4 mm. wide, the testa slightly rugulose, the base flattened, the caruncle small. ROGERS—REVISION OF STILLINGIA 225 Fig. 7. Stillingia sanguinolenta. Fig. 8. Stillingia microsperma. In deep leaf mold, among rocks, dense oak woods of canyon floor, openly wooded arroyos, on lower slopes, river gravel, hills, and occasionally in desert regions, at altitudes from 500 to 1000 m. Flowers from about the middle of March through June, sometimes to October, and fruits from about the first of June through the last of August. MEXICO: TAMAULIPAS: Sierra de San Carlos, vic. of San Miguel, Bartlett 10570 (F, US); Sierra de San Carlos, vic. of El Milagro, Bartlett 11025 (F, GH, US). NUEVO LEON: Municipio de Villa Santiago, Mueller 2026 (F, MICH, MO, TEX); Canyon Diente, near Monterrey, Mueller 2668 (СН, UC); on Pan American Highway, Monterrey, Frye 9 5 ^ F, СН, MO, NY, UC, US). san Luis Potosi: Guascama, Purpus 4980 (F, GH, MO, UC, US); Sierra Tablon, Purpus 5464 (F, GH, MO, NY, UC, US); Alvarez, E. Palmer 234 (GH, NY, US). нірлісо: Zimapán, Kenoyer s. n. (MICH, МО); foot of Chipique (Hidalgo?), Kenoyer 266 (Е). cHrAPAs: Comitan, Goldman 902 (US). PUEBLA: vic. of San Luis Tultitlanapa, Puebla, near Oaxaca, Purpus 3523 (UC). STATE AND LOCALITY UNKNOWN: Coulter 1502 (GH); Pavon s. n. (D). The identity of the collector of the type specimen of S. sanguinolenta В. angustifolia is doubtful. Mueller cites "Schiede? 1245", but Pax in Engl. Pflanzenr. V, Fam. 147, V:191. 1912, cites Ebrenberg 1245, with the same locality as Mueller’s. Since this specimen was in the Berlin Herbarium, it is safe to assume that Pax cited the correct collector. Although the specimen cited by Mueller for B. angustifolia was unavailable, it is evident from his descriptions that the varieties are based on leaf variations. — In the material available for this monograph are specimens which show that there is no definite break from the smallest to the largest leaf, and no geographical basis [Vor. 38 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN for separating the varieties from the parent species. In addition, the floral struc- ture is rather stable in all specimens examined, without sufficient evidence for maintaining any subsidiary entities. 5. STILLINGIA MICROSPERMA Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, V:187. 1912. (Т.: Heyde 9 Lux 4265!). Open shrubs 1.0-2.5 m. tall; stems woody, sparsely branched, the branches opposite to fascicled, terete, slender, sparsely lenticellate, the sap milky. Leaves opposite to approximate, widely spaced, petiolate, membranaceous, elliptic, 7.5-12.8 cm. long, 2.3—4.0 cm. wide, apex acuminate to acute, base broadly acute to obtuse, usually with 2 cyathiform glands, finely serrate, the midrib and secondary venation prominent; petiole 0.4—1.1 cm. long, pale green above, paler beneath. Inflorescence 3.0-8.7 cm. long, pedunculate and somewhat thickened below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, not distinctly separate upon the peduncle, the pistillate distant, the staminate overlapping; bracts of pistillate and staminate cymules elliptic, mucronulate, about 1.5 mm. long. Sepals of the pistillate flower 3, elliptic; ovary sessile, 3-carpellate; styles about 2 mm. long. Staminate cymules 7- to 9-flowered, the flowers subsessile, about 1.5 mm. long; calyx deeply 2-lobed, the lobes serrulate; pollen spheroid to ellipsoid, circular in cross-section, with three pores, the exine finely punctate. Fruit about 8 mm. wide, globular, not deeply 3-lobed, the lobes of the gynobase about 4 mm. long; seeds ellipsoid, about 5 mm. long, 4 mm. wide, smooth or slightly rugulose, the micropylar end slightly flattened, the caruncle small. In open creek beds, oak woods, mountains, at altitudes from 1400 to 3000 m. on south-facing slopes, scrub-oak hillsides. Flowers from July through August, and fruits from August through November, and the first of January. Shrub said to have a disagreeable odor. Вкітізн HoNpunas: El Cayo District, in open creek bed, San Augustin, Lundell 6740 (C, F, GH, MICH, NY, S, TEX, US). Gu ALA: HUEHUETENANGO: along Aguacatan road cast of Huchuetenango at km. 13-14, Standley 82033 (F); Cumbre Papal, on south-facing slopes between Cuilco and Ixmoqui, Steyermark 50010 (F). ЈАТАРА: La Laguna, at base of Volcan Jumay, 1 mi. north of Jalapa, Steyermark 32201 (F); mountains along the road between Jalapa and Paraíso, Standley 77230 (F); no locality, Ruano 1358 (F). SANTA ROSA: Santa Rosa, Heyde & Lux 4265 (GH, US). 6. Ѕтпллмҹстл diphtherina D. J. Rogers, spec. nov. (Т. Williams & Molina 10003!). Frutices diffusi 1—2 m. alti, caulibus lignosis saepe ramosis; rami oppositi vel fasciculati teretes graciles rigidi, cortice moderate lenticellato cineraceo saepe nigro. Folia opposita breviter petiolata lanceolata vel anguste elliptica coriacea pallide viridia, odore resinae, 1.8-3.5 cm. longa 0.6-1.8 cm. lata, apice acuto vel rotundato base anguste obtuso, margine crenulato base glandulis 2 cyathiformibus, nervio medio prominenti, venis secundariis obscuris, petiolo 1-2 mm. longo. Inflorescentia 3-5 cm. longa parce flexuosa, pedunculo elongato sub cymula pistillata ima 1951] ROGERS—REVISION OF STILLINGIA 27 Fig. 9. Stillingia diphtherina. aliquando crassato, cymulis superioribus staminalibus et cymulis inferioribus pistil- latis spiralibus remotis; bracteae late ellipticae mucronulatae integrae, glandibus patelliformibus breviter pedicellatis. Sepala floris pistillati 3 late elliptica ca. 1 mm. longa subtiliter serrulata. Ovarium sessile 3-carpellatum, stylis ignotis. Cymulae staminales 7- vel 9-florae, tenuiter 2-lobatae lobis serrulatis; pollen sphaeroideum, foraminibus 3 exosporiis granularibus. Fructus ignotus. Lobi gyno- basis 3-4 mm. longi; semina ellipsoidea vel ovoidea ca. 4 mm. longa 3 mm. lata extremo micropylari parce complanato testa laevi basi rotundata caruncula parva. In barrancos in pine barrens, rocky limestone slopes at altitudes from 800 to 1400 m. Flowers in July, and fruits from August to the first of October. [Vor. 38 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN GUATEMALA: HUEHUETENANGO: Miis Nenton and Las Palmas, via Yalisjao, Rin- con EUN Chiaqui al, Guaxacana, in Sierra de los Cuchumatanes, S/eyermark 51648 (F). E: without loc «t Adler "6 (Е), 1106 (Е). Ном NDURAS: MORAZAN: west of Guinope, Williams 9 Molina 10603 (Е, MO ного- түре, UC); Tanque, Valerio 860 (F); Zamorano, Valerio 2158 (F); San Antonio de Oriente, Valerio 3130 (F, MO). This species is most closely related to $. microsperma, from which it may easily be distinguished by its small, leathery, short-petiolate leaves, the slender open inflorescence, and the stiff terete branches. Although none of the specimens cited have an attached fruit, it is apparent from the fragments contained in packets that the fruit is shallowly 3-lobed. м” imaa 8 Fig. 10. Sżillingia diphtherina. Fig. 11. Stillingia bicarpellaris. 7. STILLINGIA BICARPELLARIS 5. Wats. in Proc. Am. Acad. 21:455. 1886. (Т.: Pringle 128!). Shrubs 1.5-3.0 m. tall; stems woody, profusely branched, the branches alter- nate to approximate, terete, the bark slightly roughened, sparsely lenticellate, leaf scars prominent. Leaves alternate, closely spaced, shortly petiolate to subsessile, membranaceous, lanceolate to linear, 1.7-5.5 cm. long, 0.2-0.8 cm. wide, apex acute, the base acute, with occasional cyathiform glands, irregularly serrulate to entire, the midrib prominent, the secondary venation obscure; petiole about 1 mm. long. Inflorescence 1.5-2.9 cm. long, sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, not dis- tinctly separate upon the peduncle; bracts of pistillate and staminate cymules broadly elliptic, serrulate, mucronulate, the glands cyathiform, sessile. Sepals of pistillate flower 2, rarely 3, broadly cuneate to rhombic, serrulate; ovary sessile, 2-, rarely 3-carpellate; stigmas 2, rarely 3, style and stigma about 2—3 mm. long. Staminate cymules 6- to 7-flowered, the flowers subsessile, about 1 mm. long; calyx 1951] ROGERS—REVISION OF STILLINGIA 229 deeply 2-lobed, the lobes entire, mucronulate; pollen nearly spheroid, with 3 seg- ments, 3 pores, the exine granular to reticulate. Fruit slightly 2-, rarely 3-lobed, 6-8 mm. wide, the lobes of the gynobase 3—5 mm. long; seeds nearly spheroid, about 6 mm. long, 5.5 mm. wide, the testa smooth, shiny, the micropylar end very slightly flattened, the base rounded, the caruncle small. Thick underbrush in woodland associations of pines, oaks, and junipers, in mountains at altitudes up to 3100 m. Flowers and fruits from the last of June through October. EXICO: COAHUILA: summit of Mt. Jimulco, 13 kilo. east of Jimulco, Stanford, eiat kel - etii 94 (MO, NY, UC); mountains, Jimulco, Pringle 128 (A, BR, F,G . HIDALGO: near Ixmiquilpan, fate Painter & н $053 (US); ee nts pes 1454 (F, GH, MO, NY, UC). GUANAJUATO: "sur la montagne prés de Guanajuato," Duges 242 (F, US). PUEBLA: peris places, сап, Purpus 1320 (UC). If the dimerous structures of the pistillate flowers of S. bicarpellaris were con- stant features, a third subgenus might well be established for this species. Since, however, both dimerous and trimerous pistillate flowers are found on the same plant, I have not considered this procedure to be advisable. 8. STILLINGIA AQUATICA Chapm. Fl. South. U. S. 405. 1865. (T.: Chapman s. n. lectotype, US!). Erect shrubs 5.0-11.6 dm. tall. Primary root very short, the secondary fili- form, closely set on primary, forming a dense mat. Stems single, terete, tapering gradually from the base, leafless below the branches, the bark lenticellate, light gray to brown, the wood very light, the branches fascicled or dichotomous near the apex of the main axis, the leaf scars prominent on the older branches. Leaves alternate, crowded, mostly confined to the branch apices, petiolate, membranaceous, lanceolate to linear, 3.5—8.5 cm. long, 0.4—1.7 cm. wide, apex acuminate to acute, base narrowly obtuse, without cyathiform glands, irregularly crenulate to serrulate, the midrib prominent, the secondary venation obscure; petiole 0.1-0.6 cm. long. Inflorescence 3.0—3.8 cm. long, sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, distinctly separate upon the peduncle; bracts of pistillate and staminate cymules entire, sharply cuspidate, about 2 mm. long, the crateriform glands shortly pedicellate. Sepals of pistillate flower 5, rounded, slightly crenulate; ovary sessile, 3-carpellate; styles and stigmas about 4 mm. long. Staminate flowers 11-13 in a cymule, subsessile, about 1.0 mm. long; calyx shallowly 2-lobed, the lobes entire; pollen ellipsoid to ovoid, with 3 pores, the exine coarsely punctate. Fruit deeply 3-lobed, 5—6 mm. wide, the lobes of the gynobase 2.5-3.0 mm. long; seeds spheroid, 2—3 mm. in diameter, testa strongly rugose, the flattened micropylar end hippocrepiform, the base rounded, the caruncle small. In shallow ponds, wet pine barrens, edges of flatwoods ponds, cypress swamps, wet sandy soils, moist ditches, in the Everglades. [Vor. 38 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN / Fig. 12. Stillingia aquatica. UNITED STATES: GEORGIA: Sumter Co., FLORIDA: Brevard, ia Dade, Franklin, ntie Gadsden, Hendry, Lee, Palm Beach, Wakulla, and Washington cou t many of the specimens examined from Florida are of hybrid origin, but since No dou these specimens were not include e my own collection of S. aquatica owing introgression with S. sylvatica are жағы іп the herbarium of the Missouri аі G NITED сь FLORIDA: HENDRY CO.: between LaBelle and Immokalee, in edge of uen swamp, Rogers 8; near Immokalee, ME реи сургеѕѕ lrg R COLLIER CO.: 3 mi. se. of Naples, Rogers ACH CO 5 sw. Worth in small cypress pond, Rogers 14-2, oP h MARTIN CO.: Е "и $. ol Stuart, in ress pond, oe 15-4, 15-8, 15-0. (See also specimen citations of S. sylvatica ssp. 41). Ж айса, р. Vernacular name: Corkwood. Series 2. DICHOTOMAE D. J. Rogers, ser. nov. $ Pachycladae Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. V:186. 1912. Frutices; caules plerumque succulenti; folia succulenta; pollen ellipsoideum, foramine uno laterali.. Braziliae orientali. KEY TO THE SPECIES A. Stems um leaves crowded, spiral, with 2 cyathiform glands at the ase; staminate cymules 7- to 9-flowered AA. Stems ovni d leaves widely spaced, or whorled when closely set, 9. S. Uleana 1951] ROGERS—REVISION OF STILLINGIA 231 dn glands, or the glands scutelliform at base of leaf; staminate mules 3- to 7-flowered. B. "Banche alternate or fascicled; leaves long-petiolate, mostly alter- te, trapezoid 10. S. trapezoidea BB. Ph opposite; leaves short- dicas opposite or verticillate, narrowly to broadly elliptic or spathulat C. Stems enlarged at the ДА» opaque; enm broadly elliptic, bic" out glands; sepals disti . S. dichotoma CC. Stems ЛЕ: enlarged at the nodes, vernicose; leaves narrowly ы! o spathulate, with 2 basal scutelliform glands; sepals united into a truncate annulus 12. S. saxatilis 9. STILLINGIA ULEANA Рах & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, V:187. 1912. (T.: Ule 71351). Shrubs 2-6 m. tall; stems woody, branches infrequent, fascicled, slender, terete, sparsely lenticellate, opaque, gray-brown, frequently blackened. Leaves alternate, crowded, petiolate, succulent, elliptic to spathulate, 4—5 cm. long, 1.2-1.5 cm. broad, apex acute, base narrowly acute, crenulate, with 2 basal scutelliform glands and infrequent, irregularly spaced elongate-crateriform glands toward the tips, the midrib prominent, the secondary venation sunk in the fleshy tissue; petiole about 0.3—0.5 cm. long. Inflorescence 5—7 cm. long, slightly flexuose, pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate regions distinctly separate upon the peduncle, the cymules spiral, crowded, bracts of pistil- late cymule unknown, the staminate broadly cuneate to rhombic, entire, the glands clliptic, flattened, tightly clasping the peduncle. Sepals of pistillate flower un- known; ovary sessile, 3-carpellate; styles unknown. Staminate cymules 7- to 9- flowered, the flowers subsessile, about 2-3 mm. long; calyx shallowly 2-lobed, the lobes entire; pollen grains ellipsoid, with 1 pore, the exine coarsely reticulate. Lobes of the gynobase about 3 mm. long. Fruit and seed not seen. BRAZIL: BAHIA: Sincorá, Serra de Vendinha, 800 m., Ule 7135 (K, photos in F, GH, MO, US). 10. SrILLINGIA TRAPEZOIDEA Ule, in Engl. Bot. Jahrb. 42:233. 1908. (T.: Ule Shrubs 1—4 m. tall; stems succulent, angular, infrequently branched, the branches alternate or fascicled, slender, sparsely lenticellate, light gray- to reddish- brown. Leaves alternate, scattered, or crowded near insertion of inflorescence, petiolate, succulent, elliptic-trapezoid, 3—4 cm. long, 1-2 cm. broad, apex abruptly acuminate, base narrowly acute, without cyathiform glands, crenulate, the midrib and secondary venation immersed in the fleshy tissue; petiole 0.5—1.0 cm. long. Inflorescence 1.2-2.0 cm. long, slender, pedunculate below the lowest pistillate cymule, the cymules spiral; bracts of the pistillate cymules broadly elliptic, mucronulate, concave, the staminate broadly caudate-acuminate; the glands elliptic, convex. Sepals of pistillate flower 3, broadly elliptic to obovate, serrulate; ovary sessile, 3-carpellate; styles 2-3 mm. long. Staminate cymules 3- to 5-flowered, the flowers subsessile; calyx 2-lobed. Fruit about 7 mm. broad, deeply 3-lobed. Seed not seen. (Vor. 38 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN Details of the staminate flower have been taken from the descriptions given by Ule and Pax & Hoffmann. The specimen from Kew had no mature staminate flowers. BRAZIL: PIAUHY: іп der ca£inga^" in der Serra Branca, Jan., 1907, Ule 7160 (K, photos in F, GH, MO, US). 11. STILLINGIA DICHOTOMA Muell. Arg. in Linnaea 32:88. 1863. (T.: Riedel 1851). Shrubs to 1 m. tall; stems succulent, thickened, enlarged at the nodes, leaf scars prominent, branches opposite, without lenticels, opaque, gray- to reddish- brown, frequently blackened. Leaves verticillate, succulent, petiolate, broadly elliptic, 5-12 cm. long, 3-5 cm. broad, apex acute to shortly acuminate, base broadly acute, without cyathiform glands, serrulate to crenulate, the midrib prom- inent, the secondary veins obscure; petiole 0.5-1.5 cm. long. Inflorescence 4—8 cm. long, slightly flexuose, pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate regions distinctly separate upon the peduncle, the cymules widely spaced, spiral; bracts of pistillate and staminate cymules con- cave, obovate, truncate, mucronulate; the glands patelliform, somewhat auriculate. Sepals of pistillate flowers 3, about 2 mm. long; ovary pedicellate, exserted beyond the bract and sepals, 3-carpellate; styles about 3 mm. long. Staminate cymules 10- to 12-flowered, the flowers subsessile, about 1.5-2.0 mm. long; calyx shallowly 2- lobed, the lobes serrulate; pollen ellipsoid, triangular in cross-section, with 1 lateral pore, the exine finely to coarsely punctate. Fruit deeply 3-lobed; lobes of the gyno- base about 3 mm. long; seeds ovoid to slightly ellipsoid, about 4 mm. long, 3.5 mm. wide, tuberculate-striate, gray-brown, the caruncle small. ZIL: RIO DE JANEIRO: Forterepe du Pic de Sta. Crus prés de Rio de Janeiro, Gc 6126 (C, G, S) ; in montosis saxosis prope Rio de Janeiro, Riedel 185 (C, F, G ОТ.ОТҮРЕ, GH, NY). 12. STILLINGIA sAXATILIS Muell. Arg. in Mart. Fl. Bras. 11?:539. 1874. (Т.: Riedel 1172!). Gymnostillingia loranthacea Muell. Arg. in Mart. Fl. Bras. 112:541. 1874. (T.: Blanchet ! Stillingia loranthacea (Muell. Arg.) Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. 85. 1912. Shrubs to 1.2 m. tall; stems succulent, infrequently branched, the branches opposite, thickened, leaf scars prominent, without lenticels, vernicose, red-brown to frequently blackened. Leaves alternate, crowded, sessile to petiolate, somewhat succulent, elliptic to spathulate or oblanceolate, 3—6 cm. long, 0.8—2.3 cm. broad, apex acute to acuminate, base narrowly acute with 2 suborbicular inset glands, irregularly serrulate to serrate, with frequent irregularly spaced scutelliform glands toward the tip, the midrib prominent, secondary venation immersed in the fleshy ??"Catinga" refers to a region of open scrub forest, with mostly deciduous species and very low rainfall. 1951] ROGERS—REVISION OF STILLINGIA 233 tissue; sessile or the petiole 0.1-0.8 cm. long. Inflorescence 2.5—6.4 cm. long, slightly flexuose, pedunculate below the lowest pistillate cymule, the upper stam- inate and lower pistillate regions distinctly separate upon the peduncle, the cymules widely spaced, spiral; bracts of pistillate and staminate cymules concave, obovate, truncate, mucronulate, the glands flattened, auriculate. Sepals of pistillate flower united into a truncate annulus, or greatly reduced; ovary subsessile, 3-carpellate; styles about 3 mm. long. Staminate cymules 5- to 7- flowered, the flowers sub- sessile, about 1.5 mm. long; calyx very shallowly 2-lobed, the lobes unevenly ser- rulate; pollen ellipsoid, with one lateral pore, the exine granular. Fruit deeply 3-lobed; lobes of the gynobase 3-4 mm. long; seeds ellipsoid, about 3.5 mm. long, 2.5 mm. wide, smooth, brown, the caruncle small. In rocky, hilly areas. Flowers and fruits from October through January. ZIL: BAHIA: Blanchet 271 (С, hg F, MO); Serra das Ew 1700 m., Luet- wa 172 (NY). MiNas GERAIS: Serra da Lapa, Riedel 11ў2 (С, photos F, MO Municipio Diamantina, Bom Successo, pe 9675 (F). WITHOUT LOCALITY: Cabo 19952 (C). The only known specimen of S. lorantbacea, Blancbet 271, is a mere fragment on which no pistillate flowers are present. However, the great similarity of vege- tative and floral characters, including the morphology of the pollen grain, leads me to the conclusion that the Blanchet specimen is but a variant of the species S. saxatilis. Series 3. SYLVATICAE (Pax & Hoffm.) D. J. Rogers, stat. nov. $ Sylvaticae Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. V:191. 1912. Subshrubs arising from a woody underground base; stems herbaceous or sub- herbaceous, green to reddish-brown, without lenticels. KEY TO THE SPECIES AND SUBSPECIES A. Glands of the subtending bracts of the cymules elongate, tubiform or urceolate; leaf margins with a broad calli Paraguay, Argentina and Bolivia. B. Leaves generally narrowly elliptic to linear, without glands at base of the blade; pistillate Vic broadly rhombic; caruncle hippocrepi- form, attached below the BB. Leaves generally pardo to “rondi ipus, with 2 scutelliform or cyathiform glands at of t de wes ган elliptic to obova er micropyle 14. S. scutellifera AA. Glands of the subtending bracts of the piae flattened, UN p iform or ста only 13. S. salpingadenia C. Inflorescence peduncle bo the lowest ин cymule; uis usually with 2 or e cyathiform glands of blade, rowly callose, Mice pue: C 5” and М ехїсо. 0: те sessile, usually . long or less; staminate cymules 10- 12-flowered, 15 Се аса Brazil 15. S. Dusenii DD. is petiolate, usually 3 cm. long or longer; staminate cymules 3- to 5-flowered, the idea. Medie: Mexic o and са Central Americ 16. S. zelayensis GG. poe Т. IM the lowest pistillate cymule; leaves without glan 5? of the blade; United States and extreme кру фар лий Mexi [Vor. 38 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN E. »- linear; seeds truncate at base; secondary roots fibri- form; southern Oklahoma, — central Texas to north- easter exico, in limestone soils...... 17. S. texana EE. Leaves elliptic, or spathulate to obovate (except ssp. tenuis, wit ry narrowly elliptic to же: leaves); seeds rounded at the base; — ary roots fusiform; southern Virginia to Florida to Texas New haces mostly on sands............... 18. S. sylvatica F. Leaves eas. hin or obovate; southern Virginia to Florida to Texas and New Mexico, exclusive of extreme southeastern Florida, in sands......... .. 18a. S. sylvatica ssp. sylvatica FF. Leaves very narrowly elliptic to linear; extreme south- eastern Florida, on limestone оцсгорѕ. 18b. S. sylvatica ssp. Zenuis 13. STILLINGIA SALPINGADENIA (Muell. Arg.) Huber, in Bull. Herb. Boiss. Il, 6:452. 1906. Sapium salpingadenium Muell. Arg. in Linnaea 32:121. 1863. (T oo 918!). Be salpingadenia (Muell. Arg.) Muell. Arg. in DC. Prodr. 157; 12 1866. Sapium см matospermum sensu v E in Bull. Herb. Boiss. II, 1:399. n non Muell. rg. in Linnaea 34:217. $. Sapium Parir preas Hemsl. in Hook. Icon. Pl. 28:/. 2679. 1901. (T.: Hagenbeck s, п.). — salpingadenium Muell. Arg. var. р» Chodat & Hassler, in Bull. Herb. Boiss. : 05. Hassler 6346! к n Sais Arg.) Huber, sp. anadena Pax & Hoffm. in Engl. Pflanzenr. a . V:190. 1912. (No type cit Stillingia 5. 55р. pie Ж var. д, NRI (Hemsl.) Pax & Hoffm. loc. cit. Stillingia salpingadenia ssp. anadena var. e. salicina (Chodat & Hassler) Pax & Hoffm. loc cit. 1912. Subshrubs 3-10 dm. tall; stems subherbaceous, woody toward the base, in- frequently branched, the branches alternate to approximate, leaf scars on older parts prominent, sparsely lenticellate to elenticellate, reddish- to gray-brown. Leaves alternate to approximate, subsessile to petiolate, membranaceous, narrowly elliptic to slightly lanceolate, 4-7 cm. long, 0.8-2.5 cm. broad, apex acuminate, mucronulate, base acute, without cyathiform or scutelliform glands, the margin serrulate, broadly callose, the midrib prominent, secondary venation obvious, not prominent; petiole 0.1—0.4 cm. long. Inflorescence 3.5-8.5 cm. long, sessile or shortly pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, net distinctly separate upon the peduncle, the pistil- late cymules widely spaced, the staminate crowded; bracts of pistillate and stam- inate cymules broadly rhombic, mucronulate, the glands tubiform to urceolate, about 2.5 mm. long. Sepals of pistillate flower 3, elliptic, mucronulate; ovary sessile, 3-carpellate; styles about 4-5 mm. long. Staminate cymules 7- to 9- flowered, the flowers subsessile, about 2 mm. long; calyx shallowly 2-lobed, the lobes entire to slightly serrulate; pollen ovoid to ellipsoid, with 3 pores, the exine unevenly striate to granular. Fruit not seen. Lobes of the gynobase 3—4 mm. long; seeds ellipsoid, 6 mm. long, 4.5 mm. wide, slightly rugulose, light gray, the caruncle small, hippocrepiform, attached below the micropyle. 1951] ROGERS—REVISION OF STILLINGIA 235 In rocky areas, hill regions, high plains above flowing rivers, generally below 1000 m. alt. Flowers from September through December and fruits from October through February. n IVIA: SANTA CRUZ: Chiquitos, D’Orbigny 018 (F, P, photos E US). RAGUAY: Cordillera de Altos, Fiebrig 7 "a F, G, GH, LIL), same locality, m 2110 (G) (fasciated) , Hassler 3304 (A, F, G); Fort Lopez, Hassler 888 (С); Cerros de Tobaty, Hassler 6346 (A, F, G, К); in regione cursus superioris fluminis Apa, Hassler 7782 (G) 14. Ѕтпллмсіл scutellifera D. J. Rogers, nom. nov. Stillingia saxatilis sensu Chodat & Pup in Bull. Herb. Boiss. II, 5:676. 1905, non Muell. rg. in Mart. Fl. Bras. 117:5 йн ко уаг. ТІ Cie & Hassler, loc. cit. 1905. (T.: Hassler 4360!), non § cifolia (Torr.) R Stillingia sexo var. salicifolia f. latior Chodat & Hassler, loc. cit. 1905. (T.: dn m Stillingia saxatilis var. salicifolia f. angustior Chodat & Hassler, loc. cit. 1905 CE 4). Stillingia saxatilis var. gr P Chodat & Hassler, loc. cit. 1905. (T.: Hassler 4794!). Stillingia sal pingadenia (Muell. Arg.) Huber м. saxatilis (Chodat & Hassler) Рах & Ioffm. in Engl. Pflanzenr. IV. Fam V: 912 Stillingia salpingadenia ssp. saxatilis var. a. b Pax & Hoffm. loc. cit. 1912. (T: Hassler 6790!). SEE е ssp. saxatilis var. B. grandifolia (Chodat & Hassler) Pax & Hoffm., 9 А ‘ulhingaden ssp. saxatilis var. ү. angustior (Chodat & Hassler) Pax & Hoffm., loc Subshrubs 3-7 dm. tall; stems herbaceous, arising from a perennial, woody underground base, single or several together, unbranched, or, if branched, the branches alternate to fascicled, leafy to the base, without lenticels, red-brown. Leaves alternate to approximate, subsessile to shortly petiolate, stiffly membrana- ceous, broadly to narrowly elliptic, 3-8 cm. long, 1-3 cm. broad, apex acute, mucronulate, base acute, with 2 scutelliform or cyathiform glands, the margin serrulate, broadly callose, the midrib and secondary venation prominent; petiole 0.1-0.7 cm. long. Inflorescence 2.8-11.8 cm. long, sessile to shortly pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules relatively widely spaced, the staminate crowded, bracts of pistillate and staminate cymules elliptic to broadly rhombic, serrulate, sharply mucronulate, the glands infundibuliform, 2-3 mm. long. Sepals of pistillate flower 3, rhombic, serrulate, bluntly mucronulate; ovary subsessile, 3-carpellate; styles about 3—5 mm. long. Staminate cymules 3- to 5-flowered, the flowers shortly pedicellate, about 3 mm. long; calyx deeply 2-lobed, the lobes serrulate; pollen spheroid with 3 pores, the exine granular. Fruit not seen. Lobes of the gynobase 4—5 mm. long; seeds ovoid to slightly ellipsoid, about 5 mm. long, 5 mm. wide, slightly rugulose to smooth, dark gray, the caruncle large, waxy, rhombic, surrounding the micropyle. Grassy campos, near margins of forest, high campos, in river regions at altitudes usually below 500 m., but occasionally to 1000 m. Flowers from October to February and fruits from November through March. [Vor. 38 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN ARGENTINA: MISIONEs: Dept. Cainqua, Campo Grande, Pierotti 5365 (LIL), 5384 (LIL); Dept. Candelaria, ба Ana, Schwarz 3520 (LIL) ; Dept. San Ignacio, El — Schwerz 1354 (LIL); Zob. Roca, Schwarz 5105 (LIL); Santa ere Schwarz (LIL); Maria Antonia, Schwarz 8301 LM Posadas, Ekman 506 (S, US). PARAGUAY: Pirayu, Balansa 1708 (К); С Spy prés de Villa-Rica, dans les prairies, Balansa 1711 (G); Cordillera de Altos, Fiebrig 99a (A); Dept. Piribebuy, Colonia Pedro P. Caballero, cdi `Ао$6? (LIL); Villa Rica, j sedi 3985 (A, C, F, LIL, MO, 5, US); Sierra de Maracayu, in regione fluminis Tapiraguay, Hassler 4360 (F, GH, S); in regione fluminis Capibary, Hassler 4446 (G, S) ; in regione fluminis Corrientes, Hassler 4494 (G); ec in regione vicine "Igatimi", Hassler 4704 (С), 5612 (С); Cordillera Centralis, in regione ursus superioris fluminis Y-aca, Hassler 6790 (С); in campo montano pro. Cholola (Chulolo), Hassler 6790 (С); auf nasse Stellen, Arecipe, Rio Apa, Anisits 2435 ur There is little doubt in my mind that S. scufellifera is sufficiently distinct from S. salpingadenia to be considered as a separate species. It is apparent, however, that rather free hybridization is taking place between the two species, with the result of confusion when an effort is made to key some of the individuals of either species. 15. SriLLINGIA. DusENm Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, XVII:202. 1924. (T.: Dusén 10440!). Subshrubs 1-3 dm. tall; stems herbaceous, arising from a perennial, woody, underground base, single or several together, unbranched, or, if branched, the branches alternate to fascicled, leafy to the base, without lenticels, red-brown. Leaves alternate to approximate, widely to closely set, sessile, membranaceous, ovate to elliptic, 1.0-2.5 cm. long, 0.5-1.3 cm. broad, apex acute to rounded, base obtuse to broadly acute, with 2 cyathiform to patelliform glands, the margin serru- late, narrowly callose, the midrib prominent, the secondary venation obscure. In- florescence 8-12 cm. long, pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate regions spiral, distinctly separate upon the peduncle, the cymules relatively widely spaced; bracts of the pistillate cymules foliaceous, linear, about 5 mm. long, acute, the base auriculate, serrulate, the stam- inate squamaceous, ovate to broadly elliptic, serrulate, mucronulate, the glands circular, flattened. Sepals of pistillate flower 3, broadly triangular, serrulate, with- out mucro; ovary sessile, 3-carpellate; styles unknown. Staminate cymules 10- to 12-flowered, the flowers pedicellate, the pedicels about 2 mm. long; calyx shallowly 2-lobed, the lobes serrulate; pollen irregularly spheroid, with 3 pores, the exine е to coarsely punctate. Mature fruit and seed not seen. RAZIL: PARANA: Jaguariahyva, in campo, Oct. 25, Dusén 10440 (MO, S); Ma ( vd Jan. 25, Dusén 0143 (5); in campo limpo, Oct. 11, Dusén 13179 (G, GH, S); campo, 740 m., Jonsson 204a (А). 16. STILLINGIA ZELAYENSIS (HBK.) Muell. Arg. in Linnaea 32:87. 1863. Sapium zelayense HBK. Nov. Gen. et Spec. 2:51. 1817. (T.: Humboldt 8 Bonpland s. n.), Erect to spreading subshrubs 5—15 dm. tall; stems subherbaceous, arising from a perennial, woody, underground base, terete, older parts hollow, woody, the sec- ondary branches opposite or fascicled, solid or with a small pith, without lenticels, 1951] ROGERS—REVISION OF STILLINGIA ye fà green- to reddish-brown. Leaves petiolate, membranaceous, elliptic to oblanceolate to spathulate, 2.8—12.5 cm. long, 1.2—4.0 cm. wide, apex acute to acuminate, base acute, usually with 2 cyathiform glands, the margin serrulate, narrowly callose, the midrib and secondary venation prominent; petiole 1-9 mm. long. Inflorescence 5.0—16.0 cm. long, pedunculate and somewhat thickened below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, distinctly separate upon the peduncle, the pistillate cymules distant, the staminate crowded; bracts of the pistillate and staminate cymules broadly elliptic, mucronulate, serrulate, the glands patelliform, sessile. Sepals of pistillate flower 3, narrowly elliptic, about 3 mm. long, mucronulate, serrulate; ovary sessile, 3-carpellate; styles about 3 mm. long. Staminate cymules 3- to 5-flowered, the flowers sessile, about 3 mm. long; calyx deeply 2-lobed, the lobes serrulate; pollen spheroid, with 3 pores, the exine reticulate. Fruits shallowly 3-lobed, 0.6-1.8 cm. wide, the lobes of the gynobase 3-9 mm. long; seeds nearly spheroid, about 5.5-6.5 mm. long, 5.5-7.0 mm. wide, testa smooth, shiny, flattened at the micropylar end, the basal end rounded to slightly flattened, the caruncle small. Pine woods and barrancas, rocky hills, shady open slopes on edges of cultivated areas, from about 1200 to 2800 m. altitude. Flowering mostly from about the last of March through the last of August, but occasionally in January and in October, and fruiting from June through October and occasionally in January. Mexico: cHiAPAs: Hacienda Monserrate, Purpus 9174 (F, GH, MO, NY, UC, US), 0323 (UC); without locality, Purpus s.n. (US); pine and oak forest, Fenix (Chiapas?), Purpus 10334 (NY). DISTRITO FEDERAL: east of Transfiguración, Russell 9 Souviron 205 (US). JALIsco: Santa Cruz, M. E. Jones 22 (МО, US); Rio Blanco, E. Palmer 73 (NY, US); near Guadalajara, Pringle s. n. (C, F, LIL). MEXICO: Dist. Temascáltepec, Penon, Hinton 4400 (GH); Dist. Temascáltepec, Temascáltepec, Hinton 1127 (F, СН, , US). MoreLos: rocky hill near El Mirador, road from Cuernavaca to Mexico, Williams 3814 (GH); Valle del Tepeite, Lyonnet 088 (US); barranco near Cuernavaca, Pringle 7244 (СН); pine woods about Cuernavaca, Pringle 6886 (BR, F, GH, MO, NY, UC, US). micHoacan: vic. of Morelia, Arsène 6679 (MO, US); vic. of Morelia, Arséne 5236 (GH, MO, US); Tancitaro region, above Acahuaato, Munic. Apatzingan, Leaven- worth & Hoogstraal 1821 (F, СН); hills near Tepitenga, between Tolura and Morelia, Gregg 727 (MO). PurnLA: Cerro Tepoxuchil, prés Puebla, Nicolas s.n. (LIL, NY); vic. of Puebla, Cerro Tepoxuchil, Arsène 1400 (GH, MO, US); Rancho Losada, vic. of Puebla, Arsène ё Nicolas 282 (F). saN Luis Potosi: 22° n. lat., alt. 6000-8000 ft., Parry 9 Palmer 823 (GH, MO, US). TLAXCALA: Huamantla, alt. 8500 ft., Balls B.5640 (A). STATE AND LOCALITY UNKNOWN: Pavon s.n. (G, photos in F, 5 UATEMALA: GUATEMALA: without locality, Aguilar 93 (F). QUICHE: without locality, Aguilar 1314 (F). sANTA rosa: Zamorora, Heyde © Lux 4579 (US); without locality, Deam 6074 (F, GH, MICH, MO, NY, US). PANAMA: CHIRIQUÍ: Boquete Distr., savannah, alt. 4000 ft., Davidson 750 (A, P, MO, US). The type specimen of Stillingia zelayensis, collected by Humboldt & Bonpland, was labelled for locality “prope Zelaya”. The actual locality is Celaya, Guanajuato, Mexico. [Vor. 38 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN Р, - 4 ——— Мег Fig. 13. Stillingia zelayensis. Fig. 14. Stillingia texana. 17. STILLINGIA TEXANA I. M. Johnston, in Contr. Gray Herb. n. s. 68:91. 1923. — unii var. linearifolium Torr. in Emory, Rept. U. S. & Mex. Bound. Surv. 2:201. 1859. ( Stillingia sylvatica ò, linearifolia (Torr.) Muell. Arg. in DC. Prodr. 157:1158. 186 — pic (Muell Arg.) "pne ex S. Wats. in Proc. Am. Acad. p 154. о specimen cited, not as to basin лімін linearifolia п) Small. Fi. 5. E. "United States, 704. 1903; not S. Wats. in 4:297 res 5 var. ed Waterfall, in Rhodora 50:95. Stillingia texana var. latifolia Waterfall, loc. cit. 1948. (Ta pow 6523!). Loose erect subshrubs, 2.5—6.0 dm. tall; roots fibriform; stems herbaceous, arising from an enlarged woody root crown, terete or slightly striate, unbranched, or, if branched, the branches fascicled near the apex of the main axis, without lenticels, light green or brown. Leaves alternate, sessile, membranaceous, linear to slightly lanceolate, 1-7 cm. long, 0.3-1.0 cm. wide, apex acute, base narrowly obtuse, without cyathiform glands, serrulate to crenulate, the midrib prominent, slightly decurrent. Inflorescence 3.0-8.0 cm. long, sessile below the lowest pistil- late cymule, the upper staminate and lower pistillate cymules spiral, not distinctly separated on the peduncle; bracts of pistillate and staminate cymules narrowly elliptic, mucronulate, sharply denticulate, the patelliform glands sessile. Sepals of pistillate flower 3, elliptic, serrulate; ovary sessile, 3-carpellate; styles about 3 mm. long. Staminate cymules 3- to 5-flowered, the flowers subsessile, about 2.5 mm. long; calyx deeply 2-lobed, the lobes slightly serrulate; pollen mostly spheroid, with 3 pores, the exine granular to reticulate. Fruits shallowly 3-lobed, 4—6 mm. wide, the lobes of the gynobase 2—3 mm. long; seeds mostly ellipsoid, 3—5 mm. long, 2.5-4.0 mm. wide, testa slightly rugulose, with a very slight depression at the micropylar end, the base truncate, the caruncle small. On limestone soils, mostly in open prairie lands, occasionally on loamy black soils, in rolling country. Flowers from about the last of April through the middle of July, and fruits from the first of June through September. 1951] ROGERS—REVISION OF STILLINGIA 239 UNITED STATES: OKLAHOMA: Murray Со. rEXAs: Bell, Bexar, Bosque, Burnet, Comal, Conche, Erath, Fort Bend, Gillespie, Hays, Kendal, Kerr, Lampasas, Mills, Nolan; Parker, Real, Schackelford, Tarrant, Taylor, Travis, Washington, Williamson, Wilson, and Valverde counties. MEXICO: COAHUILA: Munic. de Muzquiz, near Puerto Santa Ana, Wynd & Mueller 225 (GH, MO, NY, US); Hacienda zo Rafael, along the Sabinas River, Wynd 705 (M NY, US); Muzquiz, Marsb бо (F, TEX); Muzquiz, Yerda Spring, Marsb 291 (TEX). One specimen was collected by C. Wrigbt in New Mexico, without locality. Although Watson gave a description for S. angustifolia Engelm., he placed S. sylvatica var. angustifolia in synonymy. It scems best to follow Johnston’s interpretation (in Contr. Gray Herb. n. s. 68:91. 1923), that Watson was making a new combination and the name must apply to the Florida plant (Mueller's type) not, as was done for many years, to the Texan plant. Since no other name was available, Johnston renamed the species, basing it upon var. linearifolia Torr. 18. STILLINGIA sYLvATICA Garden, ex L. Mant. 19. 1767. (T.: Garden s. n.). Erect or semi-erect subshrubs 2-12 dm. tall; roots fusiform; stems herbaceous to subherbaccous, arising singly or several together from a woody rhizome, terete, unbranched, or, if branched, the secondary branches dichotomous, approximate or fascicled near the apex of the main axis, without lenticels, bark of older parts frequently with transverse and longitudinal cracks, red to reddish-brown or mottled gray and reddish-brown. Leaves alternate, sessile or petiolate, membranaccous, green or sometimes bright red, narrowly elliptic, elliptic, obovate, spathulate or oblanceolate, 2—12 cm. long, 0.3—4.8 cm. wide, apex acute, rounded or emarginate, base narrowly acute to narrowly obtuse, without cyathiform glands, serrulate to crenulate, the midrib prominent, secondary venation obscure; petiole when present 0.1-0.8 cm. long. Inflorescence 2.5-13.0 cm. long, slender or stout, sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, not distinctly separate upon the peduncle; bracts of the pistillate cymule caudate- acuminate to broadly elliptic, serrulate, mucronulate, the staminate broadly elliptic, mucronulate, the glands patelliform or cyathiform, sessile; peduncle and bracts red or yellow-green. Sepals of pistillate flower 3, elliptic, slightly cuspidate, serrulate, 2-3 mm. long; ovary sessile, 3-carpellate; styles about 2—5 mm. long. Staminate cymules 5- to 13-flowered, the flowers subsessile, about 1.5—2 mm. long; calyx shallowly 2-lobed, the lobes entire to slightly serrulate; pollen ellipsoid to spheroid, with 3 pores, the exine reticulate to granular. Fruit shallowly 3-lobed, 0.6—1.2 cm. wide, the lobes of the gynobase 3-6 mm. long; seed ellipsoid 4-8 mm. long, 3—7 mm. wide, testa rugose, the micropylar end flattened, the base rounded, mottled dark gray, the caruncle large, waxy. 18a. STILLINGIA SYLVATICA Garden ssp. SYLVATICA. Stillingia sylvatica var. salicifolia Ps i Ann. Lyc. N. Y. 2:245. 1826. (T,: James s, n.!). Stillingia salicifolia 1. ) Raf. . Jour. 1:146. 1832. Stillingia lanceolata Nut THE Phil. Soc. n. s. 5:176. 1837, (Т,: Pitcher s. E Stillingia sylvatica a. Pr M e in е Ргойг. 152:1158. 1866. (T.: Bose s,n,). Stillingia sylvatica B. genuina Muell. Arg. loc. cit. 1866. Stillingia sylvatica у. angustifolia Т Arg. Те cit. 1866. (T.: Mitchell s. т,), [Vor. 38 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 15. Stillingia sylvatica. Stillingia керісін pueda ge Engelm. ex S. Wats. in Proc. Am. Acad. 18:154, , as to basinym, no о specimen cite жд» қышы (Muell. ripen течь Fl. S. E. United States, 704. 1903. Stillingia salicifolia Small, loc. cit. 3 (no type cited), non S, salicifolia (Torr.) Raf. Stillingia Smallii Wooton & St ur a in Contr. U. S. Nat. Herb. 19:405. 1915. (Based on the preceding). Leaves elliptic, spathulate or obovate, 3.2-11,5 cm. long, 0.3—4.8 cm. wide, apex acute, rounded or emarginate. Bracts of the pistillate cymule elliptic. Coastal plain, except extreme southeastern Florida, well-drained sands, in open pine woods and sometimes on limestone, occasionally on roadsides and railroad banks. Flowers from March through June and fruits from April through Sep- tember. UNITED STATES: VIRGINIA: Isle of Wight, Nansemond, Sussex, and Southampton counties, NORTH CAROLINA: Buncombe, Harnett, Iredell, Мен, Richmond, and Scotland counties. SOUTH CAROLINA: Aiken, Beaufort, Berkeley, Charleston, Clicendon, Georgetown, Hampton, Horry, and Lancaster counties. GEORGIA: Bull k, Columbia, Gwinnett, Haber- sham, Jasper, con, Meriwether, Richmond, yne counties. FLORIDA: Alachua, Baker, Bradford, Brevard, Broward, Charlotte, Citrus, Clay, Columbia, Dade " Dixie, Duval, Flagler, Gadsden, с Hardee, ndry, Hernando, Highlands, Hills- borough, Jefferson, Lafayette, Lake, ‚ Leon, Levy, Liberty, Manatee, Marion, Monroe Nassau, Orange, Palm Beach, Pasco, 512 Polk, Putnam, St. John's, Sumter, Posi Walton, Wakulla, Union, and Volusia counties. ALABAMA: Dale, Henry, Lee, and Mobile counties. LOUISIANA: Natchitoches, and Rapides parishes. М1551551ррІ: Holmes Co. TEXAS: Anderson, Angelina, Aransas, Austin, Bailey, Bastrop, Bexar, Brazos, Caldwell, Childress, Dallas, Denton, DeWitt, Dimmit, Erath, Fayette, Fort Bend, Goliad, Gonzales, Kenedy, Lamb, Lee, Leon, Lubbock, Medina a, Montgomery, Nacogdoches, Newton, Nueces, Parker, Tarrant, Terry, Travis, Upshur, Walker, Waller, Washington, Wichita, Wilson, 1951] ROGERS—REVISION OF STILLINGIA 241 and Wood counties. ARKANsAs: Crawford, Jefferson, and Miller counties. KANSAS: wley, Harper, Montgomery, Morton, Sedgwick, and Stafford counties. OKLAHOMA: Beaver, Beckham, Blaine, Caddo, Comanche, Cleveland, Ellis, Greer, Kingfisher, Kiowa, Logan, McClain, McCurtain, Muskogee, Oklahoma, Osage, Payne, and Roger Mills counties. NEW MEXICO: Chaves, Harding, and Roosevelt counties. The taxonomic confusion concerning S. sylvatica has resulted in part from the lack of knowledge of habitat and in part from a failure to recognize that some of the variability possibly is due to hybridization. The narrow-leaved specimens from Arkansas, Oklahoma, Texas and New Mexico, to which the epithets S. sylvatica var. salicifolia or S. salicifolia previously have been applied, is thought to be due to the influence of hybridization with S. texana, and as a result no nomenclatural status has been given them. The typical subspecies is largely confined to sandy areas within the coastal plain of the southern United States. It occurs only on limestone as a result of its putative hybridization with S. texana, and its occurrence on limestone soils is rather infrequent. Stillingia sylvatica ssp. tenuis, on the other hand, has become a stable entity on the limestone outcrops of extreme southeastern Florida. Hybridization of S. sylvatica ssp. sylvatica with S. aquatica has been discussed under the heading of Subgeneric Categories. The following specimens of S. sylvatica in the herbarium of the Missouri Botanical Garden demonstrate the influence of introgressive hybridization with S. aquatica. UNITED STATES: FLORIDA: OSCEOLA COUNTY: 5 mi. s.w. of Kissimee, dry sandy pineland, Rogers 4. HIGHLANDS COUNTY: п. of Highland, Hammock State Park, dry sands, Rogers 5; V4 mi. e. of Sebring, sandy flats, cleared field, Rogers 6; 2-3 mi. s.e. of Sebring, near shallow, marshy pond, Rogers 7. PALM BEACH COUNTY: 15 mi. s. w. of Lake Worth, sands near cypress swamp, Rogers 14-1, 14-3. MARTIN COUNTY: 1 mi. s. of Stuart, 100—200 yards from cypress pond, on a low sandy ridge, Rogers 15-1, 15-2, 15-3, 15-5, 15-7, 15-10, 15-11. (See also specimen citations of S. aquatica showing introgression with S. sylvatica, p. 230.) The medicinal values of S. sylvatica have been explored and reported in several journals. An alcohol or water extract of the root was, and occasionally still is, used by physicians and laymen in South Carolina particularly, in the treatment of syphilis, in cutaneous diseases, chronic hepatic infections and in the composition of diet drinks.?. As late as 194424, this plant was mentioned as of minor drug importance, particularly as a sialagogue and expectorant. Vernacular names: Queen's Delight, Queen's Root, Silverleaf, Nettle Potato, Yaw-root, Marcory, and Cockup-hat. 18b. SrIL.LINGIA sYLvATICA Garden, ех L. ssp. tenuis (Small) D. J. Rogers, stat. nov. Stillingia tenuis Small in Bull. М. Y. Bot. Gard. 3:429. 1905. (T.: Small 9 Wilson 1580!). Leaves very narrowly elliptic to linear, 2-10 cm. long, 0.3-1.0 cm. wide, apex acute. Bracts of the pistillate cymule caudate-acuminate. ?3Millspaugh, C. F. Medicinal Plants 1:151. 1892. 24 Allport, Noel L. The Chemistry and Pharmacy of Vegetable Drugs, p. 235, Tab. XVII. 1944. [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN 242 Stillingia sylvatica ssp. tenuis. Fig. 16. 1951] ROGERS—REVISION OF STILLINGIA 243 Confined to the extreme southeastern coast of Florida, usually growing only where limestone outcrops occur at the surface, with none or only a very slight amount of sandy soil mantle. Flowers from January to May or June, and fruits from March through June. UNITED STATES: FLORIDA: Dade Co. The subspecies occupies a distinct geographical and ecological region from its closest relative $. sylvatica ssp. sylvatica, but it has not sufficiently diverged to make it a separate species. This is perhaps due to the fact cither that there is little possibility of further migration, it being limited to a very narrow strip on the southeast coast of Florida, or that insufficient time has elapsed since its formation, This part of Florida is one of the most recently exposed land masses on this continent, Subgen. П. GvMNosrILLINGIA (Muell. Arg.) D. J. Rogers, stat. nov. Gymnostillingia Muell. Arg. in Linnaea 32:89. 1863, as genus. Series 4. ACUTIFOLIAE D. J. Rogers, ser. nov. $ Gymnostil'ingia (Muell. Arg.) Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. V:193, 1912; Frutices aut arbores parvi; sepala floris pistillati minuta vel inchoata; flores staminales in cymula solitaria, polline sphaeroideo vel ellipsoideo; semina ecaruncu- lata. Mexico australi et Guatemalae. 19. STILLINGIA ACUTIFOLIA (Benth.) Benth. ex Hemsl. Biol. Centr.-Amer. Bot. 33135: 1697, Sapium acutifolium Benth. Pl. Hartweg. 90. 1842. (T.: Hartweg ч Gymnostillingia acutifolia (Benth.) Muell. Arg. in Linnaea 32:89 о тастапіра Muell. Arg. loc. cit. 1863. (Т.: Povon 5.” Stillingia macrantba (Muell. Arg.) Benth. ex Hemsl. Biol. Centr.-Amer. Bot. 35. 1883, Stillingia propria Brandg. in Univ. Cal. Publ. Bot. 6:185. 1915. (T.: i езй} 21 Compact shrubs or small trees 1-7 m. tall; trunk to 12.5 cm. diameter below the lowest branches, the branches alternate to fascicled, terete below, angled above, the older branches sparsely lenticellate, the sap milky. Leaves alternate, petiolate, thinly membranaceous, dark green above, paler beneath, ovate, lanceolate, or elliptic, 2.8-15.0 cm. long, 1.0—4.3 cm. broad, acuminate, base acute, without cyathiform glands, the midrib and secondary venation prominent; petiole 2—5 mm. long. Inflorescence 2.5-4.2 cm. long, sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, not distinctly separate upon the peduncle; bracts of the pistillate and staminate cymules elliptic, cuspidate, about 1.5 mm. long, the glands patelliform, sessile. Sepals of the pis- tillate flower 3, minute or rudimentary; ovary sessile, 3-carpellate; styles about 2 mm. long. Staminate flowers solitary, very shortly pedicellate, about 2 mm. long, the calyx lobes entire; pollen ellipsoid to spheroid, triangular in cross-section with [Vor. 38 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3 pores, the exine finely punctate. Fruit 3-4 mm. wide, deeply 3-lobed, the lobes of the gynobase 1.5—2.0 mm. long; seeds about 3.5 mm. long, 2.5—3.0 mm. wide, with a slight depression on each side of the raphe at micropylar end, the base rounded, the testa smooth, the caruncle absent. Mountainous regions of Chiapas and Guatemala in damp wooded regions, open banks, barrancas in pine forests, white sand slopes, at altitudes from 1400 to 3000 m. Flowers from late June to the first of September, and fruits from the first of Т. to the last of December. ХІСО: CHIAPAS: Cerro del Boqueron, Purpus 7343 (Е, СН, MO, NY); Siltepec, ама; 168; (А, МІСН, U ALA: CHIMALTENANGO: Volcan Acatenango, Kellerman 6576 (F); сеи Skutch 20 (US), 554 (A); west of Patzun, Williams & Molina 11844 (Е, MO); А J. R. Johnston 051 (F, NY); near Rio Pixcayo, between Сеново and San an 1 lotepeque, S/andley 64330 (F); Barranco de La Sierra, southeast of Patzun, Standley 61510 (A, Е, NY); near Parramos, Standley 50870 (А), 50880 (F); region of Los Positos, above Las Calderas, region of Las Calderas, ендің 57803 (А, Е), 60012 (Е), 80138 (А). GUATEMALA: Ruano 1256 (Е); Aguilar 2 O (F). HUEHUETENANGO о Santos, Melbus t$ Good man 3617 (F); San Juan rei Skutch 1165 (A, F, US). JALAPA: between Jalapa and Montana Miramundo, — 32695 (F). QUEZALTENANGO: Aguas Calientes, Seler 2741 (A, GH, NY, US); Cumbre de Tuilacan, s. w. of San Martin Chile Verde, Standley 67787 (F); region of a s.e. of San Martin Chile Verde, Standley 83827 (F). sACATEPEQUEZ: along Rio Guacalate, on е between Antigua and Chimal- tenango, Standley roro (F). Да MARCOs: mountains along the road between San Marcos and Serchil, Standley 85327 (F); El Boqueron, n near I of Dept. Quezaltenango, Standley 66284 (F); El Boqueron, in the ntains at the mit of the road between зап Antonio Sacatepequez and Palatina, Standley 85280 (P); parria above Rio Tacana, near San T PH 66165 (F). SANTA ROSA: Santa Rosa, Heyde & Lux 3473 (F, GH, NY, US). LOLA: о Г. О. Williams аА (Е); Los Encuentros, Seler a (GH, Us). . TOTONICAPAN: along road between San Francisco El Alto and Momostenango, Standley 84033 (F). AZCATEPEQUE: Bernoulli & Cairo 2499 (K, S). Dept. unknown, Hacienda de Argueta, Hartweg 614 (F, К). According to information from a labeled specimen, the acrid milky sap of S. acutifolia is used as a caustic in the treatment of sores and boils. Fig. 17. Stillingia acutifolia. Fig. 18. Sfillingia Treculiana. 1951] ROGERS—REVISION OF STILLINGIA 245 Series 5. TRECULIANAE D. J. Rogers, ser. nov. $ Leptostachyae Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. V:194. 1912. $ Gymnostillingia Wats. in Pax & Hoffm., loc. cit. 1912, in synonymy. Herbae perennes; flores pistillati sine sepalis (S. Treculiana excepta sepalis minutis et fagaceis) ; flores staminales in cymula solitaria, polline sphaeroideo seg- mentis 3 foraminibus 3; semina aut ecarunculata aut carunculis minutis et fugaceis. KEY TO IHE SPECIES A. оа Ва дел of pistillate кий nearly sessile. B. Leaves obovate-spathulate, rounded or broad А acute, dr ра in- DINE pei minute, fugacious; kin bout 1 mm. long...... 20. S. Treculiana BB. =? linear, acuminate or acute, sparsely setulose- xr ace pn pals absent; Min about 4 mm. long 1. S. paucidentata AA. eus cle absent; glands of ретке bracts long-pedicellate С. Leaves clip на арех ас ate, finely че dentate, conspicuously 3-costate; stems striate 22. S. spinulosa CC. Leaves Gast ы acute, entire or sparsely denticulate, venation not prominent; stems Тегесе........................................... а ы... 23. S. lincarifolia 20. SriLLINGIA TRECULIANA (Muell. Arg.) I. M. Johnston, in Contr. Gray Herb. 10$ O7 Oke POD. Sapium annuum var. dentatum Torr. in Emory, Rept. U. S. & Mex. Bound. Surv. 2:201, 1859. (T.: Bigelow Gymnanthes Treculiana Muell. A Arg. in Linnaea 34:216. 1865. (T.: Trecul 1458!). Sebastiania Treculiana (Muell. Arg.) Muell. Arg. in DC. Prodr. 152:1160. 1866, itc Torreyana S. Wats. in Proc. Am. Acad. 14:298. 1879. (Based on Sapium var. dentatum Torr.). Stillingia dentata (Torr.) Britt. & Rusby, in Trans. N. Y. Acad. 7:14. 1887. Spreading perennial herbs 1.0—4.5 dm. tall; elongate tap root woody; fascicled branches arising from the ground level, striate, green. Leaves alternate, sessile, membranaceous, obovate-spathulate, 1.0-3.8 cm. long, 0.6-1.5 cm. wide, apex rounded to broadly acute, base narrowly cuneate, without cyathiform glands, coarsely incised-dentate, the midvein prominent, secondary venation obscure. In- florescence 2.5—7.0 cm. long, pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, not distinctly sep- arate upon the peduncle; bracts of the pistillate and staminate cymules elliptic or cuneate, concave, with 1—3 mucros, the patelliform glands nearly sessile. Pistillate flowers solitary, sepals minute, fugacious; ovary sessile, 3-carpellate; styles about 1 mm. long. Staminate flowers solitary, subsessile, about 1.5 mm. long; calyx deeply 2-lobed, the margins involute, entire; pollen spheroid, deeply 3-segmentéd, with 3 pores, the exine coarsely punctate. Fruit deeply 3-lobed, the lobes of the gynobase 0.8—1.0 mm. long; seeds about 2.5 mm. long, 1.5-2.0 mm. wide, with a slight depression on each side of the raphe at the micropylar end, the base rounded, the testa smooth or slightly rugulose, the caruncle present. Sandy soils with limestone base, gravel, on hills, mesas, grasslands, along rail- roads and roadsides in southwestern Texas and northeastern Mexico. Flowers from the last of February to the first of June, and fruits from the last of March through the first of August. [Vor. 38 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN UNITED STATES: TEXAs: Bexar, Cameron, Concho, Crockett, Presidio, Star, Tom Green, Val Verde, and Webb counties. Mexico: coaHuiLA: Allende, Marsh 1768 (F); Muzquiz, Marsh 1116 (F); Cuatro Cienegas, Shreve 8454 (UC); Sabinas, М. E. Jones 20115 (MO, UC). NUEVO LEON: Mesas, Garcia, Pringle 2504 (F, GH, MO, NY, UC, US); 100 kilo. s. of Nuevo Laredo, on road to Monterrey, Frye & Frye 2375 (GH, MO, NY, US); near Santa Catarina, Alonis, Johnson & Barkley 15180М (Е, GH, MO, TEX); 23 mi. n. of Sabinas Hidalgo, Barkley & Webster 14407 (MO, TEX, UC); canyon 12 mi. w. of Monterrey, Barkley & Painter 14253 (GH, MO, TEX); 10 mi. w. of Monterrey, Shreve 9 Tinkbam 0572 (GH); Rancho Resendez, Lampazos, Edwards 389 (F, MO, TEX, UC); Monterrey, E. Palmer 1258 (GH, US). rAMAULIPAs: San Miguel, Pringle 2071 (F). Mexico orientali ad Rio Bravo del Norte, Trecul 1458 (MO, D). There has been some confusion as to whether S. Treculiana is an annual or perennial. This confusion arose from the original description of Sapium annuum var. dentatum made by Torrey. He could easily have called this plant an annual since the specimen used as a type was a small fragment, without roots. Mueller, however, definitely states in his description of Gymnanthes Treculiana that this is a perennial species. Examination of material from several herbaria amply cor- roborates his observation. Vernacular name: Yerba del Sapo (Mexico). 21. STILLINGIA PAUCIDENTATA S. Wats. in Proc. Am. Acad. 14:298. 1879. (Ea E. Palmer 517!). Stillingia linearifolia var. paucidentata (S. Wats.) Jepson, Fl. Cal. 21:422. 1936. Compact, erect perennial herbs 2.5-3.5 dm. tall; elongate tap root woody; the fascicled branches arising from ground level, secondary branches alternate to ap- proximate. Leaves alternate to approximate, sessile, crowded, membranaceous, linear, 2-4 cm. long, 0.1-0.3 cm. wide, acuminate or acute, not narrowed at the base, without cyathiform glands, sparsely setulose-denticulate to entire, the mid- vein prominent, decurrent, secondary venation obscure. Inflorescence 2.3-7.0 cm. long, usually sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, not distinctly separate upon the peduncle; bracts of the pistillate and staminate cymules elliptic or cuneate, mucronulate, the patelliform giands short-pedicellate (less than 1 mm.). Pistillate flowers sclitary; sepals absent; ovary sessile, 3-carpellate; styles about 4 mm. long. Staminate flowers solitary, subsessile, about 1.5 mm. long; calyx shallowly 2-lobed, the lobes slightly serrulate; pollen spheroid, shallowly 3-segmented, with 3 pores, the exine coarsely punctate. Fruit deeply 3-lobed, the lobes of the gynobase 1.5—2.0 mm. long; seeds about 3 mm. long, 2 mm. wide, testa smooth, the caruncle small. In sandy washes, plains, bare arid hills and gravel, up to 4000 ft. altitude. Flowers from the first of April through the end of May, and fruits about the first of May through the middle of June. UNITED STATES: CALIFORNIA: Kern, Los Angeles, and San Bernardino counties. 1951] ROGERS—REVISION OF STILLINGIA 247 А “ 1 ыы Fig. 19. S/illingia paucidentata. Fig. 20. S/illingia spinulosa. 22. STILLINGIA SPINULOSA Torr. in Emory, Notes Milit. Rec. 151. 1848. (T.: Emory s. n.!). Sapium annuum Torr. in Emory, Rept. U. S. & Mex. Bound. Surv. 2:201. 1859. (Based n S. spinulosa). Stillingia annua (Torr.) Muell. Arg. in DC. Prodr. 15?:1160. 1866. Compact perennial herbs 0.5-2.0 dm. tall; elongate tap root woody, the fas- cicled branches arising from ground level, the secondary branches opposite or ap- proximate. Leaves opposite or approximate, sessile, crowded, membranaceous, elliptic-spathulate, 1.5—4.0 cm. long, 0.5—1.7 cm. wide, apex acuminate, base nar- rowly cuneate, without cyathiform glands, finely callose-dentate, decurrent, promi- nently 3-costate. Inflorescence 1.2-2.0 cm. long, usually sessile below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, crowded, not distinctly separate upon the peduncle; bracts of pistillate and staminate cymules broadly elliptic, concave, cuspidate, somewhat 3-lobed, about 1 mm. long, the patelliform glands pedicellate, 1.5-2.0 mm. long. Pistillate flowers solitary, sepals absent; ovary sessile, 3-carpellate; styles about 3 mm. long. Staminate flowers solitary, subsessile, about 1.5 mm. long; calyx shallowly 2-lobed, the lobes entire; pollen ellipsoid, triangular in cross-section, with 3 pores, the exine finely to coarsely punctate. Fruit deeply 3-lobed, the lobes of the gynobase about 2 mm. long; seeds 3.0—3.5 mm. long, the testa smooth, sometimes mottled, the caruncle absent. Sandy open desert. Flowers from late December to late March, and fruits from March to June. UNITED STATES: ARIZONA: Yuma Co. CALIFORNIA: San Diego, Riverside, Imperial, and San Bernardino counties. [Vor. 38 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN MEXICO: BAJA CALIFORNIA: mesa near Cerro Breto, Cucapa Mts., MacDougal 206 (NY). sonora: Colonia Lerdo, MacDougal s.n. (NY); Buena Mesa near Col. Riv., MacDougal s. n. (NY); Pimeria Alta, Schott s. n. (NY). Torrey described Szillingia spinulosa in 1848, but changed the name іп 1859 to Sapium annuum, using the same specimens. No explanation was given for this change. 23. STILLINGIA LINEARIFOLIA S. Wats. in Proc. Am. Acad. 14:297. 1879. (T.: E. Palmer 449!). Stillingia gymnogyna Pax & Hoffm. in Engl. Pflanzenr. IV. Fam. 147. V:196. 1912. ased on S. linearifolia S. Wats.). Diffuse perennial herbs 1.5—9.0 dm. tall; elongate tap root woody, the fascicled branches arising from ground level, the secondary branches alternate, approximate, or fascicled. Leaves alternate or approximate, sessile, membranaceous, linear, 1.2—3.2 cm. long, 0.1-0.3 cm. wide, apex acute, the base narrowly acute, without cyathi- form glands, entire or sparsely denticulate, shortly decurrent, the venation not prominent. Inflorescence 1.6-7.0 cm. long, sessile or shortly pedunculate below the lowest pistillate cymule, the upper staminate and lower pistillate cymules spiral, scattered, not distinctly separate upon the peduncle; bracts of the pistillate and staminate cymules rhombic or cuneate, mucronulate, flattened, about 1 mm. long, the cyathiform glands pedicellate, about 1 mm. long. Pistillate flowers solitary; sepals absent; ovary sessile, 3-carpellate; styles about 2 mm. long. Staminate flowers solitary, subsessile, about 1 mm. long; calyx shallowly 2-lobed, the lobes entire or slightly serrulate; pollen spheroid to irregularly ellipsoid, broadly triang- ular in cross-section, with 3 pores, the exine finely punctate. Fruit deeply 3-lobed, the lobes of the gynobase about 1 mm. long; seeds about 1.8-2.5 mm. long, 1.51.8 mm. wide; testa smooth, sometimes mottled, the caruncle absent. Desert sands, lava rocks, chaparral, dry sandy washes, and sandy roadsides. Flowers from the first of March to the end of April, and fruits from the middle of March to the middle of June. UNITED STATES: CALIFORNIA: Imperial, Los Angeles, Riverside, San Bernardino, and San Diego counties. NEVADA: Clark Co. ree о: е propi sandy arroyo margin, San Augustin, Gentry 4004 (GH, UC); 28.6. of Augustin on road to Catavina, Car E lexander & Kellogg ре MO); еы head, Guidi Island, i Palmer 785 (F, GH, US); 37 mi. s. of CH, Pozo Aleman, Shreve 7018 (F, MICH, US); sandy plains, eda Purpus 25 (F, US); Yecate, Orcutt s.n. (F, ), Socorri, биси. s.n. (F, MO); coast near Ascension Island, ) й amas Epling & Stewart s. n. (F, US); south slope 8 mi. from Rosario on road to El Marmol, Wiggins 4337 (GH, US); sandy wash at junction of El Marmol and San чамы roads, 25 mi. from El Marmol, Wiggins 4354 (GH, MICH, NY, UC, US); sandy flats of Pozo Pride Wiggins 7880 (F, MICH, NY, UC, US); sandy wash 11 mi. е. or Ignacio, Wiggins 7013 (F, MICH. NY, UC, US); along Rio Santo Domingo about 4 ids above the Mission, Wiggins 8 Demaree 4778 (F, GH, MICH, NY, UC, US). sonora: 30 mi. s. Ww. of ен on road to Punta Penasca, Sbreve 7501 (F, MICH, MO, US); llano 30 mi. from Sonoyta, паре. $350 (MICH, UC, US); 28 mi. s. of Sonoyta on road to Punta енн Keck 4183 (NY). 1951] ROGERS—REVISION OF STILLINGIA 249 Fig. 21. Séillingia linearifolia. DOUBTFUL SPECIES Stillingia cruenta Standl. & Steyerm. in Field Mus. Publ. Bot. 23:125. 1944. (T.: Standley 91207!). Without exception, the available specimens of this species are sterile. It will be impossible to assign the plant to a genus until flowering or fruiting specimens are available. EXCLUDED SPECIES In the following list, the species marked with an exclamation point (!) have been verified by examination either of the type specimen or of a photograph of the type, but those without an exclamation point are excluded only on the authority cf Pax and Hoffmann in Engl. Pflanzenr. IV, Fam. 147, V:197. 1912. Only those species found in the Americas are considered here. Old World species have not been included. No reference is made to Baillon’s 'Étud. Сеп. Euphorb.’ 1858, since Baillon rarely made formal transfers, although his taxonomy included many genera under Sfillingia. tu Vigne Muell. Arg. in Linnaea 32:87. 1863 — SAPIUM APPENDICULATUM (Muell. Arg.) Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, V:214. 1912! S. arborea Pav. ex Huber in synon. = SAPIUM PEDICELLATUM Huber, in Bull. Herb. Boiss. II, 6:352, f. 9. 1906. [Vor. 38 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN e eh il Ue Contr. Fl. Venez. 9. 1921 = SEBASTIANIA GRANATENSIS (Mucll. Arg.) Muell. A n DC. Prodr. 152: 1189. 1866! (Gymnantbes granatensis Muell. Arg. in Linnaea 32:1 S. babiensis (О Arg.) ere in Adansonia 5:329. 1865 (Gymnanthes bahiensis Muell. Arg. in Linnaea 32:102. 1863) = SEBASTIANIA BAHIENSIS (Muell. Arg.) Muell. Arg. in DC. Prodr. 152; 1183. 1866! S. bidentata (Mart.) Baill. in Adansonia 5:324. 1865 (Cnemidostachys bidentata Mart. Nov. Gen. & Sp. 1: J t. 43. 1824) = SEBASTIANIA BIDENTATA (Mart.) Pax in Engl. Pflanzenr. IV, Fam , V:113. 12, 5 н тэ ‘Baill. in Adansonia 5:320. i (Hippomane biglandulosa L. Sp. Pl. ed. 431. 1763) = Sapium HiPPOMANE С. F. W. Mey. Prim. Fl. Essequeb, 275, 1818. S. brasiliensis (Spreng wil in еккен 5:328. 1865 = SEBASTIANIA BRASILIENSIS Жолы Neue Entdeck. 2:118, £. 3. S. brevifolia = M Muell. үзер Baill. in Adansonia 5:328. 1865 (Gymnanthes Ll. brevifolia Kl. ex Muell. Arg. in Linnaea 32:104. 1863) = SEBASTIANIA BREVIFOLIA (K ex Muell. Arg. ell. Arg. in DC. Prodr. 15?:1186. 1866. S. Commersoniana Baill. in Adansonia 5:330. 1865 = SrBAsTIANIA KLOTZSCHIANA (Muell. Arg.) Muell. Arg. in DC. Prodr. 15?:1178. 1866 pides фа Klotzschiana З i ў. S. concolor о Baill. in Adansonia 5:327. 1865 (Gussonia concolor Spreng. Neue Entdeck. 2:120, /. 1821) = AcTINOSTEMON CONCOLOR (Spren а) Muell. Arg. Var. К. GENUINUS Muell. FR in Mart. Fl. Bras. 117:595, £ 505, f, П, S. coriacea. (Mart.) Baill. in Adansonia 5:323. 1865 VEM Bep aide i odi hig Nov. Gen. & Sp. 1:71. 1824) = .SEBASTIANIA MARGINATA (Mart.) Muell. n DC. S. corniculata (Vahl) Baill. Étud. Gén. pis Atlas, pl. 8, fig. 1-12. 1858 (Tragia corniculata Vahl, Eclog. Amer. 2:55, 1, 10. 1789) = SEBASTIANIA CORNICULATA (Vahl) Pax in Engl. hen IV, Fam, 147, ak 1912, SEBASTIANIA GLANDULOSA (Mart.) Pax, loc. cit. 100. 1912, SEBASTIANIA penis (Mart.) Pax, е cit. 105 12. S. cremostachya Baill. in Adansonia 5:322. 1865 = SAPIUM KLOTZSCHIANUM (Muell. Arg.) Huber, in Bull. Herb. Boiss. II, 6: s P 30. 1906 (барш fine a Neca var, є, Klotz zschienum Muell. Arg. in Рен 32:117. S. daphniphylla Baill. in Adansonia 5:326. 1865 = SEBASTIANIA DAPHNIPHYLLA (Baill.) Muell. Arg. in DC. Prodr. 157:1180. 1866. 27 (Spreng. Baill. in Adansonia 5: 327. 1865 (Gussonia discolor Spreng. Neue Entdeck. 2:1 1.2, f. 7-10. 1821) = SEBASTIANIA DISCOLOR (Spreng.) Muell. in DC. Prodr. 15°; ТРН 1866! S. divaricata Kl. ex Pax & Hoffm. іп synon. in Engl. Pflanzenr. eo E" 147, V:230. S. d Baill. in Adansonia 5:321. 1865 — rea КРЕНА var. p. DRACUNCULOIDES (Baill.) Muell. Arg. in DC. Prodr. 152:1207. 1866. 5, ке Эш жаны Rich. in Sagra, Hist. Fis. Cuba 11:202. 1850 = GrIMMEODENDRON gara ee osuM (Rich.) pos Symb. Antill. 5:398. 1908! S. frutescens Bosc. ex Steud. Nomencl. Bot. 1:815. 1821 — SEBASTIANIA FRUTICOSA ue Fera n. in Nodo 46: 45. 194 S. fru ulicosa ore ex 17% Syst. 3: 805. 1826 = SEBASTIANIA FRUTICOSA (Bartr.) Fern. in Rhodora 46:45. 5. Gaudichan (Muell. TR Baill. in Adansonia 5:332. 1865 = SEBASTIANIA GaupricHAUDII (Muell. Arg.) Muell. Arg. in DC. Prodr. 15?:1177. 1866! S. glabrata (Mart.) Baill. Étud. Сеп. Euphorb. Atl. 17, bl. 8, figs r3—16, 1858 (Cnemidostachys glabrata Mart. Nov. Gen. & Sp. 1:70. 1824) — SEBASTIANIA MULTI- uell. Arg. var. В. GLABRATA (Mart.) Pax in Engl. Pflanzenr. IV, Fam. I47, V:120. 1912! S. glandulosa Dombey ex Juss. in Ж” Sci. Nat. Bot. 25:24. 1832 = ApENOPELTIS COLLIGUAYA Bert. ex rud loc. cit. 183 1951] ROGERS—REVISION OF STILLINGIA 251 S. paeem ( Aubl.) Eu in ooa 5:332. 1865 — MAPROUNEA GUIANENSIS Aubl. Hist. РІ, Guian. 2:895, 2. 342. S. ыыы Standl. in e Mo. 122 Gard. 27:314. 1940 = SAPIUM MORITZIANUM Kl. in Seem. Bot. Voy. Herald, 100. 1852! S. hastata Kl. ex Baill. in od 5:324. 1865 = SEBASTIANIA po (Didrichs.) Muell. Arg. var. 8. GLAB uell. Arg. in DC, Prodr, 157:1174, 186 S. beterodoxa Muell. Arg. in Ex ph 89. 1863 — SEBASTIANIA HETERODOXA TNT Arg.) Benth. in Benth. & Hook. f. Gen. Pl. 3:334. 1880 S. Hilariana Baill. in Adansonia 5:332. 1865 — MAPROUNEA BRASILIENSIS St. Hil. Pl. n Bresil, /. 65. 1824-18 S. bypoleuca (Benth.) Baill. in Adansonia 5:330. 1865 = GYMNANTHES HYPOLEUCA Benth. in Hook. Jour. Bot. 6:325. S . jacobinensis (Mucll. Arg.) Baill. in Adansonia 5:329. 1865 == SEBASTIANIA JACOBINENSIS (Muell. Arg.) Muell. Arg. in ОС, Prodr. 152:1188. 1866. S. laureola Baill. in Adansonia 5:327. 1865 = SEBASTIANIA LAUREOLA (Baill.) Muell. Arg. in DC. Prodr. 157:1180. 1866. S. laurifolia Rich. in "a Hist. Fis. Cuba 11:201, pl. 69. 1850 =SAPIUM JAMAICENSE Swartz, Adnot. Bot S. ligustrina 56.15 Fl. ‘Bor Amer. 2:213. 1803 = SEBAsTIANIA FRUTICOSA (Bartr.) Fern. in Rhodora 46:45. S. marginata (Muell. Не Baill. in ME RET 5:321. 1865 — SAPIUM MARGINATUM var. а, LANCEOLATUM Muell. Arg. in Linnaea 32:120. 1863! S. multiramea (Kl.) Baill. in UE $: 325. 1865 (Sarothrostachys multiramea К}. in Wiegm. (Erichs.) Arch. 7:185. 1841) = SEBASTIANIA MULTIRAMEA (Kl.) Muell. Arg. var. 8. GLABRATA (Baill.) Pax in fu Pflanzenr. IV, Fam. 147, V:120. 1912! S. myrtilloides (Mart.) Baill. in Adansonia 5:323. 1865 (Cnemidostacbys myrtilloides Mart. Nov. Gen. & Sp. 1:67, £. 40. 1824) = SEBASTIANIA MYRTILLOIDES (Mart.) Pax in Engl. Pflanzenr. IV, Fam. 147, V:93. 1912, SEBASTIANIA OLEOIDES (Mart.) Muell. Arg. in S. nervosa (Muell. Arg.) Pu in Adansonia 5:328. 1865 (Gymnanthes nervosa Muell. Arg. in Linnaea 32:102. 1863) = SEBASTIANIA NERVOSA (Muell. Arg.) Muell. Arg. in DC. Prodr. 15?:1183. 1866! S. obovata (Kl. ex Muell. Arg.) Baill. in Adansonia 5:321. 1865 — SAPIUM OBOVATUM Kl. ex Muell. Arg. in Linnaea 32:120 N S. pachystachya (Kl.) Baill. in Adansonia | 5:330. 1865 (Adenogyne pachystachys КІ. in Wiegm. (E Erichs.) Arch. 7:184. 1841) — аА PACHYsTACHYS (Kl.) Muell. = in DC. Prodr. 152:1182. 1866! S. patagonica (Spegazz.) Pax & Hoffm. in Engl. Pflanzenr. Ad Fam. 147, V:188. 1912 (Colliguaye mt. Spegazz. in Revist. Facult. Agron. T Veter. La Plata 3:572. 1907) UM patafonicum (Spegazz.) D. J. Rogers, comb. This snecies b very few affinities with any other id. species of Sapium, Stillingia ог Colliguaya. However, the absence of a gynobase, the axillary inflorescence, the struc- ture of the staminate flower, the shape of the glands at the base of the leaf blade all point to characters of Sapium, not Stillingia. In one character, the texture of the seed coat, this species shows no affinities with the American species of Sapium. All of the new world species of Sapium have an arillate seed coat, but S. patagonicum has a firm, hardened testa, without evidence of an aril. S po Baill. in Adansonia 5:331. 1865 = SEBASTIANIA SCHOTTIANA Muell. Arg. var. 8. ANTHIFORMIS (Baill.) Pax & Hoffm. in Engl. Pflanzenr. IV, Fam. 147, Mis 127: S. prostrata (Mart.) Baill. in 52 5:324. 1865 (Cnemidostachys prostrata Mart. Nov. Gen. & Sp. 1:70. 1824) — ASTIANIA CORNICULATA (Vahl) T var. Р. PROSTRATA (Mart.) Mus Arg. in DC. Ы 152:1172. 1866!, SEBASTIANIA GLANDULOSA (Mart.) dem in Engl. Pflanzenr. IV, Fam. 147, V:100. 1912, and Eid. HIsPIDA (Mart.) ах, loc. cit. 105: 1912 [Vor. 38 524 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN S. pteroclada (Muell. Arg.) -— in Adansonia 5:329. 1865 (Gymnanthes pteroclada Muell. Arg. in Linnaea 32:107. 1863) = SEBASTIANIA PTEROCLADA (Muell. Arg.) Muell. Arg. in DC. Prodr. il ee om S. ramosissima (St. Hil.) Baill. in "ON 5:328. 1865 (Microstacbys ramosissima St. Hil. Hist. Pl. Remarq. me 242. 1824) = SEBASTIANIA BRASILIENSIS Spreng. Neue 182 S. rigida (Muell. Аг), ` Baill. in Adansonia 5:330. 1865 (Gymnanthes rigida Muell. S. in Linnaea 32:99. 1863) = SEBASTIANIA RIGIDA (Muell. Arg.) Muell. Arg. in DC. Prod. 157:1180. 1866. S. rufescens Moritz ex Pax & Ho сш” in synon. in Engl. Pflanzenr. IV, Fam. 147, V:212. 1912 = SaPIUM sTYLARE Muell. . in Linnaea 32:119. 1863. S. salicifolia Kl. ex Baill. in "матова" 5:320. 1865 = — SAPIUM HAEMATOSPERMUM "e Arg. in Linnaea 34:217 і S. Schottiana (Muell. Arg.) Ball in Adansonia 5:331. 1865 (Gymnanthes Schottiana Muell. Arg. in Linnaea 32:96. 1863) = SEBASTIANIA SCHOTTIANA (Muell. Arg.) Muell. Arg. in DC. Prodr. 15?:1176. 1866! S. sebifera (L.) Michx. Fl. Bor. Amer. 2:213. 1803 een MM L. Sp, Pl, ed, 1, 1004. 1753) = SAPIUM sEBIFERUM (L.) Roxb. Fl. In 3. rrata Kl. ex Baill. in Adansonia 3:329. 1865 dig ede ed SERRATA (Kl. ex S. serrulata (Mart.) Baill. in Adansonia 5:324. 1865 еее serrulata Mart. Nov. Gen. & Sp. HAM f. 42. 1824 — SEBASTIANIA SERRULATA (Mart.) Muell. Arg. var. a, Ка” uell. Arg. in DC. рой, 15?:1167. 1866! S. stipulacea (Kl. ex Muell. Arg.) Baill. in Adansonia 5:325. 1865 (Gymnanthes stipulacea Kl. ex Muell. Arg. in Linnaea 32:96. 1863) = SEBASTIANIA sTIPULACEA (Kl. ex Muell. Arg.) Muell. Arg. in DC. Prodr. 15?:1176. 1866! S. sylvatica Garden var. paraguayensis Morong, in Ann. N. Y. Acad. 7:226. 1893 = SAPIUM HAEMATOSPERMUM Muell. Arg. in Linnaea 34:217. 1865! S. trinervia (Muell. Arg.) Baill. in Adansonia 5:328. 1865 (Gymnanthes trinervia Muell. Arg. in Linnaea 32:101. 1863) — SEBASTIANIA TRINERVIA (Muell. Arg.) Muell. S. Weddelliana Baill. in Айко 5:329. 1865 = SEBASTIANIA WEDDELLIANA (Baill.) Muell. Arg. in DC. Prodr. 15?:1188. 66. 8, Widgreni (Muell. Arg.) pro in Adansonia 5:326. 1865 (Gymnanthes Widgreni Muell. Arg. in Linnaea 32:97. 1863) = SEBASTIANIA WIDGRENI (Muell. Arg.) Muell. = in DC. Prodr. 15?:1178. inte anemensis (Muell. Arg.) Baill. in Adansonia 5:330. 1865 (Gymnanthes ypane- mensis Muell. Arg. in Linnaea 32:100. 1863) — SEBASTIANIA YPANEMENSIS (Muell. Arg.) uell. Arg. in DC. Prodr. 15?:1179. 1866!| INDEX TO EXSICCATAE Italicized numerals refer to collector's numbers 5, s.n. (sine numero) to unnumbered vag Een numerals refer to the numbers of taxonomic entities conserved in this r Abrams, L. R. 3227 (23). Arnold, B. G., & L. E. 16026 (18a). Abrams, L. R., & McGregor, E. A. 519 (21). TN L. E. 16021, 16022, 41514 (18a). Aguilar, I. 93, 1314 (16); 220 rd 760, rnold, L. E., & West, E. 16017, 16018, 1106 (6). No Р (1 84). Albers, С. С. 32081, 34044, 34046, 30052 Arsène, ^ gro. G. 1400, 5236, 6679 и. (18а); 34043, ‚ 34047, 38017 (17). Arsène, Bro. С. & Nicolas, pis 2 (16). Alexander, A. Kellogg, L. 71377, Ashe, W. W. s.m. (18a); D 2240 ен ке; y s Ayers, H. s. n. (18a). Alonis, T., Johnson, J. C., & Barkley, F. A. Balansa, B. 1708, 444 (14). 151 89M (20). Balls, E. К. В-5640 Angier, B. 5. 19, 138 (23). Barkley, F. A. 1205 Mo. 13306, 14521C Anisits, J. D. 2435 (14). (17); 1951] ROGERS—REVISION OF STILLINGIA Barkley, F. A., & Painter, J. T. 74253 (20). ye Barkley, F. A., & Webster, С. 14407 (20 Barnhart, J. H. Го 3113 (18a). Barreto, M 75 Bartlett, H. H. Pe “11025 (4). Beck, E. s.n. (18a). Berg, N. K. s.m. anc Bernoulli, G. & Cairo, К. 2499 (19). Berry, E. A. 97 (8). Beyrich, H. C. 2 1853 (18а). Bigelow, J k ». (17); 5». (192); n 5272 (20). Biltmore, Herb. DOS 14937 (18a); ap CAE ; 35250 (8). р s." Ж (22); (23). 116 (18a); s. m. * (17); 5.7, Britton, N. L., & Wilson, Р. Burger, О. F., & West, E. 5. (18a). Bush, B. F. jo 057; 1245 (18а); 847 (20). Butler, G. D. 49, 1129b Canby, W. s.n. (8); 7510 л Carleton, M. А. 17837 (18a). Carter, ze | T. A. M., & Kellogg, L I s A rip 1: 1, ғы. (21): W. 191, S:n. b $1. (89% (18a Cohn, J. 7135 (1 eid 963 (23). Cooper, J. С. s. n. (18a); s. м. (22). E reli D. S. 5648 (8). Cory, V. L. 5725, 13510, 21776, 37486, кнд, 18а); e) 35050, 45204; 428 (17); Ar, Аг, A3, 527, 1971 (20 Coulter, Т. 1502 (4). Croft, М uH oA 20). Cummings, E. Ј. s.*». (18а). Curtiss, А, Н. 2508, 4276, 4851 (18a); 299 2509, 5892 (8). Cuthbert, А. s. n., 1316 Cutler, H. C. 3211 o 252 (17 D M.E. 750 (16). Davis, J. Н. s. n. (8). ‚С. С. 6074 (16); Ж (8). Пеат Degener, О. 4578 (21); 5053 (17). Demaree, D. RON К. г ғың 13071, 18719 Dixon, К "is 344 (18a) Dodson, К. s.n. (182). Drummond, T. 38 (8); 229 (18a). Drushel, J. А. 2562, 10437 (18а). Duckett, F. s.n. (18a). 242 (7). zs 5212 Dusén, Р. 0143, on ae (15); Gi F. 22 5 8а Eastwood, А. 2758, 2078, 3042 (23) 33057 22 SC (20). Edwards, M. T. (22). Edwards, Dr., & Eaton, EU 359 (20 5.2. (17) 5 s.m. (18a); s.n. (20). Eggert, H. Eggleston, W. W. 20107 (18 Ekman, E. L. 506 (14). Elmer, A. D. E. 3675 (21). Eman-Meyer, —. s.n. (18a) Emory, W. H. s.n. (22 ). к” apt 23I (18a); s. v. (23). Epling, C (22). Epling, C., Ellison, L., & Anderson, Н. s.n. 22 ( Epling, с., & Robison, W. s.n. (23). (18а). Eyles, D 8144 (18a). Fennell, J. L. кл (18a). Ferguson, A. M. s.n. (18a dor MN. 1... E pcd, Г: & Long, B. 7 (18a). "eui M. L., 8338, 10318, 12125, 12397 (18a). Ferris, R.S. 7099 (23); 7119 (22). R. S., & Rossbach, R. P. 9521(22). g B. 7508, 8337, оо, 10460, 12124, . ‚ (18a). ы А. na 5034 (182); 5658 (8). oer NN 2938 (1). гуе, О. E. s.p. (18a Frye, T. C., & E. M. 2375 (20); 2493 Es Gaiser, —, Butts, —, & Arnold, E. 26 (18a). Gander, F. F. 3005 (23). 254 Garber, A. P. s.n. (18a). Garrett, R. s. м. (18a). Gentry, H. S. 4004 (23). Gibbes, L. R. %.». (18а). Gilman, M. F. 379 (22). Glaziou, A. F. М. 6126 (11); 10852 (12). А R. i 4595, 5004 HE 8a). Godfrey, R. K., & Shun (18 4). Godfrey, К. K., & Tryon, R. M., Jr. L V. 4212 235, 1033 (18а). Goldman, E. А. 902 (4). Goldsmith, G. W. s.n. (18а). Goodman, G. J. 2120, 2603 (18a). Goodman, G. J., & Hitchcock, C. L. 1127 (18a). Gough, L. C. s.n. (18a) Gould, М. s.#. (18a). Graves, E. W. 6773 (18a). Gray, A n 4” Gregg, 4 ut n., 460, 551 (8); 944 Наме, Т. 614 (19). "UG Hard E. 888, 2110, 3304, 6346, 7782 (13); 4360, 4446, 4404, 4704, 5612, 6790 p^ Havard, 15 (20). Hawkins, е s.n. (8). m A. А. 021, 1530 (18a); 1714 17). Heyde, E. T., & Lux, E. (5); 4579 (16). Hieronymus, С. 617 Hilyand, E. s.n. (18a). 473 (19); 4265 ood, 24611, RS 140 24614, 24615 (18а). ooks, —. s.n. (1 Hopkins, M. 110 (18a Houghton, W. 3837 (18a) House, Н. D. 2626 (18 Howard, D. V. НЫ (17). Hubricht, L., Shoop, C., & Heinse, D. B. B1382, 81397 (18а). Huger, A. M. 23 (18а). Hume, E. G. s. ұй (18а). Metz, M. С. [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Hyams, M. e E п. (18a). James, E. Jermy, б. s. n. 7) 237 (20). ewett, А. s.n. (18a). Jórgensen, P. Es (14). кч R. 951 (19). nes, M. E. үйді 26018, 20115 (20); 23). Keeler, Н. D. 18a). Кабын, W. А. 6576 (19). Kellogg, W. H. 1 (17). Кепоуег, L. А. 266, s. n. (4). ert, М. -sone (21) Keyser, M. 6027 (18a). Killian, O, $. f. (17); s. n. (18а). Kirk, В. s. т. (18а). Knight, А ола (18а); sm (B). Leavenworth, W. C. s.n. (8). Leavenworth, W. C., & Hoogstraal, H. 1821 (16). Lehman, V. s.n. (18 emmon, J. б. 239 (22) LeRoy, P. V. s.m. (23 LeSueur, Н. s. ә. (18 ). ; 5.7, (18а); s. 2. (17). Lighthipe, L. H. 423 (18a). Lindheimer, F. 62, 169 (18a); Ór, 244, 519, 687, 712, 1152, I153 (17). . (5 Lundell, C. L., & A. A. 8756, 0482 (18a); 8868, 0152, 10614 (17); 0810 (20). Lyonnet, E. 988 (16). MacDougal, D. T. 206, s.n. (22). MacFadden, F. 73609 (21). MacKenzie, К. К. 2071 А Marsh, Е. 80, 201 (17); 1116, 1768 (2): Martin, —, Devall, —, & Arnold, E. (18a) Maso 15-3709 (23). Mathias, M. E. 860 (21) Matuda, E 6 Y .Q 3). — J. B. 868, 7614, 8132, 8210, 8322 (18a). McMullen, E. s.n. о Mearns, E. А. 3129 (23 n I. E., & Goodman, E J. 3617 (19). Mell, C. D. 8). Mellichamp, Lu s. n. (18a). 155 (12); 219 (18а). 1951] ROGERS—REVISION OF STILLINGIA 255 Miller, К. s. м. (17). Porter, T. С. s. ә. (18a). Mitchell, M. s.n. (18a). Pringle, C. С. 128 (7); 6886, 7244, s. n. Mohr, C. s.n. (8). (16); 2070, 2534, 13756 (4); 2071, Moldenke, H. N. 298, 572a, 1102, 3630, 2504 (20). 5570, 5052 dn 540 (18b); 906, II52 Purpus, C. A. 25 (23); 1320, 1454 (7); 3523, 4980, p (4); 5492 ro 7343 rse, F. P (19); 9174, 9323, 10334, s. n. (16). Mueller, C. H gor) 2668 (4). Rambo, В. 2278, 4818, 4824, 6575, 28800, H. T. 467 (4). Myers, S. Е. Sur Sa) Nash, G. V. 366, 1850, 2428 (18a); s. n. 216 (20); s. n. (18a). 6605 (4). Nicholson, A. J. s.n. (17) Nicolas, Е. 5. м. (16). Nolen, He ул (183) Norris, J. С. 500 (23). Ohlinger, L. B. 418 (18a) Olive, E. W. 110 (18a) O'Neill, H. s.n. (18a) D'Orbigny, A. 018 (13). Orcutt, C. R. 5988 (20); 6068 (17); no : Oyster, J. Н. s. n. (17) ; s. n. (18а). Palmer, E. 234, 1255 (4); 73 (16); 1256 (17); 313, 405, 406, 7515 (18a); 7512 (8); 1257, 1208 (20); 517 (21); 147, 440, 785 (23 Palmer, E. J. 5, 7976, 11645, 14120, 14277, nord (182), 6538, 6564, 13516, 13760 (17). Pammel, L. Н. s.n. (18a). Parish, S. В. 3007, 3652, 4657, s. n. (23); 4142 (22); 2458, 3731, 11856 (21). „Е. 706, 1371 (22); 1). 1027 (17): 1534 (18а). Parry, С. С. s. n. (22); s. n. (23). Parry, C. = Bigelow, J. "M, Wright, C., & d ,À. s.m". С: с. &-Lenon. ]. G. 376, t. . (23); 377 (22). Parry, C. C., & Palmer, E. 823 (16). Pavon, J. 7195, s. n. (4); s. n. (16); s.” (19). Perkins, J. R. s.m. d Pickell, L. W. 250 (18а) Pierotti, S. 5305, 5384 (14). Pilant, . (18a). Pitcher, 2 82): Plank, 20). райға, С. L., & "Collins, G. N. 234 (8). 28353, 8 44, 34775, 43136, 43828, 44828, 44955, 46236 (1). 423 (19). i ilico. Е R. I (18a). Randolph, F. R., & Small, J. K. 70 (8). Rehder, А. 071 (8). Reverchon, J. 133, 1352, 2. С 56, 866, s. n. (18а); 24 173% ‚тп. (17). ay aD; II72 (12). 4, $ 5 E9343. 1-1, 15-5, 15-7, 15-10, 15-11 (18a); 8, 9, IO, 14-2, 14-4, ША E Le 2! 15-0 (8); I2-I, 12-2, I2-3, 12-4, Rose, J. N., Painter, J. ү & Rose, J. S. 53 Rose, L. S. 34404, 36840 (23); 36704, 38343, s. n. (22); 40310 (21). Rose-Innes, R., Moon, B. 995 (18a); 1297 (20); 1317 (17 1256 (19); 1358 (5). A id s. n. (8). Russell, P. G., & Souviron, к Т Ruth, A. 178, $ "CIA ); 519, 5.%. (18a). Rydberg, P. A., & Imler, R. 304, 624, 905 (18a). 7513 (18a 1300 (18а); s. m. (22). aurman, B. F. S Schallert, P. O. Schott, A. 5. 2. (17); s. 1. (20) ; s. 1. (22). Schwarz, G. J. 1354, 2518, 3520, 5105, 8301 (14). Sce L. B. a. SUM Seibert, R. J. 9 (8). Seler, E. 2381, уя (19). Sellow, F. 4085 (1). Sesse, Мосіпо, Castillo, & M 4241, 42 (16); 4240 (4); 4283 (19). P ciun) М. 15, s.n. (18a Shreve, F. 7018, 7038, 7302, 7501 (23); 454 (20). 256 Shreve, F., & — E. R. 9572 (20). Sinclair, J. F. з.» Skutch, A. F. 29, i 1165 (1 9). Small, J. К. 406r, 8344, 8385, "e (8); 7260, 7001, 7073, 8501, s. n. (18a). Small, J. K., & Carter, J. J. 65, 515, 716, 1054, 1438, 1439, ы 2528 (18а); 1005, 1279, 1440 (8) Small, J. К., atis J. ГА & Small, С. К. е а Small, J. К 2. f B. 9163, sis 9067 (s pie (18 l, J. K., DeWinkeler, J. Е & Mosier, 11235 (8). Small, J. K., & Mosier, C. A. 5000 (18a); 6307, 2535, 2037, 3083, 53169 (8). Sma . K., Mosier, & Small, G. K. 6600, 6724 (18a); 6608 (18b) ; 5065 (8). Small, J. K., & West, E. 16034, 16035 (18a); 16008 (8). Small, J. K., & Wherry, E. T. 72012 (20). Small, J K., & Wilson, P. 1580 (18b). Smith, C. 2709 (18a Spencer, M. F. 245 (22); II (23); 1450 Ж Standley, J. P. 16, 473 (18 Standley, P. C. 57803, Stanfield, S. W. sm. (з ic Stanford, L R., Retherford, K. L., & North- 94 (7). . 3). Stevens, С. W. 855, 1106, 1022 (18a). Steyermark, J. A. 32201, 50010 (5); 32895, 34399, 36377 (19); is (6). Stork, H. & Horton, О. В. 10018 (3). Strandtmann, R. W. Taylor, K. A. Thackery, F. A. Tharp, B. C. ro 590, 627, 259 (23). 4, 2906, 44210, s. n. (17); 628, 6300, s.n. (18a); s.m. usd GO. n9 (17). EUNT OW. &» Pei Tisdale, (18a). Tolstead, Y. L. 7308, 7543 (17). Torrey, J. z n. E Tracy, S. 6555, 7855, 7856, 9411 н. dd (8); 7864 (17); 9005 je P. Trecul, M. 1346 (23 1458 d [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Trelease, W. „15, 5. п. (18а). Tweedy, Е. Ule, Е. ms эу, 7160 (10). Underwood, L. M. . (18a). Uzzell, P. B. 110 (2 she Т; ine m 0. ae 2158, 3130 (6). Von Schrenk, H. 617), ). : 1055 (17); 20677 bw Warnock, B. H., & Krodell, W. J. (17). Waterfall, Ag T ‚434%, 6523, 6741 (17). Watkins, (17). atson, T | T m Waugh, F. A 8a). Webber, H. J n D E (18a). Weber, G. F. s.m. E 41510, 41522, 41524, 41528, 41888, hd зу Ao ). White, M. 214 (18а). Whitehouse, E. 534, 535, 537 (18a); 536, Wischmann, 2 а ЖЫ; а). Manning, W. E. Wiegand, K 1835, 1836, do 23 ed Wiggins, 1. L. 7, 4354, 7880, 7913, 8350, 9705 23); : 9015 5 (22). Wiggins, I. L., & Demaree, D. 4778 (23). Wild, E. s. ". (17). Wilkinson, E. H. 17). 7 ( 127 Williams, L. О. 3814 (16); 13150 (19). Williams, L. O., & Molina, А. 10603 (6); 11844 (19). Wilmot, R. J, & Murrill, W. A. s.m. (18a). Wolf, C. B. 2284 (22); 2460 (23); 6524 (21). Wood, A. 2508 (17); s. n. (18a). Woodcock, F. С. 1813, 1814 (23). oodson, R. E. Jr., & Anderson, E. S. 1607 (18a). Woodson, К. E., Jr., & Schery, R. W. 06(18а). Wright, C. 182(23); 1792 (17) ; s.n. (20). Wynd, L. F. 705 (17 Wynd, L. F., & Mueller, C. H. 225 (17). York, C. L. 46107, 46117, 46192 (17). York, H. H. 66, s. n. (17); s. n. (18a). Young, M. 55, 5. n. (17); s. n. (18a). Young, V. 1504 (18a). 1951] ROGERS—REVISION OF STILLINGIA 257 INDEX TO SCIENTIFIC NAMES the following index, names synon 56 аге іп ед new names erences are in bold face type. which are recognized as valid are in or combinations and principal taxonomic page ref- Actinostemon concolor var. genuinus- 250 Adenogyne pachystachys ---------------------- 251 Adenopeltis colliguaya -------------- 250 Cnemidostachys bidentata -n сЕ Еа 250 COTSGGE Go лкы к са 250 glabrata 250 myrtllodes — E 251 prostrata -= S 251 serrulata 252 Colliguaya patagonica EAREN ОБК. 251 Croton sebiferum -------------- 252 M 219 Ехсоесана te anaa 20 09 АА var. dracunculoides ... 250 Bo ни EL ЗЕ эс. э быды 222 salpingadenia --- 234 cu dE eglandulosum ---------- 250 Gussonia concolor -- 250 discolor ___..--- 250 Gymnanthes abiensis - 250 brevifolia etc РИНЕН 250 granatensis -- 250 hypoleuca -.... 251 Klotzschiana 250 VOSE E: 251 pteroclada ----------- 252 irida ou ELE cscs ық 252 Schottiana —-- 252 stipulacea ——— 252 Treculiana --- - 246 trinervia M eL E 252 Widereni Ды мше 252 banemensis а. а шЕ 252 Gymnostillingia ------------------------------ 208, 243 acutifolia: eo ЖЕ ы 243 lorantbaces re 232 macrantha coe 243 Hippomane Penes MEDIA Ж 250 Maprounea —— = 209 ee NNNM Eos 251 guianensis: aee- 251 Microstachys ramosissima __--.--.------------- 252 ES QNM NNNM с> 207, 209 eculifolium —— —— ———— 243 ай Е -. 247 var. dentatum BEEN Ee ns 246 appendiculatum ——— — — — — ——— 249 ipii ctc var. Klotzschianum_.. 250 Bodenbenderi __.-..------------------------—-- 222 Mos 5 234 Roman type; haematospermum.---------------- 812234; 252 haematospermum | . 234 hippoma 250 jamaicense -------------- 251 Klotzschianum - 250 margina 211 var. lanceolatum 251 "m aga RES 220; 1251 obovatum ——— 251 opposttifolinm ae sed 221 goni _ 251 pedicellatum ————— 211, 249 salpingadenium |... 234 se um 21459252 САР ТЕТЕ 2502 e LL 252 subsessile .-------- 222 орайсин var. linearifolium ------------ 238 Eu 236 омс: multiramea ------------------ 251 Sebastiania . 209 bahiensis ---------- 250 bidentata —————— 250 7 -eaa 250, 252 2 Stent мез ы с 250 Eh Le ее =. 250 var prostrata а. 251 daphniphylla .. 250 DITE uo RM 250 uds var. Бар? а S — — 251 BEEN SAN 250, 251 Gaudichaudii -- 250 BIN .—— S A 250, 251 granatensis . 250 NL RE DDR Feet TL St REO 251 hispida ------- 250, 251 jacobinensis BAONE AES PTEE 251 Шокан... 2а... 250 um EE NO аны ыдысы 251 шанаға, eme л seca 250 mult tiramea var. glabrata... 250, 251 NORA с queni 251 соу o 2222 eS 251 oleoides 251 pachystachya — IR dee 251 pteroclada. LE e с... 252 дірдек 252 p 2500. 252 аг. уа экы; 251 БЕРГА c —- do pA ман var. КІосѕсћіапа_....- 252 Кбе .— I s. 252 Treculiana ------- 245 trinervia -- 252 [Vor. 38 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN Weddeliana — . . ---. 252 Hola НИИ TO iiit 251 242122 КЕСЕНЕ 252 ОЕ: сазды лысый 251 ypanemensis ------------- E 252 M OP су (section) — 55. TS T ee О шл: ысы осы. sirna ыы icai ai oae NEN Lee erlassen linearifolia 213, 248, a аса ой С 208, 212, 218, 243, 244 linearifolia есе ACUTIFOLIAE (series) 210, 215, 217, 218, var. paucidentata 0 246 243 loranthacea -n 232 2” саана 238, 240 crantba . ERES 243 ———— marginata - appendiculata MR ELEA microsperma s as 225, 226 aquatica... 211, 213, 215, 216, 229, E multiramea 5 BOE кишин ИУ КАНИКЕ 249 со) с Lees шы ДИ has jag, LL Lh рсе ссе 250 MIU эск eret ct ыи ДА PEN СЕП еттек rito - 250 obovata . Mon ы сыа ы нс этади —— "— HÓ 212, 228 oppositifolia c, 212, 220 bidentata -- COR PERPE СОО OPPOSITIFOLIAE | series) 521062213. 214. bislendulose STERN еск дасында 250 216, 218, 220 Bodenbenderi 0 211, 215, 222 Pachycladae Minas OTI PR hE ondes 250 pachystachya . Ад кебе NE о ы, Сз с=с 2 250 Г уст 2 cce ТІ Commersoniana _............---......-..------ 250 paucidentata 213, 214, 246, 247 о ie te REE oe . 250 peruviana (00 ыы 250 bci ina 51 corniculeta ein 250 DON „еол быы ee. cremostachya A E EEN ЖІ jx PEORES ОМ cruenta —— —À ? phorocieda- ez cer E da че, рупа RE ыан кенг Erie 250 POMOSI Lacon еы с ыз 252 ог ie ИНИСИНИН 245 Heo асе 252 di ren gO ыыы It tatis 232 оо ЕСИЕТ el 252 ас (series) 210, 212, 213, 216, BOUES. а н саана 235, 240 ‚2 salicifolia 239, 240, 252 diphtherina | -..211, 226, 227, 228 ѕаіріпвадепіа 211, 215, 234 с AERE ERROREM 250 salpingadenia divaricata -a = кр, (uM LLL LS A dracunculoides 0 — var. сири {ета 000 234 КИ a ЛИН 212, 215, 2м аг. EAE SS Lh oaa t 234 eglandulosa м sup oux S o S S S S V nns 235 p (ЖОШО) aee ы E vat, angustior — .. 235 HEN. Leine Sidi ouais ы ыы 250 var. elliptica 5 235 үдеген ECRIRE еы ен с Дед 250 var. grandifolia Fruticosae i ЖКУ УАКИЛ 220 sanguinolenta 2-2 211, 224, 225 Gaudichaudi 2—2... 250 var. angustifolia 000 224 БА БЫ ыны сары; 250 var. lanceolata |... glandulosa 2. 250 rii. эма Л ыыы доз 212, 232 RENE n ыыы м 251 saxatilis - Я &ymnogyna 2 —-— 248 var. grandifolia MONTRE. GYMNOSTILLINGIA- (subgenus) FN 210, var. salicifolia 2 215, 235 212, 213, 214, 215, 243 fe: Gymnostillingia apo --243, 245 f. latior —__ 235 baematantba |. INI ЖТА tiana ----- 252 MEG. хы du ,-- . 251 scutellifera 00 211, 215. 285 heterodoxa 2000” 251 MERE Leo eta RE e use 252 Hilerlene .. — . .. 251 MEE uec coc asado qiia eae 252 Mite — ор К... 251 serrulata ә Я 252 Mob s LS 251 mallii 240 1951] ROGERS—REVISION OF STILLINGIA 259 spinulosa шс E LEA 213, 247 tenuis - 241 SH NE (subgenus) ..210, 212, 213, texana 238 214, 220 var. latifolia 238 stipulacea сі Қ ies e iu etr var. typica --- 238 бутана 213, 215, 219, 239, 2% Torreyana 245 ssp. sylvatica_______. Pais озыне ti OL ECT ONE 217, 231 ssp. tenuis... 215, 241, 242 Tieculana соз БК 244, 245 sylvatica енер (series) ------- 2106. 215, 217, r АШЫ о. ч 239 218 уа PNE Rer ДР ВЕНЕ МААНА 239 trinervia —- 252 var. linearifolia ------------- 238 ала ыы Сал лата. 213, 231 var. paraguayensis ----------------------- 252 Weddillisia - LR a us 252 var. ЗЕЛСИОМЕ oae ue 239 Итен — —— КС Є ee ee rn sutbuldid л ee e 239 Dr vl Mt —— ан e ОСПОЙ): Lose сы 1 тэс 214, 217, 218, 236, т SYLVATICAE (series) 210,214, 215, 217, Stillingiina d 233 Tragia о Em 6.1250 THE RECENT INTRUSION OF FORESTS IN THE OZARKS AUGUST P. BEILMANN* AND LOUIS G. BRENNER** INTRODUCTION Scanlan (1950), in writing of American forests, says: "The lumber barons began to despoil and they did an excellent job." This statement might correctly be applied to a specific region but a lumber baron could not operate in a prairie region. Seemingly, it cannot be applied at all to the eastern and northern portions of the Ozarks. Within historic times this vast region was a prairie, or at least park-like in that the trees were widely spaced and confined to the water-courses and drainage-ways. The logging operations which are now so much a part of the industry of the Ozark region are but little more than a century old. The loggers have been, and still are, cutting the first crop of trees to mature there. In some sections any tree large enough to yield a 2 x 4 is cut; and the region will be scoured again and again for more trees of that size. However, there is evidence to show that insufficient time has elapsed to develop a mature second growth of either pine or hardwoods. During a study of the flora of the northeastern edge of the Ozark Uplift an effort was made to visualize the steps which are supposed to precede the formation of a mature forest—the "climax" of the ecologist. This time-elapse study of only twelve years revealed a speeding succession of plant species not at all approaching the accepted trial-and-error elimination which is supposed to set the pattern for our forest areas. In an effort to reconstruct the Ozark forest before the advent of the white settlements and before logging had progressed very far, a search was made for old trees. After considerable difficulty many old trees were found which bridged the gap between grassland and forests. In one detailed study of a relic (Beilmann, 1943), a ring count gave an estimated age of 327 years. This was a "wolf" tree with heavy lateral branches whose tips touched the ground; it had grown as an isolated specimen and only recently had it any neighbors. ECOLOGY Steyermark (1940) made the first critical study of plant successions in the Ozarks. He found and described 164 examples of "large-scale natural plots." According to him, the classical Maple-Beech Climax, as proposed by Clements and Weaver (1929) on the basis of climatic conditions, does not hold in this area. For this he would substitute five edaphic associations based on the physical, chemical, and local moisture conditions. It would seem to be quite difficult to find such ec vast variation in soils that one might find 164 "associations" based on edaphic conditions alone, unless a certain immaturity was a characteristic of the forest aspect. Steyermark devotes one chapter to "variations induced by burning, logging, * Manager, Missouri Botanical Garden Arboretum. ** Assistant Manager, Missouri Botanical Garden Arboretum. (261) [Vor. 38 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5, and other unnatural causes." For centuries, burning, aggravated by re- duced precipitation, governed the distribution of forests. As the area became settled, the decrease in wild fires and the increase in rainfall reduced the loss by fire. Logging and clearing followed, but logging reached its peak in 1900. There- fore, the unnatural causes listed by Steyermark, together with edaphic adaptations, govern the distribution of species in the Ozarks. The writings of the earlier clearing travelers indicate that not nearly enough time has elapsed for the development of either а monoclimax or а polyclimax forest, unless we are prepared to call any group of plants a climax, regardless of how ephemeral the association may prove to Бе. An ecological study of the Ozark forest must not overlook the short time which has elapsed since the area was either "sterile" or grassland. HISTORICAL ACCOUNT OF THE OZARKS VEGETATIONAL ASPECT The Ozark Highlands of Missouri is unique as one of the oldest land-masses of the North American continent. Here on a land of archaic plains, deeply dis- sected by rivers entrenched in ancient meandering courses, erosion has carvéd in high relief, the land of rugged beauty we know today. It is only fitting, then, that rooted into a land having such a singular geologic background we find a flora equally interesting. In the Cambrian and Ordovician rocks, which comprise the greater part of the Ozark highlands, the history of the area may be read, and there is little evidence today of the varied floral aspects presented by Ozarkia in times past. However, turning to the diaries and journals kept by early explorers and travelers there is considerable evidence that the vegetational aspect of Ozarkia has never been static, and the rich hardwood forests which today add so much to the beauty of the Ozark mountain region are of relatively recent development. The early Spanish and French discoverers in the Mississippi and Missouri River Valleys give us the first accounts of the early aspects of the Ozark region. Bearing in mind that the primary interest of those adventuring soldier explorers was the discovery of land rich in precious metals for the crown and that they were inter- ested more in fruits and edible herbs, wood for fuel, and the construction of boats and simple fortifications than in botany, it is understandable that only a scant record was made of the native vegetation. ‘The general structure of the Ozarks is of Cambrian and Ordovician rocks of more or less uniform composition, and a general habit for the whole region may be assumed on the basis of material reported from relatively localized areas, except where special modifications due to soil and water are noted. The Ozark area was discovered by Ferdinand DeSoto in 1541 and included in the region then known as Florida. Houck (1908) believes DeSoto entered Missouri from Arkansas and, because of the numerous swamps of the region, took advantage of the highland now known as Crowley's Ridge. It was from here that the hearts of the travel-weary soldiers of DeSoto were lightened by the sight of the village of the Casquins. We are told that the fields upon the rich alluvial bottoms were planted in maize, and the pecan, plums, and mulberry trees were abundant. That 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 263 it was possible to see the distant village can only have meant that the soldiers were in an open, very park-like country unlike the dense forest of hardwoods that cloak this region today. In the narrative of Garcilasso de la Vega, chronicler of the DeSoto expedition, we learn that DeSoto sent two of his men, accompanied by Indian guides, forty leagues to the north seeking salt and precious metals. The men returned eleven days later laden with salt and copper, but spent and famished. They reported that the country, indicated by Houck to have been the highlands and headwaters of the St. Francis River, was sterile and thinly populated. Indians of that region informed them that the country farther north was almost uninhabited, and that the interior of the Ozarks was even more sterile. Leaving the land of the Cas- quins, DeSoto marched southwest into the bottoms of the Little River. Here the gentleman of Elvas, also a chronicler of the DeSoto expedition, tells us of a land “full of good meadows on the river,” from which only recently a vast forest has been removed. Quitting the swampy lowlands country in tireless quest of gold, DeSoto and his men marched northward toward Caligoa, believed by Houck and supported by Nuttal and Schoolcraft to have been the elevated highlands between the Black and St. Francis rivers, and at that time a land devoid of timber where herds of buffalo roamed. In the swampy lowlands and on the loessial hills bordering the Mississippi River, a more park-like aspect prevailed, as indicated by Garcilasso’s description of the country of the Casquins. Coronado, in 1541, is believed to have reached the southwestern portion of Missouri (Houck, 1908). The prairie aspect is indicated there by his notation of a rolling grassland, well watered with many rivers, and Osage Indians hunting among the vast herds of buffalo. After DeSoto and Coronado, a period of French explorations begins. Enter- ing the Mississippi Valley from the Great Lakes Region and seeking a water route to the Vermillion or Indian Ocean, the explorers travelled chiefly in bark canoes upon the rivers, and acquired little knowledge of the interiors. Father Membre, accompanying LaSalle down the Mississippi in 1683, has pointed out that the groves were so open and unobstructed one could ride through them on horseback. But even in those early years the appearance of Ozarkia was changing rapidly. Joutel, writing of the vicinity of Saline Creek in 1687, says: “The country was fall of hillocks, covered with Oak and Walnut trees, and an abundance of Plum trees...” (Houck, 1908). Already we see indications of the encroaching forest. How- ever, the forest was still conspicuously park-like, and Father Vivier in 1750 writes: Both banks of the Mississippi are berdered throughout the whole of its course by two trips of dense forests, the depth of which varies, more et fire to when the grass is dry; the fire spreads everywhere and destroys mos of the young trees. This does not happen in places nearer the river, because the land being lower and conse- quently more watery the grass remains green longer and less susceptible to the attack of fire. (Houck, 1908) | Уо. 38 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN As late as 1789 Forman was told by Captain Foucher, Spanish Commandant at New Madrid, that he could drive a coach-and-four through the open woods from New Madrid to St. Louis. It was inevitable that a resident population be attracted first to the eastern border of the Ozark region, and it was about 1704 that the first settlers made their home in the vicinity of Ste. Genevieve (Sauér, 1920). Here, conveniently located to Indian tribes and the fur trade, with an abundance of salt close at hand, and with the Arcadian abundance of the land, they had an easy exist- ence. Trading in furs and mining for lead occupied most of their attention, but of this period we have only scattering impressions of the aspect of Ozarkia. It was not until the cession of the Louisiana Territory to the United States in 1803 that any real consideration was paid to the resources of the area. Then, as in earlier years, an “open aspect” of the Ozarks predominated. The uplands in the interior were a rich grassland and supported numerous species of the "broad leaved” herbaceous plants. Numerous barrens were found jn the dolomitic and porphyritic hills, and there the vegetation was sparse. At this time trees were, for the most part, found only at wide intervals bordering the water courses, in deeper and richer soils of the uplands as isolated specimens, or occasionally in small park- like groves. In a sketch of the Louisiana territory, Major Amos Stoddard, who had «ccepted the land in the name of the United States in 1803, writes: “The highlands are seldom so thickly covered with wood as to prevent the growth of grass. They exhibit more an appearance of extensive meadows than of rude and gloomy forests.” In 1819 Schoolcraft, who toured the Ozark region on foot studying the mine country, was also impressed with the openness of the area, and observed: The general aspect of the country is sterile. .... Respecting the botanical character of the mineral soil, it may be further observed that although it yields but few forest trees, and they are not of a vigorous growth, yet a botanist might find his labors well rewarded by the profusion of shrubs and wildflowers which are everywhere found on the barrens, In regard to the region between Herculaneum and the Meramec River, which today supports notable timbered lands, Schoolcraft (1819) noted: Our road this day has lain across a sterile tract of country, consisting of a succession of hills of moderate elevation, covered chiefly by oaks and without underbrush. A tall, thick, and rank growth of wild grass, covers the w ole country, in which the oaks are standing interspersed, like fruit trees in some well cultivated orchard, and giving to the scenery the most novel, pleasing, and picturesque appearance. And this was a general condition, for he describes the country in the vicinity of Bourbon thus: Our route this day has been over barrens and prairies, with occasional forests of oak, the soil poor, and covered with grass, and very little underbrush. As evenin oac rima owed in a prairie near its source. Some good bottom lands are found on its nks, but the ad- joining hills are stony and barren, covered with little timber and high grass. (Schoolcraft, 1821). Bradbury, who traveled the Ozarks concurrently with Schoolcraft, wrote: The general character of this country is that of prairie, with scattered trees and inter- spersed clumps ..... e tract of country which contains the mines is very uneven, con- sisting of high narrow ridges, separated from each other deep craggy glens; the ridges have a peculiarly bald and arid appearance... . . (Bradbury, 1819). o 4 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 265 The exploring expeditions sent out after the Louisiana Purchase were bound for the mountains, prairies, and deserts of the Far West and they give little first-hand information of the Ozark country. However, Dr. Baldwin, accompanying the Long Expedition as Botanist on its western explorations in 1820, has left some impressions of the country in the vicinity of St. Louis. Collecting plants in the region of the mouth of the Meramec River, Dr. Baldwin found a typical prairie flora, and noted: “Тһе grassy plains to the west of St. Louis are ornamented with many beautifully flowering herbaceous plants . . . . The borders of this plain begin to be overrun with a humble growth of Black-jack and Witch Hazel." (Baldwin, 1823). Twenty years later we find the openness of the Ozarks still prevalent, as Davenport (1842) tells us: “There is no part of the globe, in a state of nature where greater extents of country can be traversed more éasily, and in any direéc- tion, by carriages of any description." Featherstonhaugh (1844) describes the . at the top of which region near Herculaneum after climbing a rugged hill: “. we found ourselves in extensive barrens containing straggling trees." During a period of settlement in the Ozark area, indicated by Sauer to be about 1820—1850, we find indications of the forest slowly and surely encroaching upon the prairie grasses dominant for so long a time. Settlement and the reduction of prairie fires favored the development of a forest flora as Swallow (1859) points out: trees found in ота: a still larger part is sparsely timbered . . . . : forming the beautiful oak openings. This stunted growth is not, however, due to the poverty of the soil, but to the fires which have annually overrun this country since the earliest dates of the Indian traditions. The slopes and some of the highlands are covered with heavy forests of nearly all the the bott B Jewett (1866), reporting on the agriculture of Jefferson County, writes: "he country is entirely covered with timber, except where improved . . . . the Oaks and Hickory predominating." Elsewhere woody growth was seen invading the prairies. Broadhead (1873) reports on Barton County: “The first signs of a growth of timber on the prairies is the appearance of small Persimmon bushes. They are common nearly everywhere”; and, “In southern Missouri open prairies are rare, but in their stead are occasional large tracts of barrens, or hilly districts „э covered with tall grass, on which are scattering stunted Oaks ....” Sargent, in 1884, notes: “А gratifying improvement in the condition of the forest in the parts of the state first settled has followed the enactment of a fence law pre- venting the general ranging of stock through the timber land." Toward the close of the nineteenth century we find growing concern over future timber supplies for extensive lumbering, and the widespread use of wooden ties for a rapidly expanding railroad system was making noticeable reductions upon a forest which was still somewhat open. However, Shepard (1898) reporting on the geology of Greene County writes: pply of timber in this region is decreasing; on the con- It is a mistaken idea that the su trary it is largely increasing. Attention was first called to this fact by Mr. J. W. Blankin- ship who has given a great deal of study to the flora of the region. Не learned from the testimony of old settlers that seventy years ago there was probably not one-half the timber in the country that there now is..... It has probably increased one-third within the last forty years. [VoL. 38 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN When Henry Shaw first visited the area which is now the Missouri Botanical Garden in 1820, he observed that "for a distance of two miles no trees were grow- ing .... The prairie was grown over with a tall, natural grass." (Shaw, 1880). Thus we see that the encroachment of forest upon the ancient prairies has been rapid indeed and that our forests as we know them today are of relatively recent origin. The barrens which were conspicuous in earlier times in their paucity of vegetation have remained as "islands" included within the forests, and are observed as botanical and geologic oddities. PRESENT FOREST RESOURCES King, Roberts, and Winters (1949) point out that "Missouri's forests are largely composed of immature stands" and that only 14 per cent of the "land bearing or capable of bearing commercial timber" supports stands of saw-timber value. In a summary they state that 43 per cent is in pole timber, 32 per cent in seedlings and saplings, and 12 per cent is poorly stocked. Thus, nearly 90 per cent of the commercial forest land is in immature stands. The saw-timber averages only 789 bd. ft. per acre; 5 per cent of the forest land supports 3,000 ft. per acre; and an additional 10 per cent, 2,000 ft. per acre. They also point out that one- third of the gross value is in cull trees and unmerchantable species. In addition, the net growth per acre is estimated at 38 bd. ft. in the Ozark region and reaches only 59 bd. ft. in the river-border region. They suggest that the commercial forest land might ultimately produce three times the present value of growth under good management. The Ozarks as a forest region is disregarded by Cheyney (1942), and the species of trees growing there are included among those of the forests of the Lake states and of the Atlantic and the Gulf Coastal Plains. Apparently, the bd. ft.-produc- tion per acre is so low that logging has not developed to the magnitude found in other areas. Even today, it is a region from which much of the output of the saw-mill goes into specialized products. Logging is reported to have reached its peak in Missouri about 1900, when 1,169 saw-mills produced three-quarter billion board feet of lumber, and approximately three million cross-ties. From such figures it is apparent that within the space of fifty years most of the merchantable timber in the state was logged. At present, the net growth of the growing stock exceeds the cutting drain by only 1.8 per cent annually. At this rate, it would take 225 years to support five times the present volume of saw-timber. The logging in- dustry revolves around a vast number of small mills operating seasonally. There is no evidence that the type of logging has changed greatly in the last fifty years. The backbone of the lumber industry at present is the 1,585 mills or 56 per cent of the total in the state, which produce 68 per cent of the total lumber. Many of these are under-powered and poorly equipped. Their annual cut ranges from 50,000 to 500,000 bd. ft. each, and there are only 45 mills which cut more than a million board feet annually. BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 267 The complete picture of the saw-milling operation in Missouri does not convey the impression that the Ozarks were once stocked with magnificent forests. Only fifty years have elapsed since saw-milling reached its peak, and less than 150 years since the first logging camps were established on the headwaters of the Gasconade River. Due to the transportation difficulties before the advent of the railroads and to an uncertain market, the earliest lumber operations largely served a local territory. These factors would have tended to conserve the resources in the éarly stages. Later, of course, improved transportation made the products of the Ozark saw-mills available throughout the Middle West. This would also have favored an increase in the annual cut which reached a peak 50 years ago and was not ap- proached even during the favorable years of 1940 to 1946. THE RED CEDAR At one time the scarcity of fencing material was considered a handicap to the settlement of the western country. Since such huge quantities of cedar posts are available and are so generally used in fencing today, we may wonder what position the Red Cedar (Juniperus virginiana) occupied in the early forests. Stoddard (1812) found, “it in plenty on the banks of the Mississippi and some other rivers above the mouth of the Illinois.” Brackenridge (1817) found it on the Meramec, St. Francis, Missouri, and the Mississippi—‘‘some very large islands on the Missouri are covered with this tree.” It is very likely, however, that the “cedar” of Brack- enridge was the Bald Cypress (Taxodium distichum) and not the Red Cedar (Juniperus virginiana). Bradbury (1819) found it along the Gasconade and Mis- souri River bluffs and mentions "that the tops are crowned with Cedars.” Swallow (1855) places its habitat as the “dry limestone bluffs.” Broadhead (1874), writing of Madison County, said that “Cedar Creek contained some of the finest groves in the state.” Не also mentions that great quantities have been cut for fence posts from the vicinity of Leatherwood Creek and Gray’s Mountain. Between the time when fencing material was scarce to the time of Broadhead’s observations, there are not many references to this tree. From this we might infer that the species was not nearly so conspicuous as it is today. It would have been impossible for it to have escaped the attention of Featherstonhaugh. Today Juniperus virginiana is one of the most common trees in the Ozarks, and on the northern edge it has taken over the role of pioneer and invader. It very often precedes the elm, persimmon, and sassafras trees which Swallow (1855) reported as “not common.” The Red Cedar is quite shade-tolerant and will persist in grass- land until a favorable opportunity for rapid growth arrives. It is one of the major competitors of old established trees. The seeds, voided by birds, germinate in all fence corners, open fields, and in the soil beneath the branches of any tree under which they happen to fall. This vigorous encroachment on fields and timber land by the Red Cedar appears to result from the control of fires. There are few species of trees more susceptible to fire injury than this tree. The annual burning of fields and woods which was [Vor. 38 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN so very common until just recently would have kept it within bounds. Beecher (1950) writes that the junipers (Red Cedars) were hardly known in Franklin County in 1900, and that when needed for Christmas decoration it was necessary to cross the Meramec River into Jefferson County to find suitable specimens. Today this species has gone north of the Missouri River and appears altogether too often on the grasslands in Callaway County. Unless burning is practiced, the Red Cedar has demonstrated that it can, in a short time, become the dominant tree in the entire Ozark region. SOILS AND EROSION Extraordinary efforts are being made throughout the country to prevent erosion and retain the topsoil. It is stated again and again that the early settlers “mined” the soil and then moved on to new land to repeat the process. However, travelers have left a description of the original Ozark soil, and it appears to have changed very little. Schoolcraft (1819), in his ‘Tour of the Mine Country,’ reported the “soil as a reddish coloured clay, stiff and hard, and full of fragments of flinty stone, quartz, and gravel: this extends to a depth of 10 to 20 feet, and is bottomed on limestone rock.” Bradbury (1819), traveling through the Missouri Territory, noted that “the stratum immediately below the vegetable soil is almost universally a very tenacious clay and extremely well calculated to form a material for bricks.” His prediction is borne out by the extensive clay-products industry which has developed through central and eastern Missouri. James (1823) described the Loutre and Grand prairies in Warren and Montgomery counties as having a soil which “was not very good; but mixed at the surface with so much vegetable mat- ter, accumulated by the successive growth and decomposition of the yearly prod- ucts, as to give it the aspect of fertility.” This observation is especially interesting, since these counties are still predominantly grasslands. Obviously there has never been a deep, rich topsoil capable of supporting an intensive kind of agriculture in the Ozarks. Nor, can any evidence be found to indicate that these soils ever approached typical forest soils—the podsolic soils of the north on which good forests are found. Rather these soils approach the Grood soils, the nut-structured Prairie-Forest soils (Wilde, 1946), or the “Lime Prairies” (Hilgard, 1906), described as a clay soil overlaying weathered limestone. Wilde points out that the transitional prairie soils present a picture of a struggle between grass and trees and that “this struggle does not end even when the forest canopy is closed over the prairie soil." He further states that the struggle "involves not only plants, but animals and lower organisms as well." Although it appears that the Ozark soil has always been much like the soil we know today, erosion may have increased in the past few years, if we recall Seay's (1866) observation on the absence of ditches in Crawford County. Perhaps the "newness" of erosion as a possible factor in Ozark land use is best illustrated by the numerous earth mounds thrown up by the Mound Builders. Houck (1908) was able to locate 28,000 mounds in Missouri. These earthworks, laboriously con- 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 269 structed by a prehistoric people, required the moving of millions of cubic yards of soil. To have succeeded in such a gigantic undertaking, each basket of soil carried to the top must have remained in place. The mounds could not have been built during a period of rapid erosion. The years during which they have been exposed to all weathering agencies is indicated by Houck, who states that the Indians found by the first white explorers did not recognize the mounds as their property nor did they use them or have any traditions concerning their origin. Stoddard (1812) says that they "have endured for centuries. The trees on their ramparts . . . . indicate an age of more than four hundred years." Pustmueller (1950), describing Monks Mound, states that “most of the trees have grown large in recent years, for at one time it was nearly bare." Only lately have the members of the St. Louis Archaelogical Society been concerned over the destructivé action of erosion, although the mounds have been a part of the Ozark landscape (Walker and Adams, 1946) for centuries. Now, after eight centuries, Monks Mound, rising over 100 feet above the adjoining country, is being slowly destroyed by gullies. FIRES Few travellers have written as vividly about Ozark fires as Featherstonhaugh (1844), who measured the progress of a fire at a camp eight miles from the Cur- rent River. He mentions that hunters used fire to drive game, and that in conse- quence of camp fires being left burning, "many thousands of acres were burnt over." In approaching Little Rock he expressed concern lest he be caught crossing a valley in which a fire was raging. Certainly, these fires have been common in the whole region from the earliest times. Swallow (1859), in his report following a railroad survey, describes central and southwestern Missouri as "in large part sparsely timbered, forming the beautiful oak-opemings. ‘This stunted growth is not, however, due to the poverty of the soil, but to the fires which have annually overrun this country since the earliest dates of the Indian traditions." Annual burning would have given the grasses and herbaceous plants every advantage by destroying the woody plants as well as the seedlings. Featherstonhaugh expressed the opinion "of Mr. Jefferson and others that all prairies have been produced by the Indian practice of firing annually, and thus destroying the grown timber as well as inferior plants." Fires are probably an extremely important factor in the maintenance of the openness of the country, and may have aided indirectly in the production of tre- mendous quantities of seeds and fruits. Later, as the country became populated and the settler had much more to lose, the practice of burning annually was frowned on. A fire advancing on a homestead, as described by Featherstonhaugh, could destroy the cabin, whatever stock of grain the settler had, and, by destroy- ing his fences, leave the fields open to the roving game and cattle. Even today the burning of woods is carried on only in those "backwood" areas where a kind of subsistence farming exists. [Vor. 38 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN Officials charged with fire control have long since learned that the native back- woodsman considers burning the woods his prerogative, and he cannot be convinced that he is doing harm. When apprehended he is able to offer only a few "lame" excuses; among them, he wishes "to increase the grass" or "destroy the insects." When we consider that these people, the direct descendants of the first white settlers, are almost inarticulate with strangers, and especially in a court of law, it may not seem so extraordinary that they are unable to convey their knowledge handed down through the years, of the openness of the timber and the grazing which was once available to cattle. Their insistence that they can "improve the grass" is not without foundation, since they alone have more than an academic connection with the past. There are no fires in the sections where farming has developed sufficient stability, and where there are only remote or no ancestral connections with the past. Swallow (1859) clearly indicates that a reduction in fire would result in an increase in timber. Sauer, illustrating the early opinion that fires checked tree growth, quotes the refusal by the United States of a grant of land to raise timber, on the ground that “it is only necessary to keep out the fires to cover the prairies with timber by the operations of nature.” Along the northern edge of the Ozarks, fire protection favors the woody plants to such an extent that fields are completely reseeded in just a few years. The American Elm (Ulmus americana), Red Cedar (Juniperus virginiana), Soft Maple (Acer saccharinum), and locally the persimmon (Diospyros virginiana), and sassafras (Sassafras albidum) are all invaders of abandoned fields which have been given fire protection. These species can main- tain themselves after gaining a foothold, but can be seriously injured and set back by fires at any time in their early years. If fire is allowed to enter such a field the reduction in growth rate follows a pattern so frequently described in present-day conservation literature. It is noteworthy that the light-seeded trees are the in- vaders of open ground, and not the oaks whose heavier seeds are probably planted by the smaller animals. If the oaks were the dominant trees of the area, and they are so described by both Swallow and Featherstonhaugh, they would spread rather slowly from the parent trees. This would indicate that the invasion rate a century ago was very much slower than it is today where fruiting specimens of light-seeded trees can be found in every ditch row, and where protection from fire favors their rapid spread, RAINFALL AND SALINES The early travelers spoke very highly of the healthful Missouri climate, but it is not possible to learn from the Weather Bureau records whether there has been any major change in the last century and a half. The records of the St. Louis office go back to 1837, but they do not show any pattern in the distribution of rainfall. For instance, in November, 1865, not a drop of rain fell, while the maxi- mum for November is 8.63 inches in 1847. December records show as little as .18 inches of rain and as much as 10.90 inches. July and August are usually 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 271 drier with a minimum of .25 inches for July and .07 inches for August. The maximum is 20.45 inches for August 1946 and 9.50 inches for July 1875. This wide variation in precipitation serves only to demonstrate the limited value of precipitation records in so far as plant growth is concerned, and all efforts to dis- cover trends toward an improvement or deterioration in climate have been unsuc- cessful. There must have been an appreciable increase in annual rainfall to have favored the explosive invasion of the grassland by the trees. Much less rainfall is needed to grow good grass than trees, and it is only necessary to observe the area 400 miles west of St. Louis to see a first-hand struggle between grass and trees for the avail- able moisture. Raup (1937) is convinced that the prairies at one time extended as far east as the Hudson Valley. If there has been a great increase in precipitation, it probably occurred just before the establishment of Spanish colonies in the Mississippi Valley. Of Mar- quette's description of the Mississippi in the vicinity of Grand Tower, Houck (1908) writes: I about Grand Tower, although it does not exactly describe the present condition. But 225 years will effect great changes, and that the river has widened in that period and that some rocky obstacles have been washed away is also certain. The seasons at that time appear to have become wetter. Since precipitation records are of no value in furnishing the needed information, it becomes necessary to rely on the early writers for proof. Featherstonhaugh (1844) wrote that "at one time voices could be heard across the Mississippi at St. Louis"—a far smaller stream than the Mississippi of today! One of the important occupations of all settlers and travelers was the procure- ment of salt. From the time DeSoto sent some of his party north for this purpose until almost two centuries later, when Moses Austin (1797) reported that the salt works, “when extended, might furnish all the upper settlements on the Mississippi," the evaporation of salt was a major project. In 1799, the works on Saline Creek in Ste. Genevieve County, produced 956 bushels of salt. Daniel Boone operated a salt lick which became sufficiently famous to lend its name to a major highway west—Boone's Lick Road. The fastidious Featherstonhaugh often refused the rough fare made available to travelers because much of the salt in use was gathered with a spade from a "lick," and both the salt and the soil was added to the cooking. Schoolcraft reports a salt lick near the present site of Fenton and two salt-manu- facturing operations nine miles from Herculaneum. He mentions another salt-lick in the Bellevue Valley, and he describes a buffalo lick called Bates' Lick covering about twelve acres and worn by game to a depth of ten or twelve feet. Although modern industry has supplanted the old iron kettle and evaporating pans and now supplies a very high-grade salt, the “licks” have vanished. It would seem that there are only two possible causes for their disappearance: (1) The rainfall may have increased so much that the brackish water is being diluted, and [Vor. 38 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN the water of today could not be used in the crude evaporating pans; (2) The growth of timber has greatly reduced the flow from springs, with the result that those which were used as sources of salt have now dried up or flow intermittently only during wet weather when the “salty” character would go undetected. The smaller springs, perhaps flowing seasonally, furnished the supply of salt for the “licks” used by both the game and the travelers. It is well known that the maximum amount of percolation occurs on a good sod where run-off is reduced to a minimum, while a very appreciable part of the rainfall striking the forest cover is evaporated. Edward Clark, in a personal com- munication, expressed the thought that the disappearance of the salt licks might be partly due to increased precipitation. Wolff (1948), working in the drier climate of Oklahoma, has shown that restoration of grass cover increased the flow of springs which had been checked by the advancing timber line. In his work with “cedar brakes” on the Edwards Plateau he was able to restore intermittent springs to full-time flowage by removing the cedars from the watershed and re- seeding to grass, It would seem that a climatic change tending toward increased precipitation has occurred in the Mississippi Valley and the near-by Ozark highlands. In a report on Crawford County, Seay (1866) states: The valleys. . . . . are frequently wide, scarcely ever rocky, covered with grass or hazel, with a deep, loose, sandy soil, and generally no definite channel to them. The water, if it accumulates rapidly, washes over the whole ground, but hardly ever so as to do any serious damage; and [in] an ordinary wet spell the water never finds its way out, but sinks. It would hardly seem necessary to point out that there are few if any valleys in Crawford County today that are not bisected by a drainage-way in the form of a fast eroding ditch. The salt “licks” have disappeared and the forest has in- vaded the grass land. The additional rainfall may have been all that was needed to support trees. It has been demonstrated (Wolff, 1948) that a tree cover re- duces percolation and reduces the flow of springs, while a grass cover, checking run-off, favors percolation and ground storage of water. SEED PRODUCTION Assuming that the Ozark highland experienced less rainfall two centuries ago than today, the leaching of soluble plant food would have been reduced to a minimum. This would have assisted in maintaining fertility at a fairly high level, which could be expected to stimulate seed production. Foresters have long been familiar with the "good" seed years which follow a severe drought. Kraus and Kraybill (1918) have shown that fruit production is correlated with the carbo- hydrate-nitrogen ratio. Klebs (1918) points out that during a hot, dry summer there is an excess production of carbohydrates in relation to the nutrient salts. This "in turn increases the probability of flower formation.” Heyward and Bar- nette (1934), investigating the effect of fires on the chemical composition of forest soils, stated: 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 273 higher percentages of replaceable calcium and total nitrogen. An indication was found that these burned soils were also characterized by larger quantities of organic matter as judged by loss on ignition. The soils subjected to frequent fires were found to be consistently less acid, and to have nd The above investigations would serve to show the seed production potential of the Ozark area. With the trees widely scattered, a reduced precipitation, a mini- mum of leaching—all factors favoring flower formation—the stage was set for the production of heavy seed crops. That seed production was very heavy is con- vincingly shown in the early records. Audubon (Wilson and Bonapart, 1831) observing a flight of Passenger Pigeons, estimated that they consumed more than 17 million bushels of grain daily. In 1700 Father Gravier (Houck, 1908) killed two bears in Scott County and saw fifty more cross the Mississippi River during one day. In the first statistical report of the Spanish settlement at St. Louis and Ste. Genevieve, Piernas (Houck, 1908), in 1772, listed 905 packs of furs shipped to New Orleans. These few cita- tions, among many, indicate that the game “carrying capacity” of the region was tremendous. It follows, then, that the production of food must have been in keeping with the needs of the game, also that a vast assortment of food was avail- able. Few of these game birds and animals can be found in the wooded Ozarks today. Not even the turkey, when given full protection, has staged a comeback. Even though the Ozark woodland is rated as "understocked," we find trees growing so close together that very few individuals produce a crop of seed. The plum thickets at the edge of the woodlot and some of the smaller shrubby plants may seed annually, but the heavy production of mast on which the pioneers fattened their hogs is a thing of the past. Only those oaks which are growing as isolated individuals bear seed with any degree of regularity. Lawn trees and scat- tered oaks near the Arboretum nursery mature seed quite regularly, some indi- viduals bearing heavy crops in alternate years. Many oaks in the forested areas have not produced seed in ten years, although they have been carefully watched since seed collections were needed to complete certain taxonomic work. Most of the trees submerged in the Ozark forest bear very infrequently, and some indi- viduals apparently never set fruit. This would appear to be in contrast to the behavior of these same species a century or more ago. In a region of reduced rainfall and no leaching, one would expect a favorable balance of carbohydrate to nitrogen. Annual burning would have made plant food readily available. Widely spaced trees would have produced a maximum crop of fruit. Tree fruits were not the only source of food. Many of the larger animals grazed as much as they browsed on the trees, or pawed the forest floor for acorns as do the deer today. This would suggest that the grassland contained a vast variety of species other than the Gramineae. Probably many of these were legumes stimulated by the same conditions which produced heavy "mast" yields from the scattered trees which were the early forests. [Vor. 38 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN BEES Native flower pollinators in the form of Bumble and Solitary bees of the genus Megachile, Nomia, Osmia, Andrena, and Bombus must have been present from the earliest times until destroyed by modern agriculture. However, Bradbury (1819) reports that the Honey Bee ( Apis mellifera) crossed the Mississippi River in 1797 and moved westward 600 miles in the succeeding fourteen years. Accord- ing to him, this "extraordinary progress in these parts is probably owing to a por- tion of the country being prairie, therefore yielding a succession of flowers during the whole summer which is not the case in forests." He points out that the Indian believed that when the Honey Bee appeared, the white man was not far behind. The flora of the prairie must have contained many kinds of plants other than the grasses to have favored the astonishing increase and progress of the bee. Grass alone would not have supported the bee, nor would a forested region have furnished more than suitable colony sites. The vast quantities of game, both animals and birds, indicate that there was a heavy production of fruits and seeds. Modern beekeeping depends very largely upon the legumes as a source of nectar. To havé produced the vast quantities of high protein foods required by the game, the country could not have been heavily forested, and the legumes must have made up a very appreciable part of the flora. HISTORIC FLORA Much has been written of the early vegetational aspects of the Ozark area, but there is only a scant record of specific plant material. However, the material available substantiates the vegetational aspect recorded of that time. For earliest records we must lean heavily upon Schoolcraft (1819, 1821) and Bradbury (1817) whose interests in natural history enabled them to record many valuable observa- tions. Following them, the botanists Nuttall, Baldwin, and James, attached to westbound exploring parties, have left valuable notes on this early flora. How- ever, as their route of travel lay along the Missouri River, they recorded little of the interior of the Ozarks. Tracy (1886) wrote the first flora of Missouri but admitted that, due to difficulties of travel and lack of sufficient field material, he had not included all the plants which probably grew in Missouri. Naturally the attentions of the first travelers were attracted to the arboreal flora, for in a region of extensive prairies the presence or absence of trees for fuel and lumber was a deciding factor in the settlement of the region. Stoddard (1812) pointed out that the city of St. Louis may never grow to the west because the ex- panse of prairie there did not provide enough wood for fuel and fencing. Al- though the composition of the forest flora has not changed to any great extent, change is noted in a greater number of trees. Early travelers found nut-bearing trees and hardwoods on the ridges and prairies where the heavy growth disfavored the light-seeded trees. In the lowlands, rich alluvium and inundation permitted the growth of light-seeded trees, Bradbury (1817) found: 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 275 On the summits of the ridges, the timber is generally red cedar; on the prairie, post oak, black-jack oak, black walnut and shell bark hickory. The alluvion of d rivers contains a great variety, of which the principal are cotton wood, sycamore, over-cup oak, nettle ^ hoop ash, honey locust, black 5 coffee tree, pecan, and many of 4% bien common i the states east of the Alleghanie Stoddard (1812) found the Sugar Maple abundant near Cape Girardeau and the people making large quantities of sugar from the trees. He observed persimmon, mulberry, chestnut oak, iron wood, and crabapple growing on the "high ground" and noted: "Common to both the high and low grounds are sugar trees in abun- dance, several kinds of walnuts, several kinds of hackberry, cherry, buckeye, black and honey locust, three kinds of elm, gum tree, lyn, sassafras, nine bark, spice and leatherwood, two kinds of ash, and the coffee tree." All the early settlers agreed that the Post Oak (Quercus stellata) was the most common tree scattered about the prairies. Schoolcraft (1819) writes of the Post Oak: "They are seldom found to grow higher than 30 feet, and 40 is the highest, seldom exceed a foot in diameter, and stand scattering.” The Black-Jack and Shingle Oak also scem to have been common trees of the uplands. Wild plums in thickets were common and prized for their fruit, as Bracken- ridge (1817) noted: с st is была fruits of Louisiana, the plum has been celebrated . . . .. ther re than the prairie plum, Prunus Chickasa dr today our Prunus angustifolia], which ШЕ covers tracts of ground of many acres in extent, and produc it so abundantly as to bend down to the earth po its weight. Frequent references to the crabapple and hawthorns are to be found. As late as 1908 Sargent found the Ozark region a rich field in which to work as he mono- graphed the genus Crataegus. These plants are not so conspicuous today and the recent invasion of the Red Cedar may be a factor in the failure of this genus so susceptible to the cedar rusts. Wild grapes were common everywhere, and the vines were far larger than any we know today. Swallow (1859) measured vines from 22 to 27 inches in circum- ference, and 55 to 120 feet in length. Bradbury (1817) found a vine, near the Meramec River, 37 inches in circumference. The rich prairie flora drew little comment from the earliest travelers. School- ec craft (1819), however, remarks: . a botanist might find his labours well rewarded by the profusion of shrubs pe wild flowers which are everywhere found on the barrens.” He was impressed with "sensitive brier,’ probably Schrankia Nuttalli, and mentions the peterswort, upland dock, and smartweed. Bradbury’s list of plants found in the lead mine country included 75 species (Bradbury, 1817), and Thomas Nuttall was introducing "interesting plants" to Fraser's Nursery out of the Upper Louisiana Territory in 1813. Baldwin’s (1823) notes are among the earliest specific references to the flora of the region, and he collected near the mouth of the Meramec: "Rudbeckia hirta, and К. purpurea, a small white flowering species of Houstonia, Galium tinctorium, Smyrnium aureum, a Phlox, a new species of Potentilla, a Conyza, the Trifolium reflexum, . . . . Campanula [Vor. 38 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN perfoliata, Diospyros virginiana, Rhus glabra, and many others." Оп the grassy plains west of St. Louis Dr. Baldwin observed "Aristolochia Sipho, Sypripedium spectabile, Lilium catesbeiana, Bartsia coccinea, Triosteum perfoliatum, Cistus canadensis, Clematis viorna, and Tradescantia virginica.” A notable feature of these early lists of plants is the consistent inclusion of the Leguminosae. Consid- ering the great abundance of wildlife dependent on plants of this family for food, and the rapidity with which the Honey Bee spread, it is believed that the legumes were an important constituent of the early prairie flora, as they are today in most successful range growths. Swampy lowlands of the southern Ozarks supported the impenetrable "brakes" of the giant cane (Arundinaria macros perma), popular retreat of game of all sorts. Featherstonhaugh found it to be one of the favored haunts of the black bear. The settlers generally thought that the tender new spring shoots of the cane formed a superior pasture for livestock. (Schoolcraft, 1821; Featherstonhaugh, 1844). The cane has a very limited distribution today. The joint-reed or rush (Equisetum byemale) was common in the more northern parts of the Ozarks. Bradbury (1817) observed of it: The rushes, P hyemale, were so I and tall that it was both ара and difficult to wal along, even at a very slow pace..... the rushes are... . . valuable, affording to the first sth er winter food for "his cattle for several years, after which they perish, being destroyed if fed on during the winter The tall, lush grasses of the prairies were інінің to all the travelers. School- craft described them as “often as tall as a man on a horse,” yet we find few records of their identity. Tracy (1886) wrote: "When the state was organized, our hills and prairies produced an abundant growth of buffalo grass which soon gave place to the blue joint which is, in turn, being driven out by the more valuable blue grass" It is significant that the buffalo grass is not listed in Tracy’s flora, though C. S. Jeffries (1888), in a letter to Dr. G. Hardeman of Gray Summit, states that it was common in the area about 1876. Broadhead (1873) points out that the blue grass "grew well after the prairies have been grazed down." GAME BIRDS AND ANIMALS The open and park-like aspect of the Ozark region was further evidenced by the great herds of buffalo, elk, deer, and the flocks of turkeys, grouse and quails which the area supported. Certainly this was an important factor in the early settlement of the region, for nowhere in the middle-western country was a human existence made easier than here in the abundance of game, wild fruits, and clear springs and streams. These habitants of a prairie or park-like region bear vivid testimony of its high virginal fertility and productivity—a striking contrast to the low-carrying capacity we experience today. That the wildlife was one of the dominant features of this region is indicated in some of the earliest writings. Garcilasso (Houck, 1908), recording the DeSoto expedition in 1541, tells us that in the Ozark country the buffalo were so num- erous that the Indians did not cultivate corn but rather lived upon game. Con- 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 277 sidering the crude hunting arms of that time, game must have been abundant. Father Membre, accompanying LaSalle, wrote of this area: “The fields are full of all kinds of game, wild cattle, stags, does, deer, bears, turkeys, partridges, parrots, quails, woodcock, wild pigeons, and ringdoves.” (Houck, 1908). In 1710 Father Vivier writes of the game, “the plains and forests contain wild cattle, which are found in herds; deer, elk, and bear . . . . Nowhere is game more abundant; from mid-October to the end of March the people live almost entirely on game, especially on wild ox and deer." (Houck, 1908). The "fields" of Father Membre, and the "plains" of Father Vivier clearly reflect the close relation of these vast animal numbers to the prairie habitat. The very important part that trade in furs played in the economics of the early colonies may further illustrate the ease with which pelts were secured. Piernas, in his first and third Detailed Statistical Reports for St. Louis and Ste. Genevieve, reported 914 packs of furs in 1772, and 2,888 packs in 1774, sent to the capital, then at New Orleans. (Houck, 1908). Nor was game pursued for furs alone; much of the meat was “salted down" in the centers of large canoes, then known as "pettyaugers," hollowed from large cottonwood trees and often 50 feet long. Much meat was supplied to New Orleans in this manner (Houck, 1908). Birds, too, were present in overwhelming abundance. Bossu traveled in the neighborhood of the St. Francis River in 1764 (Houck, 1908) and found it difficult to sleep at night on account of the noise made by the great numbers of ducks, geese, swans, and cranes in these marshy areas. Perhaps no wildlife spectacle has ever equalled the prodigious numbers of the passenger pigeon described by Bossu as often eclipsing the sun. Audubon reckoned the number of passenger pigeons in one flock to be 1,115,136,000 (Wilson and Bonapart, 1831). Featherstonhaugh (1844) remarks about the “.... : a sandbar in the Mississippi River near Herculaneum. Townsend (1839), travel- ` whirring and croaking of tens of thousands of cranes... ." on ing westward toward the Rocky Mountains in 1834, noticed on the prairies near St. Charles, ". . . . thousands of golden plovers; the ground was often literally covered with them for acres." With an apparently unending supply of game on every hand it was inevitable that much needless waste and slaughter of animals and birds took place. Indeed, the effects were noticed at an early date. Houck reports that the reduction of game in lower Louisiana was conspicuous as early as 1750. This condition became general as the area became more populated with settlers. James (1823) wrote of his journey across the prairie west of St. Louis in 1820: “The elk, the deer, and the bison, the indigenous inhabitants of these delightful meadows, had long since been driven away by the incursions of the white settlers...” Audubon (Audubon & Bachman, 1851), in his travels up the Missouri River in 1843, did not come upon the elk and buffalo until he was a great distance up the river. The settlers and the trader-trappers have always been pointed out as the cause of this great waste. Much of this opinion is doubtless justified. However, in the light of the [Vor. 38 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN current importance given to habitat in relation to animal populations, cannot similar principles be applied, in part, to the great historic reductions of wildlife? Vegetational aspect is the direct expression of geo-, climato-, and bio-relations, and conspicuous changes in floral aspect necessarily are the index of alteration of the habitat. We have noted already that the encroachment of white settlements upon the prairie lands was concurrently accompanied by the reduction of annual burning of the prairies which permitted the establishment of forest trees. A general reduction in fertility and productivity of principal game food sources has also been pointed out and together present such marked changes in the prairie habitat as to reduce seriously the carrying capacity of the area. Reduced in num- bers by a decadent habitat, the extinction of major game animals and birds by an ever-increasing settler population was inevitable. DISCUSSION An attempt to apply conservation measures of any sort to the Ozarks must take into account that the transition from grassland to trees has been achieved in less than a century and a half. We cannot hope to develop a Bunyanesque kind of forestry in the region so recently invaded by trees. The failure to recognize thé non-forest character of the area can only result, as Wilde (1946) indicates, in “a struggle that does not end even when the forest canopy is closed over the prairie soil.” It is possible that many of our epidemic tree diseases may stem directly from their growing on a prairie and not a forest soil. Forestry in the Ozarks can look forward to a multiplicity of problems until adequate time for trial-and-error selection points the way to dependable practices. The successful forestry project will be one which recognizes the fluid conditions prevailing, and is geared to profit by each and every advantage—no matter how unorthodox. Laws have been enacted which make the firing of timber a criminal action. Fire, perhaps more than any other factor, maintained the prairie and park-like aspect of the Ozarks. But to be effective, the fire season must be long and dry. An increase in precipitation would lessen the effectiveness of fire, and the ad- vancing tree line would suffer fewer disastrous burns. With the advent of white settlements, more and more portions of the area were protected from fire and these in turn furnished a haven for the advancing woodland. During every step of the invasion an increase in precipitation served to reduce the incidence and the damage which fire might cause. The native backwoodsman (derisively called "ridge runners," "brush apes," etc.), who sets the woods ablaze, is a direct descendant of the first white settler. Without holding a brief for the woods-burner, we may well ask some questions. Is it wise to attempt commercial forestry in an area that averages less than 800 bd. ft. per acre, and when only 10 per cent of the forest supports 2,000 bd. ft. per acre—especially when at least one-third of this forest is in cull trees and trees of non-commercial species? Perhaps a large part of the Ozarks should remain in grass. 1951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 279 The earth works of the Mound Builders have demonstrated the absence of erosion through as much as eight centuries. They were reported at first as almost without trees, then supporting trees, and now in serious danger from erosion. The rainfall which was adequate for grass has increased to the point required by trees. Conservation programs of any character, whether engineering works designed to control floods, efforts to increase game, or simply to farm on a more even keel, cannot ignore the fact that the Ozarks are becoming milder and wetter. No flood- control project can succeed if it does not recognize this climatic change. Major floods in the Mississippi River occur with increasing regularity. The river at St. Louis is much wider than a century ago, and it may be significant that the spring floods in 1950 occurred on a north-to-south line from the Dakotas to Oklahoma. The loss of life and the property damage were aggravated by the fact that this area, without a history of heavy precipitation, was completely unprepared for the heavy rainfall. An ecological study has shown clearly that the Ozark flora is immature, with the observed associations changing quite rapidly. In addition, the historical ac- counts of the vegetational character of the area bear out the contention that this is a young and vigorous flora not easily classified. Perhaps many species have dis- appeared, but one, the Red Cedar, has become more conspicuous by its aggressive pioneering in old fields. The encroaching forest sounded the death knell of the big game and the game birds. Regardless of hunting pressure, the habitat had deteriorated so rapidly that the herds of game and flocks of birds were doomed. Gun pressure admittedly has increased from the day of the first white settlement. The hunter appeared on the scene and added just the necessary weight to force the game from a habitat which was rapidly becoming untenable. Game management, whether aimed at the increase of deer or the improvement of turkey or quail range, will always be con- fronted with the present instability of the Ozark weather as expressed by its effect on the vegetational cover. The "Aux Arcs" Mountains, once the habitat of buffalo, elk, and bear, cannot now support turkey under complete protection. А com- pletely stocked forest of pole timber, which makes up 85 per cent of the forest area, is not a habitat for big game or game birds. From a mechanical standpoint, it isn't suitable even for buck deer; a buck with a trophy head will range іп the pole timber only under fear and compulsion. SUMMARY An imposing list of writings, dating from the earliest travelers, clearly shows the predominance of grass in the Ozark landscape. Featherstonhaugh traveled by horse and buggy from St. Louis to Hot Springs, Arkansas, and the border of Mexico (now Texas). Today such a trip would be impossible except over estab- lished highways. Insufficient time has elapsed to permit any plant association to be termed a climax. [Vor. 38 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN A study of forest resources indicates that today's lumber production, in a favorable market, is far less than the peak which was reached in 1900. Eighty- five per cent of the commercial forest land is classified as "immature," and there is little likelihood of an increase in production in the foreseeable future. Although very little information is available from Weather Bureau records, some pertinent observations are included which indicate that the Ozarks today enjoy a much milder and wetter climate. The disappearance of salt licks is deemed of considerable significance. Apparently all factors favoring heavy seed production were present—heavy seed production is not of common occurrence today. Sufficient information has been gathered to show the extremely important role of fire in the perpetuation of the grassland at the expense of the trees. As early as 1830 the United States Government recognized that the control of fires would clothe the prairies with trees. The Red Cedar (Juniperus virginiana) has shown itself to be a most aggressive invader of the grassland and run-down fields. The beginning of its explosive invasion coincides roughly with reduced burning as a result of an increased popu- lation. Bradbury's observations concerning the Honey Bee leaves little doubt of the open aspect of the country, with the legumes occupying an important position in the flora. The tremendous quantity of game, easily exceeding that of the domestic cattle and fowl of today, indicates the vast amount of food which was available. Typical podosolized soils, deemed to be characteristic of forest areas, are un- known in the Ozarks. Soil erosion, rated as the worst enemy of fertility in the Ozarks, has only recently become important. Ninety years ago the fields їп Crawford County were without gulleys. The Indian Mounds stood for centuries before showing signs of erosion. ACKNOWLEDGMENTS The authors wish to express their sincere appreciation to Mr. Charles Van Ravenswaay, Director, and the library staff, of the Missouri Historical Society, through whose efforts many important historical volumes were made available; to Mr. Trifon von Schrenk from whose library the authors have gained much valuable information; and to the library staff of the Missouri Botanical Garden for their kind and untiring assistance. 951] BEILMANN & BRENNER—FOREST INTRUSION IN THE OZARKS 281 BIBLIOGRAPHY Т n s and Bachman, Rev. John (1851). The viviparous quadrupeds of North America. New FM is (1797). A memorandum of Moses Austin's journey from the lead mines in the County of Wythe in the State of Virginia to f lead mines in the Province of Louisiana west of the Mississippi—1796—1797. Amer. Hist. Rev. 5:518—542. 1900. Baldwin, William (1823). Manuscript notes. In Edwin James’ Accou f [Long’s] expedition fr Pittsburgh to the Mountains performed in the years pr —1820. London (їп Thwaites Early Western Travels, Mad 4. 1905 А Beecher, Edward (1950). Personal communication od Beilmann, August P. (1943). А Whit Po over d on 11. 31:147-149 eilma old. Mo. Bot. Gard. шулен, H. M. (1817). 1. of Louisiana фе ка го гары, ау and historical ice. Baltimore. Bradbury, John (1817). A description of the minerals and е found at the lead mines in the ri Territory. Med. Rep. Orig. Essays & Intell. М. S. 3%135-138. 9). Travels in the Ee of America in the years 1809-11. London. (In Thwaites' Early W 1904.) nson, E. B. (1944). The geology of Missouri. Univ. Mo. Studies 19:270—336. кые С. C. (1874). Report of the geological survey of the State of Missouri, including field work of 1873-1874. Jefferson City. Cheyney, Ее kj (1942). American silvics and еее Clements, Е. E., and J. E. Weaver (1929). Ecology. r Davenport, Bishop (1842). А History and New ча. ог Geographical Dictionary of North merica and West Indies. Yor Featherstonhaugh, G. W. (1844). Excuision y dd the Slave States. New Forman, General (1789). Forman's journey down the Ohio and Mississippi. Ta s Editi Heyward, Frank, and R. M. Barnette (1934), Effect of frequent fires = p chemical ды at forest soils in the Longleaf Pine region. Fla. Agr. Exp. Sta. Bull. 2 i New Y Publ. Univ. Minn. Minneapolis. Hilgard, E. W. (1906 sid ог Houck, Louis и History of Missouri. 4 vols. Chic James, s don (1823) ue of [Long's] "apo ran Pittsburgh to the Rocky Mountains med in TE years 1819—1820. London. (In Thwaites Early Western Travels, Vol. 14. 19 Jeff ries, C 8. (1888). Letter to Dr. С. О. Har deman of Gray ез (In History of Franklin, efferson, Crawford, Gasconade Counties in Misso uri). Chica Jewett, W. S. (1866). Report on Jefferson County. Second rom Rept. Mo. State Bd. Agr. 27 { 275. King, D. B., Roberts, E. V., and or R. K. (1949). Forest resources and industries of Mis- souri. Mo. Agr. Exp. Sta, Bull. 452. Klebs, G. (1918). Uber die ТҮН von Sempervivum. Flora 111—112:128— Kraus, E. J., and Kraybill, H. R. (1918). Ман ion and reproduction with special ш to the tomato. Oregon Agr. Exp. Sta. Bull. ыыы T (1813). Fraser's Catal dies F5 catalogue of new and interesting n collected r Louisiana “ S pally on the River Зане, North America. London. (К үй Pittonia 2:114—119. s Palmer, Ernest J. (1922). The forest flora of the Ozark region. Jour. Arnold Arb. 2:2 Pustmueller, A. E. (1950). Cahokia brought to life. Publ. Greater St. Louis ee Ha Society. Raup, Hugh M. (1937). Recent changes of с res vegetation in southern New England and adjacent New York. Jour. Arn old Arb. 18:79-11 Sargent, C. S. (1884). Report on the jue pe North America. U. S. Dept. Int., 10th Census, Forestry. "EC 12 Crataegus in Missouri. Pt. 1. Ann. Rept. Mo. Bot. Gard. 19:35-126, 1908; : lbid.. 22:67-83. 191 Sauer, Carl O. (1920). The geography of the Ozark Highland of Missouri. Geog. Soc. Chicago, Scanlan, Edward H. (1950). [wena Trees С е К. (1819). view of the lead mines of Missouri. New York. ————, fon А a tour into the interior of Missouri and Pu in the years 1818 а Td Lon Seay, E. A. (1866). iG on Crawford County. Second Ann. Rept. Mo. State Bd. Agr. pp. Shaw, Henry (1880). Henry Shaw's idea of a botanical garden. Mo. Bot. Gard. Bull. 31:135-145. 1943. [Vor. 38, 1951] 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN Shepard, Edward M. zm" 8). A report on Greene County. Geol. Surv. Mo. 12:1 Steer, Henry B. (1948). Lumber production in the United States. U. 5; De. roe Misc. Publ. Steyermark, чеч А. M odia Studies of the vegetation of Missouri—I. Field Mus. Nat. Hist., . Ser. 9:349-4 bins rene 18127; Sketches, — and descriptive, of Louisiana. Philadelphia. olo ong the line of the southwestern [77 z Z2 = о 24 o: O = © ae? С) о S i m. 3 © et > oO © © с 5 et "t L4 = » (1855). С of the trees and shrubs af Va Ann. Rept. Mo. Geol. Surv. 2:221-226. Townsend, John К. (1839). Narrative of а pe across the Rocky Mountains to the Columbia River. Philadelphia. (In Thwaites' Early Western Travels, Vol. 21. 190 Tracy, S. M. (1886). Catalogue of the гв речен апі vascular cryptogamous plants of Mis- souri. Publ. Univ. Mo. Columbia -— Winslow M., and Adams, “көн McC. re ). Excavations in the Matthews Site, New adrid County, ME rans. Acad. Sci. St. Louis 31:71-12 946. Wilde, s. A. (1946). Forest soils and forest growth. Publ. Chron. Bot. Waltham, Mass, Wilson, A., and Bonapart, C. L. (1831). American DIM Edinburgh. 18 Wolff, Simon E. (1948). Evaluation of some we eedy Texas Junipers. Soil Conserv. 1111, THE CHANGING FOREST FLORA OF THE OZARKS AUGUST Р. BEILMANN* Амр LOUIS С. BRENNER** In this work the first time-lapse study on the forest flora of the northern Ozarks is reported. Brenner (1942) has shown the influence of the soils and the under- lying rock strata upon the forest flora within a portion of the area covered, but the present study includes additional plant habitats and may be more representa- tive. Recently the current associations were compared with those of an earlier map (Anderson, 1938), and the observed changes are here recorded. GEOLOGY OF THE AREA The area under discussion consists of about 650 acres in the Arboretum of the Missouri Botanical Garden at Gray Summit, Franklin County, R. 2 E., T. 43 N; Mo. It is a rectangular area lying within the rugged hills bordering the Meramec River, in the northern limits of the Ozark uplift, a region of dolomitic limestones weathered into a topography of moderate relief. A correlation of the logs from deep wells on the area has placed the underlying rocks in the Jefferson City, Cotter, and Powell formations of the Canadian Series, in the lower Ordovician Period REX RY ХЕХТ ХХОМХХХХАҚҚЫ 9555 00, x 59552 MAU ESO OOO М МН УУУ, 2245 ОООО SO "cece, P 2. ро [> оо ӨРГӨ PIS, А, Xo T 2% OPER ROKR CBS Хо EBM ХАС оо Экс бо у ЖЫ D ES оо V0.0 9, обо бето 99707097, DOO OX 4 О Уос) SEK «KX 2555442 SRA 2 950040600400; ообо 000060 5257 PRESSION Poe) D EIU TTA КЫ Ур о бы "er. d atr 9.0 9 9.0 99,9, ', FOR ELS о Miet (065525605099 900 9990900 Š 5, терер" 2? б, 55 т, e 7559; % 1275 69052100 MS 54 р o D 54 <) с? 955450 55555 5 Ф, 22 > Se. a e S Ё 5 2 Жы; 5% 9 ҰҒЫ сомто К, “т. Ш ктү VL 9 Ж | ШШ Dolomitic Limestone — Jefferson City Formation 9) ШШШ І i ШШ ШШ) DE А О | (|) = ІШ | РАМЕ е SCALE IN FEET 0 500 1000 INTOUR INTERVAL 20 FEET Ер Fig. 1. The rocks underlying the Forest Preserve of the Arboretum. Jefferson City Formation.—This formation forms the lowest outcrop to be found in the area, and is represented by a phase of dense gray-brown, oolitic, dolo- mitic limestone. This stratum outcrops at comparatively low elevations, and the * Manager, Missouri Botanical Garden Arboretum. ** Assistant Manager, Missouri Botanical Garden Arboretum. (283) [Vor. 38 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN greater portions of it are covered with talus debris, considered to have only a small influence on the forest cover. Cotter Formation.—The Cotter, lying unconformably upon the Jefferson City formation, is composed primarily of dolomitic limestones with sandstone phases of one to several feet in thickness, and thin shale lenses. The base of this formation is a gray-brown sandstone, 37 feet in thickness, and of interest because several perennial springs have their origin in this stratum. The sandstone is succeeded by a phase of dense, massive, slightly dolomitic limestone, 34 feet in thickness. The remaining 91 feet is composed of succeeding strata of medium-grained, loosely cemented sandstones; dense, oolitic, slightly dolomitic limestones; and relatively thin strata of pure dolomite, locally known as "cotton rock." In chemical content (a double carbonate of magnesium and calcium), in high porosity, and in degree of fractibility, the cotton rock may have a more direct influence upon the flora than any other stratum. The occurrence and distribution of glade floras have been shown to be directly related to these cotton-rock strata (Erickson, Brenner, Wraight, 1942). | Powell Formation.—The Powell is represented by somewhat thinly bedded strata of limestone, chert, dolomite, and sandstone. There are but few outcrop- pings, since the formation is almost completely overlain by the mantle of Union Silt Loam. The predominance of cotton rocks which permit the rapid percolation of water creates a "dry" appearance in the forest during most of the growing season. The ease with which water moves through these rocks is shown by the numerous springs, and by the great amount of seepage water to be found in the valleys in wet seasons. This rapid percolation of ground water does not encourage the deep penetration of tree roots into the rocks in search of water. The influence of the rocks is further reflected in the slow growth of the trees. The dolomitic rocks, because of their slow rate of decomposition (Hilgard, 1910; Lutz and Chandler, 1947), offer only small quantities of the minerals necessary for the growth of plants. Feeding roots seem to be concentrated primarily in the shallow soil mantle, where the decomposing organic matter offers greater quantities of nutrients. The effect of these two factors is evidenced in the glades which are devoid of forest trees. SOILS OF THE AREA The rocks, high relief, and the various exposures and angles of slopes, together with the pattern of land use, present a complexity of factors reflected in an equally complex "soi! picture." Based on local peculiarities, the soils are divisible into two major groups: residual and alluvial. Because of the "upland" nature of the area, the alluvial soils are of relatively little importance and will be treated only briefly (fig. 2). Union Silt Loam.—The Union Silt Loam represents the most important soil gtoup by reason of its greater areal extent and its ability to support the better timber types at considerably faster growth rates. The soil is a brown to grayish- 1951] BEILMANN & BRENNER-——THE CHANGING OZARK FORESTS 285 KOS KK SOOO DOLLS IE OOO OD? 2 2 550005606 555556560550 дай 925555 Хо дө ооу од дуд дуруу ео % ОСО ОСОО ООО ООО ОООО ЧОО Қ уу! 52 косо К 252% RIERREN SIREN УУРУ, ооо ооо OP? RISSIES AOSA R ооо ооо ООА КУ 2% EN NOS SQ Qo S eR RENSES ALA 946080) (605667055 Ау д Marte Rest A SSA ооо OA ta POKER о o SEXO xx 6 09 S ОКО роо OR ORK I OPO SOO PR CK ROCO SA 600568620, 604469; sat ооо Убу, ON RPI I POOR red Pr RRA (860060048 ооо оо ооо AR Ө So o КУСУ со 92 9746) 292460 94 ten lat 500000 ооо 5604060 ROE RRO Tre RN 2529005550500660%6605669 e OO AS Pn TASA ROS Z уу 55559562009 Мосо SCREEN RKO NING оа 028406 (02797054 /0 98756 Aq 50 994 4064004 зетот д , 2%; ооо) PREP N ROP ROO 55050620050 7 ^ о 055255 255, <<, (5о POPP ө 90426; 2520060 2% м — Рф ороо IARR э, Ps “5% 2) LR 225252; ERR RRC р TRAE ee РҚА BN SNS: ЗУ d LB SSR OU P RK 5 A 505055 5052,2 ©: LPS RTA MOE RR EK Roop SR 59 > et ^ >> 575662255695 ORES 229 max) 2% GOEL LLL LP (52 PIER = j (405 ча 44 555 <) 495% с^ 2% ©? s So LAL) > od N eel) - S S à 56; "^ [X K ode <% > A 7 aS. 255 BEERS PORN <> %% CA M E. d SCALE IN FEET 500 1000 AL 20 FEET 0 CONTOUR INTERV: Б Union Silt Loam EE tiated Ё ? Soil ШШ Talus Soit Glade Soil Huntington Silt Loam Elk Silt Loam Fig. 2. Principal soil types of the Forest Preserve of the Arboretum. brown, mellow silt loam, 6—8 inches deep, grading into a light brown or yellow- ish-brown, friable, silty clay of crumb-like structure, with а nut-like subsoil at greater depths. The fine silty nature of this soil and its great angle of slope make it an easy prey to erosion, which, however, can be controled by planned forest management. ridges. On some steeper slopes and small localized areas the soil is of a cherty nature and should be classed as Clarksville Stony Loam. However, because of the localized nature of this cherty soil and the history of land use of the area, the authors believe that it represents subsoil of the Union Silt Loam, exposed through The Union Silt Loam in its typical form is apparent as the mantle covering the erosion induced by early land use. Rougb Stony Land.—Extensive areas of the Arboretum have a soil generally referred to in soil surveys as Rough Stony Land (Vanatta and Lewis, 1911). The forest types found here have suggested the advisability of further division of this soil into four subtypes based upon site and mode of origin: Undifferentiated Forest Soil, Talus Soil, Glade Soil, and Alluvial Soil. The group is represented by soils which, because of direction and steepness of slope, have been retarded in devélop- ment. Shallow, rocky, and of high organic content, they are entirely without distinguishable horizons. The Undifferentiated Forest Soil is found chiefly on south and western hill- sides, and may be from a fraction of an inch to several inches in depth. The soil mantle lies immediately upon the bed rock, has a high organic content, and con- tains an abundance of rock fragments. Exposure of these slopes to sere late sum- [Vor. 38 286 ^ ANNALS OF THE MISSOURI BOTANICAL GARDEN mer winds has discouraged herbaceous vegetation. This, together with the high angle of slope, has caused excessive erosion, resulting in an almost permanent juvenile state. The forest type listed as "Transitional" is almost entirely confined to this soil. The Talus Soil is practically identical with the Undifferentiated Forest Soil in composition, but differs in its greater depth and in the constant movement of the mass through settling. These differences, along with a relatively low water table during drought, has discouraged the growth of good timber trees. The Glade Soil represents a distinct type of shallow soil mantle derived from dolomitic rocks. The soil has a high organic content, but the underlying highly porous cotton rock of the Cotter Formation causes it to become extremely dry during the summer and extended periods of drought. Trees have found it difficult to invade glade areas and a flora typical of rocky prairies is found there. Alluvial Soils.—As previously stated, the alluvial soils are of minor importance in this survey. Small areas of the “first bottom,” assigned to the Huntington Silt Loam, are included as well as the Elk Silt Loam which forms the soil of the ancient Meramec River terraces. Although those soils are not of sufficient extent to in- fluence the development of a distinct forest type, they are interesting additions to the complex picture of the soils of the area. LAND USE It has not been possible to determine exactly what use has been made of this land since 1850, when the first titles were granted. Certain portions show evi- dence of early timber cutting, at least for local use, and certainly quantities of firewood were also cut. Some hilltops have been clear-cut with the expectation of farming. All the area must have been heavily pastured by open-range cattle. In more recent years more intense utilization followed the fencing of pastures and the change from open-range conditions. It is not possible to furnish dates at which Certain areas were pastured or cut over. On the whole, the history has been one of intense usage. Very few mature, sound trees can be found, and it is assumed that many have been cut. Usually, following such operations the land was grazed and quite frequently it was burned. The extent and purpose of the timber cutting depended largely upon the topography of the area. The valleys and the more gently sloping hillsides were within the capacity of the equipment available to the early settlers. Опе area is reported to have been a sawmill site, and the condition of the adjacent timber indi- cates that it may have been clear-cut (indicated on fig. 3 as “Logged more than fifty years ago”). A number of older trees escaped the axe, probably because of their low value. Some of the high hill land may have been clear-cut "Less than fifty years ago.” The areas so designated are without old trees, and much of the younger growth appears to have originated from stump sprouts. One fairly large valley was cultivated prior to 1925, when the area was acquired by the Missouri Botanical Garden, and a similar area is to be found near one of the ridge tops. 1951 BEILMANN & BRENNER—THE CHANGING OZARK FORESTS 287 > > e <2 Ясы SR t Ж; = 555 OD 252506, Ж 52555555 се SRR <<) б 22 Ж > ғ: X 5255, SEEKS 5050 2 > 55 20; 55 <> >; >< ^ 4 Pg Х : ағ. . қ se] р ғ. „угу Bo б, A = Sy 225 d < 9056 hey ^ Sis peewee Ср 26924 С А] LE] d > 97929,0 RPL -a e > Q H ооо LAK See % Sepe Оо; <> CLAPP M o vee oS V 27 5559529 Ке D ~ POR P 9 = Hj = OA SP <> -€-— — = | SCALE IN FEET о зоо 1000 CONTOUR INTERVAL 20 FEET Cut ov er more than 50 grs aqo =] E Logged 9 Logged less than 50yrs. 900 ET] Cultivated prior fo 1924 Fig. 3. History of land use of the Forest Preserve of the Arboretum. The presence or absence of old trees may prove misleading when used as a measure of logging practices and land use. A few scattered old trees may indicate either that “selective” logging had been practiced—selective in the sense that the early settler cut certain trees for specific purposes—or merely that tremendous changes have occurred in the arboreal flora of the Ozarks in the last hundred years. Apparently, the Ozark forest today is more heavily stocked than it was a century ago, and the few relic trees to be found are those left from what was a park-like prairie (Schoolcraft, 1819) which seems to have been the outstanding character- istic of the region before 1850. DEVELOPMENT OF TREE ASSOCIES DURING A TEN YEAR PERIOD Anderson’s forest-tree classification (1938) has been modified to conform to present usage (fig. 4). The tree species involved in each association were covered by Brenner (1942). The extent and location of the present forest tree associations are shown in fig. 5 Post Oak-Black Jack Oak (Quercus stellata - Q. marilandica) Association.— This association is limited to isolated stations on the ridges and to the Union Silt Loam, and appears to be highly unstable. The decadent nature of this association and the rapidity with which it has been absorbed by the more vigorous Oak- Hickory association indicate the speed with which the hardwood forests are en- croaching upon the early prairie associations. White Oak-Sugar Maple (Quercus alba- Acer saccharum) Association.—Dis- tribution of the Sugar Maple appears to be controlled by the presence of old fruit- ing trees, and definite edaphic relations of this member of the association are not [Vor. 38 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2)j ^ <5) 27 2 SN — c-——-- O _ ESO —. 7 заа м ooa „Фм Ф, OKO „Жэ, о ASS > 2.5, 54 (A EAEAN: ро ооо МАМ у 4 oA ERINNERN A АСУ CC OK OC d POPS NOE So ре уг SCALE IN FEET 0 E 1000 : А CONTOUR INTERVAL 20 FEET (ZA Post Oak / Black Jack Oak Brush E Osk Z Hickory Ped Cedar / Chinquapin Oek ШІ White Oak Coppice Glace EC White Oak / Sugar Maple Transitional Forest Fig. 4. Principal tree associations in 1938, Forest Preserve of the Arboretum. (From an unpublished field survey in 1938 by Edgar Anderson.) S SS MOS АО N AO EUN ў Y aa LPP AN ТП. [| Sh ў M x к rH hy ^ 5 А А В аыл 52 iY 7 KAN 4 <. HA А Хә * е, 47 f SCALE IN FEET o 500 1/000 CONTOUR INTERVAL 20 FEET Post Oak / Black Jack Oak Red Cedar / Chinquapin Oak Oak / Hickor БОЙ White Ook / Sugar Maple Glade < Transitional Forest Fig. 5. Principal tree associations in 1948, Forest Preserve of the Arboretum. 1951] BEILMANN & BRENNER—THE CHANGING OZARK FORESTS 289 shown. The rapid and widespread regeneration of the Sugar Maple in the deep moist valleys is apparently conditioned by the more favorable microclimate of those sites. In 1938 this association included several valleys where some selective cutting had been done in the last fifty years, but no clear-cutting for at least a hundred years. Such cutting and the resultant openings in the forest favored the rapid growth of the Sugar Maple. At present, however, it is losing its position as a codominant and may eventually disappear from this association. Above the valley floor the changes have been more rapid. The Sugar Maple in the White Oak- Sugar Maple Association of 1938 has been succeeded by hickory, and these areas are now classed as belonging to the Oak-Hickory Association. Oak-Hickory (Quercus alba, Q. borealis, О. stellata-Carya ovalis, C. ovata, C. tomentosa, C. Buckleyi) Association—The high cdmmercial value of these species, their rapid regeneration, and good annual increment make this forest association particularly important to foresters. The marked adaptibility of these trees permits almost cosmopolitan distribution, for little preference is shown toward rock, soil, or direction of slope. However, the association has been most consist- ently mapped on areas of the Union Silt Loam where greater depth of soil and concurrent fertility have favored rapid growth and excellent regeneration. Perhaps no other tree association has had more continual exploitation. There is ample evi- dence that the trees have long been regarded as a source of firewood, rough lum- ber, and railroad ties, and such great use was made of them that in 1938 the Oak- Hickory Association was one of the least extensive in the area. However, it 1s becoming increasingly vigorous under the present system of land management and at present about 60 per cent of the area falls in this class. The Red Oak (Quercus borealis) and the important Shumard Red Oak (Q. Sbumardi) are invading these associations and are becoming codominant with the White Oak. White Oak Coppice (Quercus alba).—Yhe White Oak Coppice is shown оп the early map to cover about 20 per cent of the area. Nearly all of this is now included in the Oak-Hickory Association, with the Red Oak an important tree of the understory. Red Cedar-Chinquapin Oak (Juniperus virginiana-Quercus Mublenbergii) As- sociation.—This association is unique in being the only опе of the area which is closely correlated with the underlying rock strata. Оп the accompanying maps (figs. 1, 4, 5) the Red Cedar-Chinquapin Oak Association is seen immediately bordering the cotton rock both above and below its extensive outcroppings on the western slopes. In this narrow belt, often a mere rod or two broad, the influence of the cotton rock still excludes other forest types. This association is shown to be more extensive on the map for 1938, since at that time accelerated erosion in- duced by earlier logging and pasturing only served to cause a more widespread effect of the conditions induced by the cotton rock. Rapid growth of the Red Cedar and its remarkable powers of regeneration have caused it to be exploited [Vor. 38 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN greatly as a source of fence posts and telephone poles within the past fifty years. Today the association has a more limited distribution, and the Oak-Hickory and White Oak-Sugar Maple associations have apparently invaded much of its outlying portions. Glade Association.—Here the inherent properties of the cotton rock in con- junction with a western exposure of slope have combined to form a critical zone for the growth of trees. Trees are conspicuously absent, an herbaceous, prairie-like flora prevailing. Transitional Forest.—Because of its heterogenous composition and lack of aged specimens, the Transitional Forest is considered to be in a state of flux. Intensive exploitation of these sites through pasturing and logging has created a long- enduring juvenile state where the soil mantle is poorly defined. Comparison of the maps of early and present tree associations (figs. 4, 5), and the soil map (fig. 2) shows this association to be closely correlated with the Undifferentiated Forest Soil. Recent survey of the area shows the invasion of the Transitional Forest by Oak-Hickory and White Oak-Sugar Maple associations as the site matures and a deeper soil mantle protected by a layer of forest litter is developed. Brusb.—Several small areas classed as “Brush” іп 1938 are now included in the White Oak-Sugar Maple Association. In 1938 those areas contained much sassafras (Sassafras albidum), elm (Ulmus sp.), and sumac (Rhus copallina), and were considered as not likely to persist. They are now stocked with some of the best young timber. SUMMARY Because studies in the natural succession of forest trees in the Ozark region of Missouri have been based largely upon site-to-site comparisons it is felt that this work, founded upon actual observations of the same area over a ten-year period, will give a better understanding of forest tree succession in the northern Ozarks region. Considerable attention has been given to both the rocks and the soils, since these have been observed to play important roles in the distribution of certain tree associations which may obtain only in similar areas. The history of land use of the area is reviewed, since the activities of man, in his pursuit of a livelihood in forestry and agriculture, may have altered the natural succession of forest trees. This time-lapse study indicates a rapid invasion of valuable timber species such as Red Oak, White Oak, and White Hickory. The Black Jack Oak-Post Oak, and White Oak-Sugar Maple associations have been invaded by these Vigorous species and will be dominated by them, indicating a future invaluable source of high- grade timber species in this area. 1951] BEILMANN & BRENNER—THE CHANGING OZARK FORESTS 291 LITERATURE CITED Баи үг (1938). Field survey map showing the аха tree associations of the Forest of the Arboretum, Missouri Botanical Garden 38. Unpublishe fidis ү С. (1942). The en Pay m variables of p "Missouri Botanical Garden Wild- flower Reservation at iro Summit. . Mo. Bot. Gard. 29:103-13 Erickson, Ralph O., L . Brenner, ind gen Wraight (1942). A YS glades of east- central Missouri. Thid. 2 89—101. 19 : ils. p. 42. New s York. Lutz, Н. J., and R. F. Chandler (1947). Forest Soils. pp. 56—57, 358. New York. Schoolcraft, Н. R. (1819). A view of the lead mines of Missoati. New York Steyermark, J. A. (1940). Tos of the vegetation of Missouri. Part I, Natural rw associa- tions ES succession in the Ozarks of Missouri. Field Mus. Nat. Hist. Bot. Ser. 9:349—375. dried E. and H. G. Lewis ча 911). Soil - of Franklin: County, Missouri. U. S. Dept. Feat ad Field Operations Rept. 13:1603—16 HOW KENTUCKY BLUEGRASS GROWS* ALFRED GORDON ETTER** INTRODUCTION Human cultures of widely varying origin have been treading sods of Poa pratensis for thousands of years, long before it became known as Kentucky blue- grass. Indifference to the plant's personality has been bred by this long familiarity, with the result that while bluegrass may often be thought of in terms of sod, turf, herbage, forage, or just plain grass, it is only rarely considered in terms of an indi- vidual plant. It is the purpose of this paper to describe in simple words what a bluegrass plant looks like and how it grows. In general, only that part of the organization and development will be considered which can be seen with the naked eye or low-power binocular microscope. The importance of understanding the structure of a plant has become especially apparent in crop plants such as maize where solutions to practical problems— picking, detasseling, cultivating, spraying, cutting, and prevention of lodging— depend for their success on a knowledge of the manner in which the plant is put together. Anderson (1949) has stressed the need for this knowledge in maize and has raised many pertinent questions concerning the maize plant of the future. Bonnett (1935, 1940), in describing the development of the flowering head in barley and maize, has suggested the practical applications of such studies. In spite of the ever-increasing number of grass farms and suburban homes with ample lawns, little effort has so far been made to relate developmental and morpho- logical studies of forage and turf grasses to practical problems. Knowledge of the individual plant is fundamental to proper analysis of variation in any species, and consequently to the proper appreciation of its genetic possibilities; successful prediction and selection of strains to fit specific environmental conditions depend on such knowledge. Lawn and pasture species are constantly being subjected to management practices of various sorts. An understanding of the basic pattern of development of the plants involved is essential to any attempt to discern the effects of such treatment. Proper statistical planning and analysis of forage production and plot tests must necessarily be based on an appreciation of the variables in- volved in the normal growth processes of the individual plant. The widely varying and inconsistent results which are frequently obtained in pasture and turf research are probably partly due to the lack of information available on grass mechanics. * The work reported in this paper was a on while the author held a special fellowship under a bequest of the late Dr. Malvern B. Clopton. The field work was done at Me khill Farm, Clarksville, Mo., Dr. Clopton's former yd өрі the laboratory vM the Henry Shaw School of Botany of Washington University. It seemed a peculiarly ec gne an she ds Ес it concerned the bluegrass pastures at Brookhill which Dr. Clopton knew and lov Та Gratefu ^ bcn d P edgment is made to Chancellor Compton and to members 5 the Bro okhill С mittee n ou unusual opportunity and to the staff of the Doane Акпы] rwr ice n friendly eration in the field. This investigation was submitted to the Board of Graduate Studies in а fulfillment of the requirements for the degree of бе a "phi losophy at Washington University. ** New Mexico Military Institute, Roswell, N. M. (293) (Vor. 38 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN METHODS Problems Involved.—Shortage of data devoted to growth of sod grasses may reflect the subtle difficulties involved in the measurement and description of their growth. Analysis of a wild population of grass is complicated by the fact that plants of different strains and different ages are all mixed up and growing together. Study of these grasses in the laboratory, greenhouse, or even the nursery plot, involves the introduction of artificial conditions which have been shown in several studies (Musgrave, 1940; Ahlgren, et al, 1945) to influence strongly the growth characteristics of the plants. Direct measurement of growing parts can scarcely be accomplished without damaging or removing protecting organs or soil, and consequent introduction of complicating factors. Indirect measurement of growth by examination of a number of plants in a developmental series is based on a number of assumptions which may be difficult to prove. (See van de Sande- Bakhuyzen, 1937). Preliminary Studies.—Observations on bluegrass growth and morphology were made over a period of two years to provide a general understanding of the problem and the plant. Sods were occasionally dug up, washed, and studied and diagram- med in detail. Special effort was made to discern seasonal trends. From these preliminary investigations sufficient familiarity with the plant was acquired to allow plot tests or laboratory experiments to be made on a small scale. Although the anticipated results were not always obtained, every new condition to which bluegrass was subjected yielded information of some sort. For example, in the course of measuring leaf growth on bluegrass grown in nutrient solutions, it be- came apparent that all plants of the same origin under the same conditions produced leaves at practically the same rate. Plants could thus be roughly arranged as to time of origin by referring to the number of leaves they bore. Specific periods in the life of a plant could be identified, since each plant that lived through the winter had one leaf shorter than all the rest of the leaves; also certain types of shoots were limited to the cool fall and winter period, while summer buds fre- quently remained dormant. It soon became apparent that while the chief concern of the study was to shed light on problems of pasture and lawn, such places were not the best for studying bluegrass behaviour, for an excessive number of complicating factors were in- volved. Constant cutting maintains the plant in a vegetative condition and seasonal activity is not clearly defined. It was found that the most favorable place to study basic behaviour was in a relatively unproductive old meadow where mowing, treading, and fertilizing were minimum complications and where the grass could be expected to respond only to the strongest seasonal stimuli. Measurement of Meadow Populations—A project was then begun on which most of the conclusions in the following work are based. It was decided to measure periodically the last completely exserted leaf on ten plants with respect to blade length, sheath length, and blade width (measured 5 mm. above the ligule). Other data which later came to be recorded in detail included the lengths of the inter- 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 295 nodes and panicles on the flowering shoot, the number of leaves which were green, the nodes which bore roots, and the number of tillers and rhizomes. Measurements were begun in early March of 1949 and extended to the following March. During the spring measurements were made weekly, while during the summer, fall, and winter they were made bi-weekly or monthly. Dissection of plants under a binoc- шаг microscope was necessary to provide much of this data. То obtain the necessary rhizome material, blocks of sod were subjected to water under moderate pressure until the soil was washed away. In order to gain preliminary information on variation with habitat, identical studies were made of grass in four situations: the unmowed meadow; a closely grazed pasture; a meter-square plot of meadow grass which had been burned over in mid-February; and a similar-sized plot which had been cleared of all surface vegetation with a sharp hoe while the ground was frozen in February. Although information on all of these plots is not included in the present study, an under- standing of the basic trends was facilitated by these measurements and observations. Supplementary Observations.—Data obtained from the above measurements were supplemented in several ways. Exclosures! were placed in pastures being grazed at various times of the year to determine the repercussions of different types of grazing management. Also, a special box was made which had glass sides that could be darkened by inserting a tight-fitting panel This box was filled with dirt and several plants of bluegrass were placed against the glass. By removing the panel their underground activity could be watched throughout the year. Information has also been derived from examination of grass plants growing under unique environmental conditions. Plants that grew in soil on which urine or manure had fallen, or that sent rhizomes into mole tunnels, or which had been grazed closely by mice in their winter pathways, or had been covered with alluvium, or grew in deep shade or in very dry or wet localities, have all con- tributed small bits of important data. Laboratory Investigations—In general, laboratory experiments were designed with the idea of finding out how rhizomes grow and what makes them turn up. Some data on leaf growth were also obtained in connection with this work. Blue- grass rhizomes were grown in two ways: separated from the parent plant and grown in water or various solutions in test tubes or Petri dishes; or left attached to the plant and allowed to develop in a natural manner except that they were enclosed in glass tubes so that they could be measured, protected, and identified. Some experiments were made in which these tubes were filled with nitrogen, watér, or air. To obtain as much control over bluegrass plants as possible, some were grown in bottles containing nutrient solution. These were kept under constant light and temperature conditions. In nature, the base of the bluegrass plant, together with its roots and rhizomes, is underground and thus invisible. In searching for some n exclosure is a part of an experimental area which is fenced to eliminate the influence of certain animals, in this case livestock. [Vor. 38 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN manner of exposing these underground parts without too greatly disrupting their natural relationships it was discovered that ordinary "Even-Flow" baby bottles had peculiar possibilities. The nutrient solution could be placed in the bottle and the plant could be held in place above by inserting it through a slightly enlarged hole in the nipple (pl. 2, fig. 1). This would allow the basal part of the plant to be within the nipple and thus protected from desiccation. At the same time, by coating the nipple with opaque paint the inside was kept quite dark, thus simulating soil conditions. The roots could extend into the nutrient solution below and the leaves into the light above. Rhizome growth was provided for by making five perforations at the base of the nipple, just above the plastic cap, and in these inserting short pieces of 14-inch glass tubing which were connected at the other end to a -inch glass tube, about 7 inches long, with a 1-hole stopper in the end (pl. 2, fig. 2). As a bud on the basal part of the grass plant was seen to be developing into a rhizome it could be guided into the small glass tube and then would grow out into the larger tube where various experiments and observations could be conveniently made. A per- foration was made down through the cap and nipple so that an air-supplying tube could be inserted into the bottle. Saturated air under low pressure was bubbled into the nutrient solution through a fine glass capillary. This air, in escaping, flowed around the roots and base of the plant and out through the rhizome tubes and served to keep the water aerated and the atmosphere in the tubes moist at all times. A light-tight box high enough to accommodate the bottles was then con- structed and in the top surface were cut twelve holes spaced on a 9-inch grid (pl. 2, fig. 3). These holes were just large enough to receive the bottle caps. One side of the box was removable so that bottles full of nutrient solution could be inserted into the openings from underneath. The height was made just suffi- cient so that when the cap was screwed on the bottle, the rhizome tubes inserted into the nipple would rest on the top surface of the box (pl. 2, figs. 2—4). New rhizomes which had been turned up long enough to form two good leaves and several roots were obtained from a single clone of bluegrass and were inserted into the nipples in such a way that their roots contacted the solution while the bud region of the plant was in the air opposite the openings for rhizomes. In order to keep light away from any rhizomes which might develop, it was neces- sary to construct a false top of removable panels (pl. 2, figs. 4—5). Ву loosening а thumb screw these panels could be removed so that access could be had to the rhizome tubes and bottle caps. A fluorescent fixture containing six 15-watt tubes was placed at a height of 30" above the plants (pl. 2, fig. 3). This battery of bottles was kept first at a constant temperature averaging 70^, and then later at 40°. Variation was not ordinarily more than 5° in either direc- tion. Day length was fourteen hours during most of the experiment but was reduced to twelve hours during the last three weeks of the 40? temperature. The nutrient solution used was that given in Meyer and Anderson (1941), and was changed every two weeks. А 5 per cent iron solution, made up of equal 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 297 quantities of FeCl, and tartaric acid, was added weekly. No difficulty was had in getting plants to grow, though the light intensity was apparently insuffi- cient for normal growth, and the leaves were rather lax. The low light intensity at the 70? temperature was not conducive to elongation of buds or roots, while at 40? roots grew well but budding was largely restricted to aerial shoots instead of rhizomes. There was no opportunity for further experiments under more optimum conditions for rhizome production. LITERATURE The importance of Kentucky bluegrass as a lawn and pasture grass in northern Europe (especially in Germany and Sweden) and in the northeastern part of the United States has led to many studies on various aspects of the plant. Most of this research has been concerned with seed and forage yields and chemical content under different treatments and seasons. A great deal of intensive genetic work (see Brittingham, 1943) and histological work (see Nielsen, 1946) has been done in connection with its apomictic method of reproduction. Considerable juu on bluegrass variation (Smith et al., 1946), taxonomy (Mecenovié, 1939), a germination (Gassner, 1930) has also been published. Somewhat less information is available on the life history of the plant, its seasonal activities, and its manner of growth. Nishimura (1923), in a compara- tive study of the morphology and development of Poa pratensis, Phleum pratensis (timothy), and Setaria italica (millet), anticipated by many years subsequent work on the species. Close observation and detailed drawings of the seedlings, the basal region of the plant and of the developing inflorescences were made. Some discussion of germination and roots and the first mention of polyembryony were also included. Wieland (1926), in connection with an intensive study of variation, neces- sarily devoted some attention to the gross morphology of the grass, but was not concerned with how it got that way nor how it changed with the season. W. L. Brown (1940), also dealing with variation in the species, traced the seasonal development of two strains from March to October. Musgrave (1940), in an unpublished thesis on the life history of bluegrass, discussed various aspects of germination, leaf, shoot, rhizome, and inflorescence development, also differences between plants grown as spaced seedlings and in un- disturbed sod. Evans (1949) discussed his investigations on bluegrass over a long period of years, including various aspects of its life history and of the influence of environmental factors on its growth. He summarizes information on rhizomes, roots, inflorescences, photoperiodic relationships, and annual cycles of growth, development and reproduction. Work on the physiological aspects of bluegrass growth has been done by Brown (1939, 1943), Harrison (1934), Peterson (1946), Peterson and Loomis (1949), Phillips (1943), Darrow (1939), Naylor (1939), and others. Practically no information exists on the developmental anatomy of bluegrass. [Vor. 38 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN The best work of this sort has been done by Sharman on maize (1942) and on Agropyron repens (1945 and 1947). He has given close attention to the relation- ship between the progress of anatomical and morphological events, has emphasized the existence of a common fundamental ground plan in grasses, and has shown how slight changes can produce plants which appear wholly different. McCall (1934) has discussed the developmental anatomy and homologies of various struc- tures in wheat with special reference to the seedling. Evans and Grover (1940) reviewed much of the literature on the develop- mental morphology of grass and discussed terminology and general concepts of growth with special reference to the inflorescence in a number of species. Bonnett (1935, '36, "37, '40) and Noguchi (1929) have described and illustrated various aspects of the development of the inflorescences of the small grains and maize. Prat (1934, 735) examined the anatomy and morphology of the grass culm and discussed certain theoretical aspects of its growth. He expressed various correla- tions, and discussed maturation of tissues. Certain aspects of the developmental physiology of grass growth have been discussed by such workers as De Ropp (1946), Weintraub and Price (1947), and van de Sande-Bakhuyzen (1937). Important life histories of grass plants include those of Percival (1921) on wheat, Weatherwax (1923) on maize, and Evans (1927) on timothy. BLUEGRASS IN SIMPLEST TERMS The Sod.—When we look down at the leaves of grass they merge into a con- fusion of green (pl. 3, fig. 6). What does an individual bluegrass plant really look like? If we take up a 6-inch square piece of sod the question remains un- answered, for, even after the dirt has been washed away, there is left a confusion of roots and strong wiry underground stems that criss-cross in every direction (pl. 3, fig. 7). These stems seem to connect the various aerial leafy shoots in a haphazard manner. Occasionally they are seen to branch, and some may be white at the tip, showing that they are still pursuing their subterranean destinations. These wiry interwoven stems are called rhizomes, and they are one of the reasons why sod can be rolled up in a ball. They are also one of the reasons why studying an individual bluegrass plant is difficult. A Single Plant.—If we start with a seedling the problem is simple, but even- tually we shall have to deal with the vegetatively spread individual. For our purpose only the gross features of the seedling are significant. Pull a six-weeks- old plant from the soil. It is about as simple as bluegrass ever gets (text-fig. 1A). It has a beginning and an end; it has shed its nourishing grain and become auton- omous. It will have four or five narrow spreading leaves which are bundled together toward the base for about one-half of their length. At the bottom of the bundle are roots. If we are used to plants which have stems we shall be dis- appointed when we pull the leaves apart, for they enclose only the bases of other leaves. At the center of the bundle, at its base, with its tip barely visible, is a 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 299 GROWING POINT RIMORDIAL NUBBIN (CROWN LEAVES ROOTS. / PRIMARY ROO GN aie ROOT Fig. 1. А six-weeks-old bluegrass plant—about as simple as bluegrass ever gets: A, seedling, Х2; B, crown, X 13. young, rather yellowish green leaf which is in the process of growing out into the light. Its tip is darker green but it is not yet bent outward as are the other leaves. Within this elongating leaf is a younger one only a few millimeters in length and inside of it is the minute youngest leaf which protects the nested primordia of two or three future leaves. At the center of this nest is the small bulbous growing point of undifferentiated cells, from which the primordia arise. These structures are shown in detail in text-fig. 1B. The only stem the plant possesses is the nubbin of solid material to which the leaves, the growing point, and the roots attach. This nubbin, or crown, as it will be called, may be only a millimeter or two in length. Under magnification it is found to be a systematically organized structure with bilateral symmetry, holding alternate leaves and enclosing at the base of all but the lowest one an axillary bud (text-fig. 1B). These leaves and buds seem to arise from successive horizontal partitions on the crown which are termed the nodes. Separating the nodes are the internodes, sections of stem which, while sometimes much elongated, here are scarcely distinguishable. From each of the lower two or three nodes a pair of roots arises, one root on each side of the axillary bud. These are the adventitious roots which play a major role in feeding and anchoring the plant. In all young seedlings there is also a primary root, the first root of the newly germinated seed, which persists for a short time at the basal tip. In many species of plants this primary seedling root develops [Vor. 38 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN into a taproot by which such plants are permanently fixed to one spot. But blue- grass is not tied down for a lifetime of sitting on top of a taproot; instead, its liberated stem can develop roots wherever an axillary bud exists, and can thus gradually shift position and still maintain itself. y N A i үү FROM A i RHIZOME INTRAVAGINAL Ч SHOOT INTRAVAGINAL : SHOOT ж-е” RHIZOME I \ ~ к, “у= “у - 77 N RHIZOME Fig. 2. А spring seedling in its first fall: A, the plant, X 4; B, crown, X about 5. The Appearance of Brancbes.—N/hen we examine the spring seedling late in the fall of the same year (text-fig. 2A and B), we find that the story has begun to be a little more complicated. The apical growing point is no longer the only active опе. Some of the axillary buds, of which there may be 10 or 12 by now, have begun to grow. Thus bluegrass, like most plants, produces branches. A bud has a choice of two destinies. Which it follows depends upon where it is situated. In the fall those buds which are enclosed at the base of the most recent two or three green leaves will send up vertical leafy shoots within the secrecy of the re- taining leaves of the parent plant. A few of the lower, older buds borne in the axils of leaves which are dead will develop, or may already have developed, into non-leafy horizontal shoots. The upward-growing branches, or fillers, are some- times called “intravaginal” shoots because they arise within the confines of the sheathing base of living leaves. The horizontal shoots are "extravaginal" and pro- trude through the base of the dead sheathing leaf if it is still attached. These are the rbizomes which, as we have seen, grow underground. Sooner or later they turn up to form new plants quite similar to the parent. The differences between tillers and rhizomes are easily seen. Опе risés in the axil of a green, living leaf, one in the axil of a dead or dying leaf. The most 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 301 distinguishing feature of the rhizome is that it has short colorless scale-like leaves and has a distinct stem made up of a series of fairly long internodes, while the upright intravaginal shoot, or tiller, bears normal green leaves and has a very much condensed stem or crown, just as did the parent plant which it resembles in almost every respect. The end result of this system of branching is that the slender static seedling becomes a mobile plant, capable both of increasing its number of leafy shoots and of extending itself over considerable area, sometimes as much as two square meters in two years time under ideal conditions. (Kannenberg and Wrede, 1934). The intravaginal shoots thicken the plant by making available more new leaf-producing places, often as many as 10, sometimes up to 50 in a season when there is very active secondary and tertiary branching. Rhizomes spread the plant horizontally, and for all practical purposes once a rhizome has turned up and developed a new leafy shoot it can be considered a new individual, even though genetically it is identical with the parent plant. It may remain attached to the parent for a con- siderable period, though apparently it derives practically no nourishment from it. An interesting demonstration of this independence was provided by an experi- ment primarily designed for other purposes (pl. 4, fig. 8). A flower pot was filled to capacity with soil. A bluegrass plant with long, well-developed roots was placed in the pot so that the roots were covered by the soil but the crown area was left above the surface. A shallow tray with a hole in the center was placed on top of the pot, the leaves of the grass plant being arranged so that they stuck up through the hole. The tray was then filled with vermiculite, a micaceous cutting medium with practically no nutrient value. The buds on the crown were thus surrounded by mica. The soil was kept moist by setting the pot in a shallow dish of water. The mica was watered separately. After the plant became estab- lished, rhizomes began to grow out into the vermiculite and turn up along the edges of the tray. It was consistently noticed that after turning up, these new plants grew only very weakly and their leaves were very short, yellowish, and narrow. The parent plant, in the meantime, was deep green, long-leaved, and healthy. Eventually most of the offspring in the tray succumbed. There was no evidence that the vigor of the parent could be conveyed to the second vegetative generation through the rhizome, once it had turned up. Flowers Develop.—Some time during winter, after intravaginal shoots have been initiated, metabolic changes occasioned by cool weather and short days in- duce "mature" growing points to begin a process of proliferation which eventually ends up in the appearance of a flowering head. This is a slow sequence of events which begins with a slight elongation of the growing point, the suppression of leaf development, and formation of many small buds which don't elongate (pl. 4, fig. 9). This is the secret winter beginning of the seed-bearing “panicle” which becomes so prominent a feature of bluegrass during the month of May. With the advent of longer days and warmer weather multiplication of panicle buds ceases, flower development begins at the end of each bud, and elongation of the panicle [Vor. 38 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN branches takes place. When this is finished, the whole panicle is thrust out of its retaining leaves atop a flowering stalk (pl. 4, fig. 10). This stalk is called a “culm” and is part of the main axis which has become greatly elongated. Finally the plant has achieved a conspicuous stem, but only at the price of its own future, since the panicle terminates the growth of the crown’s growing point. A plant in its second spring will thus be slightly more complicated than its fall progenitor, because, besides having attached to it various satellite plants in the form of tillers and rhizomes, it will be developing a flowering shoot. The mature individual can thus be said to show three types of shoots: the fall rhizome, the tiller, and a central culm, topped with a panicle (text-fig. 3). All these derive from the basal nubbin of telescoped internodes, the crown. After the inflorescence dies down, the rhizomes and tillers assume the role of perpetuating the plant. Each of these pur- sues an essentially independent course from then on, and eventually are separated by the decay of the parent. THE STRUCTURAL UNITS Leaves.—The leaves of the bluegrass plant have been described as narrow and spreading, but bundled together toward the base. It is useful to distinguish the green spreading parts as the blades (or laminae) from the lower colorless sheaths which in clasping each other form the bundle. The sheaths are slightly compressed tubes, while the blades are flat or folded straps. Successive sheaths enclose each other, but the blades bend outward at their junction with the sheath (pl. 5, fig. 11). A characteristic structure called the ligule, a membranous, valve-like flap of tissue, is found at this union and looks like a brief continuation of the sheath beyond the base of the blade (pl. 5, fig. 12). There are no obvious anatomical differences between sheath and leaf, although, according to Sharman (1942), there is some consolidation of the many small (basipetal) strands of the blade at the ligule and consequently fewer strands in the sheath. In general, however, he minimizes the differences between the two parts. Prat (1935), on the other hand, states that the epidermis of the sheath is more differentiated than that of the blade, and that there is a striking similarity between the epidermis of the sheath and of the internode. In any case, it is ap- parent that the blade is the primary photosynthetic organ and is always green, while usually the sheath is green only in the vicinity of the veins, if at all. Just how such a structure as a leaf arises and becomes differentiated into sheath and blade has been well told in detail by Sharman (1945). Although his work was done on Agropyron repens, his description of leaf development agrees with that of bluegrass given by Musgrave (1940). Mention has been made of the nest of leaf primordia and the central growing point which lie hidden and protected within enclosing leaves at the apex of the bluegrass crown. The activ- ities of this critical region must necessarily be understood if we are to interpret correctly the growth of the plant as a whole. The bulbous, translucent growing point (pl. 5, fig. 13) is the source of life, 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 303 BLADE LEAF peer FLOWERING SHOOT. y n E \ E RHZOME | Ж A mature bluegrass plant showing the three types of shoots which develop Fig. 3. from the crown. [Vor. 38 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN the source of new undifferentiated cells from which the plant is fashioned. The first evidence of differentiation occurs when the leaf primordium begins to develop. This starts out as a number of localized cell divisions on one side of the growing point. These divisions produce a small protuberance which soon spreads out laterally, eventually becoming a crescent and finally a complete collar around the growing point. Two of these primordial leaves can be seen in pl. 5, fig. 13. Since the first formed midsection of the collar is able to maintain its advantage over the wings, the collar has an eccentric appearance with the midsection somewhat higher. This incipient leaf grows primarily by cell division at its base and to some extent by a short-lived meristematic band along its tip. As growth continues the collar soon forms a sort of loose-fitting, parka-like hood or cowl over the apex (pl. 5, fig. 14). Through the face opening of this parka the growing point can be seen, and by this time it will have given rise to one or two new crescents. This opening is at first quite minute, .2 mm. or less, just large enough to allow the collar to slip over the growing point. At about this time the leaf is supplied with its primary vascular bundle, and the thin fold of epidermis, the ligule, first makes its appearance at the junction of leaf and crown, thus indicating that sheath tissue is beginning to develop. Growth of the leaf blade (the part above the ligule) soon becomes very rapid. The face opening of the hood is so stretched out as the blade elongates that it becomes unrecognizable in the mature leaf. Between the time that active elon- gation begins and the leaf grows to maturity only two or three new leaf primordia develop.” Since the hood stage itself represents about the third primordium in a series it is evident that under ordinary vegetative conditions there are about five or six leaves in various stages of development from the newest collar to the newly matured leaf. "These leaves would measure roughly about .1 mm. (crescent), .2 mm. (collar), 1 mm. (hood), 4 mm. (blade beginning to elongate), 50 mm. (sheath beginning to elongate), and 120 mm. (mature). The last two measure- ments are given only for easy comparison and might be much longer. The first three leaves are waiting in line while the fourth and fifth elongate. The blade and sheath of the sixth have fully matured. De Ropp (1946), in an analysis of the growth of the first blade and sheath of a rye seedling divided the very young leaf, while it was still in the embryo, into equal sections by marking them with a mixture of lamp black and vaseline. He then made daily measurements of each section for a period of a week. His data showed very well how sheath-growth was delayed until leaf-growth had begun to slow down. It was apparent, however, that the change from leaf-growth to sheath- growth was not an abrupt one, and the growth curve of the entire leaf remained relatively unaffected. This growth was primarily one of cell elongation, since no new cell division was noted after the third day. De Ropp concludes that ultimate leaf length depends on the activity of the basal meristem during the first three "The interval between the appearance of successive leaf primordia is commonly termed a “plastochrone.” 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 305 days. This period of cell division undoubtedly varies greatly with species, environ- ment, and season. De Ropp has also called attention to the difference between the manner of elongation of the blade and the sheath. In the sheath it was found that each marked segment contributed about equally to the mature organ. In the blade, however, the basal fourth of the marked primordium contributed three- fourths of the total growth, while the tip segment scarcely enlarged. This may in part demonstrate why the tip of grass leaves is frequently slightly boat-shaped. Apparently, the primordial hood elongates so slightly that its original structure is more or less preserved. A mature vegetative leaf is thus the product of two growth processes, the elongation of blades and elongation of sheaths. The two processes do not happen to the same leaf at the same time. What actually happens is that an elongating blade (leaf No. 4, for example,) and the sheath of the enclosing older leaf (leaf No. 5) elongate simultaneously and at the same rate so that there is no necessity for the tender young blade point to push up alone through the close confines of the sheath bundle. Instead, the two grow up together. Upon dissection of leafy shoots, the tips of young leaves will generally be found to be just about even with the ligule of the preceding leaf. This is additional testimony to the essentially similar growth rate of blade and sheath. But the sheath does not grow as long as the enclosed blade, and consequently as the sheath matures and its growth stops, the blade within continues its growth alone out into the light. It often happens that the tender new blade is not strong enough to start pushing out of the orifice of the ligule, and the tip of the leaf becomes cramped and crinkled into one or more small folds. These folds remain impressed in the leaf after exsertion and are com- monly seen on rank-growing bluegrass. Under vegetative conditions no blade-sheath pair begins to elongate until the previous blade has stopped growing. This has some significant repercussions which will be discussed in detail later. Phytomers.—Bluegrass is not built of leaves alone. A leaf is only the most obvious part of a more fundamental structural unit called a рруѓотет, or plant segment. This unit consists of a piece of stem with a leaf on the upper end and a potential bud or shoot at the lower end. If the leaf is on the left side of the stem at the top, then the bud will be on the right side at the base. There are complementary sections with the leaf on the right and the bud on the left. The two types of sections alternate with each other and produce the characteristic bilateral symmetry of the grass plant (text-fig. 4A and B). Another result of this alternation is that the bud of the segment above rests within the concavity of the leaf below, just above the point where the leaf is attached. This concavity is called the axil of the leaf and the bud is therefore commonly called an axillary bud. Though this name is a convenient one, it encourages the idea that a leaf mer, a term according to Evans and Grover (1940) used as early as 1879 by Asa Gray Phy’ А to designate a structure which, when produced in a series, makes a plant of a higher grade. The term was also employed and described by Weatherwax (1923). [Vor. 38 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN and its axillary bud are closely related, when actually from an anatomical and physiological standpoint the association between a bud and the leaf above, on the opposite side, is much closer. Sharman (1942) has provided anatomical evi- dence supporting this. Fig. 4. А and В, M ve phytomers or grass-building units, A from a vegetative crown, B, a rhizome; C, the hood leaf which protects the growing point. , Where two phytomers join together a node is formed. The phytomer axis thus becomes an internode which may be long or almost undiscernible. Each phytomer, in addition to its leaf, internode, and bud, is theoretically capable of producing roots at its lower end, usually one on each side of the bud Phytomers have their beginning at the growing point. А leaf primordium is the first visible evidence that a new phytomer is forming. The small hood leaf which has been described is not just a shell of tissue tacked on the apex but actually can be visualized as a sort of kettle-shaped structure with a bottom made up of undifferentiated cells produced by the growing point. Inside this kettle new phytomers continue to begin their differentiation as cells in the meristematic point (text-fig. 4C). The course of differentiation and maturation in a phytomer, as Sharman (1942) has shown, is a diagonal one which begins with the development of the leaf at one side at the top, then proceeds to the internode, and passes on to the bud on the opposite side at the base. When roots are present they are the last organs to be formed (text-fig. 5). This course of phytomer development has its basis in the maturation of various anatomical units, especially the vascular system. By the time one phytomer has matured, several others will have begun their dif- 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 307 BLADE SHEATH INTERNODE Ac cdd . 5. Diagram illustrating the wave “ elongation and maturation passin d je blade, sheath, and associated internode in Zea Mays. So lid black indicates the more actively dividing ubi. poem Sharman, 194 2) ferentiation. The exact number depends on the stage of maturity of the plant. Plants ready for flower initiation may have as many as eight or nine segments in various stages of elaboration, while, as we have seen, a vegetative plant may have only five or six readily discernible. The diagonal course of development is outwardly apparent when we measure the elongation of the different phytomer elements, as van de Sande-Bakhuyzen (1937) has done for wheat. It has been said that the blade elongates first, and is followed by its sheath. In text-fig. 6 the whole leaf has been measured as a unit; consequently successive leaves overlap (or would overlap if the growth curve were extended to its origin). This overlap represents that period of time when a sheath and the new blade which it encloses are growing up together. Where there is no overlap, the leaf has grown out of the sheath and continued its growth alone. The internode may or may not elongate, depending on where it is located on the plant. If it does elongate, it does so only after the leaf has matured. Under vegetative conditions, as in the rhizome, and in the early reproductive phase as shown in text-fig. 6, only one internode grows at once. Under the influence of the inflorescence, however, there is a tendency for some overlap of elongation in those internodes just below the flower head. In bluegrass the last three internodes elongate more or less simultaneously. An overlap has also been recorded by Shar- man for maize (1942) and Agropyron (1947), by Prat for rye (1935), and is suggested in the last two nodes of wheat (text-fig. 6). A grass plant thus develops by virtue of its ability to produce new phytomers and to elaborate the plastic organs situated thereon. The remarkable plasticity and independence of these units make possible the varied appearance of different grasses and the different parts of the same grass. This will become apparent with the following description of the three main shoot types in bluegrass. [Vor. 38 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN LENGTH IN CM. 440 T T T T T T T T T T T T T 440 1400 400r Leaves and звон sheaths 4360 (Ist to 5th) 320r 1320 а 4280 — 2m] э] ат 240} 1240 200r 1200 Ist /¥ leor 1160 120г ИТ” 1120 [e] + 80 ы ) Ix u/ Internodes 40 I "i (Тоу) 440 “ , ^ 1 1 1 ug қ, 1 fy EL.) 1 1 1 1 о ~ l (5 17 19 2i 23 25 27 2 l 5 7 9 M 3 DAYS OLD 9 э 33 35 е wth in length of leaves = нды s M whea (From H. L. van de Sande- Bakhuyzen, "Studies Whea n Under Constant Con й м " Food Res. Inst. Misc. Pub. No. 8, p. dg pe University, Calif., March 1937). THE RHIZOME What Does It Look Like?—A rhizome is by definition an underground stem. In bluegrass this means from a few millimeters to several centimeters underground. It may grow horizontally below the surface for 6—12 inches; it may merely swing out and up in a short inverted arch; or occasionally it may turn up abruptly with- out making any lateral growth, іп which case it should probably be considered merely an extra-vaginal shoot. Plate 6, fig. 15, shows rhizomes of these three types on a single plant. The fact that the rhizome is a stem imposes upon it certain re- sponsibilities. It cannot pursue its underground path by simply adding cells at its tip as does a root. Its growing point is obliged to produce phytomers, which in turn produce leaves, internodes, nodes, buds, and roots. The rhizome thus becomes a jointed structure with various appendages rising from the vicinity of its nodes. Internally the separate phytomers are bound together by a complicated vascular system The most prominent parts of the rhizome are the elongate white internodes, which, in series, make up the axis. These average 1 or 2 cm. in length, are solid, and are frequently rather flattened in cross-section. The tip of this underground stem is like a sharp white awl and is well suited for penetrating soil (pl. 6, fig. 16). This tip is actually a much-reduced leaf which has no blade. It is essentially a closed tube although a minute pore is present at the tip. It serves well as a sort of caisson within which the subterranean growing point can work to lay down building material and elaborate new structures. When a new rhizome leaf has been fashioned inside, it penetrates the caisson, which then splits and dries up. On 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 309 the older parts of the rhizome these leaves hang loosely at each node and become weak brown remnants scarcely recognizable as foliar structures. They are given the special name of cataphylls, or scale leaves. These cataphylls are arranged on opposite sides at successive nodes, and partially conceal the internode of the next phytomer (text-fig. 7). In the axil of most cataphylls is an axillary bud, though occasionally no such structure can be found, especially at the first few nodes. A pair of roots usually appears at each node, one root on either side of the bud. The first two or three internodes nearest to the parent plant are quite short, less than 1 or 2 mm. The next few are intermediate in length, while subsequent ones may range from 1 to 30 mm., apparently depending largely on the amount of water available to the plant. The sequence of cataphyll lengths is essentially similar to that of internodes. At the junction of the main shoot and the rhizome is found the first scale leaf, if such it may be called (text-fig. 8). It has only two prominent veins and arises from the upper side of the rhizome with its back to the crown. It is minute, less than 1 mm. in length, and slightly heart-shaped when it has been broken open. Presumably, it is the undeveloped prophyll which will be more completely discussed with reference to intravaginal shoots. The next scale leaf is a short structure, about 2 mm. in length, and is the first true cataphyll. It is well supplied with vascular strands as any legitimate leaf should be. This cataphyll and its axillary bud are not located on the opposite side of the rhizome from the prophyll but are turned only one-fourth of the way around so they are on the side instead of underneath (text-fig. 9). Succeeding cataphylls then alternate first оп one side of the rhizome axis and then on the other. If this quarter-twist were not put into effect the axillary buds on the rhizome would point up in the air or straight down in the soil. The first few cataphylls are rather short, but later ones average just slightly longer than their internodes. Cataphylls do not vary nearly so much in length as do the internodes, nor do they appear to be so sensitive to water shortages. How Does the Rhizome Grow?—The scale leaves are produced from the grow- ing point of the rhizome in the same manner as described for ordinary leaves, except that blades do not develop. When a crown bud has been stimulated to grow into a rhizome, the small enclosing prophyll is broken and successive cataphylls elongate, though only one at a time. Each new scale leaf is a whitish cylinder which tapers to a strongly pointed tip. Since the new cataphyll develops within the previous one (text-fig. 10) there comes a time when penetration must occur. Consequently, the retaining point of the rhizome is broken open at the tip permitting the next cataphyll to appear. Two main processes occur on every phytomer: one is leaf growth, one is inter- node growth. On the rhizome, elongation takes place first in the cataphyll, then in the stem behind it, thus following the pattern described for phytomers in gen- eral. The scale leaf is extended not only by its own growth but by that of the internode below. It can be seen, therefore, that the penetration of one cataphyll by another is not simply a matter of one leaf growing up inside the other but is the product of two separate forces, leaf elongation and stem elongation. [Vor. 88 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN TRANSITIONAL LEAF OLD CATAPHYLL GROWING POINT MATURE CATAPHY LL ELONGATING INTERNODE ELONGATING CATAPHYLL ig. 7. A rhizome, X 4%. Fig. irst leaves of the rhizome, X 4% (P, prophyll). Fig. 9. AN - arrangement of а per buds on the bluegrass plant. 10. Diagram of a rhizome tip . 11. Diagram of a Fig. rhizome tip which has begun to produce cataphylls tipped with blades fay is 5. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 3 T1 One might describe the growth of the cataphyll as the forging of a steel tip. Once the tip is forged the developing internode serves as a shaft by which the tip is pushed out of the old cataphyll and through the soil. While this is going on, a new point is again being forged within the confines of the tip in use. Sooner or later the tips of the cataphylls become more delicate, a little less conical and a little more flattened. Upon close examination it will be seen that the point (only 1 or 2 mm. in length) is no longer a complete cone with a small pore, but is slit down one side (pl. 6, fig. 17). Furthermore, if we open this slit carefully we will find the slit point is separated from the rest of the cataphyll by a thin perforated membranous diaphragm which is the ligule. This new, longer, open point thus represents the first beginning of a leaf blade on the cataphyll. It is common to find many cataphyll leaves tipped with very short blades, but wherever this tip, by virtue of the removal of some physiological block, is allowed to reach a length of 4 or 5 mm. it is apt to be associated with a curving up of the rhizome and with its prompt appearance at the soil surface (pl. 6, fig. 18). How Does It Turn U p?—In many respects we should not speak of а rhizome's turning up at all, for very little, if any, bending originates in the internodes, nodes, or cataphylls of that structure, even though they may become secondarily curved by playing follow-the-leader with the new blade (text-fig. 11). This blade, in contrast to the indifferent sheath, has a strong objection to gravity, and as soon as it has exceeded the limitations of its enclosing cataphyll it shows this prejudice by striving to reach upward to the soil surface. This change from no blade to a fairly long blade (several cm.) is sometimes a sudden one, in which event only one leaf is required to make the break. The fact that the longer blade is usually associated with a longer sheath makes this a possibility. On the other hand, progress toward a long blade is sometimes slow, and two or three leaves may be involved in the effort to reach above ground. Once a bladed leaf reaches the light, no elongation of the internode beneath ordinarily takes place. Light, working through the blade, prevents internode growth. Weintraub and Price (1947) investigated the physiological basis of such inhibition in the oat seedling and showed which wave lengths of light cause the inhibition, but they did not isolate the photo-receptive substance. If blade-tipped cataphylls do not reach light, the associated internode will elongate. When a leaf with a long well-developed blade is prevented from reaching the light by being covered with soil, manure, or other debris the internode will become abnormally long (text-fig. 34). Where the transition from cataphyll to aerial leaf is abrupt, the rhizome shoot has a characteristic abrupt right-angled appearance, but where the transition is slow and internode growth takes place within the upbending leaf, the rhizome is gracefully curved. The manner of turning up, which varies greatly with habitat and strain, has a distinct influence on the depth and angle at which the crown of the subsequent shoot comes to rest. Every aerial leaf which develops from the underground growing point follows the course of the transitional leaves, since each new leaf blade arises within the [Vor. 38 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN previous sheath. Successive leaves increase in length, and later development of the crown is similar to that already described for the seedling. In pl. 6, fig. 18, the last roots on the top rhizome indicate the position of the growing point within enclosing sheaths. The most important result of this turning-up process is that the growing point of the shoot is usually left submerged in the soil to a depth of 1-3 cm. This is of great value to a pasture and lawn plant which must withstand close grazing, tread- ing, and mowing. Of significance also is the accumulation of a well-protected reservoir of subterranean buds in the crown area which can provide new rhizomes on a minute's notice. It will be noted in text-fig. 11 that usually the rhizome axis remains essentially horizontal and its buds point in the same direction as the rhizome. This exerts a directional influence on new rhizomes which develop. It explains in part the maintenance of a general similarity of direction from one gen- eration of underground shoots to the next. As the shoot becomes older the crown becomes more vertical and the buds more crowded; new shoots then develop in almost any direction. Why Does the Rhizome Turn Up?—The turning-up of a rhizome is usually not due to the influence of the environment on the rhizome itself, although mechanical obstruction sometimes plays a small part. Instead, turning-up seems to be correlated with some physiological development within the parent plant. Sharman (1947) has mentioned the fact that light reaching the rhizome was not the deciding factor in Agropyron repens, for rhizomes of that species will continue to grow horizontally (though they turn green) when the soil around them is re- moved. Similarly, above-ground stolons of many grasses and rhizomes of other plants such as the common Sansevieria have reduced leaves and remain prostrate regardless of their being exposed to light. With bluegrass, rhizomes artificially exposed usually begin to form blades and shorter internodes, and soon turn up. The number of transitional leaves, however, may vary considerably. That the rhizomatous habit in this species apparently hangs in a fairly delicate balance is suggested by various experimental results and observations. Harrison (1934) has found that rhizomes tend to be much shorter when grown in a high nitrogen medium. Brown (1939) has presented data that show decreasing rhizome length above a temperature of 70° F. and below 60° F. Heavy clipping and grazing also reduce rhizome length. In early spring and early fall, when above-ground growth becomes more proteinaceous and darker green and when carbohydrate reserves are drawn on heavily, practically all rhizomes below the ground begin to appear at the surface. Laboratory Experiments.—In a series of experiments, the growth of rhizomes was studied under controlled conditions. The rhizomes were separated from the parent plant and placed upright in test-tubes containing a little water, or in Petri dishes on blotter paper. These were then grown in the dark, and were subjected to the following treatments with the results indicated: 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 313 TABLE I. GROWTH OF RHIZOMES SEPARATED FROM PARENT PLANT Aver. 0 Number initial Grown Temp Days Treatment increase of lengt in (СС) їп rhizomes (cm.) length 4 3 Tubes 70 14 Tube filled with water 0 4 3 Tubes 70 14 Tube filled with nitrogen 0 4 3 Tubes 70 14 Tu lled with air 50 5 6.4 P. dish 40 14 Distilled wate 24 5 6.0 P. dis 40 14 Auxin, 1 mg./liter 20 5 6.1 P. dish 40 14 Auxin, 10 mg./liter 32 5 5.6 P. dish 40 14 Auxin, 50 mg./ liter 28 5 5.4 P. dish 40 14 Distilled water 33 5 5:7, Р. dish 40 14 Iodoacetate, .00001 т 19 5 6.4 P. dish 40 14 Iodoacetate, .0001 m 10 5 6.0 P. dish 40 14 Iodoacetate, .0005 m 15 5 6.2 P. dish 40 14 Iodoacetate, .001 m. 19 9 5.0 P. dish 40 38 Distilled water 34 18 5.0 P. dish 40 38 Glucose, 1% 88 While in general the numbers of rhizomes used were insufficient to lend sig- nificance to the small differences indicated above, the response to glucose is clear, and a slight inhibiting effect of iodoacetate is strongly suggested. All rhizomes which made any growth showed a distinct tendency to turn up. Glucose rhizomes continued alive for 314 months and eventually increased on an average of 125 per cent in length, while the distilled water controls showed almost no growth after two weeks, and were obviously dead after one month. Figure 19 of pl. 6 shows control and glucose rhizomes removed from the Petri dish for comparison. Another group of experiments utilized the rhizome while it remained attached to the parent plant. In this group test-tubes were supplied with rubber stoppers in which there were three holes. Into two of these holes right-angled glass vent tubes were inserted. The rhizome of a bluegrass plant was inserted into the test- tube through the third smaller hole and was then sealed in with a little cotton saturated with lanoline paste. The entire plant could thus be conveniently held in place by fixing it between the vent tubes. The test-tubes with their respectivé plants were then placed in a flat and covered with vermiculite so that no light could reach the rhizomes. The plants were in a perfectly normal position and could produce leaves and roots in their usual manner. The bent glass tubes en- abled the rhizome chamber to be filled with air, nitrogen, or water. Three tubes were left with free access to air; three were filled with distilled water and corkéd up; and three were flushed with gaseous nitrogen for ten minutes each week and were then sealed. All rhizomes grew well, though all showed development of leaf blades and a concomitant effort to turn up. In some cases later leaf blades became much shorter and practically scale-like after the parents became well established. Growth aver- ages of the rhizomes in each treatment over a period of four weeks were as follows: [Vor. 38 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN Tube filled with Total growth pA ОБО оте сы Я ET 7 cm. PROB Сал ыыы dabei чы RES 6.5 cm Water __. 6 cm Two of the air rhizomes continued to grow for six weeks longer, increasing in length by 23 and 29 cm. There were approximately 15 phytomers on each of these, produced over a total period of ten weeks. This average of one new phytomer every four to five days is somewhat less than that achieved under com- pletely natural conditions. In this experiment the vermiculite medium was sterile and not favorable for perfect development. These experiments were not designed with any idea of statistical demonstration of facts, but were rather trial runs to perfect techniques. The indications are fairly clear, however, that while isolated rhizomes apparently will not grow in water or nitrogen they will grow to some extent in air, and will remain alive longer and grow more when supplied with sterile glucose than with distilled water only. In every case where growth occurred rhizomes began to turn up. No treatment prevented this. Rhizomes remaining on the parent plant were able to grow equally well in gaseous nitrogen, distilled water, or air, and it happened that those in air outlived those in other media. In these rhizomes there was a strong tendency at first for leaves to develop, and for the tips to turn up. Later leaves, however, were much reduced. It is apparent that the rhizome has distinct possibilities in the field of experimental growth study, especially because of its natural independence of light. With the use of a complete nutrient solution and various hormones and inhibitors information might well be obtained on controlling factors in leaf differentiation and geotropic responses. Internode Lengths.—The internodes of rhizomes are delicate recording devices. Their lengths are highly variable and apparently indicative of some fairly funda- mental condition in the parent plant. A considerable number of rhizomes were dug up occasionally during several summers and each one was plotted on a graph on the basis of its sequence. In a given year a remarkable similarity of pattern was found to prevail in rhizomes from different plants and habitats. Text-fig. 12 shows diagrams of a few of these underground shoots dug from several localities during July, 1948. Not only is there a similarity of pattern, but also a strong correlation of phytomer numbers. The consistent reduction in inter- node length to a few mm. in all of these shoots was evidently indicative of a period of drought which occurred during May and early June. In rhizomes dug up in July, 1949, no such depression of length was seen, and the possibility that day length played a part was thus eliminated. The rainfall records given in text-fig. 12 show that there was no shortage of moisture in 1949. The conclusion that dry soil results in short internodes agrees with the observa- tion that excessively drained sites showed shorter average internode lengths (text- fig. 13). The data of Phillippe (1943) are significant here in that they show no 1951] 315 ETTER—HOW KENTUCKY BLUEGRASS GROWS 1948 1949 €— —RHIZOMES ——3 INTERNODE LENGTH it см. SHORT INTERNODE! RECOVERY ZA LONG INT RAINFALL INCHES PER WEEK NO DROUTH ED а 58:62 i 1 1 1 1 1 1 1 MAY 15 JUNE 1 JUNE 15 JULY! JULY 15 MAY 15 JUNE I JUNE 15 JULY I JULY 15 Fig. 12. Rhizome internode lengths of two years compared with rainfall records. Each curve represents a rhizome and shows the length of its successive internodes. Drought in 1948 produced short internodes; 1949 showed no such depression. [Vor. 38 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN INTERNODES NO. T І N 2 4 6 8 10 12 14 (6 20 22 24 26 28 20 32 34 INTERNODE бы (мм) Fig. The number of internodes of given lengths on rhizomes collected from two A d "ils, one of which was excessively drained. correlation between internode lengths and various levels of potassium, nitrogen, or phosphorous in his nutrient solution cultures of bluegrass. By matching and superimposing the patterns of a series of rhizomes dug up at different intervals during a year, a complete sequence of rhizome growth from some time in late May up to fall could be prepared. The sensitivity and uniformity of rhizome growth would seem to offer some interesting possibilities in interpreting the past history of a sod and related environmental and climatic conditions. When Do Rhizomes Grow?—A seasonal interpretation of rhizome growth is not simple. About the only time that one cannot find a few rhizomes still pursuing their underground path is in late winter and early spring, and there are even some exceptions to this. Wherever rich soil is kept warm and moist during winter by circumstances such as proximity to heat pipes and buildings, or in alluvial soils close to the water table, rhizomes which are especially thick and white can usually be found. Even these, however, ordinarily turn up with the advent of spring. There is one period when rhizomes are almost always found just beginning to grow, and that is when inflorescence elongation begins. From that time on through the rest of the summer rhizomes appear from subterranean axillary buds and grow at length through the soil. By late August most of these have turned up. At this time there is frequently a very brief flush of new short sprout-like rhizomes which are described in detail in the following section. For a few weeks in late Septem- ber or early October, when the leafy tops are growing rapidly, there will usually be a second period when few new rhizomes can be found still underground. A great deal depends on just how rapid the fall flush of growth is. In dry years greening may be so slow that cessation of rhizome growth is not very abrupt, and possibly may not occur at all. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 317 Production of rhizomes in late fall depends again on seasonal conditions and habitat. It is highly variable, but in general some buds do develop and grow a few inches. These late rhizomes are often caught underground, or just at the surface when the soil freezes. They then wait for spring thaws, as do the under- ground parts of many early-developing spring plants. Rhizome T ypes.—Rhizomes are not all the same. Some turn up almost im- mediately without ever making a long internode and look much more like intra- vaginal shoots except that they do not develop inside a living leaf of the parent plant (pl. 6, fig. 15). Under a few conditions rhizomes are inclined to be stolon- iferous; that is, instead of growing underground they grow along the surface of the soil and usually bear well-developed leaves instead of cataphylls. Those rhizomes which grow underground are also of various types. Some, such as those grown in the box, grow down and then up in a broad curve, while others grow horizontally just below the soil surface. Some are delicate and wiry, others fat, white, and flattened on the side. Rhizomes differ greatly in their tendency to branch. Generally, in established sods axillary buds are suppressed. If the terminal bud is injured, however, then one or more of these rhizome buds develop. Under especially favorable growing conditions vigorous rhizomes will branch voluntarily (pl. 7, figs. 20 and 21, left). Evans and Ely (1935) found that in spaced plants grown in a nursery plot about 10 per cent of the rhizomes showed some branching, and that as a result of branch- ing some clones or plants of bluegrass gave rise to as many as six vegetative genera- tions a year. Branching also occurs when an actively growing rhizome is exposed to bright sunlight accidentally, before it is prepared to turn up. In this case branching is limited to the last few nodes, and the branches do not have long white internodes but turn green and bend upward immediately. The development of branches apparently depends on some critical balance which can be altered by en- vironmental conditions. As with every other characteristic of the bluegrass plant variation with strain is also evident. There are also some fairly striking seasonal differences in rhizomes. In general, summer rhizomes pursue a deliberate underground course, extending the plant to some distance. These might well be called extensor rhizomes. They result when a developing bud feels some distinct obligation to remain underground and not to produce leaf blades except as a last resort. Figure 21 of pl. 7 shows examples of this type. Other rhizomes might well be termed sprout rhizomes, for they seem to be a response to a different combination of conditions in the parent plant. They represent a sort of desperate effort to perpetuate the parent plant in the face of some adverse condition. Such rhizomes are short and are characteristically found on plants which have been injured in some manner. They may arise from a crown or from another rhizome. Plate 6, fig. 19, shows the extreme type of sprout, de- veloped from a separate rhizome very short on food reserves. Harrison (1934) shows similar sprouts from rhizomes of plants which were injured by high tem- peratures. During a brief interval from late August to early September, just be- [Vor. 38 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN fore rapid fall growth begins, many short sprout rhizomes appear from the crowns of plants which have been burned, grazed, or mowed too close, or, weakened by rank growth, allowed to accumulate on the sod. Such sprout rhizomes play a very essential part in the perpetuation of rank meadow bluegrass, and are especially common on those plants which have bloomed during the previous spring (pl. 7, fig. 22). While rhizomes may be given names they themselves are not cognizant of any classification and may be intermediate in length, function, and appearance. Thus a pasture which has been grazed throughout the summer seldom contains any long extensor rhizomes. They are usually a sort of half-breed type, and are produced all summer long and well into fall. THE TILLER The tiller resembles a seedling or a turned-up rhizome in having green leaves and a nubbin of short root-bearing internodes at the base (pl. 7, fig. 23). This shoot arises, however, directly from an axillary bud (text-fig. 2; pl. 7, fig. 24) and not from a seed or from the tip of a subterranean stem. As pointed out earlier it develops in the axil of a living leaf. A BLADE | GROWING: | PROPHYLL SHEATH "c Å= NEXT TO LAST LEAF PROPHYLL LAST LEAF TILLER 2 1% f —TILLER 1 к" Fig. 14. The first organized structure Fig. 15. The parts of the plant which to develop from an axillary bud is the grow at the same time during the fall prophyll. It sheathes the first leaf. "shooting season." See Table II. +> 1951] í ETTER—HOW KENTUCKY BLUEGRASS GROWS 319 The Prophyll.—Morphologically the tiller is distinctive only in the possession of a well-developed prophyll, or pre-leaf (text-fig. 14; pl. 7, fig. 25). This small insignificant structure is seldom seen or examined closely, for its life is short and its purpose quickly served. It is the first organized structure which develops from an axillary bud (Sharman, 1945). It does not look like an ordinary leaf for it has no blade. It is a semi-membranous sleeve, about 2 to 3 cm. long, which is flat or somewhat grooved down the side nearest the main shoot. In cross-section it is semi-circular or somewhat crescent-shaped, and at each edge of the crescent is a prominent vascular strand which extends from the base of the prophyll to its tip. Very small strands may be visible along the outer rounded edge. The prophyll is a completely closed tube except for a small pore at its tip which represents the opening left when the primordial prophyll slipped over the growing point. In this and many other respects it is similar to the cataphyll of the rhizome. The prophyll, although the first structure to appear on the axillary growing point, does not elongate up through the confining base of its axillant leaf alone, but instead waits for the first true leaf which grows up within it. Measurements of a typical plant in process of elongating its shoots will show this synchronization. TABLE II. PARTS OF A PLANT WHICH GROW AT THE SAME TIME DOS Length in cm. of blade, sheath of leaf Parent plant Tiller sheath 1 Tiller sheath 2 and sheath Blade Sheath Blade Sheath Blade | Sheath Last leaf 1-2* 0 55 0 4* 0 Next to last 4.7 19% 2.0 25% (prophyll .4* ) 2nd to last 2.7 6.0 (prophyll 2.5) * Indicates organs in process of elongating. Here we see a repetition of a pattern of growth already discussed: that is, the simultaneous growth of a blade and its enclosing sheath. Although the prophyll does not exactly look like a sheath it seems to behave like one in most respects. It also bears a striking resemblance to the coleoptile or pre-leaf of a grass seedling (pl. 7, fig. 26) which accompanies the first true leaf of the germinating plant up through the soil and into the light. Both structures have two prominent veins and a pore at the top, are membranous, cylindrical, and have no blade. McCall (1934) has discussed this similarity in detail and has supplied anatomical evidence in support of the idea that the two organs are homologous. Since no studies of the course of prophyll growth have been made and in view of the evidence that coleoptile and prophyll are basically similar, a growth curve (text-fig. 16) is reproduced from Weintraub and Price (1947) which probably [Vor. 38 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN i 1 1 1 I І 1 1 l юор — d Ж БЕ ў MESOCOTYL / „ Ó / во 2 =a во LEAF EMERGENCE ~4 з | ü z ло > ғо d o 1 1 | 1 40 80 1 1 120 160 HOURS Fig. 16. Growth curves of the parts of a new oat seedling, in darkness at approximately 25° C. (From Weintraub and Price, 1947). expresses fairly well the relationship of the pre-leaf and its enclosed true leaf. This curve represents the growth of the oat coleoptile and first leaf in the dark. It is observed that both structures have the same growth rate, and also that as soon as coleoptile growth stops, the leaf emerges. One must assume, therefore, that both structures began growing at approximately the same time unless some drastic change of leaf growth rate occurred, which is improbable. While this curve per- tains specifically to the oat coleoptile, it undoubtedly applies roughly to the growth of the prophyll and the first leaf of an axillary bluegrass shoot. It may be interesting to explain the presence of the third curve, labeled "mesocotyl" (text-fig. 16). The mesocotyl is an internode of the oat seedling which elongates upon germination and sends the growing point and the immature coleoptile up toward the soil surface. The curve shows this internode elongating before any of the leaves, which is not in keeping with previous statements to the effect that internodes elongate only after their associated leaf is fully developed. However, McCall (1934) in his examination of the oat seedling has shown that the so-called mesocotyl is the first internode and that another very short internode, the second, separates it from the coleoptile which can be thus considered the second leaf. The second internode does not elongate. Since the mesocotyl belongs to the first phytomer, it is under no obligation to wait for the coleoptile to elongate. The leaf of the first phytomer is minute and not readily visible. The basic system of phytomer development thus holds as true in the earliest stages of seedling growth as in more mature plants. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 321 Growth of tbe Young Tiller.—Table II and text-fig. 15 show very well the interrelations between the parent shoot and the development of intravaginal branches. On a given phytomer the bud does not begin to grow into a tiller until the phytomer leaf has matured its blade and begun its sheath. Then four organs grow at the same time, the main sheath and the next main blade, the prophyll, and the first shoot leaf. Since the prophyll and the shoot are thus synchronized, succeeding branch shoots are also synchronized. This accounts for the observation that in a sequence of intravaginal shoots on a plant, from the oldest to the youngest each shoot has one less leaf (text-fig. 17). There are exceptions to this of course. No one ку; Я n | м w [ Ыри 7 PARENT TILLERS 2 3 . On a plant which produces numerous tillers, each younger tiller is one leaf ptu W is short winter leaf. phytomer seems to be wholly compelled to operate strictly in agreement at all times with its neighbors, though for shoots produced during the optimum tillering period in fall, under good conditions, such agreement seems commonly to exist. Under less favorable situations it is only reasonable to expect that a bud might hover between the decision to grow now and become a tiller or wait and grow up to be a rhizome. The indecisive individuals explain transitional types which are not strictly synchronized. The prophyll may be longer but it is usually shorter than the leaf blade it en- closes. In any case, the first sheath raises the first leaf out of the prophyll and exposes it to light. Other leaves of the shoot follow in the usual sequence, and the tiller becomes a new plant. Its own axillary buds can develop into other shoots in the same fall provided shooting starts early enough. Three generations are not uncommon and four or five are possible (pl. 9, fig. 28). Roots grow out from each mature node of the tiller crown and give it some measure of independence. In late spring any spare buds may develop into rhizomes. An inflorescence may appear the first spring though it is usually shorter and bears fewer leaves and spikelets than an inflorescence of the main axis. An intravaginal shoot with as few as two leaves can produce a panicle, while a rhizome with less than seven or eight leaves seldom flowers. This is in agreement with the observations of Cooper [Vor. 38 322 ANNALS OF THE MISSOURI BOTANICAL GARDEN and Saeed (1949) on ryegrass where a minimum of eight leaves are required before the main shoot will flower, while some tillers flower with only a prophyll. Shoot vs. Leaf Widtb.—The above facts indicate that the flowering stimulus, once developed, can be transmitted from the main shoot to its tillers. To what extent other aspects of the growth of tiller and main shoot are interrelated is dif- ficult to tell. Von Octtingen (1930), in a study of bluegrass leaf widths, stated that on rhizomes which have turned up there is usually а gradual increase in leaf blade width up to a certain point. Wherever an intravaginal shoot appeared, how- ever, he found the leaf above to be abnormally narrow. This he attributed to a drain on the food resources of the main shoot by the branch. There are several reasons why such an explanation may not be altogether justified, though it is not so much with the idea of disproving his idea that the following discussion is presented as it is to demonstrate how information such as that included in this study may apply to a specific problem. In the first place, as we have seen, the leaf blade on the main axis above a new tiller has practically reached its full development by the time the prophyll and the first leaf of the tiller start their elongation, so that one would expect any competi- tion for food to influence the sheath above instead of the blade. It might also be added that bluegrass is an extraordinarily efficient plant and only a small amount of leaf would need be exposed to light for it to attain its independence. The en- closing leaves of winter are so short and limp that the new shoot reaches the light very soon after it starts to grow. Likewise, the first leaf is very small and would not consume extensive reserves in reaching its full extension. The efficiency of bluegrass in making maximum growth with a minimum of leaf and minimum con- sumption of stored carbohydrates or of accumulation of carbohydrates on a mini- mum of leaf has been suggested by data in Smelow (1937), Klapp (1938), Peter- son (1946), and Harrison and Hodgson (1939) among many others. Von Oettingen himself has indicated that reduced width is not correlated with a reduced number of vascular strands. In tillering varieties of sweet corn it has been shown that so-called suckers contribute to, rather than drain, food supplies of the parent (Jones et al, 1935). The explanation of Von Oettingen's observat- tion is probably much more simple and points out the need for understanding not only the course of anatomical maturation but also the seasonal sequence of develop- ment of a plant to which statistical and taxonomic techniques are to be applied. The comparison made by Von Oettingen involved rhizomes which had no shoots and those which had only one or two. As a general rule, any rhizome which appears in fall will develop at least one or two side shoots, but those appearing in winter and spring are not apt to have any such shoots. The plants with branches thus go through a winter, and during that time leaf width of short-leaved plants decreases naturally (text-fig. 18). It is on these one or two late fall and winter phytomers that most of the branch shoots appear, and they are thus automatically associated with narrow leaves. Spring leaves are longer and wider and thus the contrast is accentuated. If, then, plants are later pulled up in summer at random 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 323 and the leaf widths examined, the significance of the narrowness of leaves asso- ciated with intravaginal shoots is not apparent. Even the observation of a correlation between shoots and narrow leaves is open to some criticism. According to text-fig. 18, the relationship is actually just the reverse in rank meadow grass, where fall leaves increase in width while shoots increase in number. In grazed grass the early fall leaves increase in width while winter leaves become narrower. Shoots, meanwhile, increase in number all winter long, and virtually every new bud: develops into a tiller. Therefore if the shoot had any effect on the leaf width it should show up in the average. TILLERS LEAF (NO) LENGTH 4 35 35 TILLERS, GRAZED 3 30 LEAF WIDTH, UNGRAZED 25 25 LEAF WIDTH, GRAZED 2 “20 15 15 TILLERS, UNGRAZED 1 10 5 5 LEAF LENGTH, UNGRAZED LEAF LENGTH, GRAZED о о 16 Лс сше сг сл TN \ 1 AUG.28 SEPT.19 остё МОУ7 NOV.27 DEC.27 JAN. 24 MAR. 5 Fig. 18. Relationship of tillering, leaf length, and leaf width in meadow and pasture bluegrass in fall. There are also indications that the severity of the winter and the seasonal development of available nitrogen in the soil influence the sequence of leaf widths. If the soil is frozen much of the winter, leaves will be deprived of water and will be narrower as a result. In mild winters such as that of 1949-1950, when the leaves were measured (text-fig. 18), very little freezing weather was encountered. One might thus expect that midwinter leaves of bluegrass grown in northern localities would consistently be narrower than those of southern-grown plants. Shoot vs. Leaf Lengtb.—NW/hile the relationship between leaf width and the presence of shoots does not seem to be constant, there is one correlation which seems to be rather consistent under normal cultural conditions: that is, the asso- ciation of short leaf length with much tillering. This association is suggested in many ways. It is quite noticeable that mowing or grazing rank grass in early fall causes a great increase in the number of tillers. On unmown grass develop- [Vor. 38 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN ment of shoots is weak and delayed until late fall or early winter. This difference is expressed the following spring in a much larger number of panicles in the mown grass (Spencer, 1949) and in the thickening of the early spring stand. Some preliminary measurements of leaves on plants examined in late spring gave the following results: Blade length Number with shoots Number without shoots 13—30 cm. 3 12— 3 cm. 27 More detailed data taken on grazed and ungrazed sods in fall and winter are compiled and presented in text-fig. 18, which shows clearly the delay of branching characteristic of ungrazed grass. This delay amounts to nearly two months, and the critical leaf length would seem to be around 10 or 12 cm., which is in keeping with the above figures. On grazed grass, however, leaves average under 12 cm. for most of the summer, but no shooting occurs until September, Leaf length alone is therefore not the deciding factor. We have thus defined the permissive and obligatory limits of shoot development. They can develop as early as late August, but seldom appear in quantity until October. By December they seem more or less obliged to appear. he association between short blades and intravaginal shoots is further reflected by results obtained in some fertilization experiments made during different seasons. The fertilizer used was undiluted horse urine applied at the rate of two gallons to a square meter. Urine was used because of its quick action and its natural impli- cations. The results are shown in the following table: ABLE III LEAF LENGTH AND NUMBER OF INTRAVAGINAL SHOOTS AS RELATED TO TIME OF FERTILIZATION. (FIGURES REPRESENT AVERAGES OF 10 PLANTS MEASURED ON JUNE 4, 1949) Fertilizer applied Number of Leaf length of branches last complete leaf (cm.) Late August 5.8 6.5 Late September 10.0 21.2 Late October 8.1 33.2 Early January 3.7 55.7 Early April 3.1 33.7 Control (no fert.) 3.6 15.0 Plants typical of the above experiment are shown in pl. 9, fig. 29. With a fall application a decided increase in shooting was observed so that in spring a large number of panicles were seen, but the numerous leaves were abnormally short. Winter applications yielded only slight increase in shoots and inflorescences, but the leaf was exceptionally long. The data for August show that again shortness of leaf is not infallibly followed by prolific branching. It emphasizes, instead, that probably no correlation holds true except under specific seasonal and environ- 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 325 mental conditions. The rhythm of the plant itself must be considered in in- terpreting measurements of this sort. In August, day length, soil moisture, and temperature are all quite different from those of September or October, and as already shown the plant is essentially incapable of forming shoots during that period. It is not strange that its response then should differ considerably from that a month or two later. There was little carry-over effect of the August application of urine because of a hot dry period during which much of the nitrogen was ap- parently lost. The extreme shortness of leaf in the August plants is not readily explainable, except for the possibility that the slight burning effect which the urine had on that occasion may have depleted the plants’ reserves at a critical time. It has been observed that very close cutting or stripping of top growth at this period results in weak short-leaved plants. Tillering in bluegrass has been shown to depend to some extent on day length (Evans, 1949; Harrison, 1934). Temperatures may also be a determining factor, for in the laboratory cultures with baby bottles it was found that while no shoots were produced with a 14-hour day at temperature of 70° F., at 40° numerous shoots appeared. Shooting is thus seen to be a complicated activity. It depends on day length, temperature, nutrition, stage of development of the plant, and is associated with short leaves, though all short leaves do not produce shoots. Leaf width bears little if any relationship to shooting, except coincidentally. Death of Shoots.—There is strong evidence that the parent plant may control the destiny of the tiller at certain critical points of development. This begins to be obvious at the time of elongation of the panicle, about the 10th to 20th of April in central Missouri (Table IV). TABLE IV. DEATH OF INTRAVAGINAL SHOOTS IN SPRING Date Number of Number Per cent shoots examined dead dead March 19 12 0 0 April 13 0 0 April 17 19 6 32 April 24 15 4 27 ay 1 16 2 13 May 7 17 9 33 May 14 16 8 50 May 23 19 8 42 May 31 26 11 42 Dying of tillers is especially evident in rank, sod-bound, nitrogen-deficient grass which has not been grazed or mown, though it is probably also a common event wherever conditions are not ideal. Death of some branches is a necessity in a sod which maintains a more or less even density from year to year, even though the average number of new vege- [Vor. 38 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN tative shoots per plant is seldom more than two. The main shoot dies after bearing an inflorescence. Of the two tillers, one is generally dead before flowering is com- pleted (Table IV). Perpetuation of the grass through the rest of the season is thus left up to the one remaining intravaginal shoot and whatever rhizomes de- velop during summer or fall. By early winter, however, practically all the original tillers have died, and it can be seen that maintenance of the population from one year to the next under meadow conditions is primarily dependent on the production of rhizomes. Rhizomes which develop in summer are primarily responsible for the inflorescences of the following year. That the apparent detrimental effect which blooming has on the tiller may be related to water shortage is suggested by Sharman's statement (1947), that when the apices of the blind shoots (non-blooming tillers) of Agropyron repens are dis- sected during the flowering season they are "pliant and do not cut crisply, as though they were suffering from lack of water." As in bluegrass, these shoots of Agropyron generally die before the end of the year. It is also well known that grass crops such as maize require especially large quantities of water during the period of elongation of culms. It would seem to be to the grazer's advantage to keep these tillers alive since they provide much of the summer's forage. By mowing at the proper time (about the third week in April) and by cutting low enough to catch the flowering head, blooming can be prevented and the vigor of the tillers might well be maintained. Unless these tillers live into the summer there may be a considerable loss of forage or sufficient thinning of the stand to permit invasion by weeds. Mowing at this stage of growth is not common in the United States, but Hamilton (1942) has mentioned that it is a practice on the intensively farmed ryegrass pastures of New Zealand, and suggests that were more power-mowing equipment available it might be profitably practiced more widely. Grazing at the proper time might also be used to serve the same purpose. Ryegrass, as Cooper and Saeed (1949) point out, is more of a problem, for usually a single cutting is not sufficient to prevent later flowering. Since bluegrass can only become "ripe to flower" during winter, one cutting is all that is necessary in that species. Ahlgren (1938) has made tests simulating three conditions of spring-grazing, the first cutting of bluegrass being made: (1) to a 112-inch level at a grass height of 4—5 inches; (2) when the head had just appeared; (3) after the head was mature. His data seem to substantiate the above conclusions, for making the cutting at the 4- to 5-inch height for a period of six years gave a progressively greater yield each year. Cutting at the time of appearance of the panicle was also consistently better than cutting at maturity. The data are reproduced below: The 4- to 5-inch height just about coincides with the period when the panicle is starting to elongate. The specific time for mowing to best advantage can be determined by examining the heights of a few panicles while they are within the sheaths to see if the mower will cut them. It is quite possible that one of the more 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 327 subtle benefits of grazing а pasture involves the prevention of heading at an early stage, and thus the encouragement of the tillers which must take the plant through the summer. TABLE V PERCENTAGE YIELD OF KENTUCKY BLUEGRASS FROM PLATS GIVEN VARIOUS CUTTING AND FERTILIZATION TREATMENTS. (FROM AHLGREN, 1938) Complete fertilization No fertilization Fully Fully Year 4—5 Early headed 4—5 Early headed inches heading to mature inches heading to mature % % % % % % 1957 79 123 100 yi 93 100 1953 97 9. 100 74 81 100 1934 113 103 100 1 101 100 1955 108.2 108.2 100 118.8 110.8 100 1936 143.6 105.4 100 189.8 136,7 100 THE FLOWERING SHOOT Purpose and Design.—Bluegrass must be appreciated more for the delicate and effective way in which it presents its small flowers to the wind than for the beauty of the individual flower. There may be from 100 to 1000 flowers, or florets as they are called in grasses, on each flowering shoot. Considering that the plant has only the phytomer to build with, it does a very efficient job of display. By elimina- tion or reduction of all unnecessary structures including blades, sheaths, roots, and some buds, a flowering shoot of quite simple design is achieved. This shoot con- sists of a main stem or culm which holds a branched pyramidal inflorescence or panicle well above the ground (text-figs. 3 and 20). Clusters of florets (called spikelets) depend from the tips of the panicle branches. The average culm is merely a portion of the vegetative axis which has been obliged, because of the initiation of an inflorescence at its growing point, to follow a course of develop- ment different from that of non-flowering shoots. It is distinctive chiefly in having long internodes, reduced leaves, no buds or roots, and a hollow stem, all of which indicate the strong economies of material put into force when the plant is called upon to flower. The average culm consists of a series of 3, 4, or 5 phytomers, with each higher one being less like the normal vegetative ones at the base of the plant. For con- venience the standard number of phytomers will be considered as 4, and will be numbered from the lowest, least elongate (1) up to the highest and longest (4). Actually there is no specific phytomer at the base which one can designate as belonging to the flowering shoot rather than to the vegetative shoot from which [Vor. 38 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN Y ЗА | Ў 2 —ы = Ф 21 а” — a} 2 2 au ze > y <- = oe INTERNODE 4 ——- PHY TOMER 4 TILLER FIG.19 B LEAF 3 —— | INTERNODE З. PMYTOMER 3 —! m SHEATH 3 | PHYTOMER | — J FLOWERING SHOOT PHYTOMER 2 Fig. 19. Diagram of a typical ч зше (А), 2 one d its неу ee Me m с plant from ап old m showing an average flow shone, allen жене shoot is ... Ьу ы саа ре e on the lower side е che Bere sheat 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 329 it springs, for the transition is not always abrupt. Each of these culm phytomers is a distinctive combination of different internode, blade, and sheath lengths (text- fig. 19A). In an old unmowed, ungrazed, unfertilized, unirrigated, unshaded stand of Missouri bluegrass the lengths of successive internodes will increase toward the panicle in a rough ratio of 1, 4, 12, 30. The panicle on top would then be about 9 cm. Each of these internodes, except the last, bears a sheath at its top which in turn bears a blade. The average sheath lengths in centimeters would be about 9, 12, 13 and 0. The leaf blade lengths are in reverse order to the internodes for they average 13, 9, 4 and 0. We thus can draw the following thumbnail arithmetical sketch of the flowering shoot: TABLE VI COMPARISON OF LENGTHS (IN CM.) OF FOUR SUCCESSIVE CULM ". THOSE ON А SUCCESSION ОЕ FOUR VEGETATIVE ie а ASSOCIATED TILLERS. (SEE TEXT-FIG. 19 A & I Culm phytomer | Vegetative phytomer а — | Р 1 2 3 4 icle 1 2 3 4 Internodes 1 4 12 30 9 —* -- -- -- Sheaths 9 12 13 0 0 8 6 5 6 Blades 13 9 4 0 0 12 20 24 30 Total leaf 22 21 17 0 0 20 26 29 36 Date leaf completed Apr. Apr. Apr. Apr. Apr. May june 12 LZ, 26 12 26 16 15 * Vegetative internodes too small to be measured. The last sheath immediately below the panicle may appear as a small hook or scale at the top of the last phytomer. Rarely a complete leaf may be found. These figures give only a general picture of the culm, and it is not intended to suggest that they pertain to other strains or plants from other habitats. The first phytomer is seen to have a very short internode and a long leaf blade while the last phytomer has a long internode and no leaf at all. The first leaf is not greatly different so far as blade and sheath length are concerned from its vegetative contemporaries (text- fig. 19B, No. 1). With the second blade, however, the influence of the developing panicle is evident, for comparable vegetative blades average 10 cm. longer than those on flowering shoots. Culm leaves are consistently broader than the contemporary leaves of associated vegetative shoots. Thus measurements of the last leaves on 48 plants, 24 with inflorescences and 24 without, showed on April 21 average widths of 3.4 and 2.3 mm. respectively. On May 1, after the last culm leaf had appeared the difference was 2.7 and 2.3 respectively. This indicates that the /аѕ culm leaf is less wide than its predecessors. In general, there is little difference in the width of the first two culm leaves. The internodes of the bluegrass culm are rather delicate structures which would soon fall to the ground were they not partly encased by strengthening sheaths. [Vor. 38 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 22. А bluegrass panicle, X 44. Fig. 23. А bluegrass spikelet, X 16. The lowest two internodes are usually completely hidden from view while the third generally shows only a few centimeters at the top. The last internode is sheathed about half way up. The degree of overlap varies radically from plant to plant. In case the bluegrass culm, while elongating, happens to fall to the ground as a result of wind, rain, or trampling, a rapid reaction takes place in the vicinity of one or more of the nodes causing development of tissue on the lower side of the culm which gradually restores at least a part of its perpendicularity. On close ex- amination it will be found that in bluegrass the actual growth and straightening process depends more on the basal part of the sheath than on the internode (text- fig. 21). This sheath builds up a wedge-shaped accumulation of tissue around its lower side, which as it bends up, forces the delicate culm within to raise also. Arber (1934) states that in other species of grass such straightening may involve only the sheath, as in bluegrass, or both sheath and internode, or internode alone. The bluegrass internode, whether on the culm or the rhizome, apparently has very little to say about what position it will assume. The Panicle.—At the summit of the culm is the panicle or inflorescence (text- fig. 22). Тһе branches of this panicle, which arise at nodes, are grouped together in successive fan-shaped tiers inserted alternately on the panicle axis. Since the branches of the lower tiers are longer than those of the upper ones the outline of the inflorescence is somewhat pyramidal. There are from three to five branches at each of the lower three or four nodes, and these are almost perpendicular to the main axis at flowering time though in some strains they may droop or be appressed to the panicle axis.. The spikelets, or flower clusters of the grass, are borne at the tips of these branches and of their subsidiaries. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 531 The panicle axis, or rachis as it is sometimes called, is a continuation of the culm. It is composed of some eight or more internodes, only the lower ones being very prominent. This contrasts with the culm internodes which decrease in length toward the base of the plant. There are no leaves on the rachis phytomers. Beginnings of tbe Flowering Sboot.—A flowering shoot is a sudden, striking thing and seems so different that it sometimes prevents us from understanding its relation to the course of development in the rest of the plant. In bluegrass the initiation of the flower takes place in the cold weather of late fall and early winter, and thus development is slow and goes unnoticed. In spite of its early start, blue- grass does not display its flowers until mid-May in Missouri. Cold weather and short days have no magic in themselves, and can work only through the physio- logical systems of the plant. Under fall conditions it has been shown (Buckner and Henry, 1945; Peterson, 1946) that bluegrass carbohydrate reserves rapidly reach a peak. In general, such accumulation of carbohydrates seems to be one of the significant factors leading to the "ripe to flower" condition in grasses. That other systems play a part goes without saying, but no effort will be made here to fathom the chemical complexities associated with the inception of flowering shoots. To trace the development of a vegetative shoot into an inflorescence we shall have to go back to the growing point, for that is the beginning of everything. Musgrave (1940), Nishimura (1923), and Evans (1949) have given good ac- counts of this process. The following description attempts to synthesize the work of these three authors. The vegetative growing point runs on a rather close margin. It commonly has only three or four phytomer primordia differentiated and available for use (see pl. 5, fig. 13). As the plant approaches fall, however, either the phytomer organs differentiate less rapidly, or else new primordia are cut off from the growing point at an increasing rate, or both, for incipient phytomers begin to accumulate and as a result the blunt rounded vegetative growing tip of the youthful shoot becomes a longer, rather conical, wrinkled, translucent point (pl 8A). This is a tendency which precedes inflorescence development in practically all grasses, although the number of phytomers accumulated on the point and its actual length may vary greatly with different species. Protuberances Appear—The lower, old phytomers of this new cone appear normal enough, and bear the usual leaf primordium at their upper end, but toward the point, on the younger primordia, there may be only a faint ridge or no sign of a potential leaf. Then something happens. The deadline is up, and all phytomers which have not formed leaf primordia are obliged to get along without them, for the reproductive phase has begun. The first visible sign of this event on the growing point is the appearance of small knobby protuberances which, though arising in a similar alternate manner, are obviously different in origin from the crescentic ridges from which leaves arise (text-fig. 24A; pl. 8B). These shining protuberances appear first on those phytomers which have only the faintest trace of a leaf ridge, then soon both below and above. The protuberance is a rather [Vor. 38 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN C 5 f2, etc., successive florets; an, stámen primordium; pi, pistil showin successive leaf primordia; g, glume; fZ, etc., е lodicule; s, stamen. primordium; ғ, rachilla; p’, palea; lo, unique development for, although it obviously represents a branch of the main axis, it is the only time during the growth of a grass that a growing point is not encased in its own sheathing cone of leaves. Here is a phytomer which consists of nothing but stem. Through continued cell division it grows very slowly, almost as though it were a root. The one phytomeric function which it seems to retain is the ability to branch. Thus after about a month secondary protuberances arise (pl. 8C), and in another month tertiary ones appear. By this time most of the winter has passed, and in late March the growing point may still be only about 1.5 mm. long and yet will contain practically all the cells necessary to build the entire skeleton of the mature panicle. The Development of Spikelets.—In April on all the naked ends of protuberances of all orders small crescentric ridges begin to appear (text-fig. 24B). Just as in 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 333 the main vegetative point, these ridges are alternately arranged and later develop into minute leaf-like structures. Thus the conventional phytomer consisting of a leaf and an internode is reestablished. This is the beginning of the bluegrass spike- let, or flower cluster (pl. 8D; text-fig. 24C). The first signs of spikelet develop- ment appear on those protuberances located somewhere between the middle and upper third of the panicle, and the spikelets at the tips of the main branches develop before those on subsidiary branches. The first two leaf-like structures which develop from these primordia are called glumes. They are only 2 or 3 mm. in length when mature, and completely enclose the growing point and its developing structures. Once conceived, the two glumes elongate more or less simultaneously, which is not exactly in keeping with the usual sequence of phytomer development. As shown in text-fig. 26, the last two blades and then the last two sheaths of the culm elongate simultaneously, so that apparently when the growing point is going into or coming out of the phase of panicle formation the normal sequence of elongation is somewhat altered. Before the glumes close over the growing point a series of 3-5 phytomers with leaf primordia have begun to form (text-fig. 24C; pl 8E). These primordia elongate gradually after the glumes enclose them. They are called lemmas, and they are distinguished from the lower glumes because they bear, in their axils, an axillary bud which will soon develop into that long-waited-for phenomenon, the floret (text-fig. 24D).* Differentiation of successive lemmas and florets progresses from the base of the spikelet toward the tip. As with most axillary buds, the first structure produced from the floret bud is a membranous, two-veined, compressed sheath-like organ which, though not called a prophyll any longer, very much resembles one. It is dignified by the name palea. The growing point of the floret bud (text-fig. 24D), then proceeds with the differentiation of the critical flower parts, producing in turn three stamens from small papillae just below the growing point, the pistil from a crescentic pro- tuberance very much resembling a leaf primordium, and finally the two styles and stigmas from papillae on the pistil (text-fig. 25). The ovule or egg sac, which is perhaps the final leaf primordium, remains short inside the pistil after that struc- ture has closed around it. Within it will develop the egg cell, which would appear to be the arrested growing point. From four to seven phytomers are ordinarily found in a bluegrass spikelet. Their leaves are the glumes and lemmas, their axillary buds are the florets, and their minute internodes join together to form the spikelet axis or rachilla (text- fig. 25). The transition toward termination of the growth of this axis is usually expressed in the form of a final much-reduced phytomer which consists of a fragile internode and a minute lemma, the so-called sterile floret. P348 édiowing the phytomer concept to the end and assuming that the glumes are homologous with sheaths or blades, we should, in reality, recognize the first three leaves (two glumes and the first lemma) as glumes instead of just the first two, since the first floret is a bud at the base of the fourth spikelet phytomer, and thus belongs to the fourth leaf, not the third, which is therefore sterile.’ This is a technicality of no great importance to the matter at hand, and might cause considerable confusion a taxonomists. That there are no hard-and-fast rules in this mong grass respect is illustrated by the observations of Sharman (1947) that florets are sometimes found in the axils of the glumes. [Vor. 38 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN A, flower of a grass (schematic): lo. lodicules: r, rachilla : le, lemma; pa, palea; В, longitudinal diagram of flower ; C, spikelet; D, floral diagram. А A diagrammatic sketch of a typical grass flower. (From fig. 429, on p. 627, of Johnson's "Taxonomy of the Flowering Plants," 1931. Reproduced by permission of the Century Co., Publishers). Elongation of Panicle and Internodes.—As the process of spikelet differentiation proceeds in early April, it seems to send a pulse of life down through all the phyto- mers which have been biding their time since being inhibited when the proliferating urge assumed control months before. Their internodes begin to elongate. The actual processes involved in this revival of growth are shown in text-fig. 26. The initial quickening is limited to the panicle proper. Growth is slow at first, less than 1 cm. between March 27 and April 12, then it increases rapidly during the next ten days to full length. It may not be altogether a coincidence that this panicle elongation comes immediately after the first and second leaves of the culm have matured. It suggests that some inhibition may be involved, Not until the panicle has reached its mature length do the internodes of the culm begin to elongate to any extent. An exception to this is the first short inter- node, which begins to lengthen several weeks before the panicle matures. This course of growth seems slightly different from that characteristic of wheat as shown in text-fig. 6. In wheat the growing point at the beginning of the pro- tuberance stage has had more phytomers and culm development is more drawn out and deliberate, with each successive internode behaving as an individual. The result is a culm with more joints of more nearly equal length. Maize is a more extreme example of this same situation. Bluegrass with its short growing point presents only four internodes, only three of which reach any considerable length. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 335 It is a common observation that in wheat, corn (Sharman, 1942), and rye (Prat, 1935) the last two internodes elongate at the same time. Prat attributed this to the fact that there is no leaf on the last phytomer, therefore, the internode does not have to wait to begin its growth. The fact that in the bluegrass strain ex- amined here the last three internodes grow at the same time is probably related to the absence of the last blade and the shortness of the third one. Culm Leaves.—While it is usually considered that the bluegrass plant lies dormant during winter except for inflorescence initiation, it is actually producing a certain amount of leaf growth. On shoots destined to flower, no new leaf primordia are produced once proliferation takes place, but those primordia already completely formed do follow a normal course of development within the limits of environmental factors. The number of leaves which thus develop during the coldest weather is usually only two or three and these are very short (text-fig 26WB). The internodes below these winter leaves do not elongate, for they are apparently mature by the time spikelets begin to differentiate in spring. These leaves differ little, if at all, from comparable leaves on vegetative plants or on tillers of the same plant. QUIE 28 - o Т CENTIMETERS n го > 02 0 o Т a pat 1 MARIS —— APRI PRIS MAY! MAYIS JUNEI — JUNE 15 Fig. 26. Curves of growth made by various organs of meadow bluegrass during the period of flower-shoot development in spring. [Vor. 38 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN With the approach of warmer weather in late March leaf growth receives an impetus and new leaves grow longer. The last true winter leaf and the first culm blade apparently respond, rather abruptly, to the rise in temperature. (See sheath WS and blade WB and B-1 in text-fig. 26.) The first blade on the culm continues elongating. By early April it is joined by leaf blades 2 and 3 and sheath І, which elongate at the same time. When these mature, sheaths 2 and 3 and the panicle elongate almost simultaneously. Immediately upon their maturation, internodes 2, 3, and 4 begin to grow. This rush of growth is in decided contrast to the deliberate growth of phytomers on vegetative shoots of the same plant where only one blade and one sheath elongate at the same time and the internodes do not elongate at all. This speeding up of the development of phytomers immediately below the inflorescence has been noticed by Sharman in maize (1942) and in quack grass (1947) and is suggested by data of Prat (1935) for rye. This is one of the factors involved in the rapid appearance, in spring, of a large number of leaves in a very short time. The more flower shoots there are the more vigorous this spring flush will be. There is an interesting contrast between culm and vegetative leaves with re- spect to final length and rate of growth. Up until about the last of March the growth rate of the two types of leaves is very similar. After that each successive culm leaf elongates less rapidly and is successively shorter (text-fig. 26, leaves B-1, B-2 and B-3). Tiller leaves (T-1 to T-4), on the contrary, elongate somewhat more rapidly, and become successively longer even though on the same plant. This would suggest that if the developing panicle has any effect on leaf growth, it is not transmitted to the leaves of intravaginal shoots as the stimulus for flowering apparently can be. More probably, the shortening of culm leaves is not an in- hibitory effect at all, but dates back to the time when the growing point first changed in the direction of flowering. Leaf primordia were stopped іп their tracks and consequently when an opportunity came the following spring to elongate there were successively fewer cells in each younger primordium availablé for elongation. The absence of buds on the culm is also probably due in part to the fact that their formation was interrupted by the inception of panicle development. Sharman (1947) commented on the absence of buds on the culm of certain species, stating that microdissection showed that early stages of bud formation did take place. The failure of the bud to develop further he attributed to the effect of elongation in pulling the tissues of the bud apart. This seems improbable in view of the presence of well-developed buds on the phytomers of rhizomes which have long internodes. The influence of the developing panicle in suppressing further growth is probably of more significance. The reason for suppression of roots on culm phytomers is even less clear. In part, it may merely be that the culm internodes do not come close enough to an adequate rooting medium. Roots have been seen on culm phytomers as far ad- 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 337 P 12 H e 0r т 2 e o o CINE > 2.1. . 74 t.e? è o 5 y өө .” E ela ee << | Ж ө ш 4 ov oe ©з . aal e о e? "li > т е. $ . .. e e 9 a> ee e > * . T ux PME NN" - e e x eo өө .. eee ^ е 1 1 1 1 1 1 \ 1 E 0 2 4 6 8 10 12 14 16 18 20 22 INTERNODE LENGTH (CM) Fig. 27. The relationship between length of y internode and phytomer leaf on the culm of bluegrass. Plants were from various meadow habita vanced as No. 5, under conditions where the nodal region was exposed to moisture and darkness. The relationship of short leaves and long internodes in the upper part of the culm is apparently more than mere coincidence. Although the shortness of the leaf may be explained on a historical basis, the long internode probably involves other factors. It has been mentioned in connection with the rhizome that the internodes of phytomers whose leaves turn up and reach the light are inhibited. If the leaf does not reach the light then the internode elongates. Many other observations indicate that the bright light inhibits the elongation of internodes, while in the dark or shade elongation can take place. It is probable that the short- ness of the culm leaves reduces their inhibiting action at the same time that the developing inflorescence is producing auxin in quantity (Prat, 1935). Thus both conditions appear to favor increased internode length. The relationship between leaf and internode length on the culms of bluegrass from a variety of habitats is shown in text-fig. 27. As further evidence for the validity of this relationship, it has already been mentioned that the last internode, devoid of any leaf, is by far the longest. Also suggestive is the fact that very little elongation of the second, third, and fourth internodes takes place until the last leaf has reached almost its full extension. Unfortunately, the panicle reaches its final length about the same time, so that there is no way of determining which is the more significant releasing factor. [Vor. 38 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN LEAF GROWTH IN RELATION TO SEASON AND ENVIRONMENT Day Length, Temperature, and Growth Rate.—The casual observer probably realizes that bluegrass begins to green up in early spring, blooms in May, and gets tall and somewhat brownish in summer, then greens again in early fall and turns brown again in December. This sequence of events seems quite regular yéar aftér year. Weather variations induce only moderate fluctuations. One would gather from this that the bluegrass plant has a way of knowing what time of the year it is. Apparently that knowledge is conveyed largely through the medium of day length and temperature. Evans and Watkins (1939), Evans (1949), and Peterson and Loomis (1949) have described the influence of various changes of day length on bluegrass. The chief conclusions reached are that leaf length and tillering can be controlled by day length, though temperature may affect the leaf length to some extent at any given day length. The results of Peterson and Loomis’ experiments are indicated below: AVERAGE LEAF LENGTHS Temperature Day length 11 hrs. 19 hrs. 56? — 65? F. 9 cm. 15 cm. 61° —75? F. 16 cm. 25 cm. Day length has been widely studied because its control is so easy under experi- mental conditions, but it is only one factor among many which affect the effi- ciency and consequently the behaviour of bluegrass. Over its full range, tempera- ture can exert just as profound an influence. Thus Darrow (1939) has found that "plants grown at temperatures of 59° F., 77? F., and 95? F. produced at 59° a tall succulent bushy top growth with many leaves and at 95? an erect, non- succulent, short top growth with few leaves." Brown (1939) found that under controlled conditions maximum top growth was produced at temperatures between 80 and 90° F. when plants were clipped monthly. He also found that 50 degrees was the critical mean below which very little top growth took place in established sod, but that in new seedlings production was large at an average soil temperature of only 44? In the series of experiments where bluegrass was grown in baby-bottles, the plants were first placed under temperatures ranging from 70° to 75° F. They were later moved to a room kept at 40° F. In the beginning the day length was 14 hours; the humidity was not controlled. Measurements of the growth of indi- vidual leaves were made weekly. Average growth rates and leaf lengths are recorded in Table VII. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 339 TABLE VII Month Period Growth Length of youngest per day mature leaf in cm. іп cm. _ _(Average of 12 plants for two periods) Nov 25—30 TA 7.5 30— 7 2.43 De 7-15 94 15.0 14 hrs at 70? Е. 15-25 1.88 í Av. 221.78 сш. 25— 3 2.07 27.0 3-12 1.64 І Jan 12- 3 38 | 23.0 Feb. 3- 9 "n 14 hrs. at 40? F. 9—16 .48 23.3 Av. 42 cm. 16—22 34 22- 1 17 } 15.0 Маг 1-15 47 15-22 43 9.0 " 22-29 35 12 hrs. at 40” F. 29- 5 A 7.9 o e -1 0 Apt ж: : Temperature up to 12-19 1.07 8.2 70? F. on the 17th It can be seen that the growth rate became much slower and the leaves grad- ually much shorter after the plants were placed under the low temperatures. In general, the growing period of a given leaf remained about the same and conse- quently initiation of new leaves was not greatly retarded. Tillers were not found on any of the plants while they were kept at the high temperatures, but when plants were transferred to low temperatures tillers soon appeared and the léavés became a deeper green. These are changes very similar to those occasioned by growing bluegrass under long and then short day-lengths. Five weeks before the end of the experiment the day length was changed to 12 hours while the tempera- ture remained at 40? F. This change had only a slight effect on succeeding growth rates. During the final week, through failure of the cooling equipment, the temperature went up to 70? F., and did not come down for several days. This resulted in a striking increase in growth rate for that week. Growth Rates in tbe Field.—Approximate growth rates of bluegrass leaves under natural conditions throughout the year were calculated from two sources of data. For vegetative leaves appearing in March, April, and May, data collected in connection with the development of the tillers on the flowering shoot have been used. Since all tillers produce very short winter leaves, and then begin with surprising coordination to produce longer spring leaves about the same time, and at the same rate, successive leaves fall into definite length classes which remain fairly distinct for the first 4 or 5 leaves of new growth. It is frcm averages of 10 leaves of the same class that the curves are made. Dissection of the shoot was [Vor. 38 340 ANNALS OF THE MISSOURI BOTANICAL GARDEN £ 30 з E 9 20 z APR.24 MAY II JN7 JN.22 JY 6 Y. SEPT.23 OCT. 2 Growth rates of bluegrass leaves at various times of the year. (Data for leaves 5—12 from 444 1949). necessary in order to measure growth of the young leaves. This method has the advantage over measurement of individual leaves on the same plant, that curves on the entire leaf can be obtained (text-fig. 28, leaves 1—4). Growth rates for summer and fall leaves (text-fig. 28, leaves 5—12) were drawn from data in Evans (1949). He recorded the date on which each new leaf appeared above the previous sheath on three individual plants. Each of these plants happened to produce 14 leaves during a period of one year so that an average date of appearance could be figured, and from that the interval between the appearance of successive leaves. This recorded interval was not the elongating period of a specific leaf, however, because the first observation that a leaf was growing came after it had already arisen. from the growing point and extended itself up through the previous sheath past the ligule. About the first fifth of the growth of the blade thus takes place before its existence is recognized. The remaining % then goes on to mature. Then there is a delay until the next leaf is recorded, because it has to make the М of its growth up through the sheaths just as did the previous leaf. This growth begins when the previous blade matures. Not until the new leaf is exserted does the interval recorded by Evans end. His average interval between the appearance of each new leaf is therefore a composite figure, and instead of pertaining to the whole growth of one leaf it includes 44 of the growth of one blade and 1% of the growth of the succeeding blade. The exact proportions will vary somewhat with variation in sheath and blade length. Evans has included the average final length of each blade in his data. If we use this information carefully we can arrive at an approximate blade growth rate for the intervals of time between leaf appearance. The recorded interval pertains to parts of two blades, 44 of pui V5 of the next, but the recorded lengths are for one specific leaf. If we take % of the length of one leaf and V4 of the length of the next we shall have the «бы growth which occurred during a known period 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 341 of time and therefore be able to compute a growth rate. This has been done for the summer and fall leaves measured by Evans, and these rates are plotted in text- fig. 28, leaves 5-10. Since the fall leaf blades are proportionately shorter than their enclosing sheaths, the ratio of observed to hidden growth would change, and would be closer to 34 visible and 1⁄4 within the sheath. This correction has been applied to the last two leaves. The following shows the way in which these rates were computed: If we assume that one leaf blade when full grown was 5 inches and the next one 3 inches and the time interval between the appearance of each one at the orifice of the sheath was 10 days, the actual growth during that time must have been 45 of 5 inches (4 inches) plus 15 of 3 inches (.6 inches). Total growth would be 4.6 inches in a period of 10 days, which is a rate of .46 inches per day. There will always be some error involved in such a method but the results are so compatible between leaves measured for this study and those measured by Evans that they are con- sidered to be reasonably accurate for the present purpose. They do not agree with the data obtained in the nutrient solution experiments, where the growth rate was somewhat more than twice the natural rate at comparable temperatures. This discrepancy is probably related to the fairly high nitrogen concentration in the nutrient plus the low intensity of the light source. The complete data of text- fig. 28 do not necessarily hold for any other localities than those where the measured plants grew (Missouri and Ohio) nor to any other years than 1939 and 1949. Periods of Rapid Growtb.—Growth rate is slow in late fall, very slow in mid- winter, and sometimes rather rapid, sometimes slow іп mid-summer. Most rapid growth takes place in early spring and in early fall. The spring and fall seasons have much in common for they represent a time of vigorous growth at the expense of the plant’s reserves. It was first realized how definite and how similar these two periods were when rhizomes were being collected for the culture experiments described in an earlier section. Rhizomes needed in early October and March and April could never be found. They had all turned up in a flush of green growth which had involved the whole plant. At such times there is a tendency for sods to weaken and fall apart as the reserves are drawn out of old rhizomes and roots. New roots develop rapidly at these times and are short, thick, and white, in con- trast to the long, thin, brownish ones of summer. The period of rapid growth in the spring lasts from March to late April, while the fall rush is limited to a few weeks and depends to a great extent on when fall rains come. Recovery in each case is marked by the beginning of new rhizome production, but, as indicated elsewhere, the late fall rhizomes are shorter than those of summer. Though these periods have much in common, they are neces- sarily different, for while the carbohydrate reserves of the spring plant are greater to begin with, the progressive increase in day length and higher temperatures, to- gether with a supply of nitrogen accumulated in the soil during cool periods, favors a prolonged period of rapid growth. During fall, on the other hand, reserves are [Vor. 38 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN 40 м ы a o BLADE LENGTH (CM.) N o M AP MY JN. JY, AG. SEP ОСТ. NOV DEC. JAN. MAR. m о 8 15 и 8 l6 l8 27 27 24 5 - 29. Prevailing leaf-length through a period of a year under four conditions. Undisturbed grass was in an old meadow. A plot of this grass was burned in early February. Another plot was cleaned of all surface material with a sharp hoe at the ame time of year. The grazed grass was in a heavily grazed barnyard area immedi- ately adjacent to the meadow. low following the heat of midsummer, and day length and temperatures favor a rapid accumulation of carbohydrates which is associated with slow growth. Also during summer there is apt to be considerably less accumulation of nitrogen in the soil due to bacterial fixation. In addition to day length and temperature, there are other factors which exert considerable influence on leaf growth, either directly or indirectly. Shade increases leaf length (Watkins, 1940), though not indefinitely, since increase in lengt ultimately depends on amount of reserves available, Nitrogen fertilization makes leaves longer, the extent of lengthening depending on a complex of factors, among them especially time of application, type of nitrate ion (МН, or NOs) and pre- vailing pH of the growing medium (Darrow, 1939). Availability of water plays a part, especially in fall and winter. It has been mentioned that plants growing in soils which are prevented from freezing in winter but which are provided with adequate moisture produce very vigorous thick 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 343 rhizomes. Plants arising from these rhizomes develop remarkably fast in spring and produce exceptionally vigorous and tall plants. The relationship between fall mowing and grazing and the production of many tillers and consequent shortening of leaves has been discussed elsewhere. Denudation of soil during the period of late winter by skinning all grass from the surface of the frozen soil has been found to have a very strong effect on resulting growth of grass plants. Throughout the succeeding year leaves remain short and narrow, and lack the necessary vigor to compete with weeds. Recovery of such skinned plots is very slow. А similar effect results from skinning in August. In text-fig. 29 a complete record of bluegrass leaf lengths throughout the year in several habitats is presented. The leaf measured was the last mature one on any given date, and 10 individual leaves were measured in each habitat. A certain amount of lag existed between the actual time of leaf growth and the time when the leaf became mature enough to measure, so that the curves do not express exactly the correlation with day length and temperature that they should. They serve, however, to give comparative results and to show the pattern of effective leaf length at any time of year. Text-figure 28 gives a rough idea of the number of leaves produced during a year, though some allowance must be made for the fact that one or two winter leaves are not included. Leaf production varied from one every 10 or 14 days in fall and spring to one every 6 to 10 weeks in late fall and winter. Leaf production varies greatly with habitat. On a group of plants from different locations ex- amined at the end of August, 1949, the number of leaves produced above the last intravaginal shoot of the previous winter was as follows: TABLE VIII NUMBER OF LEAVES PRODUCED ON PLANTS FROM VARIOUS HABITATS, COUNTED FROM LAST INTRAVAGINAL SHOOT TO LAST MATURE LEAF ON AUGUST 30, 1949 Closely Old Dry sterile Dry shady grazed meadow meadow hill 11 10 9 5 10 11 3 6 10 12 7 5 11 7 8 2 14 9 8 2 16 9 8 6 10 9 8 > 12 12 10 7 12 14 10 5 10 9 9 6 Av. 11.6 10.2 8.6 3.2 Watkins (1940) has indicated that nitrogen fertilization increases the rate of leaf initiation in Bromus inermis. The number of leaves which are green at any one time also varies. Thus, on March 8, 1949, plants growing in a seepy place showed development of 4 or 5 green [Vor. 38 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN winter leaves while on adjacent pasture areas where the ground had remained frozen and dry most of the winter hardly any shoots had more than 2 green leaves. Evans' (1949) table showing the number of green leaves per shoot from the spring of 1939, when growth began, until the spring of 1940, when an inflorescence was produced, is presented below: TABLE IX AVERAGE NUMBER OF GREEN LEAVES PER SHOOT IN NON-FERTILIZED PLOTS, N 7 SHOOTS, EACH OF WHICH HAD A TOTAL OF EITHER 17 OR 18 LEAVES Average number of green leaves Year Month Entirely Partially green green Total 1939 May 18 2.0 .6 2.6 June 2 1.9 7 2.6 June 17 1:3 1.4 2.7 uly 1.7 1.9 3.6 uly 1 2.0 2.0 4.0 August 1 1.9 2.6 4.5 ugust 2.0 2.7 4.7 September 3 1.7 241 3.8 September 18 2.0 2.3 4.3 October 1.6 23 3.9 October 18 1.9 2.0 3.9 November 7-14 17 15 3.6 | 1940 April 1—2 1.1 1.6 57 April 15-16 2.1 1.6 3.7 ау 4 3.0 1.1 4.1 May 17 2.6 1.4 4.0 June 3 1.3 .9 2.2 June 17 0 42 3 Midsummer Depression.—Shortage of water is probably partly responsible for periods of slow growth of leaves during midsummer. Brown (1943) has shown that irrigation during periods of drought or low soil moisture helped considerably to increase yields, Ahlgren (1938) stated that moisture, more than any other factor, limited forage production. While summer irrigation may maintain yields, summer growth is made at the expense of stored reserves and may reduce the competitive ability of bluegrass against weeds (Brown, 1943). Irrigation during midsummer is probably less important than during a dry fall. Brown found fall droughts to have a decided adverse effect on bluegrass yields. Irrigation during fall and early winter is not commonly practiced, but the indications are that it would encourage root growth and tillering and thus greatly aid the grass in its competition with annual weeds. Numerous studies have shown that peak yields of forage occur twice during the year, usually in late spring and in late August or early September (Brown, 1943). Examination of series of leaves produced by various individual plants 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 345 show that usually the midsummer leaf is slightly shorter than those immediately preceding or following it (text-fig. 29). This may be connected with the period of maximum summer temperatures which commonly occur in July. Both Harrison (1934) and Darrow (1939) have shown shortness of leaf to result at temperatures ranging between 95 and 100° F. Since irrigation did not completely eliminate a midsummer depression of yield, Brown (1939) attributed part of the reduction to supra-optimal temperatures. Whether these midsummer depressions of forage and of leaf length are interrelated cannot be said. The spring peak can easily be understood as being a combination of factors including inflorescence development, high available nitrogen, and plenty of water, but the late August peak is more difficult to explain, unless we merely assume it indicates a return of more nearly optimum conditions. BUDS ON THE CROWN The Number of Buds Produced.—In a given habitat individual bluegrass plants will produce new leaves at almost the same rate, and at the end of the year will all have about the same number of leaves (table VII). In a meadow this number averages about 12-14; in a very dry shady locality, 7-9; while in a well-watered pasture it may well be as high as 18. Since under ordinary growing conditions only a single leaf blade grows at once, the longer the leaf the fewer will be the number of leaves in a given season. This accounts in part for the low number of leaves in dry shady places, for there the leaves are very long and growth is slow. In grazed places, leaves are short and the soil usually well fertilized so that growth is rapid. Consequently more leaves than normal appear. Such factors as this play an important role in the ultimate appearance and behaviour of a plant. The number of leaves reflects the number of phytomers formed during a given period and consequently the number of buds available for rhizomes and tillers. It is very uncommon to find that all the buds on a crown have matured into new shoots, for the buds of midsummer phytomers frequently remain dormant in- definitely, thus limiting branching to the fall and winter and early spring buds. This is especially true in the case of old meadow plants. A Conservative Crown.—Instead of speaking in abstract terms, it would be well to examine some actual plants. Text-figure 30 shows a diagram of an ex- tremely conservative three-year-old specimen from an old meadow in early Novem- ber of 1948. A photograph of the basal part of a very similar plant is shown in pl. 9, fig. 30, although only a single year's growth is included. It can be seen that in both plants only the bare minimum of buds have developed into new shoots. In text-fig. 30 two generations of inflorescences are shown, one for 1948 (I 48) and one for 1947 (1 47). On the basis of these facts, it is simple enough to put to- gether the history of this plant. It had its beginning probably as an intravaginal shoot from some plant now long since dead. That was in the fall of 1945. During the summer of 1946 no buds developed into rhizomes, but in the fall three intra- vaginal shoots developed from the last three buds of the year, the 14th, 15th, and [Vor. 38 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN 72% UPPER LEFT 147 SPRING,48 DEAD Я 99 кент І ” SUMMER, 47 ! = | RHIZOME ХІ l SUMMER, 48 FALL, 46 ЎҒАСІ., 47 SUMMER, 48 SHEATH mm 14 LOWER LEFT DT эшип, BLADE CROWN FALL, 45 Fig. 30. Diagram of a conservative old meadow bluegrass plant. Very little branching has occurred during its three years of growth. There are few tillers, few rhizomes and most of the growing points have terminated in inflorescences. Sheath and blade, X 14; crown, X 3%, léth. Buds 17 and 18 which developed after the intravaginal shooting occurred should properly be considered part of the following year's growth. This main stem ended its terminal growth by developing into an inflorescence in the summer of 1947. If we now follow the destiny of its three shoots we find that each of them pursued a somewhat similar course, at least to the extent that in 1948 they all terminated in a flowering shoot. There were, however, some interesting differences. The last shoot (text-fig. 30, upper left) did not produce any branches from its 13 buds in 1947 and thus died. The next to last shoot (right) produced 13 buds also, the last two of which developed into tillers in the fall of 1947. One of these shoots died at an early stage. In the meantime the first original shoot (lower left) produced only 10 buds of which one developed into an intravaginal shoot in the fall of 1947, and another produced a rhizome in the following summer. It will be noticed that this 1948 rhizome developed from a bud just a little below the tiller of the fall of 1947; this is the usual place of origin of most summer rhizomes when only a few develop. This rhizome is easily identified as having been produced in summer because of its length and large number of internodes. 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 347 If we follow the history of this plant into the third (1948) generation, we find that there are only three living descendants, two shoots and a rhizome. The surviving tiller on the right had produced 10 buds in 1948 up to the time it was dug up. The tiller on the left side produced only 6 buds. If we assume that only one more bud would have been added during the remainder of 1948, then the following numbers of buds were produced in each of the three years on the respective shoots: Upper left Right Lower left Main shoot, 1946 16 Tillers, 947 13 13 10 Tillers, 1948 (died) 11 7 These data strongly suggest that the longer a plant lives by intravaginal shooting only, the fewer leaves are produced each year. It is also striking that over a period of three years one growing point has only increased to three new growing points, of which one is a rhizome, and two are relatively unproductive tillers. The existence of such a conservative plant is a very precarious one. It rep- resents the bluegrass plant at its minimum, with 1 rhizome, 4 inflorescences, and 6 intravaginal shoots being produced in a little less than three years out of a total of 74 buds. A Vigorous Specimen.—In decided contrast to this old meadow bluegrass plant is the closely grazed pasture plant shown in text-fig. 31, which was also dug up in early November, 1948. Here the basic seasonal pattern is all but obscured. The main stem of 16 buds is much exaggerated in the drawing so that all the branching could be included. This stem developed during the year of 1947. In the fall of that year it gave rise to six branches, of which two have been broken off. Of the remaining four, the lower two, arising from buds 8 and 10, became short rhizomes, and the upper two, from buds 15 and 16, are intravaginal shoots. This main stem terminated its growth by flowering in 1948. During the spring and summer of 1948, when livestock were put on the pasture, considerable late- summer rhizome development resulted (branches A-H) and some fall rhizomes are present (I-K). The two can be distinguished because fall tillers have appeared only on the summer rhizomes; also the fall rhizomes have only one or two leaves. L and M are intermediate types. When the plant was dug up it was busy producing tillers in the axils of new fall leaves. Some branches show as many as four such tillers, and in branch F some of the tillers have already produced secondary shoots. Figure 31 (pl. 9) is a photograph of a crown region from a plant which was similar in all respects to the one shown in text-fig. 31. It bears an especially close similarity to the shoot just to the left of the 1948 inflorescence. [Vor. 38 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMER 48 WINTER 47-48 FALL 47 SUMMER 48 LATE SUMMER 47 чөй a RHIZOME атал | E AF ааа OR OWN —— NEW TILLERS SUMMER 47 Fig. 31. Diagram of a vigorous pasture plant. Considerable development of both tillers an rhizomes has taken place during a year and a half of growth. Rhizome, X 45; leaf, X 4; crown, х 3%. To recapitulate, from a single plant in 1947, six shoots developed, two of which were broken off. Summer and fall rhizoming in 1948 increased the number of growing points by 13, and intravaginal shooting added 28 more to make a total of 45 active growing points out of a total of 138 buds. The net results of less than two years growth were: 15 rhizomes, 30 shoots and 1 inflorescence. A Hypothetical Individual —After examining these typical plants we may be a little closer to discerning the basic bluegrass theme together with its variations. Perhaps we can now draw a hypothetical average plant if we limit ourselves to generalities. Let us begin with a rhizome which appears at the surface of the soil 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 349 sometime in late winter or early spring as rhizomes frequently do (text-fig. 32 A-D). Such a plant will not develop any tillers. Instead, its winter and early spring buds will lie dormant while rapid leaf growth takes place. About mid- April these buds will start to swell and develop into new rhizomes. The first new rhizome will probably be short; the next ones longer; and some may grow under- ground for several summer months. Most of the winter and spring buds will be thus consumed in the production of rhizomes, while the late spring and early summer buds of the parent plant are gradually suppressed so that no new branch- ing takes place. The suppression of these buds is probably related to the cessation of root initiation which Sprague (1933) has shown to take place early in May. Nishimura (1923) has indicated that no vegetative bud elongates until a root is produced. If this is true, summer dormancy is inevitable. The sprout-like rhizomes previously mentioned as appearing in late August under rank meadow conditions would be most apt to occur on old crowns, especially on those which bloomed in spring. Rhizomes of the year, such as the one we are describing, would not commonly show them. The change which comes over bluegrass in early fall brings with it shorter, greener leaves and white, short roots. The late summer buds, instead of remaining small, discolored and dormant, become increasingly white and well developed, but they do not usually produce rhizomes immediately. Instead, they are transitional in nature and anticipate the appearance of tillers from fall buds (pl. 9, fig. 32). Short fall rhizomes may, however, appear after several tillers have begun to func- tion efficiently. Frequently no rhizoming at all takes place in the autumn, and the buds are thus conserved for a rapid development of rhizomes the following spring about the time the inflorescences begin to appear. There is a conflict between rhizome and shoots for the buds of fall. Since the intravaginal shoot arises almost as soon as the bud is formed it has first choice. If conditions are right, practically all fall buds may become tillers, and thus only one or two buds will remain on the main stem from which rhizomes may appear the following spring. Where only the latest fall buds develop into tillers there is room for three or four rhizomes preceding them. If we pull these plants from the soil in late May, the former will be found to have many short leaves, and, at the most, one rhizome, and the latter will have a few long leaves and quite a few rhizomes. All rhizomes, however, do not turn up in early spring as did our hypothetical one. Some turn up throughout the summer, others in the fall. A rhizome turned up in fall is shown in text-fig. 33 А-С, for the sake of contrast. This plant begins life by tillering instead of rhizoming. Every bud at first is taken up by tillers, with the frequent exception of the lowest bud on the crown which usually remains available for production of a rhizome in the spring. In a pasture grazed closely the previous fall there are many plants of this type to be found the following spring. [Vor. 38 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN z Fig. A, a rhizome turning up in the spring will look like this in the f Summer 2 have turned up and formed new plants. А single short fall ciel is developing, X l4. B, the crown of this same plant, showing upper Ба те ing into tillers, Х 2%. С, the crown in the spring just after having turned u x 2%. D, above, the Pi. -up rhizo 3 А, a spring plant derived іші fall rhizome, with three tillers = short winter poly X М. B, the wn in spring, with only a sin ngle rhizome the base, the remaining buds лүе ris tillers, X 2. E the crown in fall ice after having turned up, X2 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 351 Some Practical Considerations —While the seasonal effect is more or less the same, the record of events as recorded by various generations of rhizomes will be slightly different. The proportion of rhizomes of a given vintage in a sod or a nursery plot may change considerably depending on the age of the stand, the treat- ment it receives, or on the weather. Evans and Ely (1935) have presented figures which show the extent of such variation within two successive years: TABLE X AVERAGE NUMBER OF NEW RHIZOMES PER PLANT IN DIFFERENT a. E 1932 AN N PLANTS TRANSPLANTED ON MAY 18, 1931, TO CULTIVA ROW PLATS. (THE PLANTS WERE NOT CLIPPED AT ALL Month 1952 1933 April 0.0 May 0.3 June 15.5 553 July 18.3 2.0 Augu 353 21.0 September 0.3 6.0 October 2.5 0.5 November 3.0 Wherever studies of bluegrass are made, the sequence of events and their pos- sible variation must be kept in mind. Undoubtedly, elaborate quantitative tests of rhizome production, top production, or other aspects of the plants’ growth have failed to obtain consistent or statistically significant results because of the failure to reckon with the flexible nature of the species. In pot experiments with nutrient solution it is doubly important to know the kind of material being grown in the pots. Such factors as the number of buds available on a transplanted rhizome or crown, the time of year such buds developed, the length of the associated leaf, and whether the rhizome is from a turf or a nursery row will all have repercussions on the results. DESIGNING A BLUEGRASS PLANT THE SPECIFICATIONS How we manage bluegrass will depend on how we intend to use it. Bluegrass is called upon to serve many purposes, including the following: Pasture Range ' Meadow 27” Turf Alternating Winter Hay Sod run-off strips Lawns Continuous Summer Silage Terrace plantings Greens Dairy Breeding Roadsides Fairways Fattening Playing fields Subsistence Landing fields With adequate study of each environment and of the factors involved in each type of use, a blueprint could be drawn showing the specifications of the plant needed. There would always be some conflicts. It might be a simple matter to [Vor. 38 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN design a plant for yield in one year, but consideration would have to be given the repercussions of management techniques on future production. There is a strong likelihood that the most efficient treatment of a bluegrass stand might involve making different use of it in successive years. Under some conditions this is not feasible so a compromise must be made. THE MATERIALS AND THE LABOR We have found that the bluegrass plant is a flexible collection of buds whose destiny, while in part controlled by inaccessible aspects of its environment, can also be molded by specific practices. Once the requirements are known, we can choose from the following alternatives: 1. Many vs. Few Buds.—The highest production of buds comes with short leaves, moist, nitrogen-rich soil, and sunny conditions. Where intensive manage- ment is necessary, usually the more buds the better. If long leaves are desirable then some sacrifice of buds is inevitable. 2. Dormant vs. Sprouted Buds.—Greatest sprouting of buds occurs in good soil with plenty of aeration and little competition, and seems to be aided by high organic-matter content and adequate water supply. (It is just such requirements which Percival (1921) specifies as necessary for maximum development of tillers in the wheat plant.) It is generally desirable to have a large proportion of the buds on a plant develop. This can be carried to extremes, however, as discussed in the next section. If the need is for a tight sod, then competition among indi- viduals will necessarily follow, and some loss of sprouting ability will be encountered. 3. Rhizomes vs. Tillers.—Tiller production is encouraged by provision of good soil conditions as described above. Treatments which specifically favor tillers include fall grazing or mowing, fall nitrogen fertilization, removal of all shading growth and dead weeds, and in dry falls the addition of moisture. Maximum tillering may not always be desirable. It can result in what one might call an excessive investment in growing points, considering the amount of capital (soil, oxygen, water, and nutrients) which the plant has within reach. Ideal conditions are necessary to support large numbers of closely packed shoots even though they are capable of forming their own adventitious roots. When such plants are sub- jected to excessive mowing or grazing or treading their reserves can be very rapidly depleted. This is especially true since strongly tillering plants seldom have very extensive underground parts. In general, it should be stressed that if shooting is to be encouraged in fall, then adequate provision should be made to provide opti- mum conditions for growth in the following spring, especially through nitrogen and potassium fertilization. Rhizome development depends in part on how extensive the fall tillering process has been. If most of the fall and winter buds have been used up, rhizomes are apt to be uncommon. The same ideal soil conditions which encourage sprouting of other buds favor the production of rhizomes. Plants which grow in open soil 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 353 as spaced seedlings or shoots in nursery plots are abnormally productive. Balanced fertilization in winter, spring, or summer favors increased rhizoming. Phillippe (1943) found that in nutrient culture, maximum rhizome production occurred under conditions of moderate nitrogen concentration (17 to 57 PPM) and by fairly high potassium concentrations (41 to 162 PPM). High nitrogen content has been found by numerous investigators to reduce rhizome production in pot and nutrient experiments. Brown (1943) has stated that summer irrigation leads to the death of many summer rhizomes, and Harrison (1934) found combinations of close clipping and temperatures around 100? F. to be fatal. This coincides with observations of Wilkins (1935) that over-grazing during drought periods when temperatures were around 100? F. reduced bluegrass recovery the following fall by 75—99 per cent. To some extent rhizome production is self-regulatory. Usually whenever high production is a possibility, as in new seedlings or broken sods, some benefit may be derived from much rhizome growth. When sods are tight, then fewer buds sprout. Under an old sod, rhizome production apparently reaches an approximate equilib- rium with the available soil nutrients, especially nitrogen, There is adequate de- composition of a certain amount of dead rhizome matter plus just sufficiently vigorous growth of the plant itself, so that no more rhizomes will be added to the soil that can be broken down the following year. Some fluctuations in this cycle will occur with weather variations. A certain amount of rhizome production is essential under most conditions because intravaginal shoots seldom persist for more than a year. It is also important, in view of the many advantages of rhizome herbage production, such as the broader, more vigorous leaves, more even distribution of roots, growing point well below the soil, shorter sheaths, and leaves close to the soil, that plants are not completely defoliated by grazing. There is little doubt, however, that too high a proportion of rhizomes in the stand may lead to low herbage productivity, for they respire and consume soil oxygen, and sooner or later die, and the micro- organisms which break them down compete with the grass itself for soil nutrients. This is especially true since rhizomes are high in carbohydrates and low in nitrogen. It is commonly found that where fields or plats are mowed during the summer for a period of years, total forage production falls off. Ahlgren (1938) states: “The data show that with the exception of 1933 (the second year) there was a progressive and significant decrease in the yield of bluegrass on all plats, regardless of the cutting or fertilizer treatment used during the period 1952-1936 inclusive." His data follow: [Vor. 38 354 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4-inch cut Hay cut Year No fertilizer Fertilizer No fertilizer Fertilizer 1932 1,619 3,113 2,050 3,957 1933 2,153 4,295 2,977 4,579 1934 1,817 2,114 1,681 2,046 1935 1,520 1,932 1,279 1,785 1936 82 1,703 5 1,617 test was begun оп The first cuts were made as above, to a level of 112". Sub- жашт сше were made to pc revel when ө grass reached a height of 4—5”. Figures аге pounds per acre.) The cause of this progressive decrease in production has not been adequately explained, but there is a strong possibility that the accumulation of rhizomes in the soil may be involved. That such accumulations occur are well shown by data from Brown (1943): Herbage yields Root Yields Rhizome Yields Year 1” 2 14” Hay 1” 2 ы Нау 1” 2 4” Нау 1937 2833 2254 5857 1927 2049 1922 122 122 108 1938 508 310 1280 1897 2137 2210 372 478 505 193 141 501 1979 2289 2281 398 519 507 19407 1286 900 1601 1946 2088 2455 660 817 1071 +NaNOsz added 100 lbs. per еи анне 5 and semi-monthly at гасе of 50 lbs. per acre from April to June 15 and from Aug. о Sept. 4 These test ced were seeded in Pong 1936. Figures are lbs./acre. 1%- and ix %- NM. cuts were made semi-monthly. Hay cuts made at full bloom and at end of growing seaso The same progressive decrease in yields of herbage are apparent heré. In both of these experiments, the removal of clippings from the plats probably exerted some influence on total yields. It is clear, however, that reduction in yield paral- led an accumulation of rhizomes. Brown has stated that death of rhizomes began in the second year, and in the fall of the third year there were so many dead ones that they were separated from the live ones and not included in the data. There- fore the figures above do not show the total accumulation of rhizome material in the soil. The data would have been even more striking had all dead material been included. Typically, maximum herbage yields were secured the first year after seeding. In that year the rhizome dry matter in the soil was less than 5 per cent of the total dry weight of the herbage and rhizomes together. By the third year herbage yields had dropped from eight to sixteen times, but rhizome weight had increased from three to five times. Rhizomes had come to make up 50 per cent or more of the total 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 355 dry weight exclusive of roots (which had a remarkably constant dry weight throughout the tests). Significantly, the least amount of rhizome dry weight was produced when the grass was mowed semi-monthly to a 1-inch level. Most of this difference probably involved rhizome length rather than number. In the fourth year heavy nitrogen fertilization in spring and summer increased herbage yields from three to six times but at the same time the rhizome dry matter was nearly doubled. Addition of nitrogen in both Ahlgren’s and Brown’s tests increased yields but did not completely restore productivity to its former level. According to the data above, rhizome production was greatly increased by fertiliza- tion in spring and summer, and the sod-bound condition could be expected to become worse the following year. Moderate mowing of pastures in early spring through summer apparently will increase the rhizomes in the soil in proportion to the amount of tops above ground, even when mowing may be semi-weekly to a height of 1 inch. Increase of rhizomes goes on until the stand is almost devoid of herbage value. Rhizomes do eventually turn up and add to the herbage total, but there is considerable delay and wasted energy in getting top production through rhizomatous plants alone. Not only that, but at the most an ordinary plant cannot be expected to bear more than four or five rhizomes while its potential of new tillers is considerably greater when properly managed. The 24-inch height such as that advised for most lawn mowing resulted, in Brown's experiments, іп the highest proportion of rhizomes to tops. ‘This would give a tough sod, but relatively little top growth, unless some counteracting practice designed to stimulate intravaginal shooting and long leaf growth were used. Kentucky bluegrass is really а plant with a split personality. It is trying to be rhizomatous, as is its relative, Canada bluegrass, and upright, as is Poa trivialis. It actually can assume either role, and it is up to the pasture or turf manager to decide which personality or complex he prefers. It is not surprising that to some extent bluegrass has lost some ground in the field of intensive pasturing to non-rhizomatous grasses and legumes, for these species do not waste energy and nitrogen on non-leafy production. Admittedly, bluegrass cannot compete with such productive non-rhizomatous types as alfalfa, perennial ryegrass, orchard grass, and ladino clover, but its quick adaptability and persistent nature are indispensable attributes under many conditions. To make maximum use of the species, however, close attention must be paid to its seasonal rhythm and its system of bud economy. 4. Long vs. Short Rhizomes.—Summer rhizomes are long, and fall rhizomes are short, but close and continuous grazing leads to short rhizomes at any time. In pot cultures Harrison (1934) found that high nitrogen content encourages short rhizomes. Length of internode has little or no relation to ultimate length of the rhizome. Dry periods shorten internodes but variation in nutrient balances has been shown by Phillippe (1943) to have no significant effect. [Vor. 38 356 ANNALS OF THE MISSOURI BOTANICAL GARDEN Short rhizomes are desirable in some cases, long ones in others. If a tight sod is required, rhizomes of moderate length must be encouraged. Stands which have only short and shallow rhizomes will have little protection under very hot and dry conditions. If properly managed, however, short as opposed to long rhizomes could be counted on to increase the efficiency of a pasture or lawn. 5. Branched vs. Unbranched Rhizomes.—Branching of rhizomes is encouraged by those conditions which favor the development and sprouting of buds. Damage to the initial growing point stimulates axillary buds to develop, and wherever summer sods are burned or mistreated, or submerged by dense weed growth or by alluvium, the ability of rhizomes to sprout in early fall from their axillary buds is of great importance to the survival of the plant. The vigor of the rhizome deter- mines how many buds develop, and whether they will form strong rhizome-like branches or merely delicate vertical shoots. For those strains of grass which are primarily rhizomatous, thickness of the stand is almost completely dependent on the ability of the rhizome to produce branches. 6. Vegetative vs. Reproductive Sboots.—The number of reproductive shoots produced in May depends on two things, the number of plants which are adequately mature and vigorous enough to undergo flower initiation in the previous fall, and the number of tillers which these plants produce. Plants which are mature and vigorous in fall will have come largely from rhizomes turned up during the previous fall, spring, and early summer. Some perennial tillers will contribute to the cause. Ultimately then, to influence the potential seed crop it is necessary to start work a year and a half ahead of harvest time. Encouragement of rhizomes, followed by encouragement of tillering, followed by encouragement of long leaves and full panicles as described below, should give maximum seed yields. This would leave the soil full of old rhizomes, and unless steps were promptly taken to deplete their reserve carbohydrates through grazing and encouragement of tillering the follow- ing fall, subsequent yields would suffer. A complete management plan would be more complex than this, but these are the main considerations. Such a plan might also provide high yields of hay or silage, but maximum development of inflorescence may not always be desirable. Limitation of flowering can be accomplished through heavy grazing in late summer and fall, and by grazing or mowing in spring when the panicles are exserting. Encouragement of a vegetative condition in bluegrass holds back the spring flush of growth which accompanies the development of the flowering shoot. It also holds the protein content of the grass at a fairly high level, while the carbohydrates are kept low (Woodman et al., 1928; Hein, 1937). This would limit the gaining of weight by fattening animals in preparation for fall markets or over-wintering. It would, on the other hand, encourage summer milk production in a dairy or breeding herd. 7. Sbort and Wide vs. Long and Narrow Leaves.—Short, wide leaves and ex- cessive tillering often go together. Long leaves are encouraged by winter and spring fertilization. When heavily tillered plants are fertilized in winter and spring a very 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 357 dense, moderately long and wide-leaved blue-green plant is produced which is highly productive. It combines the advantages of the other types. Constant grazing or mowing keeps grass leaves short, probably for two main reasons. First, without the over-burden of shading tops, the blades reach the light rapidly and do not become attenuated. Second, such treatment encourages the production of many short rhizomes the early leaves of which are commonly some- what abbreviated. Infrequent mowing or grazing has little effect on leaf length. While long-leaved plants are commonly considered more productive of herbage, the fact remains that short, wide-leaved foliage is more palatable to livestock. They will barely consider the rank growth of mid-summer meadows. 8. Heavy Seed Production vs. Little Seed.-—There is a complex relationship between number of panicles and number of seed produced. In general, number of panicles is increased by mowing or high nitrogen fertilization in fall (Spencer et al., 1949; Nillson-Leissner, 1937). This treatment, however, usually reduces the panicle length and the culm length and reduces the number of seed set per panicle when compared to spring fertilization (Phillippe, 1943; Spencer et al., 1949; Nill- son-Leissner, 1937). Winter or very early spring fertilization tends to give tall panicles and inflorescences, but does not increase the number significantly. Seed production is generally good. Late spring applications in April, when the panicle is growing, tends to encourage leaf growth rather than inflorescence development, and results in a weakening of the culm and reduction in quality of the seed. 9. Sod vs. Open Growth.—Bluegrass, like most other plants, grows most pro- fusely when grown alone. When it is allowed to sod over, then we inevitably lose production, whether it be of forage or seed. In this respect it is significant that bluegrass plants growing in alluvium produce by far the greatest amount of seed per panicle, bear a goodly number of panicles, have large seeds, and long and wide leaves. A year-round supply of water, rich soil, and sun combine to produce maximum yields. Inevitably in a sod we have to sacrifice some greenness, some vigor, some thickness of stand. From a productivity standpoint the best that can be done is to provide management practices which make the sod-grass think it is down by the river. Discing and harrowing aid in this simulation by ‘providing loosened and bare soil areas. To avoid weediness treatment should be done only in fairly late fall when tillering, rooting, and fall rhizoming together can recover the vacant areas while weed growth is at a minimum. Another way of getting the productivity of the river bottom up on the pasture is by using alluvial ecotypes for seed. This process is going on more or less unconsciously in many grass-breeding experiments. So-called high-producing strains of many species invariably bear a striking resemblance to river-bottom types. [Vor. 38 358 ANNALS OF THE MISSOURI BOTANICAL GARDEN THE Parts OF THE BLUEGRASS PLANT COMPARED? WHERE DO THEY ORIGINATE? Rhizome.—From underground, axillary, mature buds whose ық зегі ад are dead, or dying. vtri May extension type predominantly from buds of early fall a rly spring. August sprout type from spring and summer buds, and fall rhizomes from С rg "ein Crown.—From underground terminal iub f rhizomes, seedlings, or intravaginal shoots, May Meus pa above ground, especially in seedlings and shaded shoots. a oe axillary buds below or above ground o n late fall and early winter P. Cu om terminal bud of crowns of sufficient maturity. If from a rhizome, the cro will not ы ii than 6—8 leaves; if a tiller may have only 2. anicle.—Main axis or rachis is continuation of auis. "Side branches arise as secondary, tertiary, and quarternary нме са this rachis. Spikelet.—Some time in April. WHEN DO THEY BEGIN TO FORM? izome. a nerui psi ү concomitant with floweri ring, but to some extent throughout early summer; also in fall, early or late, depending on condition of grass. Sprout type in late August or early sees m on ne meadow type or burned or i areas. In ыга grazed d soi t p inter, n.—When rhizome turns up, especially late summer and early fall, ved каш fall and early ун From tillers іп late fall and winter and from seedlings in fall and s I тагу fall to late winter, principally late fall. M Hime in late fall and winter following initiation of intravaginal shoots. One ter and early spring; may not form until early March (Musgrave, 1940 )% ergo i-i time in April. NUMBER ON A PLANT OF 16 PHYTOMERS PER YEAR Rhizome.—Varies greatly, 0-8, average probably 2, but including secondary and tertiary plants may а ыр 6. own, plus average number of turned-up rhizomes (2) plus average number E de (3, see pé gives average of 6; may be кат amplified by secondary and tertiary ing. таныса around 3, less than 1 or more than 10 infrequent; with secondary branching may reach 50 Culm.—Usually 1 on main axis and 1 on each of 1 or 2 tillers. May be up to 12 per plant with multiple peru icle Е e ре er culm; number of protuberances may vary greatly; avera age 3—5 at each node of ra E There are from 5—10 rachis nodes, therefore 15—40 branches on main axis. and — bes yide fewer. bikelet.—Average number not counted. Secondary and tertiary tillers have fewer. Secondary TO WHAT EXTENT DO THE PARTS BRANCH? pean hepa eid limited; can be exten Crown.—Branching includes rhizomes ог Кш Of 16 buds, not more than half usually Tiller.—Branches as does crown; may form rhizomes or secondary ии but only the latter in first fall and winter. Rhizomes rarely appear until following May, from any left-over buds. Culm.—Does not branch (no buds present). Panicle.—Almost all buds develop. Spikelet.—bBasal phytomers чу msi the two glumes and lowest lemma do not usually bear branches. Upper phytomers bear flor Mn ра аге а егес of information from authors previously credited, pod usd —— the co of the present study. They are in 8 variation plc not vr xn be en into consideration due to lack of data. to bluegrass growing in Missou s very and Most of (On rm irm 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 359 THE LENGTH OF THE INTERNODES Rhizome.—Highly variable in length, 1-50 mm., average 10-15. Shorter in dry soils or dry ipe Crown. by E or ое е. wit —Same as for wn. 1 —Gradationul, М first a few e last up to 30 cm. or more. Su r cale fall dm a straight j^ under normal conditioni, (See Prat, 1934.) rnodes of culm. Range from 30 to —Mi inute, somewhat less oe 1 mm.; individual internodes can be induced to elongate ccessive internodes Са on rachis, in геу onde er to 2 mm., but do not form a straight line on atlas natural or весе pee Spikelet. —Longest EN first glume, 1—3 mm.; rest are minute. THE NUMBER OF INTERNODES e.—Many in summer extensor rhizomes—may be up to 30. Fall rhizomes seldom show r6. Rbiz as many as ite erri 50 Crown.—Depends on habia and season. a Tiller.—Same as for crown, but few Millers 4. e a vibe year. If they do, there tend to be fewer phytomers sie year. Culm.— us e Panicle.—Rachis 5—10, main MES about the same, branches of lower order successively fewer. Spikelet.—Averages 6, varies from 4—10 (includes 3 sterile and 3 or more fertile). HOW FAST DO ITS INTERNODES ELONGATE? p 5 mm. per day. Qwn.—No соо Tiler —No elonga —Varies wrth pon internode number. Internode 1, 0.5 mm. per day; Internode 2 and 3, = y fo Spikelet. --Уегу little if. any elongati THE SHAPE OF THE INTERNODE; IS rt SOLID OR HOLLOW? Rhizome.—Compressed dorsoventrally, solid. ollow Panicle. үү somesbat ribbed and grooved at extremities; hollow, solid at extremities. Spikelet.—Not examine WHAT KIND OF LEAVES, IF ANY, DOES IT BEAR? Rhizome.—Cataphylls only; very short blades may appear (up to 2 or 3 mm.) without First leaves via d short blade, shorter than T later, blade is longer. Tiller. so ves as for c ‚ but first leaf is enclosed in prop Culm.—Normal leaf at (ries upper leaf has very short blade ind long sheath. Others tran- sitional. No leaf on 8 internode. Panic —No leave I Presumably glumes are sheath part of leaf; palea is prophyll of the shoot which Spike becomes the flor WHAT POSITION DOES EACH PART ASSUME? ae ee al, nd arch, almost upright, or may show some geotropism at first. cus to vertical. Ss MEC өн prostrate to erect. Culm.—Usually vertical, though under special conditions may be prostrate or procumbent Е p of rachis same as culm. Branches may be appressed to drooping, ы Рап usually at right а Spikele riae usually fairly closely appressed. [Vor. 38 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN WHAT IS THE NORMAL LIFE OF EACH PART? Rhizom —May turn up within a few days after it Ке ins to grow, or may stay әлегі налаш 60 Е ог more in summer. During cold weather rem anie nd while soil is fro —Matures with developmen of о; shoot; shortest igiri is on fall аы shoots, m 6—7 months. Rhizomes rA n up in late summer may bloom in 9 or 10 months, and intravaginal shoot which not bloom p rst year may bloom in 1 ені and 6 months. Fall dis s take г ех 8 т ған ийе spring TM 1 year and 3 mo iller.—6 im r ] yea and 6 mo Culm. абери з the унаа ети d takes vnd a short time in late fall; elongation in month spring takes about 1 month. Total life is about 6 Panicle.—5 months for elaboration of branching uid 9 or 10 days E elongation of the panicle proper, and 45 days for its complete exsertion including bald elongati ү эө Вы rst it e ied starts to form when panicle is about М inch Е At 1 inch there are at least 3 — T florets. Glumes enclose entire spi ikelet at 1% inches DAMEN 1940). This Vnd be 5—7 days. To al. maturity time, to exsertion of anthers, about 5—6 weeks. HOW FAST ARE NEW PHYTOMERS INITIATED? "see Ми averaging nearly one every two days during summ Crow giis around 13-14 a year, exclusive of er elim, Slow ii in winter and hot or dry мае ad summ rapid in spring and for a brief period in early fall. А new leaf every 15-20 days in т p early fall; every 30 to 50 days in summer and late fall; i every 60 Ti wed as xm Culm.—4 x оа peg ha culm produced some time in late fall but - not kno anicle-—Assuming 8 internodes on each rachis іп period pug to April 1, rate is about 1 phytomer every 11 days. Comparable rate on vege Hime m part is 1 about ind 60 days. Rate of branching is different, depending on strain and time of year; кый» about one new generation Spikelet.—A sp ie with 3 rudim entary dor оме develops in about 6 days (Musgrave, 1940); HE T кең ns kiat is about 1 phytomer per day THE FREEDOM OF THE PHYTOMER A grass plant is a community of phytomers. No phytomer exists autonomously, but is necessarily involved in the total effort of the plant. At the same time, in its extreme youth an individual phytomer is quite plastic and may, as it ages, develop into any one of a number of things. It may or may not take full advantage of its potentialities to produce leaf blade and sheath, internode, root, and bud. It may consist only of an internode, as in the case of the last phytomer of the culm. It may be only an internode with a branch as in the panicle. It may be an inter- node and a sheath and blade, as in the lower culm internodes, or an internode, sheath, bud, and root, as in the rhizome. Sometimes, there is no well developed internode, and only the sheath, blade, bud, and root are prominent. Rarely, an internode, sheath, blade, root, and bud are all present as in stoloniferous shoots. We can summarize the various main combinations of the five structural elements on phytomers of various parts of the plant as follows: Phytomer Internode Sheath Blade Bud Root location 4th culm 0 0 0 0 Lower culm Medium Medium-long Medium 0 0 Panicl Medium 0 0 0 Spikelet Minute Short 0 No floret 0 Crown Minute Medium Short tolong Usua Usual Rhizome Medium Medium 0 to minute Usual Usual 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 361 { = 0» Ж» T^ 4 12 2 > 2 = с) — 5 TIT DAT У № Toa a W, Ñ ак. > Me OC Y v y UP г> р n ^ 2 М 54 4 > Fig. 34. In fall and spring when vegetative shoots are covered over by alluvium or manure, or by soil excavated by gophers, the internodes on the phytomers whic e just maturing will develop long internodes such as A, B, C, and D, even though ue leaf blades of the phytomers (LL) are long; OC, the old crown, now buried; NC, the new crown Intermediate combinations of the above structures may occur under special cir- cumstances or in areas transitional between different parts of the plant. А dis- tinctly different ratio exists between the three prominent features of phytomers (blades, sheaths, and internodes) when they are located on a rhizome, flowering shoot, and vegetative crown. Under certain circumstances these ratios can be altered by subjecting the plant to changed external conditions. If the plant is shaded, internodes will elongate in spite of the presence of leaves. This occurs in vegetative shoots which are shaded by heavy mown grass or covered by soil, as often happens around gopher mounds (text-fig. 34). Where such growth follows mowing, the growing point is raised out of the ground with the result that it is easily killed by drought, cold, or trampling. A somewhat similar departure from normal behaviour can be noticed on rhizomes grown under special conditions. We have discussed the fact that rhizomes turn up when leaves begin to develop. If the rhizome is enclosed in a glass tube [Vor. 38 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4 SHEATHS 6r BLADES IN CM. LENGTH ю I INTERNODES SUCCESSIVE PHYTOMERS Fig. 35. А rhizome was prevented from turning up by directing it into a glass tube buried in vermiculite. The rhizome grew for several months, and was then removed and its phytomer parts measured. These measurements are shown above in the form of curves of growth. and its leaves prevented from reaching the light, as was done in an experiment (see pl. 2), the sheaths and internodes become abnormally long. Such a rhizome is illustrated in text-fig. 35. From this diagram it can be seen that the response of the blade, sheath, and internode to the stimuli coming from the parent plant are different. While the internodes tend to respond with considerable sensitivity to prevailing conditions, the leaf blades increase in length only when the stimulus passes a certain threshhold. Sheaths as usual appear to be somewhat intermediate in their reaction, embodying features of the two other organs. The physiological basis of this manner of growth was not investigated, but the data indicate, in part, how correlations between plant parts can appear to exist at particular times and at other times to have no apparent basis in fact. It also shows that a process which normally seems irreversible can be reversed upon oc- casion. The trend from rhizome to shoot is commonly observed. The reverse trend is seldom witnessed or imagined possible. The reversal of differentiation jn more advanced parts of the plant, as in the development of so-called viviparous shoots in the spikelet or in the appearance of leaves on the panicle, has been dis- cussed by Sharman (1947). The mere appearance of glumes and lemmas after all leaf forms had been suppressed is also a form of reversal. The development of any given phytomer apparently is a complicated matter which involves a great many variables operating on the plant through its physio- 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 363 logical systems. While phytomers are plastic and can be affected by environmental factors, they nevertheless eventually resolve themselves into a consistent pattern which becomes the mature plant of the proper species. This argues for some sort of internal control. Such control, however, must of a secondary nature, since the achievement of such features of the mature plant as the inflorescence ultimately depend on environmental factors. Thus while the development of a panicle on a growing point undoubtedly affects the internode beneath it, the panicle would never have developed at all had it not been for the fact that the plant had been exposed to a certain combination of environmental conditions. Also the panicle's effect will depend on conditions of the moment and the habitat. There are thus primary environmental influences and secondary internal situations which affect the fate of а given phytomer, but they do not operate independently. Just what role the genetic material of the plant plays in this connection is not clear. In general, it would appear that the independence of the individual structures on the grass phytomer and of the principal plant parts is more imposing than any correlations which might exist between them. Hardly any correlation can be found which can be shown to hold up under all conditions of environment. Men- tion has already been made of the lack of relation between leaf width and the presence of shoots. Brown (1940) found a relation between leaf width and num- ber of rhizomes, but he sampled only two strains. The correlation of short leaves and many tillers is not dependable beyond the limits of normal domestic types. Even the correlation between culm blade length and internode length (text-fig. 27) would break down if more extreme habitat types were included. The im- portance of this independence of the phytomer to variation in bluegrass cannot be exaggerated. It permits a given plant or group of plants to record, in detail, the variations in its environment. It leads to an inevitable confusion in field and herbarium between induced variants and ecotypes. It likewise greatly increases the problem of recognizing and classifying strains and varieties. So-called clonal differences frequently reflect different treatment rather than different inheritance. It is especially striking, and in the author’s opinion not at all accidental, that the criteria, or combinations of criteria, which differentiate one strain or ecotype from another appear to be the very ones which are included in the repertoire of characters which can be produced by environmental manipulation or variation. The important question is why, in one case, these characters should be stable enough to be passed on to offspring, while, in the other, the same characters are theoretic- ally only temporary. How is stability achieved? It does not seem reasonable that this thoroughgoing similarity of induced and inherited characters should be the result of two entirely different causes. It seems more likely that morphological (and therefore physiological) response to environ- ment and genetic change are inextricably bound together in higher plants just as they appear to be in simpler forms of life, and that drawing a line between in- heritance and environment at the finer taxonomic levels is impossible. [Vor. 38 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMARY A three-year study of Kentucky bluegrass has been made with emphasis placed on the morphology and growth of the individual plant and its variation with season and environment. In addition to extensive field observations and tests, laboratory experiments and micro-dissections have been made. Previously published and un- published work has been synthesized with these investigations in an attempt to produce as complete and readable an account of the life history of the grass plant as possible. A grass plant is built of morphological units called phytomers. These units are produced from the growing point. They potentially consist of a leaf blade and sheath, and the internode, bud, and pair of roots immediately below this leaf. Only infrequently are all structures present and fully developed. Іп various combinations they unite to produce the characteristic parts of the mature plant. Ultimate simplicity is achieved in the flowering shoot where the last phytomer of the culm consists only of a much-attenuated internode. Just as the form of the plant depends on which phytomer structures are present, so also does its growth depend on the sequence of maturation and elongation of the phytomer parts. This sequence begins with the rise of the leaf blade from the growing point, and its subsequent elongation. The leaf sheath follows, and then the internode may elongate or remain much compressed. The bud may begin to appear when the blade matures, and may develop into a shoot while the sheath is elongating or remain dormant indefinitely. Roots are usually the last structures to develop. While the individual parts of the phytomer, and the phytomers as a whole, show considerable originality and variation, they characteristically tend to organize into three main types of shoots—the rhizome, the tiller, and the flowering shoot. These structures are analyzed with respect to their morphology and manner of growth. The basic phytomer patterns are found to hold with few exceptions. The critical region of the bluegrass plant is the nubbin of short internodes just beneath the soil surface, to which the leaves attach and from which the three shoot types arise. This area contains the reserve supply of buds, the suppression or development of which determines what the plant will look like and what it will do. A bluegrass plant can be interpreted on a seasonal basis, just as though it were a winter twig or a pine bough. This requires careful study of the crown area and a knowledge of progress of basic seasonal events such as flowering, tillering, and dormancy. By examining the number of phytomers, length of leaves, whether the buds develop, the position of these buds and the kind of shoots they produce, a complete picture of the year’s activity can be drawn. Such a study can be used to provide an understanding of the type of plant to be expected in a given locality or habitat, under different management regimes, or in specific seasons. The struc- ture of the crown region is of great importance in experimental tests, both labora- tory and field, and statistical planning and analyses for the response obtained under 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 365 any situation will vary greatly depending on what type of plant is dominant in the experimental material. A plan of practical management of bluegrass must first take into account to what use we intend to put it, so that we can decide what kind of a plant we need. We are then in a position to apply specific treatments to the sod. The effects of various environmental factors and treatments on grass plants have been discussed in detail with reference to their influence on specific parts of the plant. From a taxonomic and genetic point of view it cannot be emphasized too strongly that bluegrass is flexible and that its final form is largely the result of specific environmental influences which are often brief in duration, rather remote in time, casual in distribution, and specific in effect. While under certain limited conditions correlations between various organs of the plant may exist, if popula- tions from sufficiently diverse habitats are examined most correlations will be considerably altered and may break down completely. А study on variation in bluegrass with reference to this situation is in progress at the present time. BIBLIOGRAPHY Ahlgren, H. L. (1938). Effect of и cutting treatments, and irrigation on yield of forage and chemical composition S e rhizomes of Kentucky bluegrass (Poa pratensis L.). Jour. . Smith, and E Е" Nielsen (1945). Behaviour of various selections of yd bluegrass (Poa pratensis г.) she n grown as spaced plants and in mass seedings. Ibid. 37:268— кн Edgar (1949). The ki RS a Today. Pioneer Hi-Bred Corn Co., Des 2s Iowa. Arber, Agnes (1934). The Gram tudy of cereal, bamboo and grass. Cut ы. tt, О. T. (1935). The deve meos of i barley spike. Jour. Акт. Res. 51 451-457. ---, (1936). Тһе Re CE of the wheat spike. Ibid. = ад , (1937). The development of the oat Есет Ibid. 27- Б. ‚ (1940). фе кшен of S he staminate and еы ra ы of sweet corn. Ibid. 60:25—37. Brittingham, W. Н. (1943). Type of seed formation as indicated by y oii and extent of variation in : Kentucky bluegrass and its practical implications. Ibid. —264. Brown, E. Marion (1939). rie effects of temperature on the growth 24 Eis composition of certain Kur grasses. Mo. Agr. Exp. Sta. Bull. 299. ------- (1943). Seasonal variations in the growth and chemical composition of Kentucky bluegrass. eid Brown, W. L. (1940). "Variation 1 in Poa pratensis. Master's Dissertation, Washington Univ., St. Loui Buckner, G. Davis, and Amanda H. Henry (1945). Composition and yield of Kearucky. Sivan and Korean терсе at different stages of growth. Ку. Agr. Exp. а. Bull. 473. Cooper, J. P., and S. W. Saeed (1 wee Studies on i growth icd development in Lolium: I. Relation of i АП io to head production under ous systems of cutting. Jour or 37:233-259. Darrow, Robert А. (1939). v Pin = Mi tense үс: pH, and nitrogen nutrition on the develop- ment of Poa pratensis. B 2109-127. De Ropp, R. S. (1946). ey in 5 phil ogy of leaf growth: II. Growth and structure of the first leaf of rye when cultivated in isolation or attached to ur intact plant. me he 10:31—40. Evans, Morgan W. (1927). The life een of hac U. S. Dept. Agr. Bull. -------, (1949). Mi a bluegrass. Ohio . Exp. 4% эы Buli ------, and y (1935). The Кеней. certain species of grate, Jour. Amer. Soc. Agron. p E ------ and 1. М. КОО (1939). The growth of Kentucky enim and of Canada нее іп ше ees and in autumn as affected by the caps f day. Jour. Amer. Soc. Agron 3117 and F. O. Grover (1940). Песоа: от of the point of the shoot and the 2 of grasses. Jour. Agr. Res. 61 —52 [Vor. 38 366 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gassner, а (1930). Untersuchungen über die Wirkung von Temperatur und Temperatur- nationen auf die Keimung von Poa pratensis und anderen Poa-Arten. Zeitshr. f. Bot. 3: 767- 838 Hamilton, У, M A ioa A survey of the dairy industry in New Zealand. N. Z. Jour. Sci. & Tech. Harrison, C. м. FTT Responses of Kent "in pn to variations in temperature, light, utting, and —— haps Physiol. 9:83- , and С. W. Hodgson (1939). NM PY certain perennial grasses to cutting treatments. Jour. Amer. [A Agro err 418—4 Hein, M. 9 (1937). Effect of Mire and rate of кем on beef production and plant popula- tion of pastures at Beltsville, Md. U. S. Dept. Agr. Tech. Bull. 538 nmi A. M. (1931). Taxonomy of n “Flowering Plants. New Y York. d L.C Jones, D. F., W. К. Singleton, and L. C. Curtis (1935). The — ене tillering and productiveness in sweet corn crosses. Jour. Amer. m Aa чыр Kannenberg, —, and — 2 num 25% m pen ability ol pm bluegrass of various American origins to make г rs. Pfla d aie 10:478—490. (In Pieters, Ta 7); Карр, E Пы мы уст poese pem Bis of herbage ы for hay and pasture. Herb. 57—63. McCall, M. А. (1934). Developmental anatomy and homologies in wheat. Jour. Agr. Res. 48:283- е К. (1939). Über ле stiriaca Fritsch et Hayek, und d бе Sippen aus der Ver 03. wandtschaft von Poa pratensis Linné. Osterr. Bot. schr. Meyer, B. S., and D. B у дечын (1941). — E Psion New ork. Musgrave, R. B. (1940). Life е studies of Роа pra . Diss. Ds niv. Illin Naylor, Aubrey W. (1939). s of temperature, ae a pi e arsenous acid on asm of Poa pratensis. Bot. Gaz. uu 279; Nielsen, E. L. (1946). The origin of multiple Se in Poa pratensis, Ibid. 108:41—50. Nilsson-Leissner, G. (1937). Experiments in nitrogen nuring o e plots of Festuca rubra and Poa pratensis. Svensk Frotidning 6:29—31. (In Terb, Rev. 5:91-93). са, M. (1923). Comparative morphology and Шы of Poa joe m Phleum pratense and Setaria Italica. Jap. Jour. Bot. 1:55-85 ыу? ber i (1929). Studien über die Entwicklung der DNA und der Blüten bei Pflanzen. Tokyo Imp. Univ., Coll. Agr. Jour. 10:247— M. Т. (1921). Тһе Wheat Plant. London Peterson, M. L. (1946). Physiological 2 of Kentucky bluegrass to different management treatments. Iowa State Coll. Doct. ‚ and W. E. es (1949). Effect ts of Paus and temperature on growth and ng of — 57” Plant Physiol. 2 Phillippe, т. : 43). Effects of some — гучне оп p growth and development of Kentucky Масат пш е. Doct. . Ohio State U Pieters, E i Кы $2). of some — ponds research ке U. S. Dept. Agr. Bur. Pl. Div. ace ин f Dis Prat, wi (эз 4). А biometric Mur p" the culms of cereals and grasses. Canadian Jour. Res. ү M 5). Recherches sur la structure et le mode de croissance des chaumes. Ann. Sci. Nat. (Bot.) 17:81-145. Sharman, B. C. (1942). Developmental a anatomy of the shoot of Zea s L. Ann. Bot. 6:245—282. ------, (1945). Leaf and bud initiation in the Gramineae. Bot. 106:269—289 947). The biology and developmental morphology of the pid apex in the білік New P 0—34. Smelov, S. P. (1937). "Theoretical a of grass land farming. Herb. Rev. 5:132-145. Smith DG. E L, Жер and Н. L. Ahlgren (1946). Variation in ecotypes of Poa pratensis. Bot. Gaz. 108:143-16 Spencer, J- TV HS. ae and Е. N. ras (1949). Seed production of — I as influ вавай н insects, fertilizers, and sod management. Ky. Agr. Exp. Sta. Bull. ша FH. B. 33). oot development of peer grasses A its relation to soil MEA Soil Sci T 9-203. Van de Sande-Bakhuyzen, H. L. (1937). Studies on wheat grown under constant conditions. Food Research Ins peel Stanford Uni von е Н. (1930). сое на über die Blattbreite bei Poa pratensis. Landwirtschaftl. rb. 72:59—63. Watkins, J. M. (1940). The growth habits and chemical composition x dr ier Bromus inermis г» as affected by different environmental conditions. Jour. Amer. Soc. Agro 19:511 ETTER—HOW KENTUCKY BLUEGRASS GROWS 367 Weatherwax, bes (1923). The Story of the Maize Plant. Chic Weintraub, Robert L., and Leonard Price (1947). данае а. gree ology of the grass seedling. МІ Inhibition E ences elongation in various grasses by red and by vilis light. Камын Misc. Col. 106:1—15. Wieland, Artur (1920) Beiträge zur Morphologie, Anatomie, Physiologie, und zum Chemism einiger Typ on Poa pratensis und prn perenne aus dem Gebiet der sogenannten crap Platte. ай, Jahrb. 63:2 —276. Wilkins, a z (1935). Effect of dits on Kentucky bluegrass under conditions of extreme drough ber Amer. Soc. Agron. 27:159 k^ ыты H. . B. Norman, and J. W. Bee (1928). Nutritive value of pasture: III, the in- fluence of pn intengty of grazing on the composition and nutritive value of pasture herbage. (Part I.) Jour. Agr. Sci. 18:266-296 368 [Vor. 38, 1951] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 2 Fig. 1. Baby bottles made it possible to watch how bluegrass grows. (Some of the nutrient solution was removed to show roots and capillary tube supplying air.) Fig. 2. Close-up of the top assembly with rhizome chamber attached and a rhizome extending into it The bottles were placed in a light-tight box, provided with air bubbled under nels. Fig. 3. pressure, and the rhizome chambers concealed beneath a series of pan Fig. 4. One of the panels removed exposing rhizome tube Fig. 5. View of the plants with panel in place. ANN. Mo. Вот. Слкр., Vor. 38, 1951 PLATE 2 5 ETTER—HOW KENTUCKY BLUEGRASS GROWS ANN. Mo. Bor. Слкро., Vor. 38, 1951 A enr ca s -- "TER—HOW KENTUCKY BLUEGRASS GROWS [Vor. 38, 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 369 EXPLANATION OF PLATE PLATE 3 Fig. 6. A confusion of long green midsummer leaves trodden into disarray by cattle. Underneath the summer grass is an equally confusing ie of wiry stems, the OM The soil has been washed away under a stream of water [Vor. 38, 1951] 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 4 Fig. 8. А bluegrass plant and its undernourished offspring. Large parent plant has its roots in a pot of soil underneath the tray. Small plants along the left edge are turned- up rhizomes having only sterile “vermiculite” to feed on. arly ig in the development of a flowering shoot. i cos of the “younger” leaf jM eh ‘Mead in back) has been removed, showing the youngest or “hood” leaf within, and the ит лайга growing point protruding through ү» hood. (Photo by Dr. O. T. Bonnett, of the Agronomy Department, University of Illinois.) Fig. 10. A young flowering shoot with the panicle just appearing from its enclosing sheaths. ANN. Мо. Bor. GaRp., Vor. 38, 1951 PLATE 4 ETTER—HOW KENTUCKY BLUEGRASS GROWS ANN. Mo. Bor. Ganp., Vor. ETTER 28; 1951 13 -HOW KENTUCKY BLUEGRASS GROWS PLATI 5 [Vor. 38, 1951] ETTER— HOW KENTUCKY BLUEGRASS GROWS 341 EXPLANATION OF PLATE PLATE 5 Fig. 11. Successive sheaths enclose each other, but i Vigor bend outward at their junction with the sheath. The lowest leaf on the right an Sx ag Ө shoot with one leaf bent to the left. The prophyli marked Pr is still visible between them Looking down on a bluegrass leaf. The blade is clipped with a thumb-tack, and фә КЕЗ is sii LI and the new blade coming up through it is NL 13. The bulbous translucent growing point of bluegrass, and two leaf primordia. bon by Dr. O. T. Bonnett, University of Illinois). Fig Young leaves rising from the stem apex. Hood leaf is indicated by HL. den M 1940). [Уот.. 38, 1951] 372 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 6 Fig. An August plant, showing rhizomes of extreme types. Left, a very PN б; almost Шер. like shoot; right, below, а long rhizome not yet turned up. Right, above, short arched rhizome. Fig. 16. The tip of a теме The cataphylls bend back slightly at the tip showing beginning of a very short blade. . Left, the conical point of a very young cataphyll; right, the conical point has жас and become slit, eniak blade development has begun (X 80). (From Musgrave, 1940). Fig. 18. A series of rhizomes in various stages of turning up. Upper rhizome has Sid. a single tiller on the right. The crown and growing point are located just above the last roots. Fig. 19. Rhizomes were grown in Petri dishes, some with water only (on the right), some with 1 per cent glucose (center and left). The center one died at the apex and a sprout developed from an axillary bu ANN. Mo. Вот. Ganp., Vor. 38, 1951 PLATE 6 19 ETTER—HOW KENTUCKY BLUEGRASS GROWS PLATE GARD., VOL. Mo. Bor. ANN. ROWS BLUEGRASS G KENTUCKY i) TTER—HOW I [Vor. 38, 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 379 EXPLANATION OF PLATE PLATE 7 Fig. 20. Close-up of an old branched rhizome. Fig. 21. Branched and unbranched rhizomes. The three on the left are from spaced plants in a nursery, the four on the right from a dense sod. (From Musgrave, 1940). Fig Sprout rhizomes produc ed in August on rank meadow plant which had sted A previous May. FS is old flowering ae Fig. 23. А bluegrass plant with five primary tillers. The main shoot is in the cen- west tiller on the left has given rise to a secondary shoot. The lower leaves and red se have been removed. Tillers arise from axillary buds. This plant shows a new tiller (NT), an old im e a year ago (OT), and a fairly recent short rhizome (R). 25. The first leaf of a tiller is called the prophyll (p). It has no blade. (From eee, 1940 Fig. 26. Oat seedling showing the first green leaf after it has grown out of the dira coleoptile (C) at its base. The plant on the right shows a somewhat broken MER (M) between the grain and the first minute root. 374 Fig. [Vor. 38, ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 8 Successive stages of panicle development in bluegrass. (Photos by Dr Т. Bonnett, of the Agronomy Department, University of Illinois). A. fig. 1 Ф z The growing point elongates. (From Musgrave, 1940—compare with pl. js Proliferation begins with protuberances appearing on the growing point. The primary protuberances branch and the panicle grows very slightly. Spikelets have begun to develop. Glumes of spikelets have begun to envelop the growing points. 1951] Oi 5, ANN. Mo. Вот. Сакр., Vor. 38, 1951 PLATE 8 ETTER—HOW KENTUCKY BLUEGRASS GROWS 9 PLATI GaRD., Vor. 38, 1951 Bor. ANN. Mo. SMOUD SSVYOANTA AMXO(LLNd33 AOH-—WALLZ [Vor. 38, 1951] ETTER—HOW KENTUCKY BLUEGRASS GROWS 375 EXPLANATION OF PLATE PLATE 9 Fig. 28. Bluegrass plants showing various degrees of tillering. Meadow plant at right shows two primary tillers. Pasture plant at left has several secondary shoots. e plant in the center was taken from a wet spot over a buried steam pipe and has т tillers all winter. They total about 50, and there are many of the second and third generation and a few of Fi fourth. Upper left, a much-tillered lawn plant with very short leaves. Fig. 29. Plants fertilized at different times of the year with horse urine (see Table IIT). рег left, control; lower left, late August; center, late October; right, January— note exsertion of panicle beginning in January plant. Difference in intensity of color is also apparent in the photograph, and is true to life. Control and August plants were very pale green Fig Enlarged crown (X 315) of a conservative meadow plant in fall with only three Ей, déveloged à in a period of a year. more active plant from a pasture in fall showing rhizomes and uds. 31. Crown of a ius dn late summer and fall b Fig. 32. A crown with late summer and fall buds swelling in 3k, diae for develop- ment of rhizomes the following spring. A pair of tillers at the t Annals of the Missouri Botanical Garden Vol. 58 NOVEMBER, 1951 No. 4 VALERIANA IN NORTH AMERICA AND THE WEST INDIES (VALERIANACEAE) * FREDERICK G. MEYER** The present treatment is the first taxonomic revision covering the North American and West Indian species of Valeriana since that of Hick’ (1882). The taxa within this geographical province have not suffered from lack of recognition, as may be witnessed by the rather copious synonymy, but rather from the want of a general re-evaluation of these past efforts in an attempt to bring a semblance of order and utility to the natural populations of North American Valeriana. Taxonomically, the genus as a whole remains poorly known. The European species are more often studied, although for the most part floristically. Valeriana officinalis and V. Pbu, amongst others, are known from antiquity especially for the medicinal properties of their fetid “valerian root". The most recent work on American Valeriana is that of Borsini? who treated the Argentine species. Valeriana occurs in all the continents with the exception of Australia, and in number of species, totaling perhaps 200, the genus compares with those genera largest amongst flowering plants. In the western hemisphere, Valeriana attains its greatest complexity in the South American mountains where the largest number of species is concentrated. Héck’s treatment! accounted for 155 species of Valeriana world-wide, of which 30 were of North America and the West Indies. His disposition of the North American species was limited by a paucity of material, and, while conservative, it hardly fulfilled the need of a detailed account based upon abundant field data and herbarium specimens. In the decades that followed, an increasing number of names appeared in the literature, and with well over 100 specific epithets with which to deal at the beginning of the present study, there was an urgent need for an account of the North American and West Indian Put Bot. Jahrb. 3:1-73. 1882. om et Sp. Plant. Argent. 2:275. 1944. n investigation c carried out in the graduate laboratory of the Henry Shaw School of Botany of н ton University and submitted P a thesis in partial fulfillment of the requirements for the Гавда of Doctor of mar id June 949. ** Missouri Botanical Garden, St. Louis, us Issued December 31, 1951. (377) [Vor. 38 378 ANNALS OF THE MISSOURI BOTANICAL GARDEN species as a whole. А monograph covering a selected geographical area, such as the present, limits the interpretation of extra-territorial distributions, but this method is the only means open for a full-fledged evaluation of the relationships of taxa within a natural taxonomic system. One of the results of the present study has been the determination of extra- North American affinities, and I have discovered, for instance, that nearly one- fourth of the species also occur outside the geographical limits of this paper, five extending to South America and two to Asia and Europe. The interpretation of taxa has been solely on the basis of comparative morphology, and 30 indigenous North American and West Indian species are listed in the treatment that follows. HisronicAL REVIEW The name Valeriana is derived allegedly from the Latin valere, to be strong, or from Valerius, a Roman family name, or from Valerus, the name of a Roman king. In Greece the plants of this assemblage were known as Phou (ov). Valeriana as a generic epithet appears in the works of Theophrastus, Dioscorides, and Plinius. To them this plant was important largely because of its medicinal qualities, and to this day Valeriana officinalis is listed in the United States Pharmacopoeia for use in mild cardiac therapy. The history of its medicinal use is correlated with the advance of botanical knowledge, and some of the earliest incunabula illustrate Valeriana. We find an interesting but stylized black and white wood-block print of Valeriana in the ‘Herbarius Latinus’, printed at Mainz in 1484, and a hand-colored print in ‘Gart der Gesundheit’ of 1487, printed in Augsburg by Hannsen Schónsperger. Valeriana found its place in the tomes of all the early sixteenth century herb- alists, and Cesalpino?, the Italian physician, placed Valeriana with those plants having a solitary, single-seeded fruit. But these renaissance classifications con- tributed little new information, and not until 1700 did Valeriana rise as a result of Tournefort's more modern approach. Tournefort*, in his re-evaluation of earlier classifications, incorporated Valeriana and the newly erected Valerianella in his Class II, Section III, with flowers gamopetalous, infundibuliform and rotate, with the calyx unfurled in fruit. This interpretation was a major advance, and Linnaeus, in the 1753 edition of ‘Species Plantarum’ recognized 16 species of Valeriana in the genus with the “Triandria Monogynia". His interpretation was conservative and in his list were included species of Valerianella, Fedia, and Centranthus, which have since been recognized as separate genera of Valerianaceae. In Linnaeus’ second edition (1762), the number of species reached 18, and V. scandens was described as the first American species. Jussieu” made no attempt to alter Linnaeus’ concept of Valeriana although Һе placed the genus in his newly erected family Dipsaceae. Necker® first devised the ?De Plantis. p. 147. 1583. ^Inst. Rei Herb. 1:131. 1700. 5 Gen. Pl. p. 195. 1789. 6 Elem. Bot. 1:123. 1791. 1951] MEYER— VALERIANA IN NORTH AMERICA 379 Valerianaceae and recognized five genera: Valeriana, Centranthus, Mitrophora, Odontocar pon, and Mouffetta. A. P. DeCandolle? proposed 11 genera of Valerianaceae in its first world-wide treatment. For Valeriana 82 species were recognized of which 10 were North American, and he placed the family between the Rubiaceae and Dipsaceae which is essentially the same disposition as the most recent one by Hóc 5, Endlicher? (1836-40) failed to treat the genus, although the Valerianaceae were placed between the Plumbagineae and the Dipsaceae. Bentham and НоокКег!9, like DeCandolle, placed the Valerianaceae between the Rubiaceae and the Dipsaceae. Hóck's! (1882) monograph is the last disposition of the species for North Amer- ica, and his treatment was later included in Engler and Prantl’s'* “Die Naturlichen Pflanzenfamilien’. Valeriana pauciflora was described by the younger Michaux in 1803 as the first species recognized from North America and the West Indies, and by 1850 over half the known species from this geographical area had been recorded, mainly as a result of botanical expeditions—Humboldt and Bonpland, Galeotti, and Hartweg, in Mexico; and Nuttall, Douglas, and Richardson, in western United States and Canada. Asa Gray's!? treatment of Valeriana was the last to list the species north of Mexico. GENERAL MORPHOLOGY Habit:—The North American species of Valeriana are hollow-stemmed peren- nial or annual herbs, ranging in size from that of the often diminutive napiform- rooted species, such as V. densiflora, sometimes no more than six inches tall, to that of the voluble V. scandens with branches up to twenty feet long. The erect perennial and annual species normally flower and fruit in response to seasonal fluctuations of climate, but the voluble V. clematitis and V. scandens are mostly everblooming and evergreen over much of their distribution in the warmer sections of tropical and subtropical America. The underground parts are useful for series designation and the species fall generally into two distinct groups: (1) those with /af-roofs (Mexico and Central America), and (2) those with rhizomes (north of Mexico). (Fig. 1). 1. The species with persistent primary roots may be divided into two groups, namely: (a) those with conical tap-roots, and (b) those with napiform to fusi- form tap-roots: (a) Conical tap-roots are characteristic of the three North American species in series EDULES. "Prod. 4:623. 1830 °Gen. Pl. p. 350. 1836-40. 10 Gen, P]. 2:151. 1873. 11] c., p. 1. 1882. 1? Nar. Pflanzenfam. 44:178. 1897. 13 Syn. Fl. М. Am. pp. 42-44. 1886, 380 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vor. 38 put ir en ae ERA Т еле; жаа” e Fi "i Ja tap-root of V. edulis, characteristic of the spe B, napiform tap-root of V. densiflora densiflora, characteristi C, rhizomes of V. capitata acutilob form tap-root of V. densiflora barb of the species in serics CERATOPHYL А, branched s cies in series EDULES; eristic of the species in series ORBIFOLIAE ; 4, Characteristic the species in series OFFICINA fusi- areaefolia; E, conical tap-root of V. albo-nervata LAE. LES D, , characteristic 19511 MEYER— VALERIANA IN NORTH AMERICA 381 (b) Napiform to fusiform tap-roots are characteristic in series CERA- TOPHYLLAE, CLEMATITES, DENSIFLORAE, SORBIFOLIAE, and PRATENSES. The distinctions between the perennial and annual habit in the napiform- rooted species are based primarily upon the relative differences in the degree of swelling, which perforce does not always delimit annuals from perennials satisfactorily. For instance, several of the species in series DENSIFLORAE, CLEMATITES, and SORBIFOLIAE manifest transitional forms from well-devel- oped napiform tap-roots to those in which the swelling is often quite rudimentary as in V. densiflora, apiifolia, Selerorum, and urticaefolia. Тһе Mexican and Central American species with napiform tap-roots often occur in areas of seasonal drought, and it is conceivable that the annual or perennial habit may be linked to the severe conditions of an arid habitat. The rhizomatous species are included almost wholly in series OrriciNALES in which the often much-ramified and branched rhizomes are diagnostic. In series SORBIFOLIAE, rhizomes occur in V. domingensis, and dubiously so in V. clematitis and V. scandens, in which the underground portion is rarely preserved in herbarium material. Leaves:—The leaves in all the species are opposite and decussate, and the spatu- late leaf epitomizes the basic leaf type in North American Valeriana. 'The leaf of V. edulis is the best example of this shape, and the pinnate, pinnatifid or bipinna- tifid leaf are modifications of this basic form. The leaves are the most variable of all the taxonomic criteria, although the characteristic variational pattern is often diagnostic for species determination. Undivided leaves are predominant in series EDULEs and CLEMaTITEs, but the leaves in series OFFICINALES, CERATOPHYLLAE, DENSIFLORAE, SORBIFOLIAE, and PRATENSES are more frequently pinnate to pinnatifid. The leaves in V. apiifolia and V. robertianifolia are bipinnatifid, and the laciniate leaves in the species of series CERATOPHYLLAE are the most divided of the North American species. Indument:—The kinds of pubescence, whether of the hirtellous, pilosulous, sericeous or puberulent types, offer a useful means for characterizing the series. The nodes of most species are consistently puberulent or pilosulous, and in six of the seven series, the throat of the corolla is pilosulous, but in series CERATOPHYLLAE the corolla-throat is densely short-sericeous, a character which readily marks this group of species. The disposition of the indument often contributes to the diagnostic features of the achene. In the series OFFICINALES, EDULES, DENSIFLORAE, and PRATENSES the achenes are glabrous or pubescent on the adaxial and abaxial sides, but in series CERATOPHYLLAE, CLEMATITES, and SORBIFOLIAE the achenes are more often pubescent on the adaxial and glabrous on the abaxial surface. Inflorescence:—The compound inflorescence of Valeriana has been interpreted as a thyrse, and Philipson (Ann. Bot. n. s. 11:409. 1947), on anatomical grounds, finds the thyrse in V. officinalis considerably modified as a result of arrested growth patterns of the apical meristem. This conclusion bears out my observations that [Vor. 38 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN the inflorescence in Valeriana is probably not a true thyrse of the mixed type but merely thyrsoid and actually determinate throughout. The inflorescence in North American species is manifestly of the V. officinalis type. Two forms may be rec- ognized: (1) An aggregate dichasium'*, which is initially more or less pyramidal. This type predominates in the North American species and is apparently produced through the reduction of decussate leafy branches from a falsely monopodial cen- tral axis; (2) A compound dicbasium, which is dichotomous, with the ultimate dichotomies closely aggregated and more or less flat-topped in anthesis, although later becoming diffuse and somewhat more elongated. The compound dichasium is found consistently in V. deltoidea, tanacetifolia, and pratensis. The terminal cymes are scorpioid, and the ultimate branches form the cincinnus. In most species the cincinni are not evident until maturity, although the heliciform branches of V. apiifolia and urticaefolia are well developed even in immature plants. Corolla:—Relative lengths of the corolla tube in North American Valeriana contribute an outstanding character for both series and species delimitation. The corolla may be: (1) infundibuliform to subsalverform with the tube usually twice as long as the lobes and gibbous at the base (except in series PRATENSES it is some- times indistinctly gibbous towards the middle), or (2) rotate, with the tube straight and much abbreviated and the lobes usually as long or longer than the tube. Modifications of these types occur in V. pauciflora, columbiana, and urticae- folia, with subsalverform corollas, and in series SoRBIFOLIAE, with campanulate- infundibuliform corollas. The corollas of V. pauciflora and V. columbiana are the longest, to 20 mm., of the North American species. Rotate corollas occur in V. occidentalis and V. dioica in series OFFICINALES and in most species in series EpuLEs and CERATOPHYLLAE. Тһе five corolla lobes are imbricate in bud and at anthesis they expand at right angles or frequently they are recurved, rarely erect. The flowers with infundibuliform corollas are more or less asymmetrical. Rotate corollas, on the other hand, are usually symmetrical. The presence of unisexual as well as perfect flowers in many species of Valeriana is expressed differentially in the length of the corolla. Unisexual flowers, for in- stance, are invariably reduced in length by at least half that of the perfect flowers. Sexual polymorphism occurs in several ways; the flowers may be completely hermaphroditic, polygamous, dioecious or polygamo-dioecious, with dioecism the least common of these sexual anomalies. Most of the species are characteristic for one or more of these sexual traits; for instance, the flowers of V. edulis are predomi- nantly polygamo-dioecious, while those of V. sitchensis are consistently hermaph- roditic. The methods of pollination are as yet unknown, although my observations on several species in western United States indicate that small insects, of undetermined kind, assist in this process. 14Woodson, Ann. Mo. Bot. Gard. 22:4. 1935. 1951] MEYER—VALERIANA IN NORTH AMERICA 383 Fig. 2. Anthers: A and B, 4-lobed type, X 10; C and D, 2-lobed type, X 8. Stamens:—The versatile anthers in North American species of Valeriana, while consistently 4-loculate, are of two distinctive types, viz. 2-lobed or 4-lobed; illustrated in fig. 2. In the 2-lobed and more common type, the anther thecae usually are somewhat lunate and opposed, with the 4-loculae equal in length. This anther type occurs in the North American series CERATOPHYLLAE, CLEMATITES, DENSIFLORAE, SoRBIFOLIAE, and PraTENSES, which include Mexican, Central American and West Indian species. The material studied for comparison shows that most of the South American species also have anthers of the 2-lobed type. In the 4-lobed type, the anther thecae are sulcate, with the ventral loculae of each theca slightly longer than the dorsal and essentially parallel. This type occurs in series OFFICINALES and EpuULEs, and was also seen in material from certain South American species and in all the European and Asian species examined. While these anther characters have no value for species distinctions, the 2- or 4- lobed anthers have been most useful for series designation. The stamens of most species are exserted and longer than the corolla lobes, although in series SORBIFOLIAE the stamens are mostly included and essentially sessile at the summit of the throat of the corolla. Achenes:—The inferior cypselate achene of Valeriana, with its epigynous setose calyx-limb and single fertile carpel, represents a reduction from a basically 3- carpellate ovary. On the abaxial side, the vestigial carpels are seen as two often indistinct protuberances near the apex of the achene, and the accompanying vascu- lar traces occur as two submedian ribs adjacent to the abaxial midrib. Variations in the length, the shape, and the indument are the most useful achene characters for species diagnosis (fig. 3). Calyx-limb:—The modified setose calyx of Valeriana is referred to as the calyx- limb. This structure is apparently derived through the progressive dividing of calyx bundle traces in the normal development of the epigynous calyx. The setose calyx-limb in Valeriana is allegedly homologous with the pappus in Compositae, although at present we lack information concerning this alleged relationship. [Vor. 38 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig nes: А and B, V. ий scandens (bax side), X 7; D side), X 4 and С, V. Selerorum (F, adaxial side, G axial side), ү en clematitis J, (H, Ta i sd I, adaxial side), & 8; side), X 6 cucurbitifolia (A, abaxial side, B В, adaxial side), V. pauciflora eran side), X 6; E, d gleba (abaxial ti V. реа ен side), X 4; К V. € (adaxial 1951] MEYER— VALERIANA IN NORTH AMERICA 385 During anthesis the calyx-limb and plumose segments are tightly inrolled, while at maturity they unfurl, the divided limb usually displaying 6-23 segments. In V. apiifolia, pulchella, and deltoidea, the calyx-limb may be either setose or merely dentate. In V. pratensis, it is consistently dentate. Graebner (1906), in his classification of Valeriana, rightly discredited the value of the setose calyx-limb as a generic or subgeneric character, although Héck (1882) and his predecessors ascribed considerable importance to this character, first for generic and later for series designation. In the North American species, the presence or absence of the setose calyx-limb is diagnostic only in combination with other characters. GEOGRAPHICAL DISTRIBUTION The international boundary between the United States and Mexico marks an obvious discontinuity for the distributions of the North American species. Northern Province: —Within this area occur the species in series OFFICINALES and EpuLEs. In addition to V. dioica and V. capitata capitata, which are found in Asia and Europe as well as in America, series OrriciNALEs includes a host of European and Asiatic species. The species in series EpULES are wholly American. Southern Province:—This region includes Mexico, Central America, and the West Indies, in which occur the series CERATOPHYLLAE, CLEMATITES, DENSI- FLORAE, SORBIFOLIAE, and PRATENsEs. ‘These series manifest an affinity with South American Valeriana, and V. clematitis, urticaefolia, scandens, robertianifolia, and sorbifolia are common to both continents. CLASSIFICATION No up-to-date general classification exists for the genus Valeriana, although the family was last treated on a world-wide basis by НӧсК! in 1882. P. Graebner’? began an extensive monograph of the family which covered mostly the South American species. Valeriana is one of seven to nine genera of Valerianaceae. The various genera are distinguished principally on the basis of stamen number as a reduction series from a basically 5-merous whorl: Nardostachys, an Asian genus, has 4 stamens, Valeriana 3, and Centrantbus and Fedia, both Mediterranean genera, each have 2 stamens. The species of Valeriana covered in the present treatment are included in seven series, five of which are used for the first time and two being upheld from Höck’s earlier classification. The synopsis that follows is a summary of the genus cover- ing the North American and West Indian species as interpreted by me. 15 Engl. Bot. oe 37:464. 1906. 16 Engl. Bot. Jahrb. 3:1—73. 1882. 11], с, 37:464 i 6 [Vor. 38 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN кнн nsi Ser. OFFICINALES arizonica pauciflora columbiana occidentalis іоіса Ser. EpuLEs texana prionophylla edulis ciniosa albo-nervata ceratophylla Ser. CERATOPHYLLAE clematitis Selerorum Ser. CLEMATITES ене ee pa, pee palmatiloba a biifolia Ser. DENSIFLORAE белде scandens Ser. SORBIFOLIAE domingensis robertianifolia Palmeri sorbifolia Ser. PRATENSES яз sse рар Яязяя 533 яяя аадар Е" ш. pus um A — — tanacetifolia pratensis Есомоміс IMPORTANCE Valeriana is of economic importance principally for the medicinal properties in the root, sometimes as an aromatic perfume, less frequently as a culinary herb (Valeriana edulis). The medicinal properties of Valeriana have long been recog- nized. The species most used commercially has been the European Valeriana officinalis, although other European and Asian species have been used at intervals. The strong fetid and aromatic odor so characteristic of V. officinalis is common to many North American species, especially in the series OFFICINALES and Ерот, although none of the American species are used medicinally, so far as known. The “fetid roots", according to Lindley!®, bring on, “as is well known а kind of intoxication in cats, and in large doses occasioning in man scintillations, agita- tion, and even convulsions”. The therapeutic value of Valerian root has a reputa- tion of long standing. It has been used externally for epilepsy and is purported to have cured this disease in Fabius Columna, a well-known herbalist of the 17th 19 m— Barton!? claims that Valeriana is very efficacious in epilepsy produced 18 Pickering, Chron. Hist. Pi. р. 518. 1879. WBritish Fl. Med. р. 391. 18 1951] MEYER—VALERIANA IN NORTH AMERICA 587 by anger and fear. It has also been used for nervousness, suffocation, asphyxiation, migraine, menopause, fevers, and parasites. Allport?? lists its uses as а nervine stimulant to the digestive organs. The culinary properties of Valeriana remain improperly known, although V. edulis in westezn United States has been used for food. This plant, with its usually large fleshy roots, is still used in this way by certain Indian tribes of northwestern United States. On his visits with these Indians in the 1820's, David Douglas recorded the following data about V. edulis (Patrinia ceratophylla) : ped io during the Aes months, are collected by the Indians, Гүн 5: on yd stones and u an article of winter or spring food. From a bitter and seemingly pernicious ние it is thus converted into а soft and pulpy mass, which has a sweet og resem- bling that of treacle, and is apparently not T ILE some.?! The only species with any current commercial use in the United States is Valeriana officinalis, and in addition to its medicinal properties, this species is still used in gardens for its ornamental value. To my knowledge, the indigenous North American species are seldom cultivated, although V. arizomica was known to European gardens at least fifty years ago. Several western American species, such as V. sitchensis and V. columbiana, should tantalize the efforts of gardeners in northern climates to grow them for their neat appearance and large showy flowers. Valeriana columbiana is a worthy rock-garden subject, while V. sitchensis would succeed well in a woodland or moist perennial border. SrUDY MATERIAL In the citation of specimens, I have, wherever possible, used the symbols for herbaria as advocated by Lanjouw (in Chron. Bot. 5:143. 1939). ARIZ—University of Arizona, те BH—Bailey Hortorium, Ithaca, N. Y. BRY—Brigham Young dee 124 Provo, Utah. G—Boissier Herbarium, Genéve m an F—Chicago Natural History [Field] Museum, Chicag CA-— Colorado Agricultural and Mechanical College, Fr. “Collins. D—Delessert Herbarium, Genéve. —Royal Botanic Garden, Edinburgh FI—Istituto Botanico dell’Universita, Firenze. GT—Gentry Herbarium, Los Angeles. GH—Gray Herbarium, Cambridge, Mass. K—Royal Botanic Gardens, Kew. M—Botanische Seaan München. MAT—Matuda Herbarium, Mexico City. MSC—Michigan State ‘College East Lansing. 20 Chem. & Phar. Veg. Drugs. p. 159. 1944. lIn Hooker, W. J. Fl. Bor. Am. 1:291. 1834. [Vor. 38 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN MU-—University of Michigan, Ann Arbor. MO—Missouri Botanical Garden, St. Louis. MIN—University of нра Minneapolis. —Division of y, Dept. of үкси Ottawa. О ; den P—Muséum National d'Histoire Naturelle Paris. PO—Pomona College, Claremont, Califor WYO—Univerity of Wyoming, Laramie. P T T—Escuela Agricultura Pan Americana, ТЕК Honduras. US—United States National Herbarium, Washing i ey. STR—Institut Botanique de l'Université, Strasbourg, WILLU—Willamette University, Salem, Oreg ACKNOWLEDGMENTS I am indeed very grateful and certainly indebted to the many individuals in the various herbaria from which specimens were either borrowed or studied and for the many courtesies and privileges extended. I wish to express thanks to the Director of the Missouri Botanical Garden, Dr. George T. Moore, for the full use of Garden facilities, and words to the library staff are inadequate as thanks for tireless assistance. I wish to thank Dr. R. E. Woodson, Jr. for his continued counsel and genuine assistance during critical moments in the study. During 1950 visits to several of the larger herbaria in Great Britain and on the continent facilitated the study of many of the older collections not available in American herbaria. Most of the drawings are from the pen of my wife whose patience and diligent assistance in this way added its full measure to the successful completion of this study. To Ellen Lissant, I extend thanks for her drawings which are included under "Morphology." The citation of specimens is in accord with the following plan: (a) Alaska and Canada: one collection per locality with the widest herbarium distribution; (b) United States: by states, one specimen per county with the widest herbarium distribution; (c) Mexico, Central America, West Indies: all specimens seen are cite TAXONOMY VALERIANA [Tourn.] L. Sp. Pl. ed. 1. 31. 1753; Gen. Pl. ed. 5. 19. 1754. DC. Prod. 4:627. 1830; Hóck, in Engl. Bot. Jahrb. 3:1-73. 1882. LECTOTYPE — V. officinalis L. Hemesotria Raf. in Ann. Gen. Sci. Phys. 6:88. 1820. Oligococe Willd. ex DC. Prod. 4:632. 1820, nomen in hb. Willd. Amplophus Raf. rem Bot. 89. 1840. (T.: based on ү. scandens L.). 1951] MEYER—VALERIANA IN NORTH AMERICA 389 Herbs, perennial or annual from rhizomes or tap-roots. Stem subscapose or leafy, fistulous, terete or occasionally more or less quadrangular. Leaves decussate, basal and cauline, spatulate and undivided or pinnate to pinnatifid or rarely bi- pinnatifid, frequently more or less decurrent on the subpetiolar and more or less clasping-patelliform base, serrate, crenate, dentate, repand, or entire, membranous to firm. Inflorescence determinate, either aggregate-dichasial and thyrsioid or cymes compound, dense and more or less scorpioid, bracteate; flowers hermaph- roditic, gynodioecious or polygamo-dioecious. Corolla infundibuliform, subcam- panulate, or rotate, the tube gibbous or straight, usually more or less hairy on the throat, the 5 lobes equal or subequal. Stamens 3, rarely 4, adnate on the throat, anthers essentially sessile and included, or filamentous and exserted, alternate with the corolla lobes; anthers 4-loculate, introrse, 2-lobed, the thecae more or less lunate and opposed, the 4 loculae equal in length, or 4-lobed, the anther thecae sulcate, the ventral loculae longer than the dorsal and parallel. Pistil inferior, ovary basically 3-carpellate, maturing 1-fertile adaxial carpel, ovule 1, pendulous, anatropous, exalbuminous, ventral raphe united; vestigal abaxial carpels 2. Sfyle 1, the stigma 3-lobed, included or exserted. Fruit a cypselate achene, adaxial veins 1-median, 2-peripheral, abaxial veins 3, oriented more or less in the median plane. Calyx initially involute, later spreading, the sessile limb concrescent and short- patelliform, hyaline and membranaceous, becoming setose in mid-plane, the setae plumose, or the limb short-cupuliform and more or less irregularly toothed or lobed. Species, 30. KEY TO THE SERIES”? A. Plants rhizomatous or from conical tap-r nthers distinctly lobed, the thecae su жы with Ж} ventral ps slightly longer ME the dorsal and essentially parallel. B. Plants from rhizomes or stolons. Leaves mostly pinnate to pinnati- fid, petiolate, and the blades of the undivided leaves more or less henes нац Ы ог £ gynodioccious OFFICINALES (p. 390) BB. Plants from conical s mostly lingulate-spatulate, gradually decurrent to gri Viral n aud clasping base, the cauline dee ently pinnate to pinnatifid and more or less decurrent. Corolla ssentially rotate. Achenes transversely à dem on or smooth, ada e ribs NEUE prominent. Flowers polygamo-dioecious AA. Plants from napiform to fusi iii Ва -roots petens usually "s EI EDULES (p. 420) lobed: the thecae usually somewhat lunate and opposed, the loculae equal in length C. Ж ЖЕҢ Рі; uniformly spreading or nearly wanting. Rhachis of the divided sg terete a, usually winged in V. Palmeri). E ibbou the tube straight. D. фы ба. Corolla (L8- ) 2.3-6.0 mm. long. ?? Dimensions and descriptions of the corolla apply to i icm flowers; unisexual flowers are reduced to approximately half the length of the perfect flower [Vor. 38 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN E. Corolla bp ip the lobes at least equaling or sometimes excee eeding the tube length, the throat manifestly clothed wi stiffis ous within. н divided, the lateral lobes more or less laciniate or palmatel а E ы е ав with the abaxial ribs usually ed CERATOPHYLLAE (p. 430) — — I EE. Corolla Забо ат то pcr uh infundibuliform, ually not пе half the tube length, the i potrei JT ti lous within. Leaves ivided or pin- na ірі crenate to dentate or irregularly "x ft to entire. Achenes i t zc baxi F. Leaves predominantly undivide 4% ог sometimes with 1 pair of lateral — Achenes frequently more or less arcuate, linear- to e-oblong, oe ered ev iy ‘hess on the ded. side or glabro IV. CLEMATITES (p. 437) FF. Leaves predomina eia Pin ate to pinnatifid or bipinnatifid (much reduced ue essentially br racteate in V. vaginata), prevailingly with more than 1 p undivided. Асһепеѕ — plane, suborbicular to es more ovate-oblong, sometim r less ovoid, uniformly DENSIFLORAE (p. 448) dense-pilosulous or glab i DD. adum "inclu ded КСК potum іп V. robertianifolia). lla VI. SORBIFOLIAE (р. 461) Coro .5-3.0 mm. lon CC. Indument usually "SE disposed on the veins and stems, the eaves ascending-ciliolate. Rhachis of the divided Been more r less winged. Corolla gibbous towards the middle i ig I. PRATENSES (р. 479) Series І. OrriciNALES Hóck,?? in Engl. Bot. Jahrb. 3:41. 1882. Perennials from rhizomes. Stem unbranched to the inflorescence. Leaves basal and cauline, elliptic- to obovate-spatulate, pinnate to pinnatifid or sometimes un- divided, petiolate, dentate to repand or entire. Inflorescence a compound dichasium, more or less flat-topped in anthesis; flowers hermaphroditic or sometimes gyno- dioecious. Corolla infundibuliform to subsalverform or rotate. Stamens and style exserted, anthers distinctly 4-lobed, the thecae sulcate with the ventral loculae longer than the dorsal. Calyx-limb 9- to 23-fid. Species, 8 Tyre Species: Valeriana officinalis L. Series OFFICINALEs includes eight closely related species in North America, all north of Mexico. In distribution, the series is circumboreal and includes many of the Valerianas in both Europe and Asia. It stands apart in being the only as- semblage of truly rhizomatous Valerianas in North America. The 4-lobed char- acter of the anthers, the usually well-developed pinnate to pinnatifid leaves, and the more or less flat-topped inflorescence are characters which combine to distin- guish the species in series OFFICINALES. KEY TO THE SPECIES A. Corolla infundibuliform to subsalverform, 3—19 mm. long, the tube gibbous. Le ore cft atulate. mens exserted, longer than the corolla lobes. Corolla lobes less m. lon Sta than half the tube length. Achenes 2.0—6.5 m C. Plants = ut stolons. Corolla white or pink, 3— 9 (—15) mm. lon enes unwinged, narrowly to broadly ovate, oblong to 5 стана 2 The epithet for this series was originally used by Hóck in the NON number as “OFFICINALIS.” The change to plural number is essential under Series designation 1951] MEYER—VALERIANA IN NORTH AMERICA 391 D. Leaves uniformly пошла е the terminal lobe linear to d or oblanceolate, cm. long, 0.4—3.0 cm. wide, the lateral lobes sometimes a ate, simulating the terminal lobe. qe lobes of the basal leaves 8—12 pairs. Corolla 3—5 mm. long. Achenes 2.5-3.0 mm. long. Adventive from Енгоре................222....... estan tasses n ‚нне енен emen innen 1. V. officinalis DD. Leaves quer or pubescence spreading. . Leaves predominantly cauline de ovate or basal and mostly spatu la 5 К ка sometimes finely ria or glabrous. Corolla to 9 mm. long. F. Plants un robust, 3—10 dm. tall. Cauline leaves evi- dently epi at least the lower, glabrous or essentially so. Alaska, British Columbia, northwestern and north- eastern кы pud Б p slender, 1-6 dm. tall. Cauline leaves essentially sessile and glabrous or “the stem and/or leaves uen Northwestern Arctic pai si western United States.c.22..:cecceccocccsesecsscscnsstnennsccnstpsapeuioesnraapsoesastonn .3. V. capitata EE. Leaves mcn basal and ovate. Stem glabrou Corolla. mm. long. Southern Colorado, New Мінсе. sitcbensis 7 "1 es I m a e - 4. V.arizonica апд А CC; Planes “ae sto Loe Corolla blue to pinkish, 13-19 mm. g ong. nes narrowly winged. Southern Illinois, Ohio Valley to VET 25. V. pauciflora BB. ae ager shorter than the corolla lobes. Corolla lobes 3-6 redde bout Lau MP шышы Achenes 5—7 mm. long. Wen atchee Mountains, Washington..........--..:s--sssssscseesseeesseeessesseeeseress . V. columbiana AA. Corolla и то borate; 2.0-3.5 mm. long, Ба tube indistinctly gibbous or straight. Leaves predominantly oblon G. Plants (3—)4.5—9.0 dm. tall, lea к robust. Achen linear- to ати sparsely to den sdy pilots, rarely glabrous. North-central Idaho orther N a and por засу ьн o Сойогафо........ ИЕ ане 7. V. occidentalis "a dm. tall, slender, somewhat sub- scapose and considerably less leafy. Achenes ovate to ovate-oblong, glabrous. Northern "fucks Mountains in the United са northern British сақы south- eastward to Newfoundland......................... n 8. V. dioica sylvatica a Q 29 582 TERT 2А мл | bag = 1. VALERIANA OFFICINALIS L. Sp. Pl. 1:31. 1753. Perennials 5-10 dm. tall, robust, from an abbreviated rhizome, 0.5-1.0 cm. thick, bearing numerous caudical rootlets. Stem leafy, 0.5-1.0 cm. in diameter, pilosulous to short-pilose towards the base, glabrescent above, the nodes densely pilosulous. Leaves predominantly cauline, 4-5 pairs, petiolate below, becoming sessile above, oblong to oblong-ovate, pinnate to pinnatifid, 9—35 cm. long, gla- brous or pilosulous to short-pilose, predominantly on the veins beneath, glabrous above, uniformly ascending-ciliate, the lateral lobes 5—8 pairs, distinct or more or less decurrent on the rhachis, linear to oblanceolate, acute, sometimes more or less falcate, 2.0-7.5 cm. long, 0.4—3.0 cm. wide, the terminal lobe % as long but as wide as the lateral lobes; basal leaves loosely tufted, 15—30 cm. long, simulating the cauline. Inflorescence 2-11 cm. wide in anthesis, later diffuse, 10—18 cm. long, 10—15 cm. wide; bracts 2—3 mm. long, relatively long-ciliate, short-apicu- late, the nodes often densely pilosulous; flowers hermaphroditic. Corolla infundi- buliform, 3-5 mm. long, white, glabrous without, the lobes half as long as the tube, the throat sparsely puberulent towards the base within. Stamens and style [Vor. 38 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN exserted. Achenes lanceolate- to ovate-oblong, 2.5-3.0 mm. long, glabrous or pilosulous, tawny or rubiginose, abaxial ribs evident. Calyx-limb 10- to 12-fid. Type Locauity: Europe. DISTRIBUTION: Introduced into gardens in the United States and Canada. Established as a garden escape from New Brunswick westward to Minnesota and Washington, apparently rarely an escape elsewhere. Flowering and fruiting May through July. CANADA: NEW BRUNSWICK: Fredericton, Groh s. n. OTB). Nova scotia: Grand Pré, Groh s. n. (OTB); Kings Co., Centreville, McLellan 718 (ОТВ). охтако: Carleton Co., Nepean Twp., Brittania, Cody & Calder 504 (OTB); Frontenac Co., Battersea, Edmonson s. n. (NY); Carleton Place, Groh s. n. (OTB); Pickering, Haight s. ». (ОТВ); Colborne, Northumberland, Victorin, Germain, Dominique 46205 (GH, OTB, WYO). QUEBEC: near St. Clements, Montgomery 000 (OTB); La Malbaie, Marie-Anselme 165 (OTB); Pontiac Co., Bristol Twp., Lindsay ё Mulligan 135 (ОТВ); Saint-Mathias, comté de Rouville, Victorin & Germain 46630 (OTB). UNITED STATES: CONNECTICUT: Hartford Co., Southington, Bissell s. n. (MO). MICHIGAN: Cheboygan Co., Mackinaw City, Dodge s.n. (MU); Emmet Co., Carp Lake, Gates 15821 (US); Presque Isle Co., Rogers, Dodge s. n. (MU). MINNESOTA: St. Louis Co., near Moriey Park, Duluth, Lakela 2560 (MO); Winona Co., Winona, Freiberg s. n. (MO). NEW JERSEY: Hudson Co., Hoboken, Schrenk s. n. (MO); Hunterdon Co., Califon, Fisher s. n. (MO). NEW YORK: Albany Co.; Tompkins Co., Ithaca, Rowlee s. m. (MO); Warren Co., Queensbury, House 28755 (MO). VERMONT: Caledonia Co., Peacham, Blanchard s. n. (MO); Franklin Co., east Berk- shire, Clausen s. n. (MO); Windsor Co., Rochester, Dutton s. n. (MO). WASHINGTON: Spokane Co., Newman Lake, Suksdorf 8702 (WTC). Valeriana officinalis has been a favorite "old fashioned" garden plant for ages past in Europe, and it has been growing in American gardens for at least 150 years, without doubt since colonial times but a definite record of this is lacking. This species is listed in the United States Pharmaecopoeia as an adulterant in certain cardiac remedies. 2. VALERIANA SITCHENSIS Bong. in Mem. Acad. St. Petersb. 29:145. 1833. Т. Mertens s. n.! (D, GH, OXF, P, W). Perennials 3-10 dm. tall, from relatively stout rhizomes 2-7 mm. thick. Stem 0.5—1.0 cm. in diameter, glabrescent, the nodes 2—5, densely puberulent or pilosulous. Leaves basal and/or cauline; the basal relatively few to many, loosely tufted with the several adventitious shoots, or essentially wanting, petiolate, un- divided or pinnate to pinnatifid, ovate to ovate-oblong or suborbicular and more or less cordate, 10-35 cm. long, dentate to repand or entire, glabrous or pilosulous on the veins below, sometimes spreading-ciliate; cauline leaves 2-5 pairs, simulating the basal, pinnate to pinnatifid, 3—20 cm. long, the lateral lobes 1—6 pairs, shorter than the terminal lobe, grading smaller. Inflorescence in anthesis 2-8 cm. wide, later diffuse, 10-12 cm. long, 6—9 cm. wide, the nodes pilosulous; bracts 5.0—5.5 mm. long, glabrous or sparsely spreading-ciliate; flowers hermaphroditic or rarely gynodioecious. Corolla infundibuliform, 4.5—9.0 mm. long, white to pinkish, 1951] MEYER— VALERIANA IN NORTH AMERICA 393 Fig. 4. Valeriana sitchensis ssp. sitchensis: Habit X l4; entire and dissected flowers, and achene (abaxial side), X 5. glabrous or sometimes pilosulous towards the base of the tube without, the limb less than half as long as the gibbous tube, rarely 1 as long, the throat sparsely to somewhat densely pilosulous within. Stamens and style exserted. Achenes linear- to ovate-oblong, 3-6 mm. long, glabrous. Calyx-limb 11- to 23-fid. [Vor. 38 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN The subalpine and low altitude variants of Valeriana sitchensis were recognized as early as 1837 when Shuttleworth in his description of V. Hookeri distinguished two distinct taxa under this name—the Rocky Mountain form merely as “in sylvis ad Montes Scopulorum”, and the Pacific coast populations as var. В foliolis sub- integeris from the Columbia River. It is clear, however, that the Rocky Mountain plant conforms to Bongard's earlier description of this plant from Sitka as V. sitchensis, which in the present treatment applies to V. s. sitchensis. Under var. B, however, Shuttleworth clearly delimits, although not by name, V. sitchensis ssp. Scouleri, as may be easily interpreted from the type specimen. In north-central United States there occurs a third taxon, V. s. uliginosa. KEY TO THE SUBSPECIES А. Cauline NEM with 1—3(—4) pairs of lateral pne н terminal lobe obovate, ovate-rhombic to suborbicular, acute or B. Plants ai , to 12 dm. tall. Leaves 1. eriline, crenate t fir to irreg ularly repand-dentate to entire or -— so, firmly membranaceo glabrous or hirtellous on the v the terminal lobe 1.5-6.5 cm. wide. Corolla 4.5— ong, есч frequently xc dm towards the base of the tube without. Achenes mostly ova long-ovate, m А ide, t purpurascent, frequently purple-maculate. Subalpine he Kenai Peninsula and the coastal mountains of Alaska to east-central and сни British Columbia; Idaho апа western Montana; I the Siskiyou Mountains, Californ NUM 2a. V. s. sitchensis т т М Pls ants sod to 7 dm. tall. Leaves jer wenns basal, entire or essentially so, thinly membranaceous, glabrous, the terminal lobe 0.9-3.8 cm. wide. Corolla 5-9 mm. long, pinkish- а ірің ous without. Achenes mostly oblong-linesz, 5-6 mm ong, wide, tawny to rubiginose, rarely purpurascent. cue British Columbia to Mendocino Co., Calif., ане $еа- бе: to abou 4000 feet altitude, west of the C саде Moun . V. s. Scouleri AA. Cauline leaves with кдын —6 pairs of Т labs, th terminal de lanceolate to elliptic, acute to acuminate, 0.9 wide. Corolla —6 mm. long, 8, gla brous Mc Michigan to Ohio, Tue York, north to New Brunswick 2c. V. s. uliginosa 2a. VALERIANA SITCHENSIS Bong. ssp. SITCHENSIS. Valeriana Hookeri Shuttl. in Flora 20?:450. 1837. T.: Drummond s. n.! (СН, К, NY). Valeriana capitata Pall. ex Link 8 Hookeri Torr. & Gray, Fl. N. Am. 2:48. 1841, as to specimens cited in part m frigidorum Савт. in Bull. Soc. Bot. Fr. 65:37. 1918. T.:Cusick 1715! (Е, К, N, MO, UC, US, YTC). зе Suksdorfii Gdgr. 1. с. 36. 1918. T.: Suksdorf бобо! (WTC). Valeriana anomala Eastw. in Leafl. West. Bot. 3:22. 1941. T.: Howell 15162! (CAS). Perennials 3.5—12.0 dm. tall, robust. Stem leafy. Leaves predominantly cauline, 10-20 cm. long, pinnate to pinnatifid, the lobes crenate to irregularly repand-dentate or essentially entire, membranaceous, glabrous or occasionally spreading-ciliate to somewhat strongly hirtellous, the terminal lobe obovate, ovate- rhombic to suborbicular, 2.5—4.5 cm. wide, acute or obtuse, the lateral lobes 1—4 pairs; basal leaves petiolate, ovate-elliptic to obovate, 11-40 cm. long, glabrous, 1951] MEYER—VALERIANA IN NORTH AMERICA 395 `7 / "n ж” are : Bi. ! 4; А Ф. e je N =] VIRTA а < F B vr $- ] | ed fU ik | мү. Ls ^h « N { íi "OW ....... (m FS TA AO А о \ ЖАЛ! =A 5) ©... da ) YF t ы ecd p Can (> ӘОЖ 22/5 2 c ; WA А2 cS N ' SJ NADA % ^ M. : Q MU Fig. 5. т of V. sitcb- Fig. 6. Distribution of V. sitch- ensis ssp. sitcben ensis ssp. Scouleri Fig. 7. Distribution of V. sitchensis ssp. uliginosa. the terminal lobe 5—10 cm. long, 2-5 cm. wide. Corolla 4.5-7.0 mm. long, fre- quently somewhat pilosulous towards the base of the tube without. Achenes ovate to oblong-ovate, 3—6 mm. long, 2.0-2.1 mm. wide, tawny or purpurascent, fre- quently purple-maculate. Calyx-limb 12- to 20-fid. Type Locatiry: "Habitat in montanis, l'Isle Sitka” [now Baranof Island], Alaska. DISTRIBUTION: Subalpine meadows and open woodlands in the mountains from about 4000 feet altitude (rarely lower) in the southern part and above 1800 feet altitude in the northern part of the range. Kenai Peninsula, southward in the [Vor. 38 396 ANNALS OF THE MISSOURI BOTANICAL GARDEN Alaska coastal mountains to east-central and southern British Columbia to south- central Idaho and western Montana; Washington and Oregon to the Siskiyou Mountains, California. Flowering and fruiting June to September. son 6503 (GH, S, WYO); Mt. Crillon, Bates 146 (GH); McDonald Lake region, Burcham 56 (NY, US); Glacier Bay, Coville & Kearney 764 (US); Egg Island, Disenchantment Bay, Coville & Kearney 1037 (BM, US); Yakutat Вау, Coville % Kearney 1150 (US); Chugash Mts., Anchorage, Dutilly, LePage, O’Neill 20530 (S); Mt. Verstonia, Emmons s.n. (US); White Pass, Enander s. n. (S); Knight Island, Prince William Sound, Eyerdam Longfield & er s. n. (вм); РЕЧ Hyder, "McCabe 8416 (UC); Deer Mt., porn Island, McCabe 8610 (UC); Skagway, Macoun 91475 (NY); Sitka, Mertens H, K, MO photo, OXF, P, W); Mt. Roberts, jme Nelson 4430 (GH, US, WYO); "Chis, Norberg s. n. (S) ; Ketchikan, Willett 30 (W TA: Loggan, Bolley то (WYO); ыва une Brown 420 (GH, 05); headwaters of Saskatchewan & Athabasca Rivers, Brown 1450 (GH); Jasper House, Burke s.n. (K); Banff, Diehl s. n. (BRY); head of Smoky River, Hollister 9 (US); Pyramid Peak, near Henry House, Hollister 59 (US); Lake Louise, Hunnewell 3845 (GH); Mt. Edith Cavell, McCabe 8343 (UC); Mt. Paget, Macoun 65383 (GH, NY); Bertha Lake, Waterton Nat. Park, Moss 476 (GH, OTB); Jasper, Moss 2675 (GH); Mt. Quincy, Osthermer 88 (ОН). BRITISH coLuMBIA: Mt. Cheam, Anderson 4. pes Mt. Garibaldi, Bennett s. n. (E, K); Asulkan Valley, Glacier, Brown 230 (С O); Burgess "Trail Field, Brown 375 (GH, MO, NY, US); Cougar Mt., Butters, хм, еен 588 (MIN); Yoho Valley, Butters & Holway s.n. (MIN); McGillivray, Crows- nest Pass, ары 15774 (ОТВ); White Pass, Eastwood 897 (GH, 1 Mission, Fletcher 814% (OTB); Yale, Fletcher 052 (BM); Stanley, Fraser s. n. (NY); Summit Lake, Fyles - (OTB); Vancouver Island, Forbidden Plateau, Greig s. n. (MBG); coast range, Van- ouver, Henry s. n. (CGE); Falkland, Hitchcock & Мана 7520 (UC); Moraine Lake, ne. Barkerville, McCabe 44a (UC); Bowron Lake rd. near Barkerville, McCabe 418 (UC); 15 mi. sw. Kleena Беи McCabe 569 (UC); 8 ті. w. Clearwater, McCabe 2254а (UC); „ 8 ті. n. Vancouver, McCabe 2744 (UC); Alta Lake, McCabe 2942 (U ch 20 mi. sw. Kaslo, McCabe 4740 ne ); 5 mi. n. Crowsnest e McCabe 4900 ы ): 1 о + © - a e z: et oo ton, McCabe 8146 (UC); 15 mi. sw. Telegraph Creek, McCabe 8900 (UC); between N. Thompson & Bonaparte HE Macoun s. n. (GH, US); Skagit Valley, between 49? and 49^15'N, and 121° and 121^20"W, Macoun 72790 (NY); Cassiar Dist., near head inn ce- -уеһ River, branch of раа River, Preble & Mixter 646 (US); Mt. Selwyn, about 56°1'N., 123?39"W., Raup & Abbe 4170 (СН, S); Cameron River Valley, Vancouver Island, Rosendahl 1990 (CGE, K, MIN, MO); Toquin Valley, Jasper Park, Sanson 900 (NY); Great Northern Mt., Scheuber s.n. (US); Big Bend Dist, about 118°20'W., 51°45'N., Shaw 933 (GH, MIN, MO, NY, US); Bluster Mt., Marble Mts., Thompson 8 Thompson 410 (US); between Mt. Field & Mt. Wapta, Walcott s.n. (US); 27 mi. n. Natal, Weber 2282 (GH, NY, UC). NORTHWEST TERRITORIEs: near the Yukon border, about 62^30'N., 129^ W., Goodwin 26 (NY). UNITED STATES: CALIFORNIA: Humboldt Co., _ Summit, Tracy 5230 (UC); Siskiyou Co., Marble Mts., Howell us (CAS, MO p IDAHO: Adams Co., between P Meadows and 55. е 1144 (СН, МО, е Blaine Co., Alturas Ls Cronquist 2578 (MO); Boi ‚ Quartzburg, Malford s. (GH, MO, NY); Bonner Co., Priest River Exp. Bata. c5 6204 (MO); chos 1951] MEYER—VALERIANA IN NORTH AMERICA 397 Co., divide between St. Joe and po due River, Leiberg 1249 (ARIZ, ОН, К, MO, NY, UC, US, WYO); Custer Co., 25 mi. nw. Stanley, Cronquist 2772 (MO); Bore Co., 20 mi. n. Pine, Meyer & Meyer E (MO); Idaho Co., Hibbs Cow Camp, Dry Diggins, T23N, R2W, Packard 335 (UC); Kootenai Co., Wiessner’s Peak, Sandberg, MacDougal, 1335 (ARIZ, GH, MO, NY, UC, US, WYO); Valley Co., Sawtooth Mts., Thompson 13793 (D, GH, ee NY, UC, US); Washington Co., Cuddy Mts., Jones s. n. (PO). NTANA: Beaverhead Co., Bitter Root Range, 3 mi. above O'Neill' s е Hitch- cock d Mublick ІЛ (СН, МО, S, UC, WYO); Clark Co., Helena, Starz (MO); Deerlodge Co., Anaconda Mts., shore of Storm Lake, Hitchcock © Muhlick “Ж (МО); Flathead Co., Columbián Mt., Rogers © Rogers 1164 (MO, NY); Glacier National Park, Osterbout s. п. (MO); Granite Co., 2 mi. w. Skalkaho rd. summit, Hitchcock 9 Mublick 14464 (MO, UC); Lake Co., Mission Range, se. McDonald Lake, Hitchcock 16281 (UC); Madison Co., Tobacco Mts., Hitchcock 17048 (WTC); Meahger Co., Little Belt Mts., Flod man 804 (MIN, MO, NY, US); Missoula Co., Flathead 27% near Upper Holland Lake, Hitchcock 18440 (UC); Powell Co., Gordon Mt., 6 mi. s. Big Prairie Ranger Station, Hitcbcock 18842 (UC). OREGON: Baker Co. Anthony Lake, Blue Mts., Maguire & Holmgren 26015 (NY, UC, US); Crater Lake Nascondi Park, Heller 12956 (D, GH, MO, NY, UC, US); Crook Co., along a brook, Whited 648 (СН, К, MO, NY); Grant Co., Strawberry Mt., Hender- son 5688 (GH, MO); Hood River Co., Elk River, Mt. Hood, Benson 2540 ( US); в Co., Huckleberry Mt., Coville 9 Applegate 371 (NY, US); Feferon Co., 4 mi. s. Mt. Jefferson, Nelson 2813 (GH); Josephine Co., Siskiyou Mts., Whittaker 55100 (WTC); Klamath Co., Union Peak, Applegate 4770 (WILLU) ; Lane Co., McKenzie Pass, 5 mi. w. summit, Peck 9799 (GH, MO, WILLU); Marion Co., Jefferson Park, Т105, R8E, Sec. 11, Peters 170 (MO); Umatilla Co., near Langdon Lake, Peck 22281 (WILLU); Union Co., Wenaha National Forest, Bo Springs, Lawrence 208 (ARIZ, US); Wallowa Co., Buckhorn Springs, Peck 18315 (W WASHINGTON: Asotin Co., Blue Me near Big Butte, 9 mi. se. Anatone, Meyer 414 (MO); Chelan Co., Mt. Stuart, W hited 774 (US); Clallam Co., Deer Lake, Meyer 1042 (MO); Columbia Co., Blue Mts., Goodman Springs, Constance, Сї, Staats, Van Vleet 1163 (MIN, MO, P, WYO); Ferry Co., Graves Lookout, R35E, T36N, Boner & Weldert 234 (GH, MO, NY, UC, WYO); Ga che ld Co., Stentz Spring, T9N, R42E, Constance 9 оов 1735 (MO); Grays Harbor Со., tra 4% о Mt. Colonel Bob, Meyer 000 (GH O); Jefferson Co., Constance Ridge, Meyer 710 (MO); Kittitas Co., Table Mt., Meyer 3 Meyer 2234 (MO); Mason Co., Mt. Ellinor, Eyerdam 1262 (BM, D, MO); Okanosin Co., Big Craggy, Thompson 10851 (MO, NY); Pend Oreille Co., Pass Creek Pass, Laskey s.n. (WTC); Pierce Co., Chinook Pass, Eyerdam s. n. (MO); Mt. Rainier National Park, Paradise Valley, Meyer & Meyer 2240 (MO); Skagit Co., Mt. Baker Forest, Neff 506 (WTC); Skamania Co., Mt. St. Helens, Coville 762 Si Snohomish Co., alpine meadows of Mt. Dickerman, Thompson 8875 (MO); Spokane Co., Mica Peak, Suksdorf 8817 (WTC); nurap Co., Black Hills, Meyer 1637 (GH, MO, OTB); Walla Walla Co., Blue Mts., Piper s. (WTC); Whatcom Co., Mt. Baker ee Heller 14751 (D, MO, NY, US); Yakima Mt. Adams, Suksdorf "бобо (WTC Shuttleworth quite adequately recognized in his original description of V. Hookeri two categories which are in the present treatment recognized as subspecies under V. sitchensis. The synoptial chart below reveals the salient characters that distinguish these taxa: V. s. sitchensis V. s. Scouleri rura subalpine, on үл e = xd ке, 0—4000 ft., on basalt bluffs along ows, open or dens eams; flowering in spring. PAGS Mp "pese in summer. ABIT: robust, 3.5—12.0 dm. tall. slender, 3.5-7.0 dm LEAVES: л ш crenate to predominantly Бы, entire or essentially so. irregularly repand-dentate. CHENEs: ovate to oblong-ovate. oblong-linear. [Vor. 38 398 ANNALS OF THE MISSOURI BOTANICAL GARDEN Valeriana s. sitchensis is one of the conspicuous and quite attractive subalpine plants, especially in the Olympic and Cascade mountains of Washington state where it is commonly referred to as "Mountain Heliotrope". 2b. VALERIANA SITCHENSIS Bong. ssp. Scouleri (Rydb.) F. С. Mey., stat nov. Valeriana Hookeri var. 8 Shuttl. in Flora 20?:450. 1837. T.: eode к cá Pu NY). Valeriana capitata Pall. ex Link 8 Hookeri Torr. & Gray, Fl. 48. 1841. T.: Douglas s. n.! (BM , K). n Scouleri Rydb. in Mem. N. Y. Bot. Gard. 1:377. 1900. T.: based on V. Hookeri г. В Shuttl. Veleriane sitchensis Scouleri (Rydb.) Piper, in Contr. U. $. Nat. Herb. 11:533. 1906, Valeriana Adamsiana Eastw. in Leafl. West. Bot. 2:196. 1939. T.: Eastwood & Howell 1334! (CAS) Valeriana Follettiana Eastw. l. c. 197. 1937. T.: Follett s. n.! (CAS). Valeriana humboldtiana Eastw. l. c. 198. 1939. T.: Eastwood & Howell 4868! (CAS). Perennials 3.5—7.0 dm. tall, slender. Stem sparsely leafy. Leaves predomi- nantly basal, 3—10 cm. long, pinnate to pinnatifid or undivided, the lobes entire or essentially so, thinly membranaceous, glabrous, the terminal lobe of the divided leaves and blade of the undivided leaves obovate, ovate-rhombic to suborbicular, 0.9—3.0 cm. wide, the lateral lobes 2—5 pairs. Corolla 5.0—8.8 mm. long, glabrous without. Achenes oblong-linear, 5—6 mm. long, 0.9-1.8 mm. wide, tawny to rubiginose, rarely purplish. Calyx-limb 12- to 18-fid. Type Locariry: “Оп moist rock and islands of the Columbia River near Oak Point". DISTRIBUTION: Sea-level to about 4000 feet altitude on bluffs along forest streams west of the Cascade Mts. from southwestern British Columbia to Mendo- cino Co., Calif. Flowering and fruiting March to July. CANADA: BRITISH COLUMBIA: between Alexandra Bridge and Yale, McCabe 2504 (UC); Doyle wee Gordon Channel, beet: e 7122 (UC); Yale, Macoun s. n. (NY); Chilliwack Valley, 49^—49^10'N., 121725'-122^W., Macoun 64902 (NY); Skagit Valley, 49*—49^15'N., ае" Macoun d (NY); Vancouver Island, vicinity of Nanaimo, Macoun 88005 (NY); Vancouver Island, vicinity of Ucleulet, Mr. Fraser's garden, Macoun 88006 (NY); exposed cliffs along Cameron Lake, Vancouver Island, Meyer & Meyer 2246 (MO); Vancouver Island, Sookes River area, Nelson & Nelson 716 (MO, WYO); banks of Gordon River, Vancouver Island, Rosendahl & Brand s. n. (MIN); Port Renfrew, Vancouver Island, Rosendabl 1744 (MIN); Fraser River Canyon near Yale, Thompson & Thompson 10 (MO, NY, UC, US) UNITED STATES: CALIFORNIA: Del Norte Co., Mary Adams Peacock Bridge, along Smith River, East- wood 9 Howell 1334 (CAS); Humboldt Co., 5 mi. e. Berry Summit, Eastwood & Howell 4868 (CAS); Mendocino Co., along the creek at Elk, Follett s. n. (CAS). OREGON: Columbia Co., rocky bluffs below Clatskanie, Thompson 2450 (MO); Clackamas Co., Clackamas Lake, Peck 15850 (WILLU); Coos Co., Middle Fork Coquille River, Detling 4087 (UC); Curry Co., shady bank of Rogue River near Gold Beach, Peck 8005 (WILLU); Douglas Co., west fork Cow Creek Canyon, Peck 7424 (WILLU); Hood River Co., Cascades, Harford & Dunn 350 (MO); Jackson Co., Rogue River, Wood- 1951] MEYER—VALERIANA IN NORTH AMERICA 399 ville, Peck 7423 ddr eA Josephine Co., woods near Savage Rapids, 6 mi. from Grants Pass, Henderson 6058 (MO, WYO); Lane Co., Castle Rock Trail, Detling 2818 (UC); аса Co., cliff above the гел, Otter Crest, Peck 16333 (UC, WILLU); Linn Co., Willa- mette watershed, Cusick 4545 (WTC); Multnomah Co., Warrendale, Thompson 11870 (MO, NY); Polk Co., Mill Creek, 4 mi. sw. Buell, Peck 16221 (WILLU); Tillamook Co., Tillamook, Ferguson 5. п. (WI n WASHINGTON: Clallam Co., lower slopes of Mt. Angeles, Tbompson 7574 (GH, K, MO, UC); Grays Harbor Co., Montesano, P Heller © Heller 3937 (E, GG, GH, MO, NY, P; E E. Co., moist Woodi along Бс оре River, Thompson 6551 (К, МО, МҮ ; King Co., ‚ Green River Canon, Mosier s. ». (US); Klickitat Co., Larm River, os, s. n. (BM, D , E, GH, MO, P, US); Mason s Hoodsport, Otis 2359 (WTC); Pierce Co., Estonville, Flett 2204 (WTC); Skamania Co, C Chenowith, Suksdorf 1658 oe Snohomish Co., Perry Creek Trail, Thompson 14526 (GH, MIN, MO, NY, S, US); Thurston Co., Tumwater, Townsend s.n. (MO, UC, WTC); Wahkiakum Co., S St. John 8748 (GH, MO, UC); Whatcom “Со, “trail from Glacier to Mt. Baker, Mason 3894 (UC) The subspecies Scouleri is a low-altitude forest plant of the moist Pacific coastal climate west of the Cascade Mountains from British Columbia to northern California where it prefers Middle Tertiary basalt or sedimentary bluff habitats principally along forest streams from the sea-coast to the foot-hills of the Cascades. These specialized habitats occur, for the most part, locally through- out the distribution of the subspecies. The facies of these bluffs consist of masses of bare-faced or sparsely wooded rock cliffs, with a thin soil cover and accompany- ing conditions of reduced water supply—semi-drought in mid-summer. Under these specially provided conditions these bluffs support a characteristic "bluff flora" where they afford special means for isolation and may be alluded to as islands within the coniferous belt throughout the area where this plant occurs. Although the distribution of V. s. sitchensis lies within the realm of V. s. Scouleri west of the Cascade Mountains, these taxa remain effectively isolated from each other by marked habitat and altitudinal discontinuities. The ecological barriers ef- fectively isolate these subspecies so that interbreeding between them can be virtually ruled out; differences in flowering time increase the difficulty in this respect. At least I have seen no material that would suggest hybridization between these taxa. 2c. VALERIANA SITCHENSIS Bong. ssp. uliginosa (Torr. & Gray) F. G. Mey., stat. nov. Valeriana sylvatica B uliginosa Torr. & Gray, Fl. М. Am. 2:47. 1841. T.: Sartwell s. n.! ut A (Torr. & Gray) Rydb. ex Britton, Man. 878. 1901. Valeriana dubiosa Gdgr. in Bull. Soc. Bot. Fr. 65:37. 1918. T.: Beal s.n. Perennials 3—10 dm. tall, relatively slender. Stem leafy. Leaves predominantly cauline, pinnate to pinnatifid, 5.5—21.0 cm. long, the lobes dentate to repand or entire, membranaceous, spreading-ciliate, glabrous or sparsely pilosulous below, the terminal lobe lanceolate to elliptic, acute to acuminate, 0.9—2.5 cm. wide, the latera! lobes 5—6 pairs; basal leaves 20—35 cm. long, petiolate, undivided or with 1 pair of lateral lobes, ovate-elliptic to obovate, dentate to repand or entire, the blades and termina! lobe of the divided leaves 6.5—14.5 cm. long, 3.5—6.0 cm. [Vor. 38 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN wide. Corolla 5-6 mm. long, glabrous without. Achenes lanceolate, elliptic to ovate-oblong, 3—4 mm. long, glabrous. Calyx-limb (11-)15- to 23-fid. ТҮРЕ Locarirv: Lake Ontario, Wayne Co., М. Y. DisTRIBUTION: Marshy meadows, swamps, and bogs, Michigan, northern Ohio, New York and adjoining Canada, northward to Vermont and New Brunswick. Flowering and fruiting end of May through August. DA: NEW BRUNSWICK: Gloucester Co., Petit Rocher, Blake 5502 (GH, K, NY, P, US); Restigouche Co., Dalhousie, Poit 9 Alm 175 (S). ONTARIO: Proton, Quinn s.n. (OTB). QUEBEC: oo Co., New Richmond, Chrysler 1264 (US); Rimouski Co., Bic, Forbes s.n. (GH, WYO); St. Жн Williamson 1376 (NY); Lower Canada, Somerset, Brunet s. n. ( y NI d AINE: oostook Co., Perham, Steinmetz 782 (US); Penobscot Co., Staceyville, Allard 7210 (GH, US). GAN: Charlevoix Co., ne. Clarion, McVaugh 9409 (MU); ——— Co., Ecker- n, Dodge s. n. (MU); Baton Co., Olivet, herb. Coville (US); Emmet Co., Carp Lake, пие 15431 (US); Сепеѕее Co., Flint, Trencb s.n. (M с ы” © 55 ы I с с S ы ч a о" et e Р Jackson, Dodge s. ». (MU); Lapeer Co., Lang Lake, Dodge s. ». (MU); Livingston Co., Whitmore Lake, Ehlers 4747 (US); Luce Co., Lake Manistique, Hermann 8303 (MU, NY); Mackinac Co., Naubinway, McVaugh 10928 (MU); Wo ree Co., Negaunee, Rydberg s.n. (NY); Montcalm Co., . CM Davis s. n. (MU); Oakland Co., n Pontiac, Chandler s. n. (MSC, US); Otse о., 5 mi. e. Vanderbilt, McVaugh 10890 ; Schoolcraft Co., p pA Choix, ies 9 Bean 26347 (GH); Washtenaw Co., Dexter, Hermann 0455 (GH EW YO Columbia Co., A McVaugb 3118 (GH); Dutchess Co., Pine Plains, Hoysradt s. n. (GH, NY, UC, US); Genesee Co., Bergen Swamp, near visa, Johnson 4078 (US); Herkimer Co., Jordanville, Haberer 402 (GH, K, MO, NY); Madison Co., Syracuse, Sheldon 1144 (NY); Monroe Co., Mendon, Metcalf 8872 E Амы Co., зч Wiegand 5 Wrigbt 7208 (СН); рм Co., Sartwell s. п. (BM, GH, NY). о: Stark Co., Canton, Steele 2 (MO, US). VERMONT: Fairhaven, Eggleston 1275 өлі; K, MIN, МО, NY, P). Subspecies uliginosa prefers wet meadows and swampy areas іп the deciduous forest regions north of the southern extent of Pleistocene glaciation. "The broad (about 1200 miles) discontinuity that separates ssp. sitchensis and ssp. uliginosa may be likened to the distributions of other species in Hultén?*, such as Scheuch- zeria palustris, Festuca rubra, Equisetum scirpoides, and Gnaphalium uliginosum among others. Hultén attempts to outline a history of the modern distribution of these species on the assumption that the pre-Pleistocene distribution was con- tinuous and by the onset of the Wisconsin phase of the Pleistocene epoch the distributions became disjunct. Thus, many widely distributed species were left as remnants of the former distribution as modern inhabitants in western and eastern North America. Ншігеп essentially accepts the view that refuge areas existed within the glaciated regions of northeastern United States; or perhaps these species existed to the south of the glaciated areas and moved northward in post-glacial times. Modern glacial geologists do not give very much support to the refugia concept, and Flint?" does not locate any large refuge areas in north- 24 Outline of the History of Arctic and Boreal s dias the Quartenary Period. p. 123. 1937. 25 Glacial Geology and the Pleistocene Epoch. 1951] MEYER—VALERIANA IN NORTH AMERICA 401 eastern United States. The modern distribution of V. s. wliginosa is entirely within glaciated areas. Perhaps V. s. uliginosa sought its way to northeastern United States and adjoining Canada from western America contemporaneously with the final stages of Wisconsin glaciation. Subspecies uliginosa and sitchensis may be distinguished as follows: Ves е5 У. s. sitchensis Cauline pri with (3-)4-6 pairs of lateral Cauline leaves with 1-3(-4) pairs of lateral lobes; terminal lobe E дер to elliptic, lobes; terminal lobe obovate, ovate-rhombic acute to acuminate. to suborbiculsr; acute or obtuse 3. VALERIANA CAPITATA Pallas, ex Link, Jahrb. d. Gewachsk. 18:66. 1820. T.: Pallas 789 (in hb. Willd., photo in hb. MO). Perennials 1—6 dm. tall, from slender to relatively stout rhizomes 1—6 mm. thick. Stem leafy to more or less subscapose, 1—6 mm. thick, glabrous to uniformly pilosulous, the nodes consistently pilosulous. Leaves forming a loosely tufted rosette with the several adventitious shoots at the base, or obsolete, undivided, ovate to obovate-spatulate, 2-20 cm. long, 0.6—3.0 cm. wide, acute or obtuse, irregularly dentate to entire or essentially so, glabrous to puberulent; cauline leaves 3—5 pairs, undivided or pinnate to pinnatifid, ovate- to oblong-oblanceolate or obovate-spatulate, the uppermost much reduced, becoming bract-like, the blades and terminal lobe cf the divided leaves ovate to elliptic or suborbicular, 1-5 cm. long, 1.0—2.6 cm. wide, acute to obtuse, the lateral lobes 1—5 pairs, less than half as long as the terminal lobe, grading smaller, the petioles to 4 cm. long, grading shorter to nearly obsolete, more often spreading-ciliate towards the base. In- florescence dense in anthesis, 1.5—5.0 cm. wide, later diffuse, 3-14 cm. long, 3-9 cm. wide, nodes and internodes pilosulous or glabrous; bracts glabrous to sparsely spreading-ciliate; flowers hermaphroditic. Corolla infundibuliform, 3-8 mm. long, white to pinkish, glabrous to pilosulous without, the lobes half as long as the gib- bous tube, the throat sparsely pilosulous within. Stamens and style longer than the corolla limb. Achenes ovate, oblong to oblong-linear, 3.0—6.5 mm. long, glabrous or pilosulous, smooth, tawny, rubiginose, or purpurascent, often purple- maculate, abaxial ribs evident. Calyx-limb 10—17-fid. The modern distribution of Valeriana capitata is marked by a series of discon- tinuous distributions. With this as a basis, I have recognized four subspecies in the present interpretation of V. capitata, with V. c. capitata in the Arctic north of 60^, and three subspecies in western United States, V. c. acutiloba, californica, and pubicar pa. Hultén in his critical work on Arctic plants lists certain species in western America with disjunct distributions essentially like V. capitata. He mentions Anemone Drummondii, Potentilla rubricaulis, Thlaspi alpestre purpurescens, and Carex nudata in this category. An explanation is sought by suggesting for these species a former continuous distribution which extended from unglaciated areas, particularly in northern and central Alaska, through western Canada to areas in western United States that fall south of the maximum effect of Pleistocene glaci- [Vor. 38 402 ANNALS OF THE MISSOURI BOTANICAL GARDEN ation. The present disjunct distribution is thought to have been brought about during the maximum thrust of the continental ice mass which completely covered western Canada. Flint's?" map offers further evidence that during the maximum extent of glaciation in western Canada the Pleistocene ice field engulfed the whole of western Canada with the exception of the extreme northwest. The present distribution of V. capitata coincides adequately with geological data. KEY TO THE SUBSPECIES A. төзді со predominantly ovate and cauline, pong nd more or less inate, irregularly iun or toothed. es = eaves few, ae ym lateral offshoo Rhizome slender, 1—3 thick, rarely branched. Kodiak Mond Unalaska, _ ie the Bering Sea; interior Alaska to the Yukon Territory north of 60°; also in Arctic Asia to the Kola Peninsula, Scandinavia 3a. У. с. capitata AA. Leaves puberulent or glabrous, predominantly spatulate (sometimes ovate in V. c. pubicarpa), basal and cauline. Rhizome stouter, 3—7 B. Whole plant or the stem only puberulent. гез more чу roadly ovate- to obovate- aed ulate or sometimes ovate, obtuse, ap- proximately | n as broad as long. Lateral lobes of the divided leaves 1—4 C. Plants i мәдән то к — rudis ge terminal lobe of the divided leaves more often ncate retuse, regularly 3- to 7-toothed or ied — 4542 mm. long. Southern Oregon and the Sierra = mountains of California to Tulare Co.; also in reri Neva da 3b. V. c. californica CC. Stem puberulent, leaves essentially glabrous. — leaves and terminal lobes -— keseng Achenes 3 ong. Southwestern Monta yoming, жас "Idaho to т ы калы. DN to the Charleston Moun- tains, 3c. V. c. pubicarpa BB. hiv, Tied pestes glabrous. Leaves more often oblanceolate- tulate, twice to several times as long as broad, acute to more or 74е apiculate. Lateral lobes of Ки divided leaves 2—5 pairs, much reduced, long-acuminate d. V. c. acutiloba 3a. VALERIANA CAPITATA Pall. ex Link ssp. CAPITATA. Valeriana bracteosa Britt. in Bull. N.Y. Bot. Gard. 2:183. 1901, non Philippi. T.: Williams Y). s. n.! Valeriana v Pall. ex Link var. bracteosa (Britt.) Hult. Fl. Alaska & Yukon, pt. 53 Perennial 1-6 dm. tall. Rhizome slender, 1—3 mm. thick. Stem glabrous or pilosulous towards the base, glabrescent above. Leaves predominantly cauline, 3.0—5.5 cm. long, 1.3—5.0 cm. wide, ovate to obovate, acuminate, the lowermost pair more often undivided, petiolate or subsessile, obtuse, pinnate above, sessile or subsessile, lateral lobes 1—2 pairs, usually much reduced, sharply and irregularly dentate or essentially entire, glabrous, the terminal lobe ovate to suborbicular, 1.5-6.0 cm. long, 1.0-2.6 cm. wide, the lateral lobes less than twice as small, grading smaller; petiole 0.5-1.5 cm. long or nearly obsolete. Inflorescence 1.5—3.5 cm. wide in anthesis, subglobose; bracts exserted midst the flowers and fruits, *6Flint, R. F. 1. с. p. 23. 1951] MEYER—VALERIANA IN NORTH AMERICA 403 filiform to linear, glabrous, usually purpurascent. Corolla 5—7 mm. long, glabrous without. Achenes ovate- to elliptic-oblong, 3-4 mm. long, 1.5-2.5 mm. wide, glabrous, rubiginose or purpurascent. Calyx-limb 12- to 15-fid. Tyre Locauity: "vom hochsten Gipfel der Sajanischen Alpen.” I was unable to locate the Sajanischen Alpen on Pallas's пар”! to his ‘Reise’, but the name “Sajan Mountains" still exists for a mountain range separating Tannu Tuva and Siberia in lat. 52—54^N., long. 100°E. вер» оарі баба Fig. 8. Distribution of V. capitata ssp. capitata. DISTRIBUTION: Tundra in the extreme north and in woods in the southern part of the range. Islands of the Bering Sea and the whole of Alaska, except the outer Aleutian Islands and southeastern Alaska; Yukon Territory to the Mac- kenzie River and north of 60°. This plant occurs across Arctic Asia to the Kola Peninsula, Scandinavia. Flowering and fruiting June to August. ALAsKA: Tofty, Adeney s.m. Eo Humboldt Harbor, Albatross Exp. s.n. (US); Herendeen Bay, Albatross Exp. s. n. (US); Fairbanks, Anderson (NY); Ft. St. Michaels, Bannister s. n. (GH oe Kodiak Tand, Bean s. n. (US); Agiapok River, ba s.n. (МҮ); Cape Nome, Blaisdell s.n. (GH, NY); between Cook Inlet and Rampart City, Brooks 9 Prindle з.п. (US); Teller, ibe 71 (US); St. Lawrence Island, Chambers 35 (US); Upper Matanuska Valley, Chaney 90 (MO); Anvik, Chapman 65 (GH, NY); York, 27 Reise durch verschiedene Russischen Reiche. St. Petersburg. 1771. [Vor. 38 404 ANNALS OF THE MISSOURI BOTANICAL GARDEN ward Peninsula, Coats І (US); Chunik, Seward Peninsula, Collier s. n. (US); Eagle, Colier 37 (05); Nunivak Island, Collins s.n. (US); Port т Coville 9 Kearney 0 (US); Hall Island, Coville 9 Kearney 2020 (US); Mt. McKinley National Park, Dixon 57 (UC, US); Glen road, mile 124, Dutilly, LePage, O’Neill 20423 (OTB); so. Anchorage, Dutilly, LePage, ONeill 20727 pd White River valley, Eaton s. n. (US); Ladue Valley, Eaton s. п. (US); no. Mt. St. Elias, Eaton 40 (US); Kussiloff, Evans 631 (US); Fox Bay, Evermann s. n. (US); St. Paul Island, Evermann 84 (US); St. George А Н); St. Matthew, Gabrielson s. п. (СН); ge Lake, Gabrielson s. п. (GH); Unga, Golder 87 (US); Шатпа River, Gorman 37 (P, US); Katmai Region, Hagelbarger 77 (US); Good- news Bay, гооо 34 (US); SRuifiagin Island, Harrington s. n. (MO); Ft. Gibbon, Karluk, Horne s. n. (NY); Seward, Hutchison $6 (BM); Unimak Island, Hutchison 333 (BM); p Jordal 2076 (05); Bettles, Jordal 2454 (US); Popof Island, Kincaid s.n. (US); Kotzebue Sound, Lay 9 Collie s. n. (BM); McGrath, Layden 16 (US); Norton rn Жасаш s.n. (UC); Nushagak, е s.n. (US); Kowak River, McLevegan s. n. (US); Cape Lisburne, Mason s. n. (GH, K UC); TM 6 Menden- ball s. n. (US); Sheenjek к ни s.n. ed 65 “65° 30'N., 2°W., Merti ie Point Barrow, Murdock s. n. (US); Nelson Island, Palmer 183 (US) ; Ft. Hamlin, Piper s.n. (US); 1415: Meridian, 3 mi. from Arctic Ocean, Pope 3 (US); Little Diomede Island, Porsild 1719 (GH, US); between Point Barrow and HR River, Pullen s. n. CGE); кн, Rader 83 ( US); Yukon Delta, Russell s. ». (US); Kuskokwim Basin, Schmitt 44 (US); St. Michael, Setchell s.n. (UC); Putnam River, Stoney s. n. (US); Piels River, Taylor s. n. (US); 68°50'-69°15'N., 160 52 Tbompson s. n. (US); Porcupine River, Turner s. n. (UC); Point Hope W bite s. ). ANADA: NORTHWEST TERRITORIES: Arctic Red “tea "Taklovik; Gates & Mellenby s. п. (К); MacPherson, Smith 103 (к). YUKON TERRITORY: King Point, Anderson s. n. (NY); Dawson, MP 315 (GH, UC, US); Moosehide, Eastwood 488 (GH, UC, US); 63°57'N., 135° , Gillett 6 2. 4324 (ОТВ); Herschell Island, Lindstróm s. n. (NY); Сі Duy Macoun 01474 (NY); Ft. Selkirk, Tarleton Тота (NY); Dawson, Williams s. n. (NY). Valeriana c. capitata is wholly arctic or subarctic in distribution, occurring in the most northerly latitudes of any member of the genus. This taxon may be readily distinguished by its nearly sessile leaves, comparatively slender habit, and showy white flowers. Its distribution includes the whole of Arctic Asia, but in North America I have seen no material from the glaciated region east of the Mackenzie River delta, although it is apparently quite common over the whole of Arctic Alaska. This subspecies is separated by a geographical discontinuity of nearly 1500 miles from V. capitata in western United States where the history of the intermediate area is most intimately associated with Pleistocene glaciation. No collections of V. capitata from the intermediate area are known at this time. 3b. VALERIANA CAPITATA Pall. ex Link ssp. californica (Heller) F. С. Mey., stat. nov. Valeriana californica Heller, in Muhlenbergia 1:60. 1904. T.: Heller 7156! (BM, D, E, G O, NY, P, S, UC, US). Velerine енды Piper, in Smiths. Miscel. Coll. 50:202. 1907. T.: Coville & Applegate ! (GH, MO, US). 1951] MEYER—VALERIANA IN NORTH AMERICA 405 Valeriana seminuda Piper, in Proc. Biol. Soc. Wash. 37:95. 1924. T.: Coville & Funston 1486! (K, NY, US). Valeriana sylvatica Banks var. glabra Jepson, Man. Fl. Pl. Calif. 970. 1925. T.: Culbert- son 4376! (GH, K, MO, NY, UC Valeriana W biltonae Eastw. in Leafl. West. Bot. 3:24. 1941. Т. Winblad ѕ.п.! (CAS). Perennials 2—6 dm. tall. Rhizome relatively stout, 3—7 mm. thick. Stem uniformly puberulent to pilosulous towards the base, glabrescent above. Leaves predominantly basal, usually forming a rather loosely tufted rosette with the adventitious offshoots, mostly undivided or pinnate to pinnatifid, oblanceolate- to obovate-spatulate, 4.5—15.0 cm. long, 0.8—2.5 cm. wide, truncate or retuse at the tip, frequently 3- to 7-toothed or essentially entire, acute, firmly membranaceous, uniformly puberulent, or merely ciliate or glabrous, the lateral lobes of the divided leaves 1-4 pairs; cauline leaves petiolate, pinnate to pinnatifid, 2.5-8.0 cm. long, rarely simple; petioles (1.5-)2-4 cm. long. Inflorescence 1.5—2.5 cm. wide in anthesis; bracts sometimes sparsely ciliate. Corolla 3.0—5.5 mm. long, glabrous or sometimes pilosulous towards the base of the tube without. Achenes elliptic to ovate-oblong, 4.0—6.5 mm. long, 2.0-2.8 mm. wide, glabrous or pilosulous. Calyx- limb 12- to 17-fid. Tyre Іослілтү: "Ridge south of Donner Pass at 8500 ft., August 12, 1903”, Nevada Co., Calif. NN ssp. californica {ззр. pubicarpa ssp. acutiloba e Fig. 9. Distribution of the subspecies of V. capitata in western United States. [Vor. 88 406 ANNALS OF THE MISSOURI BOTANICAL GARDEN DISTRIBUTION: Subalpine slopes, meadows, and along creek banks, also in conif- erous woods, 5000—10400 ft. alt. Southern Oregon and the Sierra Nevada Mts. of California to Tulare Co.; also in adjoining Nevada. Flowering and fruiting June to August. UNITED STATEs: CALIFORNIA: Alpine Co., Hansen 725 (BM, D, GG, K, MO, NY, P, US); n Co., Jonesville, Copeland 481 (ARIZ, BM, D, СН, К, MIN, MO, NY, P, UC, W); Eldorado Co., Echo Summit Pass, Robbins 1228 (05); Fresno Co., dd Creek, Bot E d 413 (E, GH, K, MIN, MO, NY, UC, US); Humboldt Salmon EIUS dens 14381 (UC); Modoc Co., n. end Warner Mts., Alexander 8 m 46 50 (UC, US); Mono Co., ua Grade, Winblad s. n. (CAS, MO photo); Nevada Co., Donner Pass, "Heller 7156 (ВМ, Р ‚ E, ОН, К, MIN, MO, NY, P, S, UC, US); Placer Со.; Mt. NEVADA: Douglas Co. ‚ Spooner, Baker ey (D, GH, MO, NY, UC, US, W); Ormsby Co., Marlette ag) Train P (NY, UC); Washoe Co., so. Slide Mountain, Heller 10020 (GH, MO, NY, UC, US OREGON: Crater Lake National Park, Coville & Applegate 340 (GH, MO, US); Jack- son Co., Siskiyou Mts., Whittaker 259 (WTC); Lake Co., Crane Mt., Thompson 13235 (MO, NY, UC, WILLU). Plants of the subspecies califormica are usually puberulent throughout, with leaves usually obovate-truncate to retuse and frequently 3- to 7-toothed. These characters, along with the geographical discontinuity, sharply distinguish the Sierran subspecies, and the Californian populations might well be maintained under full specific rank, but to do this would obscure the natural relationships of the Californian, the Rocky Mountain, and the Arctic elements of V. capitata. 3c. VALERIANA CAPITATA Pall. ex Link ssp. pubicarpa (Rydb.) F. G. Mey., stat. nov. Yero pubicarpa Rydb. in Bull. Torr. Bot. Club 36:697. 1909. T.: Rydberg t$ Carlton 7717! ,US, WYO). Valerlene buberulenta Rydb. l.c. 1909. T.: Rydberg & Carlton 7065! (GH, MIN, NY, Valeriana Cusickii Gdgr. in Bull. Soc. Bot. Fr. 65:36. 1918. T.: Cusick 2131! (D, E, GH, N, M TC) Valeriana utahensis Gdgr. l. c. 37. 1918. T.: Linford s.n, Valeriana maculata Eastw. in Leafl. West. Bot. 3:23. 1941. T.: Alexander бтос! (CAS, UC). Perennials 1.5-5.4 dm. tall. Rhizome relatively stout, 2-6 mm. thick. Stem puberulent to densely pilosulous or glabrescent. Leaves predominantly basal, un- divided or rarely divided, ovate or oblong to obovate-spatulate, 3-21 cm. long, 0.6-3.0 cm. wide, acute or obtuse, essentially entire, glabrous or sometimes spread- ing-ciliate towards the subpetiolar base; cauline leaves petiolate, the lowermost undivided those above pinnate to pinnatifid, 4.2-10.0 cm. long, the lateral lobes 1—3 pairs, lanceolate to elliptic, to 8 mm. wide; petioles about 0.4—3.0 cm. long. Inflorescence 1.5-5.0 cm. wide in anthesis; bracts glabrous, sometimes purpur- 1951] MEYER— VALERIANA IN NORTH AMERICA 407 ascent. Corolla 4—7 mm. long, the tube mostly pilosulous, rarely glabrous without. Achenes linear- to lanceolate-oblong, 3.5-5.5 mm. long, 1.5-2.0 mm. wide, pilosulous or glabrous, frequently purple-maculate. Calyx-limb 11- to 17-fid. Tyre Іослілтү: Mount Nebo, Utah. August 15, 1905. DISTRIBUTION: On limestone granitic soils, stony sagebrush slopes or beneath pines and on talus slopes in the mountains, 6200—11800 ft. alt. Southwestern Montana, western Wyoming, central Idaho to northern Utah; southeastern Oregon to the Charleston Mts., Nevada. Flowering and fruiting June to August. cotorapo: Hohus Peak, Tweedy 4555 (NY, US). IDAHO: Blaine Co., Lost River Mts., Macbride & Payson 3154 (GH, MO, NY, US, WYO); Bonneville Co., Caribou Mt., Payson & Armstrong 3512 (GH, MO, WYO); Clark Co., Kilgore, Cronquist 1428 (MO); Custer Co., Mackay, Nelson & Macbride 1405 (GH. MIN, MO, WYO); Fremont Co., Henry Lake, Payson & Payson 1088 (GH, MO, NY, WYO); Lemhi Co., Eighteenmile Peak, Davis ЕА (UC); Owyhee Co., Silver City, Mac- bride 932 ae E, GH, MIN, MO, NY, UC, US, WY MONTANA: Beaverhead Co., Lima, Rydberg 2704 (NY); Madison Co., Midway Station, Nelson & o 5455 (MO, NY). NEVADA: Clark Co., Charleston Mts., Lee Canyon, Clokey 7734 (ARIZ, BM, BRY, CA, GH, MAT, MIN, MO, NY, OTB, US); Elko Co., Jarbridge, Nelson & Macbride 1046 (GH, MIN, МО, МҮ, S, US, WYO); Esmeralda Co., David Davis Ranch, Shockley 518 (GH); Eureka Co., Eureka, Train 255 ied Lender Co., Toiyabe Range, Kennedy 4182 (MO); Nye Co., Тоз Range, Maguire 9 Holmgren 25076 (ARIZ, GH, NY, UC, US); White Pine Co., Ruby Range, Hitchcock % Martin 5662 (NY, UC, OREGON: Crook Co., Ochoco Forest, Peck 17181 (WILLU); Harney Co., Steens Mts., pd 2131 (D, E, GH, K, MIN, MO, P, UC, US, WTC). AH: Cache Co., w. ridge Spring Hollow, Maguire 13803 (GH, NY); Juab Co., Dus Creek Range, Нот? 37446 (NY); Piute Co., Marysvale, Rydberg & Carlton 7065 (GH, MIN, NY, US); Salt Lake Co., Sunset Mts. „ Maguire 17441 (NY); Utah Co., Mt. Timpanogos, Harrison 0358 (MO); Wasatch Co., Provo, Goodding 1148 (D, GG, GH, MO FN; 2, UG) ШӘЛ ҮС)» WYOMING: eee Co., Afton, Payson & Armstrong 3339 (GH, MO, WYO); Sub- lette Co., Gros Ventre Mts., Payson & Payson 3026 (GH, MO, NY, UC, US, WYO); Teton Co., Teton Pass, Hall 11460 (UC). Valeriana c. pubicarpa may be distinguished by its uniformly puberulent stem and predominantly entire and glabrous leaves. It is confined to the mountains of the Great Basin of the intermountain region west of the Continental Divide. On its western perimeter ssp. pubicarpa nearly makes contact with V. с. californica, and the following variant bears mention: (Syn. V. maculata Eastw. in Leafl. West. Bot. 3:23. 1941). ra depauperate, 1.5— 2.5 dm. tall; leaves mostly undivided, 3.5—9.0 cm. long, 0.6-1.1 cm. wide, Charleston Mts. and Toiyabe Range, Nevada. 3d. VALERIANA CAPITATA Pall. ex Link ssp. acutiloba (Rydb.) F. G. Mey., stat. nov. Valeriana acutiloba Rydb. in Bull. Torr. Bot. Club 28:24. 1901. T.: Rydberg 9 Vreeland (NY 5576! Valeriana Crandallii Gdgr. in Bull. Soc. Bot. Fr. 65:36. 1918. Т,: Crandall 2080! (CA, MO photo, NY, US, WYO). Valeriana glacialis Gdgr. І. c. 1918. T.: Crandall s. n.! = n. 2076 (CA, NY, US). [Vor. 38 408 ANNALS OF THE MISSOURI BOTANICAL GARDEN Perennials 1.5—6.0 dm. tall. Rhizome relatively stout, 2-4 mm. thick. Stem glabrous or glabrescent. Leaves predominantly basal, more often numerous and forming a rather loosely tufted rosette, undivided or rarely divided, predomi- nantly oblong-, oblanceolate- to obovate-spatulate, 3.5—36.0 cm. long, 1.2-2.3 cm. wide, short-acuminate or acute to more or less apiculate, glabrous or some- times spreading-ciliate towards the subpetiolar base; cauline leaves essentially sessile, pinnate to pinnatifid, the lowermost sometimes undivided, 1,5-7.0(-12) cm. long, the lateral lobes 2—5 pairs, linear to oblong-linear, grading smaller, becoming more or less filiform, 1-3 mm. wide, petioles obscure. Inflorescence 1.5—3.5 cm. wide in anthesis; bracts glabrous. Corolla 4-8 mm. long, glabrous or sometimes pilosu- lous towards the base of the tube without. Achenes ovate to oblong-lanceolate, 2.5-5.0 mm. long, 1.5-2.0 mm. wide, glabrous, frequently purple-maculate. Calyx-limb 10- to 14-fid. ТҮРЕ LocaLmY: coLorapo: Near Gray-Back Mining camps and Placer Gulch, Sangre de Cristo Range, Custer Co., 2600—2800 m. alt. June 25-27, 1900. DISTRIBUTION: Wet meadows, open woods, along stream sides, grassy and rocky slopes, 8000-13000 ft. alt. Southeastern Wyoming, Colorado, New Mexico and Arizona. Flowering and fruiting May to August. ARIZONA: Apache Co., White Mts., е 6 Lone 3260 (ARIZ); Coconino Co., San Francisco Mts., Peebles 6 Smith 13506 (М coLorapo: Boulder Co., Fourth July ted rd 12154 (UC); Chaffee Co., 5 n above St. Elmo, Rollins 1371 (D, GH, MO, NY); Clear Creek Co., Brookvale, Churchill Co es Peak road, Wiegand © Upton 4345 ( ; Co., Sierra Sangre de Cristo, Brandegee 700 (NY, UC); Garfield Co., near Trappers Lake, Hanna 142 O); Gilpin ‚ James Peak, Cox 3 O); nnison Co., Carson, Baker 313 (D, E, GH, Co., Mt. Иш, Baker, Earle & Tracy 258 (BM E, GH, К, MIN, MO, NY, US, W, WYO); Park Co., sg Mile Canyon, Killip ban. (MO, de Routt Co., Anita Peak, Goodding 1760 (E, GH, MO, NY, US, WYO); San Juan ‚ Silverton, Crandall 2080 (CA, MO photo, NY, Us. WYO); Summit Co., .. Brandegee 102 (MO, NY, UC). Piles MEXICO: Catron Co., mi. e. Mogollon, Meyer & Meyer 2201 (MO); county own, Pecos а Bailey Pe (US); county unknown, Pecos River National [oed nh 4182 (US). WYOMING: Albany Co., Laramie Mts., Porter 1062 (MO). Valeriana c. acutiloba is most easily distinguished by its leaves, which are oblong-, oblanceolate- to obovate-spatulate, short-acuminate or acute to apiculate. This is the only subspecies of V. capitata in which lanceolate leaves predominate. Leaf shape, which basically offers the best character for subspecies differentiation in V. capitata, is summarized as follows: V. с. capitata V. c. pubicarpa V. c. californica V. c. acutiloba Ovate to obovate , Ovate or oblong to E ог —_ Oblong- to oblanceo- obovate-spatulate late-spatulat late-spatulate 1951] MEYER— VALERIANA IN NORTH AMERICA 409 4. VALERIANA ARIZONICA Gray, in Proc. Am. Acad. 19:81. 1884. T.: Palmer s.n.! (GH, MO). Ші ovata Rydb. in Bull. Torr. Bot. Club 31:645. 1904. T.: Clements & Clements ІІ b. ‚ MIN, MO, NY, OXF, US, WYO). Valeriano acutilobe Rydb. var. ovata (Rydb.) A. Nels: Man. Bot. 476. 1909, Perennials 1—3 dm. tall, from rather slender rhizomes 1—4 mm. thick. Stem 2-5 mm. in diameter, glabrous, the nodes puberulent. Leaves predominantly basal, forming a loosely tufted rosette with the several adventitious shoots, petiolate, un- divided or sometimes pinnate, ovate to suborbicular, rarely subcordate, 2.5-17.0 cm. long, entire or essentially so, glabrous, acute or obtuse, the blades and terminal lobe of the divided leaves 0.8—6.5 cm. long, 1.5—3.0 cm. wide, the lateral lobes 1—3 pairs, distinct, grading smaller, the petioles 1.5—5.5 cm. long; cauline leaves 2—3 pairs, pinnate to pinnatifid, 2.0—4.6 cm. long, sessile, the uppermost much reduced, the terminal lobe elliptic, oblanceolate to obovate, acute to subobtuse, 1.2-3.2 cm. long, 0.9—1.9 cm. wide, the lateral lobes 1-3 pairs, grading smaller. Inflorescence dense in anthesis, 2.5—3.0 cm. wide, later expanding to about 5 cm. wide; bracts 5-7 mm. long, glabrous; flowers hermaphroditic or rarely gynodioecious. Corolla infundibuliform, 5—15 mm. long, white to pinkish, glabrous without, the tube gibbous, the lobes 2.0-2.5 mm. long, the throat sparsely pilosulous towards the base within. Stamens and style exserted. Achenes ovate to oblong-lanceolate, 2—5 mm. long, glabrous, tawny or purpurascent. Calyx-limb 10—12-fid. 2 Қыс... ^ --. Ж ө МЕГ Fig. 10. Distribution of V. arizonica. [Vor. 38 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN Type Гослілтү: Mountains near Prescott, Arizona. April 20, 1876. DistRIBUTION: Dry rocky ravines or shady damp banks in the mountains, 5400—9000 ft. alt. Southern Colorado, New Mexico to Arizona; also in northern Chihuahua, Mexico. UNITED STATES: ARIZONA: Cochise Co., Chiricahua Mts., Goodman © Hitchcock 1183 (MO, NY, WYO); Coconino Co., San Francisco Mts., Purpus 7056 (UC, US); Gila Co., Matzatzal Mts., Collom 257 (ARIZ, E, GH, MO, NY, US); Graham Co., Graham Mts., Wiegand % Mis е 2264 (GH); Greenlee Co., 35 mi. n. Clifton, Maguire, Richards, Moeller 11801 (СН. NY); Maricopa Co., mts. near Phoenix, Norville s. п. (MO); Navajo Co., between Carrizo and Showlow, Foster & Arnold 280 (GH, MO, US); Pima Co., Santa Catalina Mts., Shreve 5428 (ARIZ, GH, UC); Yavapai Co., Prescott, Palmer s.n. (GH, MO). COLORADO: Fremont Co., between Cafion City and Cripple Creek, Maur 8 Wiegand 2265 (GH); Huerfano Co., Cameron's Cone, Clements & Clements 241 (E, e MIN, MO, NY, OXF, US, WYO); Las Animas Co., Stonewall, Johnston 826 (GH : NEW MEXICO: Catron Co., vp Mts., Meyer = Meyer 2201 p. p. (MO); Colfax En г St. Jobn s. n. (GH); D na Ana Co., Organ Mts., Wooton s. n, (US); Grant mi. n. Pinos Altos, McVaugh 8054 (MU); ; Hidalgo бо, Animas Mts., McVaugh Boos "tMU); Lincoln Co., Eagle Creek, Wooton s. n. (US) ; Otero Co., Ruidosa, Hinckley s.n. (GH); Sandoval Co., Pueblo Canyon near Lot Alamos School, xx 20040 С MIN, МО, NY, P, US); San Miguel Co., Pecos Valley, Eggleston 10008 (GH, МО, NY, US); Sierra Co., Kingston, Metcalfe 1580 (MO); Socorro Co., Mogollon Mts., Wooton s. n. (US); Valencia Co., San Mateo Mts., Herrick 532 (US). MEXICO: CHIHUAHUA: Sierra Madre, Townsend ©] Barber s. п. (US). Valeriana arizonica may be distinguished essentially on the basis of the ovate leaves and the deep pink corolla which may attain a length up to 15 mm. This species has been much confused with V. capitata acutiloba, but these taxa may be consistently distinguished by leaf shape. The populations of V. arizonica in Colo- rado and New Mexico consistently maintain a corolla length of 5—8 mm., and this form was described by Rydberg as V. ovata. This variant also occurs in Arizona, but there it has corollas 11-15 mm. long. This form includes the type of V. arizonica. 5. VALERIANA PAUCIFLORA Michx. Fl. Bor. Am. 1:18. 1803. T.: Michaux s. n.! (MO photo, P). Perennials 5—10 dm. tall, from an abbreviated rhizome 2—5 mm. thick; stolons 2-3, radiating from the caudex, 3—7 dm. long. Stem leafy, 2-5 mm. in diameter, glabrescent, the nodes sparsely pilosulous. Leaves predominantly cauline, 4—6 pairs, petiolate towards the base, sessile towards the inflorescence, pinnate, ovate to ovate- cordate, 5—18 cm. long, the segments crenate- to serrate-dentate to repand or essentially entire, spreading-ciliate or scattered-pilosulous, the terminal lobe ovate to suborbicular, short-acuminate, cuneate to subcordate, 1.5-5.5 cm. wide, the lateral lobes distinct, more or less rhombic, 1—4 pairs, 1.8—5.5 cm. long, 0.7-2.7 cm. wide, grading smaller towards the internodes; leaves of the stolons undivided, ovate-cordate or pinnate and obovate, 2-25 cm. long, crenate-dentate, sometimes scattered-pilosulous or merely spreading-ciliate. Inflorescence 2.5—5.0 cm. wide in anthesis, later diffuse, 3-10 cm. long, 4—6 cm. wide, the nodes densely tufted- 1951] MEYER—VALERIANA IN NORTH AMERICA 411 pilosulous; bracts 5-7 mm. long, relatively long spreading-ciliate; flowers hermaph- roditic. Corolla subsalverform, 13—19 mm. long, more or less filiform towards the base of the tube, bluish-pink, glabrous without, the tube gibbous, the limb 2.0-2.5 mm. long, the throat sparsely pilosulous within. Stamens and style exserted. Achenes elliptic to ovate-oblong, about 5 mm. long, 2.0-2.5 mm. wide, tawny, sparsely pilosulous on the midveins, the margins narrow-winged, abaxial ribs evi- dent. Calyx-limb 14- to 15-fid. Түрк Locarrrv: “in montosis nemoralis Tennessee prope Flinn-Creek. Junio florens". DISTRIBUTION: Moist rich woods along streams and in river flats in alluvial soil or rich wooded hillsides. Southern Illinois eastward to the Ohio Valley, Ken- tucky, Tennessee, Pennsylvania, Virginia, and Maryland. Flowering and fruiting April to June. Fig. 11. Distribution of V. pauciflora. UNITED STATES DISTRICT OF COLUMBIA: boh of the Potomac, Morong s. п. (MO). ILLINOIS: Johnson Co., Tunnel Hill, Palmer 15158 (MO); Richland Co. Ridgway 1508 (MO, US); Union Co., Meyer 2171 (MO); Vermillion Co., Danville, Gleason s. n. (D, GH); Wayne Co., Hilgard s. n. (MO INDIANA: Allen Co., St. Mary's River, Deom IOI7 (E, MIN); Decatur Co., Greens- burg, Grover s.n. (M IN); Jefferson Co., Madison, Barnes s. n. (GH); Knox Co. Bieg cennes, Ridgway 731 (GH, K, P); Lawrence Co., Oolitic, Sbinners 1572 (UC, О): Marion Co., Brendonwoods Friesner 18852.2 (GH , UC); Montgomery Co., ДАНА Evans s. п. (MIN) ; Morgan Co., Bradford Farm, Wynn 20 (MIN); Orange Co., Orange- ville, Deam i (MIN); Putnam Co., Bainbridge, Dawson 1261 (MIN); Tippecanoe mi. n. Soldier’s Home, Lafayette, Hermann oe (MIN); Warren Co., Independence, Deam 53807 (MIN). KENTUCKY: Carter Co., Tygart’s River, Gilbert 964 (GH); Hardin Co., Fort Knox, Wadmond 7337 (MIN); Laurel Co., Mershons, McFarland 88 (GH, MIN, MO, UC, о Mammoth Cave National Park, Lix 212 (US); Meade Co., Tioga Springs, Curry s. m. (GH); Powell Co., Natural Bridge, Azderson 113 (GH); Warren Co., Barren River, Price [drawing] (MO). AND: Cecil к Octoraro Station, Tatnall 4515 (GH); Montgomery Co., Glen Echo, Painter 1331 (MO, WY OHIO: Belmont Co., oe ani Laughlin 967 (GH); Champaign Co., Urbana, collec- tor unknown (MO); Clermont Co., Loveland, James s.n. (NY, UC); Hamilton Co., 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ду, Ns A 35 P dA Fig. 12. Valeriana pauciflora: Habit, X 18; inflorescence, X 74; single flower, X 1%. е» Lloyd s. n. (FI, GH, MO, NY, Р, W); Lorain Co., Oberlin, Dick s. n. (GH); Meigs Co., Jones s. п. (NY); Scioto Co., Otway, Demaree 10646 (GH, MIN, MO, UC, US). NSYLVANIA: Allegheny Co., Aspinwall, Booth & Shafer 1901 (UC, US); mo - Millersville, Heller s. n. (D, E, GH, MIN, MO, P, US); Washington Co., New Eagle Fahr 18508 (UC). Fairfax Co., Scott's Run, Maxon 6321 (US); Loudoun Co., Short Hill, US). WEST VIRGINIA: Monongalia Co., Morgantown, Myers s. n. (MIN, NY, UC); Ohio Co., Wheeling, Mertz s. n. (US). [Vor. 1951] MEYER—VALERIANA IN NORTH AMERICA 413 Valeriana pauciflora is the only blue-flowered stoloniferous Valeriana in North America, and the corolla is the longest, 13-19 mm., of any of the North American species. In general habit and leaf shape, V. pauciflora manifests closest affinities with V. sitchensis. The modern distribution of V. pauciflora lies over a relatively broad east-west contour from southern Illinois to Maryland where the populations occur more or less locally at rather widely separated localities. However, the mode of variation does not suggest subspecific differentiation, the populations as a whole remaining quite homogeneous. Curiously, I have seen no modern material of this species from Tennessee, although Michaux first collected it in that state at the end of the 18th century. 6. VALERIANA COLUMBIANA Piper, in Bot. Gaz. 22:489. 1896. T.: Whited 140! (US). Perennials 0.8—3.0 dm. tall, from stout rhizomes 3-7 mm. thick. Stem leafy, 2-4 mm. in diameter, uniformly puberulent below, glabrescent above. Leaves basal and cauline; the basal imbricate, forming a loosely tufted rosette with the several adventitious shoots, petiolate, undivided, broadly ovate to ovate-oblong or sometimes suborbicular, 6—15 cm. long, irregularly dentate, sinuolate to entire or essentially so, glabrous or the veins sometimes pilosulous, the blades 2.5—5.5 cm. long, 1.3—3.6 cm. wide, obtuse, the petioles 2.0—6.5 cm. long, spreading-ciliate towards the base or uniformly pilosulous; cauline leaves 1—4 pairs, 4—12 cm. long, the lowermost petiolate, the uppermost sessile and much reduced, pinnate to pinnatifid, acute, irregularly repand-dentate or essential entire, the terminal lobe oblong to oblanceolate or obovate, 2.2—6.2 cm. long, 1.1-2.7 cm. wide, occasionally 3-lobed, the lateral lobes 1—2 pairs, sometimes equalling the terminal lobe in length or grading smaller. Inflorescence 3—6 cm. wide in an- thesis, corymbose, later diffuse, 7.5—15.0 cm. long, 5.5—10.0 cm. wide, internodes sometimes sparsely pilosulous; bracts 8—10 mm. long, glabrous, reduced above; flowers hermaphroditic. Corolla infundibuliform to subsalverform, 11—18 mm long, white, glabrous without, the tube gibbous, the lobes 3—6 mm. long, the throat densely pilosulous especially towards the base within. Stamens and style shorter than the corolla lobes. Achenes oblong-linear, 5-7 mm. long, 1.5-2.0 mm. wide, tawny, smooth, sometimes purple-maculate, abaxial ribs meagre. Calyx- limb 11- to 16-fid. Tyre Locatity: "Side hill above Farwell’s house west of Wenatchee [Wash- ington], June 9, 1896”. DISTRIBUTION: Open to forested rocky slopes in the mountains, 2500—6000 ft. alt. Flowering and fruiting May to July. UNITED ae WASHINGTON: Chelan En Lookout Mt. near Leavenworth, Thom p- son 0530 (GH, MO); Kittitas Co., Table , Thompson 9273 (MO, NY); Okanogan Co., Little Ji Creek, branch Twi River, ы. 278 (WTC). [Vor. 38 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 13. Valeriana columbiana: Habit, X 55; flower and achene, X 10. Valeriana columbiana is ап endemic species to the Wenatchee Mountains of north-central Washington, and is distinguished by the large, showy, corolla. This is the only North American species in series OFFICINALES with stamens shorter than the corolla lobes. It is most closely related to V. sitchensis. 1951] MEYER—VALERIANA IN NORTH AMERICA 415 7. VALERIANA OCCIDENTALIS Heller, in Bull. Torr. Bot. Club 25:269. 1898. T.: Heller 3353! (D, E, GG, G, MIN, MO, NY, P, UC, US, WTC). Valeriana micrantha E. Nels. in Erythea 7:166. 1899. T.: Nelson 793! (E, GH, MIN, MO, NY, US, WYO). Perennials (3—)4.5-9.0 dm. tall, relatively robust, from stoutish rhizomes, 3-6 mm. thick. Stem 3-6 mm. in diameter, glabrous or glabrescent, the nodes consistently white tufted-pilosulous. Leaves at the base forming a loosely tufted rosette with the several adventitious shoots, petiolate, undivided or pinnate to pinnatifid, oblong to narrowly ovate or more or less spatulate, rarely suborbicular, 12-30 cm. long, petiolate, entire or essentially so, glabrous, the blades and terminal lobe of the divided leaves 2—10 cm. long, 1.3—6.0 cm. wide, short-acuminate or obtuse, the lateral lobes 1-2 pairs, grading smaller; petiole 1-1/2 times the blade length, spreading-ciliate towards the base, sometimes spreading to the blade and lateral lobes; cauline leaves 2-4 pairs, the lowermost short-petiolate, pinnate to pinnatifid or sometimes undivided, 4.5-14.5 cm. long, the uppermost much reduced and sessile, the terminal lobe oblong-linear, ovate to obovate, 1.9-6.8 cm. long, 0.9—4.0 cm. wide, acute or obtuse, the lateral lobes 1-6 pairs, grading smaller. Inflorescence 3.5—5.0 cm. wide in anthesis, later diffuse, 8—60 cm. long, 4.5—15.0 cm. wide, nodes pilosulous, internodes glabrous or sometimes scattered-pilosulous; bracts 5-6 mm. long, glabrous; flowers gynodioecious. Corolla rotate to subrotate, 3.0-3.5 mm. long, white, glabrous without, the throat scattered-pilosulous within. Stamens and style exserted. Achenes linear- to ovate-oblong, 3-5 mm. long, 1-2 Fig. 14. Distribution of V. occidentalis. [Vor. 38 416 ANNALS OF THE MISSOURI BOTANICAL GARDEN mm. wide, sparsely to densely pilosulous or glabrous, tawny, abaxial ribs often rather prominent. Calyx-limb 11- to 16-fid. Type Послілтү: “Near western end of the Craig Mt. Plateau above Lake Waha, Nez Perce County [Idaho], alt. 3500 ft. July 2, 1896”. DISTRIBUTION: Aspen glens and yellow pine woods, wet meadows or grassy places among willows, in rich loam or on open rocky hillsides in the mountains, 4700—11000 ft. alt. Montana to Nevada and Utah; Wyoming and Colorado; also in the Black Hills of South Dakota. Flowering and fruiting May to September. UNITED STATES ARIZONA: Grand Canyon National Park, Neal Springs, north ca Collom 1056 (G). CALIFORNIA: Modoc Co., Willow Creek, Austin s. n. (B , US). coLoRADO: Delta Co., Hotchkiss, Cowen s.n. (US); Eagle Co., Pando, McDonald C289 (US); Garfield Co., Trappers Lake, Hermann 5451 (MO); Th Co., Tolland, Palmer 31300 (MO); Grand Co., Tabernash, Whitehouse 19003 (US); Gunnison Co., Mt. Carbon, Eggleston 5884 (US); Larimer Со» Rabbit Ears, Goodding 1560 (D, E, СН, Ho: Bannock Co., ісек Cronquist 2300 (МО); Benewah Co., De Smet Mis- sion, Leiberg 1010 (ARIZ, GH, K, MO, UC, WYO); Clark Co., Spencer, Rust 259 (US); Fremont Co., Henry Lake, Payson 9 Payson 1007 (GH, MO, NY, UC, WYO); Kootenai Co., Coeur d'Alene, Rust 259 (US); Lewis Co., Winchester, Meyer 1448 (MO); Nez Perce Co., Craig Mts, Henderson 2658 (GH, US); Twin Falls Co., Shoshone Falls, MONTANA: Flathead Co., Columbia Falls, Williams 105 (MIN, NY, US); Gallatin o., Bridger Mts., Jones s. n. (MO, UC, US, WYO); Fergus Co., Big Snowy Mts., Hitch- cock 16003 (WTC); Madison Co., Taylor Mts., Hitchcock & Mublick 15200 (MO); Mineral Co., Savenac Creek, Moore 308 (GH, MO, US); Park Co., Elton, Eggleston 7006 (US); Sweet Grass Co., MacLeod, Pope s.n. (NY). NEVADA: Elko Co., Jarbridge Mts., Train 605 E MO, mia OREGON: 72 Co., Lapine, Peck 0606 (GH, MO); G 4. ЖЕ" tin, Hender- son 5523 (G O); Harney Co., Myrtle Park, ws 21028 WIL ; Jefferson Co., Camp B m I0754 (WILLU) : Xlemsth Co., Johnson here jovis 2468 (US). SOUTH DAKOTA: Lawrence Co. Dumont, Over 15001 (US); Meade Co., Piedmont, Pratt s. n. (MIN); ii ipii Co., Oroville, и 752 jráe NY, US). AH: Beaver Co., Beaver, Pone er 189 (GH, NY); Cache Co., Mt. Naomi Region, Maguire, Hobson, Maguire 14020 (NY); өз, Co., 10 mi. c. Sutiiyside, Grabam 9604 (GH, МО); Grand Co., La Sal Mts., Maguire © Maguire 21708 (NY); Iron Co., 10 mi. e Cedar City, Grabam 8655 (MO); Juab Co., Mt. Nebo, Harris C2877 (MIN, MO); Piute Co., Marysvale, Jones 5366 (GG, MO, NY, UC, US); Salt Lake Co., Red Butte Canyon, Clistóni s. n. (D, Е, GH, МО); San Juan Co., Abajo Mts., Rydberg & Garrett 0724 (NY, О); San Pete Co., Ephraim Plateau, Harris C27707 (MIN, MO); Uintah rd Little Brush Creek, Harrison 9 Larsen 7704 (BRY, MO, WYO); Utah Co., n. Provo Canyon, Harrison 7244 (BRY, MO); Wasatch Co., Wolf Creek Pass, Grabam Pes (MO, US); Wayne Co., La Sal Mts., Rydberg & Garrett 8716 (GH, NY, US, WYO). WYOMING: Albany Co., Centennial, Nelson 8716 (D, GH, MO, Р, US, WYO); Big Horn Co., Worthley 15 (US); Carbon Co., T. B. Ranch, Goodding 71 (D, GG, GH, MO, NY, P, UC, US, WYO); Fremont Co., Wind River, Nelson 703 (E, GH, MIN, MO, NY, US, WYO); Lincoln Co., Afton, Payson 9 Armstrong 3306 (GH, MO, WYO); Sheridan 1951] MEYER—VALERIANA IN NORTH AMERICA 417 Co., Big Horn Mts., Williams & 1. 3204 (D, МО, WYO); Sublette Co., 15 m Merna, Payson & Payson 2786 (GH, MO, NY, UC, US, pond Teton e Hoback Canyon, Williams & Pierson 656 Td MO, NY, S, WYO); Uinta Co., La Barge, Steven- son 5 (US); Yellowstone National Park, Mammoth Hot «Ане gps s.n. (MIN). Inasmuch as Valeriana occidentalis and the next species, V. dioica sylvatica, have been perennially confused in herbaria and in the field, I had hoped for con- siderable respite in the present treatment. However, even with the large suite of study specimens, it has not been possible to eradicate all the attendant difficulties. The distributions of V. occidentalis and V. d. sylvatica overlap on their peripheries in western Montana, central Idaho, northern Wyoming and the Black Hills of South Dakota, and it becomes practically impossible to distinguish these taxa in this tension zone. Very frequently the differences are subtle ones and it is difficult to assign names to fit the variants, although these taxa may be easily distinguished outside the area where their distributions overlap. V. occidentalis V. dioica sylvatica ABIT: relatively vigorous, to 9 dm. tall, relatively slender, to 4.5 dm. tall, less leafy. and leafy. AcHENEs: sparsely to densely pilosulous or glabrous, 2.8—5.2 mm. long. glabrous, 3—5 mm. ng. DISTRIBUTION: Wyom Utah, eastern ssoutheast to north-central Canada, western Oregon, northern КЕНДЕ Со1огадо. Montana, central Idaho, sien Wyoming. 8. VALERIANA DIOICA L. ssp. sylvatica?? (Sol. ex Richards.) F. G. Mey., stat. nov. қастан it Sol. ex 1. іп Frankl. 1st Jour. App. 7:730. 1823. Т.: Richard- M, CGE, MO p Valeri riana cnn var. sylvatica "Sol. ex х Кісь а) Сгау, іп Ргос. Асай. iae 63. Valeriana NE E. Nels. іп Erythea 7:167. 1899. T.: A. 6 E. tus 5686! D, E, GH, K, MIN, MO, P, UC, US, WYO). Wiles septentrionalis Rydb. in Mem. N. Y. Bot. Gard. 1:376. 1900. T.: Drummond 7; b о Hr Nels. var. wyomingensis (E. Nels.) A. Nels. in Coult. & Nels. Man. ot. Rocky Mts. 476. 1909. Valeriana psilodes Gdgr. in Bull. Soc. Bot. Fr. 65:37. 1918. T.: Rydberg & Bessey 5001! , E, GG, GH, K, NY, WYO). Perennials 1.5—3.0(—4.5) dm. tall, relatively slender, from slimmish rhizomes about 5 mm. thick. Stem sparsely to moderately leafy, 2—4 mm. in diameter, glabrous to glabrescent, more often pilosulous in a line decurrent from the nodes. Leaves at the base sometimes forming a loosely tufted rosette with the several ad- ventitious shoots, petiolate, undivided, oblong, ovate-oblong, or spatulate, rarely suborbicular, 3—27 cm. long, entire or essentially so, the blades 1.5—8.0 cm. long, 0.7—2.9 cm. wide, short-acuminate, acute or obtuse; petioles 1—1%%(—3) times as long as the blades, spreading-ciliate towards the base; cauline leaves 3-4 pairs, the lowermost short-petiolate, pinnate to pinnatifid, rarely undivided, becoming sessile and much reduced above, oblong, ovate to ovate-oblong, 2.7-11.5 cm. long, gla- 28In the Sherardian herbarium at Oxford University a specimen collected by Mr. Tilden, “ех 1 1728" [Hudson's Bay Кере] is the oldest collection of Valeriana Б from North rica. [Vor. 38 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN EN H \ ИСА | Me у. oV Ў г жай A~ л ЖА т. ы -f Fig. 15. Distribution of V. dioica ssp. sylvatica. brous, the terminal lobe linear- to ovate-oblong, acute or short-acuminate, 1.1-5.1 cm. long, 0.2-2.4 cm. wide, the lateral lobes 1-7 pairs, grading smaller. Inflores- cence 1.5-3.0 cm. wide in anthesis, later diffuse, 5-18 cm. long, 3.5-10.0 cm. wide, the nodes pilosulous, internodes glabrous or sometimes scattered-pilosulous; bracts 5-6 mm. long, glabrous, flowers gynodioecious. Corolla rotate to subrotate, 2-3 mm. long, white, glabrous without, the throat scattered-pilosulous within. Stamens and style exserted. Achenes ovate to ovate-oblong, 3-5 mm. long, gla- brous, tawny to rubiginose, abaxial ribs evident. Calyx-limb 9- to 15-fid. Type Послілту: “On the Clearwater River". [about 56.5°N., 110°—111° W., Alberta]. July, 1820. DISTRIBUTION: Meadowlands, moist wooded hillsides in rocky duff-covered clay soil or on talus slopes in broken limestone or granite; Newfoundland north- westerly across Canada to the Rocky Mountains thence southward to Yellowstone National Park and central Idaho; 600-3400 ft. alt. in eastern Canada, and 3300— 9500 ft. alt. in the Rocky Mountains. Flowering and Mp. May to August. . (O Louise, McCabe 6343 (UC); Dry Fork, Macleod s.n. (OBT); Jumping ges Ce Macoun 20688 (NY); McLeod River, 52°80'N., 117°30'W., Ogilvie s. n. (E); cr DNE 3049 (GH); Wood Buffalo x Raup 3050 (GH, NY, UC, US); is ea Smith 3 (К); Fort Saskatchewan, Turner s.n. (OTB); Pigeon Lake, Turner 5481 (OTB). E COLUMBIA: esnel, үнімді s.n. (US); Lake Atlin, Easfwood s. n. (GH, US); about 13 mi. s. 5. 50^32'N., 120^52^W., = 8 Martin 7418 (СН, NY, UC, . Barke А тоўо (UC); ius an, McCabe 2383 (UC); "Flechead summit, McCabe 4085 Saat Kimberley, McCabe 6160 (UC); between gia and Fauquier, McCabe 6625 (UC); mi. e. Burns Lake, McCabe 7041 (UC); 30 mi. n. Ft. St. James, McCabe 7552 (UC); 159 (O 1951] MEYER— VALERIANA IN NORTH AMERICA 419 MO, NY, UC). Manirosa: Brandon, Hales s. n. (OTB); Teulon, Hunter s. n. (ОТВ); Riding Mountain National Park, Rowe 101 (OTB). NEWFOUNDLAND: Croque, Banks s. n. Жы MO photo); Table Mountain, bua ld & St. John 10864 (BM, GH, K, NY, ‚ W); Birchy Cove, Fernald & Wiegand 4064 (GH); Harry’s River, Fernald & (O Hutchinson s. n. (BM, MO photo) ; Gray Station (mile 229), ae 18006 (6 Н); Eckwan River, James Bay, Smitb 145 (US). QuEBEC: Ungava, 56^N., 75°20'-76°10'W., Риу Louis-Marie et al. 34235 (GH); Riviere Galiote, Victorin © Germain 25084 (GH); Anticosti, Crique de la Chaloupe, Victorin 4 Meiner (GH); Lac Mistassini, Rousseau 9 Rouleau 1171 (GH). SASKATCHEWAN: rkdale, Blaricom s. п. (OTB); 2. Breitung 1 (NY); McKague, Breitung 1018 MM OTB, UC); Emma Lake, Russell s. OTB). UNITED STATES: IDA uster Co., Bonanza, Macbride 9 Payson, 3404 (GH, K, MO, NY, UC, US, WYO); Lemhi Co., Salnion, Payson & Payson 1708 (GH, МО, NY, UC, WYO). MONTANA: Қатары Co., Pioneer Range, Н Hitchco ck & Mublick 12032 (MO); Carbon Co., Red Lodge, Rose 14 (GH, NY, US); Gallatin Co., Bridger Mts., Rydberg & m. 5001. (D, E, GG, on K, T WYO); Glacier National Park, Standley 15009 Flodman 802 (MIN, MO, p US); Meagher Co., Ae Bele Mts., Flodman 801 (NY, US); Missoula Co., Bonner, H tchcock & Mublick 11431 (MO, UC, WYO); Powell Co., Flathead National Forest, Lock 18636 (UC); Stillwater Co., Beartooth Mts., Kemp 16 E EGP ASHINGTON: Okanogan Co., 1 mi. e. Crawfish Lake, Fiker 2417 (WTC); Pend Oreille Co., 5 mi. n. Ione, Sz. John 6358 (MO, WTC). ig n Co., Worthley 20 (US, WYO); jos Co., Shoshone National Forest, Williams 9 Williams 3542 (ОН, MO, NY); Sheridan Co., Big Horn Mts., Тш eed y 2059 (NY); Yellowstone Sagem Park, Undine Falls, Nelson " Nelson 5686 (BM, D, E, GH, K, MIN, MO, P, UC, US$, Je Valeriana d. sylvatica throughout its Canadian distribution is practically in- distinguishable from V. dioica in western Europe. The modern distribution of V. dioica is like that of several other plants listed by Hultén as “amphi-atlantic,” or species with an eastern American and western European distribution and with- out an intermediate station in Asia. Among the amphi-atlantic species listed are Milium effusum, Liparis Loeselii and Arenaria humifusa. The following variation pattern of V. d. sylvatica bears mention as follows: The DI of V. d. sylvatica on the edge of the distribution in Montana, Idaho, Wyoming and the Black Hills come into contact with V. dp ae and in this peripheral zone Ки collections cannot be reliably determined. Тһе most aberrant popu- lations within this zone manifest the following general ee pone А. Plants more leafy than typical V. d. sylvatica, especially at the base. Leaves broader, to 2 cm. Inflorescence more often paniculate from the start. Achenes more often 5 mm. long, lance-linear. Representative specimens: IDAHO: Custer Co., Bonanza, Mac- bride & Payson 3404. MONTANA: Gallatin Co., Bridger Mts., Rydberg 6 Bessey 5001. wYoMING: Yellowstone National Park, Undine Falls, Nelson & Nelson 5686. [Vor. 38 420 ANNALS OF THE MISSOURI BOTANICAL GARDEN Series 2. EpuLEs F. С. Mey., n. ser. Perennials erect, from stout, conical, multicipital, fleshy to semi-ligneous tap- roots 0.8—3.0 cm. thick; caudex clothed with numerous imbricate, marcescent, brownish to chartaceous leaf bases of previous seasons. Stem subscapose, erect, unbranched to the inflorescence, terete or sometimes flattened and alate, glabrous or pubescent. Leaves predominantly basal, mostly undivided, rarely pinnate, lingulate-spatulate, the subpetiolar base more or less broadly clasping-patelliform, membranaceous to subcarnose, glabrous or pubescent and usually spreading-ciliate; cauline leaves simulating the basal although reduced. Inflorescence an aggregate dichasium; flowers polygamo-dioecious. Corolla rotate. Stamens and style ex- serted, anthers distinctly 4-lobed, the thecae sulcate, with the ventral loculae longer than the dorsal. Achenes linear- to ovate-oblong or oval, smooth or often transversely rugulose, glabrous or densely pilosulous to subcanescent. Calyx-limb 6- to 13-fid. Species, 3. Type Species: Valeriana edulis Nutt. ex Torr. & Gray. DISTRIBUTION: Western and north-central United States and adjoining Canada; Mexico to Costa Rica. The stout fleshy conical tap-roots and the leaves, which are mostly lingulate- spatulate and undivided, are the most salient characters that mark the species included within series EDULEs. KEY TO THE SPECIES A. Leaves undivided. В. Leaves glabrous, entire. Plants 1.0—2.5 dm. tall. Achenes oblong to oblong-linear, smooth, glabrous. Calyx-limb 6- to 8-fid. Guadalupe Mountains of Texas and New Mexico 9. V. texana о serrate-dentate and notched, the sinuses barbellate. Plants 1.0—8.5 dm. tall. Achenes ovate to ovate-oblong, often transversely rugulose, glabrous or densely pilosulous. Calyx-limb 9- to 11-fid. Southern Chiapas to Costa w |7) чы е E < б ч n "a Б ч т 2. < ч с EE б ч Р. б 3 г [^d © ч ч ы et © et 10. V. prionophylla іса assa AA. Leaves divided and undivided, somewhat repand to undulate-lamellate i 13-fid. or entire, uniformly spreading-ciliate. Calyx-limb 8- to Western and north-central United States; also in Mexico 11. V. edulis “9 VALERIANA TEXANA Steyermark, іп Ann. Mo. Bot. Gard. 19:393. 1932. T.: Moore & Steyermark 3528! (GH, MIN, MO, NY, UC). Perennials 1.0-2.5 dm. tall, from stout multicipital caudex and forked conical tap-roots 0.8-2.0 cm. thick, rugose and much contorted in age. Stem subscapose, somewhat flattened and narrowly alate, glabrous, the nodes sometimes sparsely pilosulous. Leaves predominantly basal, undivided, elliptic- to obovate-spatulate, acute, 5.5-15.0 cm. long, 1.0-2.5 cm. wide, gradually tapering to the subpetiolar base, firmly membranaceous, often with hyaline margins, glabrous; cauline leaves 1-2 pairs, 2-5 cm. long, simulating the basal, becoming bract-like above, some- times spreading-ciliate on the subpetiolar base. Inflorescence 2—6 cm. long in anthesis, later diffuse, 3.5—12.0 cm. long, 2.6—7.0 cm. wide, the internodes gla- brous or sometimes scattered-pilosulous; bracts 3—4 mm. long, reduced above, glabrous or pilosulous to spreading-ciliate. Corolla rotate, 2.5-3.0 mm. long, 1951] MEYER—VALERIANA IN NORTH AMERICA 421 Fig. 16. Valeriana texana: Habit, X 16; achene; staminate and dissected pistillate flower, Х 10. white, pilosulous towards the base of the tube without, the lobes half as long as to equaling the length of the straight tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes oblong to oblong-linear, about 2 mm. long and 1 mm. wide, smooth, brownish, more or less keeled abaxially, glabrous, adaxial ribs evident. Calyx-limb 6- to 8-fid. Type Locarrrv: “Оп boulders in creek, Upper McKittrick Canyon, Guada- lupe Mts., Culberson Co., Texas, alt. 2000 m., July 21, 1931”. DISTRIBUTION: On limestone outcrops in the Guadalupe Mountains in western Texas and adjoining New Mexico, 6000—7000 ft. alt. Flowering and fruiting April to July. UNITED STATES: NEW MEXICO: Eddy Co., Guadalupe Mts., Lee s. n. (US). TEXAs: Culberson Co., McKittrick Canyon, Guadalupe Mts., Meyer 9 Meyer 2177 (MO). [Vor. 38 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN Valeriana texana may be consistently distinguished from V. edulis, to which it is most closely related, by the oblong, glabrous and abaxially keeled achenes and the 6- to 8-fid calyx-limb. Valeriana texana is endemic to the Guadalupe Moun- tains of Texas and New Mexico, for which reason it has come to the attention of botanists only recently. The Guadalupe Mountains consist of Permian limestone of the Capitan formation, and this relatively small area is known to support a host of endemic species in other families. 10. VALERIANA PRIONOPHYLLA Standl. in Field Mus. Publ. Bot. 18:1384. 1938. T.: Stork 3040! (F). Valeriana Skutchii Standl. l. c. 22:59. 1940. T.: Skutch 1240! (F, GH). Valeriana pumilio Standl. & L. Wms. in Ceiba 1:253. 1951. T.: Williams 16729! (MO photo, T). Perennials 1.0-8.5 dm. tall, from forked conical tap-roots 0.8—2.5 cm. thick, rugose and verrucose in age. Stem subscapose, scattered-pilosulous or glabrous, the nodes consistently pilosulous. Leaves predominantly basal, imbricate, loosely or often densely tufted, undivided, oblong-linear or oblanceolate- to lingulate-spatu- late, subacute to obtuse, 2.5-35 cm. long, 0.6—3.0 cm. wide, gradually tapering to the subpetiolar base, firmly membranaceous, serrate to serrate-dentate and notched, spreading-ciliate, glabrous to uniformly pilosulous, the sinuses densely barbellate; cauline leaves 2-3 pairs, 2-25 cm. long, 0.6—3.0 cm. wide, simulating the basal, mostly sessile, or the petioles short. Inflorescence 2—6 cm. wide in anthesis, later diffuse, 15-50 cm. long, 8-12 cm. wide, the nodes tufted-pilosulous, the inter- nodes sometimes scattered-pilosulous; flower-bracts 3—4 mm. long, reduced above, glabrous. Corolla rotate, 1.5—3.0 mm. long, white to pink or pale violet, glabrous without, the lobes 1 to 117 times the length of the indistinctly gibbous or straight tube, the throat glabrous within. Stamens and style exserted. Achenes ovate to oblong-ovate, 2—3 mm. long, 1.5—2.0 mm. wide, often somewhat keeled abaxially, transversely rugulose or smooth, glabrous to densely pilosulous, sometimes purple- maculate, adaxial ribs evident. Calyx-limb 9- to 11-fid. Type LOCALITY: COSTA rica: "Cerro de la Muerte, 3000 meters, in swampy places beside streams, June 27, 1932". DisTRIBUTION: Near the summit of the Sierras from southern Chiapas to Costa Rica, 6000—12000 ft. alt., on open limestone slopes, paramos, sphagnum bogs, or in pine woods. Flowering and fruiting intermittently throughout the year. Mexico: cHiAPAs: Ghiesbreght 801 (BM, F, С, К, MO); Mt. Male, near Porvenir, Matuda 4638 (MAT, MO, NY). GUATEMALA: CHIMALTENANGO: region of Los Positos, above Las Calderas, — 80143 (F). HUEHUETENANGO: Sierra Cuchumatanes, Skutch 1240 (GH, F, MO photo); 214 mi. е. San Mateo Ixtatlan, Sierra de los Cuchumatanes, Steyermark 40882 (F); Cumbre Papal, between summit and La Libertad, Steyermark 50052 (F). QUEZALTENANGO: Volcán Santa Maria, Skutch 853 (F, GH); Volcán Zunil, Steyermark 34863 (F, NY). SACATEPEQUEZ: Volcán de Agua, Standley 65155 (F). SAN marcos: between San Sebastian and summit of Volcán Tajumulco, Steyermark 35551 (Е). soLoLA: Volcan 1951] MEYER—VALERIANA IN NORTH AMERICA 423 Fig. 17. Distribution of V. prionophylla. Santa Clara, Steyermark 46036 (F). Without definite locality: Antigua, Kellerman 4562 (US). Costa Rica: carTaco: Millsville, Holm & Iltis 540 (MO); Volcán de Turrialba, Standley 35260 (US); Irazu, ek 2807 (F); Cerro de la Muerte, Stork 3040 (F), Williams 16729 (MO photo, T). Valeriana prionophylla may be distinguished by its undivided linear- to ob- lanceolate-spatulate leaves, which are serrate to serrate-dentate with the angle at the base of the dentations barbellate. Until recently this species was barely known, although Standley and his colleagues over the past 25—30 years have added greatly to the knowledge about V. prionophylla. Variation extremes: . Plants robust, up to 10 dm. tall; leaves to 35 cm. long, 1.5—3.0 cm. wide, sharply serrate-dentate, glabrescent. Costa Rica: CARTAGO: Cerro de la Muerte, Holm 6 Iltis 468; same locality, Stork 3040 (Type of V. prionophylla). 2. Plants slender, 1-5 dm. tali; leaves 2.5-10.0 cm. long, 0.6-1.6 cm, wide, dentate, uniformly puberulent. Mexico: cHIAPAs: Mt. Male, E 4638. GUATEMALA: HUEHUETENANGO: Sierra C Cuchum atanes, Skutch 1240. Costa RicA: CARTAGO: Cerro de la Muerte, Holm & Iltis 466; same Kc E VEM 16729. These populations include V. Skutchii Standl. and V. pumilio Standl. s. 11. VALERIANA EDULIS Nutt. ex Torr. & Gray, Fl. М. Am. 2:48. 1841. T.: Nuttall s. n.! (BM, MO photo). Perennials 1—12 dm. tall, robust, from conical, often forked tap-roots 0.8—3.0 cm. thick, becoming semi-ligneous, rugose and verrucose in age; multicipital caudex covered by numerous imbricate, marcescent, brownish to blackish leaf bases of previous seasons. Stem subscapose, 2-10 mm. thick, mostly glabrous or occasion- ally appressed-puberulent, the nodes minutely puberulent. Leaves predominantly basal, imbricate, sometimes forming a rather loose rosette, linear or oblong- to obovate-spatulate, undivided or pinnate to pinnatifid, subacute to obtuse, (6—) 10— 40 cm. long, 0.3—4.2(—6.5) cm. wide, gradually tapering to the subpetiolar base, membranaceous to subcarnose, more or less repand to undulate-lamellate or entire, spreading-ciliate, uniformly pilosulous to appressed white-puberulent or glabrous, [Vor. 38 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN or the veins only puberulent; the lateral lobes of the divided leaves 1—4 pairs, distinct or more or less decurrent on the winged rhachis; cauline leaves 2—6 pairs, pinnate to pinnatifid, rarely undivided, short-petiolate or sessile below, much re- duced and bract-like above, 3—22 cm. long, elliptic- to obovate-spatulate. Inflores- cence 10—45(—75) cm. long, 2-14 cm. wide in anthesis, later diffuse, 14—65 cm. long, 2.5-17.0 cm. wide, the internodes glabrous or pilosulous; flower-bracts 3-4 mm. long, reduced above, glabrous to pilosulous or spreading-ciliate. Corolla rotate, that of the pistillate flower minute, about 0.5 mm. long, of the staminate flower 2.5-3.5 mm. long, white, glabrous or pilosulous towards the base of the tube without, the lobes half as long as to equaling the straight tube, the throat scattered-pilosulous or glabrous within. Stamens and style exserted. Achenes broadly ovate to ovate-oblong or oval, 1.8—4.5 mm. long, 1.5-3.0 mm. wide, gla- brous to densely hirtellous or subcanescent, smooth to transversely rugulose, tawny or purple-maculate, margins flat or abaxially involute, abaxial ribs prominent. Calyx-limb 8- to 13-fid. Valeriana edulis is a polytypic assemblage of three geographically disjunct sub- species: V. e. edulis in western United States and northern Mexico; V. e. ciliata in north-central United States and adjoining Canada; and V. e. procera in Mexico. Torrey & Gray?? cite the name Phyllactis obovata Nutt.?? [Valeriana obovata (Nutt.) Roem. & Schult.] as a possible synonym of V. edulis Nutt. ex T. & G. Nuttall's name has never been generally followed, although it has a priority of 23 years. Torrey & Gray” remark: “Phyllactis obovata is omitted having been de- scribed from a plant of the Upper Missouri, not yet in flower, perhaps an unde- veloped V. edulis." Та his description, Nuttall admits the plants were immature with the "flowers collected together in involucrate umbells" and with the leaves "hirsutely-pilose". This description certainly does not apply to Valeriana edulis, and I am convinced, not to Valeriana at all. I have made an exhaustive search for Nuttall's type of Phyllactis obovata but without avail. Furthermore, I have seen no material of V. edulis from the plains of north-central South Dakota where Nuttall's specimens allegedly came, e. g. "around the Arikaree village", although V. edulis occurs as near as the Black Hills. I concur fully with Torrey and Gray's doubts about pom obovata, and it is presently referred to under excluded species (p. 483 KEY TO THE SUBSPECIES > Leaves thinly ee the cauline more or less regularly pin- nate, less often pinnatifid, En lateral lobes narrow, 3—4 distinct pairs, or narrowly н uniformly spreading-ciliate. pd sem 2.5—4.5 mm. long, glabrous or hirtellous. B. Leaves more often glabrous. Wet meadows and on сенер mountain slopes; western United Be» to ‘chihuahua, Маан 11а. V. e. edulis BB. Leaves more often pilosu bogs, swamps and "— southeastern Canada, eR Eus екл DEDE C ure b. V. e. ciliata ЭБ], М. Am. 2:48. "d N. Am. Pl. eria 1818. 425 1951] MEYER—VALERIANA IN NORTH AMERICA AA. Leaves firmly membranaceous, the cauline more or less irregularly nnatifid, less often pinnate, the Dil кже Ьго ct, parted the rhachis above, bro adly nt on t te. ы Li- 3.0 mm. long, densely Durango to the state of hirtellous to occasionally short-canescent. exico llc. V. е. procera tS EN ] < А ! Y ^ LI ! N E 4 і 1 \ 2 ў Г-М П [ 1 Li —— — ГЕСЕ 2 m y П - t ә... an j I=- =- -\4 -Ò - — EJ "MEE -/- 2-1 yi ym 9 Ф Lye = x 96]: : a elm e Ё ай \ j pm s 1 25 қ - e: À “ Ж 2 E e ы 9 `» + o k ] yq ч AU rd П i M 1 4 I ' o” D С ө , 8 Мм 5 ` ^^ Ө @ А ' ач “< - ` ` 1 del .— 3 a ae = = am 7 . 6... VNQU МЕР eb. a cr РУК * i e i - 1 қ Жж Ў xL П Ф ' А ! b | o ———— ль 1 NC MT d =. dez JU ] R) ы * П 4: г” Гел NN ssp. edulis @ Те eel s К L взр. ciliata Å Ne; Ж »- Ago ELT, BSpe procer& 4 ^ KC ru i d VAS: y буы i r 4 & ж " y : 4 Fig. 18. Distribution of the subspecies of Valeriana edulis. VALERIANA EDULIS Nutt. ex Torrey & Gray ssp. EDULIS. 1834. T.: Douglas s. n.! (BM, CGE, P). 1901. T.: Nelson 7381! tla. Patrinia ceratopbylla Hook. Fl. Bor. Am. 1:290 Valeriana furfurescens A. Nels. in Bull. Torr. Bot. Club 28:232. ,NY, US). “ы trachycarpa Rydb. ibid. 31:645. 1904. T.: Underwood & Selby 352! (NY). Valeriana ceratophylla (Hook.) Piper, Contr. U. S. Nat. Herb. 11:532. 1906, non HBK. ye Aen f. glabra St. John, Fl. Southeast. Wash. & Adj. Idaho 397. 1937. T.: 2! (D, К, MIN, MO, NY, P, US, WTC). Valeriana а Standl. in Field Mus. Publ. Bot. 22:59. 1940. T.: LeSueur 1077! (F), Leaves usually numerous, the lateral lobes of Perennials 1—12 dm. tall, robust. sometimes narrowly decurrent, terminal lobe the divided leaves mostly distinct or 4.5—9.0 cm. long, 0.7-2.0 cm. wide, cauline leaves 2-3 well-developed pairs [Vor. 38 426 ANNALS OF THE MISSOURI BOTANICAL GARDEN Corolla of the perfect and staminate flower 3.0—3.5 mm. long, of the pistillate minute, 0.5 mm. long, the throat scattered-pilosulous within. Achenes 2.5—4.5 mm. long, glabrous to densely hirsutulous. Calyx-limb 9- to 13-fid. 'ТҮРЕ Locarrrv: Walla Walla, Washington. DISTRIBUTION: In moist pastures, creek bottoms, yellow pine and aspen woods, sagebrush plains, limestone cliffs, and subalpine parks, 5500—11000 ft. alt., southern British Columbia, western United States (except California) to northwestern Mexico. Flowering and fruiting May to September. CANADA: BRITISH COLUMBIA: w. of Midway, Macoun 79447 (E, NY). TA ARIZONA: ам Co., White Mountains, Gould & Robinson 5002 (ARIZ, UC); Cochise Co., Chiricahua Mts., Blumer 1353 (ARIZ, E, GH, K, MIN, MO, NY, US, DM Coconino Co., W illiam ms, Lemmon s.n. (BM, D, P, UC, US); Gila Co., Sierra ncha Mts., Gould & Hudson 3785 (ARIZ, UC); Graham Co., Pinaleno Mts., Darrow, Phillips & cde IIIQ (ARIZ); Grand Canyon National Park, Collom н СЕ 08); Navajo Co., Showlow, Hough ro (US); Yavapai Co., Prescott s. n. COLORADO: Carbon C Co., Clear Creek, Parker, McClintock & Abba: 6306 (ARIZ, WTC); Clear Creek Co., Empire, Engelmann s. n. (MO); Costilla Co., Veta Pass, Ownbey 1384 (MO, NY, WYO); Custer Co., Cusack s. n. (OXF); El Paso Co., Petrified Stump, Letterman s.n. (MO); Garfield Co., Stuart Creek, Graham 9715 (MO, US); Gilpin Co., Tolland, Palmer 31382 ( МЭН Gunnison Со., Mt. Carbon, Eggleston 5885 (US); Lake Co., Leadville, e s. * ч О); La Plata Co., voa ie Tweedy 520 (US); Larimer (MO, NY); San Miguel Co., Trout Lake, Pa " son © P Payson pu (GH, WYO); Summit Co., Breckenridge, Mackenzie 288 (MO); Teller Co., Florissant, Letterman s. n. (MO). DA quist 1705 (MO); Idaho Co., White Bird Summit, Davis 3288 (WTC); Latah Co., Mos- cow Mts., Epling 9 Hauch Ligue (МО); Nez Perce Co., Lake Waha, Heller ё Heller 3160 (E, MIN, MO, NY, P, UC, US); Teton Co., Driggs, Cronquist & Davis 2097 (MO Valley Co., 5 mi. w. McCall, NOM 9 Mublick (MO); Washington Co., Goose Creek. Jones s. n. ( 35 MONTANA: ил Co., Armstead, Payson & Payson 1750 (GH, MO, NY, bic Cascade Co., 45 mi. s.se. Great Falls, Deubenmire 48215 (WTC); Fergus Co., Big Mts., Hitcbrock & Mublick 12047 (S, UC, WYO); Gallatin Co., Belgrade, ү s.n. (МО); Lewis and Clark Co., Helena, Starz s. я. (MO); Ma die. on Со., Taylor Mts., Hitchcock & Mubhlick 15211 (MO); Meagher Co., Little Belt Mts., Hitchcock & Mublick 12241 (MO, i Park Co., Cooke City, Hitchcock & Mublick 13600 (MO, WYO). NEvADA: Elko Co., Ruby Тл; Мазоп 4665 (ОС); Humboldt Co., Summit Lake sapere Reservation, Train 3054 (NY, ОС); Washoe Co., 2 ті. w. Vya, Train 2835 (NY). MEXICO: Cat Co., 18 mi. e. Mogollon ‚ Meyer 6 Meyer 2202 (MO); Colfax Co., Ute Park, Stadien. 13685 (US); E Co. Hillsboro Peak, Metcalfe 4 (BM, D, E, GH, MIN, MO, NY, UC, US); Lincoln Co., White Mts., Wooton 320 (D, E, GG, GH E MO, NY, P, UC, US, VYO); Mora Co., Rio de la Сай, "inita з. n, (GH); Otero ‚ Mescalero Indian Reservation, Wolf 2863 (ARIZ, GH); Rio Arriba Co., Ensenada, ndi o 9 Bollman 11000 (US); San Miguel Co., Las Vegas, Studhalter 4 Сох 4137 Me ts Seun Co., west fork Gila River, Metcalfe 3I (ARIZ, BM, D, E, GH, K, MIN, Y, P UG US); Taos Co., near Taos, Mathias 570 (MO); Union Co., Sierra Grande, Pado 6149 (US). 1951] MEYER—VALERIANA IN NORTH AMERICA 427 EGON: Baker Co. Hereford, Jones 25251 (MO); Crook Co., Ochoco Ranger Rica, Peck 16000 (WILLU); Grant Co., Austin, Henderson 5525 (GH, MO); Harney Co., Narrows, Peck 13984 (WILLU) ; Klamath Co., Klamath Lake, Peck 15147 (WILLU); Malheur C Co., 24 mi. sw. Rome, Peck 21775 LEN Ыг Co., Crow ыу сыйы 8511 (MO, NY, US, WYO); Wheeler Co., Ochoco Nat. For., Weber 2911 (W soU OTA: Custer Co., Mayo, Over 1701 (US). Pennington Co., E City, Palmer 9720 “сн, МО, NY). отан: Box Elder Co., Bear River, Payson & Payson 4065 (GH, МО, US); Cache Co., Logan, Maguire 21703 (NY); Carbon Co., Willow Springs, Grabam 9541 (MO, US); Daggett Co., Uintah Mts., Williams 573 (MO, NY, WYO); Duchesne Co., Moon Lake, Graham 9327 (MO); Grand Co., La Sal Mts., Payson 9 Payson 4102 (GH, MO, WYO); Morgan Co., Echo, Jones s. n. (GG, MO, UC, US); Salt Lake Co., Salt Lake City, FAM s.n. (D, E, GH, MO); San Juan Co., Abajo Mts., е @ Redd 2136 (GH, MO); San Pete Co., Fairview, Jones 5552 (GG, MO, NY, , UC, US, WYO); Sevier Co., 40 mi. ne. Fremont, Harrison 7359 (MO); Summit Co. ы: River Valley, Hermann 5557 (GH, МО); Uintah Co., Trout Creek Ranger Station, Graham 8200 (GH, MO); Utah Co., Provo Сайоп, Palmer 38тта (СН, МО). WASHINGTON: Co., Republic, Thompson 11691 (GH, MO, NY, US); Grant Co., Grand Coulee, a ds 7656 (CA, MO); Kittitas Co., Ellensburg, Whited 60 (US); Lincoln Co., Sprague, Henderson s.n. ( ; Okanagan Co., Wauconda aee Fiker 751 (MO, US); Spokane Co., Medica! Lake, Sandberg 8 Leiberg 53 (G US); Stevens Co., Colville, Sharsmit 4041 (WTC); Whitman Co., Pullman, NN А (D, К, MIN, MO, NY, P, US, WTC). wvoMING: Albany Co., s. Sybille, Nelson 7381 (Е, MO, NY, US); Lincoln Co., nea Alpine, Payson & Armstrong 3450 (GH, MO, WYO); Park Co , Shoshone National IA Williams & Williams 3730 (D, GH, MO, NY, WYO); Sheridad Co., Big Horn Mts., Tweedy 2061 (NY); Sublette Co., Fremont Lake, o & Payson 2861 (GH, MO, NY, , WYO); Sweetwater Co., Granger, Ward s.n. (US); Teton Co., Two-gwo-tee Pi. Williams 949 (MO, NY, S, WYO); {Бадай National Park, Меке River, Nelson & Nelson 5508 (D, E, GH, MIN, MO, NY, P, US, WYO). Mrxico: CHIHUAHUA: Memelichi, Río Mayo, Gentry 2730 (ARIZ, F, GH, GT, К, MO, S, UC); Mesa Correo, LeSueur 1077 (F); Salto de Babicora, PM 1418 (ARIZ, , GH); Sierra Madre, Nelson 6008 (К, E. Pringle 1257 (E, F, GG, K, NY, US); Colonia Garcia, Townsend 9 Barber 143 (BM, D, E, F, GG, К, MO, MU, NXSUS, WYO). NUEVO LEON: Municipio de Galeana, Cerro Potosi, Schneider 964 (ARIZ, F, G, MO, NY). Valeriana e. edulis may be distinguished by the more or less regularly divided leaves with the lateral lobes mostly obtuse and distinct to the rhachis, and the achenes 2.5—4.5 mm. long, rugulose or smooth and glabrous to densely hirsutulous. This subspecies has been described under several binomials, but it seems wholly unnecessary to distinguish more than one taxon under V. edulis in western United States. The populations in the northern part of the range, north of Colorado and Utah, are more often glabrous and with broader leaves and generally of more vig- orous habit. Nuttall’s original collection from the Walla Walla region of south- eastern Washington typifies quite adequately V. e. edulis from the northern portion of the distribution. The populations in the southern part of the distribution, particularly in New Mexico, southern Arizona, and northwestern Mexico, grow under conditions of increased aridity, and have narrower and considerably more pubescent leaves. The collections described as V. trachycarpa and V. LeSueurii typify the populations from the southern part of the distribution. [Vor. 38 428 ANNALS OF THE MISSOURI BOTANICAL GARDEN llb. VALERIANA EDULIS Nutt. ex Torr. & Gray, ssp. ciliata (Torr. & Gray) F. G. Mey., stat. nov. mee иси 2” іп Edinb. М. Phil. Jour. 19:59. 1835. T.: Macnab s.n., ngifolia H ай и Torr. k y Fl. N. Am. 2:49. 1841. T.: Samples 149! (NY). Perennials 3-10 dm. tall, relatively slender to robust. Leaves few to numerous, the lateral lobes of the divided leaves mostly distinct or sometimes narrowly de- current, terminal lobe 2.3-6.8 cm. long, 0.7-1.1 cm. wide; cauline leaves 2—3 well-developed pairs. Corolla of the perfect and staminate flowers 2.8—3.2 mm. long, of the pistillate about 0.5 mm. long, the throat scattered- pilosulous within. Achenes 3-4 mm. long, glabrous. Calyx-limb 9- to 13-64. ТҮРЕ Locarrrv: Urbana, Ohio. DISTRIBUTION: In low peaty hummocks, marly bogs, prairies and tamarack swamps or wooded hillsides. Minnesota to Illinois, Michigan to Ohio. Flowering and fruiting May to October. CANADA: ONTARIO: London, Saunders s. n. (OTB); Goderich, Victorin, Germain & Dominique 45080 (GH, OTB). NITED STATES: ILLINOIS: Cook Co., Chicago, Babcock s.n. (FI, MO, P, US); Henry Co., Geneseo, Dobbs 20 (GH); Kane Co., Aurora, collector unknown (GH); Lake Co., Sand Barren Beach, Gates 1666.3 (GH); McHe nn pr Fox River Grove, Benke 6473 (GH); Stephenson Co., Freeport, Johnson s. n. (MIN, US). NDIANA: Cass Co., Hoovers, Friesner 20607 (GH, MO, S). Iowa: Clayton Со. Edgewood, pane s.n. (MIN); В Tt Hawk Co., low prairie, Burk 274 ge Fayette Co., Fink s. n. (GH, MIN, US); Floyd Co., Nora Springs, mes n. (MIN); Winnehiek Co., б. Twp., Sec. 26, Tolstead s. n. (UC), HIGAN: зе Co., Pawpaw Lake, Haines 447 (СН); Oakland Co., Rochester, Chandler s. n. (MSC, US); W sehtespsr Co., Ann Arbor, Almendinger s. n. MINNESOTA: Dakota Co., Nicols, Moore, Butters & Jenkins 15116 (MIN); File jt , Etna, Moore & Phinney 12485 (MIN); Goodhue Co., Cannon Falls, Sandber 5 05); Je ap Co., Fort Snelling, Mearns 426 (US); Houston Co., Spring Grove, RE Jabi 275 (GH, MIN); Mower Co., Austin, Moore & Phinney 12457 (MIN); Todd Co., Round Prairie Landing, Parry s.n. (MO); mm Co., Wilton, Sheldon 638 (MIN); жо с. bere Holzinger s.n. (MIN, US). 'ampaign Co., cedar swamp, Bartley & Pontius 812 (NY, US); Franklin Co., “Жаныл Sullivan? s. n. (G, GH, MO, OXF). WISCONSIN: Columbia Co., 5 mi. e. Portage, Wadmond s.n. (MIN); Dane Co., Madison, Hermann $058 (NY); Grant "a Mosca Pringle s. n. (GH); Iowa Co., ени Fassett 8120 (MIN); La Cross a Crosse, collector unknown (MO); Racine Co., Racine, Davis s. ». (MO); Т. em Bloomfield, Wadmond 4226 (MIN). Valeriana edulis is essentially a western American species, the modern distribu- tion of which may have developed its pattern in close association with the effects of Pleistocene glaciation. At the present time, however, the western and eastern populations are completely out of contact, so that interbreeding between these taxa is quite impossible. Purely on a morphological basis, the populations in the West and those in the East are barely distinguishable, but I am convinced of the intrinsic value of the ecologic factors as a basis for distinguishing these taxa. Valeriana e. ciliata is essentially a plant of bogs and swamps throughout the dis- tribution where it occurs wholly within glaciated areas, except in the driftless area in Wisconsin. 1951] MEYER—VALERIANA IN NORTH AMERICA 429 à 99 T dy 9. Valeriana edulis ssp. ciliata: Habit, X l4; pistillate and staminate flower, X 7; Fig. 1 achene (abaxi al side), X 4. [Vor. 38 430 ANNALS OF THE MISSOURI BOTANICAL GARDEN llc. VALERIANA EDULIS Nutt. ex Torr. & Gray ssp. procera (HBK.) F. С. Mey., stat. nov. Valeriana procera HBK. Nov. Gen. et Sp. 3:329. 1819. T.: Humboldt & Bonpland s. n.! Valeriana knautioides Graebn. in Engl. Bot. Jahrb. 26:427. 1899. T.: Ebrenberg 172. Perennials 3-10 dm. tall. Leaves predominantly basal, few, the lateral lobes mostly distinct above, broadly decurrent on the rhachis below, the terminal lobe (1.2-)1.4(-5.0) cm. wide; cauline leaves 3-6 well developed pairs. Corolla of the staminate flower 2.5—3.0 mm. long, of the pistillate about 0.5 mm. long, the throat glabrous within. Achenes 1.8-3.0 mm. long, densely hirtellous to short- canescent. Calyx-limb 8- to 12-fid. "ТҮРЕ Locatity: near Pázcuaro, ас 6000-7000 ft., Mexico. DISTRIBUTION: Open rocky slopes, oak-pine forest, moist sandy soil, 8000- 10400 ft. alt. Durango to Puebla. Flowering and fruiting July to September. MEXICO: DURANGO: 5 mi. n. Coyotes, 45 mi. w. C. ini sh 7146 (MO, MU); near El 8 гоз 4539 (NY, US), 4581 (S, US); С. ngo, Palmer (BM, E, F, GG, GH, К, MO, NY, UC, US); Otinapa, Palmer 380 A "GH, K, MO, NY, UC, US). HIDALGO: Regla, Galeotti 2550 (BR); El Chico, Lyonnet 1042 (F, US); Tula, Rose, Painter & Rose 8331 (US). мехісо: Comunidad Temascaltepec, Hinton 505 (К); Meson Veijo, Temascaltepec, Hinton 1319 (GH, К); Valley of Toluca, Pringle 4206 (D, F, GH, MO, NY, S, UC, US); Vallée vi Mexico, Schaffner 193 (MO, P); Llano Grande above Rio Frio 54 km. from Mexico City, Sharp 4468 (MO). MICHOACAN: Pázcuaro, Humboldt & Bonpland s. n. (MO photo, P). MORELOs: road to Cuernavaca, Kenoyer A416 (F, MO). PuzsaLa: vicinity of Puebla, Mayorazgo sur l'Atoyac, Arsène 1044 (US); entre les haciendas Sta. Barbara et Cristo sur l'Alseseca, Arséne 1363 (P, US); Esperanza, Purpus 2741 (F, MO, NY, UC, US). sonora: Cerro Saguarivo, east of San Bernardo, Pennell 19592 (US). Valeriana e. procera may be recognized by the leaves, the lateral lobes of which are broadly decurrent on the rhachis. This is in contrast to V. e. edulis with the lateral lobes more often distinct or when pinnatifid not apparently decurrent. Series ІП. CERATOPHYLLAE Hóck, in Engl. Bot. Jahrb. 3:52. 1882. Perennials from napiform or fusiform tap-roots, to 5.5 cm. thick. Stem subscapose to leafy, branched or unbranched, glabrous or puberulent to pilosulous. Leaves basal or cauline, petiolate, pinnate to bipinnatifid, elliptic- to obovate- spatulate, glabrous or hirtellous on the veins, the lateral lobes distinct, 2-10 pairs, laciniate or sometimes palmately lobed. Inflorescence a compound or aggregate dichasium; flowers hermaphroditic or gynodioecious. Corolla subrotate, 3—4 mm. long, the lobes usually equalling the length of the gibbous or nearly straight tube, the throat short-sericeous or scattered-pilosulous within. Stamens and style ex- serted, the anthers distinctly 2-lobed, thecae entire, the loculae equal in length. Achenes elliptic to ovate-oblong, 2.0—6.5 mm. long, abaxial ribs becoming carinate or at least relatively prominent. Calyx-limb 9- to 20-fid. Species, 3. Tyre Species: Valeriana ceratophylla НВК. DISTRIBUTION: Mexico. 1951] MEYER—VALERIANA IN NORTH AMERICA 431 The napiform roots, the laciniate leaves, the corolla with sericeous throat, and the achenes usually with carinate ribs combine to distinguish this series. KEY TO THE SPECIES A. Lateral leaf —— trigonal in outline, 2-9 cm. long, 0.2—5.0 wide, most frequently long-laciniate, associa pp on the rhachis with 1-3 ligular appendages. East-central to south-central Мехїсо............ 12. V. laciniosa ages mple. жр зд palmately 3- to 7-lobed cleft, 3.0-4.5 cm. long, 2.5—5.5 cm. wide, obtuse, С. -ciliate, the veins often licis S к below. Achenes cq Calyx-limb 14- to 20-fid. Northeast to aep -central Mex 13. V. albo-nervata BB. Stem branched. Leaf segments short- еа! $320 long 0.1—0.5 cm. wide, acute, ШЕ. ous. Achenes densely {күй to short-sericeous. Calyx-limb 11- to 13-fid. South-central Mexico... 14. V. ceratopbylla 12. VALERIANA LACINIOsA Mart. & Gal. in Bull. Acad. Brux. 11':121. 1844. T.: Galeotti 2548! (BR, К, MO photo, P). Valeriana calcicola Greenm. in Proc. Am. Acad. 41:252. 1905. T.: Pringle 9622! (GH). Valeriana macropoda Greenm. I. c. 1905. T.: Pringle 8998! (BM, Е, Е, FI, GG, GH, MIN, , NY, P, $, UC, US). Valise Nelsonii Greenm. I. c. 253. 1905. T.: Nelson 4574! (GH, US). Perennials 1.5—8.5 dm. tall, slender, sometimes divaricate at the base, from napiform to somewhat fusiform tap-roots, 1-4 cm. thick, rugose to somewhat verrucose; caudex covered with a succession of marcescent, brownish, chartaceous leaf bases. Stem subscapose, unbranched to the inflorescence, 1-5 mm. in diameter, glabrous or scattered-pilosulous, the nodes consistently pilosulous. Leaves pre- dominantly basal, imbricate, forming a loosely tufted rosette, petiolate, pinnate- bipinnatifid, elliptic- to oblanceolate-spatulate, 9—40 cm. long, 3-8 cm. wide, glabrous or hirtellous, especially below; lateral lobes approximately 6—10 distinct pairs, irregularly disposed, relatively long-laciniate, 1- to 8-cleft, or dilated and somewhat trigonal towards the tip, acute or apiculate, the divisions ascending or divaricate, frequently recurved, 2.2-9.0 cm. long, 0.2—5.0 cm. wide, usually asso- ciated at the rhachis with 1—3 ligular appendages, rudimentary or equalling the lateral segments, also similarly cleft, or absent, the terminal lobe simulating the lateral lobes, 1.1-10.0 cm. long, 0.1-0.3 cm. wide; petioles grading to the rhachis, sparsely hirtellous to papillate or glabrescent, spreading-ciliate towards the sub- petiolar base; cauline leaves 1-2 pairs, simulating the basal, or bract-like. Inflores- cence an aggregate dichasium, 1.5—36.0 cm. long in anthesis, later diffuse, 2—27 cm. long, 2-14 cm. wide, the nodes often tufted-pilosulous, internodes glabrous, or pilosulous; bracts 2.0—3.5 mm. long, reduced above, glabrous, somewhat erose; flowers hermaphroditic or gynodioecious. Corolla subrotate, that of the perfect flowers 3-4 mm. long, of the pistillate 1.0-1.1 mm. long, white to pink, glabrous without, the lobes nearly as long as the gibbous tube, the throat densely white- sericeous or scattered-pilosulous within. Stamens and style exserted. Achenes ovate to ovate-oblong, 4.0-6.5 mm. long, 2.5—3.5 mm. wide, glabrous or uniformly [Vor. 38 432 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 20. Distribution of V. laciniosa. pilosulous sometimes only on the adaxial side, abaxial ribs more or less carinate. Cal yx-limb 9- to 20-fid. Type Locarrry: Santa-Maria, prés de Morelia de Michoacán, 6500-7000 pieds, Mexico. DISTRIBUTION: Fields, limestone hills in cut-over oak woods on mountain sides among rocks, 4500—6000 ft. alt. Durango and Hidalgo to Puebla. Flowering and fruiting June to August. MEXICO: DURANGO: without locality, Garcia 317 (US); El Oro to Guanacevi, Nelson 4744 (GH, US); El Salto, Nelson 4574 (GH, US). nHipALGo: mountain side among rocks, Chase 7346 (F); Ixmiquilpan, Purpus s. n. (UC); Dist. Jacala, between Jacala and Zimapan, Moore & Wood 3023 (BH); Dist. Atotonilco el Grande, Cerro Colorado, Moore 5 Wood 4176 (MO); Dist. Zimapan, Moore © Wood 4274 (BH). мехісо: есімді Station, Pringle 8008 (BM, E, F, FI, GG, GH, MIN, MO, NY, P, S, UC, U E MICHOACÁN: Morelia, La Huerta, yide s. n. (D, F); и ч 2548 (BR, К, P). OAXACA Mitla, Andrieux 326 (D, К, P). PurBLA: Te , Pringle 9622 ( GH); ; Esperanza, Purpus s. n. (ОС). san LUIS POTOSI: without definite. locality, Virler 1813 (P). Valeriana laciniosa is clearly related to V. ceratopbylla, but V. laciniosa may be readily distinguished by its subscapose, unbranched stem and relatively long- laciniate leaves which are subtended by ligular appendages. With a paucity of study material, leaf variation in V. laciniosa was difficult to interpret, and the collection of Andrieux 326 from Oaxaca exhibits much broader to more or less trigonal and apiculate leaves. This suggests the polymorphic tend- encies of V. laciniosa, and with more copious material it is likely that subspecific taxa under this species would become evident. 1951] MEYER— VALERIANA IN NORTH AMERICA 433 Fig. 21. Valeriana laciniosa: Habit, X 25; achene and flower, X 7. [Vor. 38 434 ANNALS OF THE MISSOURI BOTANICAL GARDEN wan ARY Pa RN 27 seitens V / uw. f Fig. 22. Valeriana albo-nervata: Habit, X 16; dissected and entire flower, X 10. 13. VALERIANA ALBO-NERVATA Robins. in Proc. Am. Acad. 27:170. 1893. T.: Pringle 3612! (F, GH, US). Perennials 6—11 dm. tall, slender, from napiform to more or less fusiform, often forked tap-roots 2.0-5.5 cm. thick, transversely rugose to verrucose; caudex 2—6 cm. long, clothed with few marcescent, chartaceous leaf bases. Stem sub- scapose, 3—6 cm. in diameter, uniformly pilosulous or puberulent mostly on the 1951] MEYER— VALERIANA IN NORTH AMERICA 435 upper portion, or glabrescent, the nodes consistently puberulent. Leaves predomi- nantly basal, rather closely imbricate and often loosely tufted, petiolate, pinnately divided, elliptic- to oblanceolate-spatulate, 12-40 cm. long, 4.5-13.0 cm. wide, spreading-ciliate, the veins hirtellous, sometimes glabrous above, the lateral lobes 5-8 pairs, distinct, irregularly disposed, 3.0—4.5 cm. long, 2.5-5.5 cm. wide, palmately 3- to 7-lobed or cleft, obtuse and often again short-cleft; petioles grading to the rhachis, sparsely to densely hirtellous, spreading-ciliate towards the base; cauline leaves 1—2 pairs, simulating the basal, 2.5—23.0 cm. long, the uppermost much reduced. Inflorescence an aggregate or sometimes a compound dichasium, the terminal dichotomies 3.0—3.5 cm. wide in anthesis, later diffuse, 7-30 cm. long, 3.5—21.0 cm. wide, the nodes often tufted-pilosulous, internodes glabrous or pilosulous; bracts 2-4 mm. long, reduced above, glabrous, papillate-ciliate; flowers hermaphroditic. Corolla subrotate, 3.0—3.5 mm. long, white, glabrous without, the lobes half as long as to equalling the gibbous tube, the throat densely white- sericeous within. Stamens and style exserted. Achenes ovate to ovate-oblong, 3,5-6.0 mm. long, 2-3 mm. wide, glabrous, abaxial ribs subcarinate. Cal-yx-limb 14- to 20-fid. Type Locarrrv: Hillsides, бап José Pass, San Luis Potosi, Mexico. July 12, 1890. DISTRIBUTION: On open oak-studded limestone slopes, 2300-6000 ft. alt. Sierra Madre Oriental of Mexico. Flowering and fruiting March to July. albo-nervata A ceratophylla @ Fig. 23. Distribution of V. albo-nervata and V. ceratopbylla. [Vor. 38 436 ANNALS OF THE MISSOURI BOTANICAL GARDEN MEXICO: NUEVO LEON: Dist. Aramberri, Dulces Nombres, Meyer & Rogers (K, MO); Monterrey, Orcutt 571 (US). sAN Luis Poros: San José Pass, Pringle 3612 (F, GH, US). TAMAULIPAs: Jaumave Rd. 13 mi. sw. C. Victoria, McVaugh 9855 (MO); Jaumave, Runyon 36 (US); Huisachal, Stanford, Lauber 8 Taylor 2005 (US); Miquihuana, Stan- ford, Lauber 8 Taylor 2437 (US); between Marcella and Hermosa, Stanford, Lauber & Taylor 24374 (US). Valeriana albo-nervata is one of the most distinctive of Mexican Valerianas. It may be easily recognized by the leaves, the lateral lobes of which are palmately 3- to 7-lobed or cleft, and with the veins white-hirtellous. Until recently this species remained very imperfectly known, although I found it abundant over a rather wide area in the Sierras of Tamaulipas and Nuevo Leon. 14. VALERIANA CERATOPHYLLA HBK. Nov. Gen. et Sp. 3:333. 1819. T.: Hum- boldt & Bonpland s. п.! (MO photo, P), non V. ceratophylla (Hook.) Piper (Patrinia ceratopbylla Hook.). Valeriana Napus Lindl. in Bot. Reg. Misc. 76. 1840. T.: Hartweg s. n. (CGE, K). Valeriana ramosissima Mart. & Gal. in Bull. Acad. Brux. 11:122. 1844. T.: Galeotti 2552! (BR, MO photo, P). Perennials 1.7—5.7 dm. tall, slender, from napiform to somewhat fusiform, often forked tap-roots 1.5-3.0 cm. thick, transversely rugose to verrucose; caudex 2-4 cm. long, covered with a succession of imbricate, marcescent, brownish, chartaceous leaf bases and dried broken stems from previous seasons. Stem branched, 1—3 mm. in diameter, glabrous, the nodes minutely papillate to pilosulous. Leaves predominantly basal, rather closely imbricate, often loosely tufted with the several basal shoots, petiolate, bipinnate-pinnatifid, elliptic- to oblanceolate-spatulate, 8—30 cm. long, 1.0-3.8 cm. wide, glabrous, the lateral lobes 5-8 distinct pairs, irregularly disposed, 0.8—2.0 cm. long, 0.1—0.5 cm. wide, rather abruptly dilated towards the tip, short laciniate, 1- to 3-lobed or cleft, and again short-cleft, mucronate, the terminal lobe similar; petioles grading to the rhachis, sparsely hirtellous; cauline leaves 1—3 pairs, simulating the basal but smaller, 5.0-11.2 cm. long, the upper- most much reduced. Inflorescence an aggregate or compound dichasium, the terminal dichotomies 1.4-2.8 cm. wide in anthesis, later slightly diffuse, 1.0—3.5 cm. long, 1.4—3.5 cm. wide, the internodes densely puberulent in a shallow groove decurrent from the nodes; bracts 2.0-2.5 mm. long, somewhat erose, reduced above, spreading-ciliate, frequently scattered-hirtellous; flowers hermaphroditic. Corolla subrotate, 3.5—4.0 mm. long, white to pinkish, sparsely to rather densely pilosulous without, the lobes half as long as to exceeding the length of the gibbous or straight tube, the throat densely white-sericeous. Stamens and style exserted. Acbenes elliptic to ovate-oblong, 2-5 mm. long, 1.5-2.0 mm. wide, densely hirtel- lous to short-sericeous, abaxial ribs prominent, subcarinate. Calyx-limb 11- to 13-fid. Type Locarrrv: Chapoltepec, 7000 ft., Mexico. DisrRIBUTION: Dry rocky hills, 8000-8500 ft. alt. Sierra Madre Oriental, San Luis Potosi and Hidalgo to Puebla. Flowering June to September. 1951] MEYER—VALERIANA IN NORTH AMERICA 437 EXICO: HIDALGO: Cerro Ventosa, between Pachuca and Real del Monte, Galeotti 2552 (BR, MO photo, Р); Metepec Station, Pringle 13020 (ARIZ, F, СН, К, US); Ixmiquilpan, Purpus s. n. (US); Pachuca, Purpus 401 (UC, US). MEXICO: $e: a la desierta Nieja, снае s. n. (Р); Chapoltepec, Humboldt 9 Bonpland s. п. (MO photo, P). MICHOACÁN: orelia, E Arséne s. n. (E). PUEBLA: зе Luis Tultitlanapa, ds 3336 (F, G, Mo NY, UC, US); between Tepeaca and Santa Rosa, Rose 6 Hough 4736 (US). san LUIS POTOSI: without definite жна Vier p (P) Valeriana ceratophylla may be readily distinguished by its profusely branched stems, narrow unsubtended laciniate leaf lobes, and hirtellous flowers, fruits, and bracts. Series IV. CrEMarITES F. С. Mey., n. ser. Perennials or annuals, voluble or erect, from subnapiform to fusiform tap-roots. Stem leafy, branched or unbranched, terete or sometimes quadrangular, glabrous or pubescent. Leaves cauline or basal, petiolate, undivided or 3-parted, elliptic, ovate or suborbicular, cordate to reniform or palmately lobed, irregularly dentate to repand or entire, sparsely to densely spreading-hirtellous or pubescence restricted to the veins. Inflorescence an aggregate or compound dichasium; flowers hermaph- roditic or gynodioecious. Corolla infundibuliform, the tube gibbous, the throat scattered to densely pilosulous within. Stamens and style exserted; anthers 2-lobed, the thecae entire, the loculae equal in length. Achenes linear- to ovate-oblong, usually somewhat falcate to more or less ampulliform, 2—4 mm. long, 0.9—1.5 mm. wide, glabrous or sometimes pilosulous on the adaxial side. Calyx-limb 11- to 14- fid. Species, 5 Type Species: Valeriana clematitis НВК. DISTRIBUTION: Mexico, except V. urticaefolia which occurs throughout Cen- tral America and at least to Peru in the Andes. The undivided leaves and the more or less arcuate and linear- to ovate-oblong achenes are the predominant characters that mark the species of this series. KEY TO THE SPECIES A. not d or clambering, rarely erect. Leaves 2—3 times as lon wide, ovate to elliptic or suborbicular, acini to subcaudate, n or slightly dentate, the petioles mostly shorter than the blades. Northeastern Mexico to Central America; also in Colombia 15. V. clematitis AA. Pla ct. Leaves about as long as wide, acute to obtuse, serrate to obe un abs hed at phe ase. с ive mostly т апа Mesi p gos е the than the blades. m manifesting pubescen upper portion, mini aids below. Ре вм нб а кн аЬ- axial veins more or less submedian. Corolla campanulate-infundi- buliform. саа 25. 16. V. Selerorum CC. Leaves mostly ovate and truncate at the base, p көнеді shorter than the Pp Stem uniformly pubescent gla bro en en I al. САК infundibuliform to subsalverform. pend to 17. V. urticaefolia [Vor. 38 438 ANNALS OF THE MISSOURI BOTANICAL GARDEN BB. m reniform-cordate to palmately- per: undivided or with 1 Е lateral lobes. P from a thic x Әле branched e s 6.8—13.0 dm. tall, ec Leaves reniform-cordate, m. ey itregularly and often deeply pand aed Chiapas and Guatemala 18. V. cucurbitifolia DD. Plants 1.5-2.7 dm. tall, slender. Leaves palmately 5- to 7- lobed, more or less Кайа to cordate 19. V. palmatiloba 15. VALERIANA CLEMATITIS HBK. Nov. Gen. et Sp. 3:327. 1819. T.: Humboldt & Bonpland (P). Valeriana laurifolia HBK. l. c. 328. 1819. T.: ao & Bonpland! (MO photo, P). Valeriana subincisa Benth. Pl. Hartw. 39. 1839. T.: Hartweg 303! (CGE, D, K, NY). Valeriana Pavonii Poepp. & Endl. Nov. Gen. et aa 3:16, £. 215. 1845. T.: Poeppig s. n.! W). MO, ймы bispida Turcz. in Bull. Soc. Nat. Mosc. 25?:172. 1852. T.: Jameson 794! (BM, CGE, D, FI, K, MO). Valeriana Pavonii Poepp. & Endl. var. yungasensis Briq. in Ann. Conserv. & Jard. Bot. Genéve 17:337. 1913. T.: Bang 208! (D, MO). Valeriana Ghiesbrechtii Briq. 1. c. 17:345. 1914. T.: tp c s. п! (D, К, MO, P). Valeriana laxissima Standl. & L. Wms, in Ceiba 1:252. 1951. T.: Williams 8 Allen 16524! (T). Perennials voluble to clambering or erect, 3.5—12.0 dm. tall, from a short rhizome. Stem leafy, often profusely branched, rarely unbranched, terete or some- times quadrangular, becoming suberous or subligneous in age, 2-6 mm. thick, scattered-hirtellous, the nodes more densely so, the young lateral branches hoary- puberulent, soon glabrate. Leaves cauline, petiolate, undivided, elliptic to ovate or suborbicular, usually somewhat truncate at the base, acute to acuminate or sub- caudate, 4-12 cm. long, entire or slightly dentate, sparsely to densely pilosulous or subcanescent, especially below, the blades abruptly expanding, 3.0-10.5 cm. long, 1.4-4.8 cm. wide; petioles rarely equaling the blades, 1.0-5.6 cm. long, sparsely to densely pilosulous or subcanescent. Inflorescence an aggregate or compound dichasium, terminal branches 2.6—20.0 cm. long, 4.5-16.0 cm. wide in anthesis, later diffuse, 8.5—53.0 cm. long, 7.5-30 cm. wide, nodes and internodes glabrous or spreading-pilosulous; bracts 1.5—4.0 mm. long, reduced above, glabrous or hir- tellous; flowers gynodioecious. Corolla infundibuliform, that of the perfect flower 2.0—5.5 mm. long, of the pistillate 0.9—3.0 mm. long, white to pale lavender, gla- brous or hirtellous towards the base of the tube without, the lobes half as long as the gibbous tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes linear-oblong, somewhat falcate, 2.0—3.5 mm. long, 0.9-1.2 mm. wide, glabrous, smooth, tawny to rubiginose or purpurascent, abaxial ribs evident. Calyx- limb 10- to 12-fid. Type LOCALITY: COLOMBIA: paramo de Saraguru. DīIsTRIBUTION: Thickets, damp moist woods in rocky soil, often in cloud- shrouded mountain summits, 3600—13000 ft. alt. Northeastern Mexico to Guate- mala; also in Colombia. Flowering and fruiting throughout the year. Mexico: chiapas: Volcán Tacana, Matuda 2892 (F, K, MAT, MO, MU, NY). FEDERAL DISTRICT: desierto de Los Leones, Balls 4059 qutd Eslava, Pringle ogor (F, resin MO, MU, NY, S, US); Valle de Mexico, Schaffner s. n. (MO, Р). нірлісо: Ban Hartweg 303 (CGE, D, K, NY); Encarnacion, "Peas s.n. (MO); Dist. onn. 1951] MEYER—VALERIANA IN NORTH AMERICA 439 ’ FIRES нае аса \ о Ci } Um. } p Y Г) Pd ab fe ee ee ^ | ) ----%Ф----—--- deri re O Xe ^ an, EC Қайын Ға NOR К p ж Pe, A AP) P Ns f Н 53- t “ € ^1 Ті е. o ‚№ 1% P dn 4 o E 7 М ~ LM E p^ 2 : E K F i > »- Қ Шы А fs “ узи \ 4 “ Va 50: o LJ t y m yE Dr N | e^ Fig. 24. Distribution of V. clematitis. Barranca de las Verduras, Moore & Wood 4408 (MO). ЈАІЛЅСО: nw. slopes Nevado de (M Colima, McVaugb 10063 (MO); Volcano of Colima, Siecle 4390 (BM, E, F, GG, GH, 3 MIN, MO, NY, Р, S, UC, US, WYO). mexico: са. Tolucam, Andrieux 323. = P); Popocatept, Balls 4217 (BM); foret de la рее Міеја, Bourgeau 1064 (D, СН, Р, S, US); Dist. Temascaltepec, Meson bout Hinton 2732 (D, GH, MO, NY); Dist. Tul tepec, Р йв Cruz, Hinton 8981 (СН); Cafiada de San Rafael, Lyonnet & Elcoro 1927 (US); Mt. Ajusco, between Mexico City and Cuernavaca, Mexia s. n. (UC); 222: ы 1782 (F, G, MO, NY, UC, US). MicHOACÁN: Morelia, Cerro Azul, Arséne D); п.п». C. Hidalgo, ca. 19^48'N., 100740 W., McVaugb 0053 (MO). OAXACA: dup duc p of mountain, mede 2623 (NY); Zempoaltepec, Liebmann 10825 (US); atlan, Neos 2533 (US); Alvarez, Palmer 581 (F, GH, MO, NY, US). TAMAULIPAs: 7 km. sw. Miquihuana, Stanford, Retberford, Nortbcraft 702 (GH, MO, NY, UC); Pena Nevada, Stanford, Lauber, Taylor 2548 (US). vera CRUZ: Orizaba, Liebmann 10820, v 10822 (US); Maltrata, Matuda 1349 (MAT, MO, MU); Yavezia, Galeotti 2: 3 (US, GUATEMALA: CHIMALTENANGO: Santa Elena, Skutch 150 (GH, MU, S, US); Cerro de о inii 60992, 61030, 61105 (F), 61048 (Е, NY). кі. PROGRESO: between Calera and summit Volcán Siglo, Steyermark 43090 (Е). HUEHUETENANGO: Cruz de Limon, да нөн San Mateo eei и иса, Sierra de los Cuchumatanes, Steyermark 40854 (F). QUEZALTENANGO: mt = pane Standley 66331 (F); Volcan de Santa Maria, above Palojunoj, Standley уз 67501, 67620 (F). SAN pre between San Marcos and er 1172 85436 (F); between бап o at km. 21 and km. 8, 8—18 mi. nw. San Marcos, Sfeyermark 3562 28 (Е, МҮ). sorora: Volcán Fk Keller- n „3140 (US); е Santa Clara, Steyermark 46995 I" ). TOTONICAPAN: region of u Jolom, mts. above Totonicapan, on road to Desconsuelo, Standley 84476 (F). Costa Rica: CARTAGO: Cordillera de Talamanca, near El Copey, Williams © Allen os (T. ANAMA: without definite locality, Seemann 2133 (BM, К). [Vor. 38 440 ANNALS OF THE MISSOURI BOTANICAL GARDEN & Wy V/V 4 | | | N а ae же үл RES A ОКЕ SP ARX, EH. 4 ў + Fig. 25. Valeriana clematitis: Habit, X l4; staminate and pistillate flower, X 7. The voluble and clambering habit, and the undivided, elliptic to ovate or sub- orbicular leaves are the most useful characters upon which to distinguish V. clematitis. This species stands apart from the remaining four species in series 1951] MEYER—VALERIANA IN NORTH AMERICA 441 CLEMATITES, and to interpret variation is a matter of giving the proper weight to infraspecific variants. The variability is not difficult to interpret when this taxon is viewed over its total distribution. The names in synonymy include essentially several of the biotypes which occur within the normal variation pattern of this taxon. For instance, V. laurifolia of Kunth is a common form with glabrous and thicker leaves than typical V. clematitis from northern South America. In central America this form was recently described as V. laxissima by Standley and Williams. Variation in V. clematitis concerns, for the most part, differences in leaf texture, degree of pubescence, and the variable aspect of the ovate-acuminate leaf. 16. VALERIANA SELERORUM Graebn. & Loesn. in Verhandl. Bot. Ver. Branden- burg 53:86. 1912. T.: Seler 1335! (F, MO photo). Valeriana Arsenei Briq. in Ann. Conserv. & Jard. Bot. Genéve 17:340. 1914. T.: Arsène ; F, MO photo) Annuals 2.6—10.0 dm. tall, slender, from subnapiform tap-roots 3-7 mm. thick. Stem moderately leafy, 0.2-1.0 cm. in diameter, glabrous or scattered-pilose, pilosulous in a line on the upper portion, diminishing below. Leaves cauline, 2—7 pairs, undivided, 4.5-24.0 cm. long, the petioles generally exceeding the length of the blade, 2.5-17.0 cm. long, spreading-ciliate, the uppermost sessile, the blades 2-7 cm. long, 2.9—8.5 cm. wide, broadly ovate to suborbicular, cordate, obtuse, 3.6-23.0 cm. long, crenate to dentate or essentially entire, light green to somewhat glaucous below, green above, uniformly spreading-hirtellous especially above, more or less restricted to the veins below, the margins often opaque, spread- ing-ciliate. Inflorescence a compound dichasium, the terminal dichotomies 1.2-3.2 cm. wide in anthesis, later diffuse and 6-10 cm. wide, the nodes and internodes glabrous, sometimes densely pilosulous in a line; bracts 5—6 mm. long, reduced above, glabrous or occasionally spreading-ciliate; flowers hermaphroditic. Corolla campanulate-infundibuliform, 2.8-3.5 mm. long, lavender, glabrous without, the lobes less than half the length of the gibbous tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes oblong-linear, ampulliform and un- dulate-falcate, truncate at the base, 2.9—3.5 mm. long, 1.1-1.5 mm. wide, glabrous on the abaxial, pilosulous in 2 lines adjacent the midrib on the adaxial side. Calyx- limb 10- to 12-fid. Type Locarrry: Michoacan, Dist. Pátzcuaro, Mexico. DISTRIBUTION: South-central Mexico, 6200-8200 feet altitude. Flowering and fruiting August and September. MEXICO: DURANGO: Coyotes Hacienda, 63 mi. w.sw. C. Durango, Maysilles 7481a MU). Mexico: Dist. Temascaltepec, Tequesquipan, Hinton 1342 (GH, K, US); Dist. Temascaltepec, Rincén, Hinton 4646 (K, MO); Dist. Temascaltepec, Nanchititla, Hinton 4712 (GH, К). MicHOACÁN: vicinity of Morelia, Cerro Azul, Arsène 2455 (МО, S, US); vicinity of Morelia, Rincón, Arséne 5482 (MO, US); Morelia road, Kenoyer A359 (F); Ignatio, Seler 1335 (F, MO photo). [Vor. 38 442 ANNALS OF THE MISSOURI BOTANICAL GARDEN ғ [a e е i : | \ Pron ——°%Ч 4744 “ы AS 78, S. | Б ИЕЫ ды Fig. 26. Distribution of V. Selerorum. Valeriana Selerorum may be distinguished by the pubescence on the stem, which is disposed in a line on the upper portion, the broadly ovate to suborbicular and cordate leaves, the campanulate-infundibuliform corolla, the oblong-linear, ampulli- form and undulate-falcate achenes, and the compound dichasial inflorescence. This species is most closely related to V. urticaefolia, with which it is most frequently confused, mainly on the similarity of leaf shape. However, V. Selerorum may be consistently distinguished from V. urticaefolia on inflorescence and achene char- acters alone. 17. VALERIANA URTICAEFOLIA HBK. Nov. Gen. et Sp. 3:330. 1819. T.: Hum- boldt & Bonpland 2093! (MO, Р). Valeriana scorpioides DC. Prod. 4:635. 1830. T.: Berlandier 1133! (BM, D, G, MO). Valeriana erysimoides Poepp. & Endl. Nov. Gen. et Sp. 3:16. 1844. T.: Poeppig 1670! W) (MO phot i Valeriana rbomboidea Greene, Pittonia. 1:154. 1888. T.: Forrer s. .! (Е, GH, МҮ, UC, Valeriana Sallei Brig. in Ann. Conserv. & Jard. Bot. Genève 17:339. 1914. T.: Sallé 71! (D). Annuals 1.3-7.5 dm. tall, slender, from subnapiform tap-roots 3-8 mm. thick. Stem moderately leafy, unbranched or sometimes branched, 0.5—4.0 mm. thick, minutely puberulent to densely spreading-pilosulous towards the base, glabrescent above. Leaves cauline, 3—7 pairs, sessile or short-petiolate, undivided, ovate to oblong-elliptic to suborbicular, sometimes more or less flabelliform, acute to obtuse, 0.9-9.2 cm. long, serrate to crenate, dentate to repand or essentially entire, uni- formly spreading-pilosulous especially above, sometimes more or less restricted to the veins below, blades more or less abruptly expanded, 0.7—5.0 cm. long, 0.7—4.5 cm. wide; petioles 0.2—6.0 cm. long or obsolete, rarely exceeding the blades in length, uniformly spreading-pilosulous. Inflorescence an aggregate or sometimes compound dichasium, 2-28 cm. long in anthesis, later diffuse and 13—60 cm. long, 7-28 cm. wide, the terminal scorpioid sympodia 2-6 cm. long, bracts 1-2 mm. long, glabrous; flowers hermaphroditic. Corolla infundibuliform to subsalverform, 1.8-5.0 mm. long, white to pinkish, glabrous without, the tube abruptly narrow- ing and often somewhat filiform towards the base, the lobes less than half the 1951] MEYER—VALERIANA IN NORTH AMERICA 443 length of the gibbous tube, the throat sparsely pilosulous without. Stamens and style exserted. Achenes oval to suborbicular, somewhat fabriliform (bellows- shaped), with 2 peripheral and 1 median abaxial vein, subarcuate, minutely papil- late, yellowish to purplish maculate, 1.2-2.0 mm. long, 0.9-1.1 mm. wide, glabrous or sometimes densely hirtellous on the adaxial side, glabrous on the abaxial side, the midribs prominent. Calyx-limb 10- to 13-fid. 27. Valeriana urticaefolia: Flower, X 5; achene with plumose calyx segments, Х 7; ee aba view), 14. Type Locauity: "Crescit in frigidis Novo-Granatensium, prope Almaguer et convallem Guaytarensem; item juxta urbem Loxae Peruvianorum, alt. 1000-1200 hex." DISTRIBUTION: Open oak or pine woods, dry sunny hillsides, grassy slopes and fields in loose granitic conglomerate or calcareous soil, 2600-10000 ft. alt. Mexico to Panama; also in the Andes to Peru and Argentina. Mexico: CHIAPAS: Mt. Ovando, Matuda 2187 (MU); Mt. Tacana, Matuda 2491 (F, G, MO, MU, NY); Siltepec, Matuda 4106 (GH, - NY); Hacienda Monserrate, Purpus 0165 (СН, MO, NY, UC, US). CHIHUAHUA: 65 mi. e. Batopilas, Goldman 187 (GH, МҮ, US). DURANGO: Sierra Madre w. Durango, dad s. "^. (Е, GH, NY, UC, US); EI Salto, Pennell 18337 (US). FEDERAL DISTRICT: P ada de Eslava, Lyonnet 357 (GH, MO, NY, US); Cerro Magdalena, Serrania de Ajusco, Lyonnet 9 Elcoro 1906 (US); Eslava, Pringle 9365 (GH, NY, US); Tlalpam, жылы. s.n. (US). GUERRERO: Dist. Mina, Manchon-Arroyo Hondo, Hinton 0412 (СН); Dist. Mina, с Hinton 0846 (GH); Dist. Mina, Pilas, Hinton 10701 (GH) ; Dist. Mina, Campo orado, Hinton II167 (GH); Dist. Montes de Oca, Vallecites, Hinton 11335 ( GH); Dist. Galeana, Plan del Carrizo, т 14664 (F, uic MO, NY); Dist. Mina, Petlacala, Mexia 8978 (F, GH, MO, NY .. ne. Chilpancingo on road to Chilapa, Moore 9 Wood 4658 (MO). HIDALGO: del Monte, Berlandier 494 (D). JALIsCo: e. San Sebastian, Arroyo de Santa Pd Mexia 1512 (F, GH, MO, MU, NY, UC); W. Bolanos, Rose 2963 (US); [Vor. 38 444 ANNALS OF THE MISSOURI BOTANICAL GARDEN CRB { ы Үч /ә, ht 6 P РЫЗ “АТ анан жән -- I УН СХ be ] is ~ x vA" T E . м 74 (Nemea 1 0 0 fk cm | _ b | "Ф е; ж X^ H 2 7 | фун —— — e. | Алы M Fig. 28. Distribution of V. urticaefolia. Etzatlan, Rose & Painter 7556 (US). mexico: Volcán Toluca, Heller 436 (P, STR, W); Dist. Temascaltepec, Cucha, Hinton 1677 (D, GH, MO, NY , US); Dist. Temascaltepec, Pantoja, Hinton 2849 (D, GH, NY); Flor de Maria, Pringle 3234 (BM, E, F, GG, GH, K, MO, NY, P, S, UC, US, W); кулик, LENA 1781 (F, GH, MO, NY, UC, US)! Vallée de Mexico, ее 104 (P). ÁN: vicinity of Morelia, Cerro Azul, Arséne 2458 (К, NY, P); vicinity of Morelia, posi de las Nalgas, Arséne 2563 (MO, Y, 8, US); vicinity of Morelia, Ravin Santa Maria, 27 3105 (МО, МҮ, S, US); vicinity of Morelia, Cerro San DE od 5208 (MO, US); vicinity of Morelia, Loma Santa Maria, Arséne 5424 (GH, MO, NY, S, US); 5 of Morelia, Ouest del Zapote, Arsène 9390 (MO); T Hartweg 300 a FI, G, K, NY, OXF, P ist, ‚ Pte. Las Cruces, Hinton 12189 (G H, K); Dist. Coalcomán, Pto. pe. X 3 Dist. Uruapan, асе Histon 15550 (F, GH, MO, MU, NY); Morelia, Kenoyer A358 (ARIZ, F); 7 mi. sw. Uruapan, Leavenworth t$ Hoogstraal 1254 (F); D Reiche 141 (M). MoreLos: Vallée del ени, Lyonnet & Elcoro 1839 (US) 56—57, road to Cuernavaca, Moore 120 (GH); Alarcon, Orcutt t 3871 (F, MO, US); n — ear Russell S меннн 252 (US). NAYARIT: vicinity of Jalisco, Ferris богу (US); road from Tepic to Jalcojotan, Mexia 602 (D, GH, MO, MU, NY, UC, US); Tepic, Sd; 19815 (US); between Aguacata and Dolores, Rose 3358 (US). OAXACA: mts. above Cuicatlan, Pringle 5630 (F); near Oaxaca, Pringle 5630a (US). PUEBLA; Puente del Emperador, La Venta, Sharp 44540 (МО). sINALOA: Sierra Surotato, Ocurahui, Gentry 6251 (GH, GENT, MO, MU, NY); El Batel to Pico del D Mexia 461 (F, MO, ); Cerro de las Cruces, San с Montes ka Salazar 103 (US); Santa Lucia, east of Азы. Pennell 20023 (08); Cerro Quemado, ne. of Panuco, Pennell 20143 (US). VERA CRUZ: La Joya, Balls 5515 (BM); Orizaba, Botteri 578 (P), 706 (GH, US); d'Orizaba Pul. Bourgeau 2045 (D, GH, NY, S, US) ; Amatlan, Liebmann s. n. (S). G : ALTA VERA PAZ: Santa Cruz bei Coban, Seler 2438 (GH, NY, US); Tactic, Tuerckheim 1570 (BR, F, GH, MO, NY, US). cHIMALTENANGO: Alim, Ala di; near Chimaltenango, — еи (Е, NY); barranco ds la Sierra. se. Patzum, ). 1951] MEYER—VALERIANA IN NORTH AMERICA 445 yita 31383 (F). GUATEMALA: Volcán de Pacaya, above Las Calderas, Standley 58344 HUEHUETENANGO: е. San Sebastián, Standley 81456 (F); Cerro Jolomtac, Sierra i los Cuchumatanes, Steyermark 404 M ); between San Sebastián H. and large peñasco above town, Steyermark 50492 (F). APA: vicinity of LT Standley 76543 (F); between Jalapa and Paraiso, Standley 177366 (Е); between Güiziltepeque апа Potrero Carillo, о 33083 (Е). SACATEPÉQUEZ: near Antigua, ipid 58636 (F). SANTA ROSA: Estanzuela, Heyde & Lux 3060 (GH, NY, US). soLoLÁ: Río Bravo, slopes of Volcán udi Steyermark 47972 (F, UC). zacapa: Sierra de н Minas, between Rio Hondo and summit of mt. at Finca Alejandria, Steyermark 29646 (F); along Rillito del Volcán de Monos, Steyermark 42307 (F). HONDURAS: COMAYAGUA: vicinity of Siguatepeque, Т. рь ре (F). GRACIAS: without definite locality, Нја/татѕоп s. n. (S). ZÁN: Santa Ines, Valerio 452 (F); Uyuca, Valerio 575, 640, 744, 936, 1492, "2153 (F); Piedra Herrada, Cerro de Uyuca, Standley 11088 (F); vicinity Hoya Grande, Williams 9 Molina 10179 (F); La Montaüita, Williams 9 Molina 10553 (F, MO, UC); near Siguatepeque, Yuncker, Dawson, Youse 5857 (F, GH, MO, MU, NY, S, US Costa RICA: ALAJUELA: San Piedades near San Ramón, Brenes 4434 (F); Cerros de San Rafael de San Ramón, Brenes 5807 (F); alto de Acosta de San Ramón, Brenes 16667 (F); Palmira, region of Zarcero, Smitb 270 (F); omo Alfaro Ruiz, Smitb 1294 (F, GO: Las Concavas near Cartago, Cooper 65 (F); chemin 4 Cartago, Tonduz 2030 (US). sAN Jose: Cerro de Piedra Blanca, ы Escasü, Standley 32613 (US); between Aserri and Tarbaca, EE 34077 (US); plantations de cafe d'Aserrí, Tonduz 1273 (US). PANAMA: cHIRIQUÍ: Bajo Mona, Boquete Dist., Terry 1300 (F, GH, MO). Valeriana urticaefolia exhibits relatively little variation throughout the distribu- tion, being readily distinguished by the undivided leaves, oval to suborbicular and somewhat fabriliform (bellows-shaped) achenes, and the narrowly infundibuliform corolla. These characters consistently mark V. urticaefolia, and it does not seem likely that subspecific taxa exist within V. urticaefolia, at least in Mexico and Central America. I have not studied the South American material of this species to any extent, but the specimens from Ecuador and Peru match those from North America very satisfactorily. 18. VALERIANA CUCURBITIFOLIA Standl. in Field Mus. Publ. Bot. 22:58. 1940. T.: Matuda 1709! (F, GH, MAT, MO photo, MU, NY). Valeriana cacalioides Standl. I. с. 125. 1940. T.: Matuda 1812! (F, СН, K, MAT, MO U, NY). Perennials 6.8—13.0 dm. tall, robust, from stout, more or less elongated roots at least 1.3-1.5 cm. thick, lower portion unknown; caudex covered with a suc- cession of imbricate, marcescent, brownish and chartaceous leaf bases and petioles of previous seasons. Stem moderately leafy, 2—9 mm. in diameter, glabrous or spreading-pilosulous throughout, the nodes consistently pilosulous. Leaves pre- dominantly basal, 25—30 cm. long, rather closely imbricate, forming a loosely tufted rosette, petiolate, undivided or sometimes with 1 pair of lateral lobes, the blades and terminal lobe of the divided leaves abruptly expanded, reniform-cordate to cordate, 5-15 cm. wide, irregularly and often deeply dentate to repand-dentate, [Vor. 38 446 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 29. Valeriana cucurbitifolia: Habit, X 18; flower, X 8. 1951] MEYER—VALERIANA IN NORTH AMERICA 447 light green to somewhat glaucous below, green above, uniformly spreading- pilosulous or more or less restricted to the veins above and below, spreading-ciliate, the lateral lobes petiolate, simulating the terminal lobe, somewhat smaller, 3—18 cm. long, 2-12 cm. wide; petioles usually exceeding the length of the blades, 19-40 cm. long, more or less conduplicate, glabrous to uniformly pilosulous or pubescent on the concave ventral side; cauline leaves 2—4 pairs, simulating the basal, usually smaller, 10—15 cm. long, 5.8—9.0 cm. wide, petiolate towards the base, the upper- most sessile and much reduced. Inflorescence a compound or rarely an aggregate dichasium, the terminal dichotomies 1.5—3.5 cm. wide in anthesis, diffuse at maturity, 7.5-15.0 cm. wide, the nodes and internodes hirtellous; bracts 6—9 mm. long, reduced above, glabrous or occasionally spreading-ciliate; flowers hermaph- roditic. Corolla infundibuliform, 4.0—5.5 mm. long, white, glabrous without, the lobes less than half the length of the gibbous tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes linear- to ovate-oblong, somewhat falcate, 2-4 mm. long, 1.1-1.5 mm. wide, sometimes scattered brownish-maculate, glabrous on the abaxial, uniformly short-hirtellous on the adaxial side, abaxial ribs rather prominent. Calyx-limb 10- to 14-fid. Type Locarrry: Toblas, Siltepec, Chiapas, Mexico. August 8, 1937. DISTRIBUTION: On limestone outcrops and bluffs, 6000—10000 ft. alt., southern Chiapas to Guatemala. Flowering and fruiting April to August. Mexico: CHIAPAS: Toblas, peut Matuda 1700 (Е, GH, MAT, MO photo, MU, NY); Mt Ovando, Matuda 1812 (F, GH, К, MAT, MO photo, MU, NY); Ventana, near Siltepec, Matuda 4539 (MAT, NY); Yi "Male, near Porvenir, Matuda 4602 (F, G, MAT, MO, NY). GUATEM EHUETENANGO: Cumbre Papal, on bluffs between summit and La Libertad, ced 6 (Е). Valeriana cucurbitifolia has only been known for little over a dozen years, although it is so distinctive there should be no mistaking its identity. This species stands apart from its congenors in series CLEMATITEs by the large reniform to cordate leaves which are from 5—15 cm. wide. Its relationship would appear to be closest to V. palmatiloba. Er -2 a jen t ` KAP “53... V. cucurbitifolia € | у, palmatiloba o | | Fig. 30. Distribution of V. cucurbitifolia and V. palmatiloba. [Vor. 38 448 ANNALS OF THE MISSOURI BOTANICAL GARDEN 19. VaLERIANA palmatiloba F. G. Mey., spec. nov. Planta perennis 1.5—2.7 dm. alta tenuis erecta, radice ignota. Caudex usque ad 1.4 cm. diam. Caules moderate foliacei 1-2 mm. diam. sive glabri sive patenti- pilosuli, nodis pilosulis. Folia basilaria plerumque petiolata simplicia vel 3-partita, laminis simplicium et lobis terminalibus ternatorum 5- vel 7-lobatis acutis aliquid hastatis vel cordatis 2.0—7.5 cm. longis 1.8—3.4 latis parce dentatis patenti-hirtellis praecipue ad venas restrictis patenti-ciliatis, lobis lateralibus distinctis quam lobo terminali multo brevioribus 1.0-1.7 cm. longis brevi-petiolatis; petiolis 3.6 cm. longis, foliis caulinis 2-3 jugis folia basilaria simulantibus 3—5 cm. longis petiolatis superne multo reductis sessilibusque. Inflorescentia aggregatum aut compositum dichasium 1.5 cm. lata in anthesim nodis et internodiis hirtellis, bracteis 4—6 mm. longis superne reductis glabris aut aliquid patenti-ciliatis, floribus hermaphroditicis. Corolla infundibuliformis usque ad 4 mm. longa alba extra glabra, tubo gibbo lobis duplo vel plus longiori, faucibus intus parce vel dense pilosulis. Staminae et styla exserti. Achaeniae latere abaxiali glabrae latere adaxiali brevi-hirtellae. Calycis limbus in specimine nostro immaturus. Known only from the type collection. Flowering in May. МЕХІСО: CHIAPAS: open forest, mountains near Fenia, Purpus 410 (US, HOLOTYPE). Although the type specimen is immature, it is sufficiently well developed to indicate its distinctness from V. cucurbitifolia, to which it is related. V. palmati- loba is distinguished by the palmately 5- to 7-lobed and pubescent leaves. This species occurs within the range of V. cucurbitifolia, and additional material of both species would be very helpful in clearing up the relationships of these taxa. Series V. DENSIFLORAE F. G. Mey., n. ser. Perennials or annuals from napiform to fusiform tap-roots. Stem leafy or subscapose, unbranched to the inflorescence, glabrous or uniformly pubescent. Leaves cauline or basal, petiolate, pinnate to pinnatifid or bipinnatifid, rarely un- divided, elliptic- or oblanceolate- to obovate-spatulate, or ovate-cordate, (bract- like in V. vaginata), serrate to dentate or repand, entire or essentially so, glabrous or spreading-pilosulous, especially on the veins. Inflorescence an aggregate or compound dichasium; flowers hermaphroditic, polygamo-dioecious or dioecious. Corolla infundibuliform, glabrous or spreading-pilosulous without, the tube gibbous, the throat scattered-pilosulous within. Stamens and style exserted, anthers 2-lobed, the thecae entire, the loculae equal in length. Achenes predominantly suborbicular or ovate-oblong, sometimes more or less ovoid, uniformly dense-pilosulous to short- sericeous or glabrous, abaxial side convex or strongly keeled. Calyx-limb 6- to 12-fid or obsolete. Species, 5. Type Species: Valeriana densiflora Benth. DisTRIBUTION: Mexico, except V. pulchella which occurs also in Costa Rica and Panama. The species of this assemblage usually possess well-developed napiform tap- roots. The leaves are usually pinnate to pinnatifid. The achenes are often densely hirtellous to subsericeous. 1951] 449 MEYER— VALERIANA IN NORTH AMERICA р. Mage С CO UTE % Drawn from type specimen. Habit, X 14; flower, X 5. Valeriana palmatiloba: Fig. 31. [Vor. 38 450 ANNALS OF THE MISSOURI BOTANICAL GARDEN KEY TO THE SPECIES . Inflorescence an aggregate dichasium, paniculiform. Achenes merely convex abaxially. B. Leaves aer s -bipinnatifid. Inflorescence "EN spreading-ciliate. Achenes suborbicular to subovoid, 0.7-1.0 long, spreading- pilosulous. Calyx-limb 5- to 6-fid or cer ge Sonora and Chihuahua to Jalisco.............. . 20. V. apiifolia BB. Leaves pinnate to pinnatifid or much reduced. iere sert bracts glabrous. Achenes oblong-ovate to oval, 1.4—3 m. long, densely i "e i Я N — to Mexico . V. vaginata CC. Leaves well — Ee iptic- or oblanceolate- to obovate- pm pinnate to pinnatifid or sometimes undivided. San Luis 22. V. densiflora c © ғ с © a3 ы 5 нары от с al 1 o кә Poto AA. а a compound dichasium, dichotomous throughout. chenes more or less T mane abaxially. . Leaves cauline, 2 Achenes сту: elliptic, often rather strongly keeled, Sheik abaxial ribs indistinct. Calyx-limb 8-fid or obsolete. Durango an 23. V. deltoidea d Hin и » more numerous. Achenes oblong to broadly ovate, merely turgid or less strongly carinate, gla- brous or hirtellous, abaxial ribs e "m Calyx-limb 11- to 12-fid or obsolete. Oaxaca to Panama 24. V. pulchella е о p Ic 2 < о Ф "a т о а. © = 2 р 2 с. < = 2 € 2» 20. VALERIANA APIIFOLIA Gray, in Proc. Am. Acad. 22:417. 1886. T.: Palmer 564! (BM, GG, GH, K, MO, NY, P). Astrepbia mexicana Hook. & Arn. Bot. Beechey's Voy. 431. 1841. T.: be s.l (К), Phyllactis mexicana (Hook. & Arn.) Benth. & Hook. Gen. Pl. 2:154. Annuals 2.3—5.0 dm. tall, slender, from subnapiform tap-roots 4-7 mm. thick, sometimes forked. Stem leafy, 0.5-2.0 mm. in diameter, minutely puberulent throughout. Leaves cauline, 4-6 pairs, disposed mostly on the lower half of the stem, petiolate, pinnate-bipinnatifid, rarely undivided, ovate, usually more or less ovate-cordate, acuminate, 2.0—7.5 cm. long, 1.4—5.9 cm. wide, glabrous or some- times spreading-pilosulous mostly above, glabrescent below, the lateral lobes 3—5 pairs, the longest pair distinct, short-petiolate, abruptly expanded, acuminate, rather deeply lobed or cleft, the succeeding lobes more or less decurrent on the rhachis, ultimately 3- to 5-cleft; petioles 1.3—4.0 cm. long, more or less condupli- cate, glabrous or pilosulous to minutely puberulent, frequently restricted to the concavity. Inflorescence an aggregate dichasium, 15—45 cm. long, more or less heliciform in anthesis, the terminal scorpioid branches later 1.0-2.7 dm. long, bracts 1.0-1.2 mm. long, relatively long spreading-ciliate; flowers е dioecious. Corolla infundibuliform, of the perfect and staminate flower 2.5—4.0 m long, of the pistillate 0.9—1.5 mm. long, white, glabrous or нее сы without, the lobes less than half the length of the gibbous tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes suborbicular to subovoid, 0.7-1.0 mm. long, tawny or becoming brownish maculate, spreading- -pilosulous. Calyx-limb 5- to 6-fid or obsolete. Type Locarrrv: Rio Blanco, 10 mi. west by north of Guadalajara, Jalisco, Mexico. n 1951] MEYER—VALERIANA IN NORTH AMERICA 451 g. 32. М жн чл dnd Habit, X 18; mature achenes (adaxial side), those with ев above, thos h plumose segments below, X 13; flower and scorpioid branch of the in- x5 calyx florescence, [Vor. 38 452 ANNALS OF THE MISSOURI BOTANICAL GARDEN DISTRIBUTION: Moist shady oak or pine-covered mountain sides, canyon slopes, 3500-5500 ft. alt. Sonora, Chihuahua to Jalisco, Mexico. Flowering and fruiting September to January. MEXICO: CHIHUAHUA: Sierra Charuco, Río Fuerte, Gentry 1757 (F, GENT, GH, MO, UC); Guicorichi, Río Mayo, Gentry 1976 (F, GENT, MO, US); near Bato- pilas, Goldman 231 (GH, NY, US). puranco: Sierra Tres Picos, Gentry 5318 (GENT, GH, МО, MU, NY, US); Sianori, Ortega 5326 (US). jattsco: Rio Blanco, Palmer 564 (GH, MO, NY); near Guadalajara, Pringle 1768 (D, F, GH, NY, UC, ). NAYARIT: Tepic and San Blas, Lay s.n. (K). stNatoa: Quebrado de Mansana, Sierra Surotato, Gentry 6563 (GENT, GH, MU); Balboa, Ortega 5016 (US). SONORA: Sagauribo, Rio Mayo, Gentry 2105 (F, GENT, GH, MO, UC, US). SONORA- CHIHUAHUA boundary: La Mesa Colorado, Gentry 532m (MU); Сайоп de Rasnos, Gentry 597m (MU). Valeriana apiifolia may be readily distinguished by the cordate, ovate-acuminate, pinnate to bipinnatifid leaves, and small, 0.7-1.0 mm. long, subovoid achenes with the calyx-limb 5- to 6-fid or merely dentate. The heliciform inflorescence and bipinnatifid leaves further distinguish this species from all others in series DENSI- FLORAE. V. APIIFOLIA Ф V. VAGINATA | | Fig. 33. Distribution of V. apiifolia and V. vaginata. 21. VALERIANA VAGINATA HBK. Nov. Gen. et Sp. 3:331. 1819. T.: Humboldt © Bonpland s. n.! (MO photo, P). Valeriana denudata Benth. Pl. Hartw. 20. 1839. T.: Hartweg 150! (BM, CGE, С, GH, P, WO. Perennials 0.6—5.0 dm. tall, from smooth to verrucose napiform tap-roots 1—4 cm. thick; caudex covered with a succession of imbricate, marcescent, brownish, chartaceous leaf bases of previous seasons. Stem subscapose, unbranched to the 1951] MEYER— VALERIANA IN NORTH AMERICA 453 Fig. 34. Valeriana vaginata: Habit and single inflorescence, X 1⁄2; flower, X 8; achene (abaxial side), X 7. inflorescence, 1.0-3.5 mm. in diameter towards the base, glabrous. Leaves much reduced, bract-like, the basal 1- to several, rather closely imbricate, vaginate to subspathiform, more or less conduplicate, obtuse or acuminate, 1.3-6.0 cm. long, [Vor. 38 454 ANNALS OF THE MISSOURI BOTANICAL GARDEN glabrous; cauline leaves clasping-patelliform, 1.3—3.5 cm. long, 2- to 4-parted, the divisions 0.6-1.5 cm. long, glabrous without, spreading-ciliate towards the base of the sinus, sometimes hirtellous within. Inflorescence an aggregate dichasium, 1.0-2.8 cm. long, 1.3-2.0 cm. wide in anthesis, later diffuse, 3-15 cm. long, 3—6 cm. wide, the nodes puberulent, the internodes glabrous; bracts 5—7 mm. long, reduced above, glabrous; flowers hermaphroditic. Corolla infundibuliform, 3—6 m. long, white to pinkish, glabrous without, the lobes approximately half the length of the gibbous tube, the throat sparsely pilosulous within. Stamens and style exserted. Achenes ovate-oblong to suborbicular, 2.2-3.3 mm. long, 1.5-2.0 mm. wide, uniformly hirtellous to short-sericeous. Calyx-limb 11-64. Type Locarrry: "Crescit locis humidis Regni Novae Hispaniae, prope Real del Monte [Hidalgo], alt. 1430 hex. Floret Majo." The locality on the type speci- men is, "Moran". In his resumé of the Mexican journey of Humboldt, Sprague (Kew Bull. 20-27. 1924) relates, "Moran (Real de Moran) el. 7986 ft. May-June 1803—between Mexico City and Actopan in the State of Hidalgo". DISTRIBUTION: Dry grassy hills, Sierra Madre Oriental, 8000—10000 ft. alt. Flowering and fruiting April and May. MEXICO: FEDERAL DISTRICT: Serrania de Ajusco, Pringle 6858 (BM, D, E, F, GG, GH, К, MO, NY, P, UC, US, W, WYO). GUADALAJARA: p. Tapalpa, Jones s. n. (POM). GUANAJUATO: н 150 (BM, CGE, G, GH, K, NY, P, W). HIDALGO: Moran (Real de Moran), between Mexico City and Actopan, ANTA 6 Bonpland s. n. (MO photo, P); El Chico, урн 731 (СН, MO, NY, US); Trinidad, Pringle 13470 (US). Mexico: Dist. Temascaltepec, Las Cruces, Hinton 608 (GH). MICHOACÁN: Morelia, Cerro Azul, Arsène s. n. (D). Valeriana vaginata may be readily distinguished by the nearly naked stem and the much-abbreviated vaginate to subspathiform leaves. The vaginate leaves are the most reduced of any North American species of Valeriana. 22. VALERIANA DENSIFLORA Benth. Pl. Hartw. 39. 1839. T.: Hartweg 3or! (CGE, D, FI, К, NY, OXF, P, М). Perennials 1.1-9.0 dm. tall; tap-roots simple or forked, napiform or fusi- form, 0.6-3.5 cm. thick, subligneous to somewhat fleshy, smooth or becoming transversely rugose in age. Stem subscapose or leafy, 1—6 mm. in diameter, sparsely pilosulous to short-pilose especially towards the base, or glabrous, the nodes consist- ently pilosulous; subterranean stem 1-17 cm. long or obsolete. Leaves basal or cauline, the basal sometimes forming a loose rosette with the foreshortened inter- nodes, petiolate, pinnate to pinnatifid or sometimes undivided, elliptic-, oblanceolate- to obovate-spatulate, 4—26 cm. long, irregularly dentate to repand, entire or es- sentially so, glabrous, the terminal lobe oval to suborbicular, 1.1—5.0 cm. long, 0.5—4.7 cm. wide, acute or obtuse, the lateral lobes 1—6 pairs or obsolete, distinct or somewhat decurrent on the rhachis, simulating the terminal lobe, grading smaller; petioles 2.5-11.5 cm. long, glabrous or spreading-ciliate; cauline leaves 2—4 pairs, the lowermost petiolate, pinnate to pinnatifid, elliptic- to obovate-spatulate, 2.5— 12.5 cm. long, the terminal lobe linear, elliptic to suborbicular, acute or obtuse, 1951] ' MEYER—VALERIANA IN NORTH AMERICA 455 1.2-5.0 cm. long, 0.5—4.7 cm. wide, serrate to dentate or entire or essentially so, glabrous or spreading-hirtellous ог more or less restricted to the veins, the lateral lobes 2-5 pairs, distinct or somewhat decurrent on the rhachis, simulating the terminal lobe, grading smaller; petiole 1-3 cm. long, glabrous, or occasionally spreading-ciliate towards the base, or uniformly spreading-puberulent. Inflorescence an aggregate or compound dichasium, the terminal dichotomies 1—4 cm. wide in anthesis, later diffuse, 1.3—70.0 cm. long, 2.5-20.0 cm. wide, the nodes uniformly tufted-pilosulous, spreading to the internodes or glabrous; bracts 0.5—1.4 cm. long, reduced above, glabrous; flowers hermaphroditic, gynodioecious or dioecious. Corolla infundibuliform, those of the perfect flower 2.3-6.0 mm. long, of the staminate 1.8—3.0 mm. long, of the pistillate 1.2—1.5 mm. long, white to pink, glabrous without, the lobes usually less than half the length of the gibbous tube, the throat pilosulous within. Stamens and style exserted. Achenes linear- to ovate-oblong or oval, 1.4—3.0 mm. long, 0.9-1.8 mm. wide, uniformly hirtellous to subsericeous or glabrous, sometimes brownish-maculate, abaxial ribs indistinct or evident. Calyx-limb 11- to 12-fid. Valeriana densiflora is interpreted as a polytypic complex, and despite the efforts to pigeon-hole the several classes of variants, the only logical recourse, in view of the restrictions imposed by a paucity of material, has been to group the variants into two varieties, densiflora and affinis. As used here, this category is provisional, and its chief usefulness lies in pointing out segregating and more or less independent units of variation where genetic and geographic barriers may be acting potentially as the isolating mechanisms. KEY TO THE VARIETIES A. Leaves predeminantly cauline. Flowers aide or gyno- dioecious. Tap-root napiform, usually sim 0.8 cm. thick Subterranean stem 1—17 cm. long or e s South “ental Mexico.. 22a. V. d. densiflora Я eg: ра basal. Flowers dioecious. Тар-г subnapi- for o fusiform, usually 1- to few-branched, 87-16. cm. thick. Кышы stem ldo apparent or if so, 2—3 cm. long. Guerrero and Oaxaca 22b. V. d. affinis > > 22a. VALERIANA DENSIFLORA Benth. var. DENSIFLORA. Valeriana pilosiuscula Mart. & Gal. in Bull. Acad. Brux. 111:122. 1844. T.: Galeotti 2551! Valeriana retrorsa Fern. in Proc. Am. Acad. 36:502. 1901. T.: Pringle 8454! (BM, E, F, K, MO, NY, Р, S, UC, US, V). ee? napiform to somewhat fusiform, 0.8—2.0 cm. thick. Stem leafy, con- necting subterranean stem 1-17 cm. long or obsolete. Leaves predominantly cauline, 2—4 pairs, pinnate to pinnatifid, or sometimes undivided, elliptic- to obo- vate-spatulate, 2.5-15.0 cm. long, the terminal lobe linear, elliptic to suborbicular, 1.2—5.0 cm. long, 0.5—4.7 cm. wide, serrate to dentate, entire or essentially so, glabrous or spreading-hirtellous mostly on the veins, the lateral lobes 1—5 pairs, distinct or somewhat decurrent on the rhachis; petioles glabrous or uniformly puberulent. Flowers hermaphroditic or gynodioecious, 2.3-6.0 mm. long, the [Vor. 38 456 ANNALS OF THE MISSOURI BOTANICAL GARDEN pistillate less than half as long. Acbenes ovate-oblong to oval, 1.4-2.0 mm. long, 0.9-1.8 mm. wide, densely hirtelleus to subsericeous, sometimes glabrous on the adaxial side. Type ПГослілтү: Anganguio (Michoacán?), Mexico. DISTRIBUTION: On dry rocky or grassy slopes in oak or pine forests, 5000— 10000 ft. alt. San Luis Potosi to Puebla and Guerrero. Flowering and fruiting June to September. MEXICO: DURANGO: Coyotes Hacienda, 63 mi. Durango, Maysilles 7482a (MU). FEDERAL DISTRICT: Serrania de Ajusco, Pringle y» (D, F, GH, MO, NY, S, UC, US); Pedregal, Pringle 7315 (GH); s Pringle 0466 (GH, MO, NY, US); Contreras, went t 3642 (F); in montosis Tolluca, Sartorius s. n. (US); below Llano Grande Gap, near Rio Frio, Sharp 4475 (MO); 55 km. se. Mexico City, Weaver 729 (GH). GUERRERO: Vallecitos, Montes de Oca, Hinton 10309 (K); Dist. Mina, Aquazarca-Filo, Hinton 10448 (GH, К); Dist. Mina, Aquazarca, Hinton 10474 (GH, К); Dist. Mina, Chilazayote-Carrizal, Hinton 14386 (GH); Dist. Zimapán, Barranca de Taleman, Zimapán to Mina Loma del Toro and Balcones, Moore & Wood 4392 (MO). улїл<со: La Белі Het 237 (MO, US); Sierra Madre w. of — panj 3718 (05); e. Mamantlán, са, 15 mi. s. = Wilbur 1826 (MO,MU); ca. 11 m Autlán pp La Res Vini “Wilbur 1608 (MO,MU). Mexico: oe oto pe (GH, MO, NY,US); Dist. Temascaltepec, сее Hinton 804(NY); Dist. Temascaltepec, Volcán, Hinton 981 (GH) ; Dist. Temas- caltepec, Crucero, Hinton 1007 (Е, МО); Dist. Temascaltepec, Cerro Muñeca, Hinton 1368 (F,GH, MO,US) ; Dist. Тепер, Үрегісопеѕ, Hinton 4164 (СН); Dist. Temas- caltepec, Las Cruces, Hinton 4305 (D, GH, MO) ; Dist. Temascaltepec, Temascaltepec, Hin- ton 7051 (GH); Cuajimalpa, Lyonnet 493 (GH, MO, NY, US); Monte de Rio Frio, km. 49, road from Mexico City to Puebla, Mexia 2601 (F, MU, UC); Sierra de las Cruces, Pringle 4210 (BM, E, F, GG. GH, K, MO, MU, NY, Р, S, UC, US; W). MICHOACÁN: vicinity of Morelia, Cunicho, Arsène 3500 (US); Morelia, Trapeo, Arsène 6667 (US); vicinity of Morelia, Loma Santa Maria, Arsène 0876 (US); Anganguio, Hartweg 301 (CGE, D, FI, К, NY, OXF, P, W); Dist. Zitácuaro, Laureles, Hinton 11008 (GH, К); Dist. Coalcoman, Sierra Torricillas, Hinton 15004 (GH, NY, UC); on Pátzcuaro to Tacámbaro, Moore & Wood 3907 (MO); near Lake Pátzcuaro, Pringle 4121 (D, F, GH, MO, NY, S, UC, US, WYO). MoRELos: Sierra de "Ténoxtius. Pringle $454 (BM, E, F, GG, GH, K, MO, NY, P, S, UC, US, VW). oaxaca: without definite a ың Gbies- E 170 (P). PUEBLA: vicinity of Puebla, Cerro Tepoxuchil, ы 5176 (D, P, MO, US); Puente del Emperador near La Venta, Sharp 44360 (MO); near Río Otlati, 72 km. se. Mexico City, Weaver 052 (GH). saN 110715 potosi: without definite ecd Vire: ғ» (P). Variation in V. d. densiflora may be accounted for, in part, under the follow- ing list of variants: 1. Plants comparatively robust. S/em connecting with the napiform tap-root 1—17 3.6 dm. ta m. small, ovate-oblong to oval, 1.5-2.0 mm. long, 0.9—1.3 mm. wide, densely hirtellous, sometimes glabrous on the abaxial side. This is V. densiflora of Bentham. Representative specimens: Pringle 4210, 13021; Hinton 4305; Lyonnet 332; Weaver 720; Sharp 4475; Hartweg 301. 1951] MEYER— VALERIANA IN NORTH AMERICA 457 2. Leaves cauline, the terminal lobe narrowly elliptic, only slightly dentate to entire, lateral lobes to 4 pairs. Inflorescence mostly pyramidal in anthesis. Corolla 1.3—2.9 mm. long. Achenes ovate-oblong, 1.7-2.0 mm. long. This variant is V. pilosiuscula, in part, of Miren and Galeotti. Representative specimens: Ghiesbreght 179; Pringle 6466, 7315; Bourgeau 608; Sbarp 44360. Plants ч slender. Leaves cauline, pinnate, the terminal lobe predominantly r Corolla and achenes simulating those under varia This plant was described as V. retrorsa by Fernald, based on уг pop but pO sufficient material I eia to include it under V. densiflora. P \ Г; V. DELTOIDEA |А SUA. 2K V. D. DENSIFLORA\e D. AFFINIS . Distribution of V. densiflora var. densiflora, V. densiflora var. affinis and V. deltoidea 22b. VALERIANA DENSIFLORA Benth. var. affinis (Mart. & Gal.) F. С. Mey., stat. nov. asi affinis Mart. & Gal. in Bull. Acad. Brux. 11:123. 1844. T.: Galeotti 2555! (BR, D, K, P, US, V). Tap-root subnapiform to fusiform, simple or fascicular, 0.7-1.6 cm. thick. Stem subscapose, connecting with the tap-root at or near ground level, more or less densely pilosulous in a line on the upper portion when young, glabrescent in age; caudex covered with a succession of marcescent, brownish, chartaceous leaf bases. Leaves predominantly basal, usually forming a loose rosette near the base, pinnate to pinnatifid, oblanceolate- to obovate-spatulate, 7-26 cm. long, the term- inal lobe linear-oblong to oval or suborbicular, 1.1-4.5 cm. long, 0.6-3.5 cm. wide, irregularly dentate, glabrous, lateral lobes 3—6 distinct pairs; petioles spreading- ciliate towards the base. Flowers dioecious(?), the staminate 1.8—3.0 mm. long, the pistillate 1.2-1.5 mm. long. Achenes elliptic to ovate-oblong, 2-3 mm. long, 1.3-1.5 mm. wide, glabrous or sometimes uniformly spreading-hirtellous. Type Locarrrv: "Cerro de S. Felipe, prés d'Oaxaca, de 8 à 9000 pieds", Mexico. DISTRIBUTION: Along streams, in oak forests, and on steep mountain slopes, 9000—11000 ft. alt. South Mexico. Flowering’ and fruiting May to September. 458 ANNALS OF THE MISSOURI BOTANICAL GARDEN [^ 4A NOR va 4 * 55% | Al Ж 4 | А И н ), \ NN а "E EA | qu Fig. 36. Valeriana deltoidea: Habit, X Y$; achene with plumose segments, X 5, those with dentate calyx only, X 12; flower, X 10. [Vor. 38 1951] MEYER—VALERIANA IN NORTH AMERICA 459 MEXICO: GUERRERO: Taxco, Abbott 163 (GH). oaxaca: Mitla, Andrieux s. n. (P); nw. side of Mt. Zempoaltepec, Nelson 644, 692 (US) ; Cerro San Felpa Pringle E (D, F, GH, MO, NY, S, UC, US). Valeriana densiflora var. affinis may be distinguished by the often somewhat subnapiform to fusiform tap-root, the loosely tufted basal leaves, and elliptic to oblong-ovate achenes. The populations of this variety may be geographically distinct, although the details of distribution are as yet unknown. It appears to be most closely related to V. pulchella. 23. VALERIANA deltoidea F. G. Mey., spec. nov. Planta perennis 1.5—5.4 dm. alta tenuis erecta ex radice primaria crassa sub- napiformi vel fusiformi 1.5-1.8 cm. diam. Caules foliacei 0.5-2.0 mm. crassi nodis hirtellis, internodiis aut puberulis aut glabris. Folia caulina 2 jugata petiolata pinnata elliptica vel obovato-spatulata late acuta 2.2-11.5 dm. longa 2.2-7.0 cm. lata irregulariter dentata vel integra glabra, lobo terminali lineari sive oblongo- elliptico sive obovato 1.5-5.6 cm. longo 0.5-2.6 cm. lato, lobis lateralibus 2—3 jugis distinctis aut aliquid in rhachem decurrentibus lobum terminalem simulanti- bus; petiolis 2-3 cm. longis vel obsoletis glabris vel aliquid puberulentis. In- florescentia dichasium compositum dichotoma terminalis ad anthesim 1.5-2.0 cm. lata deinde parce diffusa, nodis saepe puberulentis, internodiis glabris, bracteis 5—6 mm. longis superne reductis glabris, floribus hermaphroditicis. Corolla infundi- buliformis 4—5 mm. longa alba vel rosea extra glabra, tubo gibbo lobis duplo vel plus longiori, faucibus intus sparse pilosulis. Stamina et styli exserti. Achaeniae oblongo-ellipticae latere abaxaliali plerumque valde carinato 2.0-3.5 mm. longae 1.0-1.5 mm. latae, glabrae aliquid fulvo-maculatae, costis abaxialibus indistinctis. Calycis limbus 8-fidus vel obsoletus. | Type Locarnrrv: ‘15 miles north of Guanecevi", alt. 7500-8500 ft., State of Durango, Mexico. August 17, 1898. DISTRIBUTION: Pine forests, 5400-8700 ft. alt. Durango and Sinaloa. Flower- ing and fruiting, August. RANGO: without definite locality, Garcia 319, 464 (US); Манн, north of b. Pennell 18398 (US); El Salto (Aserraderos), Pennell 18511 (US); mi. n. Guanacevi, Nelson 4760 (СН, K, US HororvPE). siNALOA: Cerro de la Sandia, ne. Panuco, Pennell 20075 (US). Valeriana deltoidea is closely related to V. densiflora, but it may be distinguished by the compound dichasial inflorescence, and the prominently keeled achenes which are distinctly triangular in cross-section. The calyx-limb is undivided or 8-fid. 24. VALERIANA PULCHELLA Mart. & Gal. іп Bull. Acad. Brux. 11!:123. 1844. T.: Galeotti 2560! (BR, D, MO photo). Valeriana Woodsonii Standl. in Ann. Mo. Bot. Gard. 27:346. 1940. T.: Woodson, Allen, Seibert 1043! (F, MO, NY). [Vor. 38 460 ANNALS OF THE MISSOURI BOTANICAL GARDEN Perennials 1.5-5.5 dm. tall, from simple or forked tap-roots which are often fascicular and more or less fusiform, somewhat verrucose in age, 0.7-1.5 cm. thick to 8.5 cm. long; caudex covered with marcescent, brownish-papyraceous leaf bases of previous seasons. Stem 1—2 mm. in diameter, glabrous or sparsely pilosulous, the nodes consistently pilosulous. Leaves disposed mostly towards the base, more or less imbricate, sometimes forming a rather loose rosette, petiolate, pinnate to pinnatifid, or lanceolate- to obovate-spatulate, 6.5—12.2 cm. long, 1.5—5.5 cm. wide, dentate to repand or essentially entire, glabrous or glabrescent, the terminal lobe abruptly expanded, elliptic to obovate or suborbicular, 1.3—3.4 cm. long, 1.4-2.9 cm. wide, acute to obtuse, the lateral lobes 1—5 pairs, distinct, narrower and shorter than the terminal lobe, grading smaller; petioles 1.5—7.0 cm. long. Inflorescence a compound dichasium, terminal dichotomies 1-2 cm. wide in anthesis, later about 4-6 cm. wide, the nodes usually tufted-pilosulous, the inter- nodes glabrous or scattered-pilosulous; bracts 6—9 mm. long, linear-acuminate to spatulate and 2.0-2.5 mm. wide; flowers hermaphroditic, rarely gynodioecious. Corolla infundibuliform, that of the perfect flower 4—6 mm. long, of the pistillate 2,5-3.0 mm. long, glabrous without, the lobes half as long as the gibbous tube, the throat scattered-pilosulous within. Stamens and style exserted. Achenes ob- long to broadly ovate, sometimes more or less abaxially keeled, 3.0—3.6 mm. long, 1.2—2.2 mm. wide, smooth, sometimes purplish maculate, glabrous or hirtellous, abaxial ribs rather prominent. Calyx-limb 11- to 12-64 or obsolete. Type Locarrrv: “forets de la Sierra de Yavezia, à 7500 pieds", Oaxaca, Mexico. Fig. 37. Distribution of V. pulchella. DISTRIBUTION: In the mountains, 4600-11500 ft. alt. Oaxaca to Panama. Flowering and fruiting May to October. Mexico: OAXACA: Sierra de Yavezia, Galeotti 2560 (BR, D, MO photo). CHIAPAS: without definite locality, Ghiesbreght 622 (СН, К, MO). GUATEMALA: HUFHUETENANGO: near Tunima, Sierra de los Cuchumatanes, Steyer- mark 48303 (F); Los Encuentros, Johnston 1009a (Е). QUEZALTENANGO: Sololá, Berg- wald zur Totonicapán und Los Encuentros, Seler 2340 (GH, NY, US). 1951] MEYER—VALERIANA IN NORTH AMERICA 461 Costa Rica: sAN José: Cerro de la Muerte, Barbour 1028 (F); near summit of Dos Burros Peaks, Cerro de la Muerte, Dayton 3071 (F); Cerro de las Vueltas, Pittier 10448 ined ANAMA: CHIRIQUÍ: Volcán de Chiriquí, Potrero Muleto, Davidson 1023 (F, GH o. vicinity of Finca Lerida, Woodson & Schery 377 (GH, MO); Loma Larga to sum- mit, Volcán de Chiriquí, Woodson, Allen, Seibert 1043 (F, MO, NY). Valeriana pulcbella may be distinguished by the compound dichasial inflores- cence and the oblong to broadly ovate achenes with the setose calyx-limb 11- to 12-fid or sometimes merely dentate. The distribution of this species is confined, essentially, to the mountains of Central America. It appears to be most closely related to V. deltoidea. Series VI. SorBIFOLIAE F. G. Mey., n. ser. Perennials or annuals from subnapiform tap-roots or rarely rhizomes. Stem leafy, voluble or erect, glabrous to pilosulous or retrorse-hirtellous. Leaves basal or cauline, ovate-cordate, undivided or pinnate to bipinnatifid, ovate-oblong to suborbicular or oblanceolate- to obovate-spatulate, acute or obtuse, sometimes acuminate or subcaudate, glabrous or scattered-pilosulous. Inflorescence an aggre- gate dichasium, the terminal scorpioid branches 0.6—4.0 cm. long; flowers hermaph- roditic or gynodioecious. Corolla campanulate-infundibuliform, 0.5—3.0 mm. long. Stamens and style included (more or less exserted in V. robertianifolia), anthers distinctly 2-lobed, the loculae equal in length. Achenes oblong-linear, elliptic to oval, pubescent on the adaxial side, glabrous on the abaxial, or sometimes uniformly pubescent or glabrous. Calyx-limb 6- to 11-fid. Species, 5 Type Species: Valeriana sorbifolia НВК. DISTRIBUTION: Southwestern and southeastern United States, Mexico, Central America, and the West Indies. The species within series SORBIFOLIAE are the most widely distributed of those south of the United States. They are most easily dis- tinguished by the stamens, which are usually included, and by the relatively small corolla, 0.7-2.8 mm. long. KEY TO THE SPECIES* Stem voluble. Achenes typically subfabriliform сете, shaped). . Ste South Florida; West Indies, Mexico to anama; c outh America.. 25. V. scandens B. Rhizomatous perennials. Leaves predom fiii undivided or 3- parted, crenate to crenate-dentate, firmly membranaceous. Hispa- niola 26. V. domingensis BB. Annuals Sion subnapiform to fusiform tap-roots. Leaves predomi- nantly pinnate to pin pipe rarely undivided, thinly membra- t naceous, Hint 1 lobes 1-6 p C anal ani longer ae ^ud corolla lobes. Leaves pinnate to age ifid or bipinnatifid, terminal lobe palmately 3-lobed or e or уры еш; 5- to 10-cleft or зубар Euh o 0.5-1.0 mm. long. Calyx-limb 6- (о 7-fid. South-central Mexico to Costa Rica 7. V. robertianifolia * The yin obl pone V. X Ekmanii (V. domingensis X V. s. scandens) follows the discussion under V. domin [Vor. 38 462 ANNALS OF THE MISSOURI BOTANICAL GARDEN CC. Stamens included, shorter than the corolla lobes, nearly sessile. Leaves гн to pinnatifid, occasionally undivided, terminal эз nd t la serrate to dentate or repand to entire. Corolla 1.0-2.8 mm. lon fid. D. Leaves undivided, or the — lobes of the divided leaves ually more or less decurren the winged rhachis. pec elliptic- to fe ермей бота, relatively Lon 1-3 (-4) long, 0.9-2.5 wide, the adaxial margins more or ps winged, plane or = undulate, бее limb 8- to 11-fid. North- western Mexico to Rica 28. V. Palmeri DD. Leaves pinnate to pie rts with the нун 2 — distinct. Achenes ovate to ovate-oblong, o. somew ampulliform, more or less turgid, 0.6 Р, 5 mm. wide, the adaxial сыт more or less involute. 1 limb 6- to 11-fid. Southeastern Arizona; M California to Honduras and Panama; — in South Americ 29. V. sorbifolia 25. VALERIANA SCANDENS L. Sp. Pl. ed. 2. 47. 1762. T.: unknown.?! Perennials voluble or clambering; subterranean portion not seen. S/em leafy, the lowermost becoming more or less suberous in age, much branched above, 1-3 mm. in diameter, the internodes 6—26 cm. long, glabrous or sparsely spreading- pilosulous throughout. Leaves cauline, petiolate, undivided or 3-parted, ovate- cordate, short or relatively long-acuminate, acute, sometimes mucronate, 4.5—18.0 cm. long, serrate to crenate, dentate to repand or essentially entire, glabrous or scattered-pilosulous above, glabrous below, the blades of the undivided leaves 3.2- 11.0 cm. long, 2.0-8.8 cm. wide, the terminal lobe of the divided leaves ovate to ovate-oblong, 2.2-11.0 cm. long, 0.9—6.3 cm. wide, lateral lobes smaller, often somewhat oblique; petioles 1.3-10.2 cm. long. Inflorescence an aggregate dichasium, 12—40 cm. long, glabrous or sometimes rather densely pilosulous; bracts 1.7-2.5 cm. long, glabrous or sometimes densely pilosulous; flowers gynodioecious. Corolla more ог less campanulate-infundibuliform, of the perfect flowers 1.0-2.3 mm. long, of the pistillate 0.5—1.0 mm. long, white, glabrous without, the lobes less than half the length of the gibbous tube, the throat scattered-pilosulous or glabrous within. Stamens and style exserted. Achenes oblong-linear to oval, sub- fabriliform (bellows-shape) with 2 relatively prominent subperipheral and 1 median abaxial vein, 1.9-3.1 mm. long, 1-2 mm. wide, smooth, tawny to brownish, sparsely pilosulous on the adaxial side, glabrous on the abaxial. Calyx-limb 11- to 15-fid. Valeriana scandens is provisionally separated into varieties: Candolleana with undivided leaves, and scandens with 3-parted leaves. The populations with these leaf forms occupy essentially the same geographical distribution but in spite of this overlap, these leaf types remain comparatively distinct. I have found only three or four intermediate specimens, out of a few hundred examples, in which both leaf forms occurred on the same individual. The populations of V. scandens appear to be differentiating more or less independently for leaf shape irrespective of other morphological characters. "!1Based on Loefling, Iter Hisp. 235. 1758. Collected at Cumaná, Venezuela, in January 1755, Loefling's type does not exist in the Linnean Herbarium in London. 1951] MEYER—VALERIANA IN NORTH AMERICA 463 KEY TO THE VARIETIES A. Leaves 3-parted 25a. V. s. scandens AA. Leaves undivided 25b. V. s. Candolleana 25a. VALERIANA SCANDENS L. var. SCANDENS. Valeriana volubilis Sessé & Mociño ex DC. Prodr. 4:634. 1830, momen p . 18 a .: Chrismar s. n. Valeriana scandens а genuina Mue Valeriana scandens B angustiloba Muell. е 118935. T.: Mueller бог (NY, W). Valeriana scandens 9 dentata Muell. l.c. 1885, p. p. Perennials voluble or clambering. Stem leafy, much branched above. 3-parted, the terminal lobe ovate to ovate-oblong, 2.2-11.0 cm. long, 0.9—6.5 cm. wide, lateral lobes smaller, often somewhat oblique; petioles 1.3—10.2 cm. long. Type Locarrmy: "Habitat in Cumana” [Venezuela]. Collected January, 1755. DISTRIBUTION: Open rocky or densely wooded slopes in mixed forest by edge Florida, West Flowering and Leaves of streams or on river flats, from near sea-level to 10000 ft. alt. Indies, Mexico, rare in Central America; also in South America. fruiting throughout the year. A A i -— NET 4 T — NS IL келе Сұм eme . И. КС Бу 4374 os, A e с du " 3 X e 4 | E g 4925. d 2 SI» Sw . #7: E : 264: r кө: V. S. SCANDENS COT ys (29: ‹ м Mt рУ S. CANDOLLEANA 15 | XS | | | od p.d A Fig. 39. Distribution of V. scandens var. scandens and V. scandens var. Candolleana. T STATES: LORIDA: Indian River, Palmer 222 (MO); Manatee, Simpson s.n. (MO); Turtle йы. Small 11563 (S); Pineola, Small, Small, DeWinkeler 10080 (D, MO, S); Titus- ville, Small, Small, DeWinkeler 10806 (S). MEXICO: HIDALGO: ca. 43 mi. ne. Jacala, Hitchcock 9 Stanford v. 9 (СН); Chapul- huacán, Kenoyer A620 (F); near Puerto Obscuro, SP 441307 (MO , NUEVO LEON: EM Kenoyer s. n. OAXACA: Yaveo, г Rio Yaveo, Mex 9213 (D, F, H Y, S, UC) ; vicinity ot Cafetal denda, Motto n & Makrinius p (F, US). GH, PUEBLA: ' regio Huauchinango, axae Mont Tepaxinola, Fróderstróm & Hultén 686 о. SAN LUIS Жы е Canoas, Pringle 4063 E prope ud Seler 704 ( TAMAULI ne. E Frosto м: = mi mpo, жағы p ед м РА Northcraft Eoo (GH, MO, NY). RUZ: Orizaba, Botteri 322 US) arten- berg, near Tantoyuca, Huasteca, Мут: 357 (ОН); Totozmalpa, E 7o45. (BR, 464 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 38. Valeriana scandens var. scandens: Habit, X 9%; pistillate flower (lower left) and staminate ym right), X 6. [Vor. 38 1951] MEYER— VALERIANA IN NORTH AMERICA 465 D); Cordoba, Greenman 133 (F, GH, NY); с Liebmann 10839, 10840; Cazadero, Liebmann 10841; Mirador, eee 10844 (US) ;Hacienda Mirador, Nelson ІІІ (US). BRITISH HONDURAS: El Cayo Dist., v Cayo, Bartlett 11466 (MU); El Cayo Dist., San Antonio, Bartlett 13033 (MU); Sittee River, Peck 956 (GH); “22 mile Stann Creek US ГИ 047 (D, Е, СН, МО, MU, NY, 5, UC EMALA: ALTA VERAPAZ: Finca Trece Aguas, Goll 90 (US); Patin, below Tamahu, es) 70855 (F); along Río Polochic below Tamahu, Standley 91781 (F); sw. of Lanquin, Steyermark 44112 (F, a Cubilquitz, Tuerckheim 7015 (GH, NY, US). не Сегго Тіхіхі, п. ‘Toco , Steyermark 31607 (F). ESCUINTLA: between Río Jute and Río Pantaleon, Saudia: 63371 (F). ora between Los Amates and Izabal, Kellerman 7357 (F, NY); Quebradas, Blake 7558 (US). телімі La Libertad, Aguilar 471 (F); San E to Dos Arroyos, БОШ 12828 (М L SALVADOR: SAN vADOR: San Martin to Laguna de Vna | Standley 22624 , US). Nicaracua: Volcán Mombacho, Baker 151 (GH, MO, МҮ, UC, WYO); vicinity of Casa Colorada near El Crucero, Sierra de Managua, Standley 8362 (F). Costa RICA: ALAJUELA: Villa Quesada, Smith P2545 (МО). BA: ORIENTE: Loma del Gato, Sierra Maestra, Clement 601 (NY); Sierra Maestra, above Daiquiri, Ekman 8082 (D, NY, S) ; Loma del Gato, Cobre range of Sierra Maestra, Leon, Clement, Roca Ti. E La Perla, Shafer 8487 (GH, NY); prope villam Monte Verde, Wright 278 (CGE, GH). SANTA CLARA: mts. of Trinidad (Siguanea group), valley of Río Hanabanilla, Rd 18505 (S). HisPANIOLA: HAITI: Dept. du Sud, Marne de la Hatte ad Ma, Ekman 621 (5); Dept. du Sud, prope Constant, Ekman 790 (S); Massif de la Hatte, Miragoane, near Quatre- Chemins, x an 9209 (S, US); Guimbi Galata, Marnes des Commissaires Holdridge 1365 (US); vicinity of M EEA Fonds Мез 10 3686 (GH, NY, US); vicinity of Furcy, теі 4408 (US), 4622 (GH, NY, US); vicinity of St. Michal de l'Atalaye, Dept. du Nord, Leonard 7824 (F,US); ame of Kalacroix, Dept. de l'Artibonite, Leonard 7915 (GH, US); vicinity of Marmelade, Dept. du Nord, Eon 8004 (NY, UC, US); vicinity of Ennery, Dept. de l'Artibonite, Leonard 8992 (MO, US); vicinity of Port de Paix, Leonard 12240 (US); vicinity of Bombardopolis, Leonard 13566 (US); vicinity of Port au Prince, Petionville, Leonard 15800 (US); Songer 3 Nash & Taylor 1783 (NY). DOMINICAN REPUBLIC: Prov. Barahona, Abbott 1800 (US); Montiada Nueva, se. Polo, Howard 8526 (GH); en el Corozo, а orillas del Rio Amina, Prov. de Santiago, Jimenez 1122 (US). Puerto Rico: Mt. Morales, near Utuado, Britton 9 Cowell 495 (F, NY, US); alto de la Bandera, near Adjuntas, Britton & Shafer 2116 (NY); Monte Cerrote, near Adjuntas, Britton © Brown 5401 (NY, US); along the Guayama, Cayey road, "e Britton, Earle 6453 ( NY); Maricao in sylvis montis Montos, Sintenis 302 (D, MO, S, US) ; adjuntas in sylva montis Capaes, Sintenis 4027 (F); adjuntas in monte Galsa, Sintenis 4232 (MO, NY, UC, US). The 3-parted leaves readily distinguish V. s. scandens, except in rare instances when the 3-parted and undivided leaf forms occur in the same individual, viz. Britton & Brown 5880 from Puerto Rico, and Wright 276-277 from Cuba. This variety is found abundantly in Florida and Mexico. In South America it is widely distributed over the northern half of the continent. In Hispaniola V. s. scandens hybridizes with the endemic V. domingensis, but this is discussed more fully under the latter species. 25b. VALERIANA SCANDENS L. var. CANDOLLEANA (Gard.) Muell. Fl. Bras. 6:344. Valeriana alpina Vell. Fl. Flum. 28. 1825; 1:4. 68. 182 Valeriana Candollean& Gardn. in Hook. Lond. Jour. E 4:112. 1845. T.: Gardner 461! , K, MO photo, W). [Vor. 38 466 ANNALS OF THE MISSOURI BOTANICAL GARDEN Valeriana Mikaniae Lindl. in Jour. Hort. Soc. 3:316. 1848. T.: Skinner s.n.! (CGE, Valerie! scandens L. var. ô dentata Muell. І. с. 1885, f. p. Perennials, voluble or clambering. Stem leafy, much branched. Leaves un- divided, ovate-cordate, short or relatively long-acuminate, acute, sometimes mucro- nate, 4.5-18.0 cm. long, serrate to crenate, dentate to repand or essentially entire, glabrous or scattered-pilosulous above, glabrous below, the blades 3.2-11.0 long, 2.0-8.8 cm. wide. ТҮРЕ LOCALITY: BRAZIL: in bushy places near the summit of the Organ Mountains. Flowering April and May. Fig. 40. Valeriana scandens var. Candolleana: Habit, X М, 1951] MEYER—VALERIANA IN NORTH AMERICA 467 DISTRIBUTION: Cuba, Hispaniola, Puerto Rico; also Mexico and Central America. Open rocky or densely wooded slopes in mixed forest by edge of streams or on river flats, from near sea-level to 10000 ft. alt. Flowering and fruiting throughout the year. EXICO: CHIAPAS: Siltepec, wo 562 (F); E Pasitar, Matuda 1071 (GH, MO, MU); Fraylesca, Siltepec, Matuda Q (F). GUERRERO: Dist. Galeana, Carrizo, Sto. Domingo, Hinton 14719 (GH, NY) JALISCO: west of San Sebastián, Hacienda del Ototel, Mexia 1845 (D, F, GH, MO, NY, UC, ur MEXICO: Dist. Temascaltepec, La Labor, Hinton 3240 (D, GH, MO, NY); Dist. Temascaltepec, Cumbre-Pena Blanca, Т” 8055 (GH). oaxaca: Chinantla, Villa Alta, Galeotti 2561 (NY, US). VERA RUZ: Orizaba, Botteri 568 (CGE, D); Acultizingo, Matuda 1177 (MU); ‘aban, Lieb- mann 10826; Amatlan, Liebmann 10827; Huatusco, Liebmann 10847 (US); Zacualpan, Barranca de Tanampa, Purpus 2870 (UC). Gu rp t VERAPAZ: Coban, Muenscher 12554 (Е); near San José, se. Tactic, Standley 69646 CHIMALTENANGO: Volcán Acatenango, Hunnewell 14868 (СН); Santa Elena, Rua 155 (US); above Las Cadai dE. 60083 (F). EL PROGRESO: hills between Finca Piamonte and slopes se. Finca Piamonte, езгі 45397 (Е). GUATEMALA: Finca La Aurora, Aguilar 250 (F); Volcán de Расауа, above Las Calderas, 1 QUEZ МҮ); above Santa Maria de }езїз, Standley 67273 (Е); damp thickets near Columbia, Stand! ey 67056 (F); Finca Pirineos, below Santa Maria de Jesús, Standley 68256 (F); se. Palestina, nm 84200 (F); between Finca Pirineos and Pat zulin, Standley 3658 (F); between Columba and Таз Mercedes, Standley 87963 (F); Finca Pirineos, Volcán Santa Maria, oes 33235 (F); Volcán Zunil, Steyermark 34742 (F). QUICHE: San Miguel Uspantan, Heyde & Lux 2924 (ОН, MO, NY, US). sACATEPÉQUEZ: Cuesta de las Canas, above Antigua, Standley 58061 (F); barranco above Duefias, Standley 63291 (F). marcos: Volcán Tajumulco, 7 mi. from San Sebastián, Steyermark 35866 (F); barrancos bordering Rio Vega near Volcán Tacaná, Steyermark 36339 (F). SANTA ROSA: Volcán ecuamburro, Heyde & Lux 4486 (F, GH, NY, US). sOLOLÁ: Volcán Saks Clara, н 46801 (Е). sucHITEPEQUEZ: near Pueblo Nuevo, Жын, 66048 (F). pA: between Cerro de Monos and upper slopes of Monte Virgen, Steyermark 42855(F). DOR: AHUACHAPAN: near Ataco, Standley & Padilla V. 2660 (F); Cerro de San vae d Standley 006 (СН, NY, US). san VICENTE: Volcán de San Vicente, Standley 21526 (GH, US " Номрок СОМАҮАС Río Selan, Valerio 2842 (F); vicinity of Siguatepeque, 2 ie (F,US).' BL 'PARAISO: Güi nope, Valerio 1742 (F). AGUA: Tiny of Casa Colorada, ear El Crucero, si summit t of Sierra de variada Con vicinity of x Tor UR 2177 (MO, NY, US); Vara Blanca de E. різці, between Poas and Barba volcanoes, Skutch 3541 (MO, NY); region of Zarcero, Smith A569 (F, ; os vicinity of La Verbena, Standley 32227 aU Prov. a José, Cerro de Piedra Blanca, ley above Escazu, Standley 32633 (US); Río Birris, s. slopes of Vo Ісіп de Irazu, Stand E а Prov. Ru bed La Estrella, Standley 39270 PUS). MA: CANAL ZONE: near river on island, White 153 (МО). сні сие ord of ы бш аал, central Valley of Río Chiriqui Viejo, Allen 1419 (GH, M МҮ); Baja Chorro, Boquete, Davidson 443 (F, GH , MO); Volcán de Chiriqui, id Dist., Davidson 886 (F, GH); Los Siguas, Pittier 3188 (F); Río Ladrillo, pos El Boquete, [Vor. 38 468 ANNALS OF THE MISSOURI BOTANICAL GARDEN E 3288 (US); Dist. Boquete, Salla- eripe Terry 1367 (F, GH, MO); vicinity of Finca Lerida, Woodson & Schery 225 аа О); vicinity of Callejon Seco, Volcán de Chiriquí, Woodson & Schery 488 (GH, MO); Bajo Mona, mouth of Quebrada Chiquero, along Rio Caldera, Woodson, Allen, Seibert 999 (MO, NY). гамЕм: Cana, Williams 720 Kai A: PINAR DEL RIO: Sierra de las Animas, Ekman 10513 (S); Pan de Quajaibon, cn 12756 (S); Sabien hill Leon 12026 (NY). SANTA CLARA: Trinidad Mts., Manan- tiales, Britton & Wilson 5261 (NY) HISPANIOLA: DOMINICAN REPUBLIC: Cordillera Central, Prov. de Azua, San Juan, Lomas de la Mediania, Sagana Nueva, Ekman 13580 (US). PUERTO RICO: between Aibonito and Coamo, Britton, Britton & Brown 5889 (NY). The var. Candolleana is readily distinguished from var. scandens by the un- divided ovate-cordate leaves in contrast to the 3- -parted leaves of the latter. Inter- breeding between these taxa does not appear to be common, and, although their distributions overlap in over 95 per cent of the specimens examined, this single character difference is completely diagnostic. Of the two varieties, the var. Candol- leana is the more common in Central America, and in South America it is wide- spread over much the same geographical area as V. s. scandens. An interesting element of var. Candolleana їп Guatemala may be recognized as follows: P ij dosis frequently densely pilosulous. Achenes 2.0-2.2 mm. long, 0 .0 e. ntative specimens: Aguilar 927; Heyde © Lux 4486; Matuda e Stand. к 60083 2 Толы 1247. 26. VALERIANA DOMINGENSIS Urban. Symb. Ant. 1:450. 1899. T.: Eggers 2229b. Perennials 4—9 dm. tall, erect, from relatively short rhizomes 1—3 mm. thick. Stem moderately leafy or subscapose, 1-3 mm. in diameter, glabrous or sometimes minutely puberulent on the nodes. Leaves predominantly basal, often rather closely imbricate, forming a loosely tufted rosette, petiolate, undivided or 3- -parted, obovate-spatulate, 5-20 cm. long, crenate to crenate-dentate, light green to some- what glaucous below, green above, firmly membranaceous, glabrous, the terminal lobe of the divided leaves abruptly expanded, ovate to suborbicular, sometimes cordate, subacute to obtuse, 1.0—3.5 mm. long, 0.9-3.2 cm. wide, the lateral lobes distinct, much reduced, 0.4—1.4 cm. long, 0.2-1.0 cm. wide; petioles 2.6-17.0 cm. long, glabrous; cauline leaves 2-5 pairs, shorter than the basal, the lower petiolate, the uppermost reduced and sessile. Inflorescence an aggregate dichasium, 10—38 cm. long, 5-10 cm. wide, the terminal scorpioid branches 0.8—3.0 cm. long, gla- brous; bracts 1.5-2.0 mm. long, glabrous; flowers hermaphroditic(?). Corolla campanulate-infundibuliform, 1.5-2.0 mm. long, white, the lobes less than half the length of the often obscurely gibbous tube, the throat glabrous or scattered- pilosulous within. S/amens and style included. Achenes oblong-linear to ovate, 1.5-3.0 mm. long, 0.9-1.2 mm. wide, smooth, sometimes tawny-maculate, glabrous or occasionally sparsely pilosulous on the adaxial side, abaxial ribs prominent, sub- carinate. Calyx-limb 10-fid. 1951] MEYER—VALERIANA IN NORTH AMERICA 469 b “4 ғ bal > ES Ss (o 4 m xbPANrOlA Fig. 41. Distribution of V. domingensis. Type Іосліштү: "Sto. Domingo in aqua rivuli Valle Nuevo, alt. 2270 m., gregaria crescens, m. Majo flor." DISTRIBUTION: Moist thickets, open pine forests, and limestone outcrops, 4290-8900 ft. alt. Hispaniola. Flowering and fruiting June to October. Hartt: Massif de la Selle, Marne Tranchant, Ekman 1182 (S, US); Massif de la ris Marne la Visite, Ekman 1403 (S); Guimbi Balata, Marnes des Commissaires, Holdri 1277 (US); vicinity of Furcy, Leonard 4490a (NY, U Dominican REPUBLIC: Prov. de La Vega, Valle Nes. toward Sabana Alta, Ekman 13791 (S, US); prope Cerro Saroza, Tuerckbeim (D, GH, NY, US). Valeriana domingensis may be distinguished by its erect habit and undivided or 3-parted, firmly membranaceous, crenate to crenate-dentate leaves. This species is endemic to Hispaniola. 26a. VALERIANA X Ekmanii (Valeriana domingensis X s. scandens) F. G. Mey., hyb. nov. Planta subvolubilis altitudine ignoto. Folia simplicia aut 3-partita repando- dentata aut crenato-dentata integra. Achaeniae oblongae vel subfabriliformes, glabrae vel pilosulae. Type Locauity: Hispaniola, Santo Domingo: Prov. de Azua, Loma Nalga de Meco, 1600—1800 m., June 9, 1926. DISTRIBUTION: Known only from the Dominican Republic. ` DoMiNICAN REPUBLIC: wit E biens pem Canela 357, 363 (P); Prov. Azua, Ekman 6290 (S HOLOTYPE, US); a de O ov. de Azua, San Jose de Ocoa, Loma del Rancho, Ekman 11643 (S, ur Sickel definite locality, iius 13580 (S) Valeriana X Ekmanii is the clearest example of natural hybridization that I have seen in the North American species. This hybrid was first recognized in a suite of specimens collected by Ekman in the Dominican Republic amongst a loan from the Stockholm Museum. These specimens did not present the usual subtleties of many natural hybrids, but, rather, characters almost exactly intermediate for the putative parents. The intermediates manifest the effect of diluting in a proportion of ap- proximately 1:1 the voluble habit of V. scandens and the leaf shape of V. domin- gensis, and since these are the only species in Hispaniola there is no doubt as to the (Vor. 38 470 ANNALS OF THE MISSOURI BOTANICAL GARDEN parentage. I have seen additional material of this hybrid at the Paris Museum collected by Dr. Canela. At least phenotypically, the populations of V. X Ekmanii appear to be those of an F, hybrid rather than due to the subtleties of introgression. 27. VALERIANA ROBERTIANIFOLIA Briq. in Ann. Conserv. & Jard. Bot. Genéve 17:342. 1914. T.: Sallé 70! (D, MO photo). Valeriana venezuelana Briq. l.c. 338. 1914. Т. Linden 463! (BM, E, FI, G, K, OXF, Р). Valeriana delicata Standl. & Steyerm. in Field Mus. Publ. Bot. 23:255. 1947. T.: Steyer- mark 50936! (F). Annuals 1.4—6.4 dm. tall, erect, from subnapiform tap-roots 1-4 mm. thick. Stem moderately leafy, 0.5—2.0 mm. in diameter, uniformly retrorse- or spreading: hirtellous. Leaves cauline, 5-8 pairs, petiolate, pinnate to pinnatifid or bipinnatifid the lowermost rarely undivided, cordate, ovate to ovate-oblong or obovate-spatu: late, obtuse, 1.4—6.5 cm. long, 0.8—4.0 cm. wide, uniformly hirtellous above, more or less restricted to the veins below, the terminal lobe 1.5—3.0 cm. long, 1.1-1.4 cm. wide, palmately 3-lobed, these 1- to several-cleft or the lobes more or less irregularly 5- to 10-cleft or dentate, the lobes 2(—3) pairs, distinct, oblong to obovate, 3- to 5-cleft or more or less irregularly dentate; petioles 0.5—2.7 cm. long, uniformly retrorse- to spreading-hirtellous. Inflorescence an aggregate dichasium, 9-38 cm. long, 4-12 cm. wide, the terminal scorpioid branches 0.6—1.1 cm. long, glabrous; bracts 1.0-1.5 mm. long, glabrous; flowers gynodioecious. Corolla campanulate-infundibuliform, that of the perfect flower 0.5—1.0 mm. long, of the pistillate about 0.5 mm. long, glabrous without, pinkish, the lobes less than half the length of the gibbous or straight tube, the throat glabrous within. Stamens and style more or less exserted. Achenes ovate to elliptic, 1.0-1.1 mm. long, 0.8—0.9 mm. wide, smooth, more or less rubiginose, often reddish brown-maculate, glabrous on the abaxial, densely hirtellous on the adaxial side, abaxial ribs indistinct. Calyx- limb 6- to 7-fid. Fig. 42. Distribution of V. robertianifolia. 1951] MEYER—VALERIANA IN NORTH AMERICA 471 > ILO ЛО i D Nor XN Ж К ЖАР \ Y \ N we кА „А VA {7 МЫК % 3 QU N À Ў -- y SA АТУ NS AO NS RAN Fig. 43. Valeriana robertianifolia: Achenes, X 20; pistillate flower (above) and staminate (below), X 10. Type Locarrrv: "env. d'Orizaba. 1854—55.” DISTRIBUTION: Oak-pine forests, moist rocky slopes, along roadways, 4900— 8500 ft. alt. South-central Mexico to Costa Rica. Flowering and fruiting August to December. Mina, Pilas, Hinton 10702 (GH); Dist. Galeana, Carrizo-Sto. Domingo, Hinton 14733 (F, GH, MO, NY); near Omilteni, Sharp 441549 (MO). jaLisco: without definite locality, Jony s. n. (US). Mexico: Dist. Temascaltepec, Comunidad, Hinton 1568 (GH, К, UC). міснолсАм: vicinity of Morelia, Cerro Azul, Arsène 2458 (GH, MO, NY, US); vicinity of Morelia, Arséne 3225 (US); vicinity of Morelia, Cerro San Miguel, Arséne 6071 (MO, US); vicinity of Morelia, Loma Santa Maria, Arséne 9024 (US); Dist. Tancitaro, Cerro Tancitaro, Leavenworth 725 (F, MO); Patzcuaro, Pringle 3341 (D, F, GH, MO, NY, 5, UC, US, WYO). MonELos: barranca near Cuernavaca, Pringle 5006 (MO). oaxaca: Sierras 4 8000 pieds, Galeotti 2076 (P). VERA CRUZ: Orizaba, Botteri 1192 (D, US); Vallée de Orizaba Escamella, Bourgeau 2047 (GH, P, S, US); St. Miquel, Liebmann 10855, 10856, 10857 (US); Engenio, Sierra de la Cruces, Müller 1164 (NY). GUATEMALA: CHIMALTENANGO: Alameda, Jobnston 0 F); Tecpán, Skutch 580 (MU, US); road from Chimaltenango to San Martin Jilotepeque, Standley 57891 (GH, NY). HUEHUETENANGO: Cumbre Papal, between Culico and Ixmoqui, Steyermark 50936 (F). JALAPA: Laguna de Ayarza, Heyde 6 Lux 3970 (F). [Vor. 38 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN HoNDbUnAs: MORAZÁN: Cerro de Uyuca, region of El Valle Encantado, Standley, Allen, Sbank, Padilla V. 806 (F). Costa Rica: Piedra Blanca, Brenes А4 (F); Potreros of Rancho Redondo, Dodge & MU, US); Hacienda La Esperanza, Jimenez 05 (US); region of Zarcero, Palmira, Smith A206, A538 (F); Prov. Alajuela, Palmira, Alfaro Ruiz, Smith NY1242 (NY). The distinguishing characters of V. robertianifolia are the pinnate-bipinnatifid leaves with the terminal lobe palmately 3-lobed or sometimes 5- to 10-cleft or dentate, the minute flowers, 0.5-1.0 mm. long, the exserted stamens, the small ovate to elliptic achenes, 1.0-1.1 mm. long, and the calyx-limb 6- to 7-fid. Valeriana robertianifolia occurs also in northern South America, but the material that I have seen from Venezuela (V. venezuelana Briquet) and Colombia is not sufficiently distinct from the North American populations to be placed in a sep- arate category. This species is closely related to V. sorbifolia, from which it is sometimes rather difficult to distinguish. 28. VALERIANA PALMERI Gray in Proc. Am. Acad. 22:417. 1887. Т. Palmer 754! (GH). Valeriana Langlassei Briq. in Ann. Conserv. & Jard. Bot. Genève 17:341. 1914. T. Langlassé 408! (D, GH, K, US). Valeriana fistulosa Briq. l. с. 343. 1914. Т. Galeotti 2565! (BR, D, P). Annuals 2.6-20.0 dm. tall, slender to robust, from subnapiform tap-roots 1—8 mm. thick. S/em leafy, 0.1—1.5 cm. thick, terete or sometimes quadrangular towards the base, spreading-pilosulous to pilose, mostly towards the base, glabrescent above. Leaves cauline, 6—7 pairs, petiolate, pinnate to pinnatifid or sometimes undivided, elliptic- or oblanceolate- to obovate-spatulate, acuminate, 3.5—33.0 cm. long, 0.9—20.0 cm. wide, serrate to irregularly dentate or essentially entire, spread- ing-ciliate, sparsely pilosulous above, the veins below pilosulous or glabrous, the terminal lobe of the divided leaves ovate to elliptic, acuminate to subcaudate, acute, 5.4-14.5 cm. long, 2.3-8.0 cm. wide, more or less decurrent on the rhachis, the lateral lobes 1—5 pairs, linear-oblong, relatively much narrower than the terminal lobe, 1.7-6.0(-9) cm. long, 0.3-2.0(-3.5) cm. wide, grading smaller, somewhat decurrent, the rhachis more or less winged; petioles 1-9 cm. long, spreading-ciliate or uniformly pilosulous, extending to the rhachis. Inflorescence an aggregate dichasium, 11—40 cm. long, 5—23 cm. wide, the terminal scorpioid branches 0.3—3.5 ст. long, glabrous; bracts 1-2 mm. long, glabrous; flowers hermaphroditic, rarely gynodioecious. Corolla infundibuliform to subcampanulate, 1.1—1.9 mm. long, glabrous without, white, the lobes less than half the length of the gibbous tube, the throat sparsely pilosulous within. Stamens and style included. Achenes elliptic- to oval-patelliform, thin, 1—3 (-4) mm. long, 0.9—2.5 mm. wide, smooth or some- times rugulose, often reddish brown-maculate, spreading-pilosulous on the adaxial side, glabrous on the abaxial, rarely glabrous or pilosulous on both sides, adaxial margins more or less winged, plane or undulate. Calyx-limb 8- to 11-fid. 1951] MEYER—VALERIANA IN NORTH AMERICA 473 Fig. 44. Distribution of V. Palmeri. Type Locarrrv: “Rio Blanco, on the river bank, near Guadalajara", Jalisco, Mexico. Sept. 1886. DisrRIBUTION: Cut-over and pastured slopes, on limestone outcrops, oak-pine canyons, crevices of rocks near water, fields, 2500-7000 ft. alt. Sierra Madre Occidental, Chihuahua to Costa Rica. Flowering and fruiting June to December. MER CHIHUAHUA: Sierra Canelo, Río Mayo, Gentry 2491 (F, GH, GENT, MO, NGO: without definite locality, Garcia 933 (US). GUERRERO: Dist. Mina ( pancingo on road to Chilapa, Moore & Wood 4628 ; 3 kms. past Acahuizotla to Acapulco, Moore & Wood 4678 (MO). Jjarisco: Río Blanco, Palmer 754 (GH); near Guadalajara, Pringle 4521 (D, F, GH, MO, NY, UC, US, W MEXICO: Dist. Temascaltepec, Hinton 1115 (GH, MO); Dist. Temascaltepec, Chorrera, Hinton 2630 MICHOACÁN: vicinity of Morelia, Rincón, Arséne 2538 (GH, MO, NY, S, US); vicinity of Morelia, prés Cerro de las Nee Arséne 2567 (GH, NY, US); vicinity of Morelia, a San Miguel, Arséne 5262 (MO, US); vicinity of Mo cilia Punguato, Arséne 5745 , MO, NY, US). Nayarit: La Ataries, n. Yxtlan, Mexia 676 (F). OAXACA: Interpeje, pot 2565 (BR, D, P). vera CRUZ: Orizaba, Müller з.п. (МҮ); Zacuapan, co 1 (UC). EMALA: CHIQUIMULA: Montafia Nube, between Socorro Mt. and Cerro Brujo, Шы 30005 (Е). GUATEMALA: without definite locality, Aguilar 122 (F) HUEHUETENANGO: along Río Trapichillo, Steyermark 51023 (F). JALAPA: between Jalapa and San Pedro Pinula, Standley 77103 (F); Montaña Durazno, Steyermark 32068 ). SANTA ROSA: Santa Rosa, Heyde & Lux 2924, 2024Ь (GH, US); Estanzuela, Heyde & Lux 3068 (NY, US); se. Barberena, Standley 77863 (F). EL SALVADOR: on the Guatemalan frontier near Chalehuapa, Calderón 1061, 1062, 1062a (US). Costa Rica: Cerros de San Antonio de San Ramón, Brenes 5616 (F); San Pedro de San Ramón, Brenes 16276 (F); Carrillos de Poas, Brenes 19300 (F); La Palma de San Ramón, Quiros 273 (F); Hacienda La Argentina, Grecia, Valerio 591 (F, US). [VoL. 38 474 ANNALS OF THE MISSOURI BOTANICAL GARDEN Valeriana Palmeri may be distinguished most readily by the achenes which are more or less winged and elliptic- to oval-patelliform. It is most closely related to V. s. sorbifolia but differs from that species by the pinnatifid leaves which are usually more or less decurrent on the winged rhachis. 29. VALERIANA SORBIFOLIA HBK. Nov. Gen. et Sp. 3:332. 1819. T.: Humboldt & Bonpland (MO photo, Р). Annuals 1.3—12.0 dm. tall, slender to robust, erect or rarely clambering, from subnapiform or fusiform tap-roots 0,3-1.3 cm. thick. Stem leafy, 1-10 mm. in diameter, pilosulous or sometimes short-pilose or glabrescent mostly towards the base, the upper portion glabrous. Leaves cauline or basal, the cauline 4—7 pairs, petiolate, pinnate, rarely undivided, ovate, elliptic- to obovate-spatulate, obtuse to acuminate, (1.0-)2.7-15.0(-19.5) cm. long, (0.5—) 1.2-10.0 cm. wide, serrate to subcrenate, dentate to repand or essentially entire, glabrous or occasionally scattered- pilosulous on both surfaces or only on the veins below, the terminal lobe of the divided leaves elliptic to obovate or suborbicular, obtuse to acute, sometimes more or less acuminate, 1.5-8.2 cm. long, 0.8—7.2 cm. wide, abruptly expanded or some- times decurrent on the rhachis, the lateral lobes 1—6 pairs, distinct, simulating the terminal lobe, grading smaller; petioles 2.0—6.3 cm. long towards the base, obsolete above, glabrous or pilosulous; basal leaves undivided, 9—12 cm. long, suborbicular, ү. S. BARBAREAEFOLIA ү. S. SORBIFOLIA Fig. 45. Distribution of the varieties of V. sorbifolia. 1951] MEYER—VALERIANA IN NORTH AMERICA 475 repand-dentate, petioles 7.5-10.0 cm. long. Inflorescence an aggregate dichasium, 8—55 cm. long, 4—10 cm. wide, the terminal scorpioid branches 0.2—4.0 cm. long, glabrous or sometimes tufted-pilosulous on the nodes; bracts 1.3-2.0 mm. long, glabrous; flowers gynodioecious. Corolla infundibuliform to subcampanulate, that of the perfect flower 1.0—2.8 mm. long, of the pistillate 0.5-1.0 mm. long, glabrous without, white, the lobes less than half the length of the gibbous tube, the throat glabrous or sparsely pilosulous within. Stamens and style included. Acbenes ovate to ovate-oblong, often somewhat ampulliform, 0.6-2.0 mm. long, 0.4-1.6 mm. wide, smooth or rarely scabrous, tawny to rubiginose, sometimes purple-maculate, spreading-hirtellous on the adaxial side, glabrous on the abaxial, sometimes glabrous or hirtellous on both sides, adaxial margins more or less involute. Calyx-limb 6- to 11-fid. I have recognized three varieties as provisional categories under V. sorbifolia, mexicana, sorbifolia, and barbareaefolia, in order to point out a series of semi- stabilized. variants within the range of variation of this species. These variants may actually be potential subspecies, but geographic barriers have not as yet been ascertained for these infraspecific taxa. The variability within V. sorbifolia sug- gests, furthermore, that incipient hybridization exists between V. sorbifolia, Palmeri and robertianifolia. KEY TO THE VARIETIES A. poc кы. Т Vera Cruz, Puebla, State of Mexico, to 29a. V. s. mexicana Oax AA. Moe c ap-root more or less subnapiform, unforked, 3-7 mm. thick, 0.5—1.0 cm. Leaves predominantly serrate to subcrenate, mostly with 3— Ж s of lateral mg Southeastern Arizona, Baja UM northwestern Mexico to Panama; also in northern South 29b. V. s. sorbifolia BB. Tap root more or less шры frequently forked, 0.6— cm. thick, 1.5—8.0 cm. long. predominantly dentate to ie nd or mim entire, тегін МБ 2 pairs of lateral lobes. North- eastern Mexico to Guatemala... 29c. V. s. barbareaefolia 29а. VALERIANA soRBIFOLIA HBK. var. mexicana (DC.) F. G. Mey., stat. nov. Valeriana mexicana DC. Prod. 4:640. 1830. T.: Berlandier 902! (BM, D, MO photo, F, P, W). Tap-root more or less subnapiform, 3-7 mm. thick. Leaves cauline, 4—7 pairs, more or less uniformly distributed, pinnate or occasionally undivided, predomi- nantly serrate to subcrenate, the terminal lobe to 8.2 cm. long, 7.2 cm. wide, the lateral lobes mostly 1-2 pairs. Achenes scabrous, 1.2-1.9 mm. long, 0.6-1.3 mm. wide, hirtellous or glabrous Tyre Locarrry: “around the city of Mexico.” DISTRIBUTION: Roadsides, open woodlands, 5300-9500 ft. alt. South-central Mexico to Guatemala. Flowering and fruiting August to February. MEXICO: FEDERAL DISTRICT: Vallée de Mexico, Guadalupe, Bourgeau 601 (D, С, GH P, S). oaxaca: 18 mi. sw. C. Oaxaca, Nelson 1389 (GH, US); Vallée de Etla, Smith [Vor. 38 476 ANNALS OF THE MISSOURI BOTANICAL GARDEN 787 (GH). PUEBLA: vicinity of Puebla, Cerro E acre M IQI (US); vicinity of Puebla, Teocalli de Piero Arséne 2062 (GH, MO, US); above Serdan, Cabecero, Sharp 44984 (MO). vera cruz: Orizaba, Екі, nie = la rS] Müller 290 (NY), 343 (СН); without definite locality, емін Hahn P). 2 MALA: HUEHUETENANGO: Huehu ds Standley 65732 (Е). JALAPA: Volcán Jumay, north of Jalapa, Steyermark 32436 (F). SACATEPEQUEZ: near Antigua, 2. 58632 (Е). The var. mexicana may be distinguished from var. sorbifolia by the usually broader leaves and the scabrous achenes. 29b. VALERIANA SORBIFOLIA HBK. var. SORBIFOLIA. Valeriana toluccana DC. Prod. 4:640. 1830. T.: Berlandier 1130! (BM, D, G, OXF, P, W). Valeriana gracilis Benth. Pl. Hartweg. 196. 1839. T.: Hartweg 1079! (BM, CGE, D, FI, PPM Tap-root more or less subnapiform, 3-7 mm. thick. Leaves cauline, 4—7 pairs, more or less uniformly distributed, pinnate, predominantly serrate to subcrenate, the terminal lobe 1.5—6.0 cm. long, 0.8—3.8 cm. wide, the lateral lobes mostly 3-6 pairs. Achenes smooth, 0.6-2.0 mm. long, 0.5-2.0 mm. wide, pilosulous or glabrous. ТҮРЕ Locarrry: "inter Valladolid (Morelia) de Michoacán et Pátzcuaro,” Mexico. DISTRIBUTION: Densely wooded arroyos in pine forests, oak woods, cultivated fields, 3300-8800 ft. alt. Southeastern Arizona; Baja California, northwestern Mexico to Honduras and Panama. Mrxico: BAJA CALIFORNIA: La Chuparosa, Brandegee s. n. (UC, US); Sancito, Brandegee s. n. (GH, NY, US); mts. near Todos Santos, Brandegee s. n. (UC); Sierra de Laguna, Brandegee 274 (UC); Sierra de San Francisquito, Brandegee 274 (UC); The Laguna, Laguna Mts., Jones 27846 (МО, NY, UC). cHiAPAs: Finca Fuarez, Escuintla, Matuda s. n. (MU); Siltepec, Matuda 0804 (Е, MO, MU); near San Cristobal, Nelson 3146 (СН. US); Finca Conadonga, Purpus 6702 (UC); Cerro del Boquerón, jen 6703 (F, NY, UC). снінолноА: ridge between Río Chico and Río Caballo, Barlow s. n. e Loreta, Río Mayo, Gentry 2566 (GENT); Cafion de St. Diego, qo aie. (Е, G NY, UC, US); Sierra Madre Mts., Guayanopa Canyon, Jones s. (US); Majarachie Knoblock 5284 (F); Chuhuichupa, LeSueur 1253 (F); Dist. Modus. 7 mi. w. Chu chupa, Miiller 3571 (GH, UC); Sierra ue 35 km. sw. Minaca, Pennell 18870 (US); near Guerrero, Pringle 1256 (Е. F, GG, GH, К, МО, NY. S, UC, US). FEDERAL DISTRICT: Cacubava, San Juanico, Arséne 0885 (05); $асготоге Hill, near Amecameca, M p рны s.n. (MO); Pedregal, Po 800 (GH, P); Cuajimalpa, Lyonnet 373 (US); Churu- busco, Orcutt 4288 (MO); кир М, 7085 (F, GH, WYO). GUANAJUATO: without definite нЕ peris GH). GUERRERO: Dist. Montes de Oca, Vallecitos, Hinton 11380 (СН); of bluff west of x) aps Sharp 441412 (MO). HIDALGO: Dist. Volango, vicinity үт Nola Moore 1086 (GH). ултлѕсо: trail from San Sebastian west to Mascota, Mexia 1404 (D, F, GH, MO, NY, UC, US). mexico: Dist. Temascal- Sali 4636 (MO, UC); Dist. Temascaltepec, Cuentla, Hinton 7217 (GH); Dist. Sultepec, p Gavia, Hinton 8376 (GH); Ixtacihuatl, Purpus 1783 (D, F, GH, NY, , US) via, at km. 105 on Toluca-Morelia highway, Sharp 44200 (MO). MICHOACÁN: vicinity of Morelia, Loma Santa Maria, Arsène 5, E (US); ; vicinity of Morelia, prés Cerro 1951] MEYER— VALERIANA IN NORTH AMERICA 477 de las Nalgas, Arséne 2567 (MO); vicinity of Morelia, Rto. de Mexico, Arséne 5901 (US); vicinity of Morelia, Cerro Azul, Arséne 6571 (GH, MO, NY, US); vicinity of Morelia, Trapeo, Arséne 9857 (US); Coalcomán, Hinton 12518 (К); Coalcomán, Torricilles, Hin- ton 12803 (GH, K) ; Zitacuaro-Guanoro, Hinton 13481 (СН, К); Arostes, prope Uruapan, Woronow 2682 (F). MoRELos: Vallée del Pepeite, Lyonnet 9 Elcoro 1790 (US); bar- тапса near Cuernavaca, Pringle 9837 (F, GH, МО, NY, US). OAXACA: in monte San Felipe, prope Oaxacam, Andrieux 325 (D, K, P); Mt. Zempoaltepec, Nelson 590 (US); near Reyes, Nelson 1773 (US). PUEBLA: vicinity of Puebla, Rancho Casada, Arsène © Nicolas 323 (GH, MO, NY, US); vicinity of Puebla, Manzanilla, Arsène 1635 (GH, US); vicinity of Puebla, Boca del Monte, Arsène 7000 (US); vicinity of Puebla, near Totime- huacán, Hacienda Bata, Arsène 9879 (US); bord de l'Atoyoc, Nicolas s.n. (D, E, F, GH, P). SAN LUIS POTOSI: in arenosis humidis circa urbem, Schaffner 107 (GH). SONORA: Rinconada, Lloyd 304 (GH); 4 ті. e. El Bilito, White 4781 (MU). vera CRUZ: Orizaba, Botteri s. n. (D, F, MO); Castresana, Liebmann 10831; inter S. Andres et S. Miguel, Lieb- mann 10833; Jovo, Liebmann 10838 (0 EMALA: ALTA VERAPAZ: se. Tactic, Standley 69958 (F); near Coban, Standley 71566 (F); along Río Carcha, between Cobán and San Pedro Carcha, Standley 90143 (F). CHIMALTENANGO: Volcán de Agua, Johnston 819 (F); Alameda, Johnston 044, 052 (Е); near Tecpán, Skutch 485 (GH, US). EscuINTLA: without definite locality, Aguilar 1584 (F). GUATEMALA: without definite locality, Tonduz 888 (GH, NY, US); Chillom, Rojos 51 (GH, US). HUEHUETENANGO: Cuesta de la Concepción, zur Rainen, Seler 3251 (GH, NY); Sierra de los Cuchumatanes, Steyermark 50356 (F); Cerro Pixpix, above San Ildefonso Ixtahuacan, Steyermark 50571 (F); La Sierra (Tumimach), adjacent San Juan Atitlan, Steyermark 51965 (F). JALAPA: between Jalapa and Paraiso, Standley 77331 (F); Montaña Durazno, 2 mi. e. San Pedro Pinula, Steyermark 32992 (F). SACATEPÉQUEZ: near Antigua, Standley 58611 (F); Cuesta de las Canas, above Antigua, Standley 58983 . SAN MARCOS: between Finca El Porvenir and Loma Corona, 9 mi. nw. El Porvenir, Steyermark 37751 (F). zacapa: Sierra de las Minas, Steyermark 29685 (F). HONDURAS: MORAZÁN: Cerro Uyaca, Allen 3903 (МО); Uyuca, Valerio 701 (F). The var. sorbifolia may be distinguished by the leaves, which are usually pinnate with the lobes predominantly serrate to subcrenate, and by the flowers with ex- serted stamens. Valeriana sorbifolia is closely related to V. robertianifolia and V. Palmeri, and where the distributions of these species overlap a greater incidence of variability may be recognized in the ensuing populations. I suspect that hybridiza- tion between these taxa contributes to the confusion in determining certain "difficult" specimens. 29c. VALERIANA sorBIFOLIA HBK. var. barbareaefolia (Mart. & Gal.) F. G. Mey., stat. nov. Valeriana barbareaefolia Mart. & Gal. in Bull. Acad. Brux. 111:121. 1844. T.: Galeotti 2549! (BR, MO photo). Tap-root more or less fusiform, frequently forked, 0.6-1.5 cm. thick, 1.5-8.0 cm. long. Leaves disposed mostly towards the base, more or less crowded on the foreshortened internodes, pinnate, predominantly dentate to repand or essentially entire, the terminal lobe 2.3—4.7 cm. long, 2-4 cm. wide, the lateral lobes mostly 2 pairs. Achenes smooth, 1.6-2.0 mm. long, 0.8—1.6 mm. wide, pilosulous or glabrous. Type Locarrrv: "Real del Monte et de Moran, au nord de Mexico, de 7500- 8500 pieds." [Vor. 478 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 46. Valeriana sorbifolia var. barbareaefolia: Habit, X V4; еке н side), Xx 8; „еме М (above), entire and dissected staminate lower (below), X 1951] MEYER—VALERIANA IN NORTH AMERICA 479 DISTRIBUTION: Oak-pine or mixed forests in moist soil or on boulder-covered slopes, 5500—10500 ft. alt. Sierra Madre Oriental from Nuevo Leon to southern Vera Cruz and Oaxaca to Guatemala, overlapping with var. sorbifolia in the southern part of the range. Mexico: CHIAPAS: Mt. Male, near Porvenir, Matuda 4601 (GH, MO, NY). COAHUILA: nw. Fraile, Stanford, Retherford, Northcraft 410 (MO). Hwatco: Real del 24 km. Monte, Ehrenberg 173 (GH); San Vicente, Fisher 3727 (MO, NY, US); Sierra de Pachuca, Pringle 69047 (D, F, GH, MO, MU, NY, S, UC, US, WYO); Dist. Pachuca, above Pueblo LEON: Dist. Aramberri, Cerro Lifiadero, Meyer 9 Rogers 2024 (MO); Dist. Zaragoza, Cerro del Viejo, Meyer & Rogers 2080 (МО); 15 mi. sw. Galeana, Mueller 922 (MU); 40 mi. s. Saltillo, Palmer 416 (GH, US). oaxaca: Sierra de San Felipe, Pringle 4837 (D, F, GH, MO, MU, NY, UC, US); Cerro Verde, Purpus 3337 (Е, GH, MO, NY, UC, US), 3513 (UC). PUEBLA: Cerro de Chicamole, Purpus 6405 (UC). SAN LUIS POTOSI: Alvarez, Palmer 244 (F, GH, MO, NY, US); region of San Luis Potosi, Parry & Palmer 312 (GH). TAMAULIPAS: summit Рейа Nevada, Stanford, Lauber, Taylor 2587 (US). VERA CRUZ: Engenio, Sierra de Cruz, Müller s.n. (NY); Maltrata, Mt. Orizaba, Seaton 387 (F, GH, US); near El Puerte, Sharp 44677 (MO). The var. barbareaefolia is distinguished from var. sorbifolia by the fusiform root, repand-dentate basal leaves, and broader achenes. In the northern Sierra Madre Oriental it occurs to the exclusion of var. sorbifolia, although in the southern part of the range the distributions of the varieties overlap. Series УП. PRATENSES F. С. Mey., n. ser. Perennials or annuals from fusiform to napiform tap-roots. Stem leafy. Leaves basal and cauline, pinnate to bipinnatifid, elliptic to obovate-spatulate, the lateral lobes usually more or less decurrent on the rhachis, becoming dilated and 1- to 4- cleft at the tip or short-laciniate; rhachis usually winged. Inflorescence a compound or sometimes an aggregate dichasium, flowers hermaphroditic. Corolla infundibuli- form, 2-9 mm. long, the tube gibbous towards the base or sometimes near the middle, the throat densely pilosulous. Stamens and style exserted, anthers 2- lobed, the loculae equal in length. Achenes elliptic to obovate-oblong, 2—6 mm. long, glabrous or densely hirtellous to subsericeous. Calyx-limb setose or short- cupuliform and more or less dentate. Species, 2. ТҮРЕ Species: Valeriana pratensis (Benth.) Steud. DISTRIBUTION: Mexico. KEY TO THE SPECIES A. Annuals. Stem single. Leaves cauline, with the ultimate divisions short-laciniate, the rhachis distinctly winged throughout. Corolla 2.0—2.5 mm. long. Achenes 2.0-2.5 mm. long, adaxial margins more or less іпуоімее.................. ш.. м 1..1 тр... Д.Д. 30. V. tanacetifolia AA. Perennials. Stems often several Leaves disposed towards the base, with the ultimate divisions palmately 3-lobed and cleft, the rhachis only slightly winged part way. Corolla 5.5-9.0 mm. long. Achenes 4.5—6.0 mm. long, margins plane 31. V. pratensis [Vor. 38 480 ANNALS OF THE MISSOURI BOTANICAL GARDEN 426 29. ЖАР Fig. 47. Valeriana tanacetifolia: Habit, X about V2; entire and dissected flower, and achenes (abaxial and adaxial sides), X 8. 30. VALERIANA tanacetifolia F. G. Mey., spec. nov. Planta annua 3—5 dm. alta tenuis erecta ex radice primaria crassa subnapiformi 5-7 dm. diam. Caules foliacei 1—2 mm. crassi ex nodis in lineis decurrentibus parce retrorso-hirtelli aliter glabri. Folia caulina 3—8 jugata petiolata bipinnatifida 1951] MEYER—VALERIANA IN NORTH AMERICA 481 elliptica vel obovata vel obovata-spatulata 4.8-8.0 cm. longa 2.5-3.5 cm. lata, lobis lateralibus lobos terminales simulantibus 2-4 jugis ad apicem dilatatis 1-4 fidis vel brevi-laciniatis acutis in rhachi alato decurrentibus 1.5-2.0 cm. longis 0.2-0.5 cm. latis supra glabris vel appresse hirtellis aliquando ascendenti-ciliolatis nervo medio infra sparse hirtello in internodium subtendentem decurrentibus deinde anguste alatis. Inflorescentia dichasium compositum in anthesim 0.9—1.5 cm. lata, nodis glabris vel aliquando cristato-pilosulis, bracteis 2.2-3.9 mm. longis supra reductis, glabris vel aliquando patenti-ciliatis; floribus hermaphroditicis. Corolla infundibuliformis 2.0-2.5 mm. longa alba extra glabra tubo ad medum parum gibbo lobis duplo longiori intus dense pilosulo. Stamina et styli exserti. Achaeniae ellipticae 2.0-2.5 mm. longae glabrae laeves et aliquando fulgentes brunneolae aliquid concavae marginibus plus minusve involutis costis abaxalaribus non promi- nentibus. Calycis limbus 10-fidus (?). Type Locauity: “by the crater lake," Volcán, 1500 m., District of Temascal- tepec, Mexico. September 9, 1932. DISTRIBUTION: Oak woods in thin rocky soil, 4900-5600 ft. alt. District of Temascaltepec, State of Mexico. STATE OF MEXICO: Dist. Temascaltepec, Hinton 1417 (GH); Dist. Temascaltepec, Mong Hinton 1663 (К ноготүре, GH, MO photo); Peñon Timbres, Hinton 4420 (G ). WITHOUT DEFINITE LOCALITY: Sessé 5 Mociño s. n. (G, MO, OXF). Valeriana tanacetifolia maintains a local distribution. It may be easily dis- tinguished by the short-laciniate bipinnatifid leaves, annual habit, the shallowly concave, elliptic and glabrous achenes, and the corolla tube which is usually indis- tinctly gibbous towards the middle. This species was first collected by Sessé & Mociño while on the Royal Spanish expedition to Mexico at the end of the 18th century. So far as I am aware this plant was not collected again for 130 years when George Hinton recollected it in the district of Temascaltepec. 31. VALERIANA PRATENSIS (Benth.) Steud. Nom. Bot. 22:742. 1841. Astrephia pratensis Benth. Pl. Hartweg. 39. 1839. T.: Hartweg 302! (BM, CGE, K, OXF, Valeriana Galeottiana Mart. in Bull. Acad. Brux. 111:124. 1844. T.: Galeotti 2547! (BR, K, P). Phyllactis pratensis (Benth.) Benth. & Hook. Gen. Pl. 2:153. 1873. Perennials 2.4-5.0 dm. tall, from napiform to fusiform tap-roots, becoming verrucose in age, 0.5-2.0 cm. thick. Stem moderately leafy, 1-3 mm. thick, glabrous, the nodes cften sparsely pilosulous. Leaves basal and cauline: the basal more or less imbricate, sometimes forming a rather loose rosette, petiolate, pinnate- bipinnatifid, еШіргіс- to obovate-spatulate, 9.5—27.5 cm. long, 2.7-5.0 cm. wide, ascending-ciliolate, sometimes scattered spreading-pilosulous, usualy on the veins only, the terminal lobe 2—5 cm. long, 1.5-2.2 cm. wide, decurrent on the rhachis, usually palmately 3-lobed or cleft, these 1- to 2-cleft, subacute to obtuse, the [Vor. 38 482 ANNALS OF THE MISSOURI BOTANICAL GARDEN lateral lobes 2—5 pairs, distinct or somewhat decurrent on the rhachis, shorter than the terminal lobe, grading smaller, 1- to 3-cleft, acute; petioles 5-22 cm. long, glabrous or hirtellous on the adaxial side, spreading to the rhachis, the margins sometimes retrorse-aculeate towards the base; cauline leaves 3—4 pairs, simulating the basal, 3.3—6.2 cm. long, 1.2-4.0 cm. wide. Inflorescence usually a compound or sometimes an aggregate dichasium, the terminal dichotomies 2.0-3.5 cm. wide in anthesis, later diftuse, to 36 cm. long, 15 cm. wide, the nodes usually tufted- pilosulous, the internodes glabrous; bracts 6-11 mm. long, reduced above, glabrous or spreading-ciliate; flowers hermaphroditic. Corolla infundibuliform, 5.5-9.0 mm. long, glabrous without, white, the tube gibbous towards the base or some- times indistinctly gibbous towards the middle, the lobes about 15 the length of the tube, the throat densely pilosulous within. Stamens and style exserted. Achenes ovate-oblong to oblong-elliptic, 4.5-6.0 mm. long, 2.9-4.0 mm. wide, glabrous or densely hirtellous to subsericeous, abaxial ribs more or less carinate. Calyx-limb short-cupuliform, more or less dentate. Type Гослілтү: Morelia (Michoacán), Mexico. у. TANACETIFOL IA 3 v. ENSIS Fig. 48. Distribution of V. tanacetifolia and V. pratensis. DISTRIBUTION: Wet meadows, 7000—8300 ft. alt. South-central Mexico. EXICO: DURANGO: El Salto unm Asok 18302 (UC, E GUANAJUATO: without definite locality, Dugès s. и. RERO: Dist. Mina, Aquazarca-Filo, Hinton 10501 (GH). мехісо: Dist. баш Nanchititla, Hinton 6344 Se? B Del Sn Pringle 3641 (F, GH); Valley of Toluca, Prin gle 4199 (D, GH, M MICHOACAN: Morelia, Hartweg 302 (BM ОССЕ, K, ОХЕ, P, V). NAYARIT: near 2. Ten. Ross 2120 (NY, US). VERA CRUZ: Jesus del Monte, Galeotti уу (BR. D, K, P). Valeriana pratensis is the only Mexican species restricted to wet places, and it may be distinguished by the pinnate-bipinnatifid, ascending-ciliolate leaves and by the achenes which are 4.5-6.0 mm. long. This species 1s obviously related to V. tanacetifolia, but there is no reason to confuse the identity of either species. 1951] MEYER—VALERIANA IN NORTH AMERICA 483 IMPERFECTLY KNOWN OR EXCLUDED SPECIES Valeriana cyclophylla Graebn. in os Bot. Jahrb. 37:437. 1906. T.: Scbumann 50 is robably V. sCANDENS. e type of V. CYCLOPHYLLA was destroyed at Berlin. к. latifolia Mart. & Gal. in Bull. Acad. Brux. 11!:124. 1844. T.: Galeotti 2558! (BR). — BOERHAAVIA ERECTA p obovata (Nutt.) R. & S. Mant. 1:265. 1822 (Pbyllactis obovata Nutt. Gen. m. Pl. 1:21. 1818). Description not applicable to Valeriana; type not extant. Valeriana ramosa Sessé & Mociño, in Fl. Nov. Hisp. ed. 2. 10. 1893. = V. CERATOPHYLLA BK. (?), description inadequate. 484 No w © NAv ы К) 10. 11. — N -. ” [Vor. 88 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN ENUMERATION OF TAXA V. officinalis L. V. sitchensis Bong. a. ssp. sitchensis Ь. ssp. Scouleri (Rydb.) F. С. Mey. c. ssp. uliginosa (Torr. & Gray) F. G. ey. V. capitata Pall. ex Link a. ssp. capitata b. ssp. californica (Heller) F. G. Mey. c. ssp. pubicarpa (Rydb.) F. С. Mey. d. ssp. acutiloba (Rydb.) F. G. Mey. V. arizonica Gray V. pauciflora Michx. V. columbiana Piper V. occidentalis Heller V. dioica L. ssp. sylvatica (Sol. ex Richards.) F. G. Mey. V. texana Steyermark V. prionophylla Standl. V. edulis Nutt. ex Torr. & Gray a. ssp. edulis b. ssp. ciliata (Torr. & Gray) F. G. Mey. с. ssp. procera (НВК.) F. С. Mey. . V. laciniosa Mart. & Gal. 13. V. albo-nervata Fernald V. ceratophylla HBK. 31. ore tse eee ax = “мала clematitis HBK. Selerorum Graebn. & Loesn. urticaefolia HBK. cucurbitifolia Standl. palmatiloba F. G. Mey. apiifolia Gray vaginata HBK. densiflora Benth. var. densiflora var. affinis (Mart. & Gal.) F. G. ey. deltoidea F. G. Mey. pulchella Mart. & Gal. scandens L. var. scandens var. Candolleana (Gard.) Muell. domingensis Urban X Ekmanii F. G. Mey. robertianifolia Briq. Palmeri Gray sorbifolia HBK. var. mexicana (DC.) F. G. Mey. var. sorbifolia var. barbareaefolia (Mart. & Gal.) Mey ‚ tanacetifolia F. С. Mey. V. pratensis (Benth.) Steud. T9511 MEYER—VALERIANA IN NORTH AMERICA 485 INDEX TO EXSICCATAE Italicized numbers refer to collector’s numbers, s.n”. (sine numero) to unnumbered collections; parenthetical numerals refer to the taxa conserved in this treatment which are listed in the numerical sequence indicated on opposite page. Abbott, W. H. s.n. (11b). 163 (22b); 279 (22b); Abbott, W. L 1800 ( Abrams, L. R 4765 (3b); 9017 (2a); 9842 (2a Adeney, M. s үй 3а). E JE 29b); 122 (28); 133 Uo (25b); 927 (25); 1584 (39 Aguilar, M. H. a ee Alexander, A. M. & Kellogg, L. 158 (2 а); 1849, 1859, 3885, 4075, 4302, 4850 (3b). ‚п. (5). Alexander, R Allard, H. A. Я (2с). АПеп, Ј. s. n. (2с). Allen, O. D. 243 (2b). Allen, P H E Шу Anderson, J. P. s.n., I 8,927, R1024, 1924, 5405; же e. 1936, 6488b (3а); s n. (3d). BT P. & Gassner, С. W. 7438 (3a). 814% Anderson, J. К. s^., (2а); s.m 2Ъ . 323 (15); s. n., (22b); 325 (29b); 326 (12). Anselme-Marie, Bro. TO. Applegate, E. I. 84I, 2306 (2b); 2468 (7); 4770 P oe (11a); 9078 (2a). Arnold, J. F. 46 (11a). Arrivée, David % "a (7% 1617. 2l, 292, 29b); 324 3500 (22a); 517 5262 (28); 5203 (пе); 5424 (17); 5482 (16); 5515 (29Ь); 5574 (29а); 5745 (28); 5901 (29b); 6071 (27); 6553, 6571 (29b); 6667 (22a); 6698 (17); 6748, 6981, 7000 (29b); 7274 ; 8662, 8665 (29b); 988I (27 291): 0884 ded ' 0885 (29b); 10114 (27); 18446 (6 Arséne, С. & Nicolas --. 323 (29b). . (11b). Babcock, H. H. DE Pw 4). ker, C. Е. 5.п. (11а); 136 (7); I sors 313 (34); 4ó1 TAY 620 Lu. 0 (25a); 1146 (3b); 4376 уа j e. Gn Barle,: Кл 52 s.n". (11а); 120 (7); 25. Od E la Baker, K. | 52% Ball, 7 ( eee о, Р. 8. © #. (3b). Bartlett, Н. Н. 11466, 12828, 13033 (25a). Bartley, Е. & Pontius, L. L. 872 (11b). 23). & Chapman, R. 2251(11а). Beauchamp, B. s.n. (29b). 486 Bebb, M. S. s.n. (11b). Beck, D. E. s.n. (7). Beck, ПІ. C.?] s.n. (2c) Beckett, К. Е. 11710 (4) Beckwith, F Beechey's Voyage ; eetle, A. А. 2273 (11a) Bell, F. s.n. (2c JR. 571 ; 1410 (2b); 2206, 2428, 2446, Benson, L 2540 (2a). Bereman, S. Б; „737 (11а); 738 (3d). erg, №. К. (4); 2071 (11a); 4758 (34); 4503 (112). Berghius, D Berlandier, J. L. 4 (17); 902 (29a); 1133 (17); ar ony, 120$ (15). Bernoulli, G. 297 (17). Bernoulli, e & Cario, R. ‚1950 (255), B s. n., 368 (25 d s. n. (3d) ; s. n. (11a). s. n. P s. n. (7) s. n. (11a). nu F [t] ST Bt ре соп s. n. (11b). Biltmore Herb. E (3d); 3660 (11a). Bird, J. В. s.n. (За). J, Bis, C. H. pos (1). Bissell, C. H. ` ея бей; Bitner, <5 re Blaisdell, F. 7 (3a). Blake, S. F. p pc 7558 (25a); 0416 22633 (1). Blanchard, B, ғ” (1); т (2c). Blanchard, W.H. a7 k h Bolander, H & Heller, —. s.n. (3b) Bolley, Н. L. 10 (2a). Bolt, B. 420 (3b); 783a (2a). Boner, L. & Weldert, V. 234 (2a). Bonck, L. А. s.n. (2a Booth, J. A. & 77 А, 321 (5). Booth, W. E. 1534 ( Borell, A. E. s.n. сек Botteri, M n. v . p. . (25Ь); 705 (292); mM 797 (25b); 1585 (25а); 2045 ( [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bourgeau, E. (22); (8); (14); боё (224); doo (29b); Bor (290%; 1064 (15); 1577 (25a); 2044 (27); 2045, 2047 Ma 3203 (29b). Boyce, T. Е. 114 b 5. ; sm, (29b); 102 Gd); 274 (2b); yos 790 (3d); 792, 818 (Па); 871 (4). raunton, —. 5. 7, Breitung, A. J. 1, (25b); 5897 (17); 13454 (25b); 16276 (28); 16667, 19300 (28); 22884 (25b). Brewer, W. H. ; Bridges, Thomas. 107, І Britton, E. G. & Brown, M. 8240 2. 5880 (25Ь). Britton, N.L.&B 7 (25 Вгіссоп, М. L & Shafer, J. А. 2116(25а). pg N. L. & Wilson Р. 526r (25b). ooks, A. Н, & Prindle, L. М. s. ». (3a). P H.E (3a). Brown, C. S. s.n. pd Brown, Н. E. 5. n., 448 Gb); 4 (2Ь). Brown, L. M 202, 203, 204 (2b). Brown, R. H. 2. Brown, S. 5 (2a); 426 (8); 230, (2а); той, ris (8); 1250 (8); 1450, 1544 4 Вгисе, С p 1162 (3b); 2264 (11a). Brunet, —. s.n. (2c). Bryant, P. ‚ (3a). Buffum, B. с E (7); 386 (3d); 387 d). um (11a); 358 o 1b). ‚озул. (7); s. n. (11b). 5 4 (11b). » S. n. (2а); s. n. (8) ; s. n. (11a). Burke, М. Н. 13 (7). Burnham, S. Н. s.n. (2c). Burtt-Davy, J. 3182 (3b). M Buswell, W 0539 (25a). Butler, B. T. 262, 260, 381, 604 (2a). Butler, С. D. 1655 (2a). Butters, F. K. & Holway, E. W. D. 203, 272 (2a). ER 1951] MEYER—VALERIANA Butters, F. K., Holway, E. W. D. & Rosen- dahl, C. О. 588 (2а Butters, F. К. & Rosendahl, СОО, 307 2a). Byce, M. L. 600 ain, S. A. 20 (3d); 103 (7 Calderón, 5. 1061, 1062, 1062a (28) ; 1017 (25b). Camp, S. H. & D. R. s.n. (2c). amp, W. H. 2623 (15). (3a i Canby, W. M. 2857 (11b). 288, 320, О 357 (26а); & ER A. О. 6716 Carvell, W. T. S. s." (2c) Cary, M. 25 (11a) Casebeer, L. 5. n. (4). 1153 (3d); Casteter, E. F 1804 (11a); 2150 Chai G. x 2678 (2c). Chambers, M. B. 35 (3a). Chamisso, L. A » d (3а). Chandler, B. Е. s. n, (2c); s.m. (11b). Chandler, H. P. 7583 (2a). Chandler, H. P. & Babcock, E. B. 1063 G Chaney, В. W. 90 (3a). Chapman, A. W. 2 п. E з.п. (25а). Chapman, J. W. 5 (3a Chase, V. H. 21. 8, 0392 (11b). Chickering, J. W. 5. м. (5). Choussy, F. 32 (28). IIIOÓ (11a). 1264 (2c); 5362 (25b); 5599 (28). че JR. s.m. (1); s.». Gd); s.» (11а). Clark, 3 527 Clark, M. КУЛ, E Clark, O. M. a); 4625 4576 (n (34); 5505 7: Ж aio (8); 11760 (4). » (112). Clausen, E. R. s.m. (1). Clausen, J. 1134 (3 b). Clausen, R. T., Trapido, H. & Wilson, W. c). б oi es Clement, Fr. бот (25a). IN NORTH AMERICA 2487 Clemens, J. 5.п.(2а); s. n.(7) 5 s. n. (11a). Clements, F. E. s.7.(112); 25 105(34). Clements, F. E. & E. S. б (11а); 241 (4); 328 (3d). Clinton, С. №. s.n. (2c). Clokey, І. W. 1847 (17); 2882, 3581 (34); 7342, 7734 (3c). Clokey, I. W. & Bean, К. 7341 (3c). Coats, R. R. Z (3a). Cockerell, T. D. A. s.n. (3d); s. м. (4). Cody, W. J. & Calder, J A. 504 (1). Coghill, G. E. 117 (1 Cole, E. J. s.". (2c). ollett, — С „ (3а); Соот, К. E. . (11a); Б, (4); бот (11a); 1056 (7); 1324 (11а Conard, Н. S. s.n. (7); 206 Bs. 1556 8 s. т. (2b); 2410 (3b). (8). Constance, L. F. G., Staats, W. wo L., Clarke Vleet, G. 1163 (2a). Ob is L. & Clements, Н. F. 1735 2a Constance, L. & Jacobs, C. D. 1374 (2a). QUNM L. & Rollins, А. С. 2867, 2997 (2b). Cooley, —. 5.7. (1). Cooper, б. Р. 65 (17). per, W. S. s. n. (8); s. n. (11а); 109 Eod. 138 (2a); 395, 409 (3d); 411 (11 Сем, Е.В. 481, 3841 on Copeland, H. Е. s.n., 3841 (3b). Cottam, W. P. 6075 (4). Cottam, Y. P. & Biddulph, О. 1306 (34); 2082 (7); 3188 (3d); 3506, 3061 Р. & Hutchings, —. НЕ: i a). 2219(7). Coulter, J. M. s." ; 68 (7) Coville, F. V. 576 (112); 762 (2a); 1104 (4); 1361 (2а); 1545 (11а). Coville, F. V. & Applegate, E. I. T123(793 340 (3b); 371 ( ville, F. V. & Funston, F. 1486 (3b). Coville, F. V. & Kearney, Т.Н. 264(2b); 764, 1036, 1159 (2a); 1805, 1900, 2020, 2204 (3а). Coville, F. V. & Leiberg, J. B. 397 (2a). 165 (3b); 488 Cowen, J. Н. s.n. (7, 11a); 692 (7); IIIS (11a); phe (3d); 2068, 2069, 2082 (11a). Cowles, H. C. І 0 302 (7); 282 (11а); 282 (7); 457, 603, 1114 (Па); 1117 (3d); 2066, 2067, 2070 (Па); 2072- -2080 5479. Crane, В. K. s.n. p 208 (11a). Cronquist, A. (3d); 1333 (7); 1428 зен 1681 ee 1705 (11a); 1876 (34); 2300 (7); 2512 (3d) ; 2534 (11a) ; 2576, 2772 (2a Cronquist, А. & Davis, R. ). 2007 (11a). Cros, W. 40 (3d). Crozier, A. А. s. n. (1) ‚Е. 3001 (3b). Cufodontis, б. 305, 369 (25b) Culbertson, —. 4376 (3b Curry, Lo &tw* (3). Curtis, C. С. з. м. (7, 11а) urtiss, — 26). Curtis, A. H иг н. sack, — IQ Cusick, W. C. E : (2 а); 2131 (3d); II 3813 (2а); 4060, ists (2). Dandelin, J. E. 11 e$, 2543 (4); s. n. (11а). -—— R. А. Gould. F. W., Phillips, W. Pultz, L. M. IIIQ(11a); 1452(29b). с M. E. 443, 886 (25b); 1023 4). b). Davis, R. (7, 11a); 1004, 1183 Gd); 2880 (2а); 3288 (11a). 5. 7. Dawson, —. s.n. (8); 1261 (5). Dayton, W. A. 3071, 05130 (24). Deam, C. С. 1017, 43180, 53807 (5). Debeaux, О. s.n. (5). Decaisne, D. D. s.n. (11b). Degener, О. 16350, 16462 (11a). Demaree, D 6 Й Detling, L. Е. 2878, 4087 (2b). Dewart, Е. W. s.n. (7). Dewey, L. Н. s.n. (4) Dick, W. M. s.m. (5). Diehl, I. E. ж. (2a, 3d, 7, 11а). Diguet, L. s. n. (17, 22a, 25b, 29b). Dillon, L. A. & M. 73 la). Dixon, J. 57 (3a). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Dobbs, R. J. 20 (11b). Dobbs, С. $. & Robbins, ы W. 5844(34). Dodge, C. К. £9 (1, Dodge, C.W.& Thomas, W. с. 5312(27). Donnell-Smith, J. s.n. (5, 25a); 1466 Dore, W. G. & Brietung, A. Ј. 12173(8). Doten, S. В. 160 (3b). Doubleday, —. s. ». (11b Douglas, D. s.n. (2b, 11a). Drake, — & Dickson, —. s.n. (2b) Drummond, T. s.n. (2a, 5, 8). Dudley, M. G. s.s». (2a, 11b). Duges, A. s.n. (21, 29b, 31). Dutilly, A. 4 (3a). ко A. & Lepage, E. 14303, 15657 белі A., Lepage E. & O'Neill, à 20147, 20423 (3a); 20530 (2a); 20727 (3a); 21215 (2a). Dutton, D. L. s. s. (1). Eames, A. J. & MecDaniels, L. H. 5056 1 Eames, E. H. & КЕРШ С.С. 5650 (1). Earle, F. S. 530 11561 (8); 15774 (2a). Eastwood, A. s.n. (3b, 8); 315, 407 к 455 Gb); 488 iuh 807 (2a); о Е. & Howell, J. Т. 1334, 4868 (2b) ; 8378, 14527 (3b) Faton, D. C. Wr е (114). Eaton, D. v. s.n. (3а); 4O p. p. (2a); 1275, 1276 (2c); 5884 (7); 5885, $887 (ties, 7074 (3b) ; 7906 (7); 0306 E 02 10762 (11а); Ha «Тыр 200 (2а); 14527, 18701, 18806, Pa: 1008, 20049 (4 Ehlers, кы P 3518, 4747 (2c). Ehrenberg, C. 173 (29c); 406 (14); ide, P. Ekman, E. L 1, 700 (25а); i 1403 . 621 (26); 6290 (262); 8082, 9209 (25a); 10513 (25b) ; 11643 (26a); ieri е эч, 13580 (26а); 13701 (26); 18505 (25a); Ekstam, О. 5.” ); Elliott, —. ‚ (За). Ellis, C. C. 2 w: 313, 320 (11a). 1951] MEYER— VALERIANA Elmer, A. D. Е. s. n., 440 (2a); 822(11а); 2792 (2a Elrod, M Ж. s.n. 21); 76 (11a). Elwood, W. М. 25 (2а). Emmons, С. Т. Pi п. (2а) Enander, S. |. s.n. (2a). Endrés, —. 79 (25b). Engelma һи ASG; 114) English, c. 50 (2a). Epling, C. 6204, 6205, 6206 (2a); 0105 (11a); 9516, 9553, 9757 (2a). Erlanson, E. W. 104 (2c). Ervendberg, L.D. 357 (25a). Evans, H. M. s.m. (3b). Evans, W. Н. s.m. (5); ISI (2a); 631 (32). Evermann, B. W. s.m. ; 84, IOI (Зах (3а); 263, 481, 543 с РА (11а). Ewan, J. А. 1721 а). Eyerdam, W. J. s.n., 317, 1262 (2а); 1730, 1504 Ca 3521, 5953 (2а). Fendler, А Ferguson, M. s 2b). Fernald, M. L. s.n. (8); 2335 (2c). Fernald, M. L. & Collins, ] É 240 (8). Fernald, M. L. & Pease, A. S. 3541 (2c). 10864 (8). . 4064, Ferris, R. S. бото (17). Ferris, R. S. & rn L. 776 (2a); 858 (11a); 10604 (3b). Ferris, R. S. & Lorraine, L. 10604 (3b). Fiker, C. B. 351 (2а); 751 ques 1053 (2a); 1805 2. 2417 (8) Fink, B. s.s. (11b). Fisher, G. L. 0б, (3b); 3727 (29601 Fisher, H. L. pla is itch, А. s. v. [as 2c). Fletcher, d 8141, 952 (2a). Flett, J. B. 5.п. (2a, 2b, 3а); 1203 (2a). Piodnsn, T. H. 800 A Sor (8); 802, 755 з.п. (7, 11а, 11b); 30 la). Foster, А. С. 287 (2b Foster, R. C. & Arnold, n F. Dx. ss: Francis, M. E. 280 (4). n. (3b). 128 (25a). IN NORTH AMERICA 489 Franco, —. s.m. (17, 25a, 25b). Frank, J. С. И (Ж Fraser, D. 5.п. (2a). reeman, M: s %) Freiberg, С. №. s.ni (1) Fre с, ДУС; ж жаа) 504 (3b). Friesner, R. С. 9631, 18852.2 (5); 20607 11b ). Fróderstróm, Н. & Hultén, E. 686, 700, 1191 (25a); 1298 (15). Fuertes, P. 541 (25a). Funston, F. 87 (2а); 98 (3a). Fyfe, G. & Sha p 115». (2c). Galeotti, H. (25а); 693 (15); 207 (17); 2076 (27); 2547 244 xU И 2549 (29c); 2550 (11 D (14); 2554 (17); 7 (25a); 2560 (24); 2561 (25b); е (28); 2683 ҚА 2706 (25а). СагЬег, А $9 (57236). Garcia, Р Н 317 dE 310, 464 (23); 933 ( 2.1 R. 655 (11b). Garnier, A. 1074 (25b). Garrett, 602, 1308 (34); 1691, 1087 (11а); 2143, 2483, 2011, 3638 (7); 6365, 7914 (11а); 8585 (7) SR (79. 1666.3 (11b); 15431 (2c); (1). Gates, R. R. & Mellenby, К. s.n. (3a). Gentry, Н. 5. 532M, 507M (20); 638MA (29b); 1757, 1976, 2105 (20); 2401 (28); 2566 (29b); 2739 (11a); 5318 (20); 6261 07); 20 20). Gentle, P. H. 2 (298). Geyer, C. A. i P 335, 337 (11а): Gierisch, R. К. Ghiesbreght, A. E i^ Si; 270 (224); . (22b); 623 p.p. Gillett, J. M. fos (1). ars J. M. & Boivin, B. 6024, 6034 Gi : leason, H. A. nke, S. 1. 6121 (1). ,D. R. 758 (11a). Godman, A. 220, 407, 681 (2a). Sn. 59 490 Golder, F. А. 87 (3a). Goldman, E. A. 187 (17); 231 (20). Goll, С.Р. 00, 244 (25a). Goodding, L. N. s.n. (9); 71 (7); тота (5): боо (11a); бот 4), 1148 (3с); 1140 (3d); 1329 Ди 1338, 1444, 1560 (7); 1760 (3d). Goode, R. 745 (8). Goodman, G. J. 128 (7); 193, 448 (11a); 535 (3c); 554 AUAM & Hitchcock, C. 83 (4): 1300 (7); 1307 (114); 1436 (2); 1680 (3c). Goodman, С. J. & Payson, L. В. 535 (3c). (2а). 2а Goodwin, R. Н. ie Pp. ч 826(11a). Gorman, M. W (3а); 03, 400, бог (2а); 1050 (24); 17838 (24); 5723 (2b). ouin, —. s.n. (17, 25a) Gould, F. 2610 (29b). Gould, F. W. & Haskell, H. S. 3337 (29b). Gould, F. W. & Hudson, R. K. 3785(11a). Gould, F. W. & Robinson, M. E. 5002 (11a). egeo —. s.n. (22a). Graham 8142, 8187 (7); 8200(11a); Por oun adi (7); 0327, 0374, 0541 (11a) ; 0604 (7). е С. B. 6076 (3b). nt, J. M. s.m. (a): 177 (2b). Grant. V. 834 (25b). Grassl, C. O. 4853 (2c). Greenman, T M. A3 (25a). & M. T. 4688 (11a). a). п. (2 Griffiths, D. 5358 (11a); 7216 (29b). 158 Griffiths, D. & Hunter, В. 159 (11a). Griggs, R. F. s.n. (3a Griscom, L., Metcalf, F. Р., Wright, А. Н. & А. (2с). Griswold, Н. А. s. n. (6) Groh, H. s (1, 8). Grover, E. L. s. м. (6) Gustafson, T. s.n E ahn, L. 5 . (25b, 29а, 29b). Hahn, W. L 5 (3a) Haight, C. з. м. (1). Hale, T. J. s.n. (11b) Hales, B. J. s. n. (8). Haley, G. s.n. (3a). Hall, С. Н. s. ж. (11a). Hall E. жт. (5, 11b). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN ire a М. s.n. (3d); 11469 (3c); 11586 Hall. H. M. & Babcock, H. G. 5665 “28 ars Н. М. & Chandler, H. P. 412, 3b). i^ E. & Harbour, J. Р. 123 (11a); 237 369 (29b). бу. pu Hands. —. (7); 1422 Hanscom, 5. i f. (11а). Hansen, G. 725 (3b). Hanson, s.n., A236 (11a); 623 (3d); A048 (11а). Наретап, Н. s.s. Hardin, Е. 407 (2a) Harford, G. W. & Dunn, —. 259 (2 Harrington, С. L. 34, 46 (3a Harrington, Н. D. 7599, po (7); 3654 (11a). Harrington, Н. D. & Smith, E. C. 1100 (11a); 1250 (34); 1276 Neel Harrington, M ^ 9 ( 877 b 7); 21455 (3d); 22333 (3с); C24543 (113) ; 24058 (7); 26488 (11a); C27707 (7); id 28275, 20582 (11a); 29587 Harrison, B. Е. 103 (11a); 4 (23% 7359, 7360, 7731 (11а); 7794, 8734 (7); 9358 (3c). Harrison, B. F. & Nisson, A. 8837 (11a); ам ы Жы (7) Harrison, G. J. 4860 (11a); 7872, 7882 4 Hartman, C. V. 136, 304, 736 (29b). Hartweg, T. 5. м. (14, 22a); 150 (21); 19 an jor (22a); 302 (51); 303 Hayes, e s. n. (29b). , H. E. 1814, 1087 (7); 2725 ©; ху. н. 31, a (3a). Heidenreich, V. T. 140 (2a). Heller, A. А. s.n. (5); 7156 (3b); 0118, ed (7); 10540 (3c); тоого (3b); 11020 (3c); 12070 (3b); 12632 (2a); 12928 (3b); 12956 (2a); 13043 (3b); 1951] MEYER— VALERIANA TA 14773 д ller , б. е (Па); 3353 (7); ES Оа); 3937 (2 BE Heller, A. A. & Small, J. К. s.n. (5) e s. n. ere 187 (25a); 366, 404 (22а); 438 (17). 2 а): mt (2b). А . (2a, 11а); бі (2b); 62 (2a); 426, 635 (2а); 636 (2b); 953 (2a); 2658 (7); 34084 9 b (11а); 3400 (3c); 5523 (7); 5525 (11а); 5688 (2а); 6058 (2b); gett (3b). Hendrix, T. M. 3 (b). Henry, J. A. s. E (as. Hermann, F. J 3 (110): АУЕ UJ35 5557 (34); 5857 Хы); 6075 (5); 8303 (2с); 8058 Sl 9455 (2 13 . 201, 532, 546 m (1725 103 8: 271 n 706 (25b). Heyde, E. T. & Lux, E. 2924, 3968 (28); 3069 (17); 3970 ТОА ; 2488 (25b). HEIE Hinckley, L. p? s. n. (4, 11а); 721 (4). 4 7 (15); 505 (11с); 608 (21); 628 aie, 804, 081, 1007 (22a); 1115 (28); 1319 (11c); 1342 (16); 1368 (22a); 1417 (30); 1513 (29b); 1568 (27); 1663 (30); 1677 (17); 1793 (29b); 2180, 2416, 2630 (28); 2732 (15); 2760 (29b); 2849 (17); 2889 (29b); 3240 (25b); 2104 4395 (22а); ; 4636 (29b); 505 (25b); 8081 (15); 9 ae 9468 (17) ; 9604 (27); 70846 (17); (27); 10309, 10448, 10474 (22а); Б (31); тоўот (17); 10702 (27); 10946 (28); 11167, 11335 (17); 11008 (22а); 12189 (17); 12518, 12808, 13481 (29b); 14386, 14443 (22а); 14664 (17); 14719 (25b) ; 14733 (27); 15004 (22a); 15031 (29b); 15040, 15360, 15460, 15550 (17). Hitchcock, A. E. 838, 1301, 1316, 1413 (ic) i Hitchcock, A. S. s.n. (3b, 11а); 11785, 12100 (2a). Hitchcock, C. L. 1590, 1748 (8); 1807, 1971 (2а); 15771 (11а); 15038 (8); IN NORTH AMERICA 491 (8); (8). Hitchcock, C. artin, J. 3536, 3546 (2b); bos b 5662 E. 7418 (8); 7529 (2a). Hitchcock, C. L. & Muhlick, C. V. $543 (2а); 8766, 8835 (11a); 0133, 9256 a 9405 (oer 9540 (8); 9923, 10282, ; 12047 ( (2a); 12240 (8); 12771 (2a); 12032 (8); 13135 (2a); 13600, 13036, 14002 (11а); 14011 (2a); 14061 (11a); 14158 (8); 14286, 14484, PTA 14032, 15021 (2а); 15200 (7); 15211 (11a). Hitchcock, C. L., Rethke, R. У., Raads- hoven, R. 3780 (3c); 4134, 4592 (119). Hitchcock, C. L. & Stanford, L. R. 6968 (25a); 7243 (22a) Hjalmarson, J. A. (17). obbs, A 5": Holdridge, L. К 1277 (26); 1365 (25a). Hollis, M. 5 Hollister, М. 9, ae E (2 а). Holm, К. Iltis, Н. Н. 466, 468, 540 (10); 574 (24); 501 (10). olm, T. s.m. em 463 (3d). 4 (11а); 1267 (3c); 1332 (7); 3744 f 3 Holmgren, A. H. & Hansen, S. 3475 (7). Holtz, Е. L. s.n. ( B. Holzinger, J. M. s.n. (11b). Hooker, J. D. & Gray, A. s.n. (3b, 3d, 11а). Hooper, C. L. s. н. (38). Hopkins, L. A (688 Hoover, R. F. 7461r (3b). Horne, W s. n. (32). Horner, 5.7 15): Ногпег, К. М. 244 (2а). Hough, W. s.n. (4). House, D. s.n". (2c); 837 (5); Ór51 (2€); 28755 (1). Howard, E. S. & R. А. 8526 (25a). Howden, 9 R. 63, 64 Howell, J. T. s.". 11757 (2b); 15017, 15162 (2 2) . 5.% (2а; 2D, 35, 8; 11а); 425, 833 (7); 1631 (S Hoysradt, L. H. 492 Hoyt, К. W. n. (4) Hughes, E. L. 23 (3d) Hughes, J. A. r300a (8) a . (3а); 700 (25а); 7924, 247, 8248, 8384 (24). Humboldt, А. & Bonpland, А. 14, 29b); 2003, 3330 (17 — F. W. 3845, 7045 (2a) ; 14868 (25b Hunt, R. & aver A. Hunter, J. А. s. sor ret W. "46 (24); 178, 333, 373, 0: (116; 165 (11a). 682 Jack, J ^ s. n. (7) Jacquemont, —. s.n он ames, F 5 Johnson, C. F. s.n. (11b ёл. (1, 299, 153 (11a); 1230 (2a); pork (2c Johnston, _ x P 77r. 771b (11а); 826 (4); нА, 4 г : Небо, —, aon Johnston, Н. W. Pine L. 945 (17); 254 ы); 053 (27); 0^4 (17); 1000a К 4). ‚ (5). ne (2а); 715, 8200, 10282 (2а Jones, G. М. & Е.Е. 14738 (11a). 058x (11a) 25052 (2 b); sr 27842, 27846 (29b); E Jones, W. W. s.m. X^ Jony, P.L. ;. (27 Jordal, L. H p^ 2454 (3a). Jérgensen, B J 461 (УЬ). iday, C. s. du a). Jurgensen, с” 500 (15); 870 (22b). Karwinsky, —. s.n. Kauffman, C. G., Ehlers, |. H. et al. 2260 (20). Kaye, D. 103 (3a). Kearney, Т. Н. :.п. (5). Kearney К. Н. 10070, ‚ Т. Н. & Peebles 10551 (29b); 12187, 1236; (11a). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Keeler, Н. D. s.n. (25a). шотеш АА A. 4562 (10); 5830 (15); 7357 (25a). Kellogg, A. & Harford, W. С. W. 358 (2b). Kelsey, F. D. s. s. (11a) emp, Е. к. (2a) Kenna, D. s.n irae Kennedy, Kennedy, R. B. ;L. A. ғи. (15, 17, 25а); A358 un A359 (16); А416 (11c); A620 5a). Kerber E. 200 (25a). Killip, E. P. 3777: (2c); 36411 (3d); 36421, 36424 (11a). Kincaid, T. з. м. (2b, 3a) Kirkby, W. s.n. (3a) King, А. : : Kirkwood, J. E. (2a); 6, 1300 (8) 1301 (2a); 418 (8); 1433 (112); 1786, 1873, 2008 (2a). Kittredge, J. s.n. necntel, —. 540 fS Knight, О. W. 5160 (2c). Knipe, S. W. 5. и. (6). Knipe, S. W. & Garber, A. P. s.n. (5). Knoblock, I. 5284 (29b). Kofoid, C. A. s.n. (5). Krautter, L. (3b). Kuntze, O. 4490258), 2248(25Ь); 23772 (22а). Kunze, К. E. s.n. (11а) Kusche, J. А. s. s. (3a) ake, — ull, —. 540 (2a) Lakela, O. 2560 amb, F. H. 1339, 1330a (2a). Lamm, M. 16063 (8 " Lane, С. Н. s. ж, (11a). Lankester, C. H. s.n. (10); 265 (25b); 60, (17). Langlassé, E. 335 (17); 408 (28); 57I (17); 1043 (22a). Lantis, М. s.m. (3a). Lapham, I. A. s.n. (11b). Larsen, E. 6084 (7). Laskey, C. F. s.n. (2a). Laughlin, E. E. 067 (5 awes, 814 (2a) Lay, G. T. s.m. е Lay, С. T. & Collie, А. s. м. (3a). MEYER— VALERIANA yden, R. L. 176 (3a). Leavenworth, M. C. s.n. (25a). Leavenworth, W. p^ 725 (27). C. & Hoogstraal, Leavenworth, H. 1254 (17) ее. М.т. (9) Lehmann, Е. С. 1547 (24). Leiberg, J. В. 1010 (7); 1135, 1240, P (2a); 2100 (11a); 112 86)5 25 (2а) 525307 (3b); 5бтг (3d); о. 5750 (112): Шешу J. s. 7». (7, 11а). Lemmon, J. С. 5. т. (3b, 11a, 29b); 17 4); 2713 (29b); 2727 (11a); pA ). 25Ь). 12668, 12026 (25Ь). Leon, Fr., Селе Ег. & Коса, М. 0016 (253): Leon, J. 378 (25b). Leonard, E. C. 3686, 4408 (25a); 4490а (26); 4622, 4662, 7824, 7015, $004, 8992, 12240, 13566, 15800 Leonard, F. E. %.п., 20 (7); x (3c). Lesquereux, L. s. (5) LeSueur, H. Sra (29b); I077 (11a) ; 1253 (29b); 1418 (11a). . s. 2.(3d, 11a); 65(34); Liebmann, F. M. 10820, 10821, 10822, 10823, 10825 (15); 10826, 10827 (25b); 10831, 10833, 10838 (29b); 10830, 10840, 10841, TROU: 10847 (25b); 10854 (17); 10855, 10856, 10857 (27). Linden, |. 5.т. (29b); 307 (25a) Lindsay, Р. К. & Mulligan, С. A. 135(1). Lindstrom, А. Н. 5. ». (3a Livingston, R. В. 1205а (3d) Lix, F255 Lloyd, —. 394 (29b) Lloyd, C. С. s.m». (5). Lloyd, Е. E. s- и. (2a, 2b) Long, B. 55394 (5). AS (22)* Longfield, C. & Blezard, А. s.., 219 (2a). Lo off, E SHX HB: 52, Wis 1262 (3a). a). Lyonnet, E. I (29b); 332 (22a); 357 117). 373 ies Sie сах 731 (21); 042 (11c); теве Е. m ого, К. 1700 (22b); 1839, 1906 Т7 1927 (15). 493 McCabe, T. T. 44, 444 (2а); 325, 336, 418, 569 (2a); 1070, 1190 (8); 1286, 22544, н 21 (2а); 2383 (8); 2504 (2b); 2744, 2042, 3008, 4740, 4900, 4042 (24); 4085, 4986 (8); 5403 (2а); 6160 (8); 6180 (2a); 6343 (8); 6351, 6505 (2a); 6625, 7041 (8); 7122 (2b); 7552(8); 4085, 8146, 8343, 8416, 8610, 8900, 8021 (2а). 2. у. С. 221% Ra). Cosh, A. J. 5.3. (11%): масо, А. J. & Greene, С. С. IN NORTH AMERICA s. t. (3d, 1a). MeCoilough Е. . $118). McCree 76 : 88 (5). M А.С. 235 (11a); 405 (7); 717 11 McK с. Le 52%: (Gar үле едін S. D. s.n. (2а); 4673, 4743 (3 Жош, K 10685 (34). McLellan, J. ХУ. I (1). McLevegan, —. (3а): McMillen, Е. s.n. (2b). McVaugh, R. 3116 (2c); 5818, 5903 (3d) ; 7421 (9); 8054, 8003 (4); 0400 (2c); 0855 (13); 0053, 10063 (15); 10150 b); 10028, 10980, 10003 (2c). 680 (2%); 032 (3с). Macbride, J. F. & Payson, E. B. 2055(11a); 3154 (3c); 3258, 3404 (8); 3474 (2а); 3783 (3c MacDaniels, L. H. 204 (17); 758 (29b). MacDougal, D. T. 5.п.(2а); 21, 125(4); 150 (8); 231 (3d); He "618 (2а). MacGregor, R. С. s.n. Mackenzie, К. К. 151 (7); 288 (11a); 334 (3d). MacKenzie, K. K. & Griscom, L. 10440, IIOI MacLean, J. s.n. (3a) Macleod, B. C. s м. (8) Macoun, J (2a, 2b, ); 01 (3a); 346 (2a); 650 (2b); 660 (8); 814%, 908, 033 (2a); 952 (8); 053, 10817 (2а); 20088 (8); 42708 (2a); 59973 (8); 64599, 64900 (2a); 64001 (в); 64002 (2b); 65380, 65387 (2a) 65383 (2a, 8); 65380 (22) ; 72796, 7 (2b); 72700 (2a); 79447 (11a); (2b); 91474 (3a); 91475 (2b); 04012 a). Maguire, B. 3790 (7); 13803, 13804, 494 14099 (3c); 16017 (11a); 16150 (3c); 16979 (11a); 16084 (7); 17441 (3c); 17371, 18425, 18835, 19031, 10603, 10004, 20028, 21703, 21704, 21705, 21706 (11a); 21706, 21700, 21710, 217II 7 ( Maguire, B. & R. R. & Hobson, D. A. 14014 ved 14020 (7); 14099 (3c); la). Maguire, В, & Holmgren, A. ; 22401 (7); 25608 ES шы (3c); 25704 (3c); 25076 (3c); 26457 (7); 26015 (2a). Maguire, B., R. & C. B. 15613, a ef Maguire, B. & Muenscher, W. (2a). "pens B. & Redd, J.D. 2132(7); 2136 (11 Maguire, A эк B. L. & Moeller, Т. (1). Marie-Louis, P., Fabius et Adonis, Raymond, ‚ & Paquin, J. 342 35 (8). . n. (2b). LL ‚ (3a); 3323 (3b); 3804 и 3007 (28) 55 pi окы Mason » (2a). Masters, —, . (2 ). . s.n. (29b); S-62 (25b); S-117 (15); 562, 0593 (25b); 0608, 0804 (29b); 0020, 1071, 1177 (25b); 1349 (15); 1812 (18); 2802 (15); 4106 (17); 4133 (29b); 4204, 4530, 4602 (18); 4638 (10); 4691 (29c); 5209 (25b); 5960 (225); 5987 (18); 18307 (25 Maxon, W. 526, 632 21 (5); pe TM Maysilles, 1. H. Miri (31); 7145 (11c); 7238 (23); 7481 (31); 74814 (16); 7482a (22a). Mearns, s.n. (3d, 4); 426 (11b); 1042, 1043, 1170(11a) ; 1268 (7); 1641, 1851 a); 1852, 1857, 2004, 2095 3a Merrill, E. D. & Wilcox, E. N. 542 (11a); 2, 1242 (7). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Merriman, C. Н. s.n. EM e 2a). s. n. 2110, 121, IQI, 100(3а). Mertz, Н. М. SE Metcalf, F. P. pene (1); 8872 (2c). Metcalfe, O. В. 311, 1104 (11a); 1580 (3d). Mexia, Y. s.n. (15); 461, 602 (17); 876 (28); 1404 (29b); 1512 (17); 1845 (25b); 2055, 2087 (3а); 2601 (22а); 8978 (17); 9213 (25 Меуег, Е. С. 316 (112); 414, 710, 999, 2177 (9); 2201 (3d); 2 (11а); 2222, 2223 (2b); 2234, 2240 (2a); 2241, 2246 (2b); 2261 (2a); 2425, -— Kog чн Meyer, F. С. % 2 2611 (13); 2024, 2903 (29c); 3003 ( 15). ichaux, A. n. Middleton, J. T. s.n. (3b). iller, A. E. s.n. (3a). Miller, W. B. 79e, 100с, 221с (3a). Moffatt, W. S. (ON. „м. (17, 25b). Moldenke, Н. N. 7156 (3b); 7106 (7). Montes, M. H. & Salazar, A. E. 103 (17) Montgomery, F. H. 999 (1). M oore, A. H. 1343 (2c) Moore, E. J. s.n. (11a) Moore, F. L. 416 (2a). Moore, Н. E. 720 (17); 1086 (29b); 2023 (12); 3997 (22a); 4075 (29c); 4176, 4274 (12); 4302 (22a); 4408 (12); 4502 (29c); E A" ); 4675 (29с); 4678 (28); 4845 CA & St ,. 7 ds 3526 Moore, J. W. 845 (11a). Moore, J. W., Butters, F. K. & Jenkins, D. a (11b). J. W. & Phinney, B. О. 12457, — (11b). Moore, J. W. & Thatcher, E. P. 13901, 13927 (11b). oore, (8). Morales, J. R. 1126 (29b) rong, T. s.m. (5). (5). C. V. & Makrinius, E. 2549(25a). эзе p Es А. (2b). H. » 541 (2a); обт (8); pons Зоот 20; 8150 (8). Moyer, L. К. (114), Mueller, C. H. ye (29b). 1951] MEYER—VALERIANA Mueller, С. Н. & М. Т. 922 (29c) Muenscher, W. С. 8384, 8385, 11480 (2a); 12554 ae mre her, W. & Maguire, B. 2442 (11 Иа. У. С. & M. VW. 15398 (7). Muir, J. 150 (3а); 4370 Ob Mulford, І. з. 2. (2a, 3с); 235, $56 (11a). Müller, F. s.n. (28, 29c); 173 (17); 244 (27); 290, 343 (29a); 609 (15); 647 (27); 648 (15); 769 (25a); 869 bis (25b) ; 998, Де 1164 (27). . (11b). Munroe, — Munz, P. A Tips (22). BD —. s.n. (3a). urdock, J. 3031 (7); 4618 (3d). Miu В: 8 (7). urie, О. J. 1126 (2b) Murray, E. С. s. м. (3a) Myers, J n. (5). Nash, G. V. 762 (25a Nash, С. V. & Taylor, М. 1783 (25a) Neff, G. D 2 Nelson, A б (3 d); 262, 531 (11а ); 8716 (7) Nelson, A. & 545 5 (3с); 5508 (11a); 5621 (7); 5686 (8), 5005 (11а). Nelson, A. & К 716 (2b); 2330 (7); 2127 QA): ijs RM 3735, 4129 (3a). Nelson, A. & Macbride, J. F. 1272, 1405, 1946 (3c); 1056 (7). Nelson, E. 42 (11a); 168, 4863 (7). Nelson, E. W. III (25a); 590 (29b); 630 (24); 644(22b) ; 692, 1005, 1110 (22b); 1389 (29а); 1773 (29b); 2533 (15); 3146 (29b); 4530 (11c); 4574 (12); 4581 (11c); 4744 (12); 4760 (23); ho (4). Eon. N. E. к Hd L. 96 (7); (3c). Nicolas, Fr. s.n. a 29b). 2 Ogilvie, H. s.n. (8). IN NORTH AMERICA 495 Oleson, O. M. 335 334 (112); 340 (7). 45 (11a). Olsen, J. H. lson, —. s.n. (2a). O'Neill, Н. s.n. (25a) nstot, Т. 5.7. (2a О ). Orcutt, C. R. I (49% 3049 (224); 3871 (17); 4262, 4288 (29b). Ortega, J. С. 5016, 530020 8153 (17). Osgood, V. Н. s.n. Osterhout, С. Е. 5.7. Е. 34); 47 (11а); 48, 082 (7); 1846 (34) 3308 (7); 3371, 3392, 3901, 6794 (11а); 6796 (3d Ostheimer, A. J. (2a). Ons 1. С; E 8); 785 (2a); 2359 1701, 15900 (11а); 15001, 7). шры: О. s.n. (3d, 7, 11а); 10008 Pus M. m 1384 (112). 35 331 (5). Palmer, E. s.n. (3d, 4,7): 166 (4); 168 (11а); 189 (7); 1891, (11a); а on 244 (29c); 380 (11с); 388 : 416 (29с); 526 (4); 564 (20); РЯ (15); 609 (11c); 618 uw 754 (28); 1071 (29b); 1818 (20 Palmer, E. J. 1 5158 (5); 31399 (7) ; 31382 (11а); 37349 (7); 37476 (11а); 37480 Palmer, Z. W. (2a). dcc LH. & ат К. E. 3835 Parks, H. E. & S. T. 21026 (2a). Parker, K. F., Bor E. & Robbins, G. T. 6366 Parker, S. 455 B. Parks, H. E. & Tracy T. P. 0373 (2b). Parlin, J.C. я. (1). Parry, C. С. s. м. (3d, 11а); 36, 124 (3с); 260, 2854 (3d). Parry, C. C., Bigelow, J. M., Wright, C., & Schott, A. 441 (4); 442 (11a). I C. C. & Palmer, E. nu (29b); 2 (29c). m E. B. 553 I II40 (3d); 1152 1539 (3d). Payson, E. B. & Armstrong, G. M. 3450 (11а); 3306(7); 3339, 346936); 3503 (7); go o 3665 (11 Payson B . 1708, 1842 prn 1845 (2a); 1088 а). £750 (11a); 1977 (7); 496 (3с); 2143 (11a); 2786(7); 2797 (11a); 2850 (7); 2861 (11а); 3026 (3c); 3902 (7); 4102, 4130 (11a); 4154 (3d); 4905 (11а). Pearce, О. Е. s. n. (2c). Pearmis, C. T. 5. ә. (2a). Pearson, С. A. 362 (11а). Pease, A. S. 18066 (8 Pease, A. S. & Bean, R. C. 26347 (2c). Pease, A. S. & Long, B. 22632 (1). Pease, F. N " Peck, M. E. (25 5a); 2590 (7); 7416 (3b); 7417 s 7418 (11a); 7419 (2a); 7420 (7); 7421 (2a); 7322 (2a); 7423, 7424, 7425, 8605 (2b); 9606 (7); 07 2a); 10422, 13084 (11a); 14084 (2b); 14254 (3c); 15147 (11a); 15573 (3b); 15850 (2b); 15080 (2a); 15085, 15002, 15006 (7); 15008 (3c); 16221, 16333 (2b); 16668 (2a); 16674, 16928 ( 21775 (11a); 21994 (7); 22281 (2a); 23665, 23606 (8). I R. H. 4055 (4). R. H. & Harrison, G. J. 2970 (2 ЭБ). а К. H., Harrison, С. J. & Kearney, Т.Н. 3461 (29b). Peebles, R. H. & Loomis, H. G. I (4) 13596 Gd). 797 (3b). 104 (11a); 336, 337 (3d); Penard, E 492, 528 (11a). Pennell, F. W. 17115, 17000(29с); 18302 (31); 18337 (17); 18398, 18435, 18511 iod 18879 (29b); 19815, 20023 (17); 075 ey 20143 (17). ‚ (5). 1750 (1). TUM W. S. & Humphrey, R. R. 3100 (11a). Phillips, W. S. & T. К. 3100 (11a); 3269 3d). Pickering, C. & иар V. D. 488 (2a); 909 (11а). Pickett, F. L. 292 (11a); r369 (2a); 1512 (11a). Т Е. Pi ге J. F. G., & Dillon, СА (11а). т Bra, A T han (11b). Piper, C. h (2a, 2b, 11a); 1506 п); 4367 са; 6250 (2b). Piper - (За). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Piranian, G. (25. Pittier, H. 5188, 3288 (25b) ; 10448 (24) ; 10875, 11005 (17); 14107 (25b). Pittier, H. & Tonduz, A. wes (17). Polokowsky, H. 397 (25b) Pollard, C. L. s. n., ope, M. W. 3 (3a). Popowich, P. s.n. (8). Porsild, A. E. & R. T. 04, 241, 1719 (3a). e . l062 (3d); 4655, 4880, 5056 (7); 5059 (11a). Pra қ “(7 Preble, E. ^ к Mixer С. 646 (2a). Price, S. Prince іші, 60, 609 17: Pringle, C. С. s. n. (4); 1256 (29b); 1257 (11а); pity (29b); 1765, 2750 (20); 3234 (17); 3341 (27); 3612 (13); 3641 Q1); 4063 (15); 4521 (28); 4707 (22b); (29c); 5630, 563 6466 (22а); 5858 (21); 6047 (29c); 7085 (29b); 8454 (22a); 4 (14); dons (15); soo (12); 9365 Е 0466 ( ; 0836 c (25a); 6495 (29c); 6702 (23b); 6703bp (27); 67059b (29b), 7056 (4); 7864 (28); 9165 (17). (2a Rafinesque, С. я. (5). Ramalay, F. 6754, 9240 (11a). Ramaley, F., Dodds, G. S. & Robbins, W. W. 2992 (11а). 19511 MEYER— VALERIANA Ramaley, F. & Robbins, W. W. 3353 Gd); 3366, 3546 (11a); oo 5719(11а). Randall, M. s.n. (3b Rankin, W. M. s.m. (Ho. Raup, H. M. 3040, 3050 (8). H. bbe, E. C. 3675 (8); Raup, M. A 4042, 4179 (2a). Ream, Н. 9356 (3c). Redeker, H (4); Redfeld, J. Н. 2853 (3d) Reiche, K (17). Reiter, (C Oe (Ge Rendle, A. B. & Good, R. s.n. (2a). Renspie, —. 1575 (2a). Reynolds, M. C. 11557 (25a). Rhodes, M., Newhall, P. M. & Gia- comini, A. L. s.n. (3a). hodes, W. s.m. (1). Richards, G. H. s.n. (8) Richardson, E. s.n. (5). Richardson, s. t. (2a, 2b, 3a, 8). Ridgway, К. 731, 1598 (5). Riehl, N (11 1b). де, G. T. 739 (3d); 744 (11a); 2b). 7170 (112). Sow CUED) Robinson, J. Rodeck, H. E P» (54); бо (11а). & J. M. ооз, 934 (7); mbers, Б 28. 2523 20 um 2129 (31); 2a). dni (14). nter, : оѕе, Ј. 5% 7); 8331 (11e); ios (14); 9014 (12); 9260 (22a). Rose, J. N. & Hough, ху Кове, І. 5. 35337, ek (3b); 40763 (2a); 42164, 44113 (3b). Rosendahl, С. О. 275 (11b); 972, 1000, po 1744 (2b); 1990 (2a); 2103, 7396 (11b). UE x O. & Brand, C. J. Ros, М. ия. (25b). Rossbach, С. "B. & Hodgdon, A. R. 8532 s. n. (2b). AE Td. 44 48}: Rousseau, J. & Rouleau, E. Rowe, J. S. IOI Rowland, V. H. II7I (8) (8). T20 (11з). IN NORTH AMERICA 497 Rowlee, W. W. s.m. (1). Rory, H. W. к (13). unyon, R. 36 n. (2a, 4, 5); 148 (22a); 4). (3a). Russell, Р. С. & Souviron, M. T. 252 (17). Rust, H. J. 259 (7); m А жы Rutter, 2 57% (2); қ Rydberg Vd 751 (11a); 5, 7). % 752 О); 2704 (3с); 6096, 6870 (7). Rydberg, P. A. & Bessey, E. А. 4906 (7); 4999 (2а); $ооо (7); arm аш (8); 5005 (3c); 5006, 5007 (11 Rydberg, E Garrett, A. O. 8716 (7); 8954 (из); : 9724 (7). wth. eeland, F. K. 5574, Jo 6373 к 7656 (11a); 7814 (24); 8040 р: 8748 (2Ь); 8835 (2а); dl hn, , Courtney, W. D. & Parker, Cm b. (8); 3709 (2a). , W. W., Beals, R St. John, H. & Palmer, R 9604 (2а). St. John, H., Pickett, Е. L. & Warren, F. A. 3189 (11a). St. John, H. & Smith, C. P. 8324 (2a). St. John, О. s. &. (4); 7, 151 (11a). Safford, H. T. 393 (5). alazar, Е. 5.п. (17). Salle, —. s.n. (17, 27, 29b); д (17). Salvin, O. & Godman, F. D. з. м. (29b). Samples, J. s.n., 18, 149 (1 1b). Sandberg, J. Н. sn. (11а); 3Ó1 (11b). Sampson, W. & Pearson, С. A. 17 190 5.7. (2a, 11a); 10 (2a); Sandberg, J. H. & ‘Leiber E TBs sn (28, 6, 11а); 53 (11a); ssi (6); br go. Sandberg, J. H., MacDougal, D. Heller, A. A. 240 (112); 587 (2a). Sanson, М. В. s. м. (8); 909, 1082 (2a). 7 sn. (26 Saunders, —. s.n. (11b) Saunders, deA. s.n. (3a) ag s." ; 498 Scamman, E. 441, 674, 025 (3a); 1118 (2а); 1764, 1902 (3а); 2678 (2a); 3609, 3024, 4073 (За); 4500 (За); 4631, 4724 (3a "rho" A W. s.n. (15, 17); 107 (29b); red (15); 193 (11c); 104 (17); 105 2a). E S т. (11а). Schedin, L. M. & М. T. 667, 662 (3d); 663, 664, 4 - ON Pri (3d). Schiede, C. J. W Deppe, Р. s.m. (17, EE UT с» lla); 195(2a). Schipp, W. A. и а). Schmitt, —. ` Schmitt, W. Е. u (9). Schmitz, A. 5. 15); 299 ТА 300 (29b); д, (224); 659, 085 ( Schmoll, . (3d); 1 Ent Schnec $. n. Schneider, R R. A. 064 (11 а). Schreiber, B. ER 897, 1874 (3b). S » (1). (11 а). Scribner, F. L oe aus 68a (2a). eal, Seashore, M 1b). Seaton, H. 46 yos I71 (29b); 387 eJ. 1254 (17); 1335 (16); 2340 (24); 2438 (17); 3251 (29b); 3281 a 7% Sessé, M. & Mociño, 17, 25a, 29b, 30). Sessé, M., oe: а” рр р. Maldonado, —. 2a); 100 С" 303 (17); 306 (29b М. (25а); 308 (11c); “7”, е М (29а); 454 (15). Setchell, . (3a). on Ule 15, 5. n. (5). $ 468 “a lor 4475 (22а); 44209 (29b); pH. (22a); 44549 (17); 44677 (29с); 44084 (29а); 45226 (29b); к= (25а); 441412 (29b); 441540 Sharples, S. Р. 234, 235 (2a). Sharsmith, C. W. 320, 2056 (3b); 3585 (2а); 4041 (11a). [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Shaw, Е H. 346, 908, 933 (2a). Shaw, . (2 i^ Shear, c L. s (1; lla); 4076, 4090 (3d); 5036 (7); үй? (11а); 5187 3d). Sheldon, C. S. 1.9. (2c, 7, 118). Sheldon, E. P. s.n., 638, 7456 (11b); 8511 А 8573 (2a); пб (2b). Sherwood, W. 572, 622 (2b). Shimek, B. 5. м. (11b). Shinners, L 72 (5) Shockley E 2 хе hort, C. W. . (5, 11b); 50 (5). Shreve, ү юу, 5371 ашу; 5428 (4). Simonds, VN (7, 11а): eei — £8. (25а). Sinclair, —. . (20). Sintenis, P. inr 4027, 4146, 4232 (25a); 4663 (25b). keels, Н. С. s. s. (2c). Skinner, — s к Skutch, А. Е. О (15); 155 (25Ь); 485 (29b)1 570 (17): $86 (27 )1 390 (29b); 853, 1240 (10); 1408, 2177 (25b); ҚУАНА) mall, J. К. s.m. o ); 8082, 11503 (25а). Ty TX. & G. 4224 8 age Small, J. K. & Hele, A. A. Small, J. K., K. & DeWinkeles a B. 10060, va (25a). Smiley, F. J. 47a (3b). Smith, A. A270 (17); A206 (27); H372 (25b); A538 (27); A569, A604 (25b); 1242 t 1204 (17); P2099 (25b); P2545 (25a). Smith, C. L. 766, 892 (22b). 1647 (11a); 2113 (7); 2240 Smith, C. P. (3c). Smith, D. 16 (8); 103 (3a). Smith, E. C. s.n. (3d). Smith, Е.Н. r50 (7). Smith, б. s.m". (3d) Smith, . sm. (3b Smith, J. D. 1466 (25b) Smith, L. С. 787 (29a) Smith, R. Н. 145 (8) Snell, R. rors Snell, R. & Cronquist, A. 933 (3c). Snow, E. 1 Snow, F. H. s. x. (3d); 410, 432 (11a) Sones, С. D. s 2c). Sonne, C. F (3b). Spalding, Н. s.n. (7, 11a). piegelberg, C. Н. s.n. (11a) Spillman, W. J. . (5). ргарие, К. ы 1850115); 610, б11(8). 1951] MEYER— VALERIANA Spreadborough, W. 72795(8); 72798 (2a). . 4120 (Па); 4157 (4); 6740, 7536 (11a); 8205 (25b); $362 (25a); 11440 (5); 11988 (17); 13625, 13766, 13870 (11a); 14931, 15033, 15275, 15456, 15861, 16167, 16402, 16718, 17202, 17561, 17601, 17965, 17988, 18160 (2а); 20616, 84476, 85436 (15) ; 86588, 87063 (25b); 00143 (29b); 01781 (25a). Standley, P. C., Allen, P., Shank, P., Padilla, V. 806 (27); 2660 (25b). Stanford, L. R., Lauber, —, & Taylor, —. 2548 (15); 2587 (29c). Stanford, L. R., Retherford, K. L. & North- croft, D. шо (29c); 702 (15); 1008 (25a). Starz, E. 5.п. (2a, 11а) Steele, E. 5. s. м. (2с, 5 Steinmetz, F. H. 782 (2c) Stei ТГ $5. (22) 7). J. 580, 666 (3а). Steyermark, J. A. 20646 (17); 20685 wn et e < [6] з $ В rri IN NORTH AMERICA 499 50966 (18); 51023 (28); 51965 (29b); 52248 (15). Stokes, S. С. s.n. (3c, 7, 11a). Stonchouse, —. s.n. (2a). Stone, Е. M. s.n, 78 (4); 518 (11а); Stoney, С. U. s." а): Stork, Н. Е. 2807, 3040 (10). Stork Е. & С. C. 69 (2a). rc traub, 2 575 (2с) Studhalter, R. А. & Cox, М. S4137(11a). Sturgis, W. C. s.n. (4, 11a). Sudworth, G. B. s.n». (5). . 100 (2b); 270 (8); 329 (11а); 536 (7); 550 (8); 830 (2b); 8817 (2a); 10506 (2b); 13400 (11a); 13371 (2а); 13378, 13404 (Па); 13410, з.п. (5, 11b) ; 43 (11b). s. n. (2a, 7, 8, 11a). 1014, IOIb (3а). Ў (25а). Tatnall, К. R. 4515, 4721 (5). Taylor, E. 5. м. (3a). Terry, M. E. 1300 (17); 1367 (25b). Thatcher, E. P. 67 (7); 117 (11a). Thomas, W. D. 143 (3b). Thompson, J. W. 2056, 2450 (2b); 3263, 5080, 5393 (2а); 5926 (2b); 6286 (2a); (2b); 14780, 14864 (2a) ; 14923, 14044 (6). Thompson, J. W. & E. M. 10 (2b); 410 a). Thompson, R. M. s.n. (3a). Thornber, J. J. s. 7. (3d, Па); 4723, 5605 (4). Thornber, J. J., McDougal, — & Lloyd, —. s. n. (4 Thornton, C. W. 39 (3a). Thurber, С. $. ж. (2c). Tidestrom, I. (7); 325 (11a); 348 (7); 424 (11a); 2229, 2455 (7); 3032, 3688, 4119: (11a); 10409 (3b). Tilden, —. .n. (11b). 726 (17); 826 (28); 888 2930 (17); 4508 (25b); 500 10875 (17); ч (220), Topping, D. І. orrey, J. s.m. оче E 198 Torrey, J. & "A А. s.m. Toumey, J. W. s.m. снн 3% VM 351 (11a); 360 (3d Townsend, C. H. T. & Bases, C.M, 55. Tracy, E. s.n. (2b). Tracy, J. P. 5230 (2a); 6643, 7061 (2b); 57 (фи: 9373 (2b); 10505 (2a); s.n. (11b); 455 (25a). n . 5054 (3а); 5055 (2b); 5056 (2а); Er 5058, Trench, F. D. $. n. (2с). True, F. W. & Prentiss, D. W. I (3a). Tuerckheim, H. 250 (25b); ML (29b); 903, 1247 (25b); 1265 (29b); Tek 1570 (17); 2449 (25Ь); 3442 (26 3783 (25b); 2795 (25а). Turesson sn. (11a); 135 (2c); 62 s. 5. 2, 6072, 6127 (8). ‚ (3a). Tweedy, Е. s.n. pA lla); 26, 27 (11a); 28 (7); 312 (11a); 503 (7); 504, 510, 520 (11a); 2059 (8); 2060 (7); 2061, 3086(11a); 3087(8); 4552, 4553 (11a); 4554 (7); 4555 (3c); 5833 (11a). a ch, І. M. s.m. 22.15 04 Q2); bbi D por к (2а). Underw 2c). Underwood t % М "Selby. А, TX 362, 441 (11a). Valerio, J. 452 (25b); 575, 640, 744, 779 (17); 701 (29b); 936, 1492 (17); 1742 (25Ь); d^ batt 2842 (25 Valerio, M. (27); 501 s 1036 (25b). Vasey, С. К. s. м. (2a, 5, 11a, 11b); 202 (11а); 203 (4); 235 a 290 (2a). Victorin-Marie, F. main-Rolland F. 25084, 25002, 27135, 27150, 27179 (7); 49437 (2c); 40030 (1). Victorin-Marie, Е E ALS Rolland, F. & Dominique, F. 45080 (11a); 46205 (1). Victorin-Marie, F., Germain-Rolland, F. & Jacques, E. 44543, 44912, 44917 (8). Virler, —. s.n. (22a) ; 1809 (14); 1810 (15); 1811 (29a); 1813 (12 [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2 —. 185 (17). l, J. sm. (112). Vind F. K. 620 (11a); 1054, 1133 (2а). Wadmond, S. С. s.n., 3472, 4226 (11b); . 20, 2I (3a). s. n. (5) (11b). 1505, 1800, 2055 (3a); 2210 Шан 2208 (3b); 2301 (2a). Ward, ds 72, 53 ү b Ward, І. Е. s.s. (3d, lla); 275 (7); yi ps (Па): той. (7); 1060 (11a). 2680c (3b). n. VAr Herning, H. 3552 (3а). Washburne, 2 $. 4. (За). Watson, S. б (3c); 177 (11а); 488 p.p. (3c, 7); 4% (11 la). Waugh, F. A. . (8). Wawra, H. 50(15); пн 753(25а); 1030 (25b); 1211 (22a). Weaver, J. N. 729, 952 (22а). Webb, к. J. 1544 Weber, W. A. 2282 ds 2311 (8); 2011 (11а). Wehmeyer, L. E., Martin, F. N. & Love- land, H. F. 5 (111); 5005 (7); 5127, 7 Өс); 5149 (7); 5312 (3с). Weight, K. е ). Weinmann, —. (9895 Welch, W. Н. Ee til in Т.б. 0511(5). Went, Ж W. 54 (3a). Werne з.п. (5). Wetherill, M. TE (4). m И л, Wheeler, 1. С. 3755 (3b). Wheeler, (11b). White, С. 11 (25b) White, J. T n. (3 White, P. 153, 216 (25b) i (3а). 1а); 47 7 (2а); бо (11а); s (6); 162, 648, 646a, 774, 1227(2a); 9 (6). M E. 10003 (7). Whittaker, R. Н. 5541, 58100 (2a); 250 (3b). 1951] MEYER— VALERIANA IN NORTH AMERICA 501 Whymper, E. s.n. (2a). Wolf, C. B. 2657, 2863, 3031 (11a) ; 3056 Wiegand, К. M. & M. С. 2264, 2265 (4). p 5057 (2a). Wiegand, K. M. & Upton, С. В. 2381 Wolf, C. B. & Everett, P. C. eda (11a); 4345 (3d). Wolfe J ш vd: 774 (11a). Wiegand, K. M. & Wright, A. H. 7208 Woo . (2а). (2с Wood, 3 E s. n. (2c). И I. L. 4642 (2а); 8120 (3b). lon C. N. 213 (11a); 21Ó (3c); 324 Wilbur, К.І. & C. К. І 1479, 1608, ari ! D 1826, 1850 E ЕҢ Woods, C. N. & Tidestrom, I. 2682 (2a); Wilcox 2 2745 (11а). Ta va. Woodbury, А. М. 11 (4). э Н. 2327 (3d) Woodson, R. E., Allen, P. H. & Seibert, R. ilkinson, —. s.n. 25b); 7043 (24) Willett, G. 30 (2a). 999 ( И XU':lliams : ix 105 тт К. Е. & 2” гу, c W. 225 MESE ‘ (25b); 377 (24); 498 (25b) Williams, E. F. s.n. (1, 2c). Wandw 1 Williams, E. Е, Coll, ]. F. & Fernald, Wooten, В Е. О. оп. 27” A 11а); 320 . n с (11a a). Williams, J. F. C. & S. Е. s.n. (2c). Wooton, E. О. & Standley, P. C. 3423 Dr LO UN 337 XI Mer (11a). (7): 040 (11a); 1080, 1090 (7); Woronow, С. 2682 (29b). (3c) : 1608, 2164(7); 3087 (113); 3204 Worthley, I. |. 75 (7); 20, 30 (8). > Joa а d (11). Wright, C. s.m. (3a); 276 "25b; 277, Williams, 1. О. Molina, А. R. 10170, 278 (25а). 10422, 10553 а). Wright, W. С. 1534 (2а). Moe Ri S: " Tn 11а); 105 (7); Wynn, T 20 E 9 (11a); 72 (25 b). Yates, H. S. 3056 (3b). Williamson, C. S. 1367 (2c). Yingst, D. 10 (2a). Willis, B. s.n. (2a). Ld E. Tow (3а). Willits, V. 136, 270 (7). You (5). Wilson, C. B. о 204, 356 (2а). Ml. T. © ES (2c). Wilson, N. C. . (4). Yuncker, T. С. & E. С. 12015 (7); 12016 Winblad, Y. у. s.n. (3b). (11a). Winiecki, 1. 28 (8). Yuncker, T. G., Dawson, — & Youse, —. Witt, J. G. 1223 (11а); 1253 (7). uL (17 Woelflin, —. s.n. (22a). bel, H. E. s. n. (11a). INDEX TO SCIENTIFIC NAMES Previously published names accepted here are in Roman type; new names published in this monograph are in bold face type; synonyms are in italics. Amplophus —-..-.--------- КЕСЕНЕНІ гыр 388 Phyllactis Astrepbia і 450 mexicana |. 450 TTT ee ОФ ТУ. Баст. c —— 483 pratensis aaee e ed 481 DXHMPSHUS o 31 X e scie 481 Boerbaavia erecta 483 Valeriana 388 Hemisotria 3 acutiloba 407 Oligococe 388 var. лога а ы ые ——— 409 Patrinia Adan Sand c Зараа 398 ceratobbylla 425, 42 ИК лы e 457 longifolia ааа e ЕКА NUN аа 434 [Vor. 38 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN ИЙЕ REDIERE PEERS 465 gracilis ac sir iiu NE anomala - 394 ЖЕНЕ... ааа ай ЖАЙ apiifolia _.. 450 кечи ——H— M АШ е Се 409 т. В. ЗЫ 259 ы s клы Ыы ЕЛА T 441 Биз онч ыы ы ш АТП - 477 knautioides 2-40 ОАР E pea 402 laciniosa a ЯМ Visier Nac ca ER 445 Lan glassei 472 ERE сыс „ЖО ==? 431 КИКӨНЕ-- ынаны ылар да 483 саран I S SL cmt eren 404 laurifolia А 438 andolleana д ___ 465 laxissima 2—2... 438 capitata . 401 БЕЗИНИН LLL es aee A сс 425 ssp. acutiloba - » 407 longifolia 0 _________ 428 var. bracteosa - ге 402 ИРИС Со сылы ЕНЕР 431 ssp. californica 404 MICHI eee eo es. 406 ssp. capitata __ s 02 ЖЕЛИНЕ S ee КЕЛСЕ 475 ИЫ ыры а-ы 394, 398 ШЕРЛІ; nee: - 415 ssp. pubicarpa 406 var. wyomingensis аа Cu ИРОНИ e e да 436 Mikaniae ана» ыт ceratophylla 2425, 436, 483 Мариус азық ИМЕ 436 Ceratophyllae (Series) . аа 430 "MM LL ш 431 НЕЗ сасыса... 428 obovata _ 2 483 Catan (Эвге) ыш з. 437 с, Loo е е prim 415 oio NENNEN 438 officinalis ee ORC | columbiana — — з ш бы 413 Officinales (Series) . 390 CNN Las аа с. SU MEM, анаа А - 409 cucurbitifolia 445 palmatiloba 22 448 И а ырымы Сі Ер 406 ае Lou меке E - 472 cyclophylla --- 483 КӨРМЕ. асе сс ы Оше 410 BR eu educata 470 pry зақ сал Meet et 222. 438 deltoidea TERRI ne 459 ar. yungasensis ................................. 438 77, LL мс. 454 eseoli — d 463 var ined - 457 pilosiuscula ..... 0 455 var. densiflora - 2 455 Шш. а ы ы 481 Densiflorae e Genes) mE MEE Pratenses (Series) INEEN O ND: 479 denudata орыннын Ыш 452 prionophylla -aoaaa 422 dioica ТӨГӨР 2.2, ммен рб 430 ssp. sylvatica 1. 417 ӨК A ИИИ 417 vat. syivatica ass ИУ RM cca N ы ы T 404 domingensis 468 ЖИБЕННӘНІЕ ae есе ы сы 406 HN ET 399 pubicarpa - 406 Edules (Series) 420 pulcbella eee ee ЖҮН 459 A — 423 PIED РИБЕ iui асық» 422 ssp. ciliata ——— 428 Osa ... Я 483 р. OMIM eee 425 FM UDHNE LL eere scimus 436 EM LLL sect 425 о о < 55 аы илы ie ыс 455 ENR анаа 430 rbomboidea Т. - 442 X Ekmani | . — — — 469 robertianifolia 470 erysimoides Ў 442 ei » 442 stulosa 472 scandens ... 462 Follettiana Ө 20398 B angustiloba - 463 frigidorum |. 394 var. Candolleana | iia furfurescens 00 425 B denial e e 463, 466 Galleottiana д -- 481 Ж...” Ree eer eae Deli LL sae qct 438 таг. ONE sgean erar Dens 463 DENM ыс зы EM 407 Ear Malle oae ыс ине - 442 1951] MEYER—VALERIANA IN NORTH AMERICA СОМЕН AR Lo eere reae зылады тын 398 кзг к p — 322 44 SRILA A eee лос 405 septentrionalis БҮЛ ORAT ERE 417 sitchensis —— —— 2 ee 992 COULEE EE E ee Ха 5 398 ND S000 Е кы шс 398 MD. sitchensis oe 394 өр uliginosa e 399 SEHICD (eU C RERO c Е 422 Sorbifolia — — ——— нош ый 474 var. а КОИСО aoe 477 var. mexicana -M 475 COT DONA ы ы зз AE 476 Sorbifoliae reds iocur AOL subincisa ----- ire uo 2458 SUEITOF TH а ЕЕ ee oe 394 wyomingensis -......-.---------- GENERAL INDEX TO VOLUME XXXVIII w scientific names of plants and the final members of new combinations are printed in bold-face type; synonyms and page numbers having reference to figures and plates, in italics; and all other matter in ordinary type. A Actinostemon concolor var. a a 250 Adenogyne pachystachys, E = жарыйды. 25 Apocynaceae, bkn in the, VIII, 1 ri Botanical CER 8 "adds dba Магсёгау ла anum 190% ndleri, мо ‚ 170% of the series, 128; speciation, 131; sys- 02; EY 136; ver- nacular names, 155, 2 Associes, tree: at the Meis Botanical Garden Forest Preserve, 287; in the rks, 261 Astrephia, 450 *Bauhinia, 10; reflexa, 17; Storkii, 17 Oza 74 u J. Brenner: The changing idet flora of the Ozarks, 283; The recent intrusion of forests in the Ozarks, 261 comparison of parts of phytomers, 305, 360; V UP ME of pese tillers, 3 18 Boerbaavia erecta, 483 uis G. ust P. Beilmann, and: The changing forest flora of the Ozarks, 283; The recent intrusion of forests in the Ozarks, 261 Bro Browneopsis, 36 Only t with 22 monogra C Caesalpinoideae (subfamily of Leguminosae) , Caesalpinia, 87 Californian Delphiniums, Chromosome num- rs of, 1 Cassia, 39; fruticosa var. gatunensis, 81; hispidula var. "Killipii 55; Maxonii, 77; oxyphylla var. aig wk D stenocarpa var. Stenocarpoides, 60; a, 7 Cedar, I is the Окы, ү Cercidium cee 26 FRE of Californian Del- mgr and their geographical occur- 10 Gein de Colliguaya elegi 251 Copaifera, roti bum. 251 Cynometra, 25 D Delonix, 83 DE Californian, Chromosome numbers of, and their geographical oc- currence, 101; cytological tdg, ot 101; qp distribution, meiosis in, 102, 103 Dialium, 38 Ditrisynia, 52 Dwyer, John D.: Cynometra of Panama, 25; Copaifera of Pontii, 6 E Ecology of the Ozark area, 2 Epling, Carl, Harlan Lewis, a үз Mehl- ‚ б. Wyckhoff: E ne E Etter, Alfred G How de Bluegrass grows, 293 Excoecaria, 208, Explorations in i. тойса. early, 262, 266 F Fires in the Ozarks, 269 Flora of Panama. Part V, Fasc. 3 (Legumi- nosae, second part), e new entities are given in this General Index, since a full systematic index is included raph. (505) 506 Forest flora of the Ozarks: The changing, 283; historic, 279; logging and its effects on the, 26 Forest Preserve of the Arboretum: Red , 286; principal 284, 285; ks underlying, 283; soils ai the area, 284; tree associes, u Forests in the Ozarks: The ae е” of, 261; present resources of, G Game birds and animals of the Ozark region, 276 Geology: of Missouri Botanical Garden B" Preserve, 283; of the Ozarks, 262, Gree eenman, Jesse M., ose ipe s ne NN 250 5220 Gusson ymn nthe es, 250 найы, 208, 243 Haematoxylon, 93 Hemisotria, 388 Hippomane biglandulosa, 250 Historical account of the Ozarks vegeta- tional Miss 262 T. Hymena I Indian Mounds in the Ozarks, 269 J Juniperus virginiana in the Ozarks, 267 K Kentucky bluegrass, How it grows, 293 L Leguminosae of Panama (second part), 1 Lewis, а g^ -= Epling, G. A. ehl- quist, and G. ckh d romo- some К of Ж brie барышы and their geographical occurrence, 101 Lumber production in the Ozarks, 261, 266 M Macrolobium, 21; Pittieri, 33 Harlan Lewis, Carl Epling, and C. G. Wyckhoff: Chromo- some numbers of Californian Delphiniums and their geographical occurrence, 101 [Vor. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN Meyer, F. G.: Valeriana in North America and the West Indies (Valerianaceae), 377 Meiosis i in cells o 2342 The changing forest flora of the, ч И recent intrusion of for- ests in ; 4 Missouri кедді Garden Forest Preserve: forest flora of, 283; geology of, 2 Mora, 23 a $ c ‘at О a O Oat seedling, growth 7 г" 320 Ochrosia NET Oligococe, 6 changing forest flora of, 261; The recent intrusion of forests in the, 261 P Panama, Flora of, еу M Fasc. 3 (Legumi- nosae, second p Paralyxla macrophylla, (9) Parkins Patrinia abl 425; longifolia, 428 Peltogyne, 29 Peltophorum, 84 Peltospermum, Patrisii, 194 Phyllactis mexicana, 450; pratensis, 483 Phyllocarpus, 24 Poa pratensis, 293 Poeppigia, 22 Prairies in Missouri, 261 Prioria, 27 136; latisiliquum, 194; obovata, 483; R Rainfall: in the Ozarks, 270; records of, compared with growth of bluegrass, 315 Red Cedar in the Ozarks, 267 Rogers, David: A revision of Stillingia in the New World, 207 S Salines in the "voa 270 Sapium, 207, Слет а м пан НЩ 251 Schery, Robert W., Robert E. Woodson, Jr. 3 and: Flora of Райдер. Part V, Fasc. py minosae, second part), 1 Schinopsis quebracho- ы. 198 Schizolobium m 209, 250 Seborium 1951] INDEX Seed production of Ozark plants, 272 Soils in the Ozarks, 268, 284; Indian mounds » relation to erosion of, 269; types of, "silii 219; Bodenbenderi, 222; diph- 226; peruviana, 223; scutellifera, P. с, in the New World, A revision of, 07 ; generic relationships, 209; geography ue history, 208; index to exsiccatae, 252; index to scientific names, 257; sub- generic categories, 213; systematic criteria, 10; taxonomy, Succession of forest trees in the Ozarks, 283 Swartzia, 3; simplex var. darienensis, 7 T тшшш 3/7 Tbyroma, 136 Tragia ЕРТЕ! Tree associations in the Е orest Preserve of Missouri Botanical Garden Arboretum, 287 Valeriana in North America and the West ies (Valerianaceae), 377; classification, mic importance, 386; general historical review to Ad A 485; v д scientific names, 501; Жасады 38 Only the wi i vod monogr 507 * Valeriana, 377; capitata ssp. acutiloba, 407, ssp. californica, 404, ssp. pubicarpa, 406; deltoidea, 459; E: var. affinis, var. депо 455; edulis ssp. ciliata, 428, ssp. procera, 430; х Ek- manii, 469; palmatiloba, 448; sitchensis ssp. Scouleri, 398, ssp. uliginosa, 399; sorbifolia var. barbareaefolia, 477, var. mexicana, 475; tanacetifolia, 480 ху Weather conditions in the Ozarks, Wheat, growth in length of litre idi in- ternodes of, 30 dson, Robert E., Jr.: 95; Studies Jesse More Green- o 3 (Leguminosae, second part), Wyckoff, ©. G., Harlan Lewis, Carl Epling, G. L. Me shl quist, and: Chromosome numbers of Californian Delphiniums and their geographical occurrence, 101 Z Zea Mays, 307 new entities are given in the mm Index, since a full systematic index is included aph.