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Full text of "Phytologia"



,Vol.lQ 



1964 




V,IO 



<> \ 



PHYTOLOCSJIV 



o 



Designed to expedite botanical publication ^ 

'ol. 10 March, 1964 No. 1 



CONTENTS 



MITH, L. B., Notes on Bromeliaceae, XXI 1 

10LDENKE, H. N., Materials toward a monograph of the genus 

Verbena. XVII 56 



Published by Harold N. Moldenke and Alma L. Moldenke 

15 Glenbrook Avenue, 
Yonkers 5, New York, U. S. A. 

Price of this number, $1; per volume, $5.75, in advance 

Volume 9, No. 8, was issued March 6, 1964 



NOTES ON BROMELIACEAE, XXI 
Lyman B. Smith 



PrrCAIRNIA 



The following synopsis of Pitcairnia is long overdue, espe- 
cially as there is little hope of publishing it as part of a mon- 
ograph in the near future. The last complete treatment was that 
of Mez in the "Pflanzenreich" in 1935 with some 183 species. 
Since then, over l^D species either have been transferred to Puya 
or reduced outright and more than 100 new species have been 
added. 

In preparing the key, I have not tried to make it taxononic, 
although it approaches that state in places. Rather I have tried 
to produce something that would give accurate and rapid identifi- 
cations. This is not easy with the best material and involves 
mounting the sepals to show their shape when unrolled and dis- 
secting the ovary to ascertain the type of appendage on the o- 
vule. The worst material, which unfortunately involves several 
unicate types, must have each species entered in several places 
in the key in order to cover all contingencies. 

On the other hand, geographic correlations have proved quite 
helpful. For instance primitive types with compound inflores- 
cences are not found in Mexico, while advanced types with decidu- 
ous leaves predominate there in response to a strong dry season 
but are absent in the West Indies. Subgenus Pitcairnia centers 
in the northern Andes while in the Amazon and Orinoco drainages 
it is almost wholly replaced by subgenus Pepinia . 

The species in the text are in roughly taxonomic order by sub- 
genera, but ignorance of the type of ovule-appendage makes the 
position of several conjectural. I believe that this is the 
basic division of the genus and that the broadly alate ovule is 
the more primitive, though this last is difficult to prove. 

The line between Puya and Pitcairnia subgenus Pepinia is some- 
times difficult to draw, so I am retaining P. ferruginea and P. 
nana in Pitcairnia on the basis of their slightly inferior ova- 
ries and P. mirabilis on account of its somewhat zygoraorphic flo- 
wers. They might equally well go in Pxya on accoxmt of their 
strongly twisted petads in age, but there is no point in changing 
the status quo uiiless a definite improvement is gained thereby. 

1. Inflorescence ccmpound (sometimes normally simple inflores- 
cences develope 1 or 2 lateral branches and such should be 

sought below if not found here) Subkey I 

1. Inflorescence simple. 

2. Floral bracts ample, covering most or all of the flower Just 
before anthesis (unknown in P. quesnelioides ) ; leaves per- 
sistent, usually petiolate; pedicels mostly short and stout 

Subkey II 

1 



2 PHTTOLOOIA Vol. 10, no. 1 

2. Floral bracts narrow or small, exposing most or all of the 
flower. 
3. Larger leaf -blades deciduous along a straight transverse 

line, grass-like, almost never petiolate; plants evidently 

of a strongly xerophytic habitat Subkey III 

3. Larger leaf -blades (and all others) persistent, often petio- 
late; pleints of a more mesophytic habitat. 
4. Leaf -blades narrowed at base to a definite petiole. 

Subkey H 
4. Leaf-blades little if at all narrowed at base, not defi- 
nitely petiolate. 

5 . Leaf -blades serrate at least in part Subkey V 

5 . Leaf -blades entire throughout Subkey VI 



SUBKEY I 

1. Leaf -blades (or the larger if dimorphic) uniformly spinose- 

serrate or serrulate (imknown in P. platystemon ) , persistent 
2. Ovules and seeds alate with the appendage not produced at 

base or apex (not known in P. viridis ) . species of the Ama- 
zon and Orinoco drainages and two (P. ferruginea and P. vi- 
ridis ) of the central Andes. PI. I, fig. 1. " 
3. Leaf -blades 90 mm wide; flowers reflexed. Veneiuela. 

1. P. aeavifolia 
3. Leaf-blades not over 75 nm wide, 
if. Sepals densely ferruginous- or brown-lepidote , obtuse. 
5. Sepals ferruginous-lepidote, 12-45 buh long; inflorescence 
usually ample. Ecuador, Peru, Bolivia. 

34. P. ferruginea 

5. Sepals brown-lepidote, 12 nsn long; inflorescence pseudo- 

simple with branch-buds in the axils of the lower 

bracts. Peru 35. P. viridis 

4. Sepals whitish-lepidote or glabrous. 

6. Sepals 40-50 mm long; petals naked. 

7. Floral bracts minute, much shorter than the 20-30 mm 
pedicels; flowers pendent-secund. British Guiana, 

Surinam 2. P. nuda 

7. Floral bracts exceeding the 5-12 mm pedicels; flowers 
suberect . 
8. Leaf-blades covered beneath YfLth cinereous scales. 

Venezuela 3. P. wurdackii 

8. Leaf -blades glabrous on both sides. Colombia. 

4. P. mituensis 
6. Sepals 13-35 mm long. 
9. Primary bracts equaling or exceeding the sterile bases of 
the branches. 
10. Flowers strict; branches 5-6 cm long, simple; sepals 14 

mm long. Venezuela 5. P. kunhardtiana 

10. Flowers spreading; branches 10-35 cm long. 
11. Inflorescence bipinnate; sepals acute, 35 mm long. 

Venezuela, Colombia 6. P. bulbosa 



196U Smithy Notes on Bromellaceae 3 

. 11. Inflorescence trlpinnate; sepals obtuse, 20 mm long. 

Colcmbia 7. P. hellophlla 

9. Primary bracts shorter than the sterile bases of the 
branches . 
12. Flowers sessile; sepals acute, 15-18 mm long. 

Venezuela 8 . P. armata 

12. Flowers distinctly pedicellate. ~ 

13. Flowers pendent-secund . 
14. Flortil bracts 8 mm long, exceeding the short pedi- 
cels; leaf -blades wholly glabrous. Venezuela. 

9. P. breweri 

14. Floral bracts 4 mm long, much shoi*ter than the pedi- 

cels; leaf -blades cinereous-lepidote beneath. 

Surinam 10. P. geyskesii 

13. Flowers ascending to spreading, not at all secund. 

15. Pedicels ascending; sepals abruptly acute. Venezuela 

11. P. pruinosa 
15. Pedicels stiffly spreading; sepals acuminate. 

Venezuela, Colombia, Brazil 12. P. patent iflora 

2, Ovules and seeds bicaudate or subalate with the appendage 
more extended at the ends than at the side. Chiefly 
Andean South America and West Indies. PI. I, figs. 2, 3. 
16. Sepals echinate or tuber culate, 27-45 mm long. 
17. Floral bracts acuminate, nearly equaling to exceeding the 
10-30 nt pedicels; sepals 35-45 mm long, Colombia. 

44. P. echinata 

17. Floral bracts aplculate, 5 mn long; pedicels 10 mm long; 

sepals 27 mm long. Venezuela 45. P. tuberculata 

16. Sepsils not more than lepidote. 

18. Inflorescence pseudo-simple with branch-buds in the lower 

bracts, few-flowered; sepals 12 ran long, brown-lepidote . 

Peru 35. P. viridis 

18. Inflorescence obviously branched. 
19. Petals yellow, cream or green. 
20. Pedicels 10 mm long; petals appendaged. Hispaniola. 

232. P. samuelssonii 
20. Pedicels 15-23 mm long; petals naked. 
21. Sepals 8 mm wide, asymmetric; floral bracts about half 

as long as the pedicels. Peru 46. P. puyoides 

21. SepsLLs 4.5 im wide; floral bracts much more than half 

as long as the pedicels. Bolivia.. 47. P. platystemon 
19. Petals red, appendaged. 
22. Pedicels 20-25 mm long, not exceeding the lower floral 

bracts. Jamaica 233. P. bromeliifolia 

22. Pedicels not over 15 mm long. 
23. Leaf-blades homomorphic, 10-20 mm wide, the spines to 3 
mm long; sepals abru^ly acute or obtuse. Puerto 

Rico, Lesser Antilles 234. P. angustifolia 

23. Leaf-blades dimorphic, mostly more than 20 mm wide. 
24. Leaf -blades 10 mm wide; inflorescence few-branched. 

Dominican Republic 235. P. fuertesii 

24. Leaf-blades 25-40 mm wide. 



k PHITOLOGIA Vol. 10, no. 1 

25. Sepals 25 mm long; larger leaf -blades scarcely nar- 
rowed at base. Lesser Antilles.. ..236. P. gracilis 
25. Sepals 15-18 nun long; larger leaf -blades more or less 
petiolate. 
26. Sepals densely and persistently pale-lepidote. Peru, 

48. P. pulveirulenta 
26. Sepals soon glabrous. Peru, Bolivia. 

49. P. paniculata 
1. Leaf -blades only partially spinose-serrate or serrulate or 
else entire. 
27. Sepals obtuse or rounded and apiculate; ovules or seeds 

mostly with an apical appendage (broadly winged in P. kil- 
lipiana and unknown in P. lechleri and P. verrucosaT . 
28, Leaves wholly entire, the blades all persistent. 
29. Pedicels 40 mm long; leaves petiolate; sepals verrucose. 

Colombia 50. P, verrucosa 

29. Pedicels 5-19 mm long. 
30. Lateral branches only 2 cm long; inflorescence narrowly 

thyreoid. Peru 51, P. ruiziana 

30. Lateral branches much longer. 
31. Racemes much longer than their peduncles, lax; sepals 33 

mm long, Colombia 36. P. killipiana 

31, Racemes no longer than their peduncles; sepsils 15-20 mn 
long. 
32. Plant 2 m high; leaf-blades 40 mm wide. Colombia, 

Ecuador 52. P. dendroidea 

32. Plant 7 dm high; leaf -blades 18 mm wide. Dominican 

Republic 237. P, .jimenezii 

28. Leaves partially spinose-serrate or serrulate. 
33. Leaf -blades (the larger ones) deciduous along a straight 

transverse line; pedicels 5-7 bbb long; petals appendaged 
34. Floral bracts eocceeding at least the lower pedicels; 
sepals 12 mm long, Peru. 

35. Leaves all alike; blades to 9 mm wide 53. P. rigida 

35- Leaves dimorphic; blades to 20 mm wide... 54. P, lechleri 

34. Floral bracts all much shorter than the pedicels. 

36. Leaves all alike; sepals 11-15 mm long. Peru, Bolivia. 

55. P. inermis 
36. Leaves dimorphic; sepals 22 mm long, Colombia. 

56. P. arenicola 
33. Leaf-blades all persistent; pedicels 4-30 mm long. 
37. Leaf-blades long-spreading-spinose at base, 30-40 mm 

wide; inflorescence amply bipinnate. Colombia, Ecuador 
38. Pedicels 15 mm long 57. P. commixta 

38. Pedicels not over 7 mm long 58. P. lehmannii 

37. Leaf -blades (larger) not prominently spinose at base (or 

not over 22 mm wide), 8-32 mm wide; inflorescence de- 
pauperat e-ccMnpound . 

39. Pedicels 15-30 mm long; sepals 20-30 mm long, 

40. Leaves all alike. Jamaica 238, P, platyphylla 

40. Leaves dimorphic. 
41. Petals naked. Argentina 59. P. oranensis 



I961i Smith, Notes on Eromellaceae 5 

41. Petals appendaged. Peru, Bolivia, Brazil. 

60. P, subpetiolata 
39. Pedicels 4-14 mm long. 
42. Leaf-blades 20-35 nm wide, petiolate or subpetiolate . 
43. Sepals 2? mm long; scape-bracts nearly all remote. 

Argentina 59. P. oranensis 

43. Sepals 10-18 mm long. 
44. Leaves spreading-spinose at base. Peru. 

61. P. truncata 
44. Leaves finely serrtilate at most. Argentina. 

62. P. chiriguema 
42. Leaf-bladea 8-18 Bm wide, slightly if at all narrowed 
at base. 
45. Petals yellow or cream; sepals 21 mm long; flowers 

finally recuinred. Hispaniola. ..232. P. samuelssonii 
45. Petals red; sepals 12-20 mm long. 
46. Petals naked or with 2 minute oblique calli; sepals 

12 mm long. Peru ,63. P. tarapotensis 

46. Petals bearing a large scale at base; sepals 14-20 mm 
long. 
47. Sepals 20 mm long. Dominican Republic. 

237. P. , 1imenezii 

47. Sepals 14-16 mm long. Cuba 239.~P. cubensia 

27. Sepals acute or acuminate. 
48. The sepals 30-50 mm long. 
49. Sep»ls echinate with stipitate-stellate trichomes. 

Colofflbia 44. P . echinata 

49. Sepals not more than lepidote or verruculose. 
50. Leaf-blades petiolate, persistent, 50-55 mm wide; 
pedicels 20 mm long, Colombia. 
51* Sepals 38 mm long; pedicels biangulate, exceeding the 

persistent floral bracts 64. P. chocoensis 

51. Sepals 50 mm long; pedicels terete, shorter than the 

deciduous floral bracts 65. P. haughtii 

50. Leaf-blades only slightly narrowed toward base, 15-30 mm 
wide. 

52. Racemes dense; floral bracts narrowly lanceolate; 

larger leaf -blades deciduous. Colombia. 

66. P. megasepala 
52. Racemes lax; floral bracts broad. 
53. Leaf -blades trimorphic, the Itirgest deciduous, entire. 

Colombia 67. P. trimorpha 

53. Leaf -blades apparently all alike, persistent, serrulate 
especiaO-ly toward base; sepals verruculose. 

Colombia, Ecuador 68. P. poortmanii 

48. The sepels 9-25 mm long. 
54. Floral bracts equaling or exceeding the lower pedicels. 
55. Petals white, yellow or green; sepals 12-22 mn long. 
56. Sepals 12-15 mm long; floral bracts ample, imbricate 
before anthesis, then deciduous. Colombia. 

69. P. kniphofioides 
56. Sepals 21-22 mm long; floral bracts persistent. 



P H Y T L G I A Vol. 10, no. 1 

57. Lower floral bracts lanceolate, about equaling the 

sepals. Colombia 70. P. tolimensis 

57. Lower floral bracts broadly ovate, at most equaling the 

pedicels. Bolivia 47. P. platystemon 

55. Petals red (uncertain in P. ulei . but the lower floral 
bracts narrow and about equaling the pedicels). 
58. Leaves all alike or nearly so. 
59. Sepals 10-16 mm long. 
60. Panicle aa^le; leaf -blades 13-19 mm wide. Venezuela, 
Trinidad 71. P. integrifolia 

60. Panicle depauperate; leaf-blades 10 mm wide. Cuba. 

239. P. cubensis 
59. Sepals 18-25 mm long. 

61. Ovary only 1/3 superior; panicle depauperate, subdense 

with ascending branches; sepals 25 mm long. Brazil. 

248. P. ulei 
61. Ovary 1/2-3/4 superior. 
62. Pedicels 20-25 mm long. Jamaica. 



62. Pedicels 10-15 nan long. 
63. Floral bracts elliptic, apiculate. Peini. 



233. P. bromeliifolia 

Peini. 
72. P. cassapensis 
63. Floiral bracts narix)w, acuminate. 
64. Inflorescence amply paniculate; floi*al bracts 
mostly shorter than the pedicels. Brazil. 

249. P. anthericoides 
64. Inflorescence few-branched; floral bracts mostly 
exceeding the pedicels. 
65. Leaves dilated above the sheath; sepals 19 mm 

long. Venezuela 73. P. moritziana 

65. Leaves not dilated above the sheath; sepals 25 mm 
long. Puerto Rico (?), Lesser Antilles. 

240. P. latifolia 
58. Leaves strongly dimorphic. 
66. Primary bracts about half as long as the axillary 
branches and much exceeding their short sterile 

bases. Peru 74. P. cuzcoensis 

66. Primary bracts several times shorter than the axillary 
branches and usually shorter than their sterile bases 
67. Scape 2-3 mm in diameter; foliaceous leaves not over 5 
dm long; plant to 5 dm high. Colombia. 

75. P. schultzei 
67. Scape stouter; foliaceous leaves to 7 dm long or more; 
plant over 1 m high. 
68, Larger leaf -blades deciduous, Peru. 

76. P. vargasiana 
68. Larger leaf-blades persistent. 
69. Smaller leaf -blades spiniform. Panama. 

186. P. chiriquensis 
69. Smaller leaf -blades with £in entire filiform flexuous 
apical half. Venezuela, Trinidad. 

71. P. integrifolia 



196U Smith, Notes on Bromellaceae 7 

54. Floral bracts all shorter than the pedicels. 
70. Petals yellow, green or cream; pedicels to 25 mm long; 
ovary over 1/2 superior. 
71. Floral bracts broadly ovate. Bolivia. 

47. P. platystemon 

71. Floral bracts narrowly triangular. Costa Rica. 

77. P. halophila 
70. Petals red or rarely blue (uncertain in P. ulel . but the 
pedicels only 8 mm long and the ovary only 1/3 
superior) . 

72, Panicle depauperate, few-branched. 

73. Pedicels to 8 mm long; ovary l/3 superior, Brazil, 

248. P, ulei 
73. Pedicels 15-30 mm long; ovary 1/2-2/3 superior. 
74. Leaves all alike, the blades 18-32 mm wide, Jamaica, 

238. P. platyphylla 
74. Leaves dimorphic, the larger blades only 11 mm wide, 

Guatemala 187. P. flagellaris 

72. Panicle ample, much branched. 
75. Sepals and ovary echinate; stamens exserted. Colombia, 

78. P. exserta 
75. Sepals and ovary not more than lepidote. 
76. Ovules and seeds winged; pedicels straight, spreading, 
3-4 times longer than the floral bracts. Colombia, 

Venezuela, Brazil 12. P, patent iflora 

76. Ovules and seeds caudate to apiculate; pedicels mostly 
curved. 
77. Sepals 9-17 mm long. 
78. Inflorescence amply 3-pinnate; pedicels 5-7 mm long; 
leaves all alike; sepsils 9-12 mm long, Costa Rica 

188. P. valerii 
78. Inflorescence not more than 2-pinnate or the pedi- 
cels 10-15 nm long; leaves mostly dimorphic. 
79. Floral bracts nearly equaling the pedicels, lance- 
olate; leaves alike with blades 13-19 mm wide or 
the reduced ones with soft entire filiform apices 

Venezuela, Trinidad 71. P. Integrifolia 

79. Floral bracts not much more than half as long as 
the pedicels or less. 
80. Larger leaf -blades 25-35 mm wide. 
81. Sepals densely and persistently pale-lepidote . 

Peru 48. P. pulverulenta 

81, Sepals soon glabrous. Peru, Bolivia. 

49. P. paniculata 
80. Larger leaf -blades to 15 mm wide. 

82. Branches suberect; pet«Q.s bearing a scale at base 

Ecuador 79. P. de vans ay ana 

82. Branches spreading; petals naked or with 2 

oblique calli. Peru 63. P. tarapotensis 

77. Sepals 18-25 mm long. 
83. Leaf-blades 80 ram wide; inflorescence 4-pinnate; 

sepals green. Colombia 80. P, diffusa 



8 PHTTOLOGIA Vol. 10, no. 1 

83. Leaf -blades 12-35 mm wide; Inflorescence 2-3-pinnate 
8/f> Largest pedicels 15-18 mm long; leaf-blades to 35 

mm wide. Bolivia 81. P. multiramosa 

8if. Largest pedicels 9-13 am long. 
85. Ovary 3/4 to almost wholly superior; leaf -blades 
22-35 mm wide. 
86. Leaf -blades furfuraceous beneath; petals naked. 

Bolivia 82. P. odontopoda 

86. Leaf-blades not conspicuously furfuraceous 

beneath; petals appendaged. Peru, Bolivia. 

49. P. panieulata 
85. Ovary only about 1/2 superior; leaf -blades 12-30 
mm wide. 

87. Leaf -blades to 12 loa wide; ovules very short- 

caudate . Brazil 249. P. anthericoides 

87. Leaf -blades 20-30 mm wide. *" 
88. Leaves petiolate; petals naked. Bolivia. 

83. P. divaricata 

88. Leaves not petiolate; petals appendaged. Lesser 

Antilles 236. P. gracilis 

SUBKEI II 

1. Scape very short or none; inflorescence short. 
2. Sepals carina te, 35-40 nm long; leaf -blades linear-triangular 

Bolivia 37. P. nana 

2. Sepals ecarinate, 15-20 mm long. " 

3. Plant caulescent; leaf-blades linear, 5-10 nm wide. Panama, 

Peru 84. P. aphelandriflora 

3. Plant stemless; leaf -blades oblong-sfjatulate, 30 mm wide. 

Uexico 189. P. tabulifomis 

1. Scape well developed (unknown in P. poeppi^ana . but the 
inflorescence elongate). 
4. Sepals not over 15 mm long. Colombia. 
5. Leaves dimorphic; floz*al bracts quickly deciduous; scape- 
bracts shoirter than the upper intemodes. 

69. P. kniphofioides 
5. Leases uniform, entire; floral bracts persistent; scape- 
bracts all longer than the intemodes. 
6. Floral bracts concolorous, straight; seeds long-caudate. 

85. P. capitata 
6. Floral bracts bicolorous, the upper part strongly reflexed; 

ovules obtuse 86. P. guzmanioides 

4. Sepals 18-75 mm long. 

7. Ovary 3/4 inferior, subsessile; capsule indehiscent; ovules 
winged; floral bracts deciduous, unknown. Colombia. 

13. P. quesnelioides 
7. Ovary 1/2 inferior or less. 
8. Sepals exceeding at least the upper floral bracts. 
9. Floral bracts rounded, obtuse or mucronulate, persistent. 
10. Flowers slenderly pedicellate; sepals obtuse, 40 mn 

long; leaf -blades entire. Colombia 87. P. archeri 



196U Sniith, Notes on Bromellaceae 9 

10. Flowers sessile; sepals acute, 25 mm long; leaf-blades 

serrate. Peru 88. P. poepplgiana 

9. Floral bracts acute or acuminate, deciduoas in some 
species, 

11. Sepals 60 mn long, linear-lanceolate, uncinate. 

Colombia 89. P. macranthera 

11. Sepals 20-37 nm long. 
12. Inflorescence lax toward base. 
13, Sepals oblong, rounded and apiculate; floral bracts 
even, coriaceous. British Honduras, Guatemala, 
Ueocico 190. P. recurvata 

13. Sepals elliptic, obtuse, asymmetric; floral bracts 

nerved, subehArtaceous. Central America, Guiana, 

Venesuela, Colombia 90. P. maidifolia 

12. Inflorescence dense throughout. 
lA. Posterior sepals broadly alate-carinate; flowers sub- 
sessile; inflorescence ellipsoid, 8 cm long; leaf- 
blades 20 mm wide. Peru 91. P. sandwnanll 

14. Posterior sepals not alate or if somewhat so (? P. 

sceptriforais ) then the flowers slenderly 
pedicellate. 
15, Floral bracts subcoriaceous , persistent. 
16. Sepals 20-22 mm long, pale-lepidote ; leaf -blades 

20-35 mm wide. Lesser Antilles. .. ,241. P. spicata 

16, Sepals 37 ma long, brown-lanate ; leaf-blades 90 mm 

wide, Colombia 92. P. maritima 

15. Floral bracts membranaceous, more or less deciduous. 

17. Flowers subsessile; Inflorescence 7 cm long. Peru. 

93. P. ferreyrae 
17. Flowers slenderly pedicellate for 3-8 mm; inflores- 
cence 20-60 cm long. 
18. Sepals carinate; inflorescence glabrous. Peru. 

94. P. sceptriformis 
18. Sepals ecarinate; inflorescence flocculose, often 
secund-f lowered. Costa Rica to British Guiana 

and Bolivia 95. P. brittoniana 

8. Sepals completely covered by the floral bracts. 
19. Floral bracts with divergent to spreading apices. 
20. Leaf -blades elliptic to obovate-lanceolate, only 3-4 
times as long as wide; sepals acuminate. 
21, Leaf -blades 10-14 cm wide, apiculate. Colombia, 

Ecuador 96. P. ni^a 

21. Leaf -blades 5 cm wide, acuminate. Ecuador. 

97. P. pulchella 
20. Leaf-blades lanceolate to linear, much more than 4 times 
as long as broad. 

22, Flowers refl«xed and the triangular acuminate floral 

bracts deciduous after anthesis; pedicels 8-10 um 

lon^. Colombia 98. P. brongniartiana 

22. Flowers remaining erect and floral bracts imbricate 
after anthesis. 



10 PHTTOIOGIA Vol. 10, no. 1 

• 23. Pedicels 10 urn long, slender, ferruginous-tomentulose. 
"Guatemala". (Cultivation)... 191. P. longebracteata 
23. Pedicels very short to almost none, though often 
slender. 
2k» Floral bracts serrate, 10 cm long; sepals narirowly 
triangular, ZtO mm long. Colombia, Ecuador. 

99. P. brunnescens 
2k. Floral bracts entire. 
25. Petals puberulent outside, orange; remainder of the 
inflorescence densely fenruginous-lepidote. 

Colombia 100. P. barrigae 

25. Petals glabrous. 
26. Floral bracts membranaceous, bladelessj inflores- 
cence obtuse. Meocico .192. P. matudae 

26. Floral bracts subcoriaceous or chartaceous. 
27. Petioles entire. 
28. Learf -blades entire; inflorescence subcylindric ; 
floz>al bracts nith a distinct apicsd blade. 
Central America 193. P. wendlandii 

28. Leaf -blades serrulate; inflorescence clavate; 

floral bracts without a distinct apical blade. 

Peru 101. P. clavata 

27. Petioles spinose-serrate. 

29. Sepals linear-lanceolate, acute, 32-39 mm long. 

Colombia 102. P. squarrosa 

29. Sepals oblong, truncate or broadly acute and 
apiculate, 25-3O mm long. 
30. Floral bracts with a narrowly triangular diver- 
gent blade, red-purple, red or yellowish. 

Costa Rica to Colombia 103. P. atrorubens 

30. Floral bracts acute without a distinct blade, 

only slightly divergent, pale green. Ecuador. 

104. P. campii 
19. Floral bracts straight toward apex, usually strict and 

closely imbricate but lax in P. elongata and spreading 
in P. maidifolia . ~ 

31, Sepals 75 mm long; flowers not imbricate nor concealing 
the rhachis but the apical sterile b racts doing so. 

Colombia, Ecuador 105. P. elongata 

31. Sepals 18-50 mm long. 
32. Leaves spinose-serrate either along the petiole or the 

blade. 
33. Floral bracts corrugated, soon disintegrating to a 
fine mesh of fibers; sepals broad, rounded, kO mm 

long. ColcHubia, Ecuador 106. P. bakeri 

33. Floral bracts even or finely nerved but not corrugated 
34. Leaf -blades 75-125 mm wide. 
35. Sepals 25 mm long, oblong, abruptly acute. Peru. 

88. P. poeppigiana 
35. Sepals 45-50 mm long. 



196U Smith, Notes on Bromellaceae 11 

36. Leaf -blades oblanceolate , to 125 mm widej sepals 
broadly acute and apiculate. Colombia, Panama, 

Costa Rica 107. P» oblanceolata 

36. Leaf -blades lanceolate or linear-lanceolate, 75-100 
nun wide; sepals acuminate. Colombia. 
37. Pedicels very stout; leaves all alike, 

108. P. arcuata 
37. Pedicels slender; leaves dimorphic, 

109. P. fosteriana 
34. Leaf -blades 30-55 mm wide. ~ 

38. Petiole entire; leaf -blade serrate toward apex, 

Guatemala 194. P. aacrochlamys 

38, Petiole serrate; blade entire, 
39. Flowers slenderly pedicellate, 
40, Sepals oblong, obtuse and apiculate, 

Guatemala 195 > P. hensleyana 

40. Sepals narrowly triangular, Peru. 

110. P. umbratills 
39. Flowers subsessile. 

41. Leaf -blades dimorphic. Ecuador 104. P. campii 

41. Leaf-blades all alike. 

42. Petals naked. Mexico 196, P, imbricata 

42. Petals appendaged. Guatemala, British Honduras. 

197. P. petiolata 
32. Leaves entire throughout, 
43. Sepals 40 mm long, acuminate, menibranaceous , floecose; 
floral bracts acuminate, Guatemala, 

198. P, carioana 
43. Sepals 18-30 mm long. "" 

44. Sepals vestite; leaf -blades petiolate. 
45. Leaf-blades 25 om wide; sepals 19 mm long, the pos- 
terior alate-carinate . Eciiador. 

111. P. hitehcockiana 
45. Leaf -blades 50-63 mm wide; sepals 25-28 mm long. 
46. Ovary almost wholly superior; petals naked, yellow. 

Ecuador 112. P. sceptrigera 

46. Ovary about 1/2 superior; petals appendaged, red. 

Peru 113. P, asplundii 

44. Sepals glabrous or nearly so, 
47. Leaves all ending in a stout pungent black subulus; 
sepals nariMJwly triangular. Peru. 

114. P. subulifera 
47. Leaves not modified at apeac. 
48. Leaf-blades merely narrowed toward base, not 
petiolate . 
49. Floral bracts ovate, acute, the upper ones 37 mm 
long; inflorescence racemose, lax at base, 
sepals obtuse. Costa Rica. 

199. P. membranifolia 
49. Florsd bracts foliaceous, 60-80 mm long; inflores- 
cence subcorymbose ; sepals acuminate. 
" Guatemala" . ( Cult ivat ion ) . 



12 PHITOLOGIA Vol. 10, no. 1 

, 191. P. longebracteata 

/jfi. Leaf-blades distinctly petiolate, 
50. Floral bracts broadly acute or roxmded and 
apicolate. 
51. Sepals 18 om long; floral bracts bright purple. 

Venezuela 115. P. altensteinii 

51. Sepals 24 bub long; floral bracts reddish yellow. 

Uexico 200. P. densiflora 

50. Floral bracts acuminate. 

52. Petals naked; leaf -blades 45 bod wide. Central 

America or Uexico? .201 . P . ochrolenca 

52. Petals appendaged; leaf -blades 50-85 om wide. 

Central America 193. P* wendlandii 

SUBKEI III 

1. Petals bearing a scale on the inner side at base. Northern 
Andes and Brazil, except for P. heterophylla extending to 
Uexico and P. theae in Costa Rica. 
2. Scape and inflorescence very short. Uexico to Venezuela and 

Peru i. 116. P. heterophylla 

2. Scape (unknown in P. crassa ) andj^florescence elongate. 
3. Sepals 30-42 mm long; petals mostly yellow or white (color 
unknown in P. crassa and P. theae, sometimes red in P. 
megasepala ) . 
4. Posterior sepals alate. 
5. Inflorescence dense; plant with a long erect stasm. Peru. 

117. P. august ii 

5. Inflorescence lax; plant stemless. Colombia. 

66. P. megasepala 
4* Posterior sepsils not more than carinate. 

6. Floral bracts triangular or lanceolate, acuminate. 

7. Inflorescence cinereous-furfuraceous; scape-bracts much 
exceeding the intemodes, the upper ones 5 cm long. 
Costa Rica 202. P. theae 

7. Inflorescence glabrous; scape-bracts slightly shorter 

than the upper intemodes. Ecuador. 

118. P. aequatorialis 
6. Floral bracts broadly lanceolate or elliptic, acute or 
apiculate . 

8. Pedicels slender, 11 mm long, all but the lowest exceed- 

ing the floral bracts. Colombia 67. P. trimorpha 

8. Pedicels 2 mm in diameter, 25 mm long, all shorter than 
the floral bracts; leaves unknown, species doubtfully 

belonging in this subkey. Bolivia 119. P. crassa 

3. Sepals 12-25 mm long. 
9. Pedicels obconic, alate, very short. Colombia. 

70. P. tolimensis 
9. Pedicels slenderly cylindric, not alate. 
10. Sepals emarginate, 12 mm long; pedicels 5 mm long. Peru. 

53. P. rigida 



1961i Smith, Notes on Bromellaceae 13 

10. Sepals acute or acuminate, 14-25 am long; pedicels 3-20 
am long. 
11. Leaf-blades all alike and deciduous, 10 bb wide; plant 
caulescent, branched; presence of petal-appendage 

uncertain. Colombia 120. P. lignosa 

11. Leaf -blades dimorphic or at least the inner persistent. 
12. Pedicels 10-20 mm long. Brazil. 
13. Sepals alate, 16-20 mm long 30. P. ensifolia 

13. Sepals ecarinate, 14 mm long 250. P. torresiana 

12, Pedicels 3-10 mm long. Northern Andes. 

14. Inflorescence sparsely flocculose, soon glabrous; 

leaf -blades 5-10 dm long, 5-12 mm idde. Colombia. 

121. P. stenophylla 
14. Inflorescence densely and persistently ttanentose- 
lepidote; leaf-blades 3-4 dm long, 15-30 mm wide. 
15. Leaf-blades 30 mm wide, narrowly lanceolate, 

constricted at base. Ecuador 122. P. pavonii 

15. Leaf-blades 15 nm wide, linear, scarcely constricted 

at base. Colombia to Per u 123. P. pungens 

1. Petals naked. 
16. Pedicels not over 2 mm long. 
17. Sepals 40-50 mm long. 
18, Leaf-blades 35 nn wide; inflorescence corymbiform. Peru. 

124. P. billbergioides 

18. Leaf -blades 15 lan wide; inflorescence slenderly sub- 

ellipsoid. Guatemala, Mexico. 203. P. saxicola 
17. Sepals not over 30 mm long. 

19. Inflorescence lax, at least toward the base. 

20. Sepals oblong, rounded and apiculate, 7 mm long. Pern. 

125. P. melanopoda 
20. Sepals lanceolate, acute or acuminate, 25-28 mm long. 
21. Flowers subdistichous ; sepals acute; inflorescence lax 

throughout , Mexico 204. P* purpusil 

21. Flowers many-ranked; sepals acuminate; inflorescence 
dense toward apex. Honduras, Salvador, Guatemala. 

205. P. ealderonii 
19. Inflorescence dense throughout. Mexico. 
22, Blades of the lower scape-bracts equaling the inflores- 
cence; floral bracts covered with a membrane of 

coalesced scales 206. P. roseana 

22. Blades of the scape-bracts much shorter than the inflo- 
rescence; indunent of the floral bracts of distinct 
scales . 
23. Scape-bracts lance-triangular, exposing much of the 

scape 207. P. micheliana 

23. Scape-bracts broadly ovate, ample, concejiling the scape. 

208. P. cylindrostachya 
16. Pedicels 4-30 nan long. 
24. Petals 80-100 mm long. 
25. Sepals densely vest it e; flowers secund, spreading to 
reflexed. Mexico. 



Ik PHYTOLOGIA Vol. 10, no. 1 

26. Floreil bracts about equaling the pedicels; sepals covered 
with appressed whitish scales 209. P. lepirosa 

26. Floral bracts about 4 times as long as the pedicels; 

sepals ferruginous-lanate 210. P. modesta 

25. Sepals glabrous. 

27. Sepals acuminate. Andes. 

28. Petals yellow; pedicels 5-12 mm long. Peru. 
29. Scape 8 mm in diameter; scape-bracts exceeding the 
intenaodes; flowers divergent, not secund. 

126. P. lopezii 
29* Scape 5 mm in diameter; scape-bracts shorter than the 
intemodes; flowers spreading to decurved, secund. 

127. P. decurvata 
28. Petals brownish violet; pedicels 20 Hm long. Ecuador. 

128. P. violascens 
27. Sepals acute or broadly acute. North America. 
30. Petals glabrous, red; scape-bracts entire. Guatemala. 

211. P. tuerckheimii 
30. Petals tomentose-lepidote at apex, white; lower scape- 
bracts serrate. Mexico 212. P. hintoniana 

24. Petals 34-70 ma long. ~ 

31. Floral bracts distinctly shorter than some of the pedicels 
32, Floral bracts broadly ovate. 
33. Pedicels to 23 mm long; flowers rather dense; petals 34 
mm long, yellow or green. Bolivia.. 47. P. platystemon 

33. Pedicels not over 15 mm long; flowers lax; petals 70 mm 

long, reddish green. Mexico 213. P. mooreana 

32, Floral bracts narrow, acuminate. 

34. Pedicels suberect, 10-15 mm long; axis flexudus. Mexico. 

214. P. flexuosa 
34. Pedicels spreading to reflexed. 
35, Pedicels not over 15 mm long; flowers secund. Mexico. 

215. P. palmeri 
35. Pedicels 20 mm long; flowers not secund. Peni. 

129. P. fractifolia 
31. Floral bracts equaling or exceeding the pedicels. 
36. Lower floral bracts subf oliaceous , much exceeding the 

flowers. Guatemala 216. P. puberula 

36. Lower floral bracts vaginiform, shorter than the flowers. 
37, Lower floral bracts serr\ilate; sepals oblong, rounded 

and apiculate. Brazil 31. P. limae 

37. Lower floral bracts entire. 
38. Leaves all entire or some very obscurely sernxlate 
(P. tillandsioides ). 
39. Inflorescence few-flowered, lax; scape-bracts longer 
or shorter than the internodes; posterior sepals 

subalate. Mexico 217. P. tillandsioides 

39. Inflorescence many-flowered. 
40. Scape-bracts shorter than the upper intemodes. 
41, Leaf -blades 2 mm wide, vestite with linear scales. 

Bolivia 130. P. eardenasii 



196U Smith, Notes on Bromellaceae 15 

, 41. Leaf-blades 20 mm wide, densely lepidote beneath. 

Peru 131. P. straminea 

i+0. Scape-bracts all longer than the Intemodes. 
l^2. Petals red; floral bracts about ©quailing the 

pedicels . Brazil 251 . P . decidua 

42. Petals white; lowest florjil bracts exceeding the 

sepals. Mexico 218. P. schiedeana 

38. Leaves at least partially spinose-serrate, either on 
the spiniform blades or on the persistent bases of 
the foliaceous blades or both together. 
43 • Pedicels strongly biangulate or alate. 
44. Sepals acuminate, 15-18 mm long; flowers strongly 

secund. Mexico 219. P. pteropoda 

44. Sepals obtuse or trvincate, to 40 mn long. Ecuador. 

132. P. alata 
43. Pedicels terete or faintly angled. 

45. Scape very short, exceeded by the persistent leaf- 

bases; inflorescence elongate, sublax. Mexico. 

220. P. micropoda 
45. Scape evident, much exceeding the leaf-bases. 
46. Scape-bracts distinctly shorter than the upper 
internodes. 
47- Inflorescence dense; floral bracts acute, equaling 
or slightly exceeding the pedicels; flowers not 
secund. Brazil 252. P. glaziovii 

47. Inflorescence lax; floral bracts acuminate, about 

twice the pedicels; flowers secund. Peru. 

133. P. scandens 
46. Scape-bracts all longer than the intemodes. 

48. Leaves all alike; plant long-caulescent. Colombia. 

120. P. lignosa 
48. Leaves dimorphic; plant generally stemless. 
49. Posterior sepals alate, Mexico. 
50. Sepals acuminate, glabrous; scape-bracts entire. 

221. P. karwinskyana 

50. Sepals broadly acute or obtuse, Ian ate; lowest 

scape-bracts spinose 222. P. militaris 

49. Posterior sepals not more than carinate. 

51. Petals yellow; floral bracts exceeding the 5 mm 

long pedicels. Ecuador 134. P. lutescena 

51. Petals red. 
52. Scape-bracts foliaceous, equaling the leaves. 

Mexico 223. P. foliacea 

52. Scape-bracts short, not at all foliaceous. 
53. Sepals densely and persistently lepidote. 

Mexico 224. P. sordida 

53. Sepals sparsely lepidote or apically 
barbellate, soon glabrous. 
54. Pedicels 5-6 mm long. Andes. 
55. Inflorescence dense; sepals 33 mm long; 
petals 65-70 mm long. Ecuador. 

135. P. erratica 



16 PHTTOLOGIA Vol. 10, no, 1 

55 • Inflorescence lax; sepals 22 ran long; petals 

UO HUB long. Peru 136. P. aclcolaris 

54. Pedicels 8-15 nn long. ~ 
56. Plants caulescent, stolonif erous . Peru. 

137. P. rlparla 
56. Plants stemless, without stolons. Uexico. 
57. Flowers stirongly spreading-secund ; pedicels 

15 mm long 215. P. palmeri 

57. Flowers polystichous , scarcely if at all 
secund, pedicels 8-10 nm long. 
58. Floral bracts exceeding all but the upper- 
most sepals 225. P. aontieola 

58. Floral bracts all much shorter than the 

sepals 226. P. ringens 

SUBKET IV 

1. Floral bracts shorter than the upper pedicels. 
2. Sepals 1^2-60 am long; pedicels 40~70 am long; leaves entire. 
Colombia. 

3. Sepals veirrucose, obtuse 50. P. verrucosa 

3. Sepals even or nerved, acute or acuminate. 
4. Leaf -blades 65-90 mm wide; pedicels 50-70 mm long. 

138. P. spectabilis 
4. Leaf -blades not over 30 mm wide; pedicels 50-60 mm long. 

5. Sepals densely ferruginous-tomentose I39. P. laxissima 

5. Sepals glabrous 140. P. longipes 

2. Sepals not over 35 mm long; pedicels 5-40 mm long. 
6. Sepals rounded or blunt or apiculate. 
7. Pedicels only 5 mm long; leaf -blades to 95 nm wide. 

Ecuador 141. P. elliptica 

7. Pedicels 10-40 mm long; leaf -blades 16-70 imn wide. 
8. Leaves spinose-serrate on the petioles or the bases of the 
blades . 
9. Scape-bracts all exceeding the intemodes; pedicels not 
over 10 mm long; leaf -blades 35 mm wide; sepals 12-18 

mm long. Argentina 62. P. chiriguana 

9. Scape-bracts shorter than -the upper intemodes; pedicels 
to 25 mm long; leaf -blades 16-24 mm wide; sepals 20-30 
mm long. 
10. Floral bracts to 5 mm long; petals naked. Argentina. 

59. P. oranensia 
10. Floral bracts 10-20 mm long; petals appendaged. Peru, 

Bolivia, Brazil 60. P. subpetiolata 

8. Leaves wholly entire. 
11. Leaf -blades 60-70 mm wide. 
12. Pedicels 10-15 nm lo^g; flowers spreading-secund. 

Mexico 227* P* chiapenaie 

12. Pedicles to 40 mm long; flowers ascending, not secund. 

Colombia ....142. P. tumulicola 

11. Leaf-blades 20-30 mm wide. 



196U Smith, Notes on Brcmellaceae 17 

13, Pedicels 10-20 mm long; ovary 2/3 superior; ovules 

caudate. Peru 143. P. biflora 

13. Pedicels 25-35 mn long; ovary I/4 superior; seeds 

winged. Colombia 38* P« alborubra 

6. Sepals acute or acuminate. 
14. Sepals uncinate-re curved; petals yellow or green, naked; 

flowers erect. Bolivia 47. P. platystemon 

14. Sepals straight or nearly so. 
15. Leaf-blades I40 mm wide, broadly oblanceolate ; flowers 
spreading to reflexed, many-ranked, dense; petals 

white. Colombia 144. P. multiflora 

15. Leaf -blades 17-70 mm wide. ~ 

16. Leaves at least partially serrate. 
17. Ovary only I/5 or 1/4 inferior; ovules or seeds 
caudate. Colombia. 
18. Plant stemless; scape~bracts spinose-serrate. 

145. P. kalbreyeri 

18. Plant caulescent for over 1 m; scape-bracts entire. 

146. P. volubilis 
17. Ovary 1/2 to 3/4 inferior; ovules or seeds alate. 

19. Larger leaf -blades evenly serrulate throiighout; 

inflorescence ferruginous-lepidote. Ecuador. 

14. P. harlingii 
19. Larger leaf -blades serrate only toward apeoc. 
20, Plant stemless; ovary 3/4 inferior. Colombia, French 

Guiana, Brazil 15. P. mbiginosa 

20. Plant caulescent; ovary 1/2 inferior. Venezuela. 

16. P. epiphytica 
16. Leaves all entire. 

21, Sepals alate-carinate . Venezuela 147. P. nubigena 

21. Sepals ecarinate. 

22, Leaf -blades not over 20 mm wide, linear-lanceolate, 
long-aciminate . 
23. Pedicels reflexed, then curved-ascending; petals 
appendaged; leaves distichous. Venezuela, 

148. P, meridensis 

23. Pedicels erect; petals naked; leaves not noticeably 

distichous. Brazil 253. P. lancifolia 

22. Leaf -blades 3O-6O mm wide, " 

24. Pedicels 9 am long; flowers strongly secund; leaf- 

blades linear-lanceolate, soon glabrous. Uexico. 

228. P. oaxacana 
24. Pedicels 15-25 nm long; flowers scarcely secund; 
leaf-blades densely brown-flocculose beneath. 

Colombia 149, P. bella 

1. Floral bracts all equaling or exceeding the pedicels. 
25. Sepals 45-75 mm long; leaf-bljwies 25-200 mm wide, 
26. Pedicels 20-60 mm long; leaves entire. Colombia. 
27. Leaf -blades oblanceolate, I40 mm wide.. ,150. P. calophylla 
27. Leaf -blades linear-lanceolate, 25-50 ma wide. 

28. Pedicels to 60 mm long 139. P. laxissima 

28. Pedicels not over 20 mm long 65.''p. haughtii 



18 PHTTOLOGIA Vol. 10, no. 1 

. 26. Pedicels not more than 10 mm long; leaves serrate on the 
petioles . 
29. Floral bracts 150 mm long, exceeding the sepals, the upper 

ones imbricate. Colombia, Ecuador 105. P. elongata 

29. Floral bracts 40-50 mm long, exceeded by the sepals. 
Colombia. 
30. Sepals densely papillose at base; leaf -blade obovate, 

nearly 200 mm wide 151, P. sylvestris 

30. Sepals even or nerved; leaf -blade elliptic or lance- 
elliptic, 90-160 mm wide 152. P. dolichopetala 

25. Sepals 10-35 nm long; leaf -blades 16-180 am wide. 
31. Pedicels U-B mm long at most. 
32. Sepals 10-12 mm long. 
33. Scape-bracts densely imbricate; pedicels reflexed after 

antheais. Ecuador 153. P. reflexiflora 

33* Scape-bracts shorter than the upper intemodes. 
34. Leaf -blades broadly elliptic, 95 mm wide. Ecuador. 

141. P. elliptica 
34. Leaf -blades linea]>-lanceolate, about 40 mm wide. 

Colombia 154. P. semaphora 

32. Sepals 15-34 mm long. ~ 

35. Ovary 3/4 inferior; ovules or seeds alate; floral bracts 

deciduous . Colombia 13 . P. guesnelioides 

35. Ovary not more than 1/2 inferior. 
36. Floral bracts much exceeding the lower flowers; leaves 
unknown, but probably not petlolate, Guatemala. 

216. P. puberula 
36. Floral bracts all exceeded by the flowers. *" 
37. Leaves wholly entire. 
38. Leaf -blades 30-40 mm wide; scape-bracts imbricate. 
Colombia. 
39. Posterior sepals strongly carinate, 28 mm long. 

155. P. macarenensis 

39. Posterior and anterior sepals ecarinate, 20 mm long. 

156. P. guaritermae 
38. Leaf-blades I5O-I8O mm wide. " 

40. Flowers spreading at anthesis; leaf -blades densely 

white-lepidote beneath. Brazil... .246. P. undulata 
/iO«. Flowers erect or divergent at anthesis; leaf -blades 

glabrous . Peru 157. P. calatheoides 

37. Leaves spinose-serrate on the petioles or on the 
bladeless sheaths (P. nobilis ) . 
41. Scape-bracts mostly or all spinose-serrate. 
42. Blades lanceolate, 80 mm wide. Colombia. 

39. P. peetinata 
42. Blades linear^-lanceolate, 22 mm wide. Ecuador. 

158. P. sodiroi 
41. Scape-bracts entire. 

43* Petioles entire; blades 23 mm wide. Ecuador. 

159. P. nobilis 
43 • Petioles serrate; blades 60-80 mm wide, Colombia. 

44. Floral bracts broadly elliptic, apiculate. 



196U Smith, Notes on Bromeliaceae 19 

160. P. adscendena 
UU. Floral bracts linear-lanceolate, acuminate. 

161. P. sneldemii 
31. Pedicels 9-30 nun long. 

45. Ovary more than l/2 inferior; ovules or seeds alate 
(unknown in P. leprieurii ) . 
/».6. Sepals to 35 nan long. 
47. Sepals oblong, broadly acute; inflorescence dense. 

Colombia 17. P. cuatrecasana 

47. Sepals triangular-ovate, acuminate; inflorescence lax. 

Ecuador 14. P . harlingii 

46. Sepals not more than 24 mm long. 

48. Blades brown-lepidote beneath; sepals ecarinate. 

Colombia, French Guiana, Brazil 15. P. rubiginosa 

48. Blades whitish-lepidote beneath; sepals carinate. 

French Guiana 18. P. leprieurii 

45. Ovary not more than 1/2 inferior; ovules or seeds alate or 
caudate . 
49. Leaves entire or minutely serrulate toward apex (P. 
cyanopetala ) . 
50. Blades 80-180 mm wide. 
51. Scape-bracts much shorter than the internodes; blades 
180 nm wide, densely white-lepidote beneath. Brazil. 

246. P. undulata 

51. Scape-bracts exceeding the internodes; blades 80 mm 

wide, subglabrous. Peru 162. P. cyanopetala 

50. Blades 20-50 mm wide. "" 

52. Flowers all secund; sepals alate-carinate; plant 2 dm 

high. Mexico 229. P. secundiflora 

52. Flowers not secund; sepals not alate; plants 4-15 dm 
high. 
53. Floral bracts coriaceous; petals yellowish white; 
leaf-blades white-furfuraceous beneath. Mexico, 

Guatemala, British Honduras 190. P. recumcata 

53. Floral bracts thin; petals red or purple; leaf-blades 

soon glabrous. Coltxnbia I63. P. brachyspenna 

49. Leaves serrate on the petioles or bases of the blades. 
54. Leaf -blades 16-22 ran wide, weakly petiolate; sepals 
obtuse; ovary only I/6 inferior. Peru, Bolivia, 

Brazil 60. P. subpetiolata 

54. Leaf -blades 30-130 nm wide, distinctly petiolate; sepals 
acute or acuminate, rarely rounded (sometimes in P. 
sprucei ) . 
55. Ovary about 1/2 inferior; ovules or seeds alate. 
56. Inflorescence prostrate on the ground with the flowers 
upwardly secund; leaf-blades entire; pedicels to 10 

ram long. Colombia, Peru 19. P. corallina 

56. Inflorescence erect, the flowers scarcely secund; 

leaf-blades serrulate toward apex; pedicels 10-30 mm 
long. 



20 PHYTOLOGIA Vol. 10, no, 1 

57. Leaf -blades lanceolate or oblanceolate , 30-90 am 

wide, soon glabrous; sepals oblong, broadly acute 
or rounded; pedicels 10-18 mm long. Colombia, 

Peru, Brazil 20. P. sprucei 

57. Leaf -blades linear-lanceolate, 35 mm wide, densely 
iriiite-lepidote beneath; sepals linear-lanceolate, 
acute; pedicels to 30 nun long. Venesuela. 

16. P. epiphytica 
55. Ovary only 1/4-1/5 inferior; ovules or seeds caudate. 
58. Sepals 12 mm long; leaf -blades 90 mm nide, covered 
beneath with a brown membrane of scales; scape- 
bracts entire. Colombia 164. P. aiw^1<« 

58. Sepals 20-25 mm long. " 

59. Leaf -blades llneaivlanceolate, attenuate, 40-70 mm 
wide; petals 65 mm long. Panama, Colombia. 

145. P. kalbreyeri 
59. Leaf-blades elliptic or obovate, abruptly acute, I30 
nm wide; petals 100 mm long. Ecuador. 

165. P. palmoides 

SDBKET V 

1. Ovary distinctly more than half inferior; ovules or seeds 
alate. 
2. Petals lepidote, naked; pedicels 3-5 nm long. Colombia. 

21. P. turbinella 
2. Petals glabrous. 
3. Sepals 40 mm long; petals naked; flowers subsessile. 

Venezuela 22. P. mapuirei 

3. Sepals 15-35 mni long; petals appendaged. 
4. Flowers subsessile. 
5. Inflorescence lax; flowers spreading; ovary wholly 

inferior. Brazil 23. P. anoinala 

5. Inflorescence dense; flowers curved ascending; ovary 3/4 

inferior. Colombia 13. P. guesnelioides 

4. Flowars distinctly pedicellate for 5-25 nm. 

6. Sepals 35 sun. long; pedicels 25 mm long. Ecuador. 

14. P. harlingii 
6. Sepals 15-27 mm long; pedicels 5-10 mm long. 
7. Leaf-blades sublinear, 6-I3 mm wide; sepals acute. 

Colombia, Venezuela, Guiana, Brazil.. 24. P. caricifolla 
7. Leaf -blades linear^lanceolate, 20 mm wide; sepals 

rounded. Guiana 25. P. incamata 

1. Ovary 1/2 inferior to almost completely superior. 
8. Sepals 30-60 mm long. 
9. Posterior sepals alate-carinate. Peru. 

10. Inflorescence dense; flowers subsessile; sepals 30 mm long 

91. P. sandemanii 
10. Inflorescence lax at base or throughout; flowers pedicel- 
late for 10 mm; sepals 46 mm long I66. P. eximia 

9. Posterior (and anterior) sepals not more than angled- 
carinate. 



196U Smith, Notes on Bromeliaceae 21 

. 11. Ovxiles or seeds alate. 

12, Floral bracts laclniate-spinosej sepals 60 nm long. 

Bolivia, Argentina 40. P. mirabilis 

12. Floral bracts entire; sepals 33-40 oim long. 
13. Petals red, appendaged; leaf -blades spinose-serrate only 

toward base. Colombia, Brazil ,26. P. uaupensis 

13, Petals pale green, naked; letif-blades spinose throughout 

Venezuela 3, P. wurdackii 

11. Ovules or seeds caudate. 
14. Sepals obtuse, 20-30 mm long; leaves subpetiolate . Peni, 

Bolivia, Brazil 60. P. subpetiolata 

14, Sepals acute or acuminate. 
15 • Petals naked; inflorescence lax; flowers secund- 
spreadixig; leaves incompletely known. Mexico. 

209. P. leprosa 
15, Petals appendaged. 
16. Flowers spreading or reflexedj floral bracts broadly 
lanceolate, acute, 35 mm long, exceeding the pedi- 
cels; axis 1 cm in diameter, Bolivia., 119. P. crassa 
16, Flowers erect or suberect. 
17 < Floral bracts to 18 mm long, equaling or exceeding the 
pedicels; inflorescence lax. Panama., 230. P. camea 
17. Floral bracts to 70 mm long, shorter than to exceeding 
the sep>als; inflorescence dense. Lesser Antilles. 

242, P, albucifolia 
8. Sepals 8-29 ma long. " 

18. Petals naked, 
19, Sepals rounded, 8-12 mm long; petals green to pale yellow 
or white, 
20. Inflorescence 5-flowered; flowers secund; s^als densely 
brown-lepidote. Peru 35 . P, viridis 

20. Inflorescence densely many- flowered; flowers not secund; 

sepals glabrous. Colombia, Ecuador, Bolivia. 

167. P. trianae 
19. Sepals acute or acuminate, 13-27 mm long. 

21, Floral bracts much exceeding the lower flowers; leaf- 

blades imknown but probably not persistent. 

Guatemala 216. P. pubenila 

21, Floral bracts all exceeded by the flowers, 
22, Leaf -blades serrate throughout; ovules or seeds alate. 
Venezuela, 
23, Pedicels stout, 6 nm long; leaf -blades 90 an wide. 

1. P. agavifolia 

23. Pedicels slender, 25 mm long; leaf-blades 12 mm wide. 

27. P. ctenophylla 
22. Leaf -blades only partially serrate (unknown in P. 

platystemon ) or the ovules or seeds caudate (unknown 
in P. cuzcoensis ) or usually both characters combined, 

24. Petals yellow, green, or cream; floral bracts shorter 

than the pedicels. 
25. Sepals uncinate-recuirved; floral bracts 15-20 mm long, 
Bolivia 47 . P . platystemon 



22 PHTTOLOGIA Vol. 10, no. 1 

25. Sepals straight; floral bracts to 12 mm long. Costa 

Rica 77. P. halophila 

2/». Petals red or violet. 

26. Scape-bracts all exceeding the intemodes; petals 

light brownish violet. Ecuador. ..128. P. violas cens 
26. Scape-bracts shorter than the upper intemodes. 
27. Leaf -blades oblong, acute; inflorescence glabrous; 

sepals 21 mm long. Ecuador. .. .168. P. oblongifolia 
27. Leaf-blades lanceolate or linear-lanceolate, acumi- 
nate; inflorescence more or less vestite; sepals 
13-17 mm long. 
28. Leaves trimorphic, some reduced to dark broad entire 

sheaths. Peru 74. P. cuzcoensls 

28. Leaves dimorphic or all alike, none reduced to con- 
spicuous sheaths. Colcanbia 75* £• schultzei 

18. Petals appendaged. 
29. Leaf-blades (or the spiniform ones if dimorphic) spinose- 
serrate throughout or entire only at the extreme apex 
(P. lanuginosa ). 
30. Leaf -blades 90 mm wide; pedicels stout, 6 mm long; 

flowers recurved. Venezuela 1. P. agavifolia 

30. Leaf-blades 4-32 mm wide. 
31. Pedicels 15-30 mm long; leaf-blades 10-32 ram wide. 
32. Ovules or seeds with annular appendages, alate. 
33. Sepals to 28 mm long; plant stemless; floral bracts 
about equaling the pedicels. Venezuela. 

28. P. fills pina 

33. Sepals not over 15 mm long; plant with an erect 15 cm 

high stem; floral bracts much shorter than the upper 

pedicels. Mexico, Guatemala 42. P. punicea 

32. Ovules or seeds with caudate appendages though the 
apical part sometimes broad (P. platyphvlla ). 

34. Petals pale yellow; sepals 15-20 mm long. Mexico. 

231. P. xant ho calyx 
34. Petals red or violet; sepals to 25 mm long. 
35. Floral bracts equaling or exceeding the pedicels; 

petals pale violet; leaf-blades dimorphic. Peru. 

169. P. lanuginosa 
35. Floral bracts much shorter than the upper pedicels; 

petals red; leaf -blades usually all alike. Jamaica. 

36. Leaf -blades 10-15 mm wide 233. P. bromeliifolia 

36. Leaf-blades to 32 mm wide 238. P. platyphylla 

31. Pedicels 3-11 mm long. 
37. Sepals subalate-carinate; pedicels 8 mm long; leaf- 
blades 8 mm wide. West Indies? 243. P. g^ymiana 

37. Sepals not at all alate. 
38. Petals bright yellow to cream. 
39. Floral bracts much exceeding the pedicels; leaf- 
blades to 24 mm wide. Paraguay. 

170. P. paraguayensis 
39. Floral bracts shorter than the pedicels; leaf-blades 
8-18 mm wide. Hispaniola 232. P. samuelssonii 



196U Smith, Notes on Bromeliaceae 23 

38. Petals red. 
ZfO. Leaves all alike. 
i*l. Leaf -blades 10-20 mm wide; flowering plant 1-2 m 
high. Puerto Rico, Lesser Antilles. 

234. P. angustifolia 
l^X• Leaf-blades 4 mm wide; flowering plant 55 an high. 

Dominican Republic 244. P. elizabethae 

40. Leaves of the fertile rosette dimorphic. 

235. P. fuertesii 
29. Leaf -blades spinose only at extreme base or apex, or th« 
sheaths only. 
42. Petal-appendages a pair of small auricles. West Africa. 

261. P. feliciana 
42. Petal-appendage a single scale the width of the petal. 
43. Scape-bracts equaling or exceeding aH of the intemodes 
44. Lower flortil bracts nearly equaling to exceeding the 
sepals; inflorescence dense. Lesser Antilles. 
45. Floral bracts broadly ovate 241. P. spicata 

45. Floral bracts lanceolate 242. P. albucifolia 

44. Lower floral bracts much shorter than the sepals; 

inflorescence subdense to lax. 

46. Petals pale yellow; pedicels 15-20 mm long. Mexico. 

231. P. xanthocalyx 
46. Petals red; pedicels 4-15 nim long. 
47. Leaves dilated above the sheath, all alike. 

Venezuela 73. P. moritziana 

47. Leaves not dilated above the sheath. 
U8. Leaf -blades covered beneath with a merabrsme of white 
scales. Puerto Rico (?), Lesser Antilles. 

240. P. latifolia 
48. Leaf -blades sparsely lepidote or glabrous beneath. 
49. Inflorescence subdense, raany-flowered. Colombia. 

171. P. macrobotrya 
49. Inflorescence lax, few-flowered. Venezuela. 

172. P. orchidifolia 
43. Scape-bracts distinctly shorter than the upper 
intemodes . 
50. Pedicels 15-30 mm long; sepals abruptly acute or 
rounded. 
51. Sepals 13 mm long, oblong; leaf-blades sparsely 

lepidote toward base. Brazil 247. P. egleri 

51. Sepals 20-30 mm long, narrowly triangular; leaf-blades 
densely white-lepidote beneath. 
52. Ovary 1/2 superior; leaves all alike. Jamaica. 

238. P. platyphyUa 
52. Ovary 5/6 superior; leaves dimorphic. Peinx, Bolivia, 

Brazil 60. P. subpetiolata 

50. Pedicels 4-10 mm long. ~ 

53. Leaf*-blades not more than 11 mm wide. 

54. Sepals 14-16 mm long. Cuba 239. P. cubensis 

54. Sepals 24 mm long. Brazil 32. P. platypetala 

53. Leaf-blades 15-30 nm wide. " 



2U PHTTOLOQIA Vol. 10, no, 1 

, 55. Inflorescence densely white-tomentose; leaf -blades 
inperfectly known, probably not persistent. Peru. 

122. P. pavonll 
55. Inflorescence sparsely flocculose to glabrous. 
56. Leaf-blades lanceolate, 25-50 cm long, 15-25 nm id.de 
57. Floral bracts elliptic, much shorter than the 

pedicels; leaf -blades densely appressed-lepidote 
beneath. Venezuela 173. P. fendleri 

57. Floral bracts lanceolate, acuminate, from shorter 

to longer than the pedicels; leaf-blades sparsely 
lepidote to glabrous beneath. Colombia. 

75. P. schultzei 
56. Leaf -blades linear, 50-100 cm long, 15-20 mm wide, 
Dominican Republic. 

58. Leaves all alike 237. P. .limenezii 

58. Leaves dimorphic 245* P* doningensis 

SUBK£I VI 

1. Ovary distinctly more than 1/2 inferior. 
2. Seeds or ovules alatej inflorescence minutely white-lepidote; 
ovary almost vrtiolly inferior. Col(»bia, Venezuela, Guiana, 

Brazil 24. P. earicifolia 

2. Seeds or ovules caudate; inflorescence densely pale- 

flocculose or arachnoid; ovary not more than 2/3 inferior. 
3. Sepals 18 imn long, carinate toward base. Ecuador. 

174. P. unilateralls 

3. Sepals 25 mm long, ecarinate. Brazil 248. P. ulei 

1. Ovary not more than 1/2 inferior. 
4. Sepals 34-45 nm long; petals appendaged. 
5. Pedicels 5O-6O mm long; leaf -blades 30 mm wide. Colombia. 
6. Sepals densely ferruginotis-tomentose 139. P. laxissiaa 

6. Sepals glabrous 140. P. longipes 

5. Pedicels 6-25 aua long, 

7. Axis of the inflorescence 1 cm in diameter; flowers 

spreading or reflexed. Bolivia 120. P. crassa 

7. Axis of the inflorescence relatively slender; flowers 
erect to divergent. 
8. Petals greenish white; floral bracts broadly ovate; sepals 
transversely ridged at junction with ovary. Colombia, 

41. P. CO Stat a 
8, Petals red, 
9. Floral bracts broadly ovate, slightly exceeding the 
pedicels; petals lepidote. Colombia. 

175. P. lepidopetalon 
9, Floral bracts lanceolate; petals glabrous. 
10. Inflorescence dense toward apex; floral bracts to 70 ma 

long. Lesser Antilles 242. P. albueifolia 

10. Inflorescence lax throughoat; floral bracts to 18 mm 

long, Panama 230. P. camea 

4, Sepals 9-30 mm long or laore (P, albueifolia ) . 
11. Petals naked. 



196U Smith, Notes on Bromellaceae 2$ 

. 12. Sepals rounded at apex, blunt or aplculate. 
13. Leaf-bladea 4-20 mn wide. 
14. Scape-bracts eocceeding the intemodes. 
15. Lower floral bracts serrulate; sepals oblong^ 21 mm 

long. Brazil 31. P. Ilmae 

15. Lower floral bracts entire; sepals 9 nm long, French 

Guiana 176. P. pusilla 

14. Scape-bracts distinctly shorter than the upper 
intemodes . 

16. Floral bracts exceeding the 5 on long pedicels. Peru. 

131. P. straainea 
16, Floral bracts much shorter than the 20 am long 

pedicels. Venezuela 29. P. .juncoides 

13. Leaf -blades 3O-6O am wide. ~ 

17. Floral bracts exceeding the pedicels. 
18. Sepals broadly elliptic, 26 mm long; leaf-blades 60 mm 
wide. Guiana, Venezuela, Colombia, Central America. 

90. P. maidifolia 
18. Sepals oblong; leaf -blades not over 35 mm wide. 
19. Inflorescence dense, many-flcrwered ; floral bracts very 

narixjwly triangular. Venezuela 177. P. tympani 

19. Inflorescence lax, few-flowered; floral bracts ovate. 

Colombia 178. P. occidentalis 

17. Florad bracts at most equaling the 8-9 mm long pedicels. 
20. Flowers erect; florad bracts ligulate, shorter than the 
pedicels, flaccid. Peru, Venezuela. 

179. P. brevicalycina 
20. Flowers spreading; floral bracts narrowly triangular, 
about equaling the lower pedicels. Colombia. 

180. P. petraea 
12. Sepals acute or acuminate. 
21. Floral bracts 4-8 cm long, exceeding the lower floweira. 
22. Pedicels slender, 10 mm long. Guatemala (?). 

191. P. longebraeteata 
22. Pedicels stout, 3-5 mm long. 
23. Sepals ecarinate; leaf -blades unknown but probably not 

persistent. Guatemala 216. P. puberula 

23. Sepals obtusely carina te. Brazil. 

254. P. encholirioides 
21. Floral bracts all exceeded by the flowers. 

24. Sepals alate-carinate. Brazil 255. P. carinata 

24. Sepals not at all alatB« 
25. Floral bracts equaling or exceeding the pedicels. 
26. Petals sparsely floccose at apex; sepals 30 mm long; 
inflorescence densely lepidote. Mexico. 

209. P. leprosa 
26. Petals glabrous; sepals 22-28 nm long. 
27. Flowering shoot about equaling to exceeding the 

leaves. Brazil 256. P. flammea 

27. Flowering shoot little more than half as high as the 

leaves . Colombia 181 . P . andreana 

25. Floral bracts all distinctly shorter than the pedicels. 



26 PHTTOLOQIA Vol. 10, no. 1 

28. Sepals uncinate-re curved ; petals yellow or green. 

Bolivia kl. P. platystemon 

28. Sepals straight. Brazil. ~ 

29. Petals regular, spirally recurving at anthesis. 
30. Rhachis of the inflorescence glabrous; sepals 15-18 
mm long; petals white to yellow. . .257. P. albifloa 

30. Rhachis of the inflorescence f urf uraceoua ; sepals 

19-22 mm long; petals red 258. P. staminea 

29. Petals zygomorphic at least by position, not spirally 
recurving at anthesis. 

31. Base of the rosette not thickened; leaves not over 

15 era long 259. P. beycalema 

31. Base of the rosette bulbous-thickened; leaves 

usually much more than 15 cm long.. 25 6. P. flammea 
11. Petals appendaged. " 

32. Sepals rounded at apeoc, blunt or s^iculate. 
33. Scape-bracts equaling or exceeding the intemodes. 
34. Inflorescence dense, especially toward apex; floral 
bracts broadly ovate. Lesser Antilles. 

241. P. spicata 

34. Inflorescence wholly lax; floral bracts linear- 

lanceolate. Mexico 231. P. xanthocalyx 

33. Scape-bracts distinctly shorter than the upper intemodes 

35. Petals red. 

36. Pedicels 20 mm long, much exceeding all the floral 

bracts. Colombia 182. P. giMbbiana 

36. Pedicels only 5 nm long, exceeding only the upper 

floral bracts. Dominican Republic. 237. P. .limenegji 
35. Petals yellow or white. 

37. Sepals 9 mm long, elliptic; ovary over 3/4 superior; 

petal-appendage of 2 vertically attached auricles. 

Venezuela (?) 183, P. caulescens 

37. Sepals 15 mm long, linear; ovary ca. 1/2 superior. 

Brazil 33. P. bradei 

32. Sepals acute or acuminate. 
38. Scape-bracts equaling or exceeding the intemodes. 
39. Scape very short or none; inflorescence globose. 

Mexico 189. P. tabullformis 

39. Scape well developed. 
40. Floral bracts 6-8 cm long, from nearly equaling to much 
exceeding the narrowly triangular sepals. 
41. Inflorescence subcorymbose, few- flowered. 

Guatemala (?) .191. P. longebracteata 

41. Inflorescence elongate, many-flowered. Lesser 

Antilles 242. P. albucifolia 

40. Floral bracts smaller, mostly shorter than the sepals. 

42. Petals red. Puerto Rico, Lesser Antilles. 

240. P. latifolia 
42. Petals yellow or white. 
43. Ovary more than 3/4 superior; ovules or seeds 

caudate. Brazil 260. P. auaveolena 

43. Ovary 1/2 superior. Mexico. 



196U Smith, Notes on Bromeliaceae 27 

44. Ovules alate; scape-bracts equaling the upper 

int emodes 43 . P . anblyosperma 

44. Ovules caudate; scape-bracts exceeding all the 

int emodes 231. P. xanthocalyx 

38. Scape-bracts distinctly shorter than the upper int emodes 
45' Sepals 16 bbi long; inflorescence dense, many- flowered. 

Venezuela 184. P. microcalyx 

45. Sepals 23-30 nun long. 
46. Posterior sepals broadly alate-carinate ; floral bracts 
broadly ovate; flowers subsessile, Peru. 

91. P. sandemanii 
46. Posterior (and anterior) sepals not at all alate, at 
Bost obtusely carinate. 
47. Floral bracts exceeding the pedicels, nairrowly 
triangular or linear-lanceolate, acuminate. 
48. Ovules subalate; ovary more than 1/2 superior. 

Brazil 32. P. platypetala 

48. Ovtiles long-caudate; ovary more than 4/5 superior; 

leaves doubtfully persistent. Ecuador. 

122. P. pavonii 
47. Floral bracts shorter than the pedicels. 

49. Petals red; floral bracts ovate, acuminate. Mexico. 

228. P. oaxacana 
49. Petals white; floral bracts elliptic, acute. 

Colombia 185 * P* .lohannis 



Subgenus PEPINIA 
(including subgenus Schweideleria ) 

Amazon-Orinoco Basin 

1. P. AGAVIFOLIA L. B. Smith, Mem. New York Bot. Card. 9:287, 
fig. 18. 1957. Venezuela. 

2. P. NUDA Baker, Joum. Bot. 19:269. 1881. British Guiana, 
Surinam. 

3. P. WURDACKII L. B. Smith, Mem. New York Bot. Gard. 10, no. 
2:17, fig. 4. I960. Venezuela. 

4. P. MITUEJISIS L. B. Smith in R. E. Schultes, PI. Austro-Am. 
XI, Rhodora 65:7, fig. 7. 1963 . Colombia. 

5. P. KUNHARDTIANA L. B. Smith, Mem. New York Bot. Gard. 9: 
289, fig. 21. 1957. Venezuela. 

6. P. BULBOSA L. B. Smith, Phytologia 5:178, pi. 1, figs. 7- 
10. 1955. Venezuela, Colombia. 

7. P. HELIOPHILA L. B. Smith, Caldasia fl], no. 5:8, fig. 
1942. Colombia. 

8. P. ARMATA Maury, Joum. de Bot. 3:270, fig. I3. 1889. P. 
cinerea L. B. Smith, Fieldiana Bot, 28, no. 1:149, fig. 23 d-f, 
1951. Venezuela. 

9. P. BREWERI L. B. Smith, Phytologia 9:246, pi. 2, figs. 1, 
2. 1963. Venezuela. 

10. P. GEYSKESII L. B. Smith, Act. Bot. Neerlandica 5:90, fig. 



28 PHITOLOOIA Vol. 10, no. 1 

1. 1956. Surinam. 

11. P. PRDINOSA H.B.K. Nov. Gen. & Sp. 1:295. 1816. Venezuela, 

12. P. PATENTIFLORA L. B. Smith, Contr. Gray Herb. 127:18, pi. 
1, fig. 4. 1939. 

Var. a. PATENTIFLORA. Plant 5-10 dm high; leaf -blades scarce- 
ly or not at all narrowed toward base, 3-I5 nm wide, serrate for 
most of their length; sepals 15-20 am long. Venezuela, Brazil. 

Var. b. SUBINTEGHA L. B. Smith, Bot. Mue. Leafl. Harvard 17: 
68. 1955* Leaf -blades distinctly narrowed toward base, to 20 mm 
wide, serrate only at extreme base; sepals 13-18 mm long. 
Colombia. 

Var. c. UACRANTHA L. B. Smith, Bot. Mus. Leafl. Harvard 17:68. 
1955. Plant 2 m high; leaf -blades distinctly narrowed toward 
base, to 25 mm wide, serrate only at extreme base; sepals 25 m 
long. Colombia. 

Var. d. ARllATA L. B. Smith, Uera. New York Bot. Gard. 10, no. 
5:37. 1964. Flowering 2 m high; leaves 14 dm long; 

blades 4 cm wide, slightly narrowed toward base, serrulate 
throughoixt; sepals to 27 am long. Venezuela. 

13. P. QUESNELIOIDES L. B. Smith, Contr. U. S. Nat. Herb. 29: 
313, fig. 33. 1949. Colombia. 

14. P. HARLINGII L. B. Smith, Phytologia 8:11, pi. 1, figs. 
18-20. 1961. Ecuador. 

15. P. RUBIGINOSA (Brongn. ex £. Morr.) Baker, Handb. Bromel. 
116. 1889. 

Var. a. RUBIGINOSA. Melinonia rubiginosa Brongn. ex E. Morr. 
Cat. Bromel. Jard. Liege 11. 1673. Leaves serrulate, at least 
toward apex; floral bracts exceeding the pedicels. Colombia, 
French Guiana. 

Var. b. AMAZONICA (Baker) L. B. Smith, Phytologia 7:109. I960. 
P. amazonica Baker, Handb. Bromel. 117. 1889. Leaves serrulate, 
at least toward apex; floral bracts shorter than the pedicels. 
Brazil. 

Var. c. INTEGRA L. B. Smith, Phytologia 7:110. I960. Leaves 
entire; floral bracts exceeding the lower pedicels, shorter than 
the upper, Brazil. 

16. P. EPIPHYTICA L. B. Smith, Mem. New York Bot. Gard. 9:289, 
fig. 20. 1957. Venezuela. 

17. P. CUATRECASANA L. B. Smith, Caldasia t l3 , no. 4:16, fig. 
1942. ColoBbia. 

18. P. (subgenus ?) LEPRIEURII Baker, Handb. Bromel. 117. 
1889. French Guiana. 

19. P. CORALLINA Linden & Andre, Belg. Hortic. 23:112. 1873. 
Var. a. CORALLINA. Leaves to 10 cm wide; inflorescence coral- 
red. Colombia, Peru. 

Var. b. VIRIDIS L. B. Smith, Bot. Mus. Leafl. Harvard 17:67. 
1955. Leaves only 3 cm wide; inflorescence yellow-green. 
Colombia. 

20. P. SPRUCEI Baker, Joum. Bot. 19:303. 1881. Colombia, 
Peru, Brazil. 

21. P. TURBINELLA L. B. Smith, Caldasia Cll , no. 4:17, fig. 
5. 1942. Colcmbia. 



196U Smith, Notes on Brcmellaceae 29 

22. P. MAGUmEI L. B. Smith, Men. New lork Bot. Gard. 10, no. 
2:17, fig. 5. I960. Venezuela. 

23. P. ANOMALA Hoehne, Conm. Linh. Telegr. Estrat. Uatto 
Grosao Publ.C47j, Annexo 5, Bot. pt. 9:9, pi. 161. 1919. Brazil. 

2if. P. CARICIFOLU Mart, ex Schult. f . In R. & S. Syat. 7, pt. 
2:12i»2. 1830. 

Var. a. CARICIFOLIA. P. kegeliana Schlecht, Linnaea 2i*:664. 
1851. P. pauciflora Baker, Joum. Bot. 19:230. 1881. P. sub- 
.juncta Baker, Handb, Brraiel. 116. 1889. Floral bracts exceeding 
the loiter pedicels, sometimes equaling the ovaries; sepals 15 mm 
long, green. Guiana, Venezuela, Colombia, Brazil. 

Var. b. MACRANTHA L. B. Smith, Mem. New York Bot. Gard. 10, 
no. 5:37. 1964. Floral bracts equaling the lowest 

pedicels, shorter than the remainder; sepals acute, 27 mm long, 
orange-red . Venezuela . 

25. P. INCARNATA (Brongn. ax E. Morr.) Baker, Harelb. Bromel. 
116. 1889. Melinonia incamata Brongn. ex E. Morr. Cat. Bromel. 
Jard. Liege for 1873:11. 1873. Pepinia Incarnata E. Morr. ex 
Baker, Handb. Bromel, 116. 1889, nomen. Pitcaimia semi.luncta 
Baker, Handb. Brtmel. 117. 1889. Guiana. 

26. P. UAUPENSIS Baker, Handb. Bromel. 93. 1889. Colombia, 
Brazil. 

27. P. CTENOPHYLLA L. B. Smith in Gleason & Killip, Brittonia 
3:161, fig. e-i. 1939. Venezuela. 

28. P. FILISPINA L. B. Smith, Men. New York Bot, Gard. 9:289, 
fig, 19. 1957. Venezuela. 

29. P. JUNCOIDES L, B, Smith, Contr, Gray Herb. 161:33, pl. 4, 
figs. 2, 3. 1946. Venezuela. 

Planalto of Brazil 

30. P, ENSIFOLIA Mez in Mart. Fl. Bras. 3, pt. 3:436, pl. 82. 
1894. 

31. P. LIMAE L. B. Smith, Phytologia 7:254, pl. 1, figs. 9, 
10. I960. 

32. P, PLATYPETALA Mez in Mart, Fl, Bras. 3, pt, 3:438. 1894. 

33. P. BRADEI Markgraf, Notizblatt 15:215. 1940. 

Andean South America 

34. P. FERRUGINEA R. & P. Fl. Peruv. 3:36. 1802. Pourretia 
ferruginea Spreng. Syst, Veg. 2:23. 1825. Pitcaimia asterotri- 
cha Poepp. & Endl. Nov. Gen. 2:42, pl. 158. 1838. Puya grandi - 
flora Hook. Bot. Mag. 87:pl. 5234. 1861. Pitcaimia conaimilis 
Baker, Joxim. Bot. 19:226, 266. 1881. Puya echlnotricha Andre. 
Rev. Hortic. 60:565. 1888. Pitcaimia echlnotricha Baker, Handb. 
Bromel. 119. 1889. P. weberbaueri Mez, Bull. Herb. Boiss. ser. 
2, 4:627. 1904. P. cotahuasiana Harms, Notizblatt 10:788. 1929. 
P. latibracteata Harms, 1. c. P. herrerae Harms, op. c. 789. P. 
imperialis Harms, op. c. 790. P. lareelana Harms, op. c. 11:58. 
1930. Ecuador, Peru, Bolivia. " 

35. P. VIRIDIS Mez, Fedde Rep. Spec. Nov. 16:8. 1919. Peru. 



30 PHTTOLOGIA Vol. 10, no. 1 

36. P. KULIPIANA L. B. Smith, Conbr. U. S. Nat. Herb. 29:308, 
fig. 29. 1949. Colombia. 

37. P. NANA (Wittm.) L. B. Smith, Phytologia 7:5. 1959. Puya 
nana Wittm. Mededeel Rijks Herb. 29:85. 1916. Bolivia. 

38. P. ALBORUBRA Baker, Handb. Bromel. 102. 1889. Colombia. 

39. P. PECTINATA L. B. Smith, Contr. Gray Herb. 98:9, pi. 3, 
figs. 6, 7. 1932. Colombia. 

AC. P. MIRABILIS Mez, Fedde Rep. Nov. Spec, 3:6. 1906. 

Var. a. MIRABILIS. Leaf -blades 60 cm long, 15 ram wide, suben- 
tire to spinosej flowering shoot to 15 dm high} inflorescence 30 
cm long. Bolivia, Argentina. 

Var. b. TUCUMANA Castellanos, Anal. Mus. Nac. Hist. Nat. 
Buenos Aires 36:53, pl. 5. 1929. Leaf -blades 22 cm long, 8 mm 
wide, strongly spinose; flowering shoot 3-4 dm high; inflores- 
cence 8-10 cm long, Argentina, 

41. P. COSTATA L. B. Smith, Caldasia 5:7, fig. 1948. Colombia. 

Mexico, Central America 

42. P. PUNICEA Scheidw. Bull. Acad. Brux. 9, pt. 1:25. 1842. 
P. .lacksoni Hook. Bot. Mag. 76:pl. 4540. 1850. Pepinia punicea 
Brongn. ex Andre, HI, Hortic. 17:33. 1870. Lamproconus .jacksoni 
Lem. ex Hemsl. Biol, Centr, Am. 3:316. 1884, Mexico, Guatemala, 

43. P. AMBLIOSPERMA L. B. Smith, Contr. Gray Herb. 117:23, pl. 
2, figs, 4-7. 1937. Mexico, 

Subgenus PITCAIRNIA 

Andean South America 

44. P. ECHINATA Hook. Bot. Mag. 79:pl. 4709. 1853. Colombia. 
Var. a. ECHINATA. Sepals and base of ovary densely and 

coarsely echinate with processes 1-2 mm lon^; petals white. 

Var. b. SUBLAEVIS L. B. Smith, Contr. U. S. Nat. Herb. 29:304. 
1949. Sepals and base of ovary minutely and sparsely stellate, 
nearly even. 

Var. c. VALLENSIS L. B, Smith, Phytologia 4:378. 1953. Petals 
rose-orange; stamens exserted. 

45. P. TUBERCULATA L. B. Smith, Fieldiana Bot. 28, no. 1:149, 
fig. 22 c, 1951, Venezuela, 

46. P. PUYOmES L. B. Smith, Field Mus. Bot. 11:147. 1936; 
Contr. Gray Herb, 114:7, pl. 1, figs. 4, 5. 1936. Peru. 

47. P. PLATYSraiON Mez in DC. Mon. Phan. 9:421, 1896, Bolivia. 

48. P, PULVERULENTA R, & P, Fl, Peruv, 3:36, pl. 259. 1802, 
Orthopetalum pulverulentum Beer, Brom, 71, 1857. Peru. 

49. P. PANICULATA (R. & P.) R. & P. Fl. Peruv. 3:36, pl. 260, 
1802, Pourretia paniculata R, & P. Syst. Veg. 1:81. 1798, Pit- 
cairnia longifolia Hook, Bot, Mag, 80:pl, 4775. 1854. P. excelsa 
E. Morr, Belg, Hortic, 25:381. 1875. P. fruticetorum Mea, Fedde 
Rep. Nov. Spec. 3:4. 1906. P. biattenuata Rusby, Bull. New York 
Bot. Gard. 4:457. 1907. Peru, Bolivia. 

50. P. (subgenus ?) VERRUCOSA L. B. Smith, Phytologia 4:380, 



196U Smith, Notes on Bromeliaceae 31 

pi. 2, figs. l-i+. 1953. Colombia. 

51. P. RUIZIANA Mez in DC. Mon. Phan. 9:420. 1896. Peru. 

52. P. DENDROIDEA Andre, Enum. Bromel. 4. Dec. 13, 1888; Rev. 
Hortic. 60:564. Dec. 16, 1888. Colombia, Ecuador. 

53. P. RIGIDA Uez, Bull. Herb. Boiss. ser. 2, 4:625. 1904. 
Peru. 

54. P. (subgenus ?) LECHLERI Baker, Joum. Bot. 19:269. 1881. 
Peru. 

55. P. INERMIS (Meyer) Meyer ex Schult. f . in R. & S. Syst. 
Veg. 7:1238. 1830. Peru. 

Var. a. INERMIS. Pourretia inermis Meyer in Presl, Rel. 
Henk. 1:123, pi. 23. 1827. Orthopetalua inerme Beer, Bromel. 72. 
1857. Flowering to 5 dm high; leaves serrulate above the base; 
petals scarlet, appendaged. 

Var. b. FLAVA L. B. Smith, Phytologia 5:46. 1954. Flowering 
to 8 dm high; leaves serrulate below the abscission line; petals 
yellow, naked. 

56. P. ARENICOU L. B. Smith, Contr. U. S. Nat. Herb. 29:299, 
fig. 21. 1949. Colombia. 

57. P. COMMECTA L. B. Smith, Contr. U. S. Nat. Herb. 29:303. 
1949. P. orgyalis sensu Andre, £num. Bromel. 4. Dec. 13, 1888, 
nomen; Rev. Hortic. 60:564. Dec. 16, 1888, nomen; Andre ex Mez in 
DC. Mon. Phan. 9:412. 1896, non Baker, 1881. Colombia, Ecuador. 

58. P. LEHMANNII Baker, Joum. Bot. 19:273. 1881. P. orgy alia 
Badcer, 1. c. P. aurlculata Mez, Bull. Herb. Boiss. ser. 2, 3: 
134. 1903. Colombia, Ecuador. 

59. P. (subgenus ?) ORANENSIS L. B. Smith, Phytologia 8:228, 
pi. 2, figs. 15-17. 1962. Argentina. 

60. P. SUBPETIOLATA Baker, Joum. Bot. 19:267. 1881. P. lati - 
folia senau Baker, Handb. Bromel. 92. 1889, in part, as to Bur- 
chell 8116 . P. caldasiana Baker, op. c. 100. P. bur c belli i Mez 
Mart. Fl. Bras. 3, pt. 3:436. 1894. P. sessiliflora Rusby, Bull. 
New York Bot. Gard. 4:457. 1907. Peru, Bolivia, Brazil. 

61. P. TRUNCATA L. B. Smith, Contr. Gray Herb. 98:10, pi. 3, 
figs. 8, 9. 1932. Peru. 

62. P. CHIRIGUANA Castellanos, An. Mus. Nac. Hist. Nat. Buenos 
Aires 36:52, pi. 4. 1929. Argentina. 

63. P. TARAPOTENSIS Baker, Handb. Bromel. 102. 1889. Peru. 

64. P. (subgenus ?) CHOCOENSIS L. B. Smith, Contr. U. S. Nat. 
Herb. 29:302, fig. 24. 1949. Colombia. 

65. P. (subgenus ?) HAUGHTII L. B. Smith, Contr. U. S. Nat. 
Herb. 29:307, fig. 28. 1949. Colombia. 

66. P. MEGASEPALA Baker, Joum. Bot. 19:229. 1881. P. araneo- 
osa Baker, op. c. 231. P. goudotiana Andre, finum. Bromel. 4. 
Dec. 13, 1888; Rev. Hortic. 60:564. Dec. 16, 1888. P. caapto - 
calyx Andre, I.e. P. caaptocalyx var. lutea, var. robusta . 

1. c. Colombia. 

67. P. TRIMORPHA L. B. Smith, Phytologia 5:179, pi. 2, figs. 
1-3. 1955. Colombia. 

68. P. POORTMANII Andre, finua. Branel. 4. Dec. 13, 1888; Rev. 
Hortic. 60:564. Dec. 16, 1888. Colombia, Ecuador. 

69. P. KNIPHOFIOIDES L. B. Smith, Proc. Am. Acad. Sci. 70:153, 



32 PHYTOLOGIA Vol. 10, no, 1 

pi. 1, fig. 18, 1935. Colombia. 

70. P. TOLIMENSIS L. B. Smith, Caldasia tlj , no. 4:17, fig. 4 
1942. Colombia. 

71. P. INTEGRIFOLIA Ker-Gairl. Bot. Mag. 36: pi. 1462. 1812. P. 
grami nif olia hort. eoc Schrad. Coram. Blumenb. 46. 1827. P. decora 
A. Dietr. Allg. Gartenzeit. 15:353. 1847. P. alta HasskT Verb. 
Naturk. Ver. Ned. Ind. 1, pt. 7:5. 1856. pT graminea Beer, Bro- 
mel, 198. 1857, nomen. 2 £♦ integrifolia var. ma.lor Regel, Ind, 
Sem. Hort. Petrop. for 1869:24. 1869. P. tenuis Uez in DC. Mon. 
Phan. 9:421. 1896. P. hartmannii Mez, Fedde Rep. Spec. Kov. 16: 
8. 1919. Venezuela, "Trinidad. 

72. P. (subgenus ?) CASSAPENSIS Mez, Fedde Rep. Spec. Nov. 16: 
8, 1919. Peru. 

73. P. MQRITZIANA K. Koch & Bouche, Ind. Sem. Hort. Berol. for 
I856, App.:4. 1857. P. klotzschiana Baker, Handb, Bromel. IO6. 
1889. P. moritziana Kl. ex Baker, 1. c, nomen. Venezuela, 

74. P. (subgenus ?) CUZCOENSIS L, B, Smith, Contr. Gray Herb. 
98:9, pi. 3, figs. 3-5. 1932. Peru. 

75. P. SCHDLTZKI Harms, Notizblatt 10:212. 1928. Colombia. 

76. P, VARGASIANA L. B. Smith, sp. nov. P, cuzcoensjs L. B. 
Smith in syStema mea proxima sed foliomm laminis majorlbus deci- 
duis, bracteis primariis quam ramis multo brevioribus differt. 

Caulescent, incomplete and immature but undoubtedly flowering 
1 m or higher; stem erect, 15 cm high; leaf-sheaths densely im- 
bricate, broadly ovate, ca. 3 cm long, entire or pectinate- 
serrate only near apex, dark castaneous, lustrous, covered with 
appressed whitish scales toward base; blades polymorphic, some 
greatly reduced and spiniform, castaneous, either entire or pec- 
tinate-serrate, others foliaceous and deciduous, either wholly 
entire or serrate below the line of abscission, linear-lanceo- 
late, acuminate, slightly narrowed toward base, 25 cm long, 15 mm 
wide, glabrous except for marginal s cales near base; scape erect, 
ca. 8 mm in diameter; lower scape-bracts densely imbricate, sub- 
follaceous with deciduous blades; inflorescence laxly confound, 
sparsely white-flocculose; primary bracts nairowly triangular, 3 
cm long; lateral branches to 18 cm long including the naked 7 cm 
sterile base, laxly flowered, terminal branch 27 cm long; floral 
bracts to 25 mm long, much exceeding the pedicels; flcwers se- 
cund, spreading to recurved at anthesis, red; pedicels 8 mm long; 
sepals lance-oblong, subacute, 2C mm long, ecarinate or nearly 
so, soon glabrous; petals naked; ovary over 1/2 superior; ovules 
caudate. PI, I, fig. 4: Flower x 1/2; fig. 5: Sepal x 1. 

PERU: Cuzco: Prov. Paucartambo: Rocky open places, K. 134. 
alt. 2000 m, July 20, 1963, C, Vargas C. no. 14716 (US, type). 

77. P. HALOPHILA L. B. Smith, sp. nov. P. platystemon Mez 
in systema mea proxlma sed bracteis florigeris an gust e triangu- 
laribus, petalis lacteis differt. 

Stemless (?), flowering 75 cm high; leaves apparently all per- 
sistent; sheaths broadly ovate, the outer 2 cm long, entire ex- 
cept near apex, dark castaneous, at first covered toward apex 
with appressed cinereous scales; blades polymorphic, some reduced 
to splnose-serrate spines, some much reduced but foliaceous and 



196U Smith, Notes on Brcmellaceae 33 

entire, the central ones foliaceous, linear, acuminate, scarcely 
narrowed at base, about equaling the inflorescence, 12 mm wide, 
entire, covered beneath with pale subappressed scales, glabrous 
above; scape slender, erect, pale-lepidote ; scape-bracts appar- 
ently exceeding the intemodes (the highest damaged and uncer- 
tain), linear-triangular J inflorescence 3U cm long with a single 
short lateral branch, lax, pale-flocculosej primary bract linear, 
attenuate, about equaling the naked sterile base of the branch; 
floral bracts linear-triangular, to 15 mm long; flowers divergent 
not secund; pedicels straight, slender, to 25 mm long; sepsds 
linear, acute, 23 mm long, scarcely carinate, soon glabrous; pet- 
als naked, over 5 cm long, cream (J Foster); ovary over 2/3 supe- 
rior; ovules short-caudate. PI. I, fig. 6: Flower x 1/2; fig. 7: 
Sepal x 1. 

COSTA RICA: Puntarenas; Base of cliffs at high-tide level, 
Quepos, December 12, 19AB, U. B. Foster no. 2669 (US, type), 

78. P. EXSERTA L. B. Smith ,"Caldasia 5:10, fig. 19i^. 
Colombia. 

79. P. DEVANSAIANA Andre ex Baker, Handb. Bromel. 93. 1889. 
P. roezlii Andre, ^um. Bromel. 4. Dec. 13, 1888; Rev. Hortic. 
^:564. Dec. 16, 1888, non E. Morr. 1885. Ecuador. 

80. P. DIFFUSA L. B. Smith, Caldasia 5:8, fig. 1948. Colombia. 

81. P. (subgenus ?) MULTIRAMOSA Mea in DC. Mon. Phan. 9:419. 
1896. Bolivia. 

82. P. ODONTOPODA Baker, Handb. Bromel. 93. 1889. Bolivia. 

83. P. DIVARICATA Wittm. Uededeel. Rijks Herb. 29:81. 1916. 
Bolivia. 

84. P. (subgenus ?) APHELANDRIFLORA Lem. 111. Hortic. 16: 
Misc. 90. 1869. Pepinia aphelandriflora Andre, m. Hortic. 17: 
32, pi. 5. 1870. Peru, Panama. 

85. P. CAP rr ATA L. B. Smith, Contr. U. S. Nat. Herb. 29:301, 
fig. 23. 1949. Colombia. 

86. P. (subgenus ?) GUZMANIOIDES L. B. Smith, Contr. U. S. 
Nat. Herb. 29:306, fig. 27. 1949. Colombia. 

87. P. ARCHEHI L. B. Smith, Contr. Gray Herb. 104:79, pi. 3, 
figs. 20, 21. 1934. Colombia. 

88. P. (subgenus ?) POEPPIGIANA Mez in Mart. Fl. Bras. 3, pt. 
3:461. 1894. Peru. 

89. P. MACRANTHERA Andre, 6num. Bromel. 5. Dec. 13, 1888; Rev. 
Hortic. 60:565. Dec. 16, 1888; emend. L. B. Smith, Contr. U. S, 
Nat. Herb. 29:310. 1949. Colombia. 

90. P. MAIDIFOLIA (C. Morr.) Dene, ex Planch. Fl. Serres 9: 
151, pi. 915. 1854. Puya maidifolia C. Morr. Ann. Soc. Agr. Bot. 
Gand 5:453, pl. 1849. P. funkiana Linden, Cat. 5:2. 1850. Pit- 
caimia funkiana A. Dietr. Allgem. Gartenzeit. 19:337. 1851. P. 
maydifolla Dene, ex Naudin, Rev. Hortic. ser. 3, 5:347. 1851. ~P. 
macro calyx Hook. Bot. Mag. 79:pl. 4705. 1853. ? P. polyanthoides 
Brongn. ex Dene. Rev. Hortic. 27:244. 1855 (! MezT. P. zeifolia 
Koch & Sello ex K. Koch, Ind. Sem. Hort. Berol. for 1854: App. 11 
1855. P. malzaifolia hort. ex Beer, Bromel. 46. 1857, nomen. 
Phlomostachys funkiana Beer, Bromel. 47. 1857. Neumannia maidi - 
folia K. Koch, Ind. Sam. Hort. Berol. for I856: App. 2. 1857. 



3U PHYTOLOGIA Vol. 10, no. 1 

Pitcairnia funkil Viaiira, It. Princ. S. Coburgi 1:174. 1883. Lam- 
proconus maldifollus Lem. ex Baker, Handb. Bromel. 109. 1889, 
nomen. Vrlesia tricolor hort. ex Baker, 1, c, nomen. Pitcair- 
nia oerstediana Mez in DC. Uon. Phan. 9:kUB. 1896. Guiana, 
Venezuela, Colombia, Central America. 

91. P. SANDMANII L. B. Smith, Contr. U. S. Nat. Herb. 29:531, 
fig. 87. 1954. Peru. 

92. P. (subgenus ?) MARITIMA L. B. Smith, Contr. U. S. Nat. 
Herb. 29:311, fig. 31. 1949. Colombia. 

93. P. (subgenus ?) FERREYRAE L. B. Smith, Phytologia 4:215, 
pi. 1, figs. 10-12. 1953. Peru. 

94. P. SCEPTRIFORMIS Mez, Bull. Herb. Boisa. ser. 2, 4:628. 
1904; Ule, Verhandl. Bot. Ver. Brandenb. 48:139. 1907. Peru. 

95. P. BRITTONIANA Mez in DC. Mon. Phan. 9:451. 1896. P. wer - 
cleana Mez, Bull. Herb. Boiss. ser. 2, 4:622. 1904. P. flaviflo- 
ra Standley, Joum. Wash. Acad. Sci. 17:247. 1927. Costa Rica to 
British Guiana and Bolivia. 

96. P. NIGRA (Carr.) Andre, finum. Bromel. 5. Dec. 13, 1888, 
incorrectly attributed to Carr.; Rev. Hortic. 60:565, Dec. 16, 
1888. Neumannia nigra Carr. Rev. Hortic. 53:390, pi. 1881, Pit- 
cairnia gravisiana Wittm. Bot, Jahrb, 11:56, 1889. Colcsnbia, 
Ecuador , 

97. P. PULCHELLA Mez in DC. Mon. Phan. 9:459. 1896. Ecuador. 

98. P. BRONCailARTIANA Andre, finum. Brcaiel, 5. Dec. 13, 1888; 
Rev. Hortic. 60:565. Dec. 16, 1888. 

Var, a, BRONGNIARTIANA, P, klabochorum hort. ex Baker, Handb. 
Bromel. 107. 1889, nomen. P, klabochiana C. Morr. ex Mez In DC, 
Mon. Phan. 9:414. 1896. Leaf-blades naiTOwly lanceolate, acumi- 
nate, 80 mm wide. Colombia. 

Var. b. LATITOLIA L, B. Smith, Phytologia 6:438, 1959. Leaf- 
blades oblanceolate, broadly acute and aplculate, 135-145 noi 
wide. Ecuador, 

99. P. BRUNNESCENS L, B. Smith, Contr, U, S. Nat, Herb, 29: 
300, fig, 22, 1949. Colombia, Ecuador, 

100. P, BARRIGAE L, B, Smith, Caldasia 5:5, fig. 1948. 
Colombia. 

101. P. (subgenus ?) CUVATA L, B, Smith, Phytologia 9:248, 
pi, 2, figs, 9, 10, 1963, Peru, 

102. P, SQUARROSA L, B, Smith, Caldasia 5:11, fig. 1948. 
Colombia. 

Var, a. SQUARROSA. Floral bracts castaneous with acuminate 
green apices; petals white, 

Var. b. COLORATA L. B, Smith, Caldasia 5:12, 1948, Floral 
bracts bright red; petals dark purple, 

Var, c, AURANTIACA L, B, Smith, Phytologia 5:396. 1956. In- 
florescence 10 cm long; florad bracts with short broadly acute 
apices; sepals 25 mm long; petals orange. 

103. P. ATRORUBENS (Beer) Baker, Joum. Bot, 19:307. 1881. 
Phlomostachys atrorubens Beer, Bromel, 48. 1857. Puya warsze - 
wiczii H. Wendl. ex Hook, Bot, Mag. 87:pl. 5225. 1861. Pitcair- 
nia lamarcheana E, Morr, ex Baker, Handb, Bromel, 111, 1889. P, 
lindeni Baker, op, c, 112, Neumannia linden! E. Morr. ex Baker, 



196U Smith, Notes on Bromellaceae 35 

1. c, nomen. N. atrorubens K. Koch ex Baker, 1. c, nomen. 
Lamproconua warszcewlczil Lem. ex Baker, 1. c, nomen. P. atro- 
rubens /3. lamarcheana Mez in DC. Won. Phan. 9:457. 1896. 
Colombia, Panama, Costa Rica, kexico (?). 

104. P. CAMPII L. B. Smith, Mem. New York Bot. Gard. 8:28, 
fig. 1, j-k. 1952. Ecuador. 

105. P. ELONGATA L. B. Smith, Contr. U. S. Nat. Herb. 29:304, 
fig. 25. 1949. Colombia, Ecuador. 

106. P. (subgenus ?) BAKERI (Andre) Andre ex iiez in DC. Mon. 
Phan. 9:460. 1896; emend. L. B. Smith, Caldasia 5:4, fig. 1948. 
Quesnelia bakeri Andre, ^um. Bromel. 4. Dec. 13, 1888; Rev. 
Hortic. 60:564. Dec. 16, 1888. Colombia, Ecuador. 

107. P. (subgenus ?) OBLANCEOLATA L. B. Smith, Contr. Gray 
Herb. 117:26, pi. 2, fig. 18. 1937. Colombia, Panama, Costa Rica 

108. P. ARCUATA (Andre) Andre, tnxm. Bromel. 5. Dec. 13, 1888; 
Rev. Hortic. 60:565. Dec. 16, 1888. Neumannia arcuata Andre, 
Rev. Hortic. 58:108, pi. 1886. Colombia. 

109. P. FOSTERIANA L. B. Smith, Contr. U. S. Nat. Herb. 29: 
305, fig. 26. 1949. Colombia. 

lie. P. UMBRATILIS L. B. Smith, Phytologia 4:216, pi. 1, figs. 
15-17. 1953. Peru. 

111. P. HITCHCOCKIANA L. B. Smith, Phytologia 5:44, pl. 6, 
figs. 1-3. 1954. Ecuador. 

112. P. (subgenus ?) SCEPTRIGERA Mez, Fedde Rep. Nov, Spec. 3: 
7. 1906. Ecuador. 

113. P. ASPLUNDII L. B. Smith, Phytologia 4:214, pl. 1, figs. 
6-9. 1953. Peru. 

U4. P. SUBULIFERA L. B. Smith, Phytologia 4:215, pl. 1, figs. 
13, 14. 1953. Peru. 

115. P. ALTENSTEINII (Lk., Kl. & Otto) Lem. Fl. des Serres ser 
1, 2:pl. 162. 1846. Venezuela. 

Var. a. ALTENSTEINn. Puya altensteinii Lk., Kl. & Otto, Pl. 
Rar. Hort. Berol. 1:1, pl. 1. 1840. Pitcairnia undulatifolia 
Hook. Bot .Mag. 72:pl. 4241. 1846, non hort. 1846. Lamproconus 
altensteinii Lem. Jard. Fleur. 2: sub pl. 127. 1852. ? Pitcairnia 
rhodostachys Hassk. Verh. Nat. Ver. Ned. I»d. 1, pt. 7:8. I856; 
cf. Uez in Pflanzenreich 4, Fara. 32:275. 1935. Phlomostachys al- 
tensteinii Beer, Bromel. 45, 1857. Neumannia siltensteinii Griseb 
Goett. Nachr. for 1864:14. I865. Flooring about 1 m high or 
less; leaves less than 12; inflorescence ca. 12 cm long. 

Var. b. GIGANTEA (Hook.) Baker, Handb. Bromel. 111. 1889. 
Puya altensteinii var. pigantea Hook. Bot. Mag. 73:pl. 4309. 
1847. P. macrostachys A. Dietr. Allg. Gartenzeit. 16:145. 1848. 
P. macrostachya Schomb. Fl. & Fauna Guyana 1068. 1848, nomen. 
Lamproconua gigaaateus Lem. Jard. Fleur. 2: sub pl. 127. 1852. 
Phlomostachys gigantea Beer, Bromel. 47. 1857. Neumannia gigan - 
tea Brongn. ex Beer, Bromel. 47. 1857, nomen. Flowering over 2 m 
high; leaves many; inflorescence ca. 30 cm long. 

116. P. HETEROPHYLLA (Lindl.) Beer, Bromel. 68. 1857. 

Var. a. HETEROPHYLLA. Leaves dimorphic. Peru and Venezuela 
to Panama and Uexico. 

Forma a. HETEROPHYLLA. Puya heterophylla Lindl. Bot. Reg. 26: 



36 PHITOLOQIA Vol. 10, no. 1 

pi. 71. 18A0. P. longlfolla C. Morr. Ann. Soc. Agr. Bot. Gand 2: 
A83, pi. 101. 1846. Pltcalmla exscapa Liebm. Ind. Sem. Hort. 
Haun. for 1848:12. 1848. P. llndleyana Lem. Jard. Fleiir. 2: sub 
pi. 151. 1852. P. morrenll Lem. op. c. 3:pl. 291. 1852. P. lon- 
^Ifolla Beer, Br^el. 6?. 1857. P. llebmannil K. Koch, Ind. Sen. 
Hort. Berol. for 1857:App. 7. 1858. Petals some shade of red. 

Foma b. ALBIFLORA Standley & L. B. Staith, Lilloa 6:383. 1941. 
Petals white. 

Var. b. EXSCAPA Mez in DC. Mon. Phan. 9:375. 1896. Pitcairnia 
cernua Kunth & Bouche, Ind. Sem. Hort. Berol. for 1848:12. 1848. 
P. exscapa Hook. Bot, Mag. 77:pl. 4591. 1851, non Liebm. 184S. 
Colombia, Ecuador. 

117. P. (subgenus ?) AUGUSTII Harms, Notizblatt 10:211. 1928. 
Peru. 

118. P. AEQUATORIALIS L. B. Smith, Contr. Gray Herb. 114:6. 
pi. 1, fig. 3. 1936. Ecuador. 

119. P. CRASSA L. B. Smith, Lilloa 14:94, fig. 5. 1948. 
Bolivia. 

120. P. LICaiOSA L. B. Smith, Contr. U. S. Nat. Herb. 29:309, 
fig. 30. 1949. Colombia. 

121. P. STENOPHYLLA Andre, Enum. Bromel. 4. Dec, 13, 1888; 
Rev. Hortic. 60:564. Dec. 16, 1888. Colombia. 

122. P. PAVONII Mez in DC. Mon. Phan. 9:386. 1896. Ecuador, 

123. P. PUNGENS H.B.K. Nov. Gen. & Sp. 1:294. 1816. P. laevis 
Willd. ex Schult. in R. & S. Syst. 7, pt. 2:1249. 1830, nomen. 

P. eoneolor Baker, Joum. Bot. 19:269. 1881. Colombia to Peru. 

124. P. BTT.T.RKRGIOIDE's L. B. Smith, Publ. Mus. Hist. Nat. 
"Javier Prado" Univ. Nac. Mayor San Marcos ser. B, no. 13:4, fig. 
6-8. 1963. Peru. 

125. P. (subgenus ?) MELANOPODA L. B. Smith, Phytologia 9:249, 
pi. 2, figs. 11-13. 1963. Peru. 

126. P. LOPEZII L. B. Smith, Phytologia 5:46, pi. 6, figs. 
4-6. 1954. Peru. 

127. P. DECURVATA L. B. Smith, sp. nov. P. lopezii L. B. 
Smith in systema mea proxima sed scapo graeili, scapi bracteis 
quam intemodiis brevioribus, floribus patentibus vel decuirvatis 
secundis differt. 

Stenless, flowering over 6 dm high; leaves subbulbous-rosu- 
late; blades dimorphic, some reduced to slender pectinate-serrate 
dark spines, others foliaceous, deciduous, linear, acuminate, 32 
cm long, 18 mm wide, entire above the line of abscission, pale- 
lepidote beneath, soon glabrous; scape erect, 5 mm in diameter, 
soon glabrous; scape-bracts linear-lanceolate, acuminate, much 
shorter than the upper intemodes; inflorescence simple, lax, 
white-flocculose, becoming glabrous; floral bracts ovate, acumi- 
nate, to 23 mm long, exceeding all the pedicels; flowers secund, 
spreading to decuirved at anthesis; pedicels slender, to 14 mm 
long, scanewhat biangulate; sepals narrowly triangular, acviminate, 
30 mm long, glabrous at anthesis, the posterior ones subalate- 
carinate toward base; petals over 8 cm long, naked, yellow; ovary 
7/8 superior; ovules caudate. PI. I, fig. 8: Flower x 1/2; fig. 
9: Sepal x 1. 



196U Smith, Notes on Br<aaellaceae 37 

PERU: Lambejeque: Prov. Lambeyeque: Rocky slope, between Bea- 
tlta de Humay and Km. 38 on the Olmos-Uaranon Hlghiray, alt. 150 
m. May 21, 1963, A. Lopez . A. Sagastegul & V. Collantea . no. 4047 
(TRP, type). 

128. P. VIOLASCENS L. B. aaith, Phytologia 6:439, pl. 2, figs. 
12, 13. 1959. Ecuador. 

129. P. FRACTIFOLIA L. B, Smith, sp, nov. P. palmeri S. Wat- 
son in systena mea proxina sed pedlcellis brevioribus, floribus 
haud secundis differt. 

Stemless (?), flowering 75 cm high; leaves bulbous-rosiilate, 
entire; sheaths very broadly ovate, centrally castaneous, densely 
pale-lepidote, becoming glabrous; blades dlmoirphic, some much re- 
duced, subulate, persistent, green, others foliaceoas, linear, 
acuminate, over 55 cm long (incoi^lete), 12 vam. wide, covered with 
appressed cinereous scales beneath, glabrous above, channeled; 
scape erect, slender, sparsely pale-lepidote; scape-bracts nar- 
rowly triangular, acuminate, much shorter than the upjier inter- 
nodes; inflorescence simple, lax, 19 cm long, subglabrous; floral 
bracts narrowly triangular, acuminate, to 10 mm long; flowers 
polystichous , not secund; pedicels spreading, slender, to 20 mm 
long, much exceeding all the floral bracts; sepals oblong, acute, 
16 mm long, short-carinate above the tumid base; petals (iiaaa- 
ture) barely exceeding the sepals, red, nsiked; ovary 4/5 superi- 
or; ovules caudate. PI. I, fig. 10: Flower x 1/2; fig. 11: 
Sepal X 1. 

PERU: Amazcmas: Prov. Chachapoyas: On rocks, Tlalango, between 
Bagua and Ingenio, alt. 800 m. May 27, 1963, A. Lopez , A. Sagas - 
tegui & V. Collantes . no. 4242 (TRP, type). 

I30T P. CARDQIASII L. B. Smith, Phytologia 8:505, pl. 3, figs. 
5-7. 1963. Bolivia. 

131. P. STRAMINEA Poeppig ex Mez in DC. Mon. Phan. 9:427. 
1896. Peru. 

132. P. ALATA L. B. Smith, Lloydia 11:304, fig. 2. 1948. 
Ecuador . 

133. P. (subgenus ?) SCANDENS Ule, Verhandl. Bot. Ver. Bran- 
denb. 48:140. 1907. Peru. 

134. P. LUTESCENS Mez & Sodiro, Bull. Herb. Boiss. ser. 2, 4: 
626. 1904. Ecuador. 

135. P. ERRATICA L. B. Smith, Lloydia 11:305, fig. 3. 1948. 
Ecuador. 

136. P. ACICULARIS L. B. Smith, Publ. Mus. Hist. Nat. "Javier 
Prado" Univ. Mayor San Marcos ser. B, no. 13:3, figs. 4, 5. 
1963. Peru. 

137. P. RIP ARIA Mez, Fedde Rep. Spec. Nov. 12:416. 1913. Peru. 

138. P. SPECTABILIS Mez in DC. Mon. Phan. 9:421. 1896. 
Colombia . 

139. P. LAXISSIMA Baker, Handb. Bromel. 101. 1889. Colombia. 

140. P. LONGIPES Mez, Fedde Rep. Spec. Nov. 12:416. 1913. 
Colombia. 

141. P. ELLIPTICA Mez & Sodiro, Bull. Herb. Boiss. ser. 2, 4: 
624. 1904. Ecuador. 

142. P. TUMULICOU L. B. Smith, Contr. U. S. Nat. Herb. 29: 



38 PHTTOLOGIA Vol. 10, no. 1 

314, fig. 3h. 1949. Colombia. 

143. P. BIFLORA L. B. Smith, Contr. Gray Herb. 127:17, pi. 1, 
figs. 1-3. 1939. Peru. 

144. P. MULTIFLORA L. B. Smith, Conbr. U. S. Nat. Herb. 29: 
312, fig. 32. 1949. Colombia. 

145. P. KALBREYERI Baker, Journ. Eot. 19:273. 1881. Colombia, 
Painama. 

146. P. VOLUBILIS L. B. Smith, Phytologia 5:33, pl. 1, figs. 
1-4. 1954. Colombia. 

147. P. NUBIGENA Planch. Fl. des Serres ser. 1, 8:265, pl. 
847. 1852-53. Venezuela. 

148. P. MERIDENSIS Kl. ex Mez in DC. Mon. Phan. 9:385. 1896. 
P. integrifolia var. "P." meridensis Kl. ex Baker, Handb. Bromel. 
97. 1889. Venezuela. 

149. P. BELLA L. B. Smith, Caldasia 3:240, fig. 1945. Colombia 
Var. a. BELLA. Upper scape-bracts shorter than the inter- 
nodes; sepals 24 mm long. 

Var. b. DENSIOR L. B. Smith, Contr. U. S. Nat. Herb. 29:300. 
1949. Scape-bracts exceeding the internodes; sepals not over 17 
mm long. 

15C. P. (subgenus ?) CALOPHYLLA L. B. Smith, Contr. U. S. Nat. 
Herb. 29:432, fig. 38. 1951. Colombia, 

151. P. SYLVESTRIS L. B. Smith, Contr. Gray Herb. 117:29, pl. 
2, fig. 27. 1937. Colombia. 

152. P. DOLICHOPETALA Harms, Notizblatt 12:530. 1935. 
Colombia. 

153. P. REFLEXIFLORA Andre, finum. Bromel. 4. Dec. 13, 1888; 
Rev. Hortic. 60:564. Dec. 16, 1888. Ecuador. 

154. P. SEMAPHORA L. B. Smith, Caldasia 3:242. 1945. Colombia 

155. P. MACARENENSIS L. B. Smith, Bot. Mus. Leafl. Harvard 16: 
192, pl. 25, figs. 1-4. 1954. Colombia. 

156. P. GUARITERMAE Andre, £num. Bromel. 4. Dec. 13, 1888; 
Rev. Hortic. 60:564. Dec. 16, 1888. Colombia. 

157. P. (subgenus ?) CALATHEOIDES L. B. Smith, Contr. Gray 
Herb. 117:24, pl. 2, figs. 8-11. 1937. Peru. 

158. P. SODIROI Mez, Bull. Herb. Boiss. ser. 2, 4:622. 1904. 
Ecuador. 

159. P. NOBILIS Mez & Sodiro, Bull. Herb. Boiss. ser. 2, 4: 
623. 1904. Ecuador. 

160. P. (subgenus ?) ADSCENDENS L. B. Smith, Contr. U. S. Nat. 
Herb. 29:298, fig. 20. 1949. Colombia. 

161. P. SNEIDERNII L. B. Smith, Phytologia 6:434, pl. 1, figs. 
5-7. 1959. Colombia. 

162. P. (subgenus ?) CYANOPETALA Ule, Verhandl. Bot. Ver. 
Brandenb. 48:139. 1907. Peru. 

163. P. BRACHYSPERMA Andre, ^um. Bromel. 4. Dec. 13, 1888; 
Rev. Hortic. 60:564. Dec. 16, 1888. Colombia. 

164. P. SIMILIS L. B. Smith, Contr. U. S. Nat. Herb. 29:434, 
fig. 40. 1951. Colombia. 

165. P. PAIAfOIDES Mez & Sodiro, Bull. Herb. Boiss. ser. 2, 4: 
626. 1904. Ecuador. 

166. P. (subgenus ?) EXIMIA Mez, Fedde Rep. Nov. Spec. 3:5. 



196U Smith, Notes on Bromeliaceae 39 

1906. Peru. 

167. P. TRIANAE Andre, £nura. Bromel. 4. Dec. 13, 1888; Rev. 
Hortic. 60:564. Dec. 16, 1888. 

Var. a. TRIANAE. Sepals obtuse; petals 35-45 mm long. 
Colombia, Bolivia. 

Var. b. RETUSA L. B. Sniith, Lloydia 11:305. 1948. Sepals re- 
tuse; petals not over 25 mm long. Ecuador. 

168. P. OBLONGIFOLIA L. B. Smith, Phytologia 6:438, pi. 2, 
figs. 9-11. 1959. Ecuador. 

169. P. LANUGINOSA R. & P. Fl. Peruv. 3:35, pi. 258. 1802. 
Puya ruiziana Uez in DC. Mon. Phan. 9:491. 1896, as to synonymy, 
not as to specimen. Peru. 

170. P. PARAGUAYENSIS L. B. Smith, Rev. Argentina Agron. ?: 
163, figs. 4-6. 1940. Paraguay. 

171. P. MACROBOTRYS Andre, Enum. Bromel. 4. Dec. 13, 1888; 
Rev. Hortic. 60:564. Dec. 16, 1888. Colombia. 

172. P. ORCHIDIFOLIA Mez, Fedde Hep. Spec. Nov. 17:114. 1921. 
Venezuela. 

173. P. FENDLERI Mez in DC. Mon. Phan. 9:387. 1896. Venezuela 

174. P. UNILATERALIS L. B. Smith, Phytologia 8:11, pi. 1, 
figs. 21-23. 1961. Ecuador. 

175. P. LEPIDOPETALON L. B. Smith, Caldasia 5, no. 21:10, fig. 
1948. Colombia. 

176. P. PUSILLA Mez in DC. Mon. Phan. 9:429. 1896. French 
Guiana. 

177. P. TYMPANI L. B. Smith, Phytologia 5:177, pl. 1, figs. 
4-6. 1955. Venezuela. 

178. P. (subgenus ?) OCCIDENTALIS L. B. Smith, Phytologia 4: 
380, pl. 1, figs. 4-6. 1953. Colombia. 

179. P. (subgenus ?) BREVICALYCINA Mez, Fedde Rep. Spec. Nov. 
16:9. 1919. Peru, Venezuela. 

180. P. PETRAEA L. B. Smith, Contr. U. S. Nat. Herb. 29:433, 
fig. 39. 1951. Colombia. 

181. P. (subgenus ?) ANDREANA Linden Cat. 1873; HI. Hortic. 
20:146, pl. 139. 1873. P. lepidota Regel, Act. Hort. Petrop. 2: 
435. 1873; Gartenfl. 22:389, pl. 772. 1873. Colombia. 

182. P. GRUBBIANA L. B. Smith, sp. nov. P. .jimenezii in sys-^ 
tema mea proocima sed pedicellis majoribus bract eas f lorigeras 
omnias superantibus differt. 

Stemless (?), flowering 45 cm high; leaves fasciculate, per- 
sistent, nearly equaling the inflorescence, entire; sheaths 
ovate, ca. 2 cm long, dark castaneous, at first covered with a 
membrane of pale coalesced scales; blades dimorphic, some much 
reduced, subulate, green, others foliaceous, linear, caudate- 
acuminate, slightly narrowed toward base but not petiolate, 14 ram 
wide, glabrous; scape er«ct, slender, sparsely pale-lepidote ; 
lower scape-bracts foliaceous, large, the highest ovate, caudate, 
shorter than the intemode; inflorescence simple, lax, the axis 9 
cm long, sparsely pale-lepidote; floral bracts ovate, aciiminate, 
to 19 mm long, thin; flowers suberect, not secund; pedicels slen- 
der, to 22 mm long, all exceeding the floral bracts; sepals ob- 
long, obtuse, 16 mm long, obscurely carinate; petals 4 cm long. 



UO PHTTOLOQIA Vol. 10, no. 1 

bearing a crenate scale at base, red; stamens included; ovary 3/4 
superior; ovules caudate, PI. I, fig. 12: Flower x 1/2; fig. 13: 
Sepal X 1. 

COLOMBIA: Boyaca: Hill sabana on path troai Bachira to Bocota 
soon -after it climbs up out of Valley bottom, Sierra Nevada de 
Cocuy, alt. ca. 2150 m, August 21, 1957, P. J. Grubb . B. A. B. 
Curry & A. Fernandez-Peres, no. blf) (US, type). 

I83T P. (subgenus ?) CAULESCEKS K. Koch ex Uez in DC. Uon. 
Phan. 9:A25. 1896. Venezuela (?). 

184. P. UICROCALU Baker, Joum. Bot. 19:228. 1881. Venezuela 
Var, a. MICROCALYX. P. lutea hort. Linden ex Baker, Handb. 

Bromel. 104. 1889. Floral bracts lanceolate, acuminate; sepals 
triangular, acuminate; petals yellow. 

Var. b. SCHLBfll (Baker) L. B. Smith, Phytologia 7:106. I960. 
P. schlimii Baker, Handb. Bromel. 100. 1889. P. caracasana 
Baker, op. c. 103. Floral bracts lanceolate, acuminate; sepals 
triangular, acuminate; petals red. 

Var. c. ELLIPTICA L. B. Smith, Phytologia 7:107. I960. Floral 
bracts elliptic, apiculate; sepals oblong, subacute; petals red. 

185. P. JOHANNIS L. B. Smith, Phytologia 5:179, pl. 1, figs. 
11-13. 1955. Colombia. 

Uexico, Central America 
(cf. nos. 77 (out of order), 84, 90, 95, 103, 107, 116) 

186. P. CHIRIQUENSIS L. B. Smith, Contr. U. S. Nat. Herb. 29: 
281, fig. 6, 1949. Panama. 

187. P. FLAGELLARIS L. B. Smith, Contr. U. S. Nat. Herb. 29: 
280, fig. 5. 1949. Guatemala. 

188. P. VALERII Standley, Joum. Wash. Acad. Sci. 17:246. 
1927. Costa Rica. 

189. P. TABULIFORMIS Linden Cat. 17:5. 1862. Mexico. 

190. P. RECUEVATA (Scheidw.) K. Koch, Ind. Sem. Hort. Berol. 
for 1857 :App. 4. 1858. Puya recurvata Scheidw. Allgem. Garten- 
zeit. 10:275. 1842. Pepinia i^curvata E. Morr. ex Baker, Handb. 
Bromel. 110. 1889, nomen. Pitcaimia taenipetala Mez in DC. Mon. 
Phan. 9:382. I896. Guatemala, British Honduras, Mexico. 

191. P. (subgenus ?) LONCEBRACTEATA Bouche ex Mez in DC. Mon. 
Phan. 9:428. 1896. Guatemala (?). 

192. P. MATUDAE L. B. Smith, Phytologia 8:220, pl. 1, figs. 5, 
6, 1962. Mexico. 

193. P. WENDLANDII Baker, Joum. Bot. 19:306. 1881. Puya sul- 
phurea Hook. Bot, Mag. 79:pl. 4696. 1853. Neumannia sulphurea K. 
Koch, Ind. Sem. Hort. Berol. for 1856:App. 2. 1857. Phlomosta - 
chys sulphurea Beer, BroD«l. 46. 1857. Pitcaimia altensteinii 
sensu J. Donnell Smith, Enum. Pl. Guatemal. 3:79. 1889-1907. P. 
sulfurea Mez in DC. Mon. Phan. 9:456. 1896, non Andr, 1802. 
Costa Rica, Guatemala, Mexico. 

194. P. MACROCHLAMYS Mez, Fedde Rep. Nov. Spec. 3:6. 1906. 
Guateaiala. 

195. P. HatSLETANA Mez in DC. Mon. Phan. 9:455. 1896, as to 
type only. P. ochroleuca Baker, Joum. Bot. 19:306. 1881; Baker 



196U Smith, Notes on Bromellaceae 111 

in Hemsl. Biol. Centr.-Am. Hot. 3:317, pl. 85. 1884, as to mate- 
rial cited, not as to basionya. P. petiolata Baker, Handb. Bro- 
mel. 112. 1889, in part, as to Salvin & Godman . Guatemala. 

196. P. IMBRICATA (Brongn.) Regel, Gartenfl. 17:135. 1868. 
Neumannla imbricata Brongn. Ann. Sci. Nat. ser. 2, 15:369. 1841. 
Phlomoatachys imbricata Beer, Bromel. 47. 1857. Pitcaimia 
imbricata Brongn. ex K. Koch, Ind. Sem. Hort. Berol. for 185 6: 
App. 2. I857, nomen. Mexico. 

197. P. PETIOLATA (Koch & Bouche) Baker, Joum. Bot. 19:307. 
1881. Netamannla petiolata Koch & Bouche ex K. Koch, Ind. Sem. 
Hort. Berol. for 1856:App. 2. 1857. Guatemala, British Honduras. 

198. P. CARIOANA Wittm. Bot. Jahrb. 14, Beibl. 32:4. 1891. P. 
cariovana kez in DC. Mon. Phan. 9:461. 1896. Guatemala. ~ 

199. P. MEMBRANIFOLIA Baker, Handb. Bromel. 109. 1889. Costa 
Rica. 

200. P. DENSIFLORA Brongn. ex Lem. Hortic. Univ. 6:228. 1845. 
P. aurantiaca Tenore, Ann. Sci. Nat, ser. 4, 2:378. 1854. Phlo- 
mostachys densiflora Beer, Bromel. 46. I857. Lamproconus auran- 
tiacus E. Morr. ex Baker, Handb, Bromel. II4. 1889, nomen. 
Mexico. 

201. P. OCHROLEUCA (Koch k Bouche) Baker, Joum. Bot. 19:306. 
1881. Neumannia ochroleuca Koch & Bouche in K. Koch, Ind. Sem. 
Hort. Berol. for 1856:App. 2. 1857. Central America or keocico ? 

202. P. THEAE Mez in DC. Mon. Phan. 9:376. 1896. Costa Rica. 

203. P. SAXICOU L. B. Smith, Contr. Gray Herb. 117:29. 1937. 
P. fulgens A. Dietr. Allgem. Gartenzeit. 19:137. May 3, 1851, non 
Dietr. 1837, nee Dene. Jan. I85I. P, splendens Warsc. ex A. 
Dietr. Allgem. Gartenzeit. 19:176. May 31, 1851, non Poir. I836. 
P. ovandensls Matuda, An. Inst. Biol. Mexico 23, nos. 1 & 2:90. 
1953. P. lymanii Matuda, op. c. 99. Guatemala, Mexico, 

204. P. PURPUSII L. B. Smith, Contr. Gray Herb. 117:27, pl. 2, 
figs. 20, 21. 1937. Mexico. 

205. P. CALDERONII Standley & Smith in L. B. Smith, Contr. 
Gray Herb. 98:8, pl. 3, figs. 1, 2. 1932. Honduras, Salvador, 
Guatemala . 

206. P. ROSEANA L. B. Smith, Contr. Gray Herb. 117:27, pl. 2, 
figs. 22, 23. 1937. Mexico. 

207. P. MICHELIANA Andre, Rev. Hortic. 73:576, pl. 1901. 
Mexico. 

208. P. CYLINDROSTACHTA L. B, Smith, Contr, Gray Herb. 117:25, 
pl. 2, figs. 14, 15. 1937. Mexico. 

209. P. LEPROSA L. B. Smith, Contr. Gray Herb. 161:33, pl. 4, 
fig. 4. 1946. Mexico. 

210. P. MODESTA L. B. Smith, Phytologia 7:417, pl. 1, figs. 1, 
2. 1961. Mexico. 

211. P, TUERCKHEIMII Donn, Smith, Bot. Gaz. 13:190. 1888. 
Guatemala. 

Var. a. TUERCKHEIMII. Leaf -blades 10 mm wide; axis of the in- 
florescence slender, sparsely floccosej floral bracts exceeding 
the pedicels; sepals 25-30 mm long, obscurely carinate toward 
base. 

Var. b. MACROLEPIS L. B. Smith, Lilloa 6:383, pl. 1, figs. 



U2 PHTTOLOGIA Vol. 10, no. 1 

5, 6. 1941. Leaf -blades to 22 mm wide; axis of the inflorescence 
stout, densely trtiite-floccosej floral bracts much exceeding the 
I>edicels; sepals to 33 mm long, alate-carinate at base. 

212. P. HINTONIANA L. B. Smith, Contr. Gray Herb. 11/^:7, pi. 

1, figs. 8, 9. 1936. Mexico. 

213. P. MOOREANA L. B. Smith, Contr. U. S. Nat. Herb. 29:523, 
fig. 79. 1954. Mexico. 

21A. P. FLEXUOSA L. B. Smith, Contr. Gray Herb. 114:6, pi. 1, 
fig. 7. 1936. Mexico. 

215. P. PALMERI S. Watson, Proc. Am. Acad. 22:456. 1887. 
Mexico . 

Var. a. PALMERI. Tillandsia secunda Sesse & Moc. Fl. Mex. ed. 

2, 81. 1894. Floral bracts about half as long as all but the 
lowest pedicels. 

Var. b. LONGEBRACTEATA L. B. Smith, Wright ia 2:64. I960. 
Floral bracts elongate, equaling or exceeding all of the pedicels 

216. P. PUBERULA Mez & Smith ax Donn. Smith, Bot. Gaz. 19:264. 
1894. Guatemala. 

217. P. TILLANDSIOIDES L. B. Smith, Contr. Gray Herb. 161:35, 
pi. 4, fig. 7. 1946. Mexico. 

218. P. SCHIEDEANA Baker, Handb. Bromel. 95. 1889. Mexico. 

219. P. PTEROPODA L. B. Smith, Contr. Gray Herb. 117:26, pi. 
2, fig. 19. 1937. Mexico. 

220. P. MICROPODA L. B. Smith, sp. nov. Ab coinibus speciebus 
adhuc cognitis foliis majoribus declduis, scapo brevissiaio sed 
inflorescentia elongata differt. 

Stemless, flowering 16 cm high; leaves bulbous-rostilate, evi- 
dently all alike; sheaths ovate, ca. 3 cm long, dark castemeous 
at least toward apex, entire; blades deciduous, xmknown abo've the 
abscission line, the base 7 nm wide, pale-lepidote beneath, spi- 
nose-serrate; scape very short, hidden by the leaf-bases; scape- 
bracts densely imbricate, lanceolate, acuminate; inflorescence 
single, sublax, glabrous; axis slender; floral bracts ovate, acu- 
minate, to 28 mm long, much exceeding all the pedicels, entire, 
thin; flowers secund, subspreading at anthesis; pedicels slender, 
subterete, to 6 mm long; sepals linear-lanceolate, acuminate, 24 
mm long, alate-carinate; petals 35 mn long, naked, r«d; ovary 
more than 1/2 superior. PI. I, fig. 14: Flower x 1/2; fig. 15: 
Sepal X 1. 

MEXICO: Mexico: Dist. Temascaltepec: On cliffs, Puerto Sali- 
tre, alt. 1300 m, February 7, 1932, G. B. Hint on . no. 22^ (US, 
type). 

221. P. KARWINSKYANA Schult. in R. & S. Syst. 7, pt. 2:1239. 
1830. P. .laliscana S. Watson, Proc. Am. Acad. 22:456. 1887. 
Mexico. 

222. P. MILITARIS L. B. Smith, Contr. Gray Herb. l6l:34, pl. 
4, fig. 5. 1946. Mexico. 

223. P. (subgenus ?) FOLIACEA L. B. Smith, sp. nov. P. sordi - 
da L. B. Smith in systema mea proxima sed scapi bracteis inferi- 
oribus foliaceis perelongatis differt. 

Short-caulescent, flowering 4 dm high; leaves subbulbous-rosu- 
late; sheaths broadly ovate, ca. 2 cm long, dark castaneous; 



196U Smith, Notes on Brcaaeliaceae li3 

blades dimorphic, some reduced to dark pectinate-serrate spines, 
others foliaceous, deciduous, linear, acuminate, slightly narrow- 
ed toward base, about equaling the inflorescence, 14 mm wide, 
sparsely white-flocculose at base, entire above the line of ab- 
scission; scape erect, slender, white-flocculose; scape-bracts 
erect, all eocceeding the internodes, the lower foliaceous and 
equaling the leaves, the upper ovate, acuminate, scarcely larger 
than the floral bracts; inflorescence simple, laxly few-flowered, 
white-floccvilose, the islender axis 7 cm long; floral bracts 
ovate, acuminate, to 22 Him long, much exceeding the slender te- 
rete 7 mm pedicels, entire; flowers suberect at anthesis, not se- 
cund; sepals linear-lanceolate, acuminate, 23 mm long, the poste- 
rior ones carinate at base; petals 5 cm long, naked, red; ovary 
3/4 superior. PI. I, fig. 16: Flower x 1/2; fig. 17: Sepal x 1. 
UEXICO: Uichoacan: Dist. Coalcoman: On tree, Naranjillo. alt. 
1250 ra, February 8, 1941, G. B. Hinton no. 15941 (US, type). 

224. P. SORDIDA L. B. Smith, Contr. Gray Herb. 161:34, pl. 4, 
fig. 6. 1946. Mexico. 

225. P. 1I0NTICOLA Brandegee, Zoe 5:197. 1905. Mexico. 

226. P. RINGENS Kl. ex Lk., KL. & Otto, Ic. PI. Rar. 63, pl. 
25. 1842. P. latifolia Wendl. Hort. Herrenh. 1:5, pl. 3- 1798, 
non Ait. 1789, cf. Hez in Pflanzenreich 4, Fam. 32:266. 1935. 

P. montalbenais hort. Linden ex Otto & Dietr. Allgem. Gartenzeit, 
19:138. 1851. P. warszewitciana Kl. ex Beer, Brooel. 66. 1857. 
P. karwinskyana Beer, Bromel. I6l. 1857. P. fulgens hort. ex 
Baker, Handb. Bromel. 101. 1889, nomen. Mexico. 

227. P. CHIAPENSIS Miranda, An. Inst. Biol. Mexico 24: L69j , 
fig. 1. 1953. Mexico. 

228. P. OAXACANA L. B. Smith, Contr. Gray Herb. 117:25, pl. 2, 
figs. 16, 17. 1937. Mexico. 

229. P. SECUNDIFLORA L. B. Smith, Contr. Gray Herb. 114:7, pl. 
1, figs. 1, 2. 1936. Mexico. 

230. P. CARNEA Beer, Oesterr. Bot. Zeitschr. 8:182. 1858. 
Puya camea Regel, Cat. Pl. Hort. Aksak. 117. I860. Panama. 

231. P. XANTHOCALYX Mart. Hort. Monac. Sem. for 1848:4. 1848; 
Linnaea 24:195. 1851. P. sulphurea sensu K. Koch, Ind. Sem. 
Hort. Berol. for 1857 :App. 5. 1858, non Andr. 1802. P. flaves- 
cens Baker, Bot. Mag. 103 :pl. 63I8. 1877, non K. Koch7 1858. 
Mexico. 

West Indies 
(cf. no. 71) 

232. P. SAMUELSSONn L. B. Smith, Contr. Gray Herb. 117:28, 
pl. 2, figs. 24-26. 1937. P. xanthocalyx sensu Mez in Urb. Symb. 
Ant. 8:87. 1920, non Mart. 1848. Hispaniola. 

233. P. BRDMELIIFOLIA L'Herit. Sert. Angl. 7. 1789. Hepetis 
angustifolia Sw. Prodr. 56. 1788, non Pitcaimia angusti folia 
Ait. 1789. H. bromeliifoUa Salisb. Prodr. 247. 1796. Pitcair- 
nia redouteana Beer, Bromel. 57. 1857, non Schult. I83O. Jamaica 

234. P. ANGUSTIFOLIA CSoland. in3 Ait. Hort. Kew. 1:401. 1789. 
P. angusti folia Redoute, Lil. 2:pl. 76. 1804, non Ait. 1789. P. 



hk PHYTOLOGIA Vol. 10, no. 1 

furfuracea sensu Jacq. f . Eclog. PI. 1:117. 1815, non Willd. 1809 
P. ramosa Jacq. f . Eclog. PI. 1:154. 1816. P. tomentosa Dietr. 
Lex. Nachtr. 6:305. 1820, nomen; Dietr. ex Beer, Brcoel. 64. 1857 
P. rfldouteana Schult. in R. & S. Sjst. 7, pt. 2:1243. 1830. P. 
angiistifolia Ryan ax -Schult. 1. c, nomen. P. ramosa K. Koch, 
Ind. Sem. Hort. Berol. for 1857:App. 5. 1858, non Jacq. 1816. P. 
intermedia hort. ex K. Koch, 1. c. P. skinneri hort. ex K. Koch, 
1. c. P. alta sensu Mez in DC. Mon."Phan. 9:406. 1896, non 
Hassk. 1856. Hepetis angustifolia sensu Mez, op. c. 973, non Sw. 
1788. Puerto Rico, Lesser Antilles. 

235. P. FUERTESII Mez, Fedde Rep. Spec. Nov. 12:415. 1913. P. 
tomentosa sensu Mez in DC. Mon. Phan. 9:389. 1896, non Dietr. ex 
Beer 1857. P. fulgens Mez in Urb. Symb. Ant. 8:87. 1920, 

non Dene, ex Dietr. 1851. Dominican Republic. 

236. P. GRACILIS Mez in DC. Mon. Phan. 9:407. 1896. Lesser 
Antilles. 

237. P. JIMENEZII L. B. Smith, Phytologia 7:1, pi. 1, figs. 
3-5. 1959. Dominican Republic. 

238. P. PLATTPHYLLA Schrad. Blumenb. 26. 1827. P. bromelii- 
folia sensu Ait. Hort. Kew. 1:401. 1789, non L'Herit. 1789. P. 
latifolia Andr. Bot. Repos. 5:pl. 322. 1803, non Ait. 1789. 
Jamaica. 

239. P. CUBENSIS (Mez) L. B. Smith, Contr. Gray Herb. U7:24, 
pi. 2, figs. 12, 13. 1937. P. latifolia var. cubensis Mez in DC. 
Mon. 9:396. 1896. Cuba. 

240. P. LATIFOLIA Ait. Hort. Kew. 1:401. 1789. Hepetis lati- 
folia Raeusbhel ex Schult. in R. & S. Syst. 7, pt. 2:1247. 1830, 
nomen. Billbergia latifolia hort. Belg, ex Heynh, Norn. 2:69. 
1846, nomen. Pitcairnia furfuracea Beer, Bromel. 59. 1857. P. 
latifolia Jacq. ex Beer, op. c. 61. P. alta sensu Baker, Bot, 
Mag. 108:pl. 66O6. 1882, non Hassk. I856. Hepetis pyrtunidata 
Rich, ax Baker, Handb. Bromel. 92. 1889, nomen. Puerto Rico (?), 
Lesser Antilles. ^ 

2a. P. SPICATA (Lam.) Mez in DC. Mon. Phan. 9:392. 1896. • 
Lesser Antilles. 

Var. a. SPICATA. Bromelia spicata Lam. Encycl. 1:146. 1783. 
Pitcairnia latifolia Redoute, Lil. pi. 74. I8O4, non Ait. 1789. 
P. bract eat a ot. Ait. Hort, Kew. ed. 2, 2:202. 1811. P. racemosa 
Woodf. ex Schult. in R. & S. Syst. 7, pt. 2:1245. 1830, nomen. 
P. fulgens Dene, ex A. Dietr. Allgem. Gartenzeit. 19:25, I85I, 
P. gireoudiana A. Dietr. Allgem. Gartenzeit. 21:105. Ap. 1853. 
P. bracteata var. fulgens Regel, Gartenfl. 2:163. 1853. ?Bill- 
bergia bifrons Lindl. Joum. Hort, Soc, London 8:54. June I853. 
P. bracteata var. gireaudiana Beer, Bromel. 51. 1857. Billbergia 
pyramidata Beer, op, c. I23. Pitcairnia conmutata Regel, Garten- 
fl. 16:289. I867. P. bracteata var. commutata Regel, op. c. 17: 
8. 1868. Petali red. 

Var. b. SULPHUREA (Andr.) Mez in DC. Mon. Phan. 9:393. 1896. 
P. sulphurea Andr. Bot. Repos. 4:pl. 249. 1802. P. bracteata /9. 
Ait. Hort. Kew. ed. 2, 2:202. 1811. P. bracteata /S. sulphurea 
Ker-Gawl. Bot. Mag. 34:pl. 1416. 18117 Tillandsia vineentiensis 
E. H. L. Krause, Beih. Bot. Centr. 32, pt. 2:337. 1914. Petals 



196U Smith, Notes on Bromellacaae U5 

yellow. 

242. P. ALBUCIFOLIA Schrad. Blumenb. 24. 1827. ? P. spec . 
fol . linearl-lanc . etc. Voigt, PI. Rar. Hort. Belved.~in Sylloge 
2:52. 1812 (?"5T^f. R. & S. Syst. 7, pt. 2:1250. I83O. P. fur- 
furacea Sims, Bot. Mag. 53:pl. 2657. 1826, non Willd. 1809. P. 
Intennedia hort. ex SIjbs, 1. c. nomen. P. ringena sensu Beer, 
Bromel. 55. 1857, non Kl. 1842. P. affinla K. Koch, Ind. Sem. 
Hort. Berol. for 1857:App. 5. I858. P. angustifolla E. Morr. ex 
Baker, Handb. Brooel. 92. 1889, nomen. Lesser Antilles. 

243. P. (subgenus ?) GLTMIANA K. Koch, Wochenschr. oSrtn. 11: 
89. 1868. Hechtia glyaiana K. Koch, op. c. 90, nomen. Puya 
glymlana K. Koch, 1. c. nomen. Pltcairnia finaa Baker, Joum, 
Bot. 19:268. 1881. P. .jaeksonl K. Koch ex Badcer, 1. c. nomen. 
West Indies (?). 

244. P. ELIZABETHAE L. B. Smith, Bromel. Soc. Bull. 8:21, 
figs. 1958. Dominican Republic. 

245. P. DOUINQENSIS L. B. Smith, sp. nov, P. .limenezii L. B, 
Smith in systema mea prcocima sed foliorum laminis dimorphis 
differt. 

Nearly stemless, flowering 6 dm high; leaves many in a fasci- 
culate rosette, the larger ones to over 8 dm long; sheaths ovate, 
3-5 cm long, dark castaneoua, entire; blades dimorphic, some re- 
duced to subulate entire dark spines, others foliaceoua lineair, 
long-acuminate, slightly narrowed toward base, 15-20 mm wide, 
glabrous, sparsely serrate toward base; scape erect, slender, 
sparsely white-flocculoae; scape-bracts erect, the lower folia- 
ceoua and exceeding the intemodes, the upper narrowly triangu- 
lar, caudate-acuminate, shorter than the intemodes; inflores- 
cence simple, subdense at anbhesis, sparsely white-flocculose, 
the axis 7 cm long; floral bracts like the upper scape-bracts, 
the lower much exceeding the pedicels, the upper shorter; pedi- 
cels alender, to 8 mm long; sepala linear, obtuse, 19 nm long, 
ecarinate; petals linear, obtuse, 4 cm long, red, bearing an ob- 
long subtruncate 5 am long scale at base; stamens included; ovary 
5/8 superior; ovules caudate (?). PI. I, fig. 18: Flower x 1/2; 
fig. 19: Sepal x 1. 

D(»iINICAN REPUBLIC: Samana: Bahia de San Lorenzo, Bahia de Sa- 
mana, ciiltivated and flowered by L. Ariza Julia in 1961, J. J. 
Jimenez no. UUUU (US, type; hb, Jimenez, isotype). 



Amazon-Orinoco Basin 



246. P. UNDUUTA Scheidw. Allgem. Gartenzeit. 10:275. 1842. 
P. undulatifolia hort. ex Lem. Fl. des Serres 2: sub pi. 162. 
I846, nomen. P. specioaisaima hort. ex Regel, Gartenfl. 23:1. 
1874» nomen. Lamproconua undulatus Lem. Jard. Fleur. 2:aub pi. 
127. I852. Amazonian Brazil (?). 

247. P. EGLERI L. B. Smith, Phytologia 8:228, pi. 2. figa. 
11-14. 1962. Brazil: Para. 



U6 PHTTOLOGIA Vol. 10, no. 1 

Planalto of Brazil 

2A8. P. ULEI L. B. Smith, Bol. Mus. Nac. Rio de Janeiro n. 
ser. no. 15:5, pl. 1, figs. h-j. 1952. 

249. P. (subgenus ?) ANTHERICOIDES Mez in Mart. Fl. Bras. 3, 
pt. 3:4a. 1894. 

250. P. TORRES lANA L. B. Smith, Bol. Mus. Nac. Rio de Janeiro 
n. ser. no. 15:4, pl. 1, figs, f, g. 1952. 

251. P. DECIDUA L. B. Smith, Arquiv. Bot. Estado Sao Paulo n. 
ser. 1:110, pl. 114. 1943. 

252. P. GLAZIOVII Baker, Handb. Bromel. 92. 1889. 

253. P. LANG IFOLIA Mez iwMart. Fl. Bras. 3, pt. 3:447. 1894. 
Var. a. LANCIFOLIA. Sepals 35 nun long.' 

Var. b. MINOR^ L. B. Smith, Bol. Mus. Nac. Rio de Janeiro n. 
ser. no. 15:4. 1952. Sepals 24 mm long. 

254. P. ENCHOLIRIOIDES L. B. Smith, Arquiv. Jard. Bot. Rio de 
Janeiro 10:146, fig. 6. 1950. 

255. P. CARINATA Mez in Mart. Fl. Bras. 3, pt. 3:448. 1894. 
P. morelii sensu Baker, Handb. Bromel, 103. 1889, in part, non 
Lem. 1846. 

256. P. FLAMMEA Lindl. Bot. Reg. 13:pl. 1092. 1827. 

Var. a. FLAMMEA. ? Tillandsia laevls Veil. Fl. Flum. 133. 
1825; Icon. 3:pl. 126T 1835, non Pitcaimia laevis Willd. 1830. 
P. olfersii Link, Verh. Gartenbauver. Berlin 7:363, pl. 3. 1831. 
? P. fulgens Poit. Rev. Hortic. 3:157. Jan. I836; Mez in DC. Mon. 
Phan. 9:430. 1896. P. morelii Lem. Hortic. Univ. 7:231, pl. 
1846. P. fulgens Dene, in Cat. Liixien 18. I85O; Dene, ex A. 
Dietr. Allgem. Gartenzeit. 19:137. 1851. ? P. rubicunda K. Koch 
& Bouche, Ind. Sem. Hort. Bgrol. for 1856:App. 3. 1857. ? P. mo- 
reliana hort. ex K. Koch & Bouche, 1. c, nomen. ? P. laevis 
Beer, Bromel. 60. 1857. P. decaisnei K. Koch, Ind. Sem. Hort. * 
Berol. for 1857:i^p. 5. 1858. ? P. I'herminieri hort. Paris ex 
K. Koch, Ind. Son. Hort. Berol.~for 1857:App. 8. 1858. P. roez- 
lii sensu Baker, Bot. Mag. 117 :pl. 7175. 1891, non E. Morr. 1885. 
? P. amaryllidiflora hort. ex Gentil, Pl. Cult. Serres Brux. 153 . 
1907, nomen, cf. Mez in Pflanzenreich 4, Fam. 32:260. 1935. ? P. 
mordli hort. ex Gentil, 1. c. Leaf -blades mostly 20-36 ram wide, 
coveired beneath with spreading scales; axis of the inflorescence 
glabrous, usually turning black on drying. 

Var. b. ROEZLII (E. Morr.) L. B. Smith, Arquiv. Bot. Estado 
Sao Paulo n. ser. 1:111. 1943. P. roezlii E. Morr. Belg. Hortic. 
35:285, pis. 18, 19. I885. P. hypoleuca Mez in Mart. Fl. Bras. 
3, pt. 3:458. 1894. Leaf-blades mostly 20-36 mm wide, covered 
beneath with spreading scales; axis of the inflorescence lepi- 
dote, usually remaining pale. 

Var. c. CORCOVADENSIS (Wawra) L. B. Smith, Arquiv. Bot. Estado 
Sao Paulo n. ser. 1:112. 1943. P. corcovadensis Wawra, Oesterr. 
Bot. Zeitschr. 12:384. 1862. Leaf -blades glabrous; inflorescence 
lax, few-flowered, remaining pale; axis glabrous. 

Var. d. GLABRIOR L, B. Smith, Arquiv. Bot. Estado Sao Paulo n. 
ser. 1:112. 1943. ? P. cinnabarina A. Dietr. Allgem. Gartenzeit. 
18:202. 1850. ? P. 'aust rails K. Koch, Ind. Sera. Hort. Berol. for 



196U Smith, Notes on Bromellaceae 1^7 

1856:App. if. 1857. Leaves glabrous; inflorescence dense at least 
toward apex, many- flowered, remaining palej axis glabrous; petals 
red. 

Var. e. PALLIDA L. B. Smith, Arquiv. Bot. Estado Sao Paulo n. 
ser. 1:112, pi. 115. 1943. Leaves glabrous; inflorescence dense 
at least toward apex, many-f lowered, remaining pale; axis gla- 
brous; petals yellowish white. 

Var. f. FLOCCOSA L. B. Smith, Arquiv. Bot. Estado Sao Paulo n. 
ser. 1:112. 1943. f. miscosa Mart, in R. & S. Syst. 7, pt. 2: 
1240. 1830. P. selloana Baker, Handb. Bromel. 100. 1889. P. 
dietrichiana Wittm. Bot. Jahrb. 13, Beibl. 29:15. 1891. P. prui- 
nosa Mez in Mart. Fl. Bras. 3, pt. 3:454. 1894, non H.B.kT 1816. 
P. claussenii Mez, 1. c. P. selloviana Mez in DC. Mon. Phan. 9: 
433. 1896. P. weddelliana Mez in DC. Mon. Phan. 9:434. 1896, non 
Baker 1889. P. minarum Mez in Pflanzenreich 4, Fam. 32:636. 
1935. Leaf -blades less than 20 mm wide, lepidote; inflorescence 
remaining pale; axis lepidote, 

257. P. ALBIFLOS Herb. Bot. Mag. 53:pl. 2642. 1826. Tilland - 
sia schuechii Beer & Fenzl, Allg, Gairtenzeit. 14:265. 1846. P. 
odorata hort. ex Beer & Fenzl, op. c. 266, nomen. P. elata 
Liebm. Ind. Sem. Hort. Haun. 14. 1849. Cochliopetalum albifloa 
Beer, Bromel. 68. 1857. C. f laves cens Beer, op. c. 69. C. 
schtlchii Beer, 1. c. Pitcaimia flavescens hort. ex Beer, 1. c, 
nomen; K. Koch, Ind. Sem. Hort. Berol. for 1857:App. 9. 1858. 
Cochliopetalum odoratum Hemsl. Biol. Centr.-Am. Bot. 3:317. 1884, 
nomen, erroneously attributed to Beer. P. xanthocalyx sensu 
Baker, Handb. Bromel. 107. 1889, in part, as to Cbchliopetalum 
flavescens Beer. 

258. P. STAMINEA Lodd. Bot. Cab. 8:pl. 722. 1823. P. speclosa 
hort. Lovan ex Schult. in R. & S. Syst. 7, pt. 2:1250."l83O, no- 
men; cf. Mez in Pflanzenreich 4, Fam. 32:262. 1935. Cochliopeta- 
lum stamineum Beer, Bronel. 70. 1857. Orthopetalum stamineun 
Baker, Handb. Bromel. 97. 1889, nomen attributed to Beer. Plt- 
cairnia caniliculata Baker, op. c. 99. P. longlcauda Homem. ex 
Mez in Mart. Fl. Bras. 3, pt. 3:445. 1894. P. staminea var. lon- 
gi Cauda Mez in DC. Mon. Phan. 9:439. 1896, attributed to Homem. 

259. P. BEYCALEMA Beer, Bromel. 63. 1857. P. muscosa sensu 
Hook. Bot. Mag. 80:pl. 4770. 1854, non Mart. 1830. P. leiolema 
hort. ex Beer, Bromel. 62. 1857, n(aaen. 

260. P. SUAVEOLENS Lindl. Bot. Reg. 13: pi. 1069. 1827. P. 
odorata Wawra, It. Sax. -Cob. 174. 1883. 

West Africa 

261. P. FELICIANA (Aug. Chevalier) Harms & Mildbr. Notizblatt 
14:118. 1938. Willrussellia feliciana Aug. Chevalier, Bull. Soc. 
Bot. France 84:503, fig. 1. 1937. [French Guinea). 

EXCLUDED AND DOUBTFUL TAXA 

262. P. alpestris (Poepp. & Endl.) L. H. Bailey, Cyclop. An. 
Hort. 1359. 1901 = PUYA ALPESTRIS Poepp. & Endl. 



U8 PHTTOLOGIA Vol. 10, no. 1 

263. P. bangll Baker, llem. Torrey Bot. Club 6:124. 1896 ■ PUYA 
STENOTHYRSA (Baker) Mez. 

264. P. brachlata Cham, in Link, Jahrb. 1, pt. 2:192. 1820, 
nomen, Unknown. 

265. P. brachyataehya Baker, Handb. Bromel. 118. 1889 ■ PUTA 
BRACHYSTACHYA (Baker) Mez . 

266. P. brevlfolla (Griseb.) R. E. Fries, Nov. Act. Reg. Soc. 
Sci. Upsal. ser. 4, 1, pt. 1:73. 1905 = ABROMEITIELU BREVIFOLIA 
(Griseb.) Castellanos. 

267. P. brocchlnia D. Dietr. Syn. PI. 2:1062. 1840 = BROCCHI- 
NIA PANICUUTA Schult. f . 

268. P. bromeliifolia var. graminifolia Griseb. Fl. Brit. W. 
Ind. 594. I864. Unknown . No species known from Jamaica has en- 
tire leaves 4 lines (ca. 8 mm) wide. 

269. P. caerulea - of. coenilea . 

270. P. chllensis Lodd. Cat. ex Loudon, Hort. Brit. 118. I83O, 
nomen = PUYA CHILENSIS Mol, 

271. P. chlorantha (Spegazz.) Castellanos, Com. Uus. Nac. 
Buenos Aires 2:142. 1925 ■ ABROMEITIF.TJA BREVIFOLIA (Griseb.) 
Castellanos. 

272. P. ehrysantha Phil. Fl, Atacam. 50. 1860 = DEUTEROCOHNIA 
CHRYSANTMA (Phil.) Mez. 

273. P. ciliaris hort. ex Paeq. Cat. Ort. Bot, N^. 81. 1867, 
nomen . Unknown . 

274. P. coarctata Pers. Syn. 1:344. 1805 ■ PUYA CHILENSIS Mol. 

275. P. coenaea Benth. eoc Baker, Handb. Bromel. 121. 1889* 
(first valid combination) • PUYA COERUIEA Lindl. 

276. P, coerulea sensu Baker, Handb. Bronel. 121. 1889, in 
part, as'to PUYA ALPESTRIS (Poepp. & Endl.) Gay and its synonym, 
P. whytei Hook. f. 

277. P. crystalina Pers. Syn. PI. 1:344. 1805 = PUYA LANUGINO- 
SA (R. & P.) Schult. f. 

278. P. darblayana Andre, Rev, Hortic. 62:33, fig. 1890 « p, 
CORALLINA X PANICULATA, 

279. P. decora hort. Linden ex Beer, Bromel. 108. 1857, 
nomen = BTT.LRF.RGIA AMOENA var. MINOR (Antoine & Beer ex Beer) L. 
B. Smith ? 

280. P. discolor Loisel. Herb. Gen. Amat. 5:pl. 345. 1821 = 
BILLBERGIA AMOENA (Lodd.) Undl. var. AMOENA. 

281. P. distacalB Beer, Bromel. 58. 1857 - BILLBERGIA DISTA- 
CHIA (Veil.) Mez var. DISTACHIA. 

282. P. dyckioides Baker, Handb, Bromel. 118. 1889 = PUYA 
DYCKIOIDES (Baker) Mez 

283. P. elygans Regel, Cat. Hort. Aksak. 112. I860, nomen. 
Unknown. 

284. P. fastuosa C. Morr. Ann. de Gand. 35411, pl. 161. 1847 = 
BILLBERGIA PYRAMIDALIS (Sims) Lindl. var. PYHAMIDALIS. 

285. P. flabelliformis hort. ex Gentil, PI. Cult. Serres Jard. 
Bot. Brux. 153. 1907, nomen. Unknown. 

286. P. floccosa Regel, Gartenfl. 23:307. 1874; Act. Hort. 
Petrop. 3:124. 1875 = PUYA FLOCCOSA (Linden) E. Morr. 

287. P. formosa Mez in Pflanzenreich 4, Fam. 32:296, 654. 



196U Smith, Notes on Brcwiellaceae ll9 

1935, erroneously attributed to Spegazzini » PUYA SPATHACEA 
(Grieeb.) Mez. 

288. P. furfuracea Willd. Enom. 1:346. 1809 - PUTA FURFURACEA 
(Willd.rL. B. Smith. 

289. P. ^andiflora Mez, Fedde I^p. Nov. Spec. 3:5. 1906, non 
Hook. 1861 = PUYA PITCAIRNIOIDES L. B. Siaith. 

290. P. guyanensis Baker, Handb. Bromel. 120. 1889 = PUTA 
FLOCCOSA (Linden) E. Uorr. 

291. P. humilis hort. ex Sweet, Hort. Brit. ed. 1, A25. 1827, 
noBien. Unknown. 

292. P. iridiflora Beer, Bromel. 51. 1857. Description inade- 
quate . 

293. P. lanata F. G. Dietr. Lexicogr. Nachtr. 6:303. 1820 ■ 
PUTA LANATA (H.B.K.) Schult. f. 

294. P. lorentziana Mez in DC. Mon. Phan. 9:373- 1896 = ABRO- 
MEITIELLA LORENTZIANA (Mez) Castellanos. 

295. P. macrophylla Willd. ex R. & S. Sjst. 7, pt. 2:1283. 
1830 = AECHMEA LATIFOLIA (Willd. ex Schult.) Kl. ex Baker. 

296. P. maronii Andre, Rev. Hortic. 56:222. 1884; 57:108, fig. 
1885 = pT CORALLINA X ALTENSTEINII. 

297. P. media hort. ex Sweet, Hort. Brit. ed. 1, 425. 1827, 
nomen. Unknown, 

298. P. megastachya Baker, Handb. Bromel. 120. 1889 = PUTA 
ROEZLII E. Morr. 

299. P. meridengja hort. ex Baker, Handb. Bromel. 121. 1889, 
nomen = PUTA FLOCCOSA (Linden) E. Morr. 

300. P. micrantha Lindl. Bot. Reg. 29:Mi8c. 44. 1843 ■ FOS- 
TERELLA MICRANTHA (Lindl.) L. B. Smith. 

301. P. monstrosa Beer, Bromel. 68. 1857. Description inade- 
quate . 

302. P. olivacea (Wittm.) Mez in Pflanzenr*ich 4, Fam. 32:247. 
1935 - PUTA OLIVACEA Wittm. 

303. P. pasto&naia Baker, Handb. Bromel. 122. 1889 = PUTA LEH- 
MANNIANa'l. B. Smith. 

304. P. pearcei Baker, Handb. Bromel. 120. 1889 = PUTA PEARCEI 
(Baker) Mez 

305. P. penduliflora Mez in DC. Mon. Phan. 9:463. 1896 = GLO- 
MEROPITCAIRNIA PENDULIFLORA (Griseb.) Mez. 

306. P. penduliflora A. Rich, in Sagra Hist. Cuba 11:262. 
1850 = HOHENBERGIA PENDULIFLORA (A. Rich.) Mez. 

307. P. philippii Baker, Handb. Bromel. 122. 1889 = PUTA VIO- 
LACEA (Brorign.) Mez. 

308. P. plumieri Baker, Handb. Bromel. 107. 1889 = GUZMAN lA 
PLUMIERr( Griseb.) Mez . 

309. P. pyramidata Link, Entim. 1:308. 1821, as to material, 
not as to basonym = PUTA FURFURACEA (Willd.) L. B. Smith. 

310. P. pyramidata (R. & P.) Pers. Syn. PI. 1:344. I8O5 = PUTA 
PTRAMIDATA (R. & P.) Schult. f. 

311. P. quetameensis Baker, Handb. Bromel. 121. 1889 " PUTA 
FLOCCOSA" (Linden) E. Morr. 

312. P. regia Witte. Tijdschr. Tuinb. Ed. Bos. 5:pl. 3, fig. 
34. 1900; Gentil, PI. Cult. Serres Jard. Bot. Brux. 152. 1907 « 



50 PHYTOLOGIA Vol. 10, no. 1 

P. CORALLINA X SPICATA. 

313. P. robusta Rusby, Bull. New York Bot. Gard. 6: ASS. I9IO = 
PUYA SANCTAE-CRUCIS (Baker) L. B. Smith. 

314. P. rusbyl Baker, Handb. Brcmel. 122. 1889 « PUYA RUSBYI 
(Baker) Uez. 

315. P. sanctae -crucls Baker, Handb. Bromel. 120. 1889 = PUYA 
SANCTAE-CRUCIS (Baker) L. B. Smith. 

316. P. secxmda F. G, Dietr. Lexicon. Nachtr. 6:301. 1820 = 
TILLANDSIA SECUNDA H.B.K. 

317. P. spathacea Griseb. Goett. Abh. 24:329. 1879 = PUYA 
SPATHACEA (Griseb.) Mez. 

318. P. spathulata hort. «x Lem. Fl, des Serres 3spl. 227. 
1847 = aScHMEA BASI-LATERALIS (Lem.) L. B. Smith 

319. P. aphaerocephala Baker, Handb. Brranel. 123. 1889 = PUYA 
VENUSTA Phil. 

320. P. spinoaa Gill, ex Baker, Handb. Bromel. I36. 1889, 
nomen = DYCKIA FLORIBUNDA Griseb. 

321. P. splendens Poir. in Rev. Hort. ser. 1, 3:157. I836. 
Not understood. May equal PITCAIRNIA FLAMMEA var. GLABRIOR L. B. 
Smith. 

322. P. atenothyrsa Baker, Handb. Bromel. 122. 1889 = PUYA 
STENOTHYHSA (Baker) Mez . 

323. P. atricta Andre, Bromel. Andr. 3I. 1889. Sterile, 
description inadequate. 

324. P. vallesoletana Leocarza in La Llave & Lexarza, Nov. Veg. 
Descr. fasc. 1:19. 1824. Description inadequate. Probably re- 
lated to P. RINGEMS Kl. 

325. P. venusta Baker, Handb. Bromel. 123. 1889 = PUYA VENUSTA 
Phil. 

326. P. violacea Brongn. Ann. Fl. & Pom. ser. 3, 1:116. 1847; 
Allgem. Gartenzeit. 15:299. 1847 = PUYA VIOLACEA (Brongn.) Mez. 

327. P. virescens K, Koch, Ind. Sem. Hort. Berol. for 1857: 
App. 4. 1858 = GUZMANIA VIRESCENS (Hook, f .) Mez. 

328. P. viridiflora Regel, Ind. Sem. Hort. Petrop. for 1866: 
81. 1867""= VRIESEA VIRIDIFLORA (Regel) Wittm. ex Mez. 

329. P. weddelliana Baker, Handb. Bromsl. 122. 1889 - PUYA 
WEDDELLlBiA (Baker) Mez . 

INDEX 
(by species numbers of Pitcaimia ) 

ABROMErriELLA brevifolia 266, 271; lorentziana 294. 

AECHMEA basi-lateralis 318; latifolia 295. 

BILLBERGIA amoena var. amoena 280; var. minor 279; bifrons 
241 ; distachia 281; latifolia 240; pyramidalis var. pyramidalis 
284; pyramidata 241. 

BROCCHINIA paniculata 267- 

BRDMELIA spicata 241. 

COCHLIOPETALUM alhiflos 257; flavescens 257; odoratum 257; 
schilchii 257; stamineura 258. 

DEUTEROCOHNIA chrysantha 272. 

DYCKIA floribunda 320. 



196U Smith, Notes on Bromellaceae 51 

FOSTERELLA micrantha 300. 

GLOMEROPITCAIRNIA penduliflora 305. 

GUaiANIA plumieri 308; virescena 327. 

HECHTIA glymlana 243. 

HEPETIS anguatifolia Mei 234; angusttfolla Sw. 2335 bromelil- 
folia 233; latifolia 240; pyramldata 240; all other combinations 
made wholesale by Uez in DC. Mon. Phan. 9: C952J, 973, 974. 1896. 

HOHENBERGIA penduliflora 306. 

LAMPROCONUS altensteinii 115; aurantiacus 200; giganteus 115b; 
Jackaoni 42; maidifoliua 90; undulatus 246; warazcewiczii 103. 

MELINONIA incamata 25; rubiginoaa 15. 

NEUMANN lA altenateinii 115; arcuata 108; atrorubena 103; gi- 
gantea 115b; imbricata 196; lindeni 103; maidifolia 90; nigra 96; 
ochroleuca 201; petiolata 197; aulphurea 193. 

ORTHOPETALUM inerme 55; pulverulent um 48; stamineum 258. 

PEPINIA aphelandriflora 84; incamata 25; punicea 42; recur- 
vata 190. 

PHLOMOSTACHYS altenateinii 115; atrorubena 103; densiflora 
200; funkiana 90; gigantea 115b; Imbricata 196; sulphorea 193. 

PrrCAIRNIA acicularis 136; adacendena 160; aequatorialia 118; 
af finis 242; agavifolia 1; alata 132; albifloa 257; alborubra 38; 
albucifolia 242; alpestris 262; alta Baker 240; alta Haaak. 71; 
alta Uez 234; altenateinii Donn. Sm. 193; altenateinii Lem. 115> 
296; var. gigantea 115b; amaryllidiflora 256; amazonica 15b; am- 
blyosperma 43; andreana 181; anguatifolia Ait. 234; anguatifolia 
E. Morr. 242; anguatifolia Redoute 234; anguatifolia Ryan 234; 
anguatifolia Soland.234; anomala 23; anthericoidea 249; aphelan- 
driflora 84; araneoaa 66; archeri 87; arcuata 108; arenicola 56; 
armata 8; aaplundii 113; aaterotricha 34; atrorubena 103; ■»ar. 
lanarcheana 103; augustii 117; aurantiaca 200; auriculata 58; 
auatralia 256d. 

bakeri 106; bangii 263; barrigae 100; bella 149; var. denaior 
149b; beycalena 259; biattenuata 49; biflora 143; billbergioidea 
124; brachiata 264; brachyaperma 163; brachyatachya 265; bractea- 
ta A . 24I; bracteata/^ . 241b; var. commutata 241; var. fulgena 
24I; var. gireaudiana 241; var. aulphurea 241b; bradei 33; brevi- 
calycina 179; brevifolia 266; breweri 9; brittoniana 95; brocchi- 
nia 267; bromeliifolia Ait. 238; bromeliifolia L'Herit. 233; var. 
graminifoHa 268; brongniartiana 98; var. latifolia 98b; brunnea- 
cena 99; bulbosa 6; burchellii. 60. 

caerulea 269; calatheoidea 157; caldaaiana 60; calderonii 205; 
calophylla I5O; campii 104; camptocalyx 66; var. lutea 66; var. 
robuata 66; canaliculata 258; capitata 85; caracaaana 184b; car- 
denaaii 130; caricifolia 24; var. macrantha 24b; carinata 255; 
carioana 198; cariovana 198; carnea 230; cassaponala 72; caulea- 
cena 183; ceniua ll6bb; chiapenaia 227; chilenaia 270; chiriguana 
62; chiriquensia 186; chlorantha 271; chocoenaia 64; chryaantha 
272; ciliaria 273; cinerea 8; cinnabarina 256d; clauaaenii 256f; 
clavata 101; coarctata 274; coerulea Benth. ax Baker 275; coeru- 
lea aensu Baker 276; ccmmixta 57; conmutata Regel 241; concolor 
123; consimilia 34; corallina 19, 278, 296, 312; var. viridis 
19b; corcovadenais 256c; coatata 41; cotahuaaiana 34; craasa 119; 



$2 PHTTOLOQIA Vol. 10, no. 1 

crystallina 277; ctenophylla 27 j cuatrecasana 17; cubensis 239; 
cuzcoensls 74; cyauiopet^a 162; cyllndrostachya 208. 

darblayana 278; decaienel 256; decidua 251; decora Dietr. 71j 
decora hort. 279; decurvata 127; dendroidea 52; densiflora 200; 
devansayana 79; dietrichlana 25 6f; diffusa 80; discolor 280; dis- 
tacaia 281; divaricata 83; dollchopetala 152; domlngensls 245; 
djckioldes 282. 

echlnata 44; var. sublaevls 44b; var. vallensls 44c; echino^ 
trlcha 34; egleri 247; elata 257; elegans 283; ellzabethae 244; 
elliptica 141; elongata 105; encholirloides 254; ensifolia 30; 
epipbytlca 16; erratlca 135; excelsa 49; eximia 166; exscapa 
Hook. Il6bb; exscapa Llebn. 116; exserta 78. 

fastuosa 284; feliciana 261; fendleri 173; ferreyrae 93; fer- 
ruginea 34; filispina 28; firma 243; flabelliformis 285; flagel- 
laris 187; flannnea 256; var. corcovadensls 256c; var. floceosa 
25 6f; var. glabrior 25 6d; var. pallida 25 6e; var. roezlii 256b; 
flavescens Baker 231; flavescens hort. 257; flaviflora 95; flexu- 
osa 214; floceosa 286; foliacea 223; formosa 287; fosteriana 109j 
fractlfolia 129; fruticetorum 49; fuertesii 235; fulgens Dene. 
241, 256; fulgens A. Dietr. 203; fulgens hort. ax Baker 226; ful- 
gens Uez 235; fulgens Poit. 256; funkiana 90; fimkii 90; furfura- 
cea Beer 240; furfuracea Jacq. f . 234; furfuracea Sims 242; fur- 
furacea Willd. 288; 

geyskesii 10; gireaudiana cf. 241; gireoudiana 241; glaziovii 
252; glymiana 243; goudotiana 66; gracilis 236; graminea 71; gra- 
minifolia 71; grandiflora 289; gravisiana 96; grubbiana 182; gua- 
ritermae I56; guyanensis 290; guzmanioides 86. 

halophila 77; harlingii lU; hartmannii 71; haughtii 65; helio- 
phila 7; hemsleyana 195; herrerae 34; heterophylla 116; forma al- 
biflora 116b; var. exscapa ll6bb; hintoniana 212; hitchcockiana 
111; humilis 291; hypoleuca 256b. 

imbricata Brongn. 196; imbricata Regel 196; imperialis 34; in- 
camata 25; inermis 55; var. flava 55b; integrifolia 71; var. ma- 
jor 71; var. meridensis 148; intermedia K. Koch 234; intermedia 
Sims 242; iridiflora 292. 

jacksonii Hook. 42; jacksonii K. Koch 243; jaliscana 221; 
jimenezii 237; johannis 185; juncoides 29. 

kalbreyeri 145; kanrinskyana Beer 226; karwinskyana Schult. 
221; kegeliana 24; killipiana 36; klabochiana 98; klabochorum 
98; klotzschiana 73; kniphofioides 69; kunhardtiana 5- 

laevis Beer 256; laevis Willd. 123; lamarcheana 103; lanata 
293; lancifolia 253; var. minor 253b; lanuginosa 169; laresiana 
34; latibracteata 34; latifolia Ait. 240; var. cubensis 239; la- 
tifolia Andr. 238; latifolia Baker 60; latifolia Jacq. ex Beer 
240; latifolia Redoute 241; latifolia Soland. 240; latifolia 
Wendl. 226; laxissima 139; lechleri 54; lehmannii 58; leiolema 
259; lepidopetalon 175; lepidota 181; leprieuri 18; leprosa 209; 
I'henninieri 256; liebmannii 116; lignosa 120; limae 31; lindenii 
103; lindleyana 116; longebracteata 191; longicauda 258; longi- 
folia Hook. 49; longifolia Beer 116; longipes 140; lopezii 126; 
lorentziana 294; lutea 184; lutescens 134; lymanii 203. 

macarensis 155; macranthera 89; macrobotrys 171; macrocalyx 



196U Smith, Notes on Bromellaceae 53 

90; macrochlamys 19i*; macrcphylla 295; maguirei 22; maidifolia 
90; maizaifolia 90; maritiaa 92; maronii 296; matudae 192; maydi- 
folia 90; media 297; megasepala 66; megastachya 298; melanopoda 
125; membranifolia 199; meridensls hort. 299; meridensis KL. 148; 
micheliana 207; micrantha 300; mlcrocalyx 184; "»ar. elliptica 
184c; var. schlimii 184b; micropoda 220; railitaris 222; minarum 
25 6f; mirabilis 40; var. tucumana 40b; mituensis 4; modesta 210; 
monstrosa 301; montalbensis 226; monticola 225; mooreana 213; 
mordil 256; moreliana 256; morelii Baker 255; morelii Lem. 256; 
moritziana Kl. 73; moritiiana Koch & Bouche 73; morrenii 116; 
multiflora 144; multiramosa 81; muscosa Hook. 259; muscosa Uart. 
25 6f. 

nana 37; nigra 96; nobilis 159; nubigena 147; nuda 2. 

oaxacana 228; oblanceolata 107; oblongifolia 168; occidentalis 
178; ochroleuca (Koch & Bouche) Baker 201; ochroleuca sensu Baker 
195; odontopoda 82; odorata hojrt. 257; odorata Wawra 260; oer- 
stediana 90; olfersii 256; olivacea 302; oranensis 59; orchidi- 
folia 172; orgyalia Andre 57; orgyalis Baker 58; ovandensis 203. 

palmeri 215; var, longebracteata 2l5b; palmoides 165; panlcu- 
lata 49, 278; paraguayensis 170; pastoensis 303; patentlflora 12; 
var. armata 12d; var. macrantha 12c; var. subintegra 12b; pauci- 
flora 24; pavonii 122; pearcei 304; pectinata 39; penduliflora 
Mez 305; penduliflora A. Rich. 306; petiolata (Koch & Bouche) 
Baker 197; petiolata sensu Baker 195; petraea 180; philippii 307; 
platypetala 32; platyphylla 238; platystemon 47; plumieri 308; 
poeppigiana 88; polyanthoides 90; poortmanii 68; pruinosa H.B.K. 
11; pruinosa Uez 256f ; pterx>poda 219; puberula 216; pulchella 97; 
pulverulenta 48; pungens 123; punicea 42; purpusii 204; pusilla 
176; puyoides 46; pyrajnidata Link 309; pyramidata Pers. 310; 

queanelioidea 13; quetameensis 311. 

racemosa 241; ramosa Jacq. f . 234; ramosa Koch 234; recurvata 
I9O; redouteana Beer 233; redouteana Schult. 234; reflexiflora 
153; regie 312; rhodostachys 115; rigida 53; ringens Beer 242; 
ringens Kl. 226, 324; riparia 137; robusta 313; roezlii Andre 79; 
roezlii Baker 256; roezlii E. Uorr. 256b; roseana 206; rubicunda 
256; rubiginosa 15; var. amazonica 15b; var. Integra 15c; ruizi- 
ana 51; rusbyi 314. 

samuelssonii 232; sanctae-crucis 315; sandemanii 91; saxicola 
203; scandens 133; sceptriformis Mez 94; sceptriformis Ule 94; 
sceptrigera 112; schiedeana 218; schlimii 184b; schultzei 75; se- 
cunda 316; secundiflora 229; selloana 256f; selloviana 256f; se- 
maphore 154; semijuncta 25; sessiliflora 60; similis 164; skin- 
neri 234; sneidemii 161; sodiroi 158; sordida 224; spathacea 
317; spathulata 3I8; spec. fol. lineari-lanc. etc. 242; speciosa 
258; specioslssima 246; spectabilis I38: sphaerocephala 319; 
apicata 241, 312, var. sulphurea 241b; spinosa 320; splendens 
Poir. 321 ; splendens Wars. 203; sprucei 20; squarrosa 102; var, 
aurantiaca 102c; var. colorata 102b; staminea 258; var. longi- 
cauda 258; stenophylla 121; stenothyrsa 322; straminea 131; 
stricta 323; suaveolens 260; subjuncta 24; subpetiolata 60; sub- 
ulifera 114; sulfurea Uez 193; sulphurea Andr. 241b; sulphurea 
Koch 231 ; sylvestris 151. 



Sk PHYTOLOOIA Vol. 10, no. 1 

tabuliformis 189; taenipetala 190; tarapotensis 63; tenuis 71 j 
theae 202; tillandsioidea 217; tolimensis 70; tomentosa Dietr. 
234; tomentosa Mez 235; torresiana 250; trianae 167; var. retusa 
167b; trimorpha 67;truncata 61; tuberculata 45; tuerckheimii 211; 
var. raacrolepis 211b; timiulicola 142; turbinella 21; tympani 177. 

uaupensls 26; ulei 248; umbratllls 110; undulata 246; \mdula- 
tifolia Hook. 115; undulatlfolia hort. 246; unilateralis 174. 

v£d.erii 188; vallisoletana 324; vargasiana 76; venusta 325; 
verrucosa 50; violacea 326; violascens 128; vlrescens 327; viri- 
diflora 328; viridls 35; volubilis I46. 

warszewitziana 226; weberbaueri 34; weddelliana Baker 329; 
weddelliana Mez 25 6f; wendlandil 193; werckleana 95; wurdackii 3, 

xemthocalyx Baker 257; xanthocalyx Mart. 231; xanthocalyx Mez 
232. 

zeifolla 90. 

POURRETIA ferruginea 34; inermis 55; paniculata 49. 

PUYA alpestris 262, 276; altensteinii 115; var. gigantea ll5b; 
brachystachya 265; caerulea 269; camea 230; chilensis 270, 274; 
coerulea 275; dyckioides 282; echinotricha 34; floccosa 286, 290, 
299, 311; funkiana 90; furfuracea 288, 309; glymiana 243; grandi- 
flora 34; heterophylla 116; lanata 293; lanuginosa 277; lehmann- 
iana 303; longifolia II6; macrostachya 115b, macrostachys 115b; 
maidifolia 90; nana 37; olivacea 302; pearcei 304; pitcaimioides 
289; pyranddata 310; recurvata 190; roezlii 298; ruiziana 169; 
rusbyi 314; sanctae-crucls 3I3, 315; spathacea 287, 317; steno- 
thyrsa 263, 322; sulphurea 193; venusta 319, 325; violacea 307, 
326; warszcewiczii 103; weddelliana 329; whytei 276. 

QUESNELIA bakeri 106. 

TILLANDSIA laevis 256; schuechii 257; secunda 317; vincenti- 
ensis 241b. 

VRIESEA tricolor 90, viridiflora 328. 

WILLRUSSELLIA feliciana 261. 



196U 



Smith, Notes on Bromeliaceae 
Plate I 



55 




^1729 



Fig. 1: Pitcairnia caricifolia ( Holt & Blake 425); fig. 2: P. 

angustifolia ( Cook & Collins 267); fig. 3: P. panicvilata 

(Killip & Smith 25074T; fig. 4-5: P. vargasiana; fig. 6-?: P. 

halophila; fig. 8-9: P. decurvata; fig. 10-11: P. fractifolia; 

fig. 12-13: P. grubbiana; fig. U-15: P. micropoda; fig. 16-1?! 

P. foliacea; fig. 18-19: P. domingensis. 



U&TERIALS TCfflAfiD A UCfKOGRAPH OF THE GBSfUS VERBENA. XVII 
Harold N. Uoldenke 



VERBENA LITORALIS H.B.K. 

Additional bibliography: F* A. Barkley, Detexmin. EJemp. Herb* 
Fac. Nac. Agron. Medellln 1 (1): 2 & U (19U9} and 5: U8 & U9. 
19U9; Boelcke & Echeverria, lilloa 18: 250, cuadro 17 & 26. 19U9| 
Uatuda Am. Ifldl. Nat. hht 576 « 1950; Moldeoke, Hijrbologia 3: 
28U, 2S6, 289, & 290 (1950) and 3: U67. 1951; Stellfeld, Trlb. 
Famac. 19 (10): 166 & 172. 1951} Moldenke in Chittenden, Roy. 
Hort. Soc. Diet. Qard. kt 2209 & 2211. 1951} Aeevedo de Vargas, 
Bol. Mus. Nac. Hist. Nat, ChUe 25: 57—58. 1951 J Moldenke, Phy- 
tologia Ut 66 & 7U (1952) and U» 267 & U5l. I953j C Skottoberg, 
Veg. Juan Fern. 922 k 9li2. 1953} Rambo, Sellowia 6: 60, 81*, & 
153. 195U; Moldenke, Phytologia $t 2h, 72, & 96. 195U} Thome, 
Av. Midi. Nat. 52: 313. 195U} Moldenke, Inform. Mold. Set U8 
Spec. [U] (195U) and U9 Spec. 3. 195U| Moldenke, Mem. N. I. Bot. 
Gard, 9: 177. 1955} Soukup, Biota li l8l. 1956; Ranibo, Selloiria 
7: 260. 1956} Moldenke, Infora. Mold. Set 51 Soec. U. 1956; Mol- 
denke in Humbert, Fl. Madag. 17U: 9. 1956i Angely, Fl. Paran. 7: 
13. 1957} Moldenke, Am. Midi. Nat. 59: 33U, 3U2, & 3U6. 1958} 
Reltz, Sellowia 11: U5 & 13U. 1959} Siranholm, St. John, &. 
Scheuer, Pac. Sci. 13: 303. 1959} Moldenke, RSsum6 11, 22, 23, 
29, 32, 33, 39, U3~U5, U7, U8, 69, 7U, 80, 81, 85, 110, 115, 
118, 119, 122, 127, 15U, 203, 210, 212, 223, 311i, 357, 359, 365, 
368, 369, 371, 1*25, & U72. 1959} Moldenke, R^sumS Suppl. 1: 2, 
6, & 8 (1959) and 2: U, 7, 10, & 12. I96O; 0. Degener, Fl. 
Hawaii. 315: Verbena: Lit oralis. I96O; Renn6, Levant. Herb. 
Inst. Agron. Minas l5l. I960} Angely, Fl. Paran. 16: 78 (I960) 
and 17: 1*6. I96I; Reltz. Sellowia 13: (13): 110. I96I} Lewis tt 
Oliv., Am. Joum. Bot. U8: 6Ul & 6U2. I96I; Moldenke, Phytologia 
8: 120, 12U. 127, & 11*8 (I96I) and 8: 200—202, 21*6, 21*7, 253, 
255—257, 280, 310, 313—318, 1*05, & 1*12. 1962} Moldenke, R6sum6 
suppl. 3: 10, 11, 13, 15, 25, 27, 36, 39, & 1*0 (1962), l*t 5 4 17 
(I962), 5: 5 (I962), and 6: U, 6, 7, 4 11. 1963} Moldenke, Phy- 
tologla 8: 1*60, 1*63, 1*87, 1*90, & 1*91 (1963) and 9» 66, 67, 113, 
121*, 126, 151, 151*, 155, 165, 206, 213, 220, 296, 38O, & 382. 
1963. 

Illustrations: H.B.K., Nov. Gen. & Sp. Pi. 2t pi. 137. 1818} 
Sanzin, Anal. Soc. Clent. Argent. Buenos Aires 88: 111*. 1919} I. 
C. Verdoom, Union of S. Afr. Dept. Agr. St Forest. Bull. 185 1 
fig. 90 [as V, officinalis ] . 1938} Augusto, Fl. Rio Grande do 
Sul fig. 102. 191*6} Troncoso & Burkart, Darwlniana 7: 211, fig. 
2b. I9U6} 0. Degener, Fl. Hawaii. 315: Verbena: Litoralis. I96O. 

Quick-growing erect or suberect perennial herb, sometimes a 
suffrutescent shrub or undershrub, 0.2—2 m. tall, stzdct, shiny, 
sometimes large and robust, woody-based, fastigiately few- to 
many-branched above, glabrous or sparingly strigillose: stems to 

56 



196U Moldenke, Uonograph of Verbena 57 

12 am* thick, tetragonal or subquadrangular in cross-section, 
with the whole intemode deeply sulcate above the insertion of 
the leaves and faintly many-solcate between, antrorsely scabridu- 
lous (on the angles) when young, gz^en, with irtiite pith, slightly 
contracted at the nodes; leaves decussate-opposite, lanceolate to 
oblong or oblanceolate, >-- 10 cm. long, 1— 1*5 cm. wide, acute at 
the apex, tapering into a very short petiole or else sessile or 
subsessile, acuminate and somewhat clasping or subanplexicaul at 
the very base, antrorsely scabridulous throughout or scabrous and 
somewhat rugose only above, the upp>er 2/3 moi^ or less sharply 
and coarsely serrate with commonly >-- 5 somewhat mucrcmate teeth 
per side, sparsely strigillose on both surfaces, the venation im- 
pressed above and prominent beneath; spikes terminal, several to 
many, pedunculate, cymose or corymbose-paniculate, slender or 
filiform, pilosxilous, at first congested and about U mm. in di- 
ameter, dense or interrupted, later loosely flowered and elongate, 
in fruit becoming 2 — 8 cm. l(mg, the fruits usually not contigu- 
ous; bractlets ovate-lanceolate, marcescent, rigid, acuminate or 
subulate at the apex, 1 — 3 mm. long, usually subequaling or some- 
irtiat shorter than the calyx, abruptly upcurved, glabrate; calyx 
often purple, 2 — 2.5 mm. long, pubescent or finely strigillose 
outside, the rim subtruncate, the 5 teeth unequal, minute, and 
subulate; corolla about 3 mni. long, varying from dark-blue, blue, 
bluish, light-blue, pale-blue, lilac-blue, lavender-blue, bright 
lavender-blue, violet-blue, or pinkish-blue to bluish-lavender, 
blue-violet, blue-purple, blue- lilac, lilac, clear- lilac, violet- 
lilac, dull-lilac, violet, pale-violet, pale rose-violet, lilac- 
violet, lavender, light- or pale-lavender, pink-lavender, 
purplish, purple, bright-purple, deep-purple, mauve, pink, or 
red, often described as "lilac with a violet tube**, "lavender 
with a red tube", "lavender with white center", or "throat pale- 
lavender or white, lobes lavender", puberulent on the outer sur- 
face, its tube greenish toward the base, variable in length, al- 
ways someifhat longer than the calyx, the limb inconspicuous, 
usually pale-lavender, 2.5—3 una* wide, puberulent on the inner 
surface, the lobes subequal, obtuse at the apex, glabrous; sta- 
mens U; anthers greenish-yellow; style about 1 mm. long, green, 
glabrous; stigma 2-lobed, the larger lobe stignatiferous; ovary 
almost 1 mm. long, green, glabrous; cocci trigonous, dark- 
stramineous, linear-oblong, 1 mm. to hardly 2 mm. long, glabrous 
and smoothish, striate, somewhat reticvilate at the ^ex, the 
commissural face about as long as the coccus, muricate-scabrous; 
chromosome number: x - 7, 2n • 28. 

An extremely widely distributed and polymorphic subtropical 
and tropical weed ranging from Arkansas, Louisiana, and Texas 
throughout Central and South America; introduced in Oregon, Cali- 
fornia, South Africa, Hawaii, Australia, and parts of Oceanica. 
It has been collected in open and waste places, grassy places and 
fields, along roadsides amd fencerows, in grassy swales and wet 
meadows, thickets, wet thickets, and pastures, savannas and 
llanos, in cultivated and wet open ground, along moist roadsides, 
on roadside banks, at the edge of coffee plantations, in clearings 



58 PHTTOLOGIA Vol. 10, no. 1 

of temperate forests and "mata", on dunes and sand bars, in gras^ 
pasturelands and ditches, in volcano craters, on moist rlverbanks 
and stream edges. In wet campos and lanns. In maz*shes and bazren 
ground along streams. In rocky calcareous fields at the margins 
of woods. In dry sandy soil on sunny roadsides. In open ground on 
lake margins, among bushes, on hillsides. In cafetsds, potreros, 
and vacant lots, on ••reddish claylsh-loam In luxuriant tropical 
forests" and "on rich claylsh-loam along lake shores", at the 
edges of cemeteries, along city streets. In dry barren scrubland 
and sunny sand, at the edge of thickets, on rocky limestone 
elopes. In Panlcum msudmum fields, on creek banks and bottoms, a- 
long grassy roadsides. In low and sandy areas, low marshy areas, 
on grassy hillsides. In springy areas and water- filled depressions, 
at the borders of cxiltivated fields, in valley land, on low 
ridges, and among low grasses in open spots, from sea-level to 
3550 meters altitude, flowering and fruiting In every month of the 
year. 

Reiche (1910) says "Planta americana, desde Mfijlco a las reji- 
ones templadas de Sud-Am4rica; en Chile tanto en la zona literal 
como en el interior; tambien en J. F. (Masatlerra) . Florece casi 
todo el afio. 

The species was found in Amador County, California, in 1896, 
and in San Joaquin Counly in 1902. King describes it as an uncom- 
mon roadside weed, growing in open stm, in San Luis Potosl, Mexi- 
co. 

The type of the species was originally described by Boi^land 
as "crescit in salsls maritimis Oceanls Paciflcl props TruxLllo, 
Santa et Lima", Peruj that of V. affinis was collected by Henri 
Ouillaume Qaleotti (no. 781) at an altitude of 6000 feet at Mor- 
elia, yichoacin, Mexico; that of V. arborea is Herb. Hort. Bot . 
Bogor. XV.K.A.XLV.17 , cultivated at Bultenzorg, Java; that of V. 
litoralls var. pycnostactys f . montana was collected by Cornelius 
Osten (no. 10615) at Villa Nougues, at an altitude of 1100 meters, 
Tucum&i, Argentina, on May 17, 1917, and is deposited in the 
herbarium of the Museo de Historia Natural at Montevideo; and 
that of V. lanceolata is Herb. Willdenow 1113U , deposited in the 
Willdenow Herbarium at the Botanisches Mviseum in Berlin. 

As to V. nudiflora , Degener (I960) says "Thomas Nuttall col- 
lected this plant on «Wahoo« [-Oahu] in 1835. Thinking it new, he 
proposed to name it Verbena nudi flora , writing this name upon 
his herbarium sheet, Nuttall died in 1859 and Nlcolaus Turczanl- 
now, a year before his own death, took up Nuttall* s binomial 
from the herbarium label and published it for him. Thanks to the 
kindness of Bayard Long of the Acadeaiy of Natural Sciences in 
Philadelphia irtio compared Nuttall 's historic specimen deposited 
at the Academy with a specimen of Verbena litoralls mailed him 
from Waialua, Oahu, in I960, we know we must relegate Nuttall 's 
name to synonymy." 

The V. littoralis var. pycnostachya Schau. sanetimes included 
in the synonymy of V. litoralls is regarded by me as V. brasil- 



196U Uoldenkey Monograph of Verbena 59 

lenais Veil., while V. Caracas ana H.B.K. and V. lltoralis var. 
caracasana Briq., also sometimes placed here, aire discussed b7 me 
\inder V. literal is var, caracasana (H.B.K.) Briq., i^ich see. V. 
nudiflora L. belongs in the synoEormy of Phyla nodi flora (L.) 
Greene, ^. parvi flora Rula is V. gracilis Desf., V. arborea H.B.K. 
is Petrea arborea H.B.K., and Lippia litoralis R. A. Phil, is 
Phyla nodi flora var, rosea (D. Don) Moldenke. 

Bentham's original description (18U6) of his var.? glabrior is 
as follows: " Verbena litto rails var? glabrior , folia hinc inde 
trifidis grosse et obtusiuscule inciso-dentatis . An species pro- 
pria? Folia V. menthaefoliae, Benth. PI. Hartw, p. 21, sed flores 
parvi V. littoralis . — Peita." 

One known hybrid of V. litoralis is with V. hispida Ruiz & 
Pav. and is discussed by me under xV. bealei Moldenke. Undoxibt- 
edly many other hybrids occur in the wild, accounting for sane of 
the anomalous intermediate specimens bandied about between this 
and related species by herbarivn workers, but more experimental 
work is needed before these can be clearly isolated. 

In his discussion of V. bonariensis L., Hooker [Bot. Misc. li 
166. 1829] says "V. littoralis Humb. seems to be a variety of 
this with shoirter spikes than usual.** On the basis of this 
statement. Gay and other authors speak of a'V. b onariensis var. 
littoralis Hook." 

Pellett (1923) reports that honey is yielded by V. litoralis 
"over an area of considerable extent frcm Baton Rouge to near New 
Orleans in Louisiana." 

Degener 17852 has leaves which greatly resemble those of V. 
brasiliensis ; Cuatrecaaas 1865U has unusually large dark-green 
leaves. Peredo s.n. [29-I-I9I46, Cabezas, Cordillera] bears this 
description on its label: "20—1^0 cm. tallj petals pale-yellow 
and stamens purple", but this is probably erroneous. 

Troncoso & Burkart (19U6) say: "La difundida V. littoralis se 

diferencia por las espigas delgadas, de y—k mm. de difim 

flores de li — $ mm. de long., oAliz de 2,3 — 2,8 ma. de long., 
cubierto de pelitos breves uniformemente repartidos sobre su 
superficie, brActeas tambi^n menores [than in V. tristachya 
Troncoso & Burkart], do 2 — 3 mm, de long, y pubescentes . . . .Qa V. 
littoralis las ramas superioi>es y los t alios J6venes son macizos, 
observandose que los tallos m&a gruesos y los entrenudos inferi- 
ores son algo fistulosos, pero con una gruesa capa de m^dula.... 
El estudio anat&aico del tallo de V. tristaujhya presents ccmo 
caracteristica interesante, que la capa de parSnquima clorofilo- 
ano es continua, es decir, que no se halla interrumpida en los 
ingulos por la columna esclerenquim&tica. Egta tiltima estructu- 
ra caracteriza a V. littoralis ." 

Herbarium material of V. litoralis has been misidentified a- 
bundantly and distributed under such names as V. angustifolia 



60 PHTTOLOOIA Vol. 10, no. 1 

Michx., V. bonarlensls L., V. braslllensls Veil., V. caracasana 
HtuDib. & Bonpl., V. caracasana H.B.K., "V. caracasana var.", »V. 
cf . caracasana HBK.", V. caracassana H.B.K., V. gracilescens 
(Cham.) Herter, V^ halel Small, V. hastata L., V, hlsplda Ruiz & 
Par., V. Isabellel Briq., »V. lltoralls Kth. vel aff .", V. liter - 
alls f . pycnostachya Schaa., V. lltoralls var. brasillenais Briq., 
V. lltoralls var. pycnostachya Schau., V. officinalis L., V. of- 
ficinalis var, gracilescens Cham,, V. panicvilata Lam,, V. poly- 
stachya H.B.K., V. stricta Vent., V. urticaefolla L,, V. urticl- 
folia L., V. aoitba Lehn., Llppia sp., Priva leptostachya A. L. 
Juss., Stactprtarpheta dichotoma Pers., and even Cyathula achyraa - 
thoides (H.B.K.) Moq., Buddleia asiatica Lour., Bjyptis pectinata 
Poit., and Wan Spartiian? " 

Perry annotated Maltby 2|2 and P. Bjj^ Kennedy 70U6 [whose label 
originally read "60116"] as "Aff. V. menthaefolla Bth." Herter s. 
n. [Herb. Osten 18U97] was aimotated by Osten as " Verbena littor- 
alis Kth. ad V. bonariensem L. speotans", while lindman A,36U7 
was annotated by Briquet as "Verbena lltoralls Kunth v. brasill- 
ensis Briq. f. aliq. ad var. caracasanam vergens." 

On the other hand, the Henschen 1.326 and Regnell 1.326 , dis- 
tributed as V. lltoralls , are actually V, alata Sweetj Bruch s. 
n. [Las Juntas, "XIIA896] , Ounckel 1686^ Harshberger s.n. tPan- 
broke's Swamp, June 21, 1905], Holway & Holway II8O & litgU , and 
Venturi 2812 are V. bonarlensls L,j M. lBang 136 , Claude- Joseph 
2616 , Cook ic Gilbert 271 & 1208 , Dus6n 8, Holway & Holway 1272 , 
Killip & Smith I73I4O , Morong 128 , Ro8e ," 'Pachano , & Rose 228U5 & 
228U6, and Sehnem Ui70 are V. brasiliensis Veil.; Heyde & Lux 
3019 , Liebmann 11339, and Renson 175 are V« Carolina L,j A. 
StewarF33l9 is V^ galapagosensis Moldenkej Anthony & Tate 333 , 
Killip & Smith 21197 , Pittier lli28 , A. Stewart 3317 , and LI, 
Williami $9U are V. glabrata H.B.K. ; M^ Bang 10^8 is V. gracil- 
escens (Cham.) Herter; JPrgensen 1021 is V. hlsplda Ruiz & Pav.j 
Liebmann 11318 , £. R. Orcutt 1371 , Purpus 3UO6, and Seler & S©- 
ler 72U are V. longlfolla Mart. & Gal.j Bourgeau 36O is V. menth - 
aefolla Benth,, as is also Arsene s.n. [Rinc6n, 2^/7 A909]; Du- 
sfa 85146 is V. minutiflora Briq.; R. Alvarez U27, Arechavaletae 
39 & 3139, Beetle 2025 . Cabrera 86I , 1710 , 2108 , 2159, & 238^ 
Collector undesignated s.n. [Dec . 1885] , JU M. Curran sja. [Oct. 
19, 1913], Du86n 7827, Grttner 1|18 , Herter 269 [Herb. Herter 
8I7I3] , Kuntze s.n. [Montevideo, 7/Xn/91] , Legrand 25U & 2009 , 
Nlcora 377, Oaten 3165 &. 3335 , Pas tore 137 , Pedersen 77U , Rambo 
U5339 , li6069 , U9723 , & 55075, Reiss 55 & 56, T. Rojas Ui8 , Rosa- 
Mato 313 , 399, & UOO, Scala 10001 , Sehnem 3519 , Seijo s.n.[l6 
Not. de I88U], Smith & Klein 11797 . Smith & Reitz 8988 & 9730 , 



196U Uoldeoke, Uonograph of Verbena 61 

Smith , Reitz, & Sxifridinl 96 2U , and Telsselre UU8I are all V. 
montevidensls Spreng.; A. H^ Moore 2939a, Herfa. Harvey s.n, [ex 
seminib. h» R. P. l8Ui], Herb. Hort. Matrlt. 3L, and Yvmcker 
1011^5 are V. officinalis L.; C. R, Orcutt 118 , 521 , & s.n. [Pin- 
ery, 7-27-1883] are V. orcuttiana Perry; Flebrig 56U$ is V. ovata 
Cham. J Kuntze sja, [Bolivian, 600 m., 1/U April 1892] is V. par- 
vula Hayek; Venturi 5 is V^ rigida Spreng.; JOrgensen 3773 is V, 
stellarioioes Cham.; H. L_. Mason 1612 and Patiflo 7138 are V . 
sphaerocarpa Perry; A. Stewart 3320 is the type collection of Y« 
Stewart 11 Moldenke; Cory US 8 78 la V. xutha Lehm.; and Herlberto 
388 is not vorbenaceous. The A. A_^ Heller 15162 , distributed as 
V. hanseni , is actually V. bonarienais L. 

H» E* Seaton 27 and Sharp hhl\ 20 are insect-galled; Liebmann 
11336 is also abnormal. Its spike is thickened like that of 
Stachytarpheta cayennenais (L. C. Rich.) Vahl and there are no 
developed flowers, only large bractlets — it may possibly repre- 
sent an intergeneric hybrid between the two taxa. Lutz 1185 is a 
mixture with V. bras ill ensis VeU,, Schultes & Reko 237 is a mix- 
ture with V. Cfirollna L., and Hanfaury-Tracy 256 is a mixture with 
something non-verbenaceous . The "V. officinalis " described and 
Illustrated by I. C Verdoorn, Union of S. Afr. Dept. Agr, & For- 
est. Bull. 185: 171, fig. 90 (1938) is actually V. litoralis . 

The H'"'™^1 s.n. cited below from Caracas, Venezuela, is an 
especially dense-flowered plant and may actually represent var. 
Caracas ana ; the same is tnie of Demaree 2l|6U9 from Arkansas . In 
fact, many specimens cited below by me as V. litoralis seem anom- 
alous in that their spikes are much more dense than in irtiat I re- 
gard as typical V, litoralis as exemplified by A_. A_^ Heller 20U6, 
P. RvisseH 52, C^ N. Forbes 188.2 , and LI. Williams 1^2 ", Meagher 
s.n. [July 7, 1937] is in part typical and in part dense. The 
dense foim is well represented by J, M. Wood 13132 , A. Forbes 
108 3n & s.n. [Tantalus Rd., 7AV30], F. E^ Egler 37-83 , D. W. 
Garfaer 339, F. R. Fosberg 8856 & 10276 , and K. Hartley s.n. [Wai- 
anae, M^ U, 1856]. Quayle 258 is rather dense but weak, while C. 
N. Forbes 2036m is very dense. C. S. Judd 11 and C.N. Forbes 
Uo5h look very much like V. braslliensls Veil. Personal observa- 
tion of the plant growing so abundantly in the Hawaiian Islands 
causes me to have misgivings about its correct determination. It 
does not appear to be the same as the coramon weed wl-dch I observed 
in so many places in Soxith America. I would not be at all sur- 
prised if further stu^y should reveal that this dense-spiked form 
belongs in var. caracasanal The JOrgensen 3767 [Herb. Osten 
222I18] (Ug), cited by me on page 322 of Phytologia, vol. 8, as V. 
brasiliensis , seesns, rather, to be V, litoralis. It is very pos- 
sible that much of the material cited as V. litoralis from south- 



62 PHTTOLOGIA Vol. 10, no. 1 

em Brazil and noz*them Argentina l8 V. montevldenals . 

August o (I9I46) says: 'H^omun no sul do Brasll (Sello). U\ilto 
comun nos campos e morros dos arredores do p6rto Alegre. (Emrlch, 
InnSo Augusto e InnSo EdSslo.)" Rosengurtt (19U6) says "Mala hi- 
erba perenne (a suf rutlctilosa? ) , de ciolo estlval. Abunda en can- 
pos vli^enes y de rastrojo". He cites his FE.$06 , $63 , 590, 696 , 
739 , 859 , and 1213. 

Halller D.589 was flr^t Identified as V. urticlfolla L., then 
as V. officinalis L., then as V. angustlfolia Michx., and finally 
as V. litoralis by various herbarium irorkersl The species is said 
to have been intrwiuced into cultivation in 1875 • 

On Indefatigable Island it is described by R. G. Taylor as oc- 
curring "in a partially deserted- clearing in the forest in moist 
zone of island" and being an "herb, 2 — 3 ft. high, fls. lilac, 
possibly an imported plant, not seen in untouched forest." 
Schulz & Schulz found it "eingeschleppt" in Berlin, Germany, In 
1898. On Easter Island it i»as found by Chapin on the inside of a 
volcano crater, as also on Maui in the Hairailan Islands. Fosberg 
found it to be a weed in yards in Cundinamarca, Colombia, while 
Mendes Uagalhaes describes it as "iruderal" in Minas Gerals, Bra- 
zil. Morel says that it is a "canmon plant on campo" at Rlacho 
Porteflo, Formosa, Argentina, irtiile jorgensen describes it as a- 
b\mdant at Andalgal^. Goodspeed asserts that it is "common and 

perhaps Introduced in rather moist soil in irrigated fields" 

at Valparaiso, Chile. Stellfeld and Roth both describe it as 
"ruderal", Smyth says that it is the "common tall verbena of the 
valley, common in iraste places", Tamayo avers that in his coun- 
try the plant is "very common in abandoned cultivated fields", 
while Yuncker, Dawson, & Youse refer to it as "a ireedy plant". 
Lutz encountered it "abundant by roadside", Stelnbach in "tienra 
greda", and Sandeman as "frequent in full exposure". Herter de- 
scribes it as "along Roadsides in dry sand^ soil e:qx>sed to fuU 
s\in" and "conmon in dry sandy soil along roadsides" at Montevid- 
eo. JOrgensen reports it "common on campo" in Paraguay, while 
Rimbach calls it a "weed in gardens and on walls" in Ecuador. 

On Masatieirra the Skottsbergs found V. litoralis in colonies 
along roads and frequent on the shore; Mandon claims that it is 
"everywhere in cultivated land, hedges, and thickets near Sora- 
ta" in La Paz, Bolivia; Macbi*ide & Featherstone descidbe it as a 
"common pasture and wayside weed" in Junln, Pezni; and Liiidman 
found it "abundant in pastures" in Rio Grande do Sul, Brazil. 
Sampalo comments "nos terranos incultos, & beira de lugares hd- 
midos". Hanbury-Tracy discovered it growing in grass meadows 
in foothills and fairly dry ground among spaced scrub with few 
trees. Steward affirms that in the Galapagos Islands it is abirn- 
dant in open country around 9000 feet altitude, common in wet 
soil near springs at 1000 feet, and rare at 1550 feet. DeWolf 
found it to be "fairly common" along trails in Costa Rica. Ewan 
describes it as a "rather common perennial" with the "appearance 
of Statice in the field". Stanford, Retherford, & Northcraft 



196U Moldenke, Monograph of Verbena 63 

found it "in broad damp river-bed with varied vegetation of large 
shrubs, small trees, and herbs'* and **on mountains with luxuriant 
vegetation". Standlej (1938) says of it "frequent in waste 
places, often in cultivated fields, tierra caliente, ascending to 
the Ueseta Centiral, mostly as 1,300 metez*8 or less, but sometimes 
at someirtiat higher elevations. Widely distributed in tropical 
America" and "weed in waste ground, dry brusly hillsides, and dry 
thickets". Northcraft says that they are "aquatic plants growing 
in canal next to road". In Honduras, according to Molina R., the 
species is a "weed in pine forests". Cook & Gilbert 2088 was 
"purchased in market" in Peru. 

Degener (I96O) records our plant from Maui and Hawaii and "on 
all the larger islands" of the Hawaiian group "from sea level to 
almost the tops of our highest mountains, preferably in pastures, 
pineapple fields, waste places and lava flows; never in dark 
forests . On Maxii and Hawaii the Hawaiians have used it as a med- 
icine for skin diseases." He also says "Mann in 186? correctly 
judged this weed introduced, while Hillebrand in 1888 described 
it as 'a troublesome weed of early introduction which has taken 
root in many parts of the Islands, most so on the highlands of 
Waimea, Hairaii, where large extents of pasture land have been 
ruined by it." Miss Neal (19U9) refers to it as a "weed at 3000 
feet" . A note on Herb. Oahu Coll. 56, however, says "once a 
troublesome weed, now nearly run out". In New South Wales, ac- 
cording to Kaspiew, it constitutes a "pest in pastures" . 

Common and vernacular names recorded for this plant in vari- 
ous paz>t8 of its range are "berbena", "erra de S8o Caetano", 
"einra do pai caetano", "escoba dura", "false vervain", "hauoi", 
"ha'uoi", "herba do Pae-Caetano", "oi", "'oi", "quininha", 
"seashore vei*vain", "titania", "verbena", "verbena amarge", 
"verbena del campo", "verbena del literal", "verbena bljuica", 
"verbena erguida", "verbena blanca serrana", "verbena cimarronaj 
"verbena negra", "verbena nigra", "verbena on parade" [meaning, 
the upright verbena], "veirbena parada", "vervena", "weed verbe- 
na", "yerba de acero", "yerba del acero", "yerba de padre 
Caetano", and "yerb6n". 

Philippi (1870) comments concerning var, leptostachya "La 
trasa es diferente de la forma comun por ser los f rutitos mucho 
mas pequefSos; pero los ejemplares, demasiado adelantados en la 
vejetacion, no dejan ver otras difei*encias". Rosengurtt (19li3) 
says "Hierba hemicript6fita a sufruticosa; florece de fines de 
prlmavera hasta principios de otofio. Habita en lugares muy vai^- 
iables de los campos, bosques, poblaciones, chaci>as, etc. Es 
comiSn, y el ganado la msmtiene i*ecortada normalmente" . He cites 
his PE.I269 , 1335, lUi7, 1576 , 1783 , I9O8 , 2059 , A .1002 , and OH. 
I8U89 fran Drugtiay. Cabrera & Dawson (19Ui) describe the plant 
as "Hemicripft6fita parecida a las anteriores, pert) de s6lo UO a 
90 cm de altura y con espigas mis delgadas . Vegeta en casi toda 
America del Sur" and cite Dawson 9ti6 and Cabrera 1710 . Calder6n 
& Standley (19lil) report it as cc3mmon in El Salvador. 

Stellfeld (1951) states "Chamada na cidad e do Rio Negro (Par- 



6U PHTTOLOQIl Vol. 10, no. 1 

ani) de 'qainlnha' e 'erva de SSo Caetano' em outros lugares e en 
SSo Paulo, reputada ccmo febrlfuga, da£ o seu nome. E* bastante 
amarga'*. Reltz also repoirbs the species as used medicinally in 
Brazil; the Hinckleys saj that it is used as a purgative in Are~ 
quipa(, irhile Archer reports it to be a "general remedy, also used 
for cough" in Paraguay. In Gtuttemala, according to Ruano, it is 
employed as a remedy for fevers, grippe, and smallpox. Qarcia &. 
Barriga report it as medicinal in Colombiaj Duque avers that it 
is cultivated there for its "bitter and medicinal properties" and 
Niemeyer also reports that it is used against fevers. Standl«y 
(I938) makes this coounent: "The plant is employed in domestic 
medicine as a remedy for fevers" in Costa Rica. In Colombia it 
is said by Fosberg to be employed as an smtimalarial and as a 
remedy for typhoid fever, irhile on the Hawaiian islands of Hawaii 
and Haul he reports the juice commonly used for skin ailments. 
Fernandez refers to it as a "shrub" and maintains that it reduces 
fever very quickly — the leaves macerated and boiled are taken 
internally as a vermifuge agsdnst hookwozu. Assis also refers 
to the plant as a "shrub". Hegi (1927) says "gegen Scrofulose 
venrendet". Hinton states that the "very bitter Juice from mac- 
erated plant [is] taken for malaria" in Mexico. Steinbach re- 
ports that in Bolivia the plant is 'taedicinal, used against con^ 
tusions" . 

Martens & Galeotti (18UU) describe their V. af finis as dif- 
fering from V. bonariensis L. in "foliis brevioribus, spicis 
filiformibus elongatis laxis vix pilosis breve peduncvilatis 
cauleque non hispido", irtiich is a fairly accurate statement of 
the differences between V. litoralis and V. bonariensis . 
Schauer (18U7) separates his two varieties as follows: " cc. pyc- 
nostachya - spicis virgineis densioribus subcylindricisj a lep - 
tostachya - spicis Jam virgineis magis filiformibus, delnceps 
eaepius valde elongadis". The former is apparently merely a 
new status and name for V. brasiliensis Veil, and must be re- 
duced to the synorQnny of that taxon. To the second of his var- 
ieties Schauer reduces V. caracasana Kunth, V. lanceolata Willd., 
and V. af finis Mart. & Gal. (which he says is a "fomia umbrosa, 
plerumque magis glabz^ta, spicis pube appressa subcanescentibus"). 
He cites a Philippi collection from Chile; Schiede 135 & 1168 
from Jalapa and Papantlam, Mexico, and C. Ehrenberg and Galeotti 
collections from "Tlocalulo alibique", Mexico; Humboldt !c Bon- 
pland s .n . , Vargas and Moritz collections from VenezuelaJ 
Leschen s.n. from Guiana; and Sellow s.n. from Brazil, all de- 
posited in the Berlin, Martius (Munich), and DeCandolle (Geneva) 
herbaria. In his 1851 publication he modifies the description 
of his variety leptostachya to "spicis Jam virgineis magis fili- 
formibus, dein saepius valde elongatis et relaxatis". He de- 
scribed V. bonariensis f litoralis as "forma campestris aprica, 
plerumque undique magis hispidula". He adds V* glabrata H.E.K. 
to the synoiiymy of has var. pycnostachya with the comment "Hujus 



196U Moldenke, Monograph of Verbena 65 

nil niai modificatio monticola, magis compacta, glabrtor est". He 
cites Harrison, Gardner, Lhotsky, Sellow, Ackermann, Quill emin, 
and Lund collections, as well as Martiua 1033 and Vauthier 192, 
all from Rio de Janeiro, and Riedel a .n. , Raben B,n. , and Regnell 
326 fran Minas Gerais. He says: "forma a[V, brasiliensis ] etiam 
provenit in Bonaria, Chile, Peruvia, Colombia, Venezuela" and 
"forma f> [V. litoralis ] in Mexico, Venezuela, Guiana," 

Briquet (190U) claims that "V. litoralis Kunth aensu 

stricto" is the same as V. brasiliensis Veil, and V. litoralis 
var, brasiliensis Briq. Of his own V. approximata (which I re- 
gard as conapecific with V. brasiliensis ) he says: "Gette espece 
est tres voislne du V. litoralis et n'est pas facile a en distin- 
guer quand les feuilles infSrieures manqaent. Le V. litoralis 
possede des feuilles infSrieures moins oblongues, grossierement 
et irrfigulierement inois6e-dent6es, tandis qu»elles aont r6gu- 
lierement et finement crSnel^es-dent^es dans le V. approximata . 
&i outre, oe dernier a des tiges a angles plus aigus, presque 
STibailles grdce au sillonnement des faces, et des fexillles In- 
fSrieurea plua allong^es dtroltement lineaires, plus raldes. 
L' inflorescence et les caracteres floraux sont tout a fait ceux 
du V. litoralis var. brasilienais " . In his 1907 publication he 
compares it with V. carollata Briq. as follows: "Nous avions 

rapporte cette espece en 1899 au V. littoralis Kvmth, con- 

cu & cette epoque dans un sens tres vaste. Une 6tude plua at- 
tentive de ce groupe, fait depuis lors, nous a amene a detacher 
du Vj. littoralis plusleurs types parfaitement distincts (V. 
leabellel , V. cordobensis ) . Le V» carollata s ' ecarte a son 
tour du V. littoralis princlpalement psu: sa corolle a tube si- 
phone longuement exsert (depaasant a peine les dents callcin- 
ales dans le V. llttoralla), son callce oblong, plus grand, see 
epls presque du double plus gros, etc." 

It should be noted here that the Loud., Hort. Brit, Suppl, 
(1839) reference given in the bibliographiy above la sometlaes 
erroneously credited to W. Baxter. 

Raimondi (19U3) says "Flores azules, ligeramente moradaa", 
"morado azulino", or "morado claro", and cites his nos. 39U8 , 
Uljlii , 6533, 6727 , and 6976 from Cajamarca, 9568 , 9581i , and 
11139 from Puno, and llltlS from province undetermined, Peru. 
Hayek (I9O8) cites two Wacket collections and a Wettsteln & 
Schiffner collection from SSo Paulo, Brazil. Herrera (I9I1I) 
cites his nos. 1505 , 166U , and 3362 fran Cuzco, Peru, and gives 
the range of the species as "El Salvador, Chile, Uruguay, and 
Argentina." Thome (195U) cites it from Decatur and Glynn 
Counties, Georgia, but describes it aa "rar«" there, Chodat 
(1902) cites Hassler U887 from Paraguay, which, however, I re- 
gard as V. brasiliensis Veil. Johnston (1931) cites Anthony 
380 , Berkeley 231, and Mason 1612 from Socoiro Island, but these 



66 PHYTOLOGIA Vol. 10, no. 1 

are actually all V. sphaerocarpa Peny. 

Rambo, in a letter to me dated August Hi, 19^5, is of the opin- 
ion that his no. 38055, Trtiich I identified as V. montevidensis 
Spreng., is actually V. litoralis because, he says, it was grow- 
ing along a roadside (typical of V. litoralis ) and not in a swamp 
(typical of V. montevidensis ) . The DiJgand & Jaramillo 3030 col- 
lection cited below may have come from either Choc6 or Valle del 
Cauca, Colombia. The name of the Venezuelan collector, Nicolas 
Funck, is misspelled "Funcke" in the Vienna herbarium. The orig- 
inal publication of Humboldt, Bonpland, and Kunth for this spe- 
cies is very often cited as "ISI?", but appears to be more cor- 
rectly dated I8l8. 

Glaziou (1911) cites his no. Ilt362 from Rio de Janeiro, but I 
regard this as V. brasiliensis . Perry (1933) cites the following 
37 additional specimens not as yet seen by me: CALIFORNIA t Amador 
Co.: G. Hansen 202$ (E) . MEHCO: Jalisco: Pringle 11093 (E, F, G). 
Michoac&a: Ars^ne s.n. [Coronilla, 8 Aug. I909] (W); Galeotti 781 
(K)j Gregg 76U (E) . Nayarit: Edw. Palmer 2011ia (G, N) . Oaxaca: 
Pringle U877 (D, E, Q); C. L. Smith 222 (E), 79U (G). San Luis 
Potosl) Edw. Palmer lUl , in part (G); Schaffner 718 (G) . Tabasco: 
J. N. Rovirosa s.n. [Maylto, 10 April I889] (D, N, W) . Vera Crua: 
Mohr s.n. [Huatusco, April 1857] (W)j Seaton 27 (F, G) . GUATEMA- 
LA: Alta Verapaz: Ttlrckheim 90U (D, F) . HONDURAS: Comayagua: P. 
C. Standley 56082 (F) . EL SALVADOR: La Ubertad: P. C. Standley 
23326 (G). San Salvador: Calder6n 729 (G), 925 (G); Renson 291 
(W)j P. C_^ Standley 20617 (G)} Velasco 881t8 (G) . San Vicente: P. 
C. Standley 211^86 (G) . COSTA RICA: Cartage: Cooper 5890 (E, F, 
G) . Province undetermined: Worthen s.n. [April I9IO] (E) . She 
says "This is a widely distributed species with very distinctive 
habit and somewhat variable inflorescence. In some specimens 
the spikes appear to remain compact, in others they tend to elon- 
gate, Schauer used this difference to separate the forms pycno- 
stachya and leptostachya , although he frankly admits the diffi- 
culty of distinguishing the two owing to the intermediate phases. 
V. af finis is characterized by a somewhat coarser floral pubes- 
cence; this, however, seems to be a variable feature, and, as 
such, does not appear to merit more than passing mention." Her 
key for distingtiishing five related species is worth repeating 
here: 
1. Leaves semiamplexicaul and subcordate. 

2. Inflorescence glandular; bractlets conspicuoxisly longer 
than the calyx; corolla-tube 2 — 3 times as long as the 

calyx V. rigida . 

2a. Inflorescence not glandular; bractlets barely equaling or 
only slightly exceeding the calyx; corolla-tube scarcely 

twice as long as the calyx V. bonariensis . 

la. Leaves not semiamplexicaul nor subcordate, tapering into a 



196U Moldenke, Monograph of Verbena 6? 

cuneate-attenuate subsesslle or petlolar base. 
3. Spikes 3 — $ nun. long, very dense and appressed-pubescent . 
U. Frui ting-calyx ascending; schizocarp longer than broad, 

raised-reticulate above, striate to»rard the base 

V. brasiliensis » 
Ua. Fruiting-calyx spreading; schizocarp as broad as long, 

practically smooth V* sphaerocarpa , 

3a. Spikes $ — 10 sun. long, dense at the apex, somewhat open be- 
low and finely strigillose V. litoralis . 

It should be noted here that her \ise of "Bm.** in the above k^ 
seems to be an error for "cm." — errors easily made in the met- 
ric syston of measurement and causing much confusion. 

Acevedo de Vargas (19^1) cites the following: CHILE: Santiago: 
Collector undesignated s.n. [HI .1877] (Sg — U2Ji60), s>n. (Sg — 
5U706)j C, G^ 108U bis (Sg — 5U703). Curic6: Collector undesigna- 
ted s.n. [Llico, XIl"Il56l] (Sg— 5^705), s.n. [Vichuqufin, HI. 
iSSl] (Sg— •U2U6U). Coquimbo: Albert 8.n. [1.1897] (Sg— 68333); 
Vitor s.n. [Xn.l897] (Sg— 6833li) . Valparaiso: Buchtien h (Sg— 
68335) • J^ian Fernandez: Collector undesignated s.n. [XI.186U] 
(Sg— U2U85, Sg— 5U700)j Edwyn Reed s.n. [X.1872] (Sg— li2U7li, 
Sg— 5U701) . 

In all, 998 herbarium specimens, including type material of 
many of the names involved, and 6 mounted photographs have been 
examined by me. 

Citations: ARKANSAS: Ashley Co.: Demaree 16^11 (N), I762I1 
(Ok), 23893 (Sm), 2U;03 (Sm); E. J. Palmer Uli270 (N) . Bradley 
Co.: Demaree 195U3 (N, Sm). Chicot Co.: Demaree 2^6 7U (Sm) . Drew 
Co.: Demaree 214619 (Gg), 3U6U9 (Bm, Sm) . LOUISIANA: Terrebonne 
Par.: Wurzlow s.n. [Hoxma, May $, I91ii] (N), s.n. [Houma, May 9, 
I9II4] (N, N). TEXAS: Newton Co.: Tharp aai. [7-2>39] (Au) . 
OREGON: Multnomah Co.: Suksdorf 2912 (Pi— 138la5), 2913 (Pi— 
1381014). CALIFORNIA: Amador Co.: Q. Hansen 2025 (Du— 2la83, W— 
338509). Boiildin Island [San Joaquin Co,]: Congdon s.n. [Sept. 2, 
1902] (Gg— 31376, W— U6667O). MEXICO: Chiapas: Matuda U38 (Mh, 
Mi, W— I689UI1I;), 836 (N), 5193 (Ld), 5302 (Ld), S.56 (Mi). Chi- 
huahua: LeSuevir 871 (Au) , Hidalgo: G. L. Fisher U6170 (W— 
I88983O); Rose , Painter , & Rose 9077 (W— U52556), 9l5l (W— 
U5263U),- A. J. Sharp Ua828 (N) . Jalisco: McVaugh I69U9 (Mi), 
I720U (Mi) J Pr ingle 11093 (Cm, Me, Me, N, Ut, W— U60U75) . Mfixico: 
Hinton 1575 (N), 3731 (N, N, W— I8UI468I) , 12156 (La), 122la (La); 
M. E. Jones 736 (Po — 7O66I); Kenoyer s.n. [Teotihuacan, 7-30-38] 
(Fs'yy Matuda 27651 (Z), -278814 (Cb) . Michoac4n: Arsene 10 3U (Br), 
3000 (W— I46I1298), s.n. [Rinc6n, 15/7A909] (Br, Br), 8.n. [Quin- 
cao, 11/11/1909] (W— U6U297) ; Barkley , Paxson , & Webster 2677 
(Au— 123277, N); Galeotti 78I (Br, F~photo, N— photo. Si— photo, 
Z— photo); Hinton 121^^kVi, N, Rf, W— 1321761), 122U1 (Mi, N, 



68 PHTTOLOGIA Vol. 10, no. 1 

Ur, W--1821762), 12520 (Au, Mi, N, N, Rf, S, ¥—1890991), 12869 
(Ca— 7Um91, N, W--1977302)j Northcraft III (La, N) . Morelos: N. 
L. H. Kraxias 70 (Ng--6575)j E. lyonnet 6$6 (W— l6U29U8)j Moldenke 
& Moldenke I9826 (Es, Lg, N)j W. Treleaae 22lt (Or). NJ^arit: Edir. 
Palmer 20lU (N), 2019 (Op, W--305272) . Oaxaca: W, H. Camp 2U33 
(N); Prlngle U877 (Br, C, Ca— 10li8Ul, Cm, lo— 38756, Me, Me, Me, 
Mft— 15390, N, P0--6389O, S, Vt, ¥—251682); Schultes & Reko 237, 
in part (Oa— 8209), 1a37 (Me)j C. L. Smith 222 (Ca— 975387, N, W— 
312570) . Puebla: Ara^ 3.n. [near Puebla, 2O/IO/I9O8] (Br, ¥— 
U6U301)j Chute M.ll;2 (Mi)j FrOderstrttn & Hult4n 105l (S)j Nicolas 
s.n. [Rancho Posadas, 2/UA909] (Br, Br, Br)} F. Salazar s.n. 
[Pahuatl4n, lU June 1913] (Me, ¥—1013227) ; A. J. Sharp 140120 
(N) . San Luis Potosl: L. Ij, Davis 239 (N) j R. M. King Ui32 (Au— 
189783, N); C. L. Lundell 12165 (LdJ; Urbina s.n. [Junio de I892] 
(Me) . Sonora: P^ B. Kenned7 70U6 (Ca— 373581, ¥—1287217) . Tabas- 
co: £. N. Rovirosa hhS (C, Pa, ¥—1323106). Tamaulipas: Kerber 
311 (BrJ} Rozynski |^ (Du— 226681) j Stanford , Retherford , & 
Northcr oft 913 (Du— 288697, N, Tu— 15152), 102U (Du— 28626l7 N, 
Tu — I62U3) . Vera Cruz: Barkloy , Rowell , & ¥eb3ter 2536 (Au — 
123273, N)j L. I. Davis 225 (N), 23U (N)7 Dressier & Jones 177 
(Bm, Ca— Ii88^, N, ¥— 23"^26); G. L. Fisher I68 (¥—1315998), 
s.n. [Aug. 13, 1926] (Du— 15U566, Mg— U2); Qaleotti 773 (Br)| F. 
¥, Johnson s.n. [Cordoba, 9-26-06] (N); Kerber 311 (¥—323103)} 
Matuda 836 (Mh, Mi)} C. T. Mohr 317 (¥—771869)} J. V. Santos 
2k6l (Mi)} H. E. Seaton 27 (C, ¥—56168). State undetermined: 
Galeotti 783 [Meratitlan] (Br)} Haenke 1575 (N); Hinton 1575 (N)} 
Liebaann 11336 [Tepiscapa] (¥—1315097). GUATEMALA: Alta Verapaz: 
Cook St Griggs IhB (¥— U07936)} Tttrckhelm 90li (C, Q, Pa, ¥—56170, 
¥—1323108). Amatitlan: Tonduz U87 (¥—1080089) . El Pet6n: Con- 
treras 2635 (Ld), 3088 (Ld)} C. L. Lundell I6386 (Ld). Guatemala: 
Bernoulli 128 (Br, C)} J. F. Brenckle l;7-257 (Gg— 353583, N), U7- 
258 (W, S)7 Tl7-259 (N)} C. C. Deam 6I8O (G, Mi, Vt, ¥—579572)} 
Degener & Degener 26U76 (N, ¥) } Moldenke & Moldenke I98I8 (Lg, N) } 
Ruano 332 (¥—1168396), U03 (¥— II68U27) } ~Te jada Agulrre U (¥— 
691762) } Tonduz 627 (¥— IO8U723) } E. ¥all s.n. [Guatemala City, 
20/U/28] (Ew). Izabal: P. C. Standley . 2l<l|0U (¥— 11500U6). Jutia- 
pa: P. C. Standlaj ]6G^{vJ, Quezaltenango: P. C. Standley 
8319^ (N) . San Marcos: G. Salas 32 (¥—1206750). Santa Rosa: 
Heyde & Lux 3019 , in part (Q), U370 (C, G, ¥—200177, ¥—1323101). 
Totonicapin: P. C. Standley 8U06I (N) . Department undetermined: 
Heyde & Lux 6IO (¥— 2U7U9U) . BRITISH HONDURAS: Gentle 6U8I (Ld), 
7119 (Ld). HONDURAS: Comayagua: P. C. Standley 56082 (¥— 
1U09O90)} Yuncker , Dawson, & Youse "^21^ (Dp, Mi, S, St). Moraz£n: 
S. F. Glassman 1^06 (Ok), ig6U (N), 2011 (N, Ok, Ur)} Molina R. 



196U Uoldenke, Uonograph of Verbena 69 

380 (Ca— 792622), 96O (W— 1975680) j Williams & Molina R. 101$9 
(Mi) . EL SALVADOR: Chalatenango: Tucker 990 (Ca— IOOO906, W— 
2088237). La Ubertad: P. C. Standi gy 23326 (W— 113901?) . San 
Salvador: Calder6n 729 Tn, ¥—1151687), 925 (W— II5189O); M. C. 
Carlson U07 (Ca— 703555); Reason 291 (N, W~39958U)j P. C. Stand- 
ley 20617 (W— 1136U52), 228U2 (W— 1138569)? L. V. Velasco 88U8 
(TT— 829120, W--I323098). Santa Ana: Tncker I308 (Ca— IOOO907, Rf, 
W— 2O88U29). San Vicente: P. C. Standley 2in5r "(N, W— 1137291). 
NICARAOnA: Managua: Garnler 301 (I), 6U2 (Mi)j Macon, Harvey , & 
Valentine 7356 (N, ¥—118106577 7U78 (W— 1181185). COSTA RICA:"" 
Alajuela: Boscilla s.n. [San Ramon, 1935] (N); Branas 11522 (N), 
m6U8 (N)j Chry3lerH9S6 (N, Ru) . Cartago: J. J. Cooper 5890 (C, 
¥—1323107); De ¥olf U25 (Tl); Moldenke & Moldenke 19 807 (N); 
0r8ted 1132^ (¥—1269919); Tonduz U220 (Br), U221 (Br); Torres 
Rojaa 97 (Du— 255758) . Guanacaste: Standley & Valeric U5U30 (¥— 
125Ul08y, Heredia: Brenes s.n, [Vol. Irazu, 1912] (N) . Puntare- 
nas: Maxon & Harvey 7951 (W— Il8l6l5); H. Pittier 3U88 (Br); 
Stork &. Morrison 8919 (Ca— 6U8929, ¥— 2216U83). San J036: Biol- 
ley 1085 (Br); H. Pittier 227 (Br), 691x (Br); Skutch 27U9 (Mi, 
N, 3, ¥— 16U26U5); P. C. Standley 32652 (¥—1225858), 33293 (W— 
I22621I1); Tondiiz 7978 (Br). Department Tindetermined: Kuntze 2109 
(N, ¥—700762); grated 11322 [Mt. Aguacote] (¥—1269918); H. 
Pittier 2389 [Agua callente] (Br). PANAMA: Chiriqul: P. H. Allen 
lla2 (N, ¥—1820068); KilUp 3510 (¥—1012167); Maurice ^7 (¥— 
l5TiTi'n7); Selbert 169 (N, N), 251 (I, N, N, N); ¥ood3on , Allen , 
& Selbert 858 (N, N) . COLOMBIA: Antloquia: Archer 307 (Fn— 1662, 
FQ— 2296, ¥— 15U1839), 105 (Fn— 1661, ¥— 15U2010), 1235 (W— 
15U2389)) Barklqy i Guti6rrez Villegas 1719 (Fn— 3216, N); Baiic- 
Iqy & Puccini B. 32 (Fn); Gavlria Nelra & Barkley 17C236 (N); 
Gutifrrea Villegas & Barkley 17C522 (Fn— 3389, N), 17C585 (Fn— 
3391, N); Gutierrez Villegas , Barkley , & Correa Velisqaez 12 
(Fn), 13 (Fn), 15 (N, N); ¥. H. Hodge 6939 (Fn— 29^8, ¥—1950659); 
Klllip , Barkley , & Daniel 3993U (N, ¥— 195U195); D. Mesa Bemal 
s.n. (Fn— 1658); Robledo & AstrAlaga R. UO (N); Tomas 81 (N); 
Toro Toro 26 (N), 97a (Fn— I66O, N), U39 (Fn— 1659, N); R. Torres 
311 (Fn). BpyacA: Cuatrecasas I698 (¥ — 1773106); Grubb , Curry , & 
Fernandez-Perez 6OI (¥—2322528). Caldas: Dryander 275l (F — 
I2938O2); Permell , Killlp , & Hazen 8593 (N) . Cauca: Asplund 10639 
(S, ¥— 222li287); Cuatrecasas 19333 (N); F. C. LehniaimT?75~ (N, N, 
S, ¥— lli20377); Sneidem 578 (S), IV^ (S, ¥—1705932), 255U (S, 
¥—2103830), U336 (F— 1273^5) . Choc6: Araqae Molina & Barkl ey 
19Chl03 (N); Dug and & Jaramillo 3030 (Hn) . Cundinamarca: Andr? 
618, in part (N); Ariste-Joseph 1003 (¥—1185559); Dugand & Jara- 
iHio 3853 (¥—1853761); Eiran 15^52" (N, W— 2105807); F. R. Fos- 



70 PHTTOLOQIA Vol. 10, no. 1 

berg 2056U (N, ¥—2108727), 21008 (N, W— 2IOI896); Grant & Gio- 
vanni 9672 (N, W~2106891)| Niemajer lU9 (W--1U212U9) ; P6rez ^ 
bel6eg 1207 (W--1517802) ; Schiefer 636 (Du--372831) ; M. Schneider 
128 (S) J Tarrag6n, Araqpie Molina , & Bariclej iSCulOl (Es, N) . Hui- 
la: F. R. Fosberg I92U9 [U. S. Nat. JLrb. 282U05] (W—2165313) ; 
Rusby & Pennell 588 (N) ; Sneidem 2$55 (S) . Magdalena; Angel 66U 
(N); Cuatrecaaaa & Castaneda 2^296 (Fg). Norte de Santander: Cu- 
atrecasas, Schultea , & Smith 12125 (Hn) . Santander: Araqae Molina 
& Barkley 18S211 (N); Barkley & Araque Molina 18S097 (N); Killip 
& Smith 16756 (N, W— 1352U53); Langenhelm 3128 (Ca— 78O89). Toli- 
ma: F. W. Pennell 3211 (N), 3362 (N). Valle del Cauca: Cuatrecas- 
as VM (N), I8052 (VI), 1857U (VI), 1865U (N), 20663 (N), 22877 
(F— I3U1699); Duqae 1605 (N, ¥--17UU5l5); Garcia & Barriga 6hh3 
(¥--17Ui953)i HoltoiTIo^ (T); Killip , Cuatrecasas7 & Dryander 
39207 (N, W— 1856731) ; Moldenke & Moldenke 19791 (Es, N, Sm) . 
Vatip6s: Gutierrez Villegas & Schultes Tifl, in part (Fn), 7U2 (N) . 
Department undeteimined: Apollinaire & Arthur 71 [Sabana] (W — 
603153) J Funck & Schlim 637 (C) . VENEZUEIA: Aragua: Chardon 182 
(W--lb01129), 7H2 (Ve— ^12839); Fendler 852 (Br, Cb, N); E. G^ 
Holt 376 (Ve), 377 (Cm), 38O (Cm), hk3 (Cm); Moldenke & Moldenke 
19550 (Es, l€, N, Sm); Yogi 938 (N). Federal District: Bailey & 
Bailey 352 (Ba, W— 1198320); Burkart I6OI6 (Ve); Collector un- 
designated 517 (S), s^ (Ve— 1281i6); Eggers 13053 (W~123U535), 
13053a (W--123li536); D. Huimnel s.n. [Caracas, 9/IV/1958] (Go, S); 
Kuntze 1263 (N); Laseer 725 (N); Linden 33li (Bm);> H. Pjttier 
9732 (Ba, N, Ve, W— 1069U97); Potter 5100 (Ms); Tamayo s.n. 
[Herb. Est. Exp. Agric. 737] (W--I8OO66I) . Larfi: Saer 329 (Ve). 
M6rida: Gehriger 219 (Ve, W— lli99053) , 533 (N, Ve); Gines U685 
(W-~2167773); Hanbury-Tracy 21* (K), 256, in part (K); Jahn 535 
(W--112025U) ; Usser hXS (Ve— 12838); E. Reed 610 (W— 1618551) . 
Miranda: Barrus s.n. [January 11, I9U0] (It); E. G. Holt hhX 
(Ve); Moldenke & Moldenke 19562 (Es, I^, N, Sm) . Sucre: Funck 5U 
(Lu, V), 325 (Lu, V), 637 (Lu, V). Trujillo: Bellard s.n. [Aug. 
1923] (¥—1189293); Burkart 16815 (Ve); Gines U312 (W— 2167652) . 
Zulia: Mocqaerys 893 (W— 2282li93) . State undetermined: Eggers 
13U53 [Los Lhorros] (W— 132311;7) ; Groaourdy s.n. [1862; Herb. 
Reichenb. f . 116382] (V); G. H. H. Tate la"[Carapas] (W— I23083U). 
ECUADOR: Azuay: Harling 625 (S); Rose , Pachano , & Rose 228U5 (N), 
228U6 (N), 23805 (W— 1023117). Cafiar: W. H. Camp E.2li95 (N, W— 
2056989). Chimborazo: Anthony & Tate U38 (W— 1192501); Asplund 
15501 (S); Penland & Summers U65 (N); Rlmbach . 671 (N, W— 
17li2069); Rose & Rose 22538 (N, W--1022188) . Cotopaxi: Harling 
U867 (S) . El Oro: Asplund 15820 (S) . Guayas: Eggers 1U372 (W— 



196U Moldenke, Monograph of Verbena 71 

1323011i). Loja: R. Eapinoaa 58 (N)j Harling STfk (S) . Los Rlos: 
Asplund 5577 (S, W—19 30550) . Napo-Pastaza: Asplund 18329 (S) . 
Pichincha: Flrmln 26U (W--li20227) j !_. Holmgren $6k (S, W— 
2059935)i Moldenke & Moldenke 19785 (N) . Santiago-Zamora : Harling 
951 (S) . Tunguragua: Pachano 11 (N, W — 10l4li505) . Province xinde- 
termlned: Andr6 6l8, in part [Facatativa] (N); Anthony & Tate 267 
[Las Maquinas, West Andes] (W — ll923U6)j Jameson s.n. [Republic of 
Ecuador; Herb. Reichenbach f. 125172 & 125205] (V, V). GALAPAGOS 
ISUNDS: Charles: N. £. Anders son 117 (Lu); A. Sterrart 3321 (Gg— 
31372), 3322 (Gg--31371), 3323 (Gg~31377) . Chatham: N. J. Andera- 
son s.n. [1853] (Br, S), s.n. (S)j Schlmpff 132 (Gg— 212ltll8, S), 
11|2 (Gg— 212U6U, N, S); A. Stewart 332U (Bi, Gg— 31379, W— 
921600). Indefatigable: R. G. Taylor T.T.125 (N) . PERU: Amazonas: 
Soukup U119 (N). Ancash: R. Ferreyra 7U51 (W— 20U9Ul6) . Arequipa: 
Cook & Gilbert U6 (W— 6033U1) ; Hinckley & Hinckley 6U (Gg— 31380, 
N, W— 1197767) . Cajamarca: Ochoa 1571 (N). HuAnuco: H. A. Allard 
20970 (N), 21013 (W— 199986U)j Asplund 13110 (S); Collector un- 
designated 3.n. [Huancayo] (B)} Ochoa 1059 (N), 1076 (N); Sande- 
man 3li98 (K)j C. Swingle 9 (W— 2058172), ^ (N, W— 2058l90)j 
Woytkowski 3U175 (Ca— 11^296), 3U503 (Ca— lla86). Junln: Killip & 
Smith 21925 (W— I35703I) ; Macbride & Featherstone 1705 (S) ; San- 
deman U566 (K) . La Libertad: ^ Castafleda 2906 (S); Ellenberg 
1783 (Ut— 115379b), 1921 (Ut— 115382b),- Eyerdam 8897 (Ca— 
?f557U); N6e 87 (Q); E. G. Snyth 25 (W— l8022Ut) ; J. West 8I9O 
(Ca— 565217). Lima: P. Agtdlar F. 1079 [27 Feb.] (N), 1079 [20 
July] (N); N. £. Andersson s.n. [S. Lorenzo, 1852] (S)j Asplund 
11096 (S); Collector undesignated s.n. [Lima] (B); Cook & Gil - 
bert 2088 (W— 703675); Ellenberg 28 (Ut— 115396b) , 2771 (Ut— 
115387b); R. Ferreyra U087 (N, Ug), U088 (N); Ochoa 1133 (N); F. 
W. Pennell lli770 (N, W— I3U0936); Rose & Rose 185U8 (N, W— 
76I23O) ; Soukup 1778 (W— I83070U) ; Stork & Vargas 9329 (Ca— 
655898); Wilkes s.n. [Callao] (W— 71951) • Loreto: Ducke s.n. 
[Herb. Jard. Bot. Rio Jan. 2559U] (N); C. Swingle 89 (W— 2058216); 
Tessmann U205 (Hb, S); U. Williams U52 (Bi), 7U3 (W— m96857), 
2588 (W— li49666U) , 265U (La). San Martin: H. A. Allard 20U00 (Ca- 
926311), 2I805 (W— 2025215); Spruce 1072 (Br, N) . Piura: Ochoa 
1757 (N). Tacna: H, H. Rusby 910 (C, Mi, Mi, Pa, Pr, W— 71997), 
913 (C, P») . Department undeterminedt L. Fernandez 16506 (Kr) . 
BRAZIL: Federal District: FrazSo s.n. [Herb. Jard. Bot. Rio Jan, 
7U63] (N); Occhioni 523 [Herb. Jard. Bot. Rio Jan. 552li6] (N) . 
Minas Gerais: Assis 239 (G, N, W— 1932536) J Mendes Magalhffes 
2219 (Be— 13790), 3151 (Be— Iii8l6, W— 212U321); Warming s.n. 
[Lagoa Santa] (N); L. 0. Williams 5650 (G, N) . Parani: Braga &. 
Lange 85 (Bm); Dusfa 2II9O (Ja— lii8U6), 7827 (N), 10856 (I, Mi, N\ 



72 PHTTOLOGIA Vol. 10, no, 1 

8«n. [Herb. Jard. Bot. Rio Jan 2U90] (N), s.n. [Herb, Jard, Bot. 
Rio Jan. lii8U6] (N)j Reiss |6 (I); Stellfeld 1109 (N), 113U [Herb. 
Mas. Paran. 2116] (N), 1260 [Herb. Mus. Paran. 2371] (N), a.n. 
[Herb, Mils. Paran. l63li] (N) . Rio de Janeiro: 0, M. Earth 1.38 
(Herb. Inst. 0. Cruz 166] (W--23U2932)j Diaa dos Santos & Frota 
Pessoa s.n. [Terezopolis, Feb, 2^, 19U2] (Ja — l46795)j Diogo 506/ 
606 (Ja — U6766)j C. V. Freire s.n. [Herb. Mus. Nac. Rio Jan, 
IIS522] (N) } G. Gardner 233 (U) J Glaziou lla62 (N) } G6es & Constan - 
tino 297 [Herb. Jard. Bot. Rio Jan. 51881] (N); A. Lutz U30 (Hk); 
B. Lutz 1185 , in part (N, N, N)j Mello FIT ho s.n, [Terezopolis, 
Feb. 19U2] (Ja— U6789)} J. T, Roig H;386 (Es); Rose & mtz 6 (Gg- 
366822); Sampaio 161;5 (Ja— U6501), U867 (Ja— U6508), 7580 (Ja— 
U6512), 7619 (Ja— It65l3), s.n. [Petropolis, Mar, 1, 1936] (Ja— 
U67U7); H. P. Velloso 10 (Ja— U6536)j J. Vidal n .507U (Ca— 
I69U87). Rio Grande do Sul: Henz 326U6 (N)j Leite 2li9 (N)j Und- 
man A.U75 1/2 (S); Moldenke & Moldenke I968O (Es, Lg, N); Raabo 
U36, in part (N), U36b (Ok), U5117 (Go), U6UI6 (w— 1997U05)j 
Reineck & Czermak 63 (Ug); Reitz C.5l6 (N); Sehnem U;36 (N), 
3519 (Gg— 356385), Santa Catarina: Ule 1067 (W— 1323100) . Slo 
Paulo: Cantos Novaes lllj (N, Sp— 15719); £. A, Krug s,n, [Herb, 
Dist, Bot. S. Paulo UOOO] (W— 1775606); Moldenke & Moldenke 19633 
(Es, Lg, N, Sm); Pickel 5301 (Sp— U5703); L. Roth~866 (Sp— 5lli77). 
State undetermined: Sellow s.n. [Brasilia] (Br) . BOLIVIA: Cocha- 
bamba: Steinbach 5901 (Cb) . El Beni: Buchtien 5889 (W— 1159358), 
5890 (W— 1159363). La Paz: M. Bang 20li (Mi, Pa, W— 719l;7); Buch - 
tien 197 (N), 351 (W— 1159362)7^97 (S); Mandon 52b [Macbride 
photos 2U68I] (Kr— photo, N, N— photo, S); H. H, Rusby 907 (Pa). 
Santa Cruz: Peredo s'.n. [Cabezas, Cordillera, 29-I-19U6] (N), sn 
n, [La Pampa de La Cruz, 2>IV-19U6] (Gg— 353261, N), s.n. [Las 
Juntas, IO-II-I9U7] (N); Steinbach 3137 (N), 6757 (Ba, Ca— 
368^83, S, Ut^91359). PARAGUAI: W. A. Archer"T788 (N, W— 
17055U2); Fiebrig U;32 (V), I4635 (Bm, V); Hassler 121^2 (N), 2585 
(N, V), 3135 (N), 5203 (Ca— 93^80, N), 11539 (Ca— 930261, N)j 
JOrgensen 3767, in part [Villarica, 10.1.1929; Herb. Osten 2221*8] 
(Du— 197905, S, Ug, TT— lU8392li); Lindman A.36U7 (S, S), A. 3651 
(N, S); Lorentz IO8 (B); T. Rojas 13556 (Go), I889 [Herb, Hort, 
Parag, 1005U; Herb. Osten 13556] (Ug), s,n. [Puerto Colonia Rlsso, 
Dec, 1916; Herb, Osten 13557] (S), URUGUAY: Caste llanos s.n, [Bel- 
la Uni6n, Jan. 28, I9U8; Herb, Inst, Miguel Lillo 15188] (N), s. 
n, [Plaja Atlintida, Dec. 29, 19U6; Herb. Inst. Miguel Lillo 
l5l95](N), 6JCX, [Punta del Este, Dec. 27, 19U6; Herb. Inst. Migu- 
el Lillo 11757] (N), s.n. [Valle EdSn, Feb. 18, 19U7; Herb. Inst. 
Miguel Ullo 15758] iW Ta .n. [San Carlos, Jan. 16, 19U7; Herb, 
Inst, Miguel Lillo l5760]"(Ny; GaTllnal , Aragone , Bergalli , Camp- 



196U Moldenke, Monograph of Verbena 73 

al, & Roaengurtt 1269 (N), PE.$lt83 (N); Herb« Herter 966I (N), 
9^078 (N), 9$$2$ (N)i Herter 269 [Herb. Herter 81713] (B, Ca— 
3IU28U, S, SP--2663U), 269b [Herb. Herter 7872U] (He), s.n. [Herb. 
Osten 181;89] (Ug), s.n. [Herb. Osten 18U91] (Ug), 8.n. [Herb. Os- 
ten I81i97] (Ug), s.n. [Herb. Osten I903U] (Ug); Moldenke & Mol- 
ded 19691 (N), 19697 (N); Osten 283O (Ug), 333^ (Ug), l^l^JJg)i 
Rosengurtt A.12la (N), 6.^07 (N) . CHILE: Atacama: Morong 120U (C). 
Colchagua: Barros Valenzuela 80li6 (N) . Concepci6n: Looser 399$ (N). 
Coquimbo: Barros Valenzuela 80I4I , in part [Terena] (N)j Blese 
2109 (N); Field 3.n. [neighborhood of Coqtdmbo] (Br); Looser $$06 
(N). Curic6: Barros Valenzuela 8OUO (N)j Herb. Mus. Nac. Chile ig 
(N). Malleco: Kunkel 2500 (Cb) . Santiago: Barros Valenzuela 8225 
(N), 8226 (N). Valparaiso: N, J. Andersson s.n. [Valparaiso, 1853] 
(S, S, S)j Buchtien s.n. [18^0/95] (B, B, S, W--ll59370)j Claude- 
Joseph 3550 (Tir--128338l)j Garaventa I66I [Herb. Looser U026] (N); 
T, H. Goodspeed 23326 (Ca — 657U79) ; W. IK Harvey s.n. [Valparaiso, 
April-July 1856] (S); Killip & Pisano 397U6 (N, If— 195U028)| 
Looser lljOl (N), li033 (N); Moldenke & Moldenke 19769 (N); R. A. 
Philippi 1302 (W--I323097) ; Wilkes Exped. s.n. [Valparaiso] (T). 
Province undetermined: Barros Valenzuela 80lil , in part [Cerro de 
la Virgen, Viufla] (N)j C. Q^ 79U (Br, N); Petr6 s.n. [Emellan] 
(S). JUAN FEIiNANDEZ ISUNDS: Masatierra: G. T. Hastings 225 (W— 
530155); £. Ingram s.n. [1938] (Bm); C. Skottsberg UOl (Go, Lu, 
S); Skottsberg & Skottsberg I80 (Go, S) . Island undetermined: D. 
Douglas 3$ (Lu), UO (Lu) . ARGENTINA: Buenos Aires: R, Alvarez I83 
(S), 311 (N); N. J_. Andersson s.n. [Buenos Ayres, 1852] (S); Cab- 
rera 1^57 [Herb. Osten 22285] (Ug), U317 (N); Chicchl 5 (N); Col - 
lector \indesignated s.n. [La Boca, 29 June '82] (C); G. Dairson 
9U6 (N); Moldenke & Moldenke 19710 (N), 19711 (N); N6e 115, in 
part (Q); Nlcora 57U (W— 2196Ii67) ; Venturi 68 (S), 93 [Herb. Inst. 
Miguel Lillo 31395] (N); Wall i Sparre sja, [EL Aroma, 27/10/li6] 
(Ew, Eir, Ew). Catamarca: A. Brizuela 986 (N); J. Brizuela I30 (N); 
Cabrera 1186 (N); JOrgensen 1295 [Herb. Osten II3U8] (Ug); Luna 
Risso 281 (N), 9^9 (Es, N), 1097 (H); T. Meyer 125U6 (N); Venturi 
7076 [Herb. Osten 20839] (Gg— I6063O, Ug, W— 1591383) . Chaco: R. 
M. Aguilar 8O3 (N). c6rdoba: Balegno 893 (N); Bmch 2635 (N, N, 
Ug); Castellanos s.n. [Valle de los Reartes; Herb. Osten 15278] 
(Ug), s.n. [Herb. Mua. Argent. Cienc. Nat. 31201] (N), a.n. [Herb. 
Mus. Argent. Cienc. Nat. 31203] (N); J. Gutierrez 113 (N); Kuntze 
a.n. [ni.1891] (N); Moldenke & Moldenke 19713 (N); Pierotti~^K 
(Es, N); Rodrigo 2310 (N); Villafane 3351 (N) . Corrientes: Cas - 
tellanos s.n. [Herb. Mus. Argent. Cienc. Nat. 3UU87] (N); Ruiz 
Huidobro WU (N), 3831 (N), 3882 (N), U19O (N, We), U27S (N), 
1;3U7 (N),~Il3^8 (N),T59U (N),"I525 (Es, N, We), U710 (N, N) . For- 



7li PHITOLOGIA Vol. 10, no. 1 

mosa: I. Morel 676 (N), 806 (N), 1392 (N), II46O (N), 1657 (N, Rf), 
1695 (N), 17^9 (N), 1831 (N), 3.n. [Riacho Porteflo, IX/I9U6] (N)j 
Sj^ A. Pierotti U200 (N, N) . Jujuy: Moldenke & Moldenke 197U9 (Lg, 
N), 197^1 (Lg, N). Mendoza: Ruiz Leal 868 (Rl); Smitt 8.n. [I89O- 
91] (S). Mislonea: Ekman 2017 (N, S), 2030 (Ml); Ullieskqid sji. 
[vie. Colonia Bonpland] (S)j Medina 128 (N)j T. Meyer 11$9U (N)j 
G. J. Schwarz 2280 (N), 3898 (Au— 122318, N); Schtrindt 7 (N). 
Salta: Garolera & Rcmero s.n. [15-I-19U7] (N)j Malvairez 137 (N)j 
Moldenke & Moldenke 19737 (N), 197U2 (Ea, Lg, N, Sm)j Venturi 
5U71 (V).'"santa f6: R. Alvarez 900 (N), 951 (N)j Job 686 (N), 
10li2 (N); Ruiz Leal lli2$2 (Rl)j Terribile 382 (N), U3$ (N), $$6 
(N) ; Wall & Sparre s.n. [Rosario, 8/II/U6] (Eir) . Santiago del Es- 
tero: Pierotti h, in part [Herb. Inst. Miguel Lillo 100888] (Bn). 
Tucumin: Moldenke & Moldenke 19725 (N), 19726 (N), 19727 (N)j 
Monet ti 185 [Herb."*Osten IO87I] (Ug), 1719 [Herb. Inst. Miguel 
Lillo 31II00] (N)j O'Donell 68 [Herb. Inst. Miguel LiUo 36206] 
(N)j Olea 198 (Ca)} F. Ortiz s.n. [30/5A5] (Ca)} Osten 10615 
(Ug); Hocha 38 30 (Vi, Vi); Ruiz Leal I2U0I; (Ss); Schreiter^7 
[Herb. Osten 150U5] (Ug), s.n. [Cadillal, Nov. 11, 1917; Herb. 
Osten 12178] (Ug); Terribile IO6 (N), 252 (N), 357 (N); L. A. 
Varela 3.n. [Maccmita, 10-in-19UU] (N) ; Venturi 50 (¥—1^91209); 
E. Villa 686 (N); Wall & Sparre s.n. [S. Javier, II/II/I46] (Ew), 
s.n. [Avenida de Lima, ll/H/hS] (Ew, Eir), a.n. [Avenida de Lima, 
I2AIA6] (Ew), 3.n. [La Famailla, li/ll/i6] (Eir, Ew) . GERMANIi 
Schulz & Schulz s.n. [Berlin, 3.10,1898] (B). REPUBLIC OF SOUTH 
AFRICA; Natal: Meebold 12826 (Mu); J^ M. Wood 12122 (Ew), 13132 
(Bi, Vi) . Transvaal: Rodin 3917 (S, W— 19911^). MIDWAY ISLAND: 
Meagher s.n. [July 7, 1933] (N), s.n. [July 7, 1937] (Bi). HA- 
WAIIAN ISUNDS: Hawaii: M. Brown s.n. [Kilauea, Jtdy 10, 1931] 
(We, We); C. N. Forbes U65h (Bi); Hosaka 2122 (Bi); A. R. Molden- 
ke 77 [H. N. Moldenke 21813] (Mi), 79 [H. N. Moldenke a8l8] (Mi); 
Rubtakoff 2615 (Mi). Kauai: C. N. Forbes 272k (Bi); A. A. Heller 
20U6 , in part (Bi, Bi, Bz— 237U0, G, Ca~50U930, Gg— I63IOU, Mi); 
Kusche 70 (Gg— 31378); A. R. Moldenke 87 [H. N. Moldenke 2l8Ui] 
(Mi); H. N. Moldenke 21839 (Mi) . Lanai: C. N. Forbes 188.2 (Bi); 
G. C. Munro Ug (Bi, N), 292 (Bi) . Maui; C. N. Forbes 1083m (Bi, 
W— 1^79108), 2036m (Bi); H. N. Moldenke 2lg2S" (Mi), 21827 (Mi) . 
Molokai; 0. Degener 17852 (N, S); A. S. Hitchcock 1516I (W— 
892U15); H. N. Moldenke 21835 (Mi) . Niihau: J^ F. G. Stokes sja. 
[Ponds on southern end, January 1912] (Bi), s.n. [South half of 
island, January 1912] (Bi, Bi), s.n. [January 1912] (Ba) . Oahu: 
N. £. Anders son s.n. [Honolulu, 1552] (N, S, S): O. Degener 1785U 
(N); Eastwood s.n. [Honolulu, August 1~16, 192li] (Gg---3U500) ; F. 
E. Egler 37-83 (Bi); C.N. Forbes s.n. [Valley above Country Club, 



196U Moldenke, Monograph of Verbena 75 

July 8, '08] (Bl), s»n. [Tantalus Rd., 7A8/30] (Bi); F, R. Fos- 
berg 8856 (Bi, N), 10276 (Bi, Du~239Ul7, N, Up)j D. W, Garber 
339 (Bi); H. W. Graham 3 (Cm); Qrunorr s.n. [Juli 188U] (V—^SUi); 
K. Harley 3»n» [Waianae, May U, 1956] (Bi) ; Hasegawa a,D. [Makaha 
Valley, April 3, 1932] (Bi); A. A. Heller 20I46 , in part (W— 
262858); Herb. Oahu Coll» 65 (Mi); A. S. Hitchcock 13726 (T»— 
898819) ; C. H. Hitchcock 8,n. [Honolulu] (Dt) ; Hoaaka 1351t (W— 
I993U15); E. P. Hume 123 (Bi); S. W. Hutchinson 6030 (Bl— 13769); 
C. S. Judd n (Bi); Kelly 12 (Gg--311i63); Rigga a.n. [Honolulu, 
July 1908] (Or— 8850) ; J^ F. C_. Rock 3007 (Bi) ; P. Russell 52 
(Bi). Island undetermined: Mann & Brigham U37 (Bi) . AUSTRALIA: 
New South Wales: Kaspiew s.n. [Moss Vale, ii.6,5l] (Ew); Valentin 
aji. [6/11/1927] (S, S), s.n. [13 Mars 1928] (S). Queensland: 
Kingston K.83 (Ng— I68U7, Ng) . AUSTRAL ISLANDS: Raivavae: Chapin 
857 (N); Quayle 258 (Bi); Saint John & Fosberg 15869 (Bi) . Rapa: 
Chapin 893 (Bi, N); £. C. Curtis 307 Tsi); Herb. Whitney Exped. 
307 (W— I968I3I), 337 (Bi); Saint John & Fosberg 15235 (Bi); A^ 
M. Stokes 195 (Bi) . Tubuai: Saint John & Fosberg I628O (Bi) . 
GAMBLER ISLANDS: Aukena: H. Saint John ll|632 (Bi) . Mangareva: Ag- 
assiz 122 (W— 1652126); H. Saint John lli570 (Bi) . EASTER ISLAND: 
Agassiz 20 (Go); Chapin 102U (N); Gusinde s.n. (Go); Herb. Eayed. 
Franco-beige s.n. [Rano-Kao, 7AoA93U] (Br), s.n. [Rsmo-Aroi, 
l8/10/3i4] (Br). CULTIVATED: Belgium: M. Martens s.n. [h. b. lov. 
1837] (Br); V^ Heurck 32 (Cp) . France: Herb. Decaisne s ji. [h. 
p.] (Br). Java: Hallier D.589 (Bz--237l8, Bz— 23719, Bz--23720), 
D.59O (BZ--2372I, Bz— 23722); Herb. Hort. Bot. Bogor. IV.KJl.ILV. 
17 (Bz — 26UU2, Bz — 26Ui3) . New York: Eggleston 8.n. [seed frcm 
Madrid, 1905; N. T. Bot. Card. Cult. PI. 23202] (N) . Spain: Herb. 
Hort. Reg. Mabrit. I46 (Q) , Sweden: Reuterman s.n. [12/9/1889] 
(Go). LOCALITT OF COLLECTION UNDETHIMINED: Collector xmdesignated 
2933 (Vi); Herb. Mus. Bot. Stockholm B.n« (sYI 

VERBENA LITORALIS var. ALBIFLORA Moldenke, Pl^rtologia 1: U32. I9UO. 

Bibliography: Moldenke, Suppl. List Common Names 23. 19iiO; Mol- 
denke, Phytologia 1: U32 (19U0) and 1: 511. I9UI: Moldenke, Known 
Geogr. Distrib. Verbenac, [ed. 1], I9 & 101. I9II2; Moldenke, Cas- 
tanea 13: 115. 19U8; Moldenke, Alph. List Git. 2: 5U2. 19li8; Mol- 
denke, Phytologia 3: 133. 19U9; Moldenke, Known Geogr. MLstrib, 
Verbenac., [ed. 2], 33, 61, & I98. 19U9; Moldenke, R6sum6 39, 69, 
& U72. 1959; Moldenke, R6svaai> Suppl. 6: 6. I963. 

This variety differs from the typical fozn of the species in 
having white corollas. 

The type of the variety was collected by George B. Hinton ( no. 
13965 ) in a llano at an altitude of 1000 meters, at Coalcauan, 
district of Coalcanan, Michoacdn, Mexico, on July 20, 1939, and 
is deposited in the herbarium of the University of California at 



76 PHITOLOGIA Vol. 10, no. 1 

Los Angeles. The foxm ascends to 2700 meters altitude in Choc6. A 
vernacular name recorded for it is »»verbena blanca", inaccurately 
given for the typical f onn of the species in my "Supplementary 
list of coomion and vernacular names....** in 19U0. 

In all, 6 herbarium specimens, including the type, and 2 moun- 
ted photographs have been examined. 

Citations: MEHCO: Michoacin: Hinton 13965 (La — type, N — photo 
of type, Z — photo of type). COLOMBIA: Antloquia: F. A. Barkley 
17C3li6 (Fn— 3223, N). Choc6: Araque Molina & Barkley 19Ch010 (N, 
W— I99962U). PEKU: San Martin: H. A. Allard~20399 (W--1999720) . 

VERBMA LITORALIS var. CARAGASANA (H,B.K.) Briq., Ann. Conserv. & 
Jard. Bot. GenSv. 7-8: 292, I90U. 

Synoaymyt Verbena caracasana H.B.K., Nov. Gen. & Sp» PI. 2: 
275 — 276. 1818. Verbena caracasana Humb. ex Spreng. in L., Syst. 
Veg., ed. 16, 2; 7U8. 1825. Verbena caracassana H.B.K. ex Cham., 
Linnaea Jt 255* 1832. Verbena caracasana Humb. & Bonpl. ex Steud., 
Nom. Bot., ed. 2, 2: 750. I8UI. Verbena caracasana Humb. & Kunth 
ex D. Dietr., Syn. Pi, 3s 601. 18U3. Verbena caracasana Kunth ex 
Schau. in A. DC, Prodr. lit 5U2, in syn, I81i7. Verbena litoralis 
var. caracasana Briq., Ann. Conserv. & Jard, Bot, Gene v. 7-8; 292, 
I90U. Verbena litoralis var, caracasana (Kunth) Briq. ex Moldenk^ 
Suppl. List Invalid Names 25, in syn, 19U7. Verbena littoral is 
var. caracassana Briq. ex Moldenke, R6sum4 369, in syn, 1959. 

Bibliography: H.B.K,, Nov. Gen, & Sp, Pi. 2: 275. 18 18; Spreng. 
in L., Syst. Veg., ed. 16, 2: 7U8. 1825; Steud., Nom. Bot., ed. 
2, 2: 750. I81a} Cham., Linnaea 7: 255. 1832; D. Dietr., Syn. Pi. 
3: 601. 18U3; Walp,, Repert. Bot, Syst, U: 19. 181^5; Schau. in 
A, DC, Prodr, 11: 5U2, 181^7; Schau, in Mart,, Fl, Bras. 9: I89, 
1851} Hook. f. & Jacks., Ind. Kew, 2: 1178, 1895; Briq,, Ann, 
Conserv, & Jard, Bot, GenSv. 7-8: 292, I90U; Briq. in Chod, & 
Hassler, Plant. Hassler. 10: U8l. 190li; Fedde in Just, Bot, Jahr- 
resber. 33 (1): 632, I906; Moldenke, Prelim. Alph. List Invalid 
Names U5 & U7. 19liO; Moldenke, Alph. list Invalid Names U6 & 1^8. 
19U2} Moldenke, Known Geogr. Distrib, Verbenac, [ed. 1], hk &. 
101. 19U2; Moldenke, Alph. List Invalid Names Suppl, 1: 23 St. 2$, 
19U7; Moldenke, Knoim Geogr, Distrib. Verbenac., [ed. 2], 61, IO6, 
& 198. 19U9; Moldenke, Alph. Ust Cit. 3: 8O8 (19U9) and U: 107U. 
19l;9; Moldenke, R6sum6 69, 127, 361, 368, 369, & U72. 1959; Mol- 
denke, R6sum6 Suppl. 3: 12, Ui, 37, & UO (I962) and li: ll^. 1962; 
Moldenke, Phytologia 8: 3ll» & 317 (1962) and 9: 67 & l5l. 1963; 
Moldenke, R^simi^ Suppl. 8: 5* 196U* 

The original description of this taxon by Kuntji (1818) is as 
folloirs: "V. caule erecto, ramoso, rarals cpiadrangularibus, glab- 
risj foliis oblongis, acutis, grosse crenato-serratis, strigoso- 
scabris; spicis f ilif ormibus , paniculatis. Crescit rarisslme in 
sylvaticis prope Caracas, alt. U30 hex. 4 Floret Januario, CAULIS 
erectus., stibquadripedalis, ramosus; ramis quadraiigularibxis, sub- 
striatis, glabris. FOLIA opposita, subsessilia, oblonga, superi- 



196U Moldenke, Monograph of Verbena 77 

ora lanceolata, acuta, basi cuneata, grosse serrata, serraturis 
rotxindatls, apice acutis, altemls minoribus, rigida, reticulato- 
venosa, nervo medio venlsque primariis subtvis prominentibus , sup- 
ra pilis minutis ac^z^sais scabra, subtus in nervo medio venisque 
strigosa, quadri- aut quinquepoHicaria, seaquipolliceoi et paulo 
latiora. SPICAE terminales, paniculatae (in specimine suppetente 
q[uinc[ue oppositae, inferiores distantes, longissime pedunculatae), 
filiformes, pedunculatae, FLORES sesailes, bracteati, minuti. 
BRACTE/LE lanceolatae acuminatae, glabriusc\ilae , calycem subae- 
quantes. CALH generis, hispidulus. COROLIA pallide violacea; tu- 
bus calyce longior; limbus quinquefidus, planus} laciniis rotun- 
datis, subemarginatis . STA.11INA ET PISTILLUM generis, FRUCTUS 
calyce persistente inclusus, elliptico-tetragonus, quadrilocula- 
ris, quadripairbibilis , sulcatus, fuacescens, glaber, magnitudine 
grani sinapis; loculis monospermis , " 

The type of this perplexing taxon was collected by Aim6 
Jacques Alexandre Bonpland at or near Gau-acas, Venezuela, and iras 
probably deposited in the herbarium at Berlin now destroyed. It 
seems veiy possible — and probable — that this variety should 
be reduced to gynonymy under typical V. litoralis H.B.K., even 
though Eunth, Chamisso, and Walpers considered the two taxa suf- 
ficiently distinct to warrant sepsorate specific designation. Un- 
fortunately, I have not yet been able to examine any of the orig- 
inal Bonpland material — if, indeed, this is still extant — 
nor even the Hassler material ( nos. 1027 and 1027a from Paraguaj^ 
cited by Briquet when he reduced the species the varietal status. 
Briquet was of the opinion that V. caracassma was the same taxon 
as Schauer's V. littoral is ^ leptostachya , which I regard as 
typical V. litoralis . Schauer also regarded the Bonpland plant 
as identical with his variety, apparently using the Herb, Will- 
denow 1 11 3U specimen labeled V. lanceolata by Willdenorf as typ- 
ifying the H,B,K. name, Chamisso (1832) also gives Willdenow' s 
name as a synonym of the H,B,K, name, commenting: "E Brasilia 
meridional! 'Campo, Rio-pardo' misit Sellowius. Diversa nostra 
ab Humboldtiana stirpe, ut planta campestris sitiens a sylves- 
tri vel palustri. Folia illi poUice saepius breviora vix unquam 
bipollicaria (nee U — 5-pollicaria) , superiora lineari-lanceolata 
integerrima, inferiora tantum serrata. Ramosior est, ramis 
gracilioribus striatis, angulis insignius nervosis, caeterum 
differentia nulla." Steudel (l8Ul) likewise reduces V. lanceolata 
Willd. to V, caracasana H.B.K. TThether this course of action is 
justified or not, I cannot as yet say, not having seen the Will- 
denow specimen myself as yet. It is possible that Willdenow's 
plant was not part of the type collection of V. caracasana. 
Schauer's variety, moreover, was based on various other collec- 
tions in addition to the Willdenow Herbarium specimen. 

All things considered, therefore, it seems best to hold the 
variety apart tentatively until these matters can be settled 
definitely. In my publications prior to the year 19U2 I reduced 
the variety to V, litoralis, but from 19U2 onwards have kept it 



78 PHTTOLOGIA Vol. 10, no. 1 

separate . 

If the specimens cited below really represent var, caracasana, 
then the corolla is said to be blue-violet, it floirers in Jxily, 
and has been found at altitudes of 2300 to 2U00 meters. 

It should be noted here that the original publication is often 
cited as "1817". 

It is possible that the Htnmnel s.n. , cited by me under typical 
7. litoralis , from Caracas, is actually var. caracasana , since 
its spikes are abnormally dense; the same is true of Pemaree 
2h6h9 from Arkansas. 

Briquet's notes irtien he established the trinomial here employ- 
ed are worth repeating: "Var, caracasana Briq, ■ V. caracasana 
Kunth in Piumb, et Bonpl. Nov. gen, et sp, II, 275 (ann. 1817) - 
V. litoralis (sphalm. littoralis ) var. leptostachya Schauer in DC 
Prodr. XI, 5U2 (ann, I81i7). Mbocaiati, pres de Villa-Rica, champs 
en friche, ffivrier 1876, fleurs d'un bleu pftle (n. 1027a); plaine 
de Pirayu-Bi, 11 f6vrier 1876 (n. 1027). Nous avons ritabli les 
ncfflis primitivement adopt^s pour ces deux formes par Kunth et ar- 
bitrairement changes par Schauer souvent trop peu scrupuleuz dans 
les questions de nomenclat\ire." 

The Kuntze 2109 , distributed as V. caracasana , seems to be 
typical V. litoralis H,B,K. On the other hand, it seems very 
possible to me that many of the specimens cited under typical V. 
litoralis which have unusually dense spikes — as, for instance, 
most of those from the Hawsdian Islands — may represent var, 
caracasana . 

Citations: COLOMBIA: El Cauca: P^rez Arbeldez & Cuatrecasas 
g98l (W— 177li2l5) . ARGENTINA: Mendoza: Caret te 3Q$0 (N), 

VERBENA LITORALIS var, MELANOPOTAMICA Hauman-Merck, Anal, Mus. 
Argent. Hist, Nat, Buenos Aires 2U: iil3 [as " littoralis "] , 
1913. 

Bibliography: Hauman-Merck, Anal, Mus. Argent. Hist, Nat, 
Buenos Aires 21;: Ul3 — hlh' 1913; Moldenke, Known Geogr. Distrib. 
Verbenac, [ed. 1] , 14; & 101 (19U2) and [ed, 2], 106 & 198, 19li9j 
Moldenke, R6sum6 127 & U72. 1959; Moldenke, R6sum6 Suppl. 3: UO. 
1962. 

This variety differs from the typical forai of the species in 
its conspicuous corolla, ^riiich is hypocrateriform, pale-blue, 
3 — U mm. long, with the limb equally as long as the conic tube. 

The type of the variety was collected by Lucien Hauman-Merck 
( no. 3U5) under trees on the shores of the Rio Negro and cei*tain 
of its islands, Rio Negro, Argentina, where he says that it is 
very abundant. He states that the variety may be recognized on 
sight by its pale-blue flowers which are larger than those of 
sill otter species and forms of this group (leading him to the 
question "An species distincta?") . He describes the plant as 
glabrous, hardly scabrous, with short spikes which are very thin, 
and the corolla-tube hardly longer than the calyx. He is of the 



196Ii Moldenke, Monograph of Verbena 79 

opinion that this plant is very closely related to V. litoralls , 
from Tihich it differs on sight by the size and fonn of its flow- 
ers, by the larger and subcordiform bractlets, and by the enlarg- 
ed conic corolla-tube. 

The author's s\imame is sometimes misspelled "Haumann-Merck" 
in literature, 

VERBENA LOBATA Veil., Fl. Flum. 18 (1825), Icon. 1: pi. k3* 1827. 

SynonyiBy: Verbena buchnera Veil., Fl. Flum. 17 (1825), Icon, 
l! pi. U2. 1827. Verbena corymbosa Cham., Linnaea 7: 255. 1832 
[not V. corymbosa Hort., I81i5» nor R\ilz & Pav,, 1798]. Verbena 
lobatA Arrab. ex Steud., Norn. Bot., ed. 2, 2j 750. l8Ul. Verbena 
corymbosa Relh. ex Moldenke, Alph. List Invalid Names Suppl. 1: 
23, in syn. I9li7. Verbena lovata Veil, ex Moldenke, Alph. list 
Cit. 2: 367, sphaljn. 19U8. 

Bibliography: Veil., Fl. Flum. 17—18 (1825), Icon. pi. Ii2 & 
U3. 1827; Cham., Linnaea 7: 255. 1832; Steud., Nan. Bot., ed. 2, 
2: 750. I81a; Walp., Repert. Bot. Syst. h: 27. 18U5; Schau. in 
A. DC., Prodr. 11: 5U0. 18U7; Schau. in Mart., FL. Bras. 9: 18U-- 
185. 1851; Hook. f. & Jacks., Ind. Kew. 2; 1178 & 1179. 1895; 
Stapf, Ind. Lond. 6: U29 & U30. 1931; Moldenke, Prelijn. Alph. 
List Invalid Names 56. I9h0} Moldenke, Alph. List Invalid Names 
U6. 19U2; Moldenke, Known Geogr. Distrib. Verbenac., [ed. 1], 39, 
Ui, & 101. 19U2; Sampaio & Peckolt, Arquiv. Mus. Nac. Rio Jan, 
37: 392 & 393. 19U3; Moldenke, Alph. List Cit. 1: 21, 255, & 290. 
I9U6; Moldenke, Alph. List Invalid Names Suppl. 1: 23. I9u7; 
Moldenke, Castanea 13: 117. 19li8: Moldenke, Alph. List Cit. 2: 
362, 36U, 366, 367, 369, 375, hhh, U48, $3h, 535, 621, & 621* 
(19U8), 3: 670, 696, 751, 772, 82I1, 8U0, 862, 921, & 922 (19li9), 
and U: 98U, 1075, IO8I, IO9U, 121*8, 121*9, 1251, & 1287. 19li9; 
Moldenke, Known Geogr. Distrib. Verbenac., [ed. 2], 9li. 106. & 
198. 19li9; Moldenke, Phytologia 3: 75 (19U9) and 3: 288 & 289. 
1950 ; Stellfeld, Trib. Farmac. 19 (10): I66. 1951; Rambo, Sel- 
lowia 6: 153. 1951*; Biol. Abstr. 27: 3735. 1955; Rambo, Sellowia 
7: 260. 1956; Angely, Fl. Paran. 7: 13. 1957; Moldenke, R6sum6 
no, 119, 127, 360, 362, & U72. 1959; Moldenke, R6sum6 Suppl. 2: 
10—12. I960; Angely, Fl, Paran, 16: 78 (I960) and 17: U6, I96I; 
Reitz, Sellowia 13 (13): HO, I96I; Moldenke, Phytologia 8: 2U7 
(1962) and 9: IiO, U*, & 292. I963. 

IllTistrations: Veil., Fl. Flimi. Icon. 1: pi. U2 & U3. 1827. 

Low-growing rather weak herb, 0.5 — 1.3 m. tall, more or less 
hispid throughout; rhizome creeping; stems slender, tetragonal, 
rooting at the lower nodes, ascending at the tips, brachiate; 
branches often elongate, ascending, very wide-spreading, florif- 
erous and ascending at the apex; leaves decussate-opposite; pet- 
ioles about 1 cm. long; leaf -blades membranous, triangular- 
ovate, to about U cm. long and 2.5 cm. wide, below the middle 
rather deeply subtrilobed with 2 lateral incisions, acute at the 
apex, truncate or subcordate at the base and cuneately attenuate 
into the petiole, rugose, more or less strigose-hispid or 



80 PHTTOLOGIA Vol. 10, no, 1 

-hirtous and subcanescent above, the margins coarsely and irregu- 
larly incised-serrate, the teeth oTate, mucronate-acviminatej 
spikes very short, temate, fonning a iride-sp reading panicle, 
compact and congested during anthesis, divaricate-cymose and some- 
what patent in fruit, but even then scarcely longer than 1 ca.; 
pedtincles filifom, U — 5 cm. long; flowers imbricate, short- 
pedicellate; bractlets ovate, about half as long as the calyx, 
acuminate at the apex, ciliate along the margins; calyx about U 
mm. long, pilose-hlrtous or -hirtellous on the outer surface, e- 
glandular, its rim short-dentate, the teeth ovate and acuminate; 
corolla small, varying from light- violet or violet to blue-lilac 
or purple (also described as "roxa" and "anil"), its tube infun- 
dibular, about one-fourth longer than the calyx, the throat vil- 
lous; fruit about half as long as the calyx; cocci striate on the 
dorsal surface, subrugose-reticulate on the upper portion. 

The type of this curious species was collected hy Jos6 Mariano 
da Conceicao Velloso probably at or near Rio de Janeiro, Brazil, 
but is now lost. Schauer (1851) says: "Species cmni nota perin- 
signis, V. corymbosae R, et P. cui adnumeravit f Chamisso in 
Coll. pi. Sellow., sane qaadammodo af finis, sed ab ea caule pan- 
iculaque brachiatis, foliis petiolatis, floribus minoribus aliis- 
que notis uberrime distincta." Walpers (18U$) erroneously re- 
duces it to synonymy under V» corymbosa Ruiz & Pav, 

Leite refers to this plant on a herbarium label as "frutex", 
but as far as I am able to ascertain it is caapletely heAaceous. 
Irwin describes it as an "occasional" herb creeping beneath low 
shrubs and among rocks. It has also been collected in shrubby 
moist fields and grassy campos, in wet sunny places in riverbank 
thickets, at the edges of marshes, in hedges, at the margins of 
small woods, and at the edge of primeval forests, as well as in 
wet thickets, at altitudes of 60 to 3130 meters, flowering from 
August to May. Smith & Klein describe it as ruderal. Rambo 
found it growing in a region of 2 — 2.5 meters rainfall and — 25** 
C. temperature. Dus6n 313 shows extra large leaves, ifcile Mttller 
92 exhibits very large (or especially well pressed?) corollas. 
"^ Herbarium material of this species has been misidentified and 
distributed as V. coiymbosa Ruiz & Pav., V. megapotanica Spreng., 
and V. strigosa Cham. On the other hand, the Weir 319 , distribu- 
ted as " Verbena sp. near V. lobata Veil.", is V. hirta var. grac- 
ilis Dus6n. 

The type of V^ corymbosa Relh. is Sellow s.n. [Brasilia] de- 
posited in the herbarium of the Jardin Botanique de I'Etat at 
Brussels, Verbena corymbosa Hort. is a synonym of V. bonariensis 
L., while V. cosymiaosa Ruiz & Pav. is a valid species, which, as 
Schauer points out, somewhat resembles this one. Stellfeld (1951) 
cites Herb. Mus. Par an. 150U &. 3053 from ParanA, Brazil, not as 
yet seen by me. 

In all, 126 herbarium specimens and 3 mounted photographs or 
illustrations have been examined by me. 

Citations: BRAZIL: Espirito Santo: Irwin 2783 (Au— 173707, N, 



196U Moldenke, Monograph of Verbena 81 

W— 2281319)} A. Lutz 1159 (Z). Mlnas Geraia: Brade lii672 [Herb. 
Jard, Bot. Rio Jan. 26211] (B); Campos Porto llU7 [Herb. Jard. 
Bot. Rio Jan, 22$65] (N)j Collector vindesignated 3.n« [Serra do 
Piefd, Dec. 10, 1886] (Ja— U6596); Dus6n 2U2 (S), 313 , in part 
(W--U99U07), a.n. [Serra do Itatiaia, 17/5/1902] (S)} Lobo 8.n, 
[Planalto do Capara6, Nov. 1922] (Ja — U6521); Schwacke a.n. [Cam- 
pos de Capara6, 9/II/I89O] (Ja — U6586) . Parani; Ciirial B.n. 
[Hatschbach U03] (N); Dusfe 2660 (Ja--U6801, N, S), 6821 (Ca— 
501689, Lu, N, S, W— Ili8l773) , 8771 (S), 9287 (S), s.n. [CubalSa, 
27.12.1911] (S); Hatschbach 337 (N), 1622 (N), 2873 (N), Ul56 
(Ok), U308 (Ita); Mattos & Moreira s.n. [11/1959] (Ih--5909); Tam- 
andar6 107 (Mp — lll;0); Tessmann 3360 [Herb. Mus. Paran. 3360] (N), 
s.n. [Herb. Mus. Paran. 3033] (N) . Rio de Janeiro: Brade 16778 
[Herb. Jard. Bot. Rio Jan. lSxQ72] (N); Dus^n 2U2 (Ja— U6562, Sp— 
20063), 313, in part {Ja—lUQUh)', Biygdio ^ (Ja— 387IO, N), 107 
(Ja— 38709, N)} Herb. Mus. Paulista U98 (N, Sp— 15722); Moldenke 
& Moldenke I96I6 (Mg, Mr, N, No, Ot, Pn, S, Sm); Segadas-Vianna 
670 (Ja), 2908 (Ja), 5035 [Herb. Brade 20319] (Ja)j Ule k3Ul (Ja- 
U6528). Rio Grande do Sul: BomntOler 602 (Ut— U7123)j Jttrgena 
U69 (B, W— lli82202)} Leite Jig (N); Rambo 2293 (N), Ui82 (Sp— 
50986), 35193 (Lg, N, S), 36U18 (S), 511462 (N, W— 21020U2), 5l505 
(N), 51912 (N), 5208U (N, S, W— 2102316); Sehnem 3ii97 (Gg— 
356200). Santa Catarina: Fritz MtOler 92 (Ja— 31555), 97 (Ja~ 
35006)} Rambo U957O (S), U9605 (N, S), 60070 (S)} Reitz 2170 (N), 
5U12 (N), C.I215 (N)j Reitz & Klein 5l50a (W— 2252077), "^ITT (Ok, 
W— 226896U), 5226 (Ok, W— 226898U), 53U2 (W— 2252081), 7691 
[Herb. Barb. Rodr. 22666] (Ito, N, S), 77U5 (Mm), 8OI8 (Mm); Smith 
& Klein 79U2 (Ok), 10508 (N, Ok, W— 2251688); Smith , Reitz , & 
Klein 79li2 (W— 2251328) . SSo Paulo: Bailey & Bailey 86I (Ba,~Ba)} 
Campos Porto 2982 , in part [Herb, Jard. Bot. Rio Jan. 3260U] (N), 
298I; [Herb. Jard. Bot. Rio Jan. 32606] (B, N); Frazfio s.n. [Herb. 
Jard. Bot. Rio Jan. 16528] (N) ; Hammar s.n. [Horto BotSnico, Ser- 
ra da Cantareira, Sept, 30, 1901; Herb, Conm, Geogr, & Geol. 58U8] 
(N, Sp— 15717)} C, A. Krug s.n. [Herb. Inst. Agron. Est. S. Paulo 
399U] (N, 7(~ni^0W)i M. Kuhlmann s.n. [Umtiarama, Jan, 26, 1935] 
(K, Sp— 32387, Sp); Leite 3U28 "(El), Iv (N)} LOfgren s,n. [S«o 
Francisco dos Campos, Dec, 21, I896} Herb, Comm. Geogr, & Geol, 
3U96] (N, Sp— 15726), s,n. [Altotiet^, Oct. 19, 1901} Herb. Coom. 
Geogr, & Geol. 58U7] (N, Sp — 15712)} Tamandar6 & Brade s.n. [Ser- 
ra da Cantareira, Jan. 11, I9IU; Herb. Brade 669I] (N, Sp— 6722); 
Usteri s.n. [Cantareira, Sept. 2U, 1905] (Sp— 15731); ViSgas, 
Franco , & Lima s.n. [Ubatuba, March 9, I9UO; Herb. Inst. Agron. 
Est. S. Paulo 5U22] (N, Sp — Ui299) « State undetermined: Sellow s_. 
n, [Brasilia; Macbride photos 3U3li8] (Br, Kr — photo, N — photo). 



82 PHTTOLOGIA Vol. 10, no. 1 

DRUGUAI: Herter 50885 (N) . ARGENTINA,: Misiones: Nlederleln s.n. 
(Ra— 23600). Salta: D. Rodriguez |5 [Herb. Inst. Miguel Lillo 
3156U] (N). MOUNTED ILLUSTRATIONS: Pohl, Icon. Plant. Brasil. 329 
(V). 

VERBENA LOBATA var. GLABRATA Moldenke, Phytologia 3: 118. 19li9. 

Bibliographer: Moldenke, Known Geogr. Dlstrib. Verbenac, [ed. 
2], 9li & 198. 19U9; Moldenke, Phjrtologia 3: 118 & 13U (19U9), 3: 
289 (1950), and 3: U5U. 1951; Moldenke, R6sum6 110, II9, & U72. 
1959. 

This variety differs frcaa the typical fom of the species in 
being completely glabrous throughout. 

The type of the variety was collected by Padre Balduino Rambo 
( no. 2816 ) at SSo Francisco de Paulo, Rio Gramde do Sul, Brazil, 
on January Ik, 1937, and is deposited in the Britton Herbarium at 
the New York Botanical Garden. The plant is said to inhabit 
thickets, stream margins, and river banks, flowering from Decem- 
ber to February. In all, U herbarium specimens, including the 
type, and U mounted photographs have been examined by me. 

Citations J BRAZIL: Rio de Janeiro: Ule 61a (Ja — 146527) . Rio 
Grande do Sul: Rambo 2816 (F — photo of type, N — type, N — photo of 
type, Rb—isotype, Sg — photo of type, Z — photo of type) , URUGUAI: 
Arechavaleta 28 (Ug). 

VERBENA LDBATA var. HIRSUTA Moldenke, Phytologia 2: U23--U2U. 
19U8. 

Bibliography: Moldenke, Phytologia 2: U23--U2U. I9I48; Moldeilce, 
Castanea 13: 117. l9U8j Moldenke, Known Geogr. Distrib. Verbenac, 
[ed. 2], 9U & 198. 19li9j Moldenke Alph. Ust Cit. 3: 70U & 8U7. 
191^9; Reitz, Sellowia 11; 57 & 13U. 1959} Moldenke, RSsumfi 110 & 
k72. 1959; Moldenke, R^sumS Suppl. 3: 111. 1962. 

This variety differs from the typical fonn of the species in 
having the branches, branchlets, and twigs, as well as the peti- 
oles, lower leaf-srurfaces, peduncles, bractlets, and calyxes very 
densely hirsute with widely spreading white or flavescent hairs. 
The upper leaf-surface is also more hirsute than in the typical 
fom. 

The type of the variety was collected by Gustaf Oskar Anders- 
son Malme ( no. 1260 ) in the grassy edges of a marsh at Pinhal, 
near Santa Maria, Rio Grande do Sul, Brazil, on Januaiy 27, 1902, 
and is deposited in the herbarium of the Naturhistoriska Rikamus- 
eum at Stockholm. The plant is described by collectors as sub- 
prostrate, with blue or violet corollas. It has been found in 
fields and thickets, hedgerows and dry grassy places^ at the 
edges of rivers, and on campos, from 8OO to 9OO meters altitude, 
flowering from November to February. Reitz records the vernacular 
names "camaradinha", "formosa sem dote", and "jurupeba" for this 
and all other members of the genxis in that area. Herbarium mater- 
ial has been misidentified and distributed as V. megapotamica 
Spreng . 



1961i Moldenke, Monograph of Verbena 83 

Ir^ all, 26 herbariiM specimens, including the type, and U 
mounted photographs have been examined by me. 

Citations: BRA.ZIL: Minas Gerais: A. Lutz 393 (Lz)j Regnell 
III .1619 [28/12A868] (N, S, S) . Rio Grande do Sul: Malme 1260 
(F — photo of type, N — isotype, N — photo of type, S — "type, Si — 
photo of type, Z— photo of type); Rambo 32302 (N), 38U2li (N), 

51U05 (s), giii62 (s), $1505 (s), 51912 (s), 519U9 (n, n, s, w— 

2102255), 53791 (S); Sehnem 3U97 (N), 3776 (B) . Santa Catarinai 
Reitz & Klein 5186 (N, Ok, W~2268971); Smith & Klein 8l05 (W— 
22513U5); Smith & Reitz 8105 (Z) . PARAGUAY: Fiebrig 588U (Bm, ¥~ 
1159390) . 

VHiBENA LOBATA var. SESSILIS Moldenke, Phytologia U: 267 & 293. 
1953. 

Bibliographer: Moldenke, Ptytologia U: 267 & 293. 1953; Molden- 
ke, Biol. Abstr. 27: 3121. 1953; Angely, Fl. Paran. 7: 13. 1957; 
Moldenke, R5sumS 110 & U72. 1959; Angely, Fl. Paran. 16: 78 
(i960) and 17: hS. I96I; Moldenke, Phytologia 9: 289. 1963. 

This variety differs from the typical form of the species in 
having its leaves sessile, the blades 1 — 2 cm. long and 5 — 13 nan. 
wide. The corolla is described as purple. 

The type of the variety was collected by Gert Hatschbach ( no. 
2876 ) at Varzea, S5o Jos^ dos Pinhaes, Parand, Brazil, on Decem- 
ber 2, 1952, and is deposited in the Britton Herbarium at the New 
York Botanical Garden. The plant has been found in bogs and pas- 
tures, on campos, and in ruderal situations, at altitudes of 750 
to 1650 meters, flowering in December and January. Smith & Reitz 
8916 is a mixture with V. hirta Spreng, 

In all, 6 herbarium specimens, including the type, have been 
examined by me. 

Citations: BRAZIL: ParanS: Hatschbach 2876 (N — type). Santa 
Catarina: Smith & Reitz 8696 (W— 22li9356), 89I6 (W— 225m66, Z), 
1031a (Ok, W— 22^1672). 

VERBENA LONGIFOLIA Mart. & Gal., Bull. Acad. Brux. 11 (2): 323. 
l8Ui [not V. longifolia Lam., 1873]. 

Synonymy: Verbena Carolina var, glabra Hultfen ex Moldenke, R6- 
sum6 361, in syn. 1959. 

Bibliography: Mart, & Gal., Bull. Acad. Brux. 11 (2): 323. 
l8Uli; Schau. in A. DC., Prodr. 11: 555. 18U7; Walp., Repert. Bot. 
Syst. 6: 687. 18U7; Hook. f. & Jacks., Ind. Kew. 2: 1179. 1895; 
Ed. Rodigas, Bull. Arboricult. Belg. 1902: llli. 1902; Perry, Ann. 
Mo. Bot. Gard. 20: 21^7, 259, 271, 272, & 355. 1933; Moldenke, 
Known Geogr. Distrib, Verbenac, [ed. 1], I9 & 101. 19li2j Moldenke, 
Phytologia 2: 331. 19U7; Moldenke, Alph. Ust Cit. 3: 919 (191^9) 
and h: II6I, 1295, & 1303. 19U9; Moldenke, Known Geogr. Distrib. 
Verbenac, [ed. 2J, 33 & 198. 19^9; Moldenke, Inform. Mold. Set 51 
Spec. h. 1956; Moldenke, R6sum6 39, 361, & U72. 1959; Moldenke, 
R«8um6 Suppl. 3: 10. 1962; Moldenke, Phytologia 8: U27 (1962) and 



8U PHTTOLOGIA Vol. 10, no. 1 

8 J U87, U88, & U92. 1963 j Moldenke, R6s\m6 Suppl. 6: U. 1963. 

Stems erect, to 1.5 m. tall, obtusely U-angled, glabrous; 
branches many, ascending; leaves decussate-opposite, short-petio- 
late; leaf-blades lanceolate to elongate-elliptic, 10—12.5 cm. 
long or the upper ones somei^at smaller, acutely serrate from be- 
low the middle to the apex, appressed-pubescent or very short- 
strigillose on both surfaces, the venation prominent beneath; 
spikes panicTilately disposed, slender, elongate, glabrous, open 
nhen in fruit; bractlets ovate, about half as long as the calyx, 
acute to acuminate at the apex, cilia te along the margins; calyx 
about 2 mm. long, practically glabrous; corolla pale-lavender or 
lilac, its tube scarcely protruding beyond the calyx, the limb 
inconspicuo\is ; fruiting-calyx with its obtuse lobes connivent o- 
ver the schizocairp; cocci trigonous, hardly 1.5 mm. long, smooth 
or faintly striate, the ccmuissural faces smooth. 

The type of this species was collected by Henri Guillaume Gal- 
eotti ( no. 791 ) in fields at Ario, Michoacfin, Mexico, at an alti- 
tude of UOOO feet, and is deposited in the herbarium of the Jai>- 
din Botanique de I'Etat at Brussels. The type of V. Carolina var. 
glabra was gathered by Harald August FrOderstrOm and Elsa Hult&i 
( no. 82) at an altitude of 1700 meters in Morelos, Mexico, and is 
deposited in the herbarium of the Naturhistoriska Riksmuseum at 
Stockholm . 

The species inhabits roadsides, brush, semixsric areas, low 
and low spiny matorral, the edges of arroyos, dry spots in gener- 
al, and heavy red clay loam on steep slopes and hillsides in 
moist pine or fir forests, at altitudes of 700 to 2050 meters, 
and has been collected in flower and fruit in January, March, A- 
pidl, and Atigust to November. McVaugh refers to it as an "abun- 
dant herb", Seler & Seler UI9I4 has galled spikes. 

Herbarium material of this species has been misidentified and 
distributed xinder the names V. litoralis Kunth, V. littoralis H. 
B.K., and V. littoralis var. b. leptostachya Schau. The Gonzalez 
Ortega 60 & 226 collections, cited below, are labeled "Elota, La 
Cruz, El Roble" and apparently coiild have cone from either Nay- 
arit or Sinaloa . Seler & Seler Ul9li was originally distributed 
as " Verbena caroliniana L. form, vel var. polystachya (Kunth) 
Loes." and 1;3U7 as "Verbena spec. aff. V. carolinianae L.", 
while of C. R. Orcutt 1371 Perry says "Aff. V. menthaefolia Bth." 

Martens & Galeotti (l8Ui) affirm that the species is related 
to V. paniculata Lam. f-V. hastata L.], irtiile Schauer (18U7) 
points out that its leaves are similar to those seen on Veronica 
longifolia L. ( Scrophulariaceae ) , with the inflorescence of Ver - 
bena urticifolia L. Perry (1933) describes it as "A rather sin- 
gular species combining the foliar characters of V. litoralis 
with the inflorescence characters of V. Carolina ." She also says 
that Nelson 752, which she considers to be V. Carolina L., "is 
almost glabrous, and the lobes of the corolla are emarginate, a 



196U Uoldenke, Monograph of Verbena 65 

rather xmusual featxire in this series. The lack of pubescence 
siiggests V. longifolia , but unfoirtxinately none of the available 
material of the species is in sufficiently good condition to re- 
veal the character of the corolla." She cites the following 9 
additional specimens not as yet seen by me: MEUCO: Morelos: Se- 
ler & Seler hl9h (G) , U3U7 (G) . Oaiaca: C^ R. Orcutt 3321 (E) . 
Puebla: Purpus 3U06 (E, F, G, N) . Vera Cruz: Seler & Seler 72U 
(G) . See under V. Carolina in these notes for her key for dis- 
tinguishing this species from its nearest relatives. The V. 
longifolia Lam. referred to in the synonymy above, is a synonym 
of V, canadensis (L.) Britton. 

In all, 27 herbarium specimens, including the type collections 
of both names involved, and h mounted photographs have been ex- 
amined by me. 

Citations: MEXICO: Coahuila: Agiiirre & Reko 132 (N) . Mexico: 
Matuda 30538 (Ss), 31307 (Ss) . MichoacSn: QaleotU 791 (Bi^-type, 
F — photo of type, N — photo of type. Si — photo of type, Z— photo 
of type). Jalisco: R. McVaugh 13077 (Mi), lli06l (Mi), 16171 (Mi). 
Morelos: FrOderstrOn & Halt6n 82 (S), U28 (S); Moldenke & Molden- 
ke 19827 (Es, Lg, N); Pipes 113 (Z); J. H. Hill kh (Mi); Seler & 
Seler la9U (W— 1205687), U3U7 (W— 1205719). Nayarit: J. Gonzalez 
Ortega 60 (Me), 226 (Me). Oaxaca: C^ R. Orcutt 3321 (V— 567312). 
Puebla: Porpus 3UO6 (N, W— 8U1139) . Sinaloa: J_. Gonzalez Ortega 
5U78 (Me). Sonora: C. R. Orcutt 1371 (W— II6806U) . Vera Cruz: 
Liebmann 11 318 (W— 1315092) j Seler & Seler 72h (Me, Me, W— 
1323102) . 

VERBENA LONGIFOLIA f . ALBIFLORA Moldenke, Phytologia 7: U30. I96I. 

Bibliography: Moldenke, Phytologia 7s U30. I96I; Moldenke, R6- 
8um6 Suppl. 3: 10. 1962. 

This fonn differs from the typical form of the species in hav- 
ing irhite corollas. 

The type of the form iras collected by Boone Hallberg ( no. 813) 
on moist clay banks of a drainage area from cleared cornfield a- 
mong Persea cloudf orests , on the east slopes near Patio de Arena, 
about 5 km. east of the summit, at about 2900 meters altitude, in 
the vicinity of CeiTO Zampoaltepetl, Oaxaca, Mexico, on August 7, 
1950, and is deposited in the herbarium of the University of 
Michigan at Ann Arbor. The plant is described as a perennial. 
It is known thus far only from the type specimen. 

Citations: MEUCO: Oaxaca: B. Hallberg 8I3 (Mi — type), 

VERBENA LUCANEIBIS Moldenke, Phytologia 3: 279—280. 1950. 

Bibliography: Moldenke, Phytologia 3: 279—280 & 286. 1950 j E. 
J. Salisb., Ind. Kew. Suppl. 11: 263. 1953; Moldenke, Infoim. 
Mold. Set 51 Spec. U. 1956j Moldenke, R6sum6 85 & U72. 1959. 

Herb, 30 — 50 cm. tallj stems and branches erect or ascending, 
slender, rather obtusely tetragonal, densely short-pubescent with 



86 PHYTOLOGIA Vol. 10, no. 1 

■whitish spreading hairs; nodes not annulate; principal intemodea 
0.5 — U.2 can. long; leaves decussate-opposite, visually with a 
dense cluster of small ones on abbreviated twigs in their axils; 
petioles obsolete; leaf-blades chartaceous, rather uniformly 
bright-green on both surfaces, the immature ones more or less 
brunnescent in drying, ovate in outline, 1 — U cm. long and wide, 
deeply 3-parted, the divisions again deeply and rather irregvilar- 
ly parted or dissected, the lamina-segments 0,5 — 2.5 Jom. wide, 
rather densely pubeinilent on both surfaces, obtuse or acute at 
the apex, subrevolute along the margins, the single vein in each 
segment impressed above, prominulous beneath; inflorescence term- 
inal and in the upper leaf -axils, short-spicate or subcapitate, 
to 3.5 cm. long, densely many-flowered; peduncles very slender, 
Ix — 15 mm. long, densely spreading-pubescent like the branches; 
bractlets lanceolate, about 3 mm, long and 1 mm. wide, gradually 
attenuate to the apex, densely puberulent; calyx tubular, 3 — h nmi. 
long, densely puberulent, its rim 5-toothed, the teeth narrow- 
attenuate; corolla hypocrateriform, varying from blue to violet 
or purple, its tube 6-- 7 mm. long, very lightly puberuloxis on the 
outside toward the apex, its limb about 6 mm. in diameter; fruit- 
ing-calyx not inflated, e asily splitting into 5 similar segments; 
cocci k, oblong, about 2 mm. long, glabrous, shiny, the dorsal 
surface vmiformly scrobiculate-ridged, the commissural surface 
white-papillose for the lower two-thirds only. 

The type of this distinct species was collected by Rara6n Fer- 
reyra ( no. 5U93) in stony habitats, at 1500 to 2000 meters alti- 
tude, between Nazca and Puquio, province of Lucanas, Ayacucho, 
Peru, on March 19, 19U9, and is deposited in the Britton Herbar- 
ivm at the New York Botanical Garden. The species has been found 
in the Baccharis and Loxopterygiimi zones, at altitudes of 2000 to 
3600 meters, blooming in March, May, and August. 

In all, 6 herbarium specimens, including the type, have been 
examined by me. 

Citations: PERU: Ayacucho: R, Ferreyra 5U93 (N — type, Ug — iso- 
type); Ravih & Hirsch P.l|07 (Hk). Cajamarca: R. Ferreyra 8U82 
(Ss). Huancavelica: Rauh & Hirsch P. 38 3 (Z). La Libertad: R^^ Fer- 
reyra 3028 (Ss). 

VERBENA MACDOUGALII Heller, Bull. Torrey Bot. Club 26: 588, 1899. 

Synonymy: Verbena macdougali Heller ex Koldenke, Suppl. List 
Invalid Names 9, in syn, 19lil, Verbena mcdougallii Heller ex 
Moldenke, Suppl. List Invalid Names 9, in syn. I9UI. Verbena 
mcdougalii Heller ex Moldenke, Resum^ 369, in syn. 1959. Verbena 
macdougali x hastata Lee ex Moldenke, RSsum6 Suppl. 3: UO, in 
syn. 1962. Verbena macdougalii Wooton ex Moidenlce, R6sum^ Suppl. 
3: UO, in syn. I962, Verbena macdouglii Heller ex Moldenke, R&- 
B\m& Suppl, 3: UO, in syn, I962, Verbena macdougali Wooton ex 
Moldenke, R6sum^ Suppl. 6; 11, in syn. I963. 

Bibliography: Heller, B\ai, Torrey Bot. Club 26: 588. 1899; 
Cockerell, Am. Nat, 36: 809. 1902; Thiselt .-Eyer, Ind, Kew. Suppl, 



196U Moldenke, Monograph of Vertena 8? 

2: 191. 190lii Tidestr., Contrib. U. S. Nat. Herb. 25: Ii69. 1925; 
A. B. Seymour, Host Ind. Fungi N. Am. 587. I929j Perry, Ann. Mo. 
Bot. Gard. 20: 260, 288—290, & 355. 1933; Moldenke, Prelim. 
Alph. List Invalid Names U7. 19U0; Moldenke, Suppl, List Invalid 
Names 9. I9UI; Wyman & Harris, Navajo Ind. Ethno-Bot. 32 & U5. 
I9UI; Moldenke, Known Geogr. Distrib. Verbenac,, [ed. 1], 10. 11, 
13 m, 7U, & 101. 19U2; Moldenke, Alph. List Invalid Names U8. 
19U2; Moldenke in Lundell, Fl. Texas 3 (1): 16 & 26. 19U2; Mol- 
denke, Am. Joum. Bot. 32: 610. 19U5; G. L. Fisher, Am. Bot. Ex- 
change List. I9U6; Moldenke, Alph. List Git. 1: Ik, 2U, 102. 126, 
182, 191, 203, 2U5, 2U6, 255, 256, 258, 265, 281, & 283. 19U6; 
Curtin, Healing Herbs Upper Rio Grande 75 & 272. 19 U7; Moldenke, 
Alph. List Git. 2: 392, 393, U38, U52, U5U, h$^, U68, U71, U72, 
U71+, U76, U77, U80, U82, U83, U89. Ii9l, 192, 506, 519, 521, 532, 
538, 539, 595, 597, 60U, 618, & 6fi0. I9l;8; Moldenke, phQrbologia 
2: 163. I9U8; H. N. & A. L. Moldenke, Pi. Life 2: 71. 19U8; Mol- 
denke, Wrightia 1: 225. 19U8; Moldenke, Castanea 13: 112 & 113. 
I9U8; Moldenke, Alph. Ust Git. 3: 681i, 697, 7U0, 7U7, 75U, 779, 
782, 831, 833, 839, 8Ul, 853, 857, 883, 88U, 890, 899, 90li, 91I4, 
952, Sc 966 (I9U9) and U: 987, IO86, 1122, II38, lllA, llli2, 
U50, 1163, 1165, 1167, 1173—1175, 1207, 1225, 1228—1230, 
1237, 12li5, 12U6, 1252, 1253, & 1289—1291. 19U9; Moldenke, 
Known Geogr. Distrib. Verbenac, [ed. 2], 19, 20, 2U — 26, 16U, & 
198. 19li9; H. N. &A. L. Moldenke, Anal. Inst. Biol. Mex. 20: Jh» 
I9U9; Moldenke in Chittenden, Roy. Hort. Soc. Diet. Gard, U: 
2209 & 2211. 1951 ; Moldenke, R^sum^ 2U, 25, 29, 31, 32- 223, 369, 
& U72. 1959; Lewis & Oliv., Am. Journ. Bot. U8: 639--6ul. 1961: 
Moldenke, R^sumS Suppl. 3: 8 & UO. 1962; Moldenke, Phytologia 8: 
12U (I96I) and 8: 213 & U35. 1962; Moldenke, R6sum6 Suppl. 5: 3, 
Ii, & 7 (1962), 6: 11 (1963), and 7: 3. 1963; Moldenke, Phytologia 
8: U87 (I963) and 9: iWi. 1963. 

Illustrations: Lewis & Oliv., Am. Joum. Bot. U8: 6UO. I96I. 

Coarse perennial herb; stems stout, 0.3 — 1 m, tall, obtusely 
U-angled, simple or occasionally branched, cinereovis-green, 
whitish-hirsute or hirsute-pubescent; leaves decussate-opposite, 
short-petiolate or narrowed into a subpetiolar base, the blades 
oblong-elliptic or elongate-ovate, 6 — 10 cm. long, coarsely and 
irregularly serrate-dentate, hirtellous, rugose and minutely 
pustulate above, densely pilose-pubescent and prominently veined 
beneath; spikes solitary or sometimes several, short-pedunculate, 
thick, comparatively dense both during anthesis and in fniit; 
bractlets lanceolate-subulate, us\aally noticeably longer than 
the calyx, pubescent on the back, ciliate along the margins; 
calyx h — 5 mm. long, rather densely pubescent on the outside, 
glandular, its lobes very obtuse, terminating in short subulate 
taeth; corolla varying firom purple or pink-purple to dark-pui^jle 
or deej>-purple, the throat white and hairy, its tube about 5 mm. 
long, scarcely protruding beyond the calyx, the limb 3.5 — 6 mm. 
wide; cocci trigonous, about 2.5 mm. long, convex on the dorsal 
surface, raised-reticulate toward the distal end, strongly or 
faintly striate below, the commissural face reaching the tip of 



88 PHTTOLOGIA Vol. 10, no. 1 

the coccus, nuriculate or almost smooth; chromosome number: n - ?• 

The type of this distinctive and handsome species iras collect- 
ed by- Daniel Trembly MacDougal ( no. 2li9 ) — in whose honor it is 
named — in the vicinity of Flagstaff, Coconino Coxmty, Arizona, 
on July 8, 1888, and is deposited in the Britton Herbarium at the 
New Tork Botanical Garden. 

The species found on flats and open flats at high altitudes, 
as 1^11 as on dry slopes, lake shores, and prairies, along road- 
sides and weedy roadsides, in moist low places and open pinelands, 
in irrigated cultivated fields, in sandy and clay^ soil, along 
drying streams, in open and dry open woods, in yellow pine forests 
and cemyons, on river floodplains, and in volcanic soil, at alti- 
tudes of 2000 to 3165 meters, blooming and fruiting from June to 
October. It was introduced into cultivation in 1927. The spec- 
ific name is often variously upper-cased. 

Hanson found the plant "frequent in open places" and "in open 
pines", and Parker says "abundant on barren disturbed area around 
dump pile", but Schallert reports it as "not common". Waterfall 
describes it from "wet flat spots" and "pine woods in the moun- 
tains" in New Mexico, Weber calls it a "common roadside plant" in 
Colorado, and Demaree reports it "common" in rocky areas and "ccja- 
mon In shade of pines" in Arizona. Goddard c£ills it a plant of 
the Upper Transition Zone. 

This species is often infected by the fungus Erysiphe cichor- 
aceanmi DC. and Ophiobolus collapsus Sacc. & Ellis. Common and 
vernacular names recorded for it are "dormilon", "dormil6n", "New 
Mexican vervain", "sleepy-head", "verbena", "vervena", and the 
Navajo name "tAdfdl.n do.X'is ncS.g£". Renner says of it "cosmion, 
wide distribution, no uses, poor forage value, grazed from June 
to October". 

Herbarium material has been mis identified and distributed under 
the names V. bracteosa Michx,, V. canescens neo-mexicana Gray, V. 
canescens neomexicana Gray, V. canescens var. neo-mexicana Gray, 
V. polystachya H.B.K., V, stricta Vent., and "V. stricta var," 
On the other hand, the G. Martin s.n. [April 12, I96O], distribu- 
ted as V. macdougalii , is actually xV. perplexa Moldenke. Benson 
9573 is a mixture with V. bipinnatifida Nutt. Riordan 1 and H. 
E. Lee s.n. [9-19-36] have short bractletsj the latter collection 
was identified by the collector as a hybrid between this species 
and V. hastata L., which is a possibility because both species 
occur in the county iriaere it was found and members of this group 
are noted for their natural proclivity toward hybridization. 

Baldirin states of the flowers of V. macdougalii that the cor- 
olla is mostly a delicate puz*ple, but the throat Is white Kid 
hairy. Wooton s^n. [Mts. west of Grant's Station, Aug, 2, 1892] 
bears a note by the collector "Near Verbena stricta but differing 
from that plant in the size and shape of leaves, length of inter- 
nodes, &c. Dr. Rusby has also collected this plant in Arizona". 



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PHYTOLOGIA 

Designed to expedite botanical publication 



V'ol- 10 April, 1964 No. 2 



CONTENTS 

lOLDENKE, H. N., Materials toivard a monograph of the genus 

Verbena. X VIU gc) 

IcCLURE, F. A., A neglected Mexican species of Arundinaria 162 

lOLDENKE, A. L., Book reviews 164 



Published by Harold N. Moldenke and Alma L. Moldenke 

15 Glenbrook Avenue 
Yonkers 5, New York, U.S.A. 

Price of this number, $1; per volume, $5. 75, in advance 



MATERIALS TOTARD A MONOGRAPH OF THE GEtTOS VERBENA . XVIII 
Harold N. Moldenke 



As of the present irriting (March U, 19Sk) 36,95U herbarium 
specimens and 80li mounted photographs and other descriptive or il- 
lustrative material have been examined and annotated by me in the 
preparation of these preliminary monographic notes. 

VERBENA (Dorst.) L. 

Additional synonymy: Verbena Toum. ex Westm. in L., Philos. 
Bot. 150. 17$1. Uwarovia Bocq., Adansonia 2: 126. 1861—1862. 
Shuttlevorthia Bocq., Adansonia 2: 126. 1861 — 1862. Verbena Endl, 
ex Bocq,, Adansonia 3: I8I, 1862. Verbena [Toum.] L. apud Robin- 
son St Fern, in A. Gray, New Man. Bot., ed. 7, 688. I908. 

Additional & emended bibliography: F. Hernandez, Rer. M«dic. 
Nov. Hisp. 399. 1628; F. Hernandez, Nov. PI, 399. 1601; Dodart., 
Mem. Acad. Sci. Paris U: 317, pi. 317. 1669; Zanoni, Hist. Bot. 
203— 20I1 fig. 77. 1675} Barrelier, Plant. Gall. Hisp. 30, pi. 
853 tc lUi6, 1711ij Westm. in L., Orat. Tellur. Habit. Incr, 6U. 
17l4lii L., Hort. Dpsal, 8—9. 17U8; L,, Philos. Bot. 63, 66, 87, 
122, 150, & 17U. 1751 J Gesn. & Gamer, in Gesb., Op. Bot. 1: II6. 
I75I; Kniphof, Bot, Orig. Herb. Viv. cent. 2, pi. [28U]. 1757; 
Rtaing, Comm. Bot. U62. 1766; J. A. Murr. in L., Syst. Veg., ed. 
13, 61—62 & 8Ui. 177U; L'H^r., Stirp. Nov. 1: 21— 2U, pi. 11 4 
12. 1786; A. L. Juss., Gen. Pi., ed. 1, 109 (1789) and ed. 2, 122 
& 123. I79I: L. C. Rich, in Michx., Fl. Bor.-Am., ed. 1, 2: 1>- 
15 & 3U0. 1803; Dum. Cours., Bot. Cult., ed. 2, 2: 622—627. I8II; 
Pursh, Fl. Am. Sept. 2: Ia5--m7, 711, & 725. I8lli; Dum. Cours., 
Bot. Cult., ed. 2, 7: 131. I8IU; L. C. Rich, in Michx., Fl. Bor.- 
Am., ed. 2, "1" [-2]: 13—15 & 3U0. 1820; Lehm., Del. Sem. Hort. 
Hamb. 1826: 16. 1826; Hook., Bot. Misc. 1: 159—173, pi. U6— US. 
1829; Lehm., Del. Sem. Hort. Hamb. 1832: 7 (1832) and I83U: 7—8. 
18 3U; Benth. in Hook., Joum. Bot. 1: 59. I83ii; Steud., Nom. Bot., 
ed. 2, 1: 205, 23li, 58U. & 687 (I81i0) and 2: 5U, 201, 397, 575, 
629, 7U9— 751, & 797. 18U1; Engelm. & Gray, Pi. Lindheijn. 1: 21. 
18U5; Benth., Pi. Hartw. 245. I8U6; Regel, Gartenfl. U: 373, pi. 
Ili2, fig. 1. 18 55 J Planch. & Van Houtte, Fl. des Serres 11 [ser. 
2, 1]: pi. 1129. 1856; J. Torr. in Bnory, Rep. U. S. & Mex. Bound. 
Surv. 2: 128. 1858; Munby, Cat. Pi. Alg. 25. 1859; Bocq., Adan- 
sonia 2: 86, 105—107, 111 113, 115, 118, 123, 12li, 126, 128, 
130—133, 136, 139, litO 1U2, Ihh. ihS, li;7, 1U9, 152— 15U, 157, & 
158, pi. 11 (1861—1862) and 3: 180, I8I, & 201— 20li. 1862; Bocq., 
Rev. Verbenac. 26, 27, 30, 31, 33, 35, U3, Uk, US~U8, 50—53, 56, 
59, 60, 62, 6U, 66, 67, 69, 72, 73, 77, & 78. 1862; Bocq., Adan- 
sonia 2: 251. 1863; Griseb., Cat. PI. Cub. 211i, 1866; J. Ball, 
Joum. Linn. Soc. Lond. Bot. 16: 607. 1878; A, Gray, Syn. Fl. N, 
Am. 2 (1): 333 & 335—338. 1878, F. Phil., Cat. PI. Vase. Chil. 
218—219. 1881; A. Gray, Proc. Am. Acad. Sci. 19: 95. 1883; Stapf, 

89 



90 PHYTOLOGIA Vol. 10, no. 2 

Denkschr. Akad. Wiss. Wien ^0: 3U— 35. I885j Batt. & Trab., KL. 
Alg. 1: 717. 1888} F. D. Bergen, Joxirn. Am. Folk-Lore 5: 102. 
1392; Hitchc. & Norton, Kans. Agr. Exp. Sta. Bull. 50: Sh, pi. 6, 
fig. 91 & 92. 1895; "Lo Spigolatore", Bull. Soc. Toscana Ortic. 
22: 77, fig. Hi. 1897; Selby, Bull. Ohio Agric. Sta. 83: 329—330. 
1897; Solered., Bull. Herb. Boiss., s5r. 1, 6: 627 & 628. I898; 
A. S. Hitchc., Fl. Kans. pi. lU & 15. 1899; Wittmack, Gartenfl. 
U9: 585. I9OO; K. Schum. in Just, Bot. Jahresber. 28 (1): U97» 
1902; Ed. Rodigas, Bull. Arboricult. Belg. 1902: 113— Uli, fig. U. 
1902: Selby & Hicks, Bull. Ohio Agric. Sta. 11^2: 118, 121. & 123, 
pi. U. 1903; Selbj, Bull. Ohio Agric. Sta. 175: 350, pi. U. I9O6; 
J. W. Blankinship in Engelm. & Gray, PI. Lindheim. 3: 186 — 187. 
1907; L. H. Bailey, Botany 26 & 372, fig. 35. 1911; N. Taylor, 
Mem. N. T. Bot. Gard. 5: [Fl. Vic. N. Y.] 525 & 526. 1915; Shreve, 
Carnegie Inst. Wash. Publ. 217: 19, 2U, & UU. 1915; Gerth van 
Wijk, Diet. Plantnames 2: 228 317, U7o, 655, 893, 1U92, & l6lli. 
1916; Rydb., Fl. Rocky Mts. 7uO. 1917; Druce, Rep. Bot. Exch. 
Club Brit. Isles I9I8: 298. 1919; Urb, in Fedde, Repert. Beih. 5: 
73. 1920; Tidestr., Contrib. U. S. Nat. Herb. 25: 1a69. 1925; 
Druce, Rep. Bot. Exch. Club Brit. Isles 1927: U52. 1928; Baeza, 
Nomb. Vulg. PI. Silv. Chile, ed. 2, U9— 50, 60 — 61, 65, 8U, 100, 
206, 212, 232, 233, & 269. I93O; Druce, Rep. Bot. Exch. Club 
Brit. Isles 1932: 3U8. 1933; Anon., Joum, Hort. Soc. Lond. 61: 
liOl. 1935; H. S. Marshall, Kew Bull. 1936: 9U. 1936; K. V. 0. 
Dahlgren, Svensk. Bot. Tidsk. 32: 231. 1938; Moldenke, Saxiflora 
pi. 16. I9UO; Of fie. Org. Malayan Agri-Horticult . Assoc. Kuala 
Lumptir [MAHA Mag.] 10: 152. 19l;0; Cain, Found. Pi. Geogr. 335. 
19hk; C.N. Jones, Ohio Joum, Sci. hki 190. 19Ui; Deam, Kriebel, 
Yuncker, & Friesner, Proc. Ind. Acad. Sci. 55s 56. 19l;6; Parodi, 
Rev. Argent. Agr. lH: 61 — 69. 19U7; Martinez Crovetto & Piccin- 
ini, Revist. Investig. Agric, kt 178, I80, 181, 225, & 226 
(I950) and U: 32, 33, & 77. 1951; Schnack & Covas, Revist. Ar- 
gent. Agron. 18: 107—108, fig. 1. 1951; Datta, Caryologia 5: 
359—370. 1952; Bally, Biol. Abstr, 30: 3351. 1956; Moldenke, 
Biol. Abstr. 30: 1092--1093 & 3551. 1956; Anon., Biol. Abstr. 
30: U36O. 1958; Ahles & Radford, Journ. Elisha Mitchell Soc. 75: 
lUit. 1959; Ratera, Not. Divulg. Inst, Munic. Bot, Carlos Thays 
1: U3. I96I; Moldenke, Biol, Abstr. 38: 865 & 1535 (1962) and 
39: 6Ui & I9U2. 1962; Troncoso, Darwiniana 12: 527—531. 1962; 
Cuf., Bull. Jard. Bot. Brux. 32: Suppl. 787—788. 1962; Langman, 
Biol. Abstr. k2: 596. 1963; Moldenke, Biol. Abstr. U2: 1519 
(1963) and Ii3: 6U3, 1278, B.llO, B.lHi, & B.II8. 1963; Soukup, 
Biota it: 260, 279, & 302. I963; Frei & Fairbrothers, Bull. 
Torret Bot. Club 90: 352. 1963; H. P. Riley, Fam. Flow. Pi. S. 
Afr. 128. 1963; J. D. Poindexter, Biol. Abstr. Ii3: 397. 1963; 
Van Steenis, Fl. Males. Bull. 18: IO69. 1963; Moldenke, R6sum6 
Suppl. 7: 1—3 & 5—10 (1963) and 8: 1—6. I96U; Pearce Seeds 
& Plants, I96U Gard. Aristocrats 20. I96U; R. A. Ludwig, Ind. 
Sem. Canada Dept. Agr. 1961;: 31. I96U; Moldenke, Phytologia 9: 
500—505. I96U. 

It should be noted here that the Blairia Houst., Kaempfera 
Houst., and Sherardia Vail., given by Linnaeus (1751) as syno- 



I96I4. Moldenke, Monograph of Verbena 91 

nyms of Vertena , are actually synonyms of Prlva Adans . , Ghinia 
Schreb., and Stachytarpheta Vahl, respectively. 

The accredition of the name Verbena to "[Tourn.] L." actually 
starts much earlier than the date of the reference given in the 
synonymy (above) of this genus in these notes. In A. L. Juss., 
Gen. PI., ed. 1, 109 (1789) the genus is accredited to "T L", the 
"T" being an abbreviation for Toumefort and the "L" for Linnaeus. 
However, since this placement of the initials is somewhat ambigu- 
ous, it is not by me jregarded as the de facto beginning of the 
accredition used by Robinson & Femald. Jussieu's symbols could 
just as well be interpreted as "Toum. & L." or "Toiuti. in L." or 
"Toum. ex L." 

According to Steudel (I8UI) the genus Verbena was classified 
in the Corytophyta by Necker, the Labiatae Verbeneae by Reic hen- 
bach, the Personatae by Linnaeus, the Ringentes Gymnospermae by 
Royen, the Ringentes Pediculares by Rttling, the Vitices by Juss- 
ieu, and, finally, in the Verbenaceae by Robert Brown. Rttling 
(1766) says of it: "Ringentes & Verticillates conjungit", 

Soukup (1963) records "huallkjapaQra" and "maycha" as vernacu- 
lar names for members of the genus in Peru. The "verbena de tres 
esqiiinas" of Chile is Baccharis sagittalis P. DC. in the Cardua- 
ceae . 

It is perhaps worth noting here that the reference Hook., Bot. 
Misc. 1: 159—173 (1829). given in the bibliography of the genus, 
is sometimes cited as "1830" in errorj the page "66" reference in 
L., Syst. Veg,, ed. 13 (177ll) sometimes cited by authors is also 
Hrroneous . 

Addenda to the list of excluded species; 
Verbena capensis L. - Lippia javanlca (Burm, f ,) Spreng. 
Verbena indica , myuros Barrel. " Elytraria imbrlcata (Vahl) Pers., 

Acanthaceae 
Verbena odorata Pers ♦ ■ Lippia alba (Mill.) N. E. Br, 

VERBQIA ABRAMSI Moldenke 

Additional & emended bibliography: Moldenke, Biol. Abstr, 38: 
865 (1962), 39: I9U2 (1962), I42: 1519 (1963), and 1|3: 1278. I963; 
Moldenke, Phytologia 9: 191. 1963. 

xVERBENA ADULTERINA Hausskn. 

Additional & anended bibliography: Moldenke, Biol, Abstr. 30: 
1093. 1956; Moldenke, Phytologia 8: 120 & lii5~lU6 (1961), 8: 
378—379 (1962), and 9: 38. I963, 

VERBENA ALATA Sweet 

Additional & emended bibliography: Steud., Nom, Bot., ed. 2, 2: 
7li9. I81il; D. Dietr., Syn. Pi. 3: 602. 18U3; Moldenke, Phytologia 
8: Ui9 (I96I) and 9: 113, Uii, 191, & 295. 1963 J Moldenke, R6sum6 
Suppl. 7: 5. 1963. 

VERBENA AMBROSIFOLIA Rydb. 



92 PHTTOLOGIA Vol. 10, no. 2 

Additional & emended bibliography: Rydb., Fl. Rocky Mts. 7U0. 
1917; Moldenke, Wiytologia 8: 123 (1961), 8: l8l, 182, 212, 213, 
231, 279, UOl, UOU, U07, & U35— U37 (1962), and 9: 15--17, 2h, 
27/ 28, 61, 63, 87, 117, 13U, 135, Ui3, 191, & 215. 1963. 

The Ehlers & Ehlers 636I distributed as V. ambrosifolia is ac- 
tually Y. tenuisecta Brig.; Baker , Earle , & Tracy 531 , F. Clark 
U60, M. K. Clemens s.n. [El Paso, Oct. 2h, 1916], Collector un- 
designated L.1-3 , Cory 31173 , Heller & Heller 3536 , G. J. Iken- 
berry 389 , C. L. Lundell 5590, E. G. Marsh 707 , E. J. Palmer 
32195, Parks & Cory 9IU8, Sturgis s.n. [May h, 1902] , Tharp 51-22, 
Waterfall 15782 , and Wooton s.n. [Devil's Park, Aug. 9, 1900] are 
all V. itrightii A. Grayj C. C. Ellis 17 appears to be a mixture 
irith V. TfTightii (the United States National Herbarium specimen 
is definitely that species'.); and £_. Wright 1503 is a mixture of 
V. tumidula Perry and V. bipinnatifida var. latilobata, Perry. 
Wooton 36Ii in most herbaria is V. wrightii, but in the Brit ton 
and the'"toited States National herbaria it is definitely V. am- 
brosifolia. 

McBreen found V. ambrosifolia on the tops of mesas, in a pine- 
piny on- juniper association at the foot of mesas, and in similar 
habitats . 

Additional citations: NEW MEXICO: Lincoln Co.; A. R. Moldenke 
638 (B). Otero Co.: A. R. Moldenke 6U0 (B) . San Miguel Co.: Mc 
Breen s.n. [June 15, 19^] (B), s.n. [near Rowe, October 1, 19631 
(Z), s.n. [near Rove, October 15, 1963] (Z). Santa Fe Co.: Mc 
Breen s.n. [June 22, I963] (B). MEXICO: Coahuila: TSjnad & Mueller 
ffTJls). 

VERBENA AMBROSIFOLIA f . EGLANDULOSA Feriy 

Additional & emended bibliograpliy: Moldenke, Phytologia 8: 
120, 12U, & 152 (I96I), 8: 177, 397, U07, U36, & U37 (1962), and 
9: 16, 17, 2li, 27, 117, 135, & I9I. 1963. 

Additional citations: NEW MEXICO: Luna Co.: A. R. Moldenke 
630 (B, Fg) . 

VERBENA AMOENA Paxt. 

Additional bibliography: Moldenke, Phytologia 8: 18 3— 186, 
UOO, & U27 (1962) and 9: 315 & 329. 1963. 

VERBENA ANDALGALENSIS Moldenke 

Additional & emended bibliography: Moldenke, Biol. Abstr. 30: 
3551. 1956; Moldenke, Phytologia 8: I86 & 1|00 (1962) and 9: 72. 
1963. 

VERBENA ANDRIEUHI Schau. 

Additional & emended bibliography: Moldenke, Phytologia 8: 
186—187 & UOO (1962), 8: I46I (I963), and 9: 16. 1963. 



196U Moldenke, Monograph of Verbena 93 

VERBENA AKAUCANA R. A. Phil. 

Additional bibliography: F. Phil., Cat. Pi. Vase. Chil. 219. 
1881; Moldenke, Phytologia 9: 191. 1963. 

xVERBENA ARGENTINA Moldenke 

Additional & emended bibliograpty: Moldenke, PhTtologia 8: 
120 (1961) and 8: 379 & U.9. 1962. 

VERBENA ARISTIGERA S. Moore 

Additional bibliography: Moldenke, Phytologia 9: 122, 192, 39l4, 
& 397. 1963. 

The A. Robert 8U9 distributed as "TYPE SPECIMEN" of this spe- 
cies is not the type collection at all, nor does it even repre- 
sent this species — it is V. teniiisecta Briq, 

VHIBENA ATACAMENSIS Reiche 

Additional bibliography: Moldenke, Phytologia 9: 192 & 39U. 
1963. 

VERBMA AUEANTUCA Speg. 

Additional bibliography: Moldenke, Phytologia 8: 187, 192—193, 
379—380, & liOO. 1962. 

xVERBENA BAILEIANA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (I96I), 
8: 380 & UOO— iiOl (I962), and 9: 219—221. I963. 

VHIBENA BALAJISAE Briq. 

Additional & emended bibliography: Moldenke, Phytologia 8: 
380 & UOl (1962), 8: U6l (1963), and 9: 192. I963. 

The Pus In 8591 & 15177 distributed as this species are actu- 
ally V. thymoides Cham. 

VERBMA BANGIANA Moldenke 

Additional & emended bibliography: Moldenke, Phytologia 8: 
200 & 202 (1962) and 9: 51. 1963. 

VERBENA BARBATA Grab. 

Additional bibliography: Ste\id., Nom. Bot., ed. 2, 2: 7li9. 
18U; D. Dietr., Syn. PI. 3: 605. 18U3; Moldenke, Phytologia 8: 
U87 (1963) and 9: 60 & 192. I963. 

xVERBENA BEALEI Moldenke 

Additional & an ended bibliography: Moldenke, Phytologia 8: 
120 (I96I), 8: 200—202 & UOl (1962), and 9: 296. I963. 

VERBENA BERTERII (Meisn.) Schau, 

Additional synorymy: Shuttelworthia berterii Meisn. ex Steud., 
Nom. Bot., ed. 2, 2: 575. I81a. Verbena bertari (Meisn.) Schau, 
ex Soukup, Biota U: 279. I963. 

Additional & emended bibliography: Bocq., Adansonia 2: 126, 



9U PHTTOLOGIA Vol. 10, no. 2 

1861— l862j Bocq., Rev. Verbenac. 1;6. 1862; F. Phil., Cat. Pi. 
Vase. Chil. 219. I88lj Moldenke, Phytologia 8: Ul7 & U20 (1962) 
and 9: 12, Ih, 67, 70, 117, 192, 388, 393, 39U, 397, 399, iiOl, & 
li03 (1963), and 9: 501. I96I1; Moldenke, R6suin6 Suppl. 8; 5. I961i. 

VERBENA BERTERII f . ALBIFLORA Moldenke 

Additional synonymy: Verbena berteri f . albiflora Moldenke ex 

Soukup, Biota kt 279. 1963. 

Additional & emended bibliography} Moldenke, Phytologia 8: 381. 
1962 J Soukup, Biota U: 279. 1963} Moldenke, R6sum6 Suppl. 8: 5. 
I96U. 

xVERBENA BINGENENSIS Moldenke 

Additional & amended bibliography: Moldenke, Phytologia 8; 120 
(1961) and 8: 205—207 & 280. I962. 

VERBENA BIPINNATIFIDA Nutt. 

Additional bibliography: Steud,, Nom, Bot., ed. 2, 2: 7U9. 
181^1; A. S. Hitchc, Fl. Kans. pi. 15. 1899} J. W. Blankinship in 
Engelm. & Gray PI. Lindheim, 3: 186. 1907} Moldenke, Phytologia 
8: 1-^1, 12li, 1I49, 150, & 152 (1961) and 8: 177, 178, 182, 2U3, 
279, 378, 397, UOO, U35— U37, & hhO. 1962} Moldenke, Biol. Abstr. 
38: 865 & 1535. 1962; J. D. Poindexter, Trans. Kans. Acad. Sci. 
65: U09 & hl9. 1962} Moldenke Phytologia 9: 1)^—18, 2ii, 27, 61, 
8U, 87, 131, 133—136, lUl, lli3, Ihh, 165, 190, 192, 193, & 220. 
1963; Moldenke, R6sum6 Suppl. 7: 2, 3, & 10. I963. 

The Aydell s.n. [Feb. 1932], H. J. Banker 3685 (in part), 
Cl^onique- Joseph 1^559 , McAtee 33^9 , Miller & Maguire 1255, L. H. 
Pammel 3.n. [Houston, 3-16-29], W. H. Rhoades s.n. [near Atal- 
tala, July 1918], s.n. [near Tipton, September 1935], s.n. [Lees- 
yille, 8-1936], & s.n. [near Attala], £. Singh I63, Tharp , Turner, 
& Johnston 51;7U6a , Ej. Wall s.n. [Nelspruit, 25^0/38], Webster & 
Wilbur 3282 , and B. Williams s.n. [Ruston, March 28, 1950], dis- 
tributed as this species, are all V. tenuis ecta Briq.j Hinton 
119m is V. teucriifolia Mart. & Gal.} and Baker , Earle , & Tracy 
531 , Collector undesignated L.1-3 , G. L. Fisher 36105, 36132, & 
s.n. [July 20, 1936], Goodman & Waterfall U8II, E^ L. Greene s.n. 
[26 July 1880], Heller & Heller 3536 , Herb. Univ. Texas s.n. 
[Marathon, 6/U/31], M. E. Jones s.n. [Rincon, 5-16-1390], R. B_. 
Livingston 3131 , Lotighridge U6I , Meams IO8 &. 109 , A. Nelson 
10358 , E.J. Palmer 32195 , F. G. Plummer s.n. [Lincoln National 
Forest, I9O3] , H. H^ Rusby 337 , E. D. Schvilz s.n. [near Alpine, 
Avig. U, 1928], Snow s.n. [Santa Fe, Aug. »80], Tharp 88UO, White- 
house 19537, Woo ton s.n. [Divide above Mescalero Agency, June 23, 
1895] and s.n. [White Sands, Aug. 25, 1399], C. L^ York U806U, 
and Zobel s.n. [Deer Creek Canyon, May 25, 193II] are all V. 
wrightii A. Gray, 

The W. H. Over 2103 (W— 582936) cited in Phytologia 8: U02 



196U Moldenke, Monograph of Verbena 95 

(1962) from 'Washington Co., South Dakota", is actually from 
Washington County, Nebraska. Likewise, the Demaree 12003 (W — 
1683782) cited in Phytologia 8: 231 & U03 from "Laoar Co., Okla- 
homa" is from Lamar County, Texas, in spite of the fact that the 
labels are plainly marked "Plants of Oklahoma" . 

Additional citations: TEXAS: Dallas Co.: Lundell & Lundell 
1131|(B). 

VERBENA BIPINNATIFIDA var. UTILOBATA Perry 

Additional bibliography: Moldenke, Phytologia 8: 177, 212— 21]^, 
231, & 397 (1962) and 9: 16, 2U, 135, & 193. 1963. 

xVERBENA BUNCHARDI Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (I96I), 8: 
U65 (I963), and 9: Ihl, 193, 219, & 220. I963. 

VERBENA BONARIENSIS L. 

Additional synonymy: Verbena quadr angulari s Arrab, ex Steud,, 
Norn. Bot., ed. 2, 2: 750. I8UI. 

Additional & emended bibliography: Kniphof, Bot. Orig. Herb. 
Viv. cent. 2: pi. [28U]. 1757; J. A. llurr. in L., Syst. Veg., ed. 
13, 62. I77U; L'H^r., Stirp. Nov. 1: 22. 1786; Dum. Cours., Bot. 
Cult., ed. 2, 2: 623. 1811; Hook., Bot. Misc. 1: 159 & 166. 1829; 
Steud., Nom. Bot., ed. 2, 2: 750 £t 751. I8UI; D. Dietr., Syn. Pi. 
3: 600. 181^3; F. Phil., Cat. Pi. Vase. Chil. 219- I88I; Hicken, 
Chloris Plat. Argent. 195—196. I9IO; N. Taylor, Mem. N. I. Bot. 
Card. 5: [Fl. Vic. N. I.] 526. 1915; Ratera, Not. Divtilg. Inst. 
Munic. Bot. Carlos Thays 1: li3. I96I; Moldenke, Phytologia 8: 
120, 121, & 12U (I96I), 8: 267, 280, 313, 3lU, 3l6~3l8, 381i, & 
U35 (I962), 8: U6U, U77, & U90 (I963), and 9: hh, 66, 67, 190, 
I9U— 197, 2IJ4, 215, 293, 295, 296, 361, 37ii— 376, 379, 381, & 
382. 1963; Moldenke, Rlsum4 Suppl. 7: 1, 2, & 6. 1963; Soukup, 
Biota h: 279. 1963; Moldenke, Phytologia 9: 502. I96U. 

Einended illustration: Kniphof, Bot. Orig. Herb. Viv. cent. 2: 
pl. [28U] (in color). 1757. 

Ratera (I96I) records the common names "verbena" and "yerba 
de los hechiceros" and comments that "Se utilizan las hojas y 
ramitas j6venes, generalmente para ciertas afecciones hepiticas 
y gastricas" . 

It should be noted here that the Kniphof, Bot. Orig. Herb. Viv. 
reference given above in the bibliography of this species is some- 
times erroneously cited as "11" and the Hook., Bot. Wise. 1: 159 
& 166 (1829) reference is often misdated "I83O". The A. R. Molden- 
ke U52 (Fg) cited in Phytologia 8: U06 (1962) from "Columbus Co., 
South Carolina" is actually from Colimibua County, North Carolina. 

Additional citations: TANGANYIKA: Drnnmond & Hensley 2092 (B) . 

VERBENA BONARimSIS var. CONGLCJiERATA Briq. 

Additional &. emended bibliography: Moldenke, Phytologia 8: 256, 
382, 383, & U07 (I962), 8: U63 (1963), and 9: 375 & 382. I963. 



96 PHYTOLOGIA Vol. 10, no. 2 

VIEBENA, BRACTEATA Lag. & Rodr, 

Additional &. emended synonymy: Verbena bracteata Cav. ex 
Steud., Nom, Bot., ed. 2, 2: 700. I8I4I. Zapania bracteosa Poir. 
ex Steud., Nom. Bot,, ed. 2, 2: 750, in syn. iSliL. Zappania 
bracteosa Poir. ex Steud., Nom, Bot., ed, 2, 2: 797. I8UI. 

Additional & emended bibliography: L'H^r., Stirp. Nov. 1: 22. 
1786; Michx., FI. Bor.-Am., ed. 1, 2: 13. 1803; Pursh, Fl. Am. 
Sept. 2: Ul6. l8lUj Michx., Fl. Bor.-Am., ed. 2, "1" [=2]: 13. 
I820j Lehn., Del. Sem. Hort. Hamb. 1826: 16. 1826; Hook., Comp, 
Bot. Mag. 1: 176. I836; Steud., Nom. Bot., ed. 2, 2: 7^0 & 797. 
18U1} D. Dletr., Syn. PI. 3: 60li. 18^3; J. Torr. in Bnory, Rep. U. 
S. & Mex. Bound. Surv. 2: 128. 1858; F. Phil., Cat. PI. Vase. 
Chil. 219. 1881; Hitchc. & Norton, Kans. Agr. E:q). Sta. Bull. 50: 
5U, pl. 6, fig. 92. 1895; Selby, Bull. Ohio Agric. Sta. 83: 329. 
1897; A. S. Hitchc, Fl. Kans. pl. 15. 1899; K. Schum. in Just, 
Bot. Jahresber. 28 (1): U97. 1902; Selby, Bull. Ohio Agric. Sta, 
175: 350. 1906; N. Taylor, Mem. N. Y. Bot. Gard. 5: [Fl. Vic. N. 
T.] 526, 1915; Tidestr,, Contrib, U, S. Nat. Herb. 25: U69. 1925; 
Moldenke, Phytologia 8: 120 & 121 (I96I), 8: 177, 206, 212, 213, 
257, 268, 316, 397—399, UOl, U28, U35, 1*37, & U39 (1962), 8: U62, 
U69, U71, U72, & Ii77 (1963). and 9: 16, 2U, 27, 53, $h, ihh, 156, 
195, 215, 219, 220, 358, & UOli. 1963; Moldenke, Biol. Abstr. 38: 
15^ (1962) and 39i 61U, I962; J. D. Poindexter, Trans. Kans. 
Acad. Sci. 65: I4I8. 1962; Moldenke, R6sum6 Suppl. 7: 1 — 3 & 9. 
1963. 

It is of interest to note that Steudel (iSIil) maintains V. 

bracteata Cav. and V. bracteosa Michx. as two sepsirate species, 
placing V. repens Spreng. and V. squarrosa Roth in the synonymy 
of the former and Zapania bracteosa Poir. and Zappania bracteosa 
Poir. in the synonymy of the latter. 

The Over 2389 (W— 58296U) cited in Phytologia 8: U08 (1962) 
from "Washington Co., South Dakota" is actually from Washington 
County, Nebraska, and the A. R. Moldenke 118 (Fg) cited on page 
UlO of the same volume from "Qratfy" County, Texas, is from Gray 
County in that state. 

Hooker (I836) cites T. Drtimmond 253 ter in the Kew herbarium. 

Additional citations: COLORADO: Boulder Co.: Ewan , Plant. Ex- 
sicc. Gray. IO9O (B, B) . 

VERBENA BRACTEATA f . ALBIFLORA (Cockerell) Moldenke 

Additional & emended bibliography: Moldenke, Phytologia 8: 309 • 
1962; Moldenke, Biol. Abstr. 39: 6lit. I962, 

VERBENA BRASILIENSIS Veil. 

Additional synonymy: Verbena brasiliensis Arrab. ex Steud., 
Nom. Bot., ed. 2, 2: 750. I8la. 

Additional bibliography: Stevid., Nom. Bot., ed. 2, 2: 750. 
I8UI; Moldenke, Phytologia 8: 121; & li^S (I96I) and 8: 256, 257, 
& U05. 1962; Moldenke, Biol. Abstr. 39: 6IU & 191*2. 1962; Mol- 



196U Moldenke, Monograph of Verbena 97 

denke, Phytologla 8: li63 (1963) and 9: 66, 101, 19U— 197, 296, & 
382. 1963| Moldenke, Biol. Abstr. U2t 1^19. 1963; Moldenke, R^su- 
ml Suppl, 7: 1—3. 1963. 

The A. R. Moldenke h$7 (Fg) cited in Koytologia 8: Ul3 (1962) 
froo "Columbus Co., South Carolina" is actually from Columbus 
County, North Carolina. 

VERBENA CABRHIAE Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (1961), 
8: 189 & Ul9 (1962), and 9: 70. I963. 

The species has been found by Meyer & Sleumer at 180 meters 
altitude, blooming in December. 

Additional citations: ARGENTINA: Santiago del Estero: Meyer & 
Sleumer s.n. [T. Meyer \$2$T\ (B) . ~ 

VERBENA CALLIANTHA Briq. 

Additional bibliography: Moldenke, Phytologia 8: 120 (I960), 
8: 189, 202, & U16 (1962), and 9: 10, 12, Hi, 67, 70, 72, 197, 
388, & 393. 1963. 

The Herb. Osten 7900 and T. Rojaa 382 & 1382 , distributed as 

this species, are actiially V. tomophylla Briq. 

VERBENA CAMERONENSIS L. I. Davis 

Additional bibliography: Moldenke, Pl^ologia 8: 12li & li36 

(1961) and 9: 62, 63, 8U, & 197. 1963j Moldenke, R4sum6 Suppl. 
7: 3. 1963. 

The specimen of Galeotti 777 photographed by Macbride as his 

photograph no. 2U699 is deposited in the Delessert Herbarium at 
the Conservatoire et Jardin Botaniques at Geneva. 

VERBENA CAMPESTRIS Moldenke 

Additional bibliograply: Moldenke, Phytologia 8: U25--U26 

(1962) and 8: U65. 1963. 

Additional citations: BRAZIL: Santa Catarina: J^ Fj. T. MBller 
s.n. (P) . 

VERBESA CANADENSIS (L.) Britton 

Additional & emended synonymy: Verbena aubletia Juss. ex 
Hook., Comp. Bot. Mag. 1: 176. I836. Billarderia explanata 
Moench apud Steud., Nom. Bot., ed. 2, 1: 205, in syn. I8U0. Ver- 
bena aubletia L. apud Steud., Noti. Bot., ed. 2, 2: 7li9. l81il. 

Additional & emended bibliography: L'H^r., Stirp. Nov. 1: 22. 
1786; Michx., Fl. Bor.-Am., ed. 1, 2: 13. 1803; Dum. Cours., Bot. 
Cult., ed. 2, 2: 623--62U. I8II; Pursh, Fl. Am. Sept. 2: la5— 
U16. iSLli; Michx., Fl. Bor.-Am., ed. 2, "1" [-2]: 13. 1820; 
Hook., Bot. Misc. 1: I69. 1829; Lehn., Del. Sem. Hort. Hamb. 
1832: 7. 1832; Hook., Comp. Bot. Mag. 1: 176. I836; Steud., Nom. 
Bot., ed. 2, 1: 205, 23U, & 637 (I8U0) and 2: 201, 7li9, & 750. 
18U1| D. Dietr., Syn. PI. 3: 69U. 18U3; J. Torr. in Bnory, Rep. 
U. S. & Mex. Bound. Surw. 2: 128. 1858; Bocq., Adansonia 2: 126. 
1861 — 1862; Bocq., Rev. Verbenac. \ik. 1862; A. Gray, Syn. Fl. N. 



98 PHTTOLOGIA Vol. 10, no. 2 

Am. 2 (1): 337. l878j A. S. Hitchc, Fl. Kans. pi. 15. 1899; Ed. 
Rodlgas, Bull. Arboricxat. Belg. 1902: 113 & 111;. 1902; L. H. 
Bailey, Botany 372. 1911; Fernald, Rhodora 38: Uli3. 1936; K. V. 
0. Dahlgren, Svensk. Bot. Tidsk. 32: 231. 1938; Cain, Found. PI. 
Geogr. 335. 19Ui; Moldenke, Phurtologia 8: 120, 121, 123, 12U, & 
151 (1961) and 8: 177, 182, 183, 187, 20U, 210, a2— 2lU, 231, 
2U3, 279, 280, UOl, U07, & li23. 1962; J. D. Poindexter, Trans. 
Kans. Acad. Sci. 65: U09 & U.9. 1962; Moldenke, Phytologia 8: U62 
& U77 (1963) and 9: 10, 16, 2li, 27, 61, 62, 82—85, 87, 115—118, 
135, 136, II43, liiU, 193, 197—198, 308—315, 352, 376, 382, 39k, 
397, UOO, Sc UOU. 1963; Moldenke, Biol. Abstr. 1^2: I5l9. 1963; 
Pearce, Seeds & Pi. 10. I963; G. N. Jones, Fl. 111., ed. 3, [Am. 
Midi. Nat. Monog. 7:] 213. 1963; Moldenke, RfisumS Suppl. 7: 9 
(1963) and 8: 5. 1961;. 

The Hook., Bot, Misc. Is I69 (1829) reference given above is 
often erroneously cited as "18 30", while J. Torr. in Emory, Rep. 
U. S. & Mex. Bound. Surv. 2: 128 is often referred to as "Torr. 
Bot. Mex. Bound. 128". 

Fernald (1936) cites Fernald & Grisccm Ui96 from Princess 
Anne County, Virginia, and notes that "In Dr. Perry's Revision ... 
recoided northward only to North Carolina but Small (Man.) ex- 
tends the range to Virginia." Hooker (I836) cites T. Drummond s. 
n. [Jacksonville], s.n. [St. Louis], and s.n. [New Orleans, 1833] 
deposited in the Kew herbarium. 

The A. D. Brubaker 22, Diener 835, S. F. Evans E.U270 , Girvin 
s.n. [Indian Res., March 15, I9U0] , Grossman 7, Parks & Cory 
22131, Rose-Innes & ffamock 21818 , and Tharp 253 & s.n. [7/23/39], 
distributed as V. canadensis , are all V. tenuisecta Briq.; H. H. 
Bartlett 10030 , Collector undesignated s.n. [Plains near Leon 
Spring, Sept. 7, 1852], and £. Wright It55 are V. tumidula Perry; 
C. Wright 1503 is a mixture of V. tumidula and V. bipinnatifida 
var. latilobata Perry; and Goodman & Waterfall 1+811, G_. J_^ Iken- 
berry 256, 0. B^^ Metcalfe IO9O , Mulford 37 & s.n. [near Albuquer- 
que, Sept. ^9^]f Parry , Bigelow , Wright , & Schott s.n. [Frontera, 
Mar. 22, 1852], Rehn & Viereck s.n. [April 16, 1902], and W, W. 
Robbins 82li2 are all V. wrightii A. Gray. 

VERBENA CANADENSIS f . CANDIDISSIMA (Haage & Schmidt) Palmer & 
Steyerm . 
Additional bibliography: Moldenke, Phytologia 8: U37 (1962) 
and 8: U68— i;69. 1963 . 

VEEBENA CANESCENS H.B.K. 

Additional synonymy: Verbena canescens Humb. & Bonpl. ex 
Lehm., Del. Sam. Hort. Hanb. 1826: 16. 1826. Verbena canescens 
Humb. & Kunth ex D. Dietr., Syn. PI, 3: 60U. 18U3. Verbena neei 
Moldenke, Phytologia 2: 2Ul. 19U7. 

Additional & emended bibliography: Lehm., Del. Sem. Hort. 
Hamb. 1826: 16. 1826; Hook., Bot. Misc. 1: I68. 1829; Lehm,, Del. 



196U Moldenke, Monograph of Veztena 99 

Son. Hort. Hamb. 1832: 7. 1832} Steud. Ncm. Bot., ed. 2, 2: 201 
& 750. I8UI; D. Dietr., Syn. Pi. 3: 60U. 18^3; J. Terr, in Emory, 
Rep. U. S. & Mex. Botind. Siirv. 2: 128. 1858; A. Gray, Syn. Fl. N. 
Am. 2 (1): 336—337. 1878; J. W, Blankinship in Engelm. & Gray, 
PI. Lindheim. 3: I87. 1907; Parodi, Rev. Argent. Agr. lli: 61— 69. 
19U7; Moldenke, Phytologia 2: 2hl, 331, & 339. 19li7; H. N. & A. 
L. Moldenke, Pi. Life 2: hh &. 73. I9li8; Moldenke, Known Geogr. 
Distrib. Verbenac., [ed. 2], 19, 23, 32, IO6, 197, & 198. 19U9; 
Moldenke, Alph. List Git. 3: 753, 767, 771, 772, 78U, 787, 799, 
802, 807, 829, Qhh, 850, 898, 93h, S. 963. 19U9; E. J. Salisb., 
Ind. Kew. Suppl. 11: 263. 1953; Moldenke, R6sum6 21;, 28, 39, 127, 
360, 361, 365, 373, ii71, & Ii72. 1959; Moldenke, Phytologia 8: lU 
(I96I) and 8: 21^3, 268, & 279. 1962; Troncoao, Danrixiiana 12: 
529—530. 1962: Langman, Biol. Abstr. k2: 596. I963; Moldenke, 
Phytologia 8: Ij77 (I963) and 95 17, 87, 155, 156, 165, 198, 215, 
& 379. 1963; Moldenke, R^sum^ Suppl. 7: 3 (I963) and 8: 2. 5. & 
6. I96U. 

It shoxild be noted here that the reference Hook., Bot. Misc. 
1: 168 (1829), cited above, is often incorrectly cited as •»1830" 
and J. Torr. in Bnory, Rep. D. S. & Mex. Bound. Surv. 2: 128 
(1858) is often cited as "Torr. Bot. Mex. Boimd. 128". 

Troncoso (I962) says "El tipo de Verbena Neei Mold....se en- 
cuentra en el Herbario del Institute Botfinico A. J. Cavanilles 
de Madrid (MA) y lleva un r6tulo que dice: 'Pampas de Buenos 
Aires, N6e, Iter IO8, Exped. Malaspina.' Examinado dicho tipo, 
pude comprobar que no es una planta argentina y que el origen 
indicado en la etiqueta es err6neo, ccmo en tantos otros ejem- 
plares de N6e que sufrieron confusi6n de r6tulos (ver Parodi L. 
R., Rev. Arg. de Agr. lU: 61— 69. 19li7) . Se trata, en verdad, 
de Verbena canes c ens H.B.K., originaria de Mexico, c<ano lo he 
podido verificar por la descripci6n y limina originales. Tambifin 
coincide muy bien con los siguientes ejemplares de herbario del 
Darwinion: MEXICO: Oaxaca, Pr ingle 1;78U . VIII-I89U (Si). COLOM- 
BIA: San Cristobal, Fr. Apollinaire , n-1908 (Si.33li3). Verbena 
Neei desaparece, pues, del catilogo de la flora argentina y pasa 
a la sinonimia de V. canescens H.B.K." This, by the way, is the 
first record of the species from Colonbia. 

The N. C. Henderson 63-96 and E. G. Marsh U9U , distributed as 
V, canescens , are actually var. roemeriana (Scheele) Perry; L. F. 
ffard 3.n. [Heame, Sept. 11, 1877] is in part V. halei Small~and" 
in part V. xutha Lehm.; Smith , Peterson , & Tejeda 3907 is V. 
menthaefolia Benth., 39UO is the type collection of V. canescens 
f . albiflora Moldenke, and 3957 is V. neomexicana (A~Gr^) Small. 

Crutchfield & Johnston describe V. canescens as "infrequent 
perennials in short brush on shale hills". 

Additional citations: MEXICO: Tamaulipas: Crutchfield & John- 
ston 5807a (Au— 1876U2) . State undetermined: Nfe IO8 (F— photoj 
F— photo, N, N— photo, Q, Z— photo) . 



100 PHTTOLOGIA Vol. 10, no. 2 

VERBENA CANESCENS f . ALBIFLORA Mcldenke, Phytologia 9: 500—501. 
196U. 

Bibliography: Moldenke, Phytologia 9: 500 — 501. I96U. 

This form differs from the typical form of the species in hav- 
ing white corollas. 

The type of the form was collected by C. E. Smith, Jr., F. A. 
Peterson, and Narclsso Tejeda ( no. 39 UO ) in gray to whitish soils 
with thom-scrub-cactus cover, among occasional calcareous out- 
crops along the Tehuacin-Orizaba highw^ just above Azumbilla, at 
1500 — 1800 meters altitude, Puebla, Mexico, on July 18, I96I, and 
is deposited in the United States National Herbariiom at Washing- 
ton. The fonn is known thus far only from the type specimen. 

Citations: MEXICO: Puebla: Smith , Peterson , & Tejeda 39l;0 (W — 
2397959--type) . 

VERBENA CANESCENS var. ROEMERIANA (Scheele) Perry 

Additional bibliography: Moldenke, Phytologia 8: 12U (I96I), 

8: U6U & li71 (I963), and 9: 36, 165, & 198. 1963? Moldenke, R6- 

sum6 Suppl. 7: 3 (1963) and 8: 1. I961i. 

Additional citations: TEXAS: Shackleford Co.: N. C. Henderson 

6>96 (Au— 2I76U6). MEXICO: Coahuila: E, G. Mar8h"Tl9ir (Au— 212lj82). 

VERBENA CAROLINA L. 

Additional & emended synocymy: Verbena biserrata Hvnnb, & Bonpl. 
apud Steud., Norn. Bot., ed. 2, 2: 7U9. I8UI. Verbena polystachya 
Hvmib. & Bonpl. apud Steud., Norn. Bot., ed. 2, 2: 750. I8UI. Ver- 
bena veronicaefolia Humb. & Bonpl. apud Steud., Norn. Bot., ed. 2, 
2: 751. I8UI. Verbena veronicaefolia Humb. & Kvmth apud D. Dietr., 

Syn. PI. 3: 601. 18U3. 

Additional & emended bibliography: J, A. Murr. in L., Syst. 
Veg., ed. 13, 62. 177U; L'H6r., Stirp. Nov. 1: 22. 1786; Dum. 
Cotirs., Bot. Cult., ed. 2, 2: 623. I8llj Hook., Comp. Bot. Mag. 
1: 176. 1836; Steud., Nom. Bot., ed. 2, 2: 7U9~75l. iSUl; D. 
Dietr. jj Syn. PI. 3: 6OI. 18U3; N. J. Anderss., Galap. Veg. 199 — 
200. I851i; N. J. Anderss., Vet. Akad. Handl. Stockh. 1853: 199—- 
200. 1855; A. Gray, Syn. Fl. N. Am. 2 (1): 335. 1878; S. ¥ats., 
Proc. Am. Acad. Sci. 18: 135. 1883; H. H. Rusby, Mem. Torrey Bot. 
Club 6: 106. 1896; Moldenke. Phytologia 8: 121, 12h, &. 1^3 (1961), 
8: 257, 317, & hn (1962), 8: U96 (1963), and 9: 52, 81, 93, 126, 
151, 165, 198—199, 211i, 215, & 219. 1963; Moldenke, R6sum6 SuppL 
8: 6. I96I4. 

Steudel (I8UI) notes that according to Sprengel V. polystachya 
Is a synonym of V. urticifolia L. This, however, is not true, 
although the two species are certainly closely related, Watson 
(I883) was of the opinion that Palmer 2037 is "probably a hybrid 
between V. polystachya or V. urticaefolia and V. xutha ", but I 
regard it as typical V. ehrenbergiana Schau, 

The Smith , Peterson , & Tejeda 3701 distributed as V. Carolina 
is actually f . albiflora Moldenke; Herb. Molliano s.n. [P.], 0. ' 
Sanders 5, and S. M^ Tracy 8037 are V. urticifolia L.; W, C. Co- 



196U Moldenke, Monograph of Verbena 101 

ker s.n. [July 8, I909] is V. urticifolia var. leiocarpa Perry & 
Feraaldj Herb. Hort. Bot. Genev. s.n, [l5 Axig . 1826] is in part V. 
urticifolia var, leiocarpa and in part V. recta H.B.K.J and T^ L. 
Andrews s.n. [Ascension, 13-1] and !_. L. Forbes s.n, [Colfax, Aug, 
20, 1927] are V, xutha Lehm, 

Hooker (18357 cites T. Drummond 253 from New Orleans, Louisiana, 
presumably deposited in the Keir herbarium. 

Additional citations: ARIZONA: Cochise Co.: A« R. Moldenke 78I 
(B, Fg). 

VERBENA CAROLINA f . ALBIFLORA Moldenke 

Additional bibliography: Moldenke, Phytologia 7: U20 (I96I), 8: 
li87 & U91 (1963), and 9: 199. 1963. 

Smith, Peterson, & Tejeda fOTind this form growing in gravelly 
gray or brown soil at the edge of a field in thom-scrub-cactus to 
mesic semi-evergreen forest formation, 100 to 1800 meters Jiltitude. 

Additional citations: MEXICO: Puebla: Smith , Peterson , & Tejeda 
3701 (W— 23977U3) . 

VERBENA CATHARINAE Moldenke 

Additional bibliography: Moldenke, Ptytologia 8: U96 (1963) and 
9: 8. 1963} Moldenke, Biol. Abstr. U3: 1278, I963. 

VERBENA CHACENSIS Moldenke 

Additional bibliography: Moldenke, Biol. Abstr. 30: 3551. 1956} 
Moldenke, Phytologia 9: 8— -9. 1963. 

VERBENA CHEITMANIANA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 1|19 (1962) and 
9: 199. 1963. 

VERBENA CHILENSIS Moldenke 

Additional bibliography: Moldenke, Phytologia 9: 122, 125, 126, 
& 199. 1963. 

VERBENA CILIATA Benth. 

Additional bibliography: Steud,, Norn, Bot,, ed, 2, 2: 750, l81ilj 
D. Dietr,, Syn, Pl, 3: 605. 18U3; A. Gray, Syn, Fl, N, Am, 2 (1): 
337. 1878} J. W, Blankinship in Engelm, & Gray, PI, Lindheim, 3: 
187, 1907; Shreve, Carnegie Inst, Wash, Publ. 217: 19 & Uh. 19l5j 
Moldenke, Phytologia 8: 12U & 152 (1961), 8: 177, 182, 202, 212, 
213, 215, 231, 2U3, 279, 397, 399, UOO, hn, U20, U23, li2li, U35, & 
U36 (I962), 8: U73 & U87 (1963), and 9: 21—28, 61, 65, 67, 70, 8U, 
87, 13U— 137, Ua, IIJ^, 156, 199—200, 388, 393, & 397. I963j Mol- 
denke, R6sum6 Suppl, 7: 3 & 9. 1963. 

Smith, Peterson, & Tejeda describe the corollas of this plant 
as magenta and fo\ind the plant in black to gray soils, common on 
roadsides and in pastures, in a formation of oak forest above giv- 
ing way to scrubby secondgrowth thickets below. Sonderstrcm de- 
scribes it as having corollais "pink, turning blue with age" , 



I 



102 PHYTOLOGIA Vol. 10, no. 2 

Shreve (1915) calls the species an ephemeral summer-active herba- 
ceous plant. 

The H. H. Rusby 12U distributed as V. ciliata is actually V. 
teucriifolia Mart, & Gal.j Schery ll;2 is in part V. menthaefolia 
Benth. and in jjart V. teucriifolia ; Pringle 3551 is V. teucrii - 
folia var. corollulata Perry j Parry & Palmer 719 is in part y. 
ciliata and in part V. teucriifolia var. corollulata ; E. J. Palm- 
er 13512 is the type collection of V. tumidula Perryj and East- 
wood 15697 , Ferril s.n. [May 11, I9O6] , Griffiths 5190 , Herb. 
State Agric . Coll. UI8U , Nelson & Nelson U983 & 5017 , & J. Skehan 
5 are V, irrightii A. Gray. 

Additional citations: MEXICO: Chihuahua: Sonderstrcm 8U7 (W — 
239611*5) . Puebla: Smith , Peterson , & Tejeda 3921 (W— 2397939) . 

VERBEMA CILIATA var. LONGIDENTATA Perry 

Additional bibliography: Moldenke, Phytologia 8: 182, 212— 211i, 
U36, & U37 (1962) and 9: 16, 19, 22—27, 61, 6U, 135, 193, 199, & 
200. 1963. 

The Waterfall 3922 distributed as this variety is actually V. 
wrightii A. Gray. 

VERBENA CILIATA var. PUBERA (Greene) Perry 

Additional bibliograpJ^jr: Moldenke, Phytologia 8: 212, 215, 
397, 399, & 1;37 (1962) and 9: 16, 2U, 26—29, 135, 136, & 200. 
1963. 

The Eastwood 15697, Meams IO9, Parks & Cory 18358 , Sperry T« 
126 & T.573 , Thurber lU3 , and £, E^ Wolff 1666 distributed as 
this variety are actually V, wrightii A. Gray. 

VERBENA CLAVATA Ruiz & Pav. 

Additional bibliograply: Steud,, Nom. Bot., ed. 2, 2: 750. 
181^; D. Dietr., Syn. PI. 3: 60U. l8U3i Bocq., Adansonia 2: 126. 
1861 — 1862 J Bocq., Rev. Verbenac. U6. 1862; Moldenke, Phytologia 
9: 29—33, 117, & 200 (I963) and 9: 501. I96U. 

VERBMA CIAVATA var. CASMENSIS Moldenke 

Additional bibliography: Moldenke, Phytologia 9s 31 — 33 & 200. 
1963. 

xVEEBENA CLEMENSORUM Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (I96I) and 
9: 33— 3li. 1963. 

VERBENA CLOVERAE Moldenke 

Additional bibliography; Moldenke, Phytologia 8: 12U (1961), 8t 
U77 (1963), and 9: 3U— 37, 6U, 198, 200, & 201. 1963; Moldenke, K&- 
sum6 Suppl. 7: 3 & 9. 1963. 

VERBENA COCCINEA Raf . 

Additional bibliography: Moldenke, Phytologia 9: 201. 1963; Mol- 



1961i Moldenke, Monograph of Verbena 103 

denke, RfisumS Suppl. 7: 3. I963. 

VERBENA COCHABAMBENSIS Moldenke 

Additional bibliography: Moldenke, Phy-tologia 9: 37 — 38, 201, 
& 39I1. 1963; Moldenke, R^sum^ Suppl. 7: 9. 1963. 

xVERBENA CONATA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 & lk$ 
(I96I) and 9: 38—39 & 166. I963. 

xVERBENA CORRDPTA Moldenke 

Additional bibliograply: Moldenke, Biol. Abstr. 30: IO93, 
1956 J Moldenke, Phytologia 8j 120 (I96I) and 9: UO. I963. 

VERBENA CORYMBOSA Rtdz & Pav. 

Additional & emended bibliography: Steud., Nom. Bot., ed. 2, 
2: 750. l8Ulj D. Dietr. Syn. Pi. 3: 602. 13U3; F. Phil., Cat. 
PI. Vase. Chil. 220. 1881^ Baeza, Nomb. Vulg. PI. Silv. Chile, 
ed. 2, 60—61, 233, & 269. I93O: Moldenke, Phytologia 8: 2U7, 2^7, 
280, & 316 (1962), 8: 1;63 (I963), and 9: IiO~U5, 201, & 296. I963. 

xVERBENA COVASII Moldenke 

Additional bibliograpty: Moldenke, Phytologia 8: 120 (I96I) 
and 9: U5— U6 & 201. I963. 

VERBMA CRITHMIFOLIA Gill. & Hook. 

Additional & emended bibliograpt^: Steud. Nom. Bot,, ed. 2, 
2: 7^0. I8I1I; D. Dietr., Syn. PI. 3: 603. 18U3; H. S. Marshall, 
Kew Bull. 1936: 9k. 1936; Moldenke, Phytologia 8: 123 (1961) and 
9: li6— li9, 120, 201—202, & 299—302. I963. 

According to the original collectors, the flowers of this plant 
"yield a honey-like smell". They describe their P> minor as 
foliis angustioribus". I regard this variety as V. hookeriana 
(Covas & Schnack) Moldenke, which see. 

Additional citations: ARGENTINA : Mendoza: Sleumer 328 (B). 

VERBENA CUNEIPOIIA Ruiz & Pav. 

Additional synonymy: Verbena cuneifolia Pers. ex Steud., Nom. 
Bot., ed. 2, 2: 750, in syn. I81a. 

Additional & emended bibliography: Hook., Bot. Misc. 1: I70. 
1829; Steud., Nom. Bot., ed. 2, 2: 750. I8UI; D. Dietr., Syn. PI. 
3: 603 & 60U. l8U3i F. Phil., Cat. PI. Vase. Chil. 220. I88I; 
Moldenke, Phytologia 9: 50—52, 202, 286, 293, & 296. I963. 

It is worth noting here that the reference Hook., Bot. Misc. 
1: 170 (1329) is often erroneously cited as "1330»». 

^® £1 hi Stevens 7 distributed as this species is actually 
V, villifolia Hayek. 

VERBENA CURTISII Moldenke 

Additional bibliography: Moldenke, Phytologia 8: U88 (1963) and 
9: 52—53. 1963. 



lOlj PHTTOLOGIA Vol. 10, no. 2 

iTKRBENA. DEAMEI Moldenke 

Additional ^ anended bibliography: Moldenke, Phytologia 8: 120 
(1961), 8: 268, 272, & 280 (1962), and 9: 53—5U, 59, & 202. 1963j 
Moldenke, Biol. Abstr. U3: 61i3. 1963. 

VHIBENA DELICATULA Mart. 

Additional bibliography: Moldenke, Phytologia 8: 201 (1962) 
and 9: 59—60. 1963 . 

VERBENA DELTICOLA Small 

Additional & emended bibliography: Moldenke, Phytologia 8: 12U 
(1961), 8: 397, U23-li25, 1;35, k36, & U39 (1962), 8: U60, I46I, & 
U65 (1963), and 9: 16, 36, 60— 6U, 8U, 87, 135, lUli, & 202. 1963j 
Moldenke, RSsumfi Suppl. 7: 3. 1963 f 

The Collector undesignated s.n. [Plains near Leon Spring, Sept. 

7, 1852], distributed as "this species ifith a question, is actual- 
ly V. tumidula Perry. 

xVEEBENA DERMENI Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (1961), 8; 
U63 (1963), and 9: 66—67 & 296. I963. 

VHIBEMA DISSECTA TITilld. 

Additional & emended bibliography: Steud.. Nom. Bot., ed. 2, 
2: 750. I81a; D. Dietr., Syn. Pl. 3: 605. I8u3j F. Phil., Gat. Pi. 
Vase. Chil. 220. I88I; Moldenke, Phytologia 8: 121 & 123 (1961), 
8: 190, 202, liOO, UOl, U17, la9, & U20 (1962), 8: U6l (1963), and 
9: 12, Hi, 67—75, 128, 202—203, 388, 393—397, & 399— liOl. 1963. 

The G. L^ Fisher 200 , R. M. Harper 25, N. Y^ Bot. Gard. Cult. 
Pi. 22J428, O'Neill 551 & s.n. [River Junction, April 9, 1925], 
P^odi 12233, R. C_. Schneider s.n. [N. Y. Bot. Gard. Cult. Pi. 
22U28], Troncoso 291, and Valeur hhQ , distributed as this species, 
are all V. tenuis ecta Briq. 

xVERBENA DISSOLUTA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (1961) 
and 9: 75—76. 1963 . 

VERBENA DUSENII Moldenke 

Additional bibliography: Moldenke, Phytologia 9: 79—80, 120, 

& 203. 1963. 

VERBENA EHRENBERGIANA Schau. 

Additional bibliography: S. Wats., Proc. Am. Acad. Sci. 18: 
135. 1883} Moldenke, Phytologia 8: 121; (I96I), 8: U87 & U88 
(1963), and 9: 80—82. I963. 

Watson (1883) says that Palmer 2037 is "probably a hybrid be- 
tween V. polys tachya or V. urticaefolia and V. xutha ", but Miss 
Perry regards it as typical V. ehrenbergiana and I see no reason 
to doubt this disposition oSTTt, 



196U Moldenke, Monograph of Verbena 105 

VERBENA ELEGANS H.B.K. 

Additional & emended synonymy; Verbena elegans Hxanb, & Bonpl. 
ex Steud., Nom. Bot., ed. 2, 2: 7$0, l8Ul. Verbena lamberti Ker 
es Steud., Ncan. Bot., ed. 2, 2: 7$0. l8Ul. 

Additional & omended bibliograply: Hook., Bot, Misc. 1: 169 . 
1829; Steud., Nom. Bot., ed. 2, 2: iSO, iQhX', Moldenke, Phytolo- 
gia 8: 213, 2li3, U23, U27, & U35 (1962), 8: W? (1963), and 9» 
16, 61, 82—87, 198, 203, 397, & 398. I963. 

It should be noted here that the reference Hook., Bot. Misc. 
1: 168 (1829) is often erroneously cited as "1830''. 

The Arsene ^9 distributed as V. elegans is V. teucriifolia 

Mart. & Gal. 

VERBENA ELEGANS var, ASPERATA Perry 

Additional & emended bibliography: Moldenke, Phytologia 8: 12U 
(1961) and 9: 16, 61, 63, 8U, 86—89, 1^, & 203. 1963. 

xVERBEWA ENGEI2IANNII Moldenke 

Additional & emended bibliography: G.N. Jones, Fl. 111. [1». 
Midi. Nat, Monog. 2:] 216. 19U5; Moldenke, phytologia 8: 120 & 
IU6 (I96I) and 8: 2U6. 1962 j J. D. Poindexter, Trans. Kans. Acad. 
Sci. 65: lao & UI7. 1962 J Moldenke, Phytologia 9: 52, 89—97, 
165, 203— 20U. 215, 219—221, 305, 315, 356, & 359. 1963i Anon., 
Biol. Abstr. k3 (2): B.112. 1963; J. D. Poindexter, Biol. Abstr. 
U3: 397. 1963; Moldenke, RSsumS Suppl. 7: 1 & 9* 1963. 

Shinners 11x79 is said to have had "fls. pale bluish-white", 
but seems otherwise to be typical V. urticifolia L. and is there- 
fore so cited by me. Fassett 20235 is also V. urticifolia and is 
said by the collector to have been growing in a pasture with V. 
hastata L. "and an apparent hybrid". J. Torrey 7U7-0 in the her- 
barium of the New York State Museum at Albany is inscribed •'hy- 
brid between V. urticifolia and V. hastata ", but I am citing it 
hereinafter a8~typical V. urticifolia . Similarly, R. Bebb 2107 
is inscribed as "probably a hybrid form", but I regard it as typ- 
ical V. urticifolia . 

Poindexter (I962) compares xV. engelmannli with its two par- 
ents as follows: Leaf index: V. hastata - 3.2 — 5 (average U.3), 
XV. engelmannii = 2.5—3.7 (average 3), V. urticifolia - 1.9—2.6 
(average 2.2); leaf-shape: V. hastata « lanceolate, xV. engelman- 
nii - broadly lanceolate to lanceolate, V. urticifolia - broadly 
lanceolate to ovate; leaf -base: V. hastata = rounded and petio- 
late, hastately lobed, xV. engelmannii - rounded, decurrent into 
the petiole, V. urticifolia « rounded, decurrent into the petiole; 
leaf-apex: V. hastata - gradually acuminate, xV. engelmannii = 
grsidually acuminate, V. urticifolia » gradually aciominate; stem 
pubescence: V. hastata = strigose, xV. engelmannii - short-hirtel- 
lous to strigose, V. urticifolia « hirtellous; nutlet length: V. 
hastata - 1.5—2 (average 1.7), xV. engelmannii - 1.7—2.2 (aver- 



106 PHYTOLOGIA Vol. 10, no. 2 

age 1.9), V. Tirticifolia » 1.6 — 1,9 (average 1.7) j markings on 
back of nutlet: V. haatata » smooth to faintly striate, xV. engel- 
mannii « moderately ribbed to faintly ribbed or striate, V. urtio- 
ifolia = moderately ribbed j pollen fertility: V. hastata « 30 — 99 
percent (average 79 .S percent), xV» engelmannii » 25 — hi percent 
(average 31.9 percent), V. xirticifolia = 83 — 99 percent (average 
9I4.2 percent); corolla-tube length: V. hastata = 2 — 3.9 (average 
3.5), xV. engelmannii » 3.2 — 3.5 (average 3.3), V. urticifolia - 
1,5 — 2.5 (average 2.1); calyx length: V. hastata = 2.U — 3 (aver- 
age 2.7), xY. engelmannii - 2.1 — 2.7 (average 2.5), V. urticifol- 
ia = 1.7 — 2,3 (average 2); petiole-lateral vein measurement: V. 
hastata = 12 — 25 (average 17.6), xV. engelmannii = 16 — 27 (aver- 
age 21.5), V. urticifolia - Hi — 25 (average 19.6). 

The G. N. Jones 22387 distributed as this hybrid is actually 
■typical V^ urticifolia L., while Eames & Wiegand 12796 distribu- 
ted as "V. hastata x V. urticifolia " is cited by me hereinafter 
under V. urticifolia var. leiocarpa Periy & Femald. 

VERBENA. EPHEDROIDES Cham. 

Additional bibliography: Steud., Norn. Bot., ed. 2, 2; 750. 
iBIa; D. Dietr., Syn. Pi. 3: 602. I81i3i Moldenke, Phytologia 6: 
ll;8 (I96I), 8: 311i St 317 (1962), and 9: 97, II3--IIU, & l5l. 
1963; Moldenke, Biol. Abstr. U3: 6U3. 1963. 

xVERBMA FABRICATA Moldenke 

Additional bibliograplf^r: Moldenke, Biol. Abstr. 30: 1093. 
1956 J Moldenke, Phytologia 8: 120 (I96I) and 9: UU & 296. I963. 

VERBENA FASCICUUTA Benth. 

Additional bibliography; Moldenke, Phytologia 9: llU — 117 & 
39U. 1963 J Moldenke, R^sum^ Suppl. 7: 5 & 9. 1963. 

xVEEBENA FECUNDA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (I96I), 
8: 182 & U37 (1962), and 9: U7— II8. 1963 . 

xVERBENA FERAX Moldenke 

Additional bibliograply: Moldenke, Phytologia 8: 120 (I96I), 
8: Ii37 (1962), and 9: 118, I963. 

VERBENA FILICAULIS Schau. 

Additional & emended bibliography: Moldenke, Phytologia 9: U7, 
80, 119—121, 151, 20I1, & 300. 1963. 

Additional citations: BRAZIL: Santa Catarina: £. £. T_l M^^^^^ 
s ji. (P) . 

VERBENA FUVA Gill. & Hook. 

Additional & eanended bibliography: Hook., Bot. Misc. 1: 170. 
I829j Steud., Nom. Bot., ed. 2, 2: 750. IQIH; D. Dietr., Syn. Pi, 



I96I1 Moldenke, Monograph of Verbena 10? 

3: 603. I81i3} H. S. Marshall, Keir Bull. 1936: 91. 1936; Moldenke, 
Phytologia 8: 123 (1961) and 9: 11 & 121—123. 1963. 

It shoxild be noted here that the reference Hook,, Bot, Misc. 
1: 170 (1829) given above is often erroneously cited as "1830". 

VERBENA GENTRTI Moldenke 

Additional bibliography: Moldenke, Phytologia 8: U87 & U96 
(1963) and 9: 12li— 125. 1963. 

VIEBENA GLABRATA H.E.K. 

Additional synonymy: Verbena glabrata Humb, & Bonpl, ex Steud., 

NCHB. Bot., ed. 2, 2: 7^0. I8IJ.. Verbena glabrata Hurab. & Kunth 
ex D. Dietr., Syn. Pi. 3: 6OI. I8II3T 

Additional bibliograply: Steud,, Nam. Bot., ed. 2, 2: 750. 
I8UI; D. Dietr., Syn. Pi. 3: 60I. I81i3; Moldenke, Phytologia 8; 
311; (I962), 8: U87 (1963), and 9: H, 125—127, & 151. 1963; Mol- 
denke, R^sximS Suppl. 8: 5. 196U. 

VERBENA GUNDULIFERA Moldenke 

Additional bibliographer: Moldenke, Phytologia 9: 127 — 129 & 
397. 1963. 

VERBENA GLUTINOSA Kuntze 

Additional bibliography: Moldenke, Phytologia 9: 122 & 129 — 
131. 1963. 

xVERBENA GONZALEZI Moldenke 

Additional & emended bibliography: Moldenke, Biol, Abstr, 39 J 
61];, 1962j Moldenke, PJ^rtologia 8: 27U & 378 (I962) and 9: 131. 
1963. 

VERBENA GOODDINGII Briq. 

Additional & emended bibliography: Tidestr., Contrib, U.S. 
Nat. Herb. 25: US?. 1925; Moldenke, Phytologia 8: 152 (I96I), 8: 
177, 213, 378, 381, 397, 399, 1;36, & U37 (1962), 8: U65 (1963), 
and 9: 16, 17, 2U, 28, 61, 85, 131— Ha, lli3, ihh, 155, & 201;. 
1963; Moldenke, R6sum6 Suppl. 8: 3. 1963. 

Additional ccmmon names recorded for this plant are "verbena" 
and "vervaim". The G. £, Ikenberry s.n. [May 1, 1937] and Nelson 
- Nelson 5025 , distributed as this species, are actually V, 
irrightii A. Gray. 

Additional citations: ARIZONA: Coconino Co.: Wiebe 27 (Z). Mo- 
have Co.: £. L. Beach 2U (Z); £• T_. Brewer ll; (Z); R. M^ Busta- 
mente 30 (Z), 3U (Z); P_. Marcus 20 (Z). Yavapai Co.: £. L. Beach 
lil (Z); R, M. Bustamente 6U (Z); Clorf 18 (Z); L. R. Fitzgerald hh 
(Z); R. Lj. Richards 19 (Z) . 

VERBENA GOODDINGII var, NEPETIFOLIA Tidestr. 

Additional & enemded bibliography: Tidestr., Contrib. U. S. 
Nat. Herb. 25: U69. 1925; Moldenke, Phytologia 8: 121; (I96I), 8: 



108 PHITOLOGIA Vol. 10, no. 2 

397, U36, & hhO (1962) 8: hSh (1963), and 9: lU, 17, 61, 62, 
135—137, llil— Ui6, 15U, 155, & 20U. 1963. 

xVERBENA. GOODMANI Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 120 (I96I) 
and 9: Ihl & I66. I963. 

ViRBEM GBACILESCENS (Cham.) Herter 

Itaended synonymy; Verbena officinalis var, Q> Hook,, Bot, Misc. 
1: 160. 1829. Verbena gracilescens Cham, ex Hicken, Chloris Plat. 
Argent. 197, in syn. 1910. 

Additional & emended bibliography: Moldenke. Phytologia 8: 317 
(I962), 8: U87 (1963), and 9: 78, 126, Ilt8-~l5li, 20U, & 382. I963; 
Moldenke, R^suml Suppl. 8: 6. 196U. 

VERBEMA GRACILIS Desf . 

Additional & emended bibliography: Steud., Norn. Bot., ed. 2, 
2: 750. iSlili Tidestr., Contrib. U. S. Nat. Herb. 25s 1*69. 1925} 
Moldenke, Phytologia 8: 279 & 1*07 (1962), 8: U61i & U71— U73 (1963), 
and 9: 16, 87, 135, Ha, 1U3, 15U~158, 195, 205, 296, 376, & 379. 
1963. 

Additional citations: ARIZONA: Cochise Co.: A. R. M oldenke 617 
(B). 

VERBENA GINOBASIS var. STRIGOSA Wedd. 

Additional bibliography: Moldenke, Phytologia 9: 159 & I60. 
1963. 

VERBENA HALEI Small 

Additional & emended bibliography: Moldenke, Phytologia 8: 120, 
121, 12li, & lli5 (I96I), 8: 212, 231, & U35 (1962), 8: U72, U77, 
U78, U87, & U88 (1963), and 9: 38, 39, 78, lli7, 160~175, 199, & 
205, fig. 9. 1963; Moldenke, R^sumS Suppl. 7: 1—3. 1963. 

The Mrs. Cottrell 87U3 , Fitzgerald 296, Mrs. A. F. Nelson 8,n. 
[11-2-la], Parks & Cory IOO68 , Tharp"6^ & s.n. [Walnut Or., 7A2/ 
20], and S. E. Wolff 829 & IOI6 , distributed as this species, are 
actually V. xatha Lehm. 

VERBENA HASSLERANA Briq. 

Additional bibliographer: Moldenke, Phytologia 9: 177—178, 367, 
& 368. 1963. 

The jorgensen 3769 collection is a mixture of this species and 
V. tomophylla Briq. 

VERBENA HASTATA L. 

Additional synonymy: Verbena hastata var. pinnatifida Pursh, 

Fl. Am. Sept. 2: I4I6, l8lli. 

Additional & emended bibliography: L., Hort. Upsal. 8 — 9. 171*8; 
J. A. Murr. in L., Syst. Veg., ed. 13, 62. 1771*; Michx., Fl. Bor^ 
Am., ed. 1, 2: H*. 1803; Dum. Cours., Bot. Cult., ed. 2, 2: 623 & 



196U Moldenke, Monograph of Verbena 109 

626. 1811; Pursh, KL. Am. Sept. 2: Ul6. I8lli; Michx., Fl. Bor.- 
Am., ed, 2, "1" [-2]: Hi. I820j Lehm., Del. Sem. Hort. Hamb. 
1826: 16. 1826; Hook., Comp. Bot. Mag. 1: 176. 18 36; Steud., Nom. 
Bot., ed. 2, 2: 750. I8ia; D. Dietr., Syn. Pi. 3: 6OO & 60U. 18U3; 
J. Torr, in Bnory, Rep. U. S. & Mex. Bound. Surv. 2: 128. 1858; 
Selby, Bull. Ohio Agric. Sta. 83: 329. 1897; L. H. Bailey, Botany 
372. I9II; N. Taylor, Mem. N. I. Bot. Gard. 5 [Fl. Vic. N. Y.] 
525. 1915; K. V. 0. Dahlgren, Svensk. Bot. Tidak. 32: 231. 1938; 
Moldenke, Phytologia 8: 120, 121, & 121; (I96I) and 8: I9U, 195, 
207, 2Ui— 2li6, 25U, 257, 279, 280, 316, 317, 380, 382, & 397. 
1962; J. D. Polndexter, Trans. Kans. Acad. Sci. 65: U09, UlO. & 
la2— UI9. 1962; Moldenke, Phytologia 8: U77 & 187— i;89 (1963) and 
9: 51i, 83, 85, 89--9U, 156, 165, 179—181, 190, 203—238, 267— 
283, 296, 356, 358, & 359, fig. 10. I963; Moldenke, R6sum6 Suppl. 
7: 1 & 9. 1963; J. D. Poindexter, Biol. Abstr. k2i 397. 1963} 
Frei & Fairbrothers, Bull. Torrey Bot. Club 90: 352. 1963; G. N. 
Jones, Fl. 111., ed. 3, [Am. Midi. Monog. 7:] 213. 1963; R. A. 
Ludirig, Ind, Sem. Canada Dept, Agr. I96U: 31. 1961i. 

It should be noted here that the reference J. Torr, in Bnory, 
Rep. U. S. & Mex. Bound. Surv. 2: 128 (1858) is often cited as 
•rrorr. Bot. Mex. Bound. 128". The Selby & Hicks, Bull. Ohio Ag- 
ric. Sta, 1*^: pi. U (1906) reference is curious — the copy of 
this bulletin in the library of the New York Botanical Garden 
does not contain such a plate nor amy illustration irhatever of V. 
hastata, but the copy in the library of the Royal Botanic Gardens 
at Kew does contain this plate, nMch is identical iiith the one 
published in Bulletin lii8 (1903). 

Hooker (I836) cites a T. Drummond s.n. [New Orleans, 1833] and 
s.n. [St. Louis] and says "From this the V. paniculata is probab- 
ly not distinct, and the V. urticaefolia seems too nearly allied^I 
The former, of course, is now regarded as a synonym, but the lat- 
ter is most abundantly distinctl 

Poindexter (I962), in his comparison of V. hastata , V. urtici- 
folia , xV. rydbergil , and xV. engelmannii , describes V. hastata 
as having a leaf- index of 3.2 — 5 (average U.3); Leaf-shape lance- 
olate; leaf -base rounded and petiolate, hastately lobed; leaf- 
apex gradually acimiinate; stem pubescence strigose; nutlet length 
1.5 — 2 (average 1.7); markings on back of nutlet smooth to faint- 
ly striate; pollen fertility 30 — 99 percent (average 79.8 per- 
cent); corolla-tube length 2 — 3.9 (average 3*S)i calyx length 
2,U — 3 (average 2.7); and petiole lateral vein measurement 12 — 25 
( average 17 .6) . 

The Brubaker 59, Carleton s ja. [July 12, 1888], E. Dnamnond s. 
n^ [October 1937T7 ii ^i Fogg I8OI3 , G. D. Fuller 5067, Herb. Un- 
iv. Okla. 103li5 , Herb. Unjv. Vermont 9581-9 , and Ripley s.n. [Sep. 
10, 1902], distributed as V. hastata, are all V. urticifolia L.; 
Ashe s.n. [Roan Mtn., July 1893] is V. \irticifolla var. lelocarpa 
Perry & Femald; Havard s.n. [San Antonio] is V. wrightii A. Gray; 
and Ecology Class Univ. Texas s.n. [Palm Grove, 3.1.30], H. B. 



no PHTTOLOGIA Vol. 10, no. 2 

Parks s«n. [Tfhite Creek, 9-7-U7], and Tharp 66? are V. xutha Lehm. 

VERBENA HASTATA f . ROSEA Cheney 

Additional bibliograply: Moldenke, Phytologia 9: 93 21li 220 
& 281—283. 1963. * 

VniBMA HASTATA var. SCABRA Moldenke 

Additional bibliography: Moldenke, Pbytologia 9: 213, 21^, &. 
283— 28U. 1963. 

VERBENA HATEKII Moldenke 

Additional bibliography; Moldenke, Pbytologia 9: 28$ — 286 & 
296. 1963. 

VERBENA HIRTA Spreng. 

Additional bibliography: Steud., Nom. Bot., ed. 2, 2: 700. 
I81a; D. Dietr., Syn. PL 3: 60U. 181^3} Moldenke, Pbytologia 9: 
288—292. 1963. 

The Regnell 1.355 [Herb. Rio Jan. Iii81i9], distributed as this 
species, is actually Lantana camara var, angustifolia Moldenke. 

Additional citations: BRAZIL; Santa Catarina: J. F. T. MtOler 

llii(P). 

VERBENA HCRTA var. GRACILIS Dus6n 

Additional bibliography: Moldenke, Pbytologia 9: 289 & 291— 
292. 1963. 

Additional citations: BRAZIL: Santa Catarina: J, F. T. Mttller 
15U(P). — 

VERBENA mSPIDA Ruiz & Pav. 

Qnended synonymy: Verbena glandulosa Morren ex Steud,, Ncm. 
Bot., ed. 2, 2: 750, ncHii. nud. l81a. 

Additional bibliography: Steud., Nom. Bot., ed. 2, 2: 750. 
I81aj D. Dietr., Syn. Pi. 3: 60U. 18U3; F. Phil., Cat. PI. Vase. 
Chil. 220. 1881: Moldenke, Pbytologia 8: 120, 121. & 121; (I96I), 
8: 200, 201, 25U, 257, 317, & U05 (1962), and 9: Uh, 50, 51, 66, 
llU, 219, 286, & 292—299. 1963; Moldenke, R6sum6 Suppl. 8: 2. 
I96U. 

The Rose & Rose 19072 , distributed as this species, is actual- 
ly V. vil 1 i folia Hayek. 

Additional citations: CHILE: Nuble: Junge 3I8I (W— 2UOU585) . 

VERBENA HOOKERIANA (Covas & Schnack) Moldenke 

Emended synonymy: Verbena crithmifolia ^ minor Gill. &. Hook, 
in Hook., Bot. Misc. 1: I69. I829, Verbena crithmifolia var. 
minor Gill. & Hook, ex Moldenke, R6sum6 363, in syn, 1959* 

Additional bibliography; Moldenke, Phjrtologia 8: 123 (1961) 
and 9: U7, I48, 71, 120, 202, & 299—303. 1963. 

The original description of this plant, by Gillies & Hooker 
(1829) — a reference, by the way, frtiich is often inaccurately 



196U Moldenke, Monograph of Verbena 111 

cited as "I83O" — is " ^ . f oliis angustioribus" . 

VERBENA HUMIFDSA Cham. 

Additional bibliograpl^: Steud,, Norn. Bot,, ed. 2, 2: 750. 
I8UI; Moldenke, PhQrtologia 9: 30>-305. I963. 

J^TIEBENA HTBRIDA Voss 

Additional bibliography: L. H. Bailey, Botany 26, fig. 35. 
I9II; Moldenke, Phytologia 8: 120, 121, 123, & lU (1961), 8: I83, 
257, 280, 316, U27, Ii35, & hh2 (1962), and 9: UO, 87, 90, I60, 
190, 305—336, 351—356, & 367—369. 1963} Moldenke, R6suin6 
Suppl. 8: 5. I96U. 

Additional illustrations: L. H. Bailey, Botany 26, fig. 35 • 
1911. 

xVERBHIA ILLICITA Moldenke 

Additional bibliograply: Moldenke, Phytologia 8: 121 (I96I) 
and 9: 89, 91— 9U, 215, 220, & 356—360. I963; G. N. Jones, Fl. 
111., ed. 3, [Am. Midi. Nat. Monog. 7:] 213, 1963. 

VHIBENA INAMOENA Briq. 

Additional bibliography: Moldenke, Phytologia 8: 255 & 267 
(1962) and 9: 360— 36I. I963. 

VERBENA INCISA Hook, 

Additional synonymy; Glandularia incisa (Hook.) Mold, ex 
Troncoso, Darwiniana 12: 530. I962. 

Additional & emended bibliography; Steud., Nom. Bot.. ed. 2, 
2: 7U9 & 750. ISUI: D. Dietr., Syn. PI. 3: 60U— 605. 18U3} Mol- 
denke, Phytologia 0: 120 & 121. I96I; Troncoso, Darwiniana 12: 
530—531. 1962; Moldenke, Phytologia 9: 70, 315, 330, 33U, 352, 
& 361—371. 1963; Langman, Biol. Abstr. U2: 596. I963; Moldenke, 
R^suml Suppl. 8: U. 1961i. 

Troncoso (I962), in an interesting article entitled "El ori- 
gen del tipo de Glandularia incisa (Hook.) Mold, (sin.: Verbena 
incisa Hook., en Bot. Mag. 65, pi. 3628. I838)" says: "Al des- 
scribir la especie seflala su autor que fue coleccionada por 
Tweedie: 'at Santa Fe, in dry pastures in Porto Alegre Bay (Nos. 
50U y 505) and on the shores of the Panama (No. U60)' . Estudia- 
dos los tres sintipos del Herbario de Kew, pude comprobar que 
la cita 'Panama' es un error de impresi6n. En la etiqueta orig- 
inal del ejemplar No. U60 dice 'this from the coast of the Paira- 
na, flower a soft pink colour' . Se trata, pues, de las costas 
del rio Parani y muy probablemente en la provincia de Entre Rios. 
Glandularia incisa es una especie bastante difundida en nuestro 
literal, principalmente en las barrancas del Paranii." She cites 
Burkart 23U55 & 23U56 in the Darwihion herbarium from Entre Rlos, 
Argentina; 0^ Boelcke U968, Burkart 1276U , Cabrera 7181, and C. 
M. Hicken s.n. [Puerto Nuevo, HI .1912] in the same herbarium frcm 
Buenos Aires. 



U2 PHYTOLOGIA Vol. 10, no. 2 

VERBEM INCISA f . ALBIFLORA Osten & Moldenke 

Additional bibliography: Moldenke, Phcrbologia 9: 366 & 371 — 
372. 1963. 

xVEEBENA INHONESTA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 121 (I96I) 
and 9: 372. I963. 

VERBENA INTEGRIFOLIA Sess6 & Moc . 

Additional bibliography: Moldenke, Phytologia 8: 12li (I96I) 
and 9: 372— 37U. I963. 

xVERBENA IHTERCEDENS Briq. 

Additional bibliograpl^: Moldenke, Phytologia 8: 121 (I96I), 8: 
257 (1962), and 9: 37li~376 & UOU. I963. 

VERBENA INTHiMEDIA Gill. & Hook, 

Additional bibliograpl^: Hook., Bot. Misc. 1: I66 & 167. 1829; 
Steud,, Nom. Bot., ed. 2, 2: 750 & 751. I8U.; D. Dietr., Syn. Pi, 
3: 602 & 603. 181;3} F. Phil., Cat. PI. Vase, Chil, 220. I88I: H. 
S. Marshall, Kew Bull. 1936: 9li. 1936; Moldenke, Phytologia 8: 
2U7, 256, 257, 267, 316—318, Ia2, U27, & UU (1962), 8: li63 & 
U77 (I963), and 9: 67, 151, 151|—156, 296, & 375--38U, I963. 

It should be noted here that the bibliographic reference Hook,, 
Bot. Misc. 1: 166 & 167 (1829) is often erroneotisly cited as 
"1830". Steudel (I8UI) reduces V, gracilis Cham, to V. tenuis 
Sterid., but it is V. gracilis Desf. irtiich is conspecific iiith 
Steudel 's plant, 

xVERBENA KONDAI Moldenke 

Additional bibliography: Moldenke, Phytologia 8: I8I (I96I) 
and 9: 386—387, I963. 

VERBENA UCINIATA (L.) Briq. 

Additional & eeiended bibliography: Hook., Bot. Misc. 1: 168— 
I69. 1829; Steud., Nom. Bot., ed. 2, 1: 581i (I8U0) and 2: 750. 
I8UI; D. Dietr., Syn. PI. 3: 60li. 18U3; F. Phil,, Cat. PI. Vase. 
Chil. 220 & 221. I88I; Baeza, Ncmb. Vulg. Pi. Silv. Chile, ed. 2. 
100, 212, & 269. I93O; Moldenke, Phytologia 8: 123 (1961), 8: 188, 
190, 192, 202, 20U, 212, 379, UOO, lil6, ^17, Ul9, ii20, & U35 
(1962), and 9: 12, lU, 16, U5, U6, 67, 69—72, 76, 87, 88, 117, 
128, 130, 156, 197, 301, 302, 30U, 386, & 388— UOU. 1963. 

Hooker in his Bot, Misc, 1: 168 — I69 (1829) — a reference, 
incidentally, vrhich is often erroneously cited as '•18 30'* — says: 
••19, Verbena erinoides ; caule ascendente ramoso hirto, foliis 
tripartito-laciniatis hirsutis , laciniis lineari-lanceolatis sub- 
dentatis, spicis axLllaribus solitariis laxis, bracteis calycem 
aequantibus patulis. Spreng, Verbena erinoides , 'WiUd. Enum. 
686. 12' . Spreng. Syst. Veget. v, 2. p, 750. Verbena multifida , 
Ruiz et Pav, Fl. Peruv. v. 1. p. 21. t. 33. c. Erinus laciniatus . 
Linn. Sp. PI. p. 879. lychnidea Verbenae temiifoliae folio, vul- 



196U Moldenke, Monograph of Verbena 113 

go Sandla-Laguen . Feill6e Per. v, 3. t. 2$. HAB. Frecjuens in 
provincils Bonarlae, Mendozae et Chile, usque ad alt. 8000 ped. 
Extremely variable in the relative length and breadth of the 
leaves: trifid or multifid, the segments nanrow and linear, or 
sometimes ovate and more or less hairy or hispid. The following 
varieties may be enumerated: a., foliis ovato-lanceolatis inciso- 
serratis vix tidfidis . ^ . foliis profunde trif idls subtripart- 
itisve, laciniis incisis, ultimis ovatis lanceolatisve . (V. mul- 
tifida, Ruiz et Pavon, v. 1. p. 22. t. 23. f. c). V. foliis trl- 
partitis, laciniis inciso-pinnatifidis ultimis lineari-oblongis 
acutis. ^, foliis bipinnatifidis, segmentis paucis linearibus 
elongatis, vel etiam foliorum segmentis angusto-linearibus . [p. 
169:] t . foliis bipinnatif idis , segmentis linearibus brevlbus, 
(bracteis plenmique calyce longioribus. An species propria?), 
g. foliis bipinnatif idis, segmentis brevibus oblongis, (caule 
prostrato foliis pubescenti-incanis. An sp. propr,?). The fig- 
ures of FeuillSe, and of Ruiz and Pavon above quoted, are very 
characteristic of many specimens of oTir plant, but the former 
author says the flowers are scarlet, the latter that they are 
pvirplish flesh colour; whereas ours are blue. Again, we have 
specimens which only differ in the leaves being as little cut, 
or very nearly so, at the margin, as those of V. chamaedrifolia. 
Both these plants are taken into the genus Erinus by Linnaeus, 
both are called Melindres by the Spaniards of South America, 
(the present one M. azules) . From all this it would appear 
that the colour of the flowers, and the more or less deeply in- 
cised leaves, are very variable circumstances. To these, again, 
is very closely allied V^ Aubletia, and still more the V, Lam- 
berti . Nay, Sir James Smith, has in Rees' Cyclopedia ^^en it 
as his decided opinion that Feuill^e's Lychnidea t. 2$, (the 
only authority for the Erinus laciniatus of Linn.) should be re- 
ferred to V. Aubletia : vrhile Mr, Ker, in his Bot, Register, re- 
fers it to V. Lamberti , (his V. Aubletia ) ,» 

Baeza (1930) records the common names "sandialahuen" and 
"hierba del incordio", the latter name also applied to V. palma- 
ta Reiche, 

VERBENA LACINIATA var. CONTRACTA (Undl,) Moldenke 

Additional bibliography: Lehm., Del, Sem. Hort. Hamb. 1826: 
16. 1826; Moldenke, Phytologia 9: 395, 396, 399, & UOl— U03. I963. 

VERBENA UCINIATA var. SABINI (Sweet) Moldenke 

Additional bibliography: Moldenke, Phytologia 9: 39li, 396, & 
U02— U03. 1963 

VERBENA LANDBECKI R. A. Phil. 

Additional bibliography: F. Phil. Cat. Pi. Vase, Chil. 220. 
1881; Moldenke, Phytologia 9: 375 & Ii03— I4OI4. I963. 

VERBENA LASIOSTACHYS Link 



llli PHYTOLOGIA Vol. 10, no. 2 

Additional bibliography: Lehm., Del. Sem. Hort. Hamb. 1826; 16. 
I826i Steud., Ncm. Bot., ed. 2, 2: 750. iBUlj D. Dietr., Syn. Pi. 
3: 602. 18U3; J. Torr. in Etoory, Rep. U. S. & Mex. Bound. Surv. 
2: 128. 1858; A. Gray, Syn. Fl. N. Am. 2 (1): 336. I878; Moldenke, 
Phytologia 8: 121, 12U, & lU2~ll*ii (I96I), 8: 26?, 268, 272, 279, 
J427, & U35 (1962), 9: 16, 83, I66, 21^, 296, 372, & 1;0U— U07 
(1963), and 9: h$9~h67 , I96U. 

It should be noted here that the bibliographic reference J, 
Torr. in Qnory, Rep. U. S. & Mex. Bound. Surv. 2: 128 (1858) is 
often cited as "Torr, Bot. Mex. Bound. 128". Steudel (l81il) af- 
firms that Sprengel regarded V. lamberti Ker as a synonym of V. 
pro strata Ait. 

VERBENA LILACINA Greene 

Additional sjmonymy; Verbena harbisonli Moldenke, Phytologia 
1: Ii38--U39. 19U0. 

Additional bibliography t Moldenke, Phytologia 1: U38 — 1^39 
(19li0) and 1: 511. I9UI; Moldenke, Alph, list Cit. 1: 256. I9I46J 
Hill & Salisb., Ind. Kew. Suppl. 10: 2U2. 19U7i H. N. & A- L. 
Moldenke, PI. Life 2: 63. 19^8; Moldenke, Alph. List Cit. 2: U88 
(19U8> and U: 12U2. 19U9; Moldenke, RSsum^ Suppl. 8: 1 & 5. 196U. 

The type of V. harbisonii is C. F. Harbison s.n. from Agna del 
Refugio, Baja California, Mexico, collected on April 1, 1935, and 
deposited in the herbarium of the San Diego Society of Natural 
History at San Diego. Hitherto I have regarded this and V. lila- 
cina as separate species, but recent material collected by Moran 
on Cedros Island and on the msiinland seems to indicate that the 
two are conspecific. Moran describes the plant as a bushy herb 
or ro\mded shrub 1 meter tall. It has been found on silty flats, 
at altitudes of 10 to 1100 meters. 

Additional citations: MEHCO: Baja California: C. F. Harbison 
s.n. [Agua del Refugio, April 1, 1935] (Du— 2i4li976, Gg— 328000, 
N, N—photo, Sd--ll808, Z— photo); R. V. Moran 8195 (Z) . CEDROS 
ISIAND: R. V. Moran IO669 (Z). 

VERBENA LITORALIS H.B.K. 

Additional bibliograpt^y: Baeza, Nomb. Vulg. Pi. Silv. Chile, 
ed. 2, 233 & 269. 1930. 

VERBENA M/LCDOUGALII Heller 

The Richardson 26 and Hinckley s.n. [June 1936] collections, 
cited below, being from the White Mountains, may have been gath- 
ered in either Lincoln or Otero Counties. Arsene I869I bears a 
printed label inscribed "PL. du MEXIQUE", irtiich is certainly an 
error because beneath it is written "Las Vegas Jardin". V. M. 
Spalding's sxirname is erroneously written "Spaulding" in the Uni- 
versity of Arizona herbarium. 

Curtin (19U7) says "The other two flowers that are called 
dormilon , meaning sleepy-head, are yellow, but this variety is 
purple, and is more often referred to as vervena. Children appar- 



196U Moldenke, Monograph of Verbena 11^ 

ently take the keenest pleasure in Inflicting minor injuries upon 
their playmates; the New Mexican boys have found that one of the 
easiest methods of accomplishing this is by striking an unsuspec- 
ting companion on the bare legs with a whip of vervena . With this, 
it won't be long before he becomes sufficiently angry for all 
purposes of amusement. The plant evidently has some irritating 
hairs that penetrate the flesh aside from the actual impact of the 
blows, and is therefore particularly fine for arousing the fury 
of one's companions. As a diuretic, a tea of the vervena is 
drunk for nine mornings j and, at Mora, for toothache, the green 
leaves are mashed and placed inside the mouth on the gvms and on 
the cheek. When not speaking of plants, the use of the name dor- 
milon applies to a bat." 

Perry (1933) cites the following 23 additional specimens not 
as yet seen by me: WICMIIK}: Laramie Co.: A. Nelson 835U (E) . UTAH: 
Juab Co.: M. E. Jones 6026 (E) . COLORADO: Archuleta Co.: C. F. 
Baker $6S Te, F, G) . NEW MEXICO: Bernalillo Co.: Ellis 2$8 (E) . 
Lincoln Co.: Wooton 208 (E) . San Miguel Co.: Anect $7 (G)j Mul- 
ford 39 (E) J P. C. Standley t;223 (E, F, G) , li927 (E, F, G) . Santa 
Fe Co.: Brandegee s.n. [1889] (E); Greene 77 (N) . County undeter- 
mined: Fendler 597, in part (E) . ARIZONA: Apache Co.: E^ C. Han- 
son A.1U8 (E, F) . Coconino Co.; MacDougal 2U9 (D — isotype, F — 
isotype, — isotype) . She says "This southwestern representa- 
tive of V. stricta is readily distinguished by the short-petiolate 
elongated leaves, the compact spikes, and the floral bracts sur- 
passing the calyx." Her " Greene 77" is probably what is cited by 
me hereinafter as "F^ S_. Earle 77 " . 

In all, 2^1 herbarium specimens, including the type collections 
of most of the names involved, have been examined by me. 

Citations: UTAH: Juab Co.: M. E. Jones 6026 (Ca — 1^9625, N, 
Po— 70877, W— 2li9917). COIiDRADO: Alamosa Co.: Ramaley 15808 (Bl— 
U23UO). Archuleta Co.: C. F. Baker |65 (B, Ca— IOU838, N, Po— 
61i$21, W— 369090); Bethel s.n. [Pagosa Springs, 7-1897] (Fc), s.n. 
[Pagosa Springs, Aug. 20, 1917] (Fc)| SohmoU 1376 (Bl— 1^2333); 
Wooton 2837 (W— 73720$) . Conejos Co.: Ramaley 12998 (Bl— U23la); 
W. A. Weber 7865a (Ca— 961i6U6, Ok, St, W— 205U906) . El Paso Co.: 
Osterhout s.n. [Palmer Lake, J\ily 22, 1895] (N) . Huerfano Co.; 
Shear 3577 (N); Vre eland 636 (N, N) . La Plata Co.: H. Hapeman s.n. 
[Durango, Aug. 29, 1918] (Hp) . Las Animas Co.: Beckwith I38 (N), 
160 (Du— 123189), 166 (Gg— 31383). Mineral Co.: Rama] ey 16531 
(Bl — ii2338) . County xmdetermined: Herb. State Agr. Coll. Colo. 
2027 (Fc). TEXAS: Culberson Co.: Grassl'~175~(Fs, I, Mi). County 
undetermined: 0. M. Clark hhlO [North Jemez Springs] (B). NEW 
MEXICO: Bemalillo~Co . : C. C. Ellis 258 [Capelin Caflon] (W— 
890739), 258 [Las Huertas Caflon] (W— 662673); Koelz s.n. [Sandia 
Mts., August 23, 1926] (Mi); Megard 7U08 [23] (S) . Catron Co.: 



116 PHITOLOGIA Vol. 10, no. 2 

Eggleston 2-260 (W— 1521M8); A. R. Moldenke 1$U (Fg), 156 (Fg), 
629 (B, Fg, S). Chaves Co.: Moldenke & Woods 609 (S). Colfax Co.: 
Berg s.n. [Vermejo River, June 29, '97] (Yc—Wf^); P. C. Stand- 
ley lli22U (N)} Tffooton s.n. [Vemejo Park, Aug. 31, 1913] (N, N, 
W~662229). Uncoln Co.: Cory 33289 (N)j Eggleston 18879 (W— 
1533602) J G. L. Fisher 36122 (Ew, W— 1679367), s.n. [Ruidoso, Aug. 
5, 1931] (Bt— 358U2, Gg— 219620), s^n. [Alto, July 21, 1936] (S1>- 
17379, St); J. H. Grant sji. [Capitan Mts., Aug. 11, 1903] (W— 
U99576); Hinckley 7U7 (Au, N), s.n. [Ruidosa, June 1936] (Fs), 
s.n. [imite Mts., June 1936] (A^, Au)} Richardson 26 (PO--202231) ; 
Steiger II89 (N)j Tucker 3208 (Ok), 3209 (Z); Wooton 208 (Ca— 
IOU839, Ka, N, Po— 70559, Po— 267630, Ur, W— 330U33), s^ [Div- 
ide, June 25, 1895] (W— 562251), s.n. [vicinity of Gilmore's 
Ranch, July 27, 1901] (W— 736878) j Wooton & Standley 3U97 (Du— 
2UI89, Fs, Mn— 689U, Or— 8867, Pi— 36306, W— 5611^30), 3597 (W— 
56211i8), 3651 (Du— 2la78, Hp, Or— 8855, W— 5621U9) . Mora Co.: 
Ar5§ne 19131 (N, Po~li;8790) ; Bacigalupi 585 (Ca— 882663, Du— 
28638U) . Otero Co.: Hershey 3.n. [Mayhill, 8A8/39] (Gg— 310866); 
A, R, Moldenke 639 (S); Moldenke & Woods 607 (S); Orcutt s.n. 
"[Cloudcroft, July 23—26, 1926] (Sd— 2326177 R. B. Randall s.n. 
[Cloudcroft, July 28, 1932] (Sd— 6251i); E. D. Schulz 255 (N), 5U9 
(Wi); V. M. Spalding 111 (Tti- 98880); E. Steams 3h2 (W— 69IOI3), 
358 (W— 690992); Waterfall 12U6i; (Mi, St, W— 2231211); C. B, Wolf 
2778 (Ba, Du— 192373, Gg— 175iA2, Rs— 999U); Wooton s.n. [Cloud- 
croft, June 30, 1899] (W— 736879, We), s.n. [Fresnal, July 21, 
1899] (Po— 70881, Tu), swi. [Toboggan, July 31, 1899] (W— 736880^ 
s.n. [Mescalero Reservation, July 21, 1905] (W— 562250) ; M. S. 
Young s.n. [High Rolls, 7/31A6] (Au) . Rio Arriba Co.: P. C. 
Standley 8239 (W— 687157). Sandoval Co.: Hershey s.n. [Jemez Mts., 
1937] (Bt— 58319); A. Nelson 11556 (Ca— 50072ii) ; A. D. Read 19 
(Tf__890i|07) ; Rousseau 35026 (Un^— 237) . San Juan Co . : KLinger & 
Floiy 265 (Tu— 111520). San Miguel Co.: Anect 211 (Gg— 31382, N), 
299 (Vi); H. S. Barber 152 (W— 56U809); Cockerell s.n. [Upper 
Pecos] (Bl— 112327); Creasy 3.n. [Sept. 27, 1951] (We); B. B. Har- 
ris s.n. [Las Vegas, 6-2U-31] (Nt); G. T^ Hastings s.n. [Las Ve- 
ias, July li, 1927] (N); Lundell & Liindell Il4lt85 (Ld); Mulford 39 
(10— 33U57, Ur); Ramaley 5217 (Bl— li2328); Mrs. St. John s.n. 
[west of Las Vegas] (Po— 70557); P. C. Standley U223 (N, W— 
li98589), U927 (N, W— U98950); Studhalter & Marr I836 (Mi). Santa 
Fe Co.: Bertaud I63 (N); F. S. Earle 77 (N); Eastwood l56l8 (Gg- 
16171;9); Edwards s.n. [Santa Fe, July 10th, I8U7] (T); D. R. 
Goddard 855 (Ca— 500921); McBreen s.n. [July 6, I963] (B); A. 
Nelson llt^6 (S, S); Osterhout 7028 (Po— I83O6I); Pringle s.n. 
[near Glorieta, 26 Aug. I886] (Vt); Renner U3U59 (Ca— 137931); 



1961i Moldenke, Monograph of Verbena II7 

Rose, Fitch , & Parkhurst 17717 (W— 76079U); SagaljTi 8 (Ms). So- 
corro Co.: Clear swu [Los Alamos, July 16, 19l;7] (Gg) . Taos Co.: 
C. C. Albers U7079 (Au); Eggleston 19277 (N, W— 1533909); Water- 
fall 122gO (St); Whitehouse s.n. [Taos, 9.7.1929] (Au); ffhltlng 
10012 (I>t) . Union Co.: P. £. Standley 6065 (W— 68^100) . Valencia 
Co.: Wooton s.n. [mts, west of Grant's Station, Aug. 1st, 1892] 
(W--2lill62) , s.n. [west of Grant's Station, Aug. 2, I892] (C, Ur, 
W— 735991). County undetermined: T. S. Brandegee s.n. [l879]*(Ca- 
16961^7); De Busk 7I69 [Bonito Dam] (Tu--10ii39U) ; laze s^n. [Cas- 
tilla Range, Aug. 16, 1899] (Gg--3138U); Vasey s.n. [N. Mex.] 
(Pa) . ARIZONA: Apache Co.: L. D. Benson 9573, in part (Po— 
267652); Eggleston 17106 (N, Vi); R. £. Ferris 1235 (Du— 9II468), 
101^8 (Ca— 882665, Du— 282088); Frisbie & Bui^ 3 (Fg--7399); S. 
W^ Hutchinson 7331^ (En) ; A^ R. Moldenke 152 (Fg) , 627 (B, S) .""" 
Coconino Co.: Borell s.n. [Sept. 22, l93lT'(Ca— 526337); Collec- 
tor undesignated s.n. [Flagstaff, 7-2li-10] (Fg— 8033); Collom 63U 
(Fg~lllli26), 1^ (W~1729l;51i); Deaver I1I63 (Fg-7332);"o7 ^gi^ 
er 10^75 (N), 1^538 (N); Demaree U2850 (Gg), I16II9 (S); Eaitw^^d~ & 
Ho^eH 6392 (Gg— 261706, w— 1733005); Eggleston 10201 (W— 767199); 
Ellison 1098 (La); Goodding ll;91 (Fg— 803li); E. I TG^eene s.n. 
[Flagstaff, U July I889] (Ca, Ca— 192903, Du— 9II3U); H. cTHanr- 
son II18 (Ur), A.lii8 (Au, Or— 201^90, Up— 8O667); M. E. Jones 605llw 

(PO— 70876, W— 2I499I6), s^n. [Flagstaff, Aug. 29, l88iJ~(P^-^^ 

71017); Kearney & Peebles 12195 (Gg— 267623, To), I3757 (Qg— 
263282, N, To); H, E. Lee sji. [2 miles west of Flagstaff, 9-19- 
36] (ca— 88266Ii, Tu); Leiberg 5882 (W— la0707); Lopmis 32li2 (To); 
MacDougal 21^9 (Ca— 101^8 37— isotype. Dp— isotype, lo— 38757^ 
isotype, N— type, Tu— isotype, W—33li3U6— isotype) ; A^ R. Molden- 
ke 150 (B, Fg, S); MOnachino s.n. [Oct. 10, 1950] (N, Qu);~eI 
"°°^; l£i- [Ft- Valley, 8.7.29] (Au); Nickell s.n. [Aug. 267" 
1956] (St); K^ F. Paricer 5931; (Ca— 736881, N); Riordan 1 (Tu); 
H. H. Rusby 780, in part (Fs, La, Mi, Mi), 6121 (W— lii7588). s.n. 
[Clark's Valley Aug. 1, 1883] (C, Ca-67978, Up-171l8, W- 
56217, W-77l81i6); p. 0. Schallert s.n. [7A8A3] (Ur), s.n. 
[west of Bellemont, l/22/h3] (N); Strickland s.n. [Grand"cIi^on 
July 31, 1951] (Fg— 7359); W. R^ Taylor 3.n. [July 29, 1926] (Gg- 
lii5330); L. P\ Ward 11 (W— IiOii5l6) . Navajo Co.: Pultz 172U (Gg— 
326019). Pima Co.: Spalding 20 (Tu, Tu) . Yavapai Co.: hTh. Rusby 
780, in part (Pr) . County undetermined: H. C. Hanson s^HTNoFth^ 
eastern Arizona] (Bl— U2388). CULTIVATED: New Me^dcoT lrsene 
■^Q^91 (B, N) . LOCALITY OF COLLECTION UNDETERMINED: Baldwin s.n. 
[June 10th, I819] (T). 

VSIBENA MACDOIGALU f . ALBIFLORA Moldenke PJ^rbologia 2: hZh 19li8 
Bibliography: Moldenke, Phytologia 2: k2\x, 191^8; Moldenke, Cas-* 



118 PHTTOLOGIA Vol. 10, no. 2 

tanea 13: 112. 19U8j Moldenke, Alph. Ust Cit. 2: 506 (19U8) and 
3: 839. 19^9: Moldenke, Known Geogr. Distrib. Verbenac., [ed. 2], 
20 & 198. 19U9} Moldenke, R6sum§ 2$ & U72. 1959} Moldenke, R^sumS 
Suppl. 3: 6 (1962) and 5: 3 & h. 1962. 

This form differs from the typical form of the species in hav- 
ing white corollas. 

The type of the fom was collected by Francis Ramaley ( no. 
1681|7) at Alamosa Canyon, Alamosa County, Colorado, at an altitude 
of 8000 feet, on September lli, 1938, and is deposited in the her- 
barium of the University of Colorado at Boulder. The collector 
notes that the plant is "rare". It has been found at altitudes 
of 7800 to 8000 feet, flowering in September. 

In all 3 herbarium specimens have been examined by me. 

Citations: COLORADO: Alamosa Co.: Ramaley 168U7 [Alamosa, Sept. 
Hi] (Bl — U2339 — type). Rio Grande Co.: Ramaley 168U7 [Monte Vista, 
i5.IX.i938] (Ca— 75U136). NEff MEXICO: Otero Co.: Wooton s.n. 
[Fresnal, July 21, 1899] (W— 736877) . 

VERBENA MACDOUGALII mut. ROSEULA Cockerell, Am. Nat. 36: 809 . 1902. 

Bibliography: Cockerell, Am. Nat. 36: 809 . 1902j Periy, Ann. 
Mo. Bot. Gard. 20: 288 & 355. 1933j Moldenke, Prelim. Alph. List 
Invalid Names U7. 19U0j Moldenke, Alph. List Invalid Names U8. 
19U2; Moldenke, R^sumS 369. 1959; Moldenke, R6sum6 Suppl. 5: U & 
7. 1962. . ^ 

This mutant differs from the typical form of the species in 
having pink corollas. 

The type of the mutant was collected by Mrs. 0. Saint John at 
the foot of Baldy Mountain, near Elizabethtown, Colfax County, New 
Mexico, in October, I898, and is deposited in the United States 
National Herbarium at Washington. The collector states that 
purple- flowered specimens were growing at the same locality. 

As yet only the type specimen has been examined by me. 

Citations: NEVT MEXICO: Colfax Co.: Mrs. 0^ Saint John s.n. 
[Foot of Baldy Mt. near Elizabethtown, Oct. 1398] (W— U0U921— 
type). 

VERBENA MACRODONTA Perry, Ann. Mo. Bot. Gard. 20: 289 — 290, pi. 
13, fig. 1— U, & D;. 1933. , . , 

Bibliography: Perry, Ann. Mo. Bot. Gard. 20: 21+9, 250, 260, 
289—290, 355, 358, & 360, pi. 13, fig. 1— U, & Hi. 1933; A. W. 
Hill, Ind. Kew. Suppl. 9: 29U. 1938; Worsdell, Ind. Lond. Suppl. 
2: U86. I9UI: Moldenke, Known Geogr. Distrib. Verbenac, [ed, 1], 
19 & 101 (19U2) and [ed. 2], 33 & 198. 19U9; Moldenke, R6sum6 39 

& U72. 1959. _ , ,. ,. 1 

Illustrations: Periy, Ann. Mo. Bot. Gard. 20: pi. 13, xig. 1— 

u, & Ui. 1933. , ^ ^ 

Coarse plant, with biennial or perennial base; stems 1 — 1.5 m. 
tall, erect, branching, hirsute-hispidulous; leaves large, thlck- 
ish in texture, decussate-opposite; petioles short, 1 — 2 cm. long, 
margined; leaf-blades elongate-ovate, 10— H; an. long, cuneate at 
the base and narrowed into the petiole, coarsely and sharply bi- 



1961i Moldenke, Monograph of Verbena II9 

serrate-dentate with mucronate teeth, rugose with impressed 
veins and scabrous-hirsute with minutely pustulate hairs above 
prominently veined and hirsute beneath; spikes panicled, subtend- 
ed by leafy bracts, dense before anthesis, becoming elongated and 
open in fruit, glandular-hirsute; floral bractlets lanceolate- 
linear, approximately equaling the fruiting-calyx, subulate at 
the apex, ciliate along the margins; calyx 5 mm. long, glandular, 
somewhat viscid-pubescent, the teeth 1.5 mm. long, subulate; cor- 
olla deep-violet, its tube protruding a little beyond the calyx, 
glabrous or pubenilent on the outer surface, the limb 5 — 6 mm. 
wide; cocci trigonous, 2 mm. long, shallowly scrobiculate on the 
upper half, tending to be sulcate toward the base, the ccmmissur- 
al face muriculate. 

The type of this little-known species was collected by Edward 
William Nelson and Edward Alphonso Goldman (no. 7h2$ ) on the road 
from Miraflores to San Bernardo Ranch in the Sierra La Laguna, at 
an altitude of about 750 meters, Baja California, Mexico, on Jan- 
uary 20, 1906, and is deposited in the herbarium of the Missouri 
Botanical Garden at St. Louis. Perry cites also an isotype in 
the United States National Herbarium at Washington. She says 
" Verbena macrodonta is a coarse plant with large thickish leaves 
and open inflorescence, in a measure similar to V. MacDougalii, 
but differing in its less strict habit, somawhatTemote fruits, 
more glandular calyxes, and shorter plumper nutlets." 

Carter describes the plant as "occasional in dry northeast- 
facing slope in decomposed granite with Quercus idonea, Lysiloma 
microphylla , and Jatropha ", at an altitude of 9OO metersT It has 
been collected flowering and fruiting in April and October. Her- 
barium material has been misidentified and distributed under the 
name V. pro strata R. Br, 

Only k herbarium specimens and 1 mounted photograph have been 
examined by me. 

Citations: MEXICO: Baja California: T. S. Brandegee s.n. [Can- 
on San Bernardo, Oct. 13, 1893] (Ca— lOlJgU^; A. Carter~2576 (Ca- 
9I6208, Du— 3Uli053, W— 2023100). MOUNTED CLIPPINGS: Perry, Ann. 
Mo. Bot. Gard. 20: 36O, pi. II4. I933 (ff— photo of type). 

VERBENA MACROSPERMA Speg., Rev. Argent. Bot. 1: 218—220. I926. 

Bibliograplv: Speg., Rev. Argent. Bot. 1: 2lO~220. I926; A. 
W. Hill, Ind. Kew. Suppl. 8: 2J46. 1933; Fedde, Bot. Jahresber. 
59 (2): UI7. 1939; Moldenke, Known Geogr. Distrib. Verbenac. fed. 
1], Uli fie 101 (19li2) and [ed. 2] , IO6 & I98. 19U9: Moldenke. Rfisu- 
m« 127 & U72. 1959. 

Nothing is known to me about this species except what is 
stated relative to it in Spegazzini's original description, nhich 
is repeated herewith: "Leptostachya, perennis; caudice gracili 
subterraneo, multicipe ramis gracilibus radiatim diffusis prost- 
ratisque, ramulis erectis v. adscondentibus, hispido-puberulis, 
foliis oppositis, subconfertis parvis petiolatis, limbo circum' 
scriptione obovato, pinnatim 3— -5-partito, pinnis 2 — 5-fidi3, lob- 



120 PHTTOLOGIA Vol. 10, no. 2 

ulis obtusiusculis planis, margine non revolutis, totis minute 
densiusculeque pubertilo-hirtis , e virescenti subcanescentibus; 
inflorescenti solitariis subseasilibus acrogenis v. alaribus, e- 
rectis, spiciformibus subcylindraceis densiusculis 10 — 20-floris, 
calycibus cylindracso-stibfuaoideis, ^-sulcato-costatis, viridi- 
purpurascentibus dense patentiusculeque hispido-pubenilis, ses- 
silibus, bracteola lineari-lanceolata triple breviore fultis, 
dentibus brevisslmis obtuse calloso-cuspidatis, inaequalibus, an- 
ticis conspicue validioribusj corolla tubulosa ochroleuca, glabra, 
sat exsertaj nuculae pro ratione magnae, subcylindraceae, obtusi- 
usculae, glabrae, dense irregulariterque rug\ilo30-reticulatae, 
castaneo-purpurascentes nitidulae, e calycis dorso longitudinali- 
ter fisso protrudentes . 

"Hab. En los faldeos de las barrancas de los alrededores de 
Talagapa [Rio Negro, Argentina]. 

"Obs, Especie que a primer vista recuerda ciertas formas d© 
la Verbena crithmifolia Gil. & Hk., de la cual se aparta por las 
lacinias de las hojas no lineares y totalmente planas y por los 
dientes del ciliz, principalmente los posteriores, calloso- 
cuspidados y de la Verb, pinnatisecta Schr. por la pubescentia 
tupida y enderezada de todas sus partes . 

"El rizcma debe hallarse plantado bastante profundamente en el 
suelo, siendo casi vertical y leKoso, produciendo a flor de tier- 
ra numerosas ramitas que se extienden radialmente sobre el suelo 
pero sin producir ra£ces adventiciasj estas ramitas ($0—100 mm 
Ing. X 0.50 — 1.2$ mm didun.) son casi cilindricas todas revestidas 
de pubescencia fina enderezada, y en la parte apical se arquean y 
tienden algo a enderezarse, ofreciendo internodios bastante regu- 
lares (8 — 12 mm Ing.) separados por nudos poco o nada hinchados y 
que suelen llevar ramitas enderezadas dibiles (2$ — UO ram alt.) 
tambi^n ceniciento-pubescentes . Las hojas todas hi spi do-pub es- 
centes, opuestas, tienen un peciolo (2,$—$ mm Ing.) semicilln- 
drico al dorso, ligeramente c6ncavo al vientre que lleva la lam- 
ina de circunscripci6n anchamente ovalada (6 — 7 mm Ing. x $ — 7 
mm lat.) generalmente tripartida (rara vez pinado-$-partida) con 
los l6bulos cortamente peciolulados, los dos laterales exterior- 
mente 1-dentados y el impar central trifido, siendo los dientes y 
la extremidad de los l6bulos cuneado-redondeada bastante obtusaj 
las inflorescencias son al principle apicales (y en muchos casos 
asl permanecen), pero lo mAs a menudo se desarrolla en su base de 
inserci6n \ina ramita mis o menos larga que las vuelven entonces 
pleur6genas y axillares, siendo desde la joventud espiciformes 
cillndricas enderezadas (30 mm Ing. x 8—10 mm diim.) llevadas 
por un corto pedicelo ($—1$ ram Ing.) robusto hlspido pubescente y 
formadas por unas 10 a 20 flores recostadas contra el eje y muy 
poco divergentes, yendo cada flor s^sil acompaflada de una brac-t>- 
eita lanceolado-lineal (3,$— U mm Ing. x 0.7$— 1 mm lat.) apli- 
cada contra el c4liz, plana y adelgazada en punta relativamente 
angosta, pero no muy aguda; el ciliz de color verde obscuro mis 
o menos ceniciento por la pubescencia, es cillndrico-subfusoideo 
(7 9 nmi Ing. x 2 mm diim.) ligeramente cuneado-redondeado en la 



196U Moldenke, Monograph of Verbena 121 

base, ligera y suavemente enangostado en la mitad superior, para 
terminar en $ dientecillos, algo diferentes de largo, muy cortos 
y pequeflos, rematados por un callo obtuse que en los dos anteiri- 
ores constituye mucr6n largo bien saliente y visible; la super- 
ficie del ciliz esti recorrida por 5 costillas obtusas (y otros 
tantos surcos) longitudinales que a la madurez suelen tomar col- 
or morado obscuro y se abre de arriba abajo a lo largo de la lln- 
ea mediana dorsal y por alii expulsa las niiculas que habi^ mad- 
urado en ^1; la corola es tubuilosa (12 mm Ing. x 0,7$ mm di5m.), 
superando durante el intesis el cSliz por una tercera parte, os- 
tentando color amarillanto pdlido y $ l6bulos obl6ngos bien ex- 
tendidos y casi igualesj los estambres estdn pegados en la misma 
garganta cuyo 6sculo obstruyen sin sobresalir, con filamentos muy 
cortos y anteras ajnarillas, de bolsas pilinicas poco desiguales y 
absolutamente sin glfindulas apicalesj el estilo es muy delgado- 
lampiflo alcanzando los estambres para dividirse en dos costas 
ramitas estagmSticas; el ovario subcillndrico es lampiflo y verde. 
Las inlculas son h, formando columna cilindrico-tetrigono y al 
henderse el ciliz, se sueltan y caen, siendo as£ cada una casi 
cilindrica (3.5 — U nm Ing, x 1 mm dilm.) convexa al dorso, donde 
ofrece de 3 * 5 estrias longitudinales y algunas reticulaciones, 
algo lustrosas, y slI vientre ostenta una angosta Hnea longitud- 
inal hundida y lisa; la extremidad superior es redondeada obtusa, 
la basal cortada en bisel." 

VERBENA MALMII Moldenke, Phjrtologia 2: klS—klS, 19U8 . 

Synonymy: Verbena malmei Moldenke ex Angely, Fl. Paran, 16: 
78. i960. 

Bibliography: Moldenke, Phytologia 2: U75~U76 . 19U8j Molden- 
ke, Known Geogr. Distrib. Verbenac,, [ed. 2], 9U & 198. 19U9; 
Moldenke, Phytologia 3: U5U. 19^1; E. J. Salisb., Ind. Ken. 
Suppl. 11: 263. 1953i Moldenke, Inform. Mold. Set U8 Spec, [h] . 
19Shi Angely, Fl. Paran. 7: 13. 1957; Moldenke, R^sumS 110 & 1^72. 
1959; Angely, Fl. Paran, 16: 78 (I96O) and 17: U6. I96I; Molden- 
ke, R^sum^ Suppl. 3: UO. I962. 

Shrubby perennial plants to 3 m. tall, much-branched; branches 
and branchlets green, acutely tetragonal, completely glabrous 
throughout, shiny, often sulcate between the angles; trdgs num- 
eix3us, ascending-erect, slender, acutely tetragonal, green and 
shiny, glabrous; principal intemodes 2 — 5 cm. long; nodes not 
annulate; leaves very tir^ and indistinct, giving the plan an a- 
phyllous appearance, sessile, oblong or linear, rather firm- 
textured, uniformly bright-green on both surfaces, decuasate- 
opposite, ascending, 3 — 8 mm. long, about 1 mm. iride, appressed- 
strigillose on both surfaces, aujute at the apex, 1- veined, the 
midrib slightly elevated beneath and subimpressed above; infloi>- 
escence spicate, very abundant, usually in groups of 3 at the tip 
of each twig, often aggregated in panicvilate fashion, the termin- 
al spikes usually short-pedunculate, the lateral ones longer- 
peduncvilate, the floriferous portion elongating to almost 3 cm, 
after anthesis; peduncles very slender, glabrous or minutely 
strigillose, 3-~15> mm, long, tetragonal, green; rachis rather 



122 P H T T L G I A Vol. 10, no. 2 

densely strigillose-puberulent with whitish hairs especially vis- 
ible after the calyxes have fallen off; bractlets lanceolate, very 
small and obscure, about 1 mm, long, strigose with subappressed 
antrorse urtiitish hairs, acute at the apexj calyx tubular, about 3 
mm. long, densely white-strigose with appressed antrorse hairs, 
the rim 5-apiculate; corolla blue, its tube about k mm. long, 
glabroiis except at the very apex where it is densely white-stri- 
gose like the calyx, its limb 3 — h mm. wide, pubervilent in the 
throat within and strigose at the base outside, the lobes glab- 
rous on both surfaces. 

The type of this species vras collected by Gustaf Oskar Anders- 
son Malme ( no. lUil ) — in irtiose honor it is named — in a swamp 
at Villa Rica7 Rio Grande do Sul, Brazil, on January 22, 1902, 
and is deposited in the herbarixan of the Naturhistoriska Riksmus- 
e\im at Stockholm. The species is obviously closely related to V. 
alata Sweet and V. ephedroides Cham, and more intensive study is 
required of these taxa when more mateidal is available. It appears 
to inhabit shrubby marshes, flowering in December and January. 
Herbarium material has been mlsidentified and distributed as V. 
isabellei Briq. 

In all, 9 herbarium specimens, including the type, and U 
mounted photographs have been examined by me. 

Citations: BRAZIL: Paranfi: Dus6n 2788 [Herb. Mus. Nac . Rio Jan. 
U6569] (N); Hatschbach 2^2 (N), 3761 (Z) . Rio Grande do Sul: 
Malme lllj (F — ^photo of type, N — isotype, N — photo of type, S — 
type. Si — ^photo of type, Z — photo of type); Rambo 3825U (N), 
39352 (N), $0026 (Le, S). 

VERBENA MARITIMA Small, Bull. N. T. Bot. Card. 3: U36. 1905. 
Synonyny: Glandularia maritima Small, Man. Southeast. Fl. 
1138 & 1508. 1933. Glandularia maritima (Small) Small apud A. W. 
Hill, Ind, Kew. Suppl. 9: 12li. 1938. Verbena aubletia var. mar- 
itima Curtiss ex Moldenke, Addisonia 21: 60, in syn. 19U2. 
GrandTJLaria maritima Small ex Moldenke, Alph. List Invalid Names 

Suppl. 1: 10, in syn. 19U7. 

Bibliography: Small, Bull. N. Y. Bot. Card. 3: ii36. 1905; Fed- 
de in Just, Bot. Jahresber. 33 (1): 632. 1906; Prain, Ind. Kew. 
Suppl. 3: 187. 1908; Small, Fl. Miami 159. 1913; Small, Fl. Fla. 
Keys 128. 1913; Moldenke, List Spec. Mold. Southeast. Set 10. 
1933; Small, Man. Southeast. Fl. II38 & 1508. 1933; Perry, Ann. 
Mo. Bot. Gard. 20: 2U8, 311, 320—321, & 355. 1933; J. A. Harris, 
Physico-chem. Prop. Plant Saps 173. 193U; A. W. Hill, Ind. Kew, 
Suppl. 9: I2U. 1938; Moldenke, Annot, & Classif . List IO8. 1939; 
Moldenke, Prelim. Alph. List Invalid Names 26. 19 UO; Moldenke, 
Alph. List Invalid Names 25. 19U2; Moldenke, Addisonia 21: 59 — 
60, pi. 702. 19ii2; Moldenke, Known Geogr. Distrib. Verbenac, 
[ed. 1], 5 & 101. 19U2; Schnack & Covas, Darwiniana 6: U75. 19hU; 
Moldenke, Bot. Gaz. IO6: 159. 19U5} Moldenke, Am. Midi. Nat. 32: 
576. 19U5; Moldenke, Am. Joum. Bot. 32: 6O9. 19li5j Moldenke, 
Alph. List Cit. 1: 22, 138, 139, 152, 155, 267, 279, 291, & 29U. 



196U Moldenke, Moiaograph of Verbema 123 

19h6; Moldenke, Alph. List Invalid Names Suppl. 1: 10 & 22. 19U7; 
Moldenke, Phytologia 2: 238— 2U0, fig. 2. 19U7; Moldenke, Alph. 
List Cit. 2: 393, UOO, U09, li69, U9U, 510, 512, 513, 5l8, 523, & 
57U (19ii8), 3: 660, 71?, 720, 721, lla, 755, 777, 778, 800, 851, 
852, 931. 9U--9Ui, & 9U8 (19l;9). and h: 1112, 1117, 1118, 1138, 
1177, 1188, 1192, 1193, 1216, 121i3, & I296. 19li9: Moldenke, Known 
Geogr. Distrib. Verbenac., [ed. 2], 10 & I98. 19U9; H. N. & A. L. 
Moldenke, Anal. Inst. Biol. Mex. 20: Hi. 19U9; Moldenke, Am. Wild 
Fls. 292 & U50. 19U9j Moldenke, Inform. Mold. Set 51 Spec, h* 
1956; Moldenke, R^sumS 13, 296, 297, 358, U2U, & U72. 1959; Mol- 
denke, R6sum6 Suppl. 1: 2 (1959). 3: 3 & 32 (1962), and U: 3. 
1962; Moldenke, Phytologia 8: li3U & U35 (1962) and 9: 19U. 1963. 

Illustrations: Moldenke, Addisonia 21; pi, 702 (in color). 
l9U2j Moldenke, Phytologia 2: 239, fig. 2. 19li7. 

Herbaceous perennial plants, decumbent or trailing; roots fib- 
rous; stems severaQ. or nimieroua, slender, decumbent or prostrate, 
creeping or trailing, branched at the base, sparingly pubescent 
or glabrate; branches smooth or spsuringly pubescent; leaves de- 
cussate-opposite, numerous, thick- textured; petioles 2 — h mm. 
long, smooth or sparingly pubescent; leaf -blades bright-green, 
cuneate to orbicular-ovate or obovate, 1—6 cm. long (usually 1 — 
h) , 1.2 — 2.5 cm. wide, tapering regvilarly in cuneate fashion to 
the margined petiole, incised-dentate or saneirtiat lobed, broadly 
obtuse or acute at the apex, sparsely pubescent or glabrate on 
both surfaces; inflorescence terminal, the spikes pedunculate, 
subcapitate or fascicle-like during anthesis, becoming elongated 
to as much as 7 cm. in fruit, densely many-flowered; flowei^ 
showy, blooming practically throughout the yesir in terminal flat- 
topped clusters; bractlets persistent, lanceolate or linear- 
lanceolate, green, about half as long as the calyx, acuminate at 
the apex, pubescent on the back, ciliate^nargined; calyx slender, 
tubular, 10 — 12 mm. long, about 2 mm, in diameter, appressed- 
pubescent with often glandular hairs, conspicuously 5-ribbed, its 
rim 5-toothed, with short, slender, subulate, unequal teeth, 2 of 
which are about 2 mm. long, the 3 others about 1 mm, long; corol- 
la hypocrateirLform, varying from blue Or blue-purple to purple, 
pinkish-purple, lilac-purple, rose-purple, or lavender, the lower 
surface much paler than the upper, its tube narrowly cylindric, 
1 1/2 to 3 times as long as the calyx, finely or densely pubes- 
cent on the outer surface, the limb wide-spreading, deeply 5- 
parted, 10 — 15 mm. wide, the lobes obovate, deeply notched at the 
apex; stamens U, inserted in pairs at two levels near the mouth 
of the corolla-tube, entirely included; filaments slender, very 
short; anthers small, oblong, with or without glands; pistil 
single, compound; style slender, smooth, 1.6 — 1.8 cm. long, short- 
ly 2-lobed at the apex, the posterior lobe smooth and non-stig- 
matiferous, the anterior lobe broader, papillose, and stigmatic; 
ovary Li-sulcate, U-celledj fruiting-calyx 10 — 13 mm. long, appres- 
sed-pubescent on the outside, often glandular, the teeth short, 
slender, subvilate, enclosing the fruit; cocci U, subcylindric , 
dark-brown, crustaceous, 1-seeded, about U mm. long, scrobiculate, 



12U PHYTOLOGIA Vol. 10, no. 2 

Tfith a broadened base, the commissural surface narrow and muri- 
cately roughened. 

The type of this handsome endemic species was collected by 
John Kunkel Small and Joel Jackson Carter ( no. 1077 ) between Cut- 
ler and Longview Camp, Dade County, Florida, between November 9 
and 12, 1903, and is deposited in the Britton Herbarium at the 
New York Botanical Garden. The species is found chief ]y in the 
coastal counties of Florida from Monroe north to Flagler and 
Saint Johns and on the west coast to Lee, vrith a record from Lake 
County in the interior. Typically it inhabits sandy ridges bor- 
dering the ocean, sand-dunes, and even the beaches themselves, 
but occurs also in coastal and dune hanmocks, low pinelands, 
flatwoods, and kitchen-middens, prefeiring sandy or dry sandy 
soil. It has been collected in virgin glade soil, in the vicin- 
ity of hammocks, in pinelands, along roadsides and sandy roads, 
on sandy beach ridges, and on rocks in tropical pinelauids, in 
anthesis from November to August, and in fruit in Janusiry, Mai^;h, 
April, Ma, and July. The only recorded popular name is "beach 
vervain" . 

O'Neill found 32 plants of this species per acre in Flagler 
County in 1929. Atwater reports it "ccramon" in pinewoods and 
glades in Dade County, while Ward & West say that it forms "mats 
on landward surface of coastal dunes 100 yards frcxn the ocean" in 
Flagler County. Young found it growing "abundantly 200 feet from 
the ocean". The specific epithet is sometimes written with a 
capital initial letter for no valid reason, 

Blazic describes the plsmt as "a native verbena brightening 
the landscape at edge of woods even in winter". Sheehan says "it 
grows in prairies and pinelands, used as an antidote [by the Sem- 
inole Indians] for the bite of the water-moccasin; a tea is made 
from the flowers and drunk hot." Hands records the species fixim 
Lake [Eldorado] and Saint Johns Counties [Matanzas], as well as 
from Mosquito Inlet, Volusia County. The specimens used by him 
(193U) in his study of the physico-chemical properties of the sap 
came from a sandy ocean beach near the Inside Route Canal at Hal- 
lendale, Broward County, 

Herbarium material of this species has been misidentified and 
distributed under the names Verbena aubletia L., V. tampensis 
Nash, Glandularia tampensis (Nash) Small, and even Phlox ampli - 
folia Britt. in the Polemoniaceae . The B^ H. Patterson s.n. 
[Daytona, Nov, 28, 1917] cited below has its upper leaves much 
like those seen in V, tampensis or some forms of V, canadensis 

(L,) Britton. 

The species is apparently closely related to the widespread 
and extremely polymorphic V. canadensis of inland portions of the 
United States, irtiich occurs in the more northern counties of 
Florida. The ranges of the two species overlap in Brevard, Flag- 
ler, and Volusia Counties, where intermediate, probably hybrid, 
specimens have been fo\md. 

The species was apparently first collected and recognized as 
distinct by A, H, Curtiss, t^io gathered it on "sand ridges border- 



196U Moldenke, Monograph of Verbena 125 

ing the ocean [near Cape Canaveral], E. Florida, July [1879]" and 
distributed it under the name " Verbena Aubletla var, maritima " . 
This name he published on the printed labels of his first distri- 
bution of North American plants (no, I963 ) and in the advertising 

leaflet concerning it, but failed ever to validate it by a formal 
description. A note on Curtiss 1963*- in the United States Natio- 
nal Herbarium says "not _V. aubletia (G. V. N[ash]), belongs to | 
Nobiles. Anther connective not appendagedj corolla lobes emargin- 
ate." Actually, Curtiss I963 is a mixture — the part from "san- 
dy ridges bordering the ocean, eastern Florida" is V, maritima , 
but the part frcan "roadsides near Charleston" is V. bonariensia 
L., while the part frtm "low open woods, Indian River, June" is 
V. tampensis Nash. 

Because of its showy inflorescences and long blooming season, 
it seems probable that this species woxild prove itself a valuable 
horticxiltural subject, especially in seaside gardens in subtrop- 
ical and tropical regions, along the coasts of Florida and our 
other Gulf States, and southern California. 

Perry (1933) records the following 8 additional specimens not 
as yet seen by me: FLORIDA: Brevard Co.: Ctirtiss I963* (E, G), 
5706 (E, G)j O'Neill 6309 (E, W) . Dade Co.: N, L. Brit ton 220 
(F) . Martin Co.: Randolph $2 (G) . She says "This native of the 
sand dvuies and the hammocks of Florida resembles V. canadensis in 
inflorescence, but is readily distinguished by the creeping habit 
and the cuneate or orbicular-ovate leaves." 

in aill, 177 herbarium specimens, including type material of 
all the names involved, and 1 mounted clipping have been examined 
by me. 

Citations: FLORIDA: Brevard Co.: F. S_. Blanton ^309 (I, N, S, 
W— lli85U67), 6U79 (I); A. B. Burgess 638 (N)j Curtiss I963* [san- 
dy ridges bordering the ocean, eastern Florida] (Be, C, Cm, I, Pa, 
Up— 17083, Vt, W— 719UO, W— I3230U8), 5706 (Al, Ca— 10U865, Dt, 
Fc, Fl— 20986, lo— 38761, N, Po— 6l46ii9, Ur, Vt, W— 280798); 0. 
Degener s.n. [Tropic, Aug. 15, 1933] (Ba)j Hotchkiss s.n. [Feb. 
5, 1935] (N); McFarlin 3871 (Gg— 237853), 66O8 (N); O'Neill s.n. 
[June 11, 1929] (I), s.n. [July 8, 1929] (I), s.n. [south of Co- 
coa Beach, August 9, 1929] (Fl~20976, I, W— 1U885U6)} W. ][. 
Rhoades s.n. [Cocoa, 12-8-27] (Fl~20993) ; P. 0. Schallert 20869 , 
in part [Cocoa Beach] (Hi — 55688, S, Ur) ; U, C. Smith s.n. 
[Georgiana, Jany. 31, '91] (Up — 17078). Broward Co.: C. C. Deam 
60837 (On, N)j J. P. Young 737 (W— 12U0563) . Collier Co.: Haw- 
kins s.n. [Royal Palm State Park, 1-25-28] (Fl~20990)j Sheehan 
s.n. [Godden's Mission, March 7, 1919] (N), s.n. [Leaning Oak, 16 
miles east of Imnokalee] (N); £. K^ Small 8123 (Ca— 79699U) . Dade 
Co.: Atwater M.225 (Hi— 167652); Bailey & Bailey 6278 (Ba, Ba), 
6388 (Ba); Blazic s.n. [Miami, Jan. 1922] (Gg— 31ii09); N. L. 



126 PHYTOLOGIA Vol. 10, no. 2 

Brltton 220 (N), ^ (N)j Busirell s.n. [April 3, 1912] (Bu)j C. 
C. Beam 60la7 (Dm, N), 609UO (Dm, N); A. A. Eaton $h6 (Rf)} A. P. 
Garber s.n. [Miami, May 1877] (Vt), 8.n. [Miami, Jvdy 1877] (Pa); 
Henderson s.n. [Cape Florida] (T); Herb. Columbia Univ. s.n. 
[Cape Fla.] (C); Htmnewell ^83$ (Ua— 36988); B. McAllister 27 (H- 
Ulli72); H. N. Moldenke ^k9 (E, Go, N, S, Up, Ur, W— l567361i), 586 
(E, Go, H— 5U7li, N, N, Ob— 83132, S, Up, Ur); Mulvania 12 (Hp); 
O'Neill 7596 (Bt— 171^5, DU--255707, En, Gg--237852, Gg— 238205, 
Hp, Hp, I, N, N, St— 9251, Um— 23, Ur, W— I6OI78I), s.n. [Jan. 30, 
1933] (I); B. H. Patterson s.n. [Feb. 7, 1918] (Cm, Cm); S afford 
& Mosier 210 (W— IO36050); J. K. Small 2100 (N), 8123 (N), 859U 
(N), 8599 (N), 8636 (Gg— 316099), s.n. [beach opposite Miami, No- 
vember I90U] (Ur); Small & Carter 1077 (It—isotype, N— type), 
299h (N), s.n. [Janviary 16, I909] (We); Small , Carter , & Small 
3311 (N), s.n. [February 1911] (H— U3l5i+, PI— 131511^); Small & 
Small $U22 (Fl— 27706, N, S, W— 1737919), s.n. [July 9, 1915] 
(N); Small & Wilson I96I (N), s.n. [May 6th to 9th, I90U] (Vi); 
Weber & Hawkins s.n. [Homestead, 3-1-28] (Fl) . Flagler Co.: 
O'Neill s.n. [August 7, 1929] (I); J. K^ Small 10372 (Hi— 7U2U); 
Ward & West 13U2 (Hi— 182203); West & Arnold s.n. [Flagler Beach, 
10/10/liO] (Fl— 32757). Indian River Co.: R. J. Lemaire I89 (Hi— 
120562); P. 0. Schallert 20869, in part [Vero Beach] (B, Ur, Ws); 
Small , DeWinkeler , & Mosier 11123 (N), s.n. [April 3, 192U] (It, 
Mi). Lee Co.: J. K. Small 831*7 (N) . Palm Beach Co.; Bailey & 
Bailey 6523 (Ba, Ba) ; A. B. Burgess 783 (N) ; W. B_. Fox 3.n. [Del- 
ray Beach, Apr. 28, 19U5] (No— 15822, We); W. H. Rhoades 6 (W— 
I53ii806), s.n. [near Palm Beach] (Hs, Hs); £. K. Small 212U (N), 
8509 (Go, lo— Iii5l63, It, N); Small , Mosier , & DeWinkeler IO89I 
(S, Up); E. West s.n. [Jupiter, 5-12-33] (Fl— 20979) . Polk Co.: 
McFarlin UI86 (Mi). Saint Lucie Co.: Brass 20397 (W— 2066056); 

A. B. Burgess 713 (N); H. N. Moldenke 21^82 (Bs, Hk, Ito, Ok, Sm, 
Ss, Z) . Seminole Co.; P. 0. Schallert I2I46 (Je — 7022). Volusia 
Co.; H. Cj. Beardslee s.n. [New Smyrna, Feb. 1925] (Ob — 97288); 

B. H. Patterson s.n. [Daytona, Nov. 28, 1917] (Cm); J. K_. Small 
B^7ir (N); Small & DeWinkeler 9856 (Mi) . Biscayne Key: Lightfoot 
s.n. [Key Biscayne, Apr. 28, 1917] (Ba) . Jupiter Island [Martin 
& Palm Beach Counties]: Cooley , West , & Daggy U8l7 (Hi— 193302); 
Small & DeWinkeler 9865 (W— l92U33ii) . Merritt's Island: Hotch - 
kiss s.n. [Feb. 5, 1935] (W— 1683053); McFarlin 3871 (Mi); H^ N. 
Moldenke 219a (N, Up, Ur) . Covinty undetermined: Curtiss s.n. 
[Fla.] (C); Herb. LeRcy 3.n. [Fla.] (C); Cj^ H^ Hitchcock s.n. 
[Oak Lodge] (Dt); "W. M. R." 3.n. [Febr. 18, '53] (Pr) . MOUNTED 
CLIPPINGS; Bull. N. I. Bot. Gard. 3: U36. 1905 (W) . 



1961; Moldenke, Monograph of Verbena 127 

VERBENA MARRUBIOIDES Cham., Linnaea 7: 269. 1832. 

Synonymy: Verbena chamaedrifolia var. melindroides Benth. ex 

Moldenke, Alph. List Invalid Names Suppl. 1: 23, in syn. 19U7 
[not V. chamaedrifolia var. melindroides (Cham.) Schau., 19U0, 
nor Schau., 19U7] . Verbena hiimifusa var. reticulata yoldenke, 
Phytologia 2: U23. 19U8. Verbena humifusa var. verticillata Mol- 
denke ex Stellfeld, Trib. Farmac . 19 (10): 166. 1951. 

Bibliography: Cham,, Linnaea 7: 269. 1832; Steud., Nom. Bot,, 
ed. 2, 2: 750. I81a: D. Dietr., Syn. Pi. 3: 602. 18U3; Walp., 
Repert. Bot. Syst. h: 27. I8h5; Schau. in A. DC., Prodr. 11: 536. 
I8ii7; Schau. in Mart., Fl. Bras. 9: 18U. l85l; Briq. in Engl. & 
Prantl, Nat. Pflanzenfam. h (3a): lii7. I89U} Jacks, in Hook. f. 
& Jacks., Ind. Kew. 2: 1179. 1895; Herter, Florul. Urug. 105. 
I93O; Herter, Revist. Sudam. Bot. 6: 97. 1939; Moldenke, Knomi 
Geogr. Distrib. Verbenac, [ed. 1], 35, 39, Ul, hh, & 101. 19U2; 
Augusto, Fl. Rio Grande do Sul 209 & 232. 19U6: Moldenke, Alph. 
Ust Git. 1: 83, 96, & 251 (I9U6) and 2: 358, hh2, U58, 532, 533, 
537, & 599. I9U0; Moldenke, Castanea 13: 117—119. 19iiO; Molden- 
ke, Phytologia 2: li23 (19U8) and 3: 75. 19^9; Moldenke, Known 
Geogr. Distrib. Verbenac, [ed. 2], 73, 9h, 100, IO6, & I98. 
19U9; Moldenke, Alph. List Git. 3: 673, 676, 688, 7U5, 780, 837, 
8liO, 873, 921, 922, & 923 (19ii9) and U: 1010, IO9O, & 1250. 19U9; 
Stellfeld, Trib. Farmac. 19 (10): 166 & I67. 1951; Rambo, Sellow- 
ia 6: 60, Qh, & 153. 195U; Angely, Fl. Paran. 7: 13. 1957; Mol- 
denke, R$sum6 85, no, 119, 127, 361, 366, & U72. 1959; Angely, 
Fl. Paran. 16: 78 (I96O) and 17: U6. I96I; Reitz, Sellowia 13 
(13): 110. I96I; Moldenke, R6sum5 Suppl. 3: 15 & 38. 1962; Mol- 
denke, Phytologia 9: 128, 289, 301;, 305, 315, & 367. 1963. 

Creeping herb; stems cespitose, tetragonal or rather terete, 
to about 1;3 cm. long, mostly simple, ascending, villous-hirsute; 
leaves subsessile or sessile, more or less approximate, variable 
in size, mostly longer than the intemodes, with a subcanescent 
aspect; petioles, when present, very short and winged; leaf- 
blades chartaceous, cxineate-obovate or obovate to elliptic or 
suborbicular, 0.7 — 3 cm. long, 0.6 — 1,5 cm, wide, usually acute 
or subacute at the apex, obtuse or rounded on smaller leaves, 
coarsely dentate or crenate-serrate frcoi the widest part to the 
apex with acute or obtuse rather regular teeth, not lobed, cune- 
ate or subacuminate and entire toward the base, narrowed into a 
very short winged petiole or subsessile, subrevolute along the 
margins, lineate-rugose, pustulate-bullate and strigose-hirtous 
above with white appressed antrorse hairs, canescent-tomentose 
beneath, strigose to rather densely spreading-hirsutulous or sub- 
sericeous-hirsutulous on the venation beneath, the midrib, sec- 
ondaries, and veinlet reticulation deeply impressed above and 
very uniformly prominent beneath; spikes terminail, solitary, of- 
ten 10 or more per plant, 5 — 7.5 cm, long, finally elongate and 
to 30 cm, long; bractlets subulate-lanceolate, half sis long as 
the calyx, ciliate-^nargined; flowers patulous, subopposite, 
paired but at least the lower ones alternately separated; calyx 
tubular, about 10 mm, long, venose, canescent-hirtous and sub- 



128 PHITOLOGIA Vol. 10, no. 2 

glandiilose, often colored, the veins broad and prcaninulent, the 
teeth somewhat iinequal, narrowly subulate, connivent but scarcely 
twisted after anthesis, the indument unequally dense, soft to the 
touch, almost shir^r in aspect, with glanduliferous hairs inter- 
spersed; corolla violet or bluish-violet to magenta or rose-purple, 
pubescent or glabrous on the outer surface, villous within, bar- 
bate in the throat, its tube slightly and gradually aB?)liats up- 
wards, three times as long as the calyx, the limb broad, 5-fid, 
the lobes rounded, undulate, emarginate at the apex; stamens suid 
style equaling or slightly surpassing the calyx; fruit as in V. 
platensis Spreng, 

The species was based on a Sellow collection from Brazil, de- 
posited in the herbarium of the Botanisches Museum in Berlin, 
where it was photographed by Macbride under his photograph no, 
I7U29, but is now destroyed. Schauer (1351) says "in canpis Bras- 
iliae meridionalisj in campo d'Utra aliisque in locis legit. 
Sellow". The type of V. humi fusa var. reticulata was collected 
by Per Karl Hjalmar Dus^n ( no. l^TlU ) in a campo between Lago and 
Desiro Ribas, at an altitude of 800 meters, Parand, Brazil, on 
October 22, 191i|., and is deposited in the herbarium of the Natur- 
historiska Riksmuseum at Stockholm. Briquet (I89U) places the 
species in Section Yerbenaca , Subsection Nobiles . The names V. 
chamaedrifolia var. melindroides Schau. and (Cham.) Schau. are 
synonyms of 7. peruviana (L.) Britton. 

Verbena marrubioides has been collected in campos, roclcy sunny 
dry campos, and dry gras^ campos, in fields, in pedragal, and in 
dry riverbeds, at altitudes of ^0 to 8OO meters, flowering from 
August to November and in January. Herbarium specimens have been 
misidentified and distributed under the names V. erinoides L., V. 
hirta Spreng., V. humifusa Cham., and V. melindroides Cham. On 
the other hand, "the Herter 1000 [Herb.~Herter 62763] distributed 
as V. marrubioides , is actually xV. osteni Moldenke. The records 
of V. marrubioides from Catamarca and Jujuy, Argentina, given by 
me in previous publications are based on mis identifications of 
specimens which prove to be V. incisa Hook, and V. peruviana (L.) 
Britton. 

Schauer (l85l) comments "Variat praeter magnitudinem folionm 
etiam crenis serraturisve latioribus magisque rotundatis et an- 
gustioribus subacutis, dein indumento modo magis patente hirto- 
que, modo magis appresso molli subsericeo. Differt a V. scordioide, 
cui proxima: foliis basi triangulari cuneata integerrima sessili- 
bus antice tantum serratis, spicis demum subdissitifloris paten- 
tibus, calyce dentibus tenuoribus longioribusque, corolla tube 
longe breviori." 

The species differs from V. hvuaifusa in having its leaf-blades 
varying from elliptic to obovate or suborbicular, 0.7 — 3 cm. long, 
0,6 — 1.5 cm, wide, usually acute at the apex, cuneate or subacum- 
inate at the base, coarsely dentate from the widest part to the 
apex with acute or obtuse rather regular teeth, not lobed, pustu- 



196U Moldenke, Monograph of Verbena 129 

late-bullate above and rather abundantly hairy -with white appres- 
sed antrorse hairs, rather densely spreading-hirsutiilous on the 
venation beneath, the midrib, secondaries, and vcinlet reticula- 
tion deeply impressed above and very uniformly prominent beneath. 
Henz collected the species in a region of 1.5 meters rainfall 
and 5 — U0° C. temperature range. Osten says for his no. 3133: 
"aff . teucrioides sed fl. violaceisj cf . humifusa? differt a di- 
agnosi in DC XI $38 caulibus tetragonis nee teretiusculis, calyce 
glanduloso (nee subglanduloso) , corolla extus glabra nee pubescen- 
te." Beetle describes our plant as a "vine creeping on ground". 
The original publication of this species is given as page "266" 
in error by Jackson (189$). 

In all, Uh herbarium specimens, including the type collections 
of all the names involved, and 11 mounted photographs have been 
examined by me. 

Citations: BRAZIL: Paran£: Ceceatto 237 [Herb, Mus. Par an. 
3333] (N); Collector undesignated s.n. "[Canpos Oerais, 187U] 
(Ja— U660ii), s.n. [Campos Geraes] (Ja— U660$), s.n. (Ja— U6$97); 
I>usgn 1571I1 (Ca— $33216, F~photo, N, N— photo, S, Si— photo, W- 
Ili8l769, Z — photo); Crurgel s.n. [Herb. Jard. Bot. Rio Jan. 37$39] 
(N)j Hatsehbach 1033 (N), 2010 (N), 2$82 (N); Tessmann s.n. [Herb. 
Mus. Paran. 2$2$] (N) . Rio Grande do Sul: Beetle I9I13 (W— 
211i3837); Collector undesignated s.n. [187U] (Ja— U66OO); Henz 
32$39 (S); Herter s.n. [Herb. Osten 20lil3] (Ug)j C. Jtfrgens 20 
(B, Ja— 17763), 119 (B); Moldenke & Moldenke 1967$ (Es, F, LgJ 
Mg, Mr, N, No, Ot, S, Sm, Ss); Rambo 27293 (N, S), 37698 (N), 
U3689 (Go), $7297 (S); Vianna 101 [Herb. Jard. Bot. Rio Jan. 
U6I68] (N). SSo Paulo: Lund s.n. [Taubat^] (Cp) . State undeter- 
mined: Raben $23 , in part [Brasilia] (Br); Sellow s.n. [Bras, 
merid.; Macbride photos 17ii29] (Br — isotype, F — photo of isotype, 
Kr — photo of isotype, N — photo of type, N — photo of type, N — 
photo of isotype. Si — photo of isotype, Z — photo of isotype). 
URUGUAY: Arechavaleta 20 (Ug); Osten 3133 (Ug)j Sellow 2800 (Vt). 
ARGENTINA: C6rdoba: Hieronymus 7$$8 (BrTT 

xVERBEIIA MATRITENSIS Moldenke, Phytologia 2: 2U0. 19U7. 

Synonyn^r: Verbena Carolina L. x V_. hastata L. ex Moldenke, 
Alph. List Invalid Names Suppl. 1: 23, in syn. 19U7. Verbena 
hastata L. x V^ Carolina L. ex Moldenke, Alph. List Invalid 
Names Suppl. 1: 21;, in syn. 19ii7. 

Bibliography: Moldenke, Phytologia 2: 2U0 t 3U8. 19U7; Molden- 
ke, Alph. List Invalid Nanes Suppl. 1: 23 £: 2ii. I9J47J Moldenke, 
Alph. List Cit. 2: 36O. I9U8; Moldenke, Jjioto Geogr. Distrib. 
Verbenac, [ed. 2] , 16U & 198. 19^9; Moldenke in Chittenden, Roy, 
Hort. Soc. Diet. Gard. h: 2209 & 2211. 19$lj E. J. Salisb.. Ind. 
Kew. Suppl. U: 263. 19$3; Moldenke, Am. Midi. Nat. $9: 3$U. 
19$8j Moldenke, R^sura^ 223, 361, 36$, ?c U73. I9$9j Moldenke, Phy- 
tologia 8: 121 (I96I), 8: hQ9 (1963), and 9: 219. 1963. 



130 PHYTOLOGIA Vol. 10, no. 2 

Apparently a natural or artificial hybrid between V. Carolina 
L. and V. hastata L., exhibiting more or less intermediate char- 
acters ;~stCTis~Tetragonal, rather densely hirsutulous with whitish 
and stiff spreading hairs; leaves lanceolate, thin-chartaceous, 
Ii.5 — 7 cm. long, l.li — 2.I4. cm. wide, sharply acute at the apex, 
cuneate-aciiminate at the base, irregularly dentate or incised- 
dentate along the margins from almost the base to the apex, stri- 
gose-scabrous above, strigillose-scabrellous beneath; inflorescei>- 
ce paniculate, or 1 or 2 spikes terminating short lateral branch- 
es; spikes narrow, elongate, to about 7 cm. long, rather densely 
flowered, apparently not setting seed, the short peduncles and 
slender rachis puberulent with appressed gray hairs; bractlets 
lanceolate, about 2 mm. long, acuminate at the apex, glabrate on 
the back, sparsely ciliolate along the margins at the widest 
part; calyx strigillose, slightly exceeding the subtending bract- 
lets; corolla-tube about U mm. long, its limb about 2 mm. wide. 

The type of this hybrid, and the only specimen of it known to 
me, was collected from a presumably cultivated plant growing in 
the Royal Botanical Garden at Madrid, Spain, and deposited in 
the Brit ton Herbarium at the New York Botanical Garden. It very 
closely resembles xV. engelmannii Moldenke, but differs in having 
rather densely hirsutulous stems, with stiff, whitish, spreading 
hairs, whereas xV. engelmannii has its stems merely appressed- 
pilosulous or puberulent. Only the type specimen and 3 mounted 
photographs have been examined by me. 

Citations: CULTIVATED: Spain: Herb. Hort. Reg. MatrLt. s.n. 
(F — photo of type, N — type, N — photo of type, Z — photo of type), 

VERBENA MEGAPOTAMICA Spreng., Syst. Veg., Cur. Post, U (2): 230— 
231. 1827. 
Synonymy: Verbena phlogiflora var. rt Cham., Linnaea 7: 266. 

1832. Verbena phlogiflora cX glabra Walp,, Report. Bot. Syst. U: 
26. I81i5. Verbena phlogiflora (3 mucilenta Schau. in A. DC., 
Prodr. 11: 538. I81i7. Verbena phlogiflora var. macilenta Schau. 
apud Briq., Ann. Conserv, & Jard. Bot. Genev. 7-8: 238, in syn. 
I90U. Verbena phlogiflora var. mucilenta Schau. ex Moldenlce, 
Suppl. List Invalid Names 9, in syn. I9I4.I. Verbena megopotamica 
Spreng. ex Moldenke, Alph. List Invalid Names ^8, in syn. 19U2. 
Glandulciria megapotanica (Spreng.) Cabrera & Dawson, Rev. Mus. La 
Plata, ser. 2, Bot. ^i 357. 19U;. Glandularia megapotamica 
(Spreng.) Cabrera & Dawson, Rev. Mus. La Plata, ser. 2, Bot. $: 
381. 19Ui. Verbena megopotomica Spreng. ex Moldenke, Alph. List 
Cit, 3: 920, sphalm. 19 U9. 

Bibliography: Spreng., Syst. Veg., Cur. Post, h (2): 230—231. 
1827; Cham., Linnaea 7: 266. 1832; Steud.. Nan. Bot., ed. 2, 2: 
750. I8UI; D. Dietr. Syn. PI. 3: 60$. 18U3; Walp., Repert. Bot. 
Syst. U: 26 & 32. 18U5; Schau. in A. DC., Prodr. 11: 538 & 555. 
18U7; Morong, Britton, & Vail, Ann, N. Y. Acad. Sci. 7: 197. 
1392; Jacks, in Hook. f. & Jacks., Ind. Kew. 2: 1179. 1895; Kunt- 



196U Moldenke, Monograph of Verbena 131 

ze, Rev. Gen. PI. 3 (2): 2^6. 1898; Briq., Ann. Conaerv. & Jard. 
Bot. Genlv. 7-8: 288—291 [Verb. Balans . Parag. l—U] . 190li} 
Briq., Arkiv Bot. Stockh. 2 (10): 7. 190U; Briq. in Chod. & Hass- 
ler, Plant. Hassler. 10: U77. 190U; Herter, Revist. Sudam. Bot. 
h: 186. 1937; Moldenke, Suppl. List Invalid Names 9. 19Ul; Mol- 
denke, Alph. List Invalid Names U8, h9, & 58. 19U2: Moldenke, 
Known Geogr. Distrib. Verbenac, [ed. 1], 39, Ul, Uh, & 101. 19U2; 
Cabrera & Dawson, Rev. Mus. La Plata, ser. 2, Bot. 5: 357 & 381. 
19hh; Cabrera, Bol. Soc . Argent. Bot. 1: 67. 19 U5; Schnack 5c Co- 
vas, Revist. Argent. Agron. 12: 222, 223, & 225—229, fig. 1 C. 
2, & 3 E— G, pi. 12, A, B, E, & H. 19U5; Moldenke, Holmbergia hi 
151. 19 U5; Schnack & Gonzalez, Rev. Argent. Agron. 12: 285, 286 
(fig. 1 C— E & G), 287 (fig. 2 F— J), 288, & pi. 15 A & E. 19U5i 
Moldenke, Alph. List Cit. 1: 26, 83—85, & 282. I9U6; Augusto, 
Fl. Rio Grande do Sul 209, 210, & 232, fig. 95. 19U6; Moldenke, 
Alph. List Invalid Names Suppl. 1: 10 & 25. 19U7; Moldenke, Phy- 
tologia 2: 337. 19U7; Moldenke, Castanea 13: 117. I9ii8; Moldenke, 
Alph. List Cit. 2: 368, 580, & 6UI (19U8), 3: 688, 798, 8U0, 920, 
922, & 923 (I9U9), and U: 1231. 19^9; Moldenke, Known Geogr. Dis- 
trib. Verbenac, [ed. 2], 9U, 99, 100, IO6, & I98. 19U9; Cabrera, 
Lilloa 20: cuadro XVII. 19U9; Moldenke, Phytologla 3: 289 (1950) 
and 3: U68. 1951; E. J. Salisb., Ind. Kew. Suppl. 11: 101. 1953; 
Moldenke, Phytologia $: I33. 1955; Rambo, Sellowia 7: 260. 1956; 
Moldenke, Biol. Abstr. 30: 1093. 1956; Moldenke, Am. Midi. Nat. 
59: 361—363 & 370. 1958; Moldenke, R^sum^ 110, II8, 119, 127, 
223, 296, 369, 372, U20, U2I, & 1;72. 1959; Moldenke, R5sum« 
Suppl. 2: 12 & 13. I96O; Angely, Fl. Paran. 16: 78 (I960) and 17: 
U6. I96I; Moldenke, Phytologia 8: 121 (1961), 9: 178, 362, 365— 
367, 387, Sc 388. 1963. 

Illustrations: Schnack & Covas, Revist. Argent. Agron. 12: 
225, fig. 1 C, 226, fig. 2, 227, fig. 3 E— G, & pi. 12 A, B, E, & 
H. 19U5; Augusto, Fl. Rio Grande do Sul fig. 95. 19ii6. 

Rather tall, subshrubby, branching, perennial herb, 1 m. or 
more tall, almost smooth, the strigose pubescence very tenuous, 
short, and sparse, not at all or at least not conspicuously vis- 
ible to the naked eye; stans herbaceous, erect, hollow, much 
branched, rather sharply tetragonal, inconspicuously pubescent to 
very shortly and obscurely retrorsely puberulent-strigillose or 
subglabrescent; branches similar but more slender, ;videly diver- 
gent; nodes plainly annulate; principal intemodes U.5 — 10 cm. 
long; leaves numerous, decussate-opposite; petioles slender, 5 — 
15 mm. long, minutely and obscurely antrorsely strigillose to 
subglabrous or subglabrescent, margined; leaf-blades thin-charta- 
ceous or submembrajious , dark-green above, lighter beneath, lan- 
ceolate or oblong-lanceolate, 3 — 7.5 can, long, 1 — 3 cm. wide, a- 
cute or subacute at the apex, cuneate or cuneately narrowed at 
the base into the petiole, rather irregularly serrate with broad 
rather roimded or slightly apiculate teeth along the margins from 
the widest part to the apex, rather sparsely and obscurely strig- 
illose with brownish antrorse hairs on both surfaces or subglab- 
rescent; inflorescence axillary and terminal at the tips of the 
branches, capitate, long-pedunculate, densely many-flowered, ap>- 



132 PHTTOLOGIA Vol. 10, no. 2 

parently not elongating after anthesis, simple, temate, or 
cymose-paniculate with 1 or 2 subsessile lateral ones at the bas^ 
pedimcles slender, tetragonal, 2 — 6.^ cm. long, retrorsely strig- 
illosej the lower pair of flowers separated from the rest; bract- 
lets ovate-lanceolate, minute or very short, about k mm. long, 
very obscurely strigillose, acute at the apex, much shorter than 
or to 1/3 as long as the calyx, ciliate; calyx tubular, elongate, 
about 10 mm. long, glabrous or subglabrescent to very minutely 
and obscurely antrorsely strigillose with closely appressed bro?m- 
ish hairs, 5-apiculate at the apex, occasionally with rather num- 
erous sessile blackish round glands; corolla hypocrateriform, 
blue or violet to lilac or rose, its tube slightly exserted, sub- 
glabrate on the outside, the limb about 10 mm. wide. 

The type of this much misunderstood species was collected by 
Friedrich Sellow ( no. 13), probably in Rio Grande do Sul, Brazil, 
and was deposited in the herbarivm of the Botanisches Museum at 
Berlin, where it was photographed by Macbride as his photograph 
no. 17ii38 (in part), but is now destroyed. The type of Ghamis- 
so's V. phlogiflora var. oC is an unnumbered Sellow collection 
frcm "Brasilia meridional!", also originally deposited in Berlin, 
photographed by Macbride under his photograph 17U38 (in part), 
and now destroyed, as is also the type of Walpers' var. glabra. 
Schauer's var. mucilenta is merely a new name for Chamisso's 
unnamed variety. Schauer says of it "Hab. in Brasiliae prov. Rio 
Grande do Siil". Walpers (18U^) placed Sprengel's V. megapotamica 
in his list of doubtful species, but unwittingly described the 
same taxon well as V. phlogiflora ^ glabra . Jackson (189^) re- 
duced Sprengel's name to synonymy under V, phlogiflora Cham,, 
while Cabrera & Dawson (19Ui) do just the reverse, reducing V. 
phlogiflora to synonyny under V. megapotamica . They describe the 
species as "Hierba perenne, erecta, con hojas lanceolado-ovadas, 
pecloladas, profundamente aserrado-cjrenadas , y flores azules dis- 
ouestas en espigas cortas, capitulif ormes . Es una especie de in- 
florescencia llamativa que algunas veces se encuentra en cultivo. 
Habita en el s\ir del Brasil, Uruguay, Paraguay y nordeste de la 
R^publica Argentina hasta el Rio de la Plata. En Punta Lara es 
un element© frecuente en las abras." 

The species has been found in hedgerows and wet thickets, on 
campos, and along rlverbanks, from 900 to 1650 meters altitude, 
flowering in October, November, and January to March. Rambo en- 
coimtered it in a region of 2 meters rainfall and — 2^° C. tem- 
perature variation. Cabrera reports it from "sabre terreno moj- 
ado en un claro del bos que". 

Herbarium specimens of this plant have been misidentified and 
distributed under the names V, megapotamica var. tweediana Kuntze, 
V. peruviana (L.) Britton, V. phlogiflora Cham,, "V^ aff . venosa 
Gill. & Hook,", and even Lantana sp. On the other hand, the Fie- 
brig 5699 distributed as V, megapotamica is actually V. kuntzeana 
Moldenke; Sehnem 3776 is V. lobata var, hirsuta Moldenke; Venturi 



196U Moldenke, Monograph of Verbena 133 

5397 is the type collection of V. moricolor Moldenke j and Cabrera 
& Corte 9610 , Kuntze s.n. [Contendas, Dec. '92], and Vattuone & 
Bianchi 60 are V. phlogiflora Cham. Parodl 122^0 closely resem- 
bles V. phlogiflora . Morong, Britton, & Vail (I892) cite Balansa 
IO2U as V. peruviana (L.) Britton, which it certainly is noti 

Verbena megapotamica is in general extremely similar to V. 
phlogiflora . Sprengel's original description (1827) is merely 
"caule herbaceo erecto pubescente, foliia petiolatis oblongo- 
lanceolatis serratis basi cuneatis glabriusculis , capitulis ter- 
minalibus, bracteis minutis cilia tis, calyce elongate glabro." 
Walpers (18U5) characterizes the taxon well when he describes it 
as "fere laevis, strigis tenuibus brevibus, oculo nudo vix con- 
spiciiis; spicis capitatis, unico florum pari inferiori remotius- 
culoj bracteolls nunc minimis, nvinc tertiae circiter csdycis 
longitudinis." Schauer (l8U7) also distinguishes it well from V. 
phlogiflora when he says '♦major, pube strigosS tenuissimd rardque 
adspectu glabra, caulibus herbaceis fistulosis ramosissimis erec- 
tis, spicis ad apices ramorum saepe ternis longe peduncvilatis 
simplicibus vel uno alterove pari ad basin primariae subsessilis 
accedente cymoso-paniculatis ," Arechavaleta 27 bears a notation 
"pistilos largos escotados en el extremidad". 

The hybrid of _V^ megapotamica with V. peruviana (L.) Britton 
is xV. schnackii Moldenke, that with V. santiaguensis (Covas & 
Schnack) Moldenke is xV. vaga Moldenke, and that with V. tenul- 
secta Briq. is xV. transitoria Moldenke . 

It should be noted here that V. megapotamica hybrida Osten is 
actually V. kuntzeana Moldenke; V. megapotamica f . truncatula 
Brlq. and var, t rune a tula Briq. are V, incisa Hook.; V. megapo- 
tamica var. pinnatiloba Kuntze and var. tweediang. f , pinnatiloba 
Kuntze are V. pinnatiloba (Kuntze) Moldenke; and V. megapotamica 
var. phlogiflora (Cham.) Kuntze, var. tweediana Kuntze, var. 
tweediana (Niven) Ktintze, and var. tweedleana Kuntze are all V. 
phlogiflora Cham, 

Kuntze (I898) says of V. megapotamica ; "Schauer citlrt zu 
dieser Art, von den ich Sprengel'sche und Chamisso' sche Origin- 
ale vergleichten konnte, noch V. cunha (err. cunea ) Veil. [182^ 
descr. 17] aber Calyx ^-dentatus dentibus acutis passt nicht, 
weder nach der Beschreibung noch nach der Abbildung tab. lil, denn 
die Kelchzahne sind ungleich, davon zwei mindesten pfriemlich." 

Briquet (190li) has a lot to say about this species, but unfor- 
tunately confuses it with V. incisa Hook. He says *^e V. megapo- 
tamica de doit pas 8tre confondu avec diverses especes voisines 
tres souvent m6l6es avec lui dans les herbiers . II est caracter- 
is6 par ses tiges dress^es, assez robustes, rameuses a rameaux 
ascendants, ses feuilles ov6es-oblongues, oblongues ou oblongues- 
lanc6ol6es, toutes tres nettement p6tiol6es, a base du limbe at- 
tenu6e ou mfime tronqu^e-subcord^e, a marges assez fortement in- 
cis6es-cr6nel6es . Les fleurs sont groupies en 5pis tres courts. 



13li PHYTOLOGIA Vol. 10, no. 2 

le plus souvent m6me r^duits a dans capitules, a calices sessiles, 
rapproch^es au sommet du pSdoncule de facon a former un h&nisphere, 
Le calice est longuement tubuletix, presque siphone, couvert d'un 
indument apprimi tres dense, a glandes rares de d^passant pas ou 
d^passant de bien peu les polls courts (longueur moyenne 1,2 — 1,5 
cm.): ses dents in^gales sont brievement acumin^es-subulees, les 
postlrieures longues de 0,5 — 0,8 mm. les ant^rieures atteignant 1 
— 1,2 ram. La corolle, tres grande possede un tube exsert et un 
limbe atteignant 1 — 1,8 cm. de diametre. Les nos. 10 2U et 102lib 
sont tres typiques. Le no. 102l|.c appartient a une forme un peu 
difflrente (var. truncatula Briq. herb.), a limbe des feuilles 
plus obtus au sommet et tronqu^-subcord^ a la base, a nervation 
plus saillante, k 5pi \in peu plus allong^, a calice l^gerement 
plus court: la corolle possede un tube moins exsert et un limbe 
de plus faible diametre. Malgr^ ces differences, cette forme 
peut rentrer dans le cycle des variations du V. megapotamica v. 
Tweediana. Nous avons longtemps hSsit^ i reprendre po\ir cette 
espece le ncan de Sprengel, parce que cet auteur attribue au V. 
megapot?mica un calice glabre, ce qui n'est le cas dans aucune 
des formes de notre espece. Mais M. 0. Kuntze, qui a pu compar- 
er des originaux de Sprengel et du Chamisso, nous apprend (I.e.) 
qu'il s'agit d'une forme glabrescente et non pas glabre. Nous 
partageons d'ailleurs tout a fait 1' opinion de M. Kuntze lorsqu- 

'il exclut de la synonymie le V. cunha Veil attendu que cette 

plante doit poss^der des dents calicinales simplement aiguBs . A 
notre avis, la figure grossiere de Vellozo ne saurait sans impru- 
dence @tre assimil^e a I'vine quelconque des ncsnbreuses especes 
af fines de ce groupe." 

August© (I9U6) cites an Isabella collection from Rio Grande do 
Sul, Brazil. In all, 38 herbarium specimens and 6 mounted photo- 
graphs, representing the type collections of all the names invol- 
ved, have been examined by me. 

Citations: BRAZIL: Parani: Hatschbach 3365 (Z) . Rio Grande do 
Sul; Ranibo 3U729 (N, S), 35729 (N)j Sellow I3 [Macbride photos 
I7U38, in part] (Kr — photo of type, N — photo of type, N — photo of 
type), Santa Catarina: Rambo 60153 (S)j Reitz 2927 (N)j Smith & 
Klein 7707 [Herb. Barb. Rodr. 22660] (Mm, N), 7765 (Ok), lOhhQ 
(Ok); Smith & Reitz lOUjS (N, W~225l679)j Smith , Reitz , & Klein 
7765 (W — 2251310) . State undetermined: Sellow s.n. [Brasil mer- 
idionali; Macbride photos 17li38, in part] (Br, Kr — photo, N — 
ibhoto, N— photo). PARAGUAI: Balansa 102U (N) . URUGUAY: Arecha- 
valeta 27 (Ug, Ug, Ug), 3128 (Ug); Collector undesignated s.n. 
(Ug); Mrs. 0. C_. James s.n. [Colonia, Jan. I909] (Du— lli9772) j 
Osten 5389 (N, Ug) ; Teisseire s.n. [Colonia, I913] (Ug— IM6) . 
ARGENTINA: Buenos Aires: G. Atkinson I6O63 (N); Burkart 3122 
[Herb. Osten 2O9l;0] (Ug); Cabrera l581i (N, Sp— 2U56U) , I6I8 (N), 
3U01 (Bt— U3217, N, Sp~389U7), 5378 (Ca— 882789, N, N); Lefebra 
s.n. [Isla Santiago, 1893] (Br); Parodi 12250 (N); Pastore & 



196U Moldenke, Monograph of Verbena 135 

Troncoso 983 (Ug— 3129). 

VERBHU MMDOCINA R. A. Phil., Anal. Univ. Chil. 3^: 191. 1870. 

Synonymy: Glandiilaria mendocina (Phil.) Covas & Schnack, Rev, 
Argent. Agron. 11: 96. I9UU. 

Bibliography: R. A. Phil., Anal. Univ. Chil. 3$: 191. I370j R. 
A. Phil., Sert. Mendoc. Alt. 33. 1370 j Jacks, in Hook, f . &; 
Jacks., Ind. Kew. 2: 1179. I895j R. A. Phil., Anal. Univ. Chil. 
90: 609. 1396; Durand & Jacks., Ind. Kew. Suppl. 1: U5l. 1906j 
Reiche, Fl. Chile $: 295 & U63. 1910j Sanzin, Anal. Soc. Cientif. 
Argent. 88: 129 — 131. 1919; Moldenke, Known Geogr. Distrib. Ver- 
benac, [ed. 1], Ui & 101. 19U2; Moldenke in Lundell, Fl. Texas 
3 (1): Ul. 19U2; Beetle, Bot. Review 9: 67U. 19U3; Covas & Schnack, 
Rev. Argent, Agron. 11: 96 & 97. 19Uij Schnack & Covas, Darwini- 
ana 7: 71, 72, 7h, & 75, pl. IB. I9I6; Covas & Schnack, Darwini- 
ana 7: 86. 19U5; Moldenke, Phytologia 2: 3U8. l9U7j Moldenke, 
Alph. List Invalid Names Suppl. 1; 10. 19U7; Moldenke, Castanea 
13: 119. 19^8; Moldenke, Known Geogr. Distrib, Verbenac, [ed. 2], 
106, 16U, & 198. 19U9; Moldenke, Alph, List Cit. 3: 688, 7U8, 770, 
& 813 (I9U9) and U: 1162 & 12U9. 1919; Moldenke, Phytologia 3: 
lla (I9U9) and 3: 289 & 290. 1950; Moldenke in Chittenden, Roy. 
Hort. Soc. Diet. Card. U: 2209 & 2211. 1951; E. J. Salisb., Ind. 
Kew. Suppl. 11: 101. 1953; Moldenke Am. Midi. Nat. 59: 356. 
1958; Moldenke, R^sum^ 119, 127, 22U, 296, & U72. 1959; Moldenke, 
Phytologia 8: 123 (1961) and 8: 396. 1962; Moldenke, R^sum^ 
Suppl. 5: 7 & 8. 1962; Moldenke, Pt^tologia 9: 39U & UOO. I963. 

Illustrations: Schnack & Covas, Darwiniana 7: pl. IB. I9I4.5. 

Annual herb, softly appressed-pilose or persistently whitish- 
strigose throughout; stems erect, sturdy, simple or branched, to 
30 cm. long; leaves decussate-opposite, ovate- triangular in out- 
line, about h cm. long and 2.U cm. wide, variable in form, some- 
times trifid or tripartite, sometimes pinnatifid or pinnatipar- 
tite with 5-parted lobules, cuneately narrowed at the base into 
the petiole, the lobules about 3 mn. wide; spikes pedunculate, 
subtemate, finally greatly elongate; bractlets lanceolate, about 
5 mm. long, subequaling the calyx; calyx about 6 mm. long, silky 
or sericeous-pubescent, not glanduliferous; corolla varying fron 
blue or pale-blue to lilac, rose-lilac, or purple, equcding the 
calyx, its tube about 1 cm. long, glabrous, the limb about 5 inni. 
wide; anthers not appendaged; pollen 87.5 percent fertile; pistil 
about 7.8 mm. long; chromosome niomber: 2n - 10, 

The type of this puzzling species was collected by Rudolf Am- 
andus Philippi at Mendoza, Argentina, in 1868, and is deposited 
in the herbarium of the Museo Nacional at Santiago, Chile; an 
isotype in the herbarium of the Botanisches Museun at Berlin was 
photographed by Macbride as his photograph no. 17U30, but is now 
destroyed . 

Covas & Schnack (19U5) discuss the relation between the 
length of the pistil and the voliime of the pollen-grains in this 
species. Beetle (19U3), through some clerical error, places 
this species in the Cruciferae! It was apparently introduced 
into cultivation in or about iSUl, although not described until 



136 PHYTOLOGIA Vol. 10, no. 2 

29 years later. It has been collected in fields and wet places 
in fields, at 2^0 meters altitude, flowering from October to Feb- 
ruary, April, and August. The only recorded common name is "Men- 
doza verbena". Herbarium specimens have beoi misidentified and 
distributed under the name V. erinoides Lam., but the H. N. Mol- 
denke 18237 distributed as V. mendocina is actually V. tenuisecta 
Briq. 

The species is very similar to V. tenuisecta , but differs in 
having its leaf-segments broader, oblong, not uniform in diameter, 
mostly 1 mm, or more wide, the corolla- tube 1 cm. long and its 
limb 5 inni* wide, the bractlets about 5 inm. long, and the whole 
plant usually more persistently iihitish-strigose throughout. 
Philippi, in his original description (I87O), says "Parece mui 
vecina a la V. incisa Schauerj que no he visto todavla; pero sus 
brae teas no son aovadas, i son mucho mayores; no haigl4ndulas en 
el cilizj la corola es azul i no 'roseo purpurascens' j las hojas 
mucho mas partidas." If my inteipretation of the species is cor- 
rect, it has very little affinity with V. incisa Hook. Sanzin 
(1919) claims that V. mendocina is intermediate between what he 
calls V. erinoides and V. erinoides var. glandulifera Sanzin [«=V. 
perakii (Covas & Schnack) Moldenke] . He says "Cerce de la ciudad 
de Mendoza, y precisamente del lado de la Cordillera a una eiltura 
de 1000 y 1200 metres, abunda una variedad (Herb. Sanzin nos. 139, 
1700 , 3099 , 3129, 3130 ) , que lleva glindulas en el cdliz y que 
tiene las hojas anchas, triangulares, de base cuneada y trlfidas 
o tripartidas con los segmentos casi enteros o con unos lobiolitos 
laterales : 

"A typo differt caule, foliis, calicibusque hirsutis, pilis 
glanduliferis mixtis. Tubo calice subduplo longiore, appendici- 
bus antherarum subexsertis clavatis violaceis. Laciniis foliarum 
lanceolatis (Osten, in litt.). Mis al sur y a las mismas alturas 
indicadas existe otre variedad que se acerca mis al tipo por sus 
hojas tripartido-pinatifidas con segmentos angostos, pero que se 
diferencia esencialmente por sus glindulas estaminales apenas 
salientes de la garganta del tubo corolar en vez de ser inclusas . 

"La V, mendocina Phil, es intermediaria entre estas dos vari- 
edadas, pues el examen de ejemplares aut^nticos de Philippi, del 
museo de Santiago, me permiti6 constatar que tiene hojas de dos 
clases, id^nticas en la forma a las hojas de las dos variedades 
citadas. El caricter de los tallos erguidos de la V. mendocina , 
no es constante, pues en la variedad glandulifera huy individuos 
erguidos y otros semirastreros . Por todo esto ma parece conven- 
iente unir en una sola las dos especies, V. erinoides y V. mendo- 
cina ." 

Covas & Schnack (19l4ii) suggest a possible hybrid between V. 
mendocina suid V. parodii (Covas & Schnack) Moldenke: "Hemos hal- 
lado en la localidad del tipo una poblaci6n formada, muy probab- 
lemente, por h£bridos (y formas derivadas de ^stos) entre esta 
especie y Glandularia mendocina Hemos podido estudiar un 



196ii Moldenke, Monograph of Verbena 137 

trozo del ejemplar tipo de esta especie (ex Herb. Mus. Nac, San- 
tiago de Chili, PHILIPPI 1868: Iter mendocinum) , y evidentemente 
se trata de una buena especie, distinta a Glandularia laciniata . . 

y no sin6nlina como lo admite SANZIN Posee tambiSn cinco pares 

de cromosomas, observados en diacinesis.) La poblaci6n hibrida 
presenta una amplia gama de variaci6n que ccanprende formas inter- 
medias y formas vecinas a ambos padres: en algunas de estas for- 
mas hemos podido observar floras con pequeflos l6bulos petsuLoides 
en la base del limbo de la corola, carficter que nunca hemos obser- 

vado anteriormente en el gSnero Glandularia kAdemis hemos ob- 

servado, en individuos de la poblaci6n hibrida mencionada, irreg- 
ularidades en la meiosis (miembros de un par de crcHnoscmas separ- 
ados en diacinesis, lo cual indica falta de hcmoligla en parte 
del material cromos6mica) En la misma poblaci6n hfbrida hem- 
os encontrado una forma con floras rosadas, color aparentemente 
debido a un derivado de cianidina." This natural hybrid is dis- 
cussed herinafter under the name xV. perturbata Moldenke. 

For a key to distinguish V. mandocina from some of its close 
allies, see under V. laciniata in these notes. 

Philippi apparently used the same specific epithet - "mendo- 
cina" - a second time (in I896) for a different plant collected 
near Mendoza in October of I87O. He describes it as follows: 
"V. fruticosa, caule erecto, crasso {k mm.); ramis virgatis 
pubemlisj foliis glanduloso-puberulis, margine revolutis, lac- 
iniis indivisis vel bi-trifidis obtusiusculisj peduncxilis longe 
nudisj spica oblongaj bracteis angustis linearibus, dimidium 
calycam aequantibusj calice glanduloso-puberxilo; corollae glab- 
rae tubo sesquies aaquanta, limbo parvo. Prope Mendozsun Octobri 
1870 lecta est. KL ejemplar que tengo a la vista, tiene la al- 
tura de 3U centimetres, i su tallo en la base el grosor de k 
millmetros, est! cubierto de una corteza blsmquizca. Sus ramos 
tienen la lonjitud de 20 centimetres, i constsm dolo de 5 a 6 
intemodiosj el pedlinculo tiene 6 a 8 centimetres de largo. Las 
hojas miden 2h millmetros de largo, las espigas, que tal vez se 
alargarin mas tarde, tienen la lonjitud de 3 a 3 1/2 centimetres, 
el cdliz mide 10 millmetros, la corola 15 millmetros. — Se par- 
ece bastcinte a la V. trachea , pero se distingue ficilmente por 
su porte, su vellosidad mui distintaj la corola mas pequefla." 
What this second species is — not having seen any type material 
of it — I do not as yet know. 

In all, 21 herbarium specimens and 7 mounted photographs, in- 
cluding photographs of the type collection, have been examined 
by me. 

Citations: URUGUAY: Berre 7U78 (N) . ARGENTINA: Buenos Aires; 
Dusgn 6308 (N, S); Rodrigo V.721 (S); Ruiz Huidobro 1330 (Bn, S). 
Chaco: Venturi 9780 (N, S) . C6rdoba; Ruiz Leal 12217 (Rl). For- 
mosa: I_^ Morel 530 (N, S) . La Rioja: Ruiz Leal 16597 (Rl), 17162 
(Rl) . Mendoza: R. A. Philippi s.n. [l868j Macbride photos 17U30] 
(Kr — photo of isotype, N — photo of isotype, N — photo of isotype); 



138 PHYTOLOGIA Vol. 10, no. 2 

Semper s.n. [Ruiz Leal 9$39] (N) . CULTIVATED: New York: Ahles s. 
n. [N. Y. Bot. Gard. Cult. PI. U77A5] (N, N) . Sweden: AM s.n. 
[l8.VIII.19li7] (S); Herb. Mas. Bot. Stockholm s.n. [HolmiaeT^lil] 
(S); E. Wall l^/TltZ (Ew, Ew, F—photo, N, N— photo, Sg— photo, Z- 
photo) . 

VERBENA MENTHAEFOLIA Benth., PI. Hartw. 21. 1839. 

Synonymy: Verbena setosa Mart. St Gal., Bvill. Acad. Brux. 11 
(2): 321. 181^1;. Verbena hintonl Moldenke, Phytologla 1: k39—hhO. 
1914.0. Verbena setosa Mart, ex Moldenke, Prelim, Alph. List In- 
valid Names U8, in syn. I9U0. Verbena menthifolia Benth. ex Mol- 
denke, Suppl. List Invalid Names 9, in syn. 19 Ul. Verbena offic- 
inalis Wats, ex Jepson, Fl. Calif. 3 (2): 380, in syn. 19i;3 [not 
V, officinalis L., 1753] . Verbena hintonii Moldenke, Alph. List 
Invalid Names Suppl. 1: 21;, in syn. 19 U7. Verbena officinalis 
var. mediterranea N6e ex Moldenke, Alph. List Invalid Names Suppl. 
1: 26, in sjm. I9I47. Verbena magdalensis Moldenke, R^suml Suppl. 
3: liO, in syn. I962, 

Bibliography: Benth., PI, Hartw. 21. 18 39 J Steud., Nom. Bot,, 
ed. 2, 2: 7^0. I81ai Mart. & Gal., Bull. Acad. Brux. 11 (2): 321. 
I8IU1; Walp., Repert. Bot. Syst. U: 32 (I8h5) and 6: 687. l8U7j 
Schau. in A. DC., Frodr. 11: 5U7 & $$$, 18U7} Coult., Contrib. U. 
S. Nat. Herb. 2: 327. I89lt; Jacks, in Hook, f . & Jacks., Ind. Kew. 
2: 1179. I895j J. K. Small, Fl. Southeast. U. S., ed. 1, IOO8 
(I903) and ed. 2, IOO8. 1913; H. J. Lam, Verbenac, Malay, Arch. 
10. 1919; Rydb., Fl. Prairies & Plains 677. 1932j J. K. Small, 
Man. Southeast. Fl. 1137. 1933; Perry, Ann. Mo. Bot. Gard. 20: 
2l;7, 259, 263—265, & 355. 1933; C017, Texas Agr, Exp. Sta. Bull. 
55O: 89. 1937; Moldenke, Prelim. Alph. List Invalid Names U8. 
I9UO; Moldenke, Phytologia 1: k39—UhO (I9U0) and 1: 511. 19ia; 
Moldenke, Suppl. List Invalid Names 9. 19U1; Moldenke, Known Ge- 
ogr. Distrib. Verbenac., [ed. 1], 1>-15, 18, 19, UU, & 101. 
19U2; Moldenke in Lundell, Fl. Texas 3 (1): 16 & 21. I9l;2j Mol- 
denke, Alph. List Invalid Names 1^8 & 50. 19U2; H. S. Gentry, 
Carnegie Inst. Wash. Publ. 527: 222 & 306. I9lt2j Moldenke, Knoim 
Geogr. Distrib. Verbenac. Suppl. 1: 2. 19ll3; Moldenke, Phytolo- 
gia 2: 68, 88, & 115. 191+5; Moldenke, Castanea 10: UO. 19U5; 
Moldenke, Alph. List Cit. 1: 3, Hi, 32, 57, 109, lli3, lUi, I69, 
178, 220, 221, 232, 233, 2I16, & 261. I9U6; Moldenke, Alph. List 
Invalid Names Suppl. 1: 2k & 26. 19li7; Moldenke, Phytologia 2: 
330 & 3U8. 19U7; Hill & Salisb., Ind. Kew. Suppl. 10: 2U2. 191+7; 
Moldenke. Castanea 13: 113. 19li8; H. N. & A. L. Moldenke, Pi. 
Life 2: Ul & 61;. 19l;8; Moldenke Alph. List Cit. 2: 36O, U65, 
U67, U71, U73, U7U, U76, li79, i;82, U83, U88, U89, U98, 5U2, 587, 
596—598, 60li, & 607 (19U8), 3: 685, 697, 72U, 7U0, 752, 775, 
785, 786, 799, 800, 803. 80ii, 829, 830, 832, 833, 905, 933, 953, 
977, & 978 (19li9), and h: 997, IO8I, 1100, 1120, 1126, II38, 
1166, 1169—1172, 1180, 1191, 1199, 1200, 1207, 1208, lai, 
1221;, 1225, 1227, 1231, 1236, 1239, 12l;l, 12l;>-12l;5, 1255, 



196ii Moldenke, Monograph of Vertena 139 

1291, 1295, 1298, & 1303. 19U9; H. N. & A. L. Moldenke, Anal. 
Inst, Biol. Mex. 20: Hi. 19ii9} Moldenke, Known Geogr. Distrib. 
Verbenac, [ed. 2], 2U, 26, 27, 33, 108, 16U, & I98. 19U9; Mol- 
denke, Phy-bologia 3: 73. 19U9; Moldenke in Chittenden, Roy, Hort, 
Soc. Diet. Gard. h: 2209 & 2211. 19^1; Rzedowski, Anal. Esc. Nac. 
Cienc. Biol. 8: 100. 19^hi Moldenke, Inform. Mold. Set h? Spec. 3 
(195ii) and 51 Spec. h. 1956: Moldenke, R6svan4 29, 32, 33, 39, 130, 
22lt, 366, 370, 371, 37U, & Ii72. 1959; Moldenke, phytologia 8: lU3 
& Ihh (I96I) and 8: 279 & U07. 1962; Moldenke, R^sum^ Suppl. 3: 8, 
38, & UO (I962) and 6: U fie 11. I963; Moldenke, Phytologia 8: U71, 
U87, U88, & U9I (I963) and 9: 39, UO, 78, 156, 165, & 167. 1963. 
Annual or perennial herb, varying from neso-ly prostrate to e- 
rect and 1.5 m. taOJL, slender, branching, often with a spread of 
60 cm,, sometimes practically leafless; stems decumbent or ascen- 
ding to erect, very slender, branched, acutely tetragonal, brown- 
ish, pilosuloug or hispid to more or less short-hirsutulous, 
sparsely and minutely hispidulous; branches similar to the stems 
in all respects; nodes annulate; principal internodes 1.5~6 cm. 
long; leaves decussate-opposite, the upper ones sessile or sub- 
sessile, the lower ones tapering at the base into a margined pet- 
iole; petioles very short and obscure or obsolete; leaf-blades 
thin-chartaceous, uniformly green on both surfaces, narrowly lan- 
ceolate or elliptic in outline to ovate or obovate-cuneate. 2.5 — 
6 cm, long (the upper ones 1.5 — 2 cm. long, 2 — 11 mm. wide; 
coarsely few-dentate along the margins, deeply cleft or subln- 
cised to incised-pinnatifid or else 3-laciniate or 3-lobed below 
the middle, acute at the apex, cuneate at the base, very ro\igh, 
more or less short-hirsutulous or strigillose on both surfaces, 
especially on the venation beneath, somewhat pustulate above, the 
divisions or segments lanceolate, acute or subacute, entire or 
remotely serrate-dentate to coarsely dentate or incised; midrib 
and the 1 or 2 secondaries, very slender, impressed aboAre, prom- 
inulous beneath; veinlet reticulation indiscernible on both sur- 
faces; inflorescence axillary and terminal, spicate; spikes ter- 
minal, panicled or fasciculate-panicxilate, numerous, elongate, 
7.5 — 23 cm. long, slender, many-flower«d, dense or else compact 
only at the apex, scabrous, the flowers densely imbricate before 
and during anthesis, loosely scattered in fruit; rachis very 
slender, sparsely pilosxilous; peduncles obsolete or extremely 
short; bractlets small, lanceolate or ovate- lanceolate, acuminate 
at the apex, variable in length, mostly 1.5 — 2 mm, long, subequal- 
ing or usually shorter than the calyx, ciliate along the margins 
and sparsely strigillose; flowers minute; cal3rx 2.5 — 3 mm. long, 
strigillose, sparsely (if at all) glandular, its teeth minute; 
corolla varying from purple, very faint piirple, light-purple, 
deep blue-purple, or pale-pink to blue, pale-blue, light-blue, 
bright-blue, blue-lavender, lavender-blue, light lavendei^blue, 
lavender, lilac, or whitish-lavender, about 7 mm. long or 3 times 
as long as the calyx, its tube only slightly longer than the 
calyx, the limb ii— -6 mm. wide, the lobes more or less truncate; 
frxiit remote; cocci trigonous, 2 — 2.5 mm. long, convex on the 
back, striate, raised-reticulate above, the commissural faces mui»- 



lliO PHYTOLOGIA Vol. 10, no. 2 

Icate . 

The type of this species was collected by Carl Theodor Hartweg 
( no. 175 ) at Leon, Guanajuato, Mexico, in 1839, and is deposited 
in the herbarium of the Royal Botanic Gardens at Kew. The type 
of V. setosa -was collected by Henri Guillaume Galeotti ( no. 778 ) 
in part in the forests of Moran, near Real del Monte, Hidalgo, 
Mexico, in I8I4.O, and in part in the forests of Sabino, near Iz- 
miquilpan, northern Mexico, at altitudes of 6OOO to 7500 feet, 
probably deposited in the herbarivm of the Jardin Botanique de 1' 
Etat at Brussels. The name, V. officinalis var, medlterranea , is 
apparently based on the Collector \inde3ignated h& [probably 
collected by N5e] deposited in the Brussels herbarium, and V. 
magdalensis on Carter , Alexander , & Kellogg 2135 in the Dudley- 
Herbarium. The type of V. hintoni was collected by George B. 
Hinton ( no. II99I ) — in whose honor it was naned — on a grassy 
bank at Zitacuaro-Bosque, district of Zitacuaro, Michoaciin, Mex- 
ico, on June 28, 1938, and is deposited in the Britton Herbarium 
at the New York Botanical Garden. 

Verbena menthaefolia is a rather puzzling species, very simi- 
lar to V. officinalis L., from which it may be distinguished by 
its leaves being densely strigiUose on both surfaces and the 
stems and brauiches being sparsely and minutely hispidulous through- 
out. It is native to the southwestern United States and most of 
Mexico. Common names recorded for it are "berctil", "tel-rebi", 
"verbena", "vervena", and "weyhooli". 

Herbarium material of this species has been misidentified and 
distributed by various workers \mder the names V. affinis Mart. & 
Gal., V. brae tea ta Lag. & Rodr., V. canescens Kvmth, V. Carolina 
L., V. ehrenbergiana Schau., V. halei Small, V. littoralis H.B.K., 
V. littoralis var. affinis (Mart. 4 Gal.) Bourgeau, V. neomexica- 
na (A. Gray) Small, V. officinale L., V. officinalis L., V. or- 
cuttiana Perry, V. polystaclya H.B.K., Vj. prostrata R. Br., V. 
recta H.B.K., V. scaberrima Cham., V. scabra Vahl, "V^ spuria in- 
edita " [Ruiz St Pa von], V. subuligera Greene, V, teucriifolia Mart. 
& Gal., V, trifida H.B.K., V. urticaefolia L., and V, xutha Lehm, 

On the other hand, the Balls 10065 , distributed as V. menthae- 
folia , is actually V. abramsl Moldenkej G. L. Fisher )iMl6 and 
Pringle s.n. [Valley Ortiz, Apr. 11, 1887] are V. halei Small; 
the Maltby 252 annotated by Perry as "Aff . V. menthaefolia Bth." 
is V. litoralis H.B.K.; the Cj, R. Orcutt 1371, similarly annota- 
ted by Perry, is V. longifolia Mart. & Gal.; Raven , Mathlas , & 
Turner 12587 is V. neomexicana var. hirtella Perry; Edw. Palmer 
lOUl is V. neomexicana var. xylopoda Perry; and Edw. Palmer 3^6 
is V» pinetorum Moldenke. Verbena setosa was reduced to syno- 
niymy~under V. officinalis L. by Schauer (181^7). 

Verbena menthaefolia has been collected in canyons, rocky can- 
yons, smd ravines, along creeks, on hills and foothills, open pine 



196ii Moldenke, Monograph of Verbena lUl 

forests and open pine-oak woods, in beds of intermittent streams 
among rocks, banks of irrigation ditches, barrancas, mesas, and 
basaltic mesas, on fields and car^ron margins, flats, dry flats, 
and adobe or grassy flats and banks, in wet meadows and river 
valley pastures, red clay soil of dense oak forests, in black 
clayish loam in scrubland, margins of dry pools, along waysides 
and arroyo margins, on dry ditch banks and steep rocky volcanic 
outcrops, heavy clay creek bottoms, on exposed vertical rocks, in 
sandy loam on grassy hillsides, in sandy or heavy alkaline soil, 
roadside bankings and moist depressions, in low growth and wet 
pinewoods, in rich moist soil in open woodlands, in msj*8hy grotind, 
arroyos and milpas, in thorn and mesquite forests, in grasslands 
with scattered pines, on playas and bottomlands, on sandy river 
banks and canal banks, in clay and limestone soil of mountainsidee^ 
in sand, along roadsides and ditches in cornland, moist adobe 
soil, the edges of fields, and in sandy arroyo margins, at alti- 
tudes of 3 to 2930 meters, flowering and fruiting from December 
to October. 

Purer reports finding it "in open valley in partial shade of 
large shrubs". Gentry (19U2) cites his no. 598 , but this proves 
to be V. Carolina L. He refers to V_. menthaefolia as "A common, 
weed-like, small-flowered lowland Verbena " found in "arroyo- 
margins in Short-tree Forest." He also calls it an "ainnual, 
gix)wing rather rankly in moist places, occurrence casual" and 
says that it grows "scattered in cienaga bottomland of grama 
grasslands". Carter, Alexander, & Kellogg encountered it "among 
mesquite in broad shallow arroyo bordered with Prosopis juliflo- 
ra". Wolf describes it as a plant of the Lower Sonoran Zone. 
Powell & Edmondson that that it is "abundant in Sinaloa". Smith, 
Peterson, & Tejeda found it growing in black to gray soils in oak 
forests giving way to scrubby secondgrowth thickets below. Stey- 
ermark avers that in Guatemala the leaves are mashed and diruik 
raw with water for curing chills and fevers . 

Jepson (I9U3) records the species from Tulare County, Califoi?- 
nia, and cites numerous specimens and literature references. 
Mueller says that it is "common in less dense oak woods of upper 
canyon". M. E. Jones s.n. [west of Uvalde, April 26, 1931] is a 
mixture with V. halei Small; the same collector's no. 2829U was 
gathered "between Kerrville and San Antonio", and so may have 
been from Kerr or Bexar Counties in Texas . Moore & Wood Ul3U is 
a mixture with V. canescens H.B.K., irtiile Schery ll|2 is a mix- 
ture with V. teucriifolia Mart. & Gal. The J. Let s.n. [San Di- 
ego, Nov, 9] was first identified as V. bracteata, then V. pros- 
trata, while C. T. Mohr s.n. [Huatusca, 1857] was first deter- 
mined as V. officinalis, then as V. recta, then V. scabra, and 
then V. canescens by various workers 1 

Jackson (l895) and Lara (1919) reduce V. menthaefolia to syn- 
onymy under V. officinalis L., as Jackson also does V. setosa , 
while Walpers (I8ii5) lists it among his doubtful species. It is 



1U2 PHYTOLOGIA Vol. 10, no. 2 

discussed by Coulter (I89U) and by Small (1903, 1913, 1933) under 
the name V. officinalis. This, however, is a case of misidenti- 
fication. Cory (1937) cites V. officinalis from the Timber Belt, 
Coastal Prairies, Rio Grande Plains, and Blackland Prairies of 
Texas, while Rydberg (1932) also records it from Texas. It is 
probable that Cory's records, at least, are based partly on V. 
halei material and partly on material of V, menthaefolia . Wiggins 
notes for his no. S$OQ that the inflorescence is not glandular. 

Perry (1933) cites the following $0 additional specimens not 
as yet seen by me: CALIFORNIA: San Diego Co.: Abrams 3i;06 (E, G); 
Carlson s. n. [San Diego, 17 April I9I8] (G); Macbride & Pay- 
son 781 (G); Edw. Palmer 308 (E); M. F. Spencer 971 (G), llilTi 
(G)j Thurber $$$ (G) . MEXICO: Baja California: Bartram s.n. 
[Tia Juana, 1 Feb. 1920] (D); L^ Schoenfeldt 2915 (G) . Chihuahua: 
Pringle lg99 (E) . Coahuila: Gregg 11 (E), 265 (E, G), 276 (E) , 
U06 (E); Edw. Palmer I9I (E, G), 10U2 (D, G) . Durango: Nelson 
li577 (E)j Edff. Palmer 153 (E, F, G) . Guanajuato: Duges s.n. [Sir- 
ena Mountain, I89U] (G)j Hartweg 175 (K — type). Hidalgo: Galeotti 
778, in part [Moran] (K); Rose, Painter , & Rose 8753 (G) . Mexico: 
Bourgeau 51f7 (W)} Pringle 853U (D, E, F, G)j Rose , Painter , & 
Rose 8382 (G). Michoacin: Arsene 2798 (G)j Gregg 823 (E) . Morelos: 
Pringle 9529, in part (F) . Oaxaca: Pringle 5715 (G); L. C. Smith 
27 (G). QuerStaro: Arsene & Agniel 102U2 (E, F, G) . San Luis Po- 
tosl; Edw. Palmer II4I , in part (E, F, G, W)j Parry & Palmer 717 
(G) . Vera Cruz: Seat on 7 (F, G) . She notes that "In the speci- 
mens cited from California south including Sinaloa, the inflor- 
escence is more densely strigillose than in the collections from 
the southern part of Mexico, the calyxes are about 1 mm. longer, 
with teeth strongly xmequal and the subtending bracts often as 
long as the calyxes . Although this apparently indigenous species 
has been known generally as V. officinalis , it has somewhat 
harsher pubescence and is scarcely, if at all, glandular. The 
fruiting calyx tends to be connivent, concealing the apex of the 
schizocarp rather than open and disclosing it. Perhaps these are 
differences only of degree and may be merely variation of V. of- 
ficinalis ; nevertheless, for the present it seems preferable to 
retain the name V. menthaefolia for the American representative." 
Her Lyonnet 3.n. [Lomas de Santa F5, July I928] is probably what 
is cited hereinafter as I^onnet 33U , and her Gregg li06 , also from 
the New York herbarium, is probably what I cite below as Gregg s. 
n. [valley of Parras, April U, '97] . The Edw. Palmer 356 which 
she cites is regarded tiy me as V. pinetorum Moldenke . 

In all, 310 herbarium specimens, including type material of 
most of the names involved, and U mounted photographs have been 
examined by me. 

Citations: TEXAS: Bexar Co.: M. E. Jones 2829U (Po— 187971) . 
Uvalde Co.: M. E. Jones s.n. [west of Uvalde, April 26, 1931] 



I96J4. Moldenke, Monograph of Verbena lii3 

(Po— 187973). ARIZONA: Pinal Co.: Peebles 1|22U (Gg—267622) . Tu- 
ma Co.: M. E. Jones s.n. [north of Yuma, April 26, I906] (Du — 
151770, Po-^0890, Po— 70891). CALIFORNIA: Calaveras Co.: J. T. 
HoTrell 30087 (Gg). Riverside Co.: T. S^ Brandegee s.n. [Indian 
Wells, March 28, 1901] (Ca — IOU836) . San Diego Co.: Abrams 3UQ6 
(Ca— U07326, Dt, Du— 2UI87, Gg— 16215U, N, Po— U007. Po— 1^6399, 
W— 613970); T. S. Brandegee s.n. [San Diego, June I89U] (Ca— 
10ii875); Carlson s.n. [San Diego, April 18, 1918] (Gg— 31395); H. 
P. Chandler 5122 (Du— 77580); D. Cleveland 1135 (Sd— 6793), s.n. 
[March, April, May 187U] (Sd— 679U), s.n. [San Diego, June 187U] 
(Sd— 6782), 3.n. [San Diego, April 21, I88I] (Sd— 6795), s.n. 
[National Ranch, Jan. 20, 188U] (Sd— 678U), s.n. [National City, 
Apr. 20, I88U] (Rs- m592), s.n. [Sweetwater Valley, May 9, I88U] 
(Rs- m311, Sd— 6783); Collector xmdesignated s.n. (Sd— 6791); G. 
W. Dunn s.n. [21; April I89I] (Ca— 25151); Gander lla»3 (Sd— 
106lii), 219.22 (Sd— 11518), 222.6 (Sd— 11620), U972 (Sd— 20U35), 
608U (Sd— 21880); H. M. Hall 3857 (Ca— 56232); Herter s.n. [Bal- 
boa Park, May 18, 1937] (Sd— 210U9); F. W. Johnson 13li7 (N); £. 
Let s.n. [San Diego, Nov, 9] (W— 7192U); Of ford s.n. [La Jolla, 
9.IV.I932] (La); £. R. Orcutt s.n. [Apr. I889] (Ca— IOU83U); Edw. 
Palmer 308 (Be); Feirson 3379 (Po— 17780); Purer 65l5 (Du— 
251i355); Mj. F. Spencer 971 (N, Po— U7697), llOM (Po~U7191), s.n. 
[li. 7.1915] (Ob— 50820), s.n. [8/1^92-] (Ob— 50819); S. G. Stokes 
s.n. [San Diego, June 1895] (Du — 95U0); Stover s.n. [Point Loma, 
April 28, 1937] (Sd— 17067); Thurber 555 (T), s.n. [San Diego, 
May 1852] (N); E. Wall s.n. [San Diego, 16/5/31] (Ew); Wiggins 
3257 (Du— I8II8II7 Du— 3S50^2); C. B. Wolf 2100 (Ca— 527652, Du— 
230975, Gg— 2378U9, Rs— 1739). MEXICO: Aguascalientes: Rose & 
Painter 7799 (W — U5lUii) . Baja California: T. S^ Brandegee s.n. 
[San Gregorio, Feb. h, I889] (Ca— 169171), s.n. [Comondu Viejo, 
Feb. 17, 1889] (Ca— 169736), s.n. [Canon Salado, June 1, 1893] 
(Ca— 169127); W. E. Bryant s.n. [1888] (Gg— 311iOO); Carter , Al- 
exander, & Kellogg 2135 (Ca— 9l6m3, Du— 3U9167, W— 2022810); 
Gander 7356 (Sd— 2l|86U); H. S. Gentry hhh^ (Du— 26U17U, Ge); D^ 
A. Johansen 6OO (Du — 206U7I;); M. E. Jones s.n. [Tia Juana, April 
13, 1925] (Po— IIU3I6); Lewis & Epling s.n. [Burn, V23/U0] (Gg— 
38OU68); MacDougal 153 (N); Raven , Lewis , & Thonyson 12180 (Ca— 
171726); L. Schoenfeldt 2915 (W— 235268), s.n. [Mearns 2915] (N); 
Schoenfeldt & Mearns 2915 (Du— 9557); Wiggins 5508 (Mi) . Chihua- 
hua: H. Sj^ Gentry 15U2 (Fs, Ge, I); Pringle s.n. [Apr. 11, 1887] 
(N). Coahulla: Gregg s.n. [valley of Parras, 11/U7] (C); Edw. 
Palmer I9I (Ca— 10U85Ii, N, W— 336191), 10li2 (Pa, W— 56169, W— 
1323113). Durango: Correll & Johnston 20159 (Rf); H. S. Gentry 
8576 (Mi, N, W— 2022223); E. W. Nelson ^^^^^"(W— 33259U) ; Edw. 



llll* PHTTOLOGIA Vol. 10, no. 2 

Palmer 1^3 (Ca~10U82U, Me, N, W— 30U2la)j Patoni & Ochoterena 
7138 (Me); Waterfall 12^73 (Gg, Ok), 1261t8 (Gg, Ok); Waterfall & 
Wallis 13lili3 (Ok, W— 22971^9), 13677 TmITT Federal District; 
Barkley & Rowell 7U6U (Au— 1700U9, Mi, N); Botirgeau 360 (Br, S, 
W— 56182) J Lj. !_. Davis 207 (N); E. lyonnet 33h (N) ; Matuda 18822 
(N), 21190 (N), 26173 (Cb)j Miranda & Barkley 16007 (Au, N), 
. l6M9l|6 (Au), 16M988 (Au, N, Si), 16M997 (Au, N, Si), 16M998 (Au, 
'U, Si) J Miranda , Barkley , & Rowell 7U67 (Au— 1701141, Mi, N); 
Pringle TlUl (Me, 7t)j Rutten & Rutten-Pekelharing 735 (Ut— - 
59226a), Guanajuato; Hartweg 175 (Lu — isotype, N — isotype); Her- 
nandez Xolocotzi , Rupert , & Guevara X»2392 (N); Spivey 175 (Ca — 
9l67U7)i Waterfall & Wallis 13905 (Ok), 13921 (Ok). Hidalgo: F. 
A. Barkley 17Mlli7 (Aur-122307, Au--170082, N)} L^ I. Davis 208 
(N), 2^ (N) , 230 (N) J M. T. Edtrards 889 (Au, Du— 275502, Tu— 
3UU0U) i Fearing & Thompson 60 (Au) j G. L^ Fisher U6172 (W— 
1889832); Galeotti 778 , in part [Moran] (Br, F~photo, N—photo, 
Si — photo, Z — photo); Gold & Eheberle 21766 (N); H« E. Moore 
2810 (N); Moore & Wood Ul3U , in part (Ba), Ul96 (Ba); Rose , 
Painter , & Rose 8753 (N, W — U522U1) ; F. Salazar s.n. [Nopala, 
Aug. 1, 1913] (W— 1013228); Schnooberger 7983 (Mi). Jalisco: 
BArcena 221; (Me); Barkley , Paxson , & Rowell 766U (Au— 167039, N). 
Mexico: Barkley , Westlrmd, & Paxson 6U9 (Au— 123259, N), 667 
(Au); E. Lyonnet 33li (W— IO3U21I1) ; MacDaniels 553 (Ba); Eatuda 
I952I; (N), 2lla5 (N), 2ll;26 (N), 21872 (N), 21885 (N), 26101 
(Cb), 26829 (Cb), 27176 (Cb), 28975 (Z), 29136 (Cb), 29220 (Cb), 
29395 (Cb), 30913 (Ss); Moldenke & Moldenke 19852 (Es, N); 
Pringle 853U (Ca— 138820, Cm, Me, Me, Mi, N, Po~63878, S, Vt, 
W— 396358); Rose , Painter , & Rose 8382 (W--li5l877); H. H_. Rusby 
181 (N); A. J. Sharp U1|326 (N); Urbina s.n. [Junio de 1882] (Me); 
Waterfall & Wallis ll;053 (Ok) . Michoac5n: Arsene 2798 (Br), s. 
n. [Rinc6n, 15/7 A909] (N, W— U61;303), s.n. [Rinc6n, 25/7/1909] 
(W— 1;6U302); Hinton 11991 (It, Mi, N, Rf); Scheiy II6 (Mi, W— 
1822728), lig, in part (Mi), lh3 (Mi). Morelos: Moldenke & Mol- 
denke 19851; (N); Pringle 9529, in part (W— li62053) . Nuevo Le6n: 
L. !_. Davis 3.n. [Chipinque, March 8, I9U6] (Au— I7I989); Heard , 
Webster , & Barkley ll;5ll (Au); C. H. Mueller 2010 (Mi); Mueller & 
Mueller 157 (Me); Edw. Palmer lOla (Pa); M. Taylor U9 (N); S. S. 
White 15U2 (Mi, Oa) . Oaxacai C_. Con2atti1;207 (Me, W— 1082270); 
E. W. Nelson 19U3 (W— 250225); Pringle 5715 (Me, Vt). Puebla: 
Kenoyer s.n« [Popocatepetl, 7-2-38] (Fs); Smith , Peterson , & Tej- 
eda 3907 (W— 2397925). Querltaro: Agniel s.n. [Arsene 102^2] (W— 
1001585); Arsene 9998 (W— IOO3638); Basile 99 (W— 1268616) . San 
Luis Potosl: Edw. Palmer lljl , in part (Cm, Me, N, W — 397685); 
Parry & Palmer 717 (1°, Pa); Urbina s.n. [Junio de 1892] (Me). 



1961i Moldenke, Monograph of Verbena H^^ 

Slnaloa: H. S. Gentry 7027 (Aic— 21738, Mi); J. Gonzalez Ortega 
7^3 (Me), h^ (W— 1083502); Edw. Palmer 268 (w— 315567) ; Powell 
& Edmondson 911i (Au~193267); J_. N. Rose 1763 (W— 3OO63I1); Rose , 
Standley , & Russell 131^22 (N, W--6362U7), 13iai7 (N, W--636270)T 
Sonora: T. S^ Brandegee s.n. [Hermosillo, May lU, I892] (Ca— 
IOI4869); H. S. Gentry 219 (Du— 263856, Fs, Mi), 13la (Ak— 19906, 
Ca— 61^6322, Fs, Ge, I), li^ (Fs, Ge, I, S, W— 1689676), 7972 (N, 
W— 1978766); D. D. Keck U228 (Du— 263793); Maltby 20U (N, W— 
311;9li9); K^ F. Parker 82^N, W— 2130585); E. A. Phjjlips 329 
(Mi); Rose, Standley , & Rose 1293l| (N, W— 63^5117; Rose , Standley, 
& Russell 12lt5l (N, W— 635258), 13130 (N, W— 6359ii8); Shreve 6I88 
(Cm, Fs); Studhalter 1399 (W— 1685700), 1^7 (W— 16857UO), 1537~ 
(W— 1685757); S. S. White 6$k (Mi), 2938 (Mi), 3790 (Mi), iOOlT 
(Mi, N, W— 2I323I8); Wiggins 6053 (Du— 253UliO),"|te8 (Ca— 59^, 
Du— 2533li2, Gg— 263811, Mi, Po— 253I13U, W— 1739925) . Tamaulipas: 
H. H. Bartlett 10021 (Mi, W— lli91366); W. H. Lewis 5385 (Nb) . 
Vera Cruz: Medellin 99 (Me); C. T. Mohr s.n« [Huatusca, 1857] 
(W— 771859); H. E. Seat on 7 (W— 56171) . State undetermined: M. 
Halsted s.n. (T); Ruiz & Pavon 3.n. (Bm); WOlflin 3.n. [1815] (M). 
GUATEMALA: Huehuetenango: Steyemark 51655 (w— 19l;999U) . SWIT- 
ZERLAND: Probst s.n. [17.8.36] (Pb). CULTIVATED: Spain: Herh. 
Hort' Matrit. 58 (Q) . LOCALITY OF COLLECTION TOIDETERMINED: Col- 
lector undesignated U8 (Q); C. T. Mohr 651 (W— 771860) . 

xVHlBENA MERETRIX Moldenke, Phytologia 5: I33. I955. 

Synonymy: Verbena hispida x officinalis Dermen, Cytologia 7: 
170. 1936. Verbena hispida Ruiz & Pav. x V. officinalis L. ex 
Moldenke, R6sum6 366, in syn. 1959. Verbena officinalis L. ex 
Verbena hispida Ruiz & Pav. ex Moldenke, R6sum6 371, in syn. 
1959. 

Bibliography: Demen, Cytologia 7: 170. 1936; Moldenke. Phy- 
tologia 3: U67 (1951) and 5: 133. 1955; Moldenke, Biol. Abstr. 
30: 1093. 1956; Moldenke, Am. Midi. Nat. 59: 3514—355. 1958- 
Moldenke, R4sum6 22li, 366, 371, & 1^72. 1959; Moldenke, Phytologia 
8: 121 (I96I) and 9: 296. I963. . v 6 

This is an artificially produced hybrid between V. hispida 
Ruiz & Pav. and V. officinalis L., produced and described by Der- 
men from cultivated material in Massachusetts in 1936. The two 
parental species grow together in Cochabamba, Bolivia, and the 
hybrid may be expected there. They both grow also in Chile, but 
V. officinalis (a European species naturalized in Chile) is known 
thus far from only one province in which V. hispida also occurs. 
Both species, however, are weedy in characTer and spread rapidly. 
It is probably only a matter of time before they both grow widely 
in association with each other in that country. The ^brid prob- 
ably has no horticultural merit other than serving as a curiosity 
in botanical or experimental gardens. 



IhS PHTTOLOGIA Vol. 10, no. 2 

VERBENA MICROPHILLA H.B.K., Nov. Gen. & Sp. PI. 2: 272, pi. 133. 
1818 [not V. microphylla R. A. Phil., 18$7] . 

Synonymy: Verbena mlcrophylla Mart, in Mart. & Spix, Reise 
Bras. 2: 792. 1823. Verbena microphylla Hianb. ex Spreng. in L., 
Syst. Veg., ed. 16, 2: 7U9. 1825. Verbena microphylla Humb. & 
Bonpl. ex Steud., Nom. Bot., ed. 2, 2: 7^0. I8itl. Verbena mic- 
rophylla Htrnib. & Kunth ex Benth., PI. Hartw. 2li5. I8I46. Verbena 
microphylla Kunth apud Schau. in A. DC., Prodr. 11: 551. 181^7. 
Lantana microphylla Mart, apud Jacks, in Hook, f. & Jacks., Ind. 
Kew. 2: 29, sphalm. l89li. Glandularia microphylla (H.B.K.) Cabre- 
ra, Revist. Invest. Agric. 11: 332. 1957. Verbena laciniata f . 
purpurea Herter ex Moldenke, Rfisum^ Suppl. 3s 39, in syn. I962. 

Bibliography: H.B.K. , Nov. Gen. & Sp. Pi. 2: 272, pi. 133. 
1818; Mart. & Spix. Reise Bras. 2: 792. 1823; Spreng. in L., Syst. 
Veg., ed. 16, 2: 7U9. I825j Hook., Bot. Misc. 1: 170— 171. 1829; 
Steud., Nom. Bot., ed. 2, 2: 750. I8UI; D. Dietr., Syn. PI. 3: 
603. 18U3; Walp., Repert. Bot. Syst. I4: 2U. I81i5; Benth., Pi. 
Hartw. 2U5. I81i6; Schau. in A. DC., Prodr. 11: 551 & 552. l8U7j 
R. A. Phil., Linnaea 29: 21. 1857; Wedd., Chloris Andina [Castel- 
nau Exped. Bot.] 2: 156. i860; Wedd., Chlor. And. 2: 156—157. 
I86I; Griseb., Abhand. Kaiser. Gesell. Wissen. Getting. 21;; 
[Symb. Fl. Argent.] 276. 1879; F. Phil., Cat. PI. Vase. Chil. 221. 
1881 J Lillo, Fl. Tucumin 9U. I888; H. H. Rusby, Mem. Torrey Bot. 
Club ii: 2l|li. 1895; Jacks, in Hook, f . & Jacks., Ind. Kew. 2: 29 
(I89I1) and 2; 1179. 1895; Kuntze, Rev. Gen. Pi. 3 (2): 256. I898; 
R. E. Fries, Nov. Act. Soc. Sci. Upsal., ser. h, 1 (1): 110. 
1905; Hayek in Engl., Bot. Jahrb. Ii2: I6U. 1908; Herzog, Bolivia 
3: li3. 1916; Sanzin, Anal. Soc. Cient. Argent. 88: 98, 129, & 1^ 
1919; Stapf, Ind. Lond, 6: U30. 1931; R. Espinosa, 6'kol. Stud. 
Kordillerenpfl . 37 & 38. 1932; Moldenke, Prelim. Alph. List In- 
valid Names U7. I9UO; Moldenke, Known Geogr. Distrib. Verbenac., 
[ed. 1], 3U, 35. UO, Ux, &. 101. 19U2; Moldenke, Alph. List Inval- 
id Names U8. 19U2; Raimondi, Bol. Mus. Hist. Nat. Jav. Prado 7: 
2h2. 19U3; Moldenke, Holmbergia U: 152. 19U5; Moldenke, Bot. Gaz. 
106: 162. 19U5; Schnack & Covas, Darwiniana 7: 71, 72, & 7U, pl. 
III. 19h$; Moldenke, Alph. List Cit. 1: 10, 77, & 266. I9i;6; Mol- 
denke, Phytologia 2: 33S» 19h7; Moldenke, Alph. List Invalid 
Names Suppl. 1: 25. 19U7; Moldenke, Castanea 13: 116. I9U8; H. N. 
& A. L. Moldenke, PI. Life 2: lU;. I9U8; Moldenke, Alph. List Cit. 
2: 375, 378, 379, 536, & 599 (19i;8), 3: 66O, 663, 705, 735, 80U, 
807, 812, 893, 901, 909, 931, 951, 952, 956, 968, & 97li (19U9), and 
ii: 1073, 1127, 1203, 12U8, & 1293. 19i;9; Moldenke, Known Geogr. 
Distrib. Verbenac, [ed. 2], 70, 73, 98, I06, & I98. 19U9; Molden- 
ke, PiQTtologia 3: 286, 289, & 290 (1950) and 5: 96. 1951i; Moldenke, 
Mem. N. Y. Bot. Gard. 9: 177. 1955; Moldenke, Inform. Mold. Set 51 
Spec. k» 1956; Soukup, Biota 1: I8I. 1956; Cabrera, Revist. Invest. 
Agric. 11: 332. 1957; Moldenke, Rlsum6 81, 85, 115, 119, 122, 127, 
296, 306, 361;, 370, & Ii72. 1959; Moldenke Phytologia 8: 123. 1961; 
Moldenke, R^sumS Suppl. 3: 13—15, 39, & UO (1962), U: 17 (1962), 



1961i Moldenke, Monograph of Verbena li^y 

and 6: 6. 1963j Moldenke, Phytologia 8: U72 (1963) and 9: 70, 128, 

Illustrations: H.B.K., Nov. Gen. & Sp. PI. 2: pi. I33. I8l8' 
Sanzin, Anal. Soc. Cient. Argent. 88: 129. 1919} Schnack & Covas 
Darwiniana 7: pi. III. I9h$, * 

Dwarf prostrate perennial herb, often suffruticose at the 
base J stems prostrate, spreading, trailing or creeping, rooting 
at the nodes, forming low mats, $—10 cm. tall, sometimes to 6.U 
m. in diameter, much branched, strigose-hispidulous, with vigor- 
ous shoots sometimes 1 m. long; branches prostrate; branchlets 
pubescent-hirtous; leaves decussate-opposite, small, pale-green, 
6—10 mm. long, to 8 mm. wide, deeply cut or tripartite, cxineate 
into a subpetiolar base, hairy, the segments obovate-oblong, ob- 
tuse at the apex, revolute along the margins, entire or the 'mid- 
dle one trifid and the lateral ones bifid; spikes terminal and 
lateral, short, subsessile, capitate, many-flowered; bractlets 
lanceolate, half as long as the calyx; flowers very fragrant, 
with a delicate perfume; calyx about 6 mm. long, very hairy, * the 
5 teeth short, ovate, acute at the apex; corolla varying from 
blue, purple-blue, pale-blue, clear-blue, "blue and lavender" or 
••whitish-blue" to pale bluish-lilac, lavender, pale-lavender * 
lilac, pale-lilac, "lilac-white", violet, red-violet, violet- 
purple, purple, Parma violet-mauve, or very pale-mauve, or even 
pale carmine-pink or rose-pink, occasionally white or turning 
white in drying, sometimes described as "white to purple", glab- 
rous, its tube scarcely twice as long as the calyx, the limb med- 
ium-large, about 6 mm. wide, pilose in the throat, the lobes e- 
marginate; anther-appendages horn-shaped, equaling the mouth of 
the corolla-tube, becoming blackish; fruit half as long as the 
calyx; cocci lightly tuberculate on the back. 

The type of this interesting high Andean species was collected 
by Friedrich Heinrich Alexander von Humboldt and Aim^ Jacques Al- 
exandre Bonpland "in frigidis Andium Quitensium alt. li|82 hex." 
Ecuador, and was deposited in the Kunth Herbarium at Berlin. The 
original publication date is often given as "1817", but according 
to the late Dr. John H. Bamhart is actually I8I8. The ty^Q of 
V. laciniata f . purpurea was collected by Wilhelm Gustav Herter 
( no. 1530) in Uruguay. 

It is not at all certain to me that the Uruguayan specimens 
cited below are actually conspecific with the remainder of the 
material cited here. One would hardly expect such a high Andean 
species as this to be found also in Uruguay. More study of the 
material is required to settle this point. It should also be 
noted that the V. microphylla or R. A. Philippi is Junellia min- 
utifolia (R. A. Phil.) Moldenke. Whether the V. microphylla~ of 
Martius is correctly placed here or not is also doubtful . The V. 
glandularia JOrgensen recorded by me in my 19$9 publication as a" 
synonym of V. microphylla is actually a sjTionym of V. laciniata 
(L.) Briq. It is possible that V. nivea Moldenke and its f . rosea 
Moldenke are not specifically distinct from V. microphylla, 



Ilt8 PHYTOLOGIA Vol. 10, no. 2 

Walpers (18U5) places V_. microphylla in his Section Verbenaca, 
Subsection Inermes , GiX)up Foliosae, Subgroup Micranthae , and Sec- 
ondary Subgroup S chl z ophyllae with 9 other species . It has been 
found by collectors in gravel, rocky clay soil, rocky sandy soil, 
and dry soil in general, in stony habitats, on sandy hills, high 
or dry hillsides and pampas, in fields and dry open sandy campos, 
on puna and dry sandy open plains, in dry meadows and volcanic 
soil, in dry or sandy places, dry disturbed roadsides in large 
open grassy pdramos, and in moist sheltered spots, at altitudes 
of 2135 to U660 meters, blooming frcan October to August, fruiting 
in February, April, May, and October. The only ccRmnon name re- 
corded for it is waltamera". 

Budin 6511 and 7U92 are described as having white corollas and 
may be worthy of nomenclatursLL designation, but Mrs. R. S. Shep- 
ard says of her no. 12 collection "flowers white to purple" and 
CArdenas affirms that the flowers are lilac, but "turning irtilte 
in drying" — his plant has almost the exact habit and appearajice 
of V. nivea Moldenke. Fosberg & Giler found V. microphylla "on 
dry overgrazed gentle slope at foot of hill", Vargas "in clay and 
rocky places", and Brook "in bleak and almost desert area partly 
due to saline and mineral deposits, among cacti and a few small 
plants". West calls it a "perennial herb, prostrate rosettes to 
30 cm. in diameter, on level open puna among shoirt grasses", 
while Sandeman says that it "forms small mats in full exposure; 
flowers usually Parma violet mauve, a pale carmine pink form not 
infrequent; very frequent in a restricted locality." Schulz 6663 
bears a notation on its label affirming that the flowers were 
purple-blue, but that some specimens have white flowers. Steyer- 
mark fovmd the species with "stems hanging down clay bsmks"; 
Haught found it "abundant on roadside" and Parodi says it is coti- 
mon in cultivated ground at Lake Titicaca. 

Herbarium material of this species has been misidentified and 
distributed under the names V. ciliata Benth., V. diffusa VJ'illd., 
V. dissecta Willd., V. erinoides Lam., and V. multifida Rxilz & 
Pav. On the other hand, the Venturi 3180 , distributed as V. mic- 
rophylla , is actually V. dissecta Willd.; JOrgensen 1026 is V. 
glandullfera Moldenke; Asplund 6862, Hartweg 1351 , A. S. Hitch - 
cock 21739 , I^ Holmgren 967, Kuntze s.n. [Oruro, 11^/3/92] , Mille 
liO, Rimbach 175, Rose & Rose 22312 , Spruce 5065, and Steyermark 
^853 are y, laciniata (L.) Brig. ; JPf gens en 1026 , in part, is V. 
tenera Spreng.; McAtee 331^9 is V. tenuisecta Briq.; Buchtien 1102 
is V. weberbauerl Hayek; and Finnin 366 , A. S. Hitchcock 21737 , 
Pachano II4J4 & 156, Rimbach I76, and Rose & Rose 22U00 & 23906 are 
Hierobotana inflata (H.B.K.) Briq. The JOrgensen 1737 cited by 
me in my Alph. List Git. 2: 599 (19U8) as V, microphylla is actu- 
ally V. tenera Spreng . 

Schauer (18U7) says that V. radio ans Gill. & Hook, is similar 
to V. microptylla in habit, but differs in its larger leaves 



1961i Moldenke, Monograph of Verbena lU9 

which are completely glabrous and more divided, beccaning yellow- 
ish in drying. 

The Linden 150 collection cited below bears a label reading 
"Mexico", but this is probably an error for Colombia. Haught 
32U2 has printed labels reading "Los Rios", but was actually col- 
lected "due west of Lataicunga", which would place it in the prov- 
ince of Leon, as is indicated on the United States Natior*al Her- 
barium specimen which has "Los Rios" crossed out and "Leon" svib- 
stituted in longhand. This collection was the basis of my record 
of the species from Los Rios in my R^sumS (1959) « Raimondi (19U3) 
cites his no. 105l6 from Santiago de Huanta, Pern, while Cabrera 
(I957) cites Keidel s»n. , Krapovickas 3lU8, and Cabrera 7772 , 
8666 , 8981 , and 91]h» 

In all, 112 herbarium specimens have been examined by me. 
Citations: COLOMBIA?: Department undetermined; Linden 150 (Br). 
ECUADOR: Azuay: Fosberg & Giler 23211 (N, W— 2109897); Wiggins 
19851 (Du— 3II6I5, Ug). Chimborazo: Hartweg 1351 (Br); Schimpff 
720 (N) . Cotopaxi: Barclay & Juajibioy 8020 (N) . Leon: Haught 
32II2 (N, W— I7O8OO6); Hartweg 176 (Br). Tunguragua: W, H. Camp 
E.2U25 (N, W— 205698U) . Province undetennined: Spruce 5065 (N, S, 
S). PERU: Ancash: Sandeman U599 (K) . Ayacucho: R. Ferreyra 5507 
(N) . Cuzco: Cook & Gilbert 5U9 (W— 603752); F^ L^ H err era s»n. 
[Cuzco, July 1923] (W— 1190005); Hicken 56 (S); C. Vargas 9827 
(Ca~6U9009). Lima: R. Ferreyra 3531 (N, Ug); Nuflez 2717 (W— 
212073U). Puno: P. Aguilar s.n. (Ss); Ellenberg 261 (Ut — 
115392b); R. Ferreyra 2605 (N); Ochoa 2029 (W— 212351i2); F. W. 
Pennell 13359 (N, S, W— 13li0553); Sandeman 3927 (K); Sharpe £2 
(K), 139 (K); Mrs. R. S. Shepard 12 (N, ¥—119777?); Soukup 70 
(Ca— 770036, N, W— I7753U8), s.n. [Nov. 2h, 1938] (Ew); R. S. 
Willians 2510 (N) . Province undetermined: Steir s.n» [Matucana] 
(Mi); C. Watkins s.n. [Tuapata, I9I6] (W— 1059580) ; Whiteley s.n, 
[Rio Casnipueta, I869] (Bm) . BOLIVIA: La Paz: Asplund 50 (S) , 
2268 (S, Us), 21U5 (S, Us), 2la5 (S, Us), 2800 (S, Us); M. Bang 
161 (C, Pa, W--7198U, W— lla67UO), I6la (c. Pa, W— 1323109); 
Buchtien U33 (N), 758 (W— II3U88U), s.n. [La Paz, 1912] (La); Man- 
don 525 (Mi, N, S, T); Parodi IOQ98 [Herb. Osten 22533] (Ug) . Or- 
uro: Asplund 3222 (S, Us); W. M^ A. Brooke 5233 (N); Kuntze s.n, 
[Oruro] (N); Troll 2919 (B). Potosi: Asplund 3001 (S, Us); Cir- 
denas 3U5 (W— 1573325); Fiebrig 2613 [Herb. Osten 15218] (Ug, W— 
1177998); J . West 6353 (Ca— 56U990). Province undetermined: 
Balls 59 3U [Villason, S. Bolivia] (K); R. S. Williams 2510 [Juli- 
aca] (W— 113U879). URUGUAT: Herter 1530 [Herb. Herter 87li30] 
(Ca— 505236); Osten 3522 (Ug) . CHILE: Malleco: Ojiva I8 (C«~ 
66U83I). ARGENTINA: Buenos Aires: Carette s.n, [Monte Hermoao, 
Afio 1916] (N) . Catamarca: Peirano s.n. [Herb. Inst. Miguel Lillo 



150 PHYTOLOGIA Vol. 10, no. 2 

328Ui] (N), s.n. [Herb. Inst. Miguel Lillo 32933] (N). Jujuy: 
Budin 7^92 [Herb. Inst. Miguel Lillo 32798] (N, Ug— U937); Claren 
11309 (S), 1132U (S), 11U68 (S); R. E. Fries 970 (S); T^ Meyer 3_. 
n, [Quebrada de Salitro, Feb, 23, l9U0j Herb, Inst, Miguel Lillo 
3Kb01] (N), 3.n. [Herb. Inst. Miguel Lillo 314+02] (Mv) . Los An- 
des: Budin 6511 [Herb. Inst. Miguel Lillo 32797; Herb. Osten 
23006] (Ca— 165679, N, Ug, Ug--U936) , Mendoza: Sanzin 132 [Herb, 
Osten 12818] (Ug). Tucumin: Bruch s.n, [Valle de Tafl, I9O8] (N)j 
Ullo U23I [Herb, Osten 8U72] (Ug), 5039 [Herb, Inst. Miguel Lillo 
31i;7li; Herb. Osten 8U70] (N, Ug), 11152 [Herb, Osten 8U73] (Ug); 
Schreiter 871U [Herb. Inst, Miguel Lillo 32902; Herb, Osten 2299ltl 
(N, Ug), 3,n, [Cumbre Alta del Chorro, Dec, 1917; Herb, Osten 
12195] (Ug), s,n. [Infiemillo, Dec, 1, 1917; Herb, Osten 12197] 
(Ug); A. G. Schulz 6663 (Z) , LOCALITY OF COLLECTION UNDESIGNATED: 
Herb, A. Gray s,n, (T). 

VERBENA MINUTIFLORA Briq, ex Moldenke, Suppl, List Invalid Names 
9, in syn, (19la), Phytologia 7: 8U~85. 1959. 

Bibliography: Moldenke, Suppl, List Invalid Names 9. 19Ulj 
Moldenke, Alph. List Invalid Names U8. 19U2; Moldenke, Lilloa 6: 
U32. 19l|2; Moldenke, RisumS U9U & h9$» 1959; Moldenke, RisumI 
Suppl, 1: 7, 23, & 25. 1959; Moldenke Phytologia 7: 8U— 85. 
1959; Angely, Fl. Par an. 16: 79 (I960) and 17: U6. I96I. 

Herb or shrub, to 3 m. tall, much branched; stems erect, 
stiff, very sharply tetragonal, glabrous or practically so, con^ 
cave bet-vreen the margins in drying; branches and twigs numerous, 
stiffly ascending, medium-stoutish, very sharply tetragonal, 
concave between the angles in drying, glabrous or practically so, 
brunne scent in drying, somewhat contracted at the nodes; nodes 
annulate; principal intemodes 3 — 8.5 cm. long; leaves decussate- 
opposite, firmly chartaceous, uniformly green on both surfaces, 
divaricate or ascending, narrowly elliptic, 1.1 — 3*3 cm. long, 
3—6 mm. wide, acute at the apex, somewhat narrowed to the ses- 
sile base, finely strigillose above and on the venation beneath, 
3-veined; midrib and the two secondaries very slender, impressed 
above, slightly prominulous beneath, the secondaries subpaurallel 
to the midrib from above the base almost to the apex, not anas- 
tomosing; inflorescence terminating the branches and twigs, sub- 
paniculate; individual spikes usually in 3's, sometimes solitary 
or paired, abbreviated, less than 1.5 cm, long, less than 1 cm, 
wide, many-flowered, pedunculate; pedvincles very slender, 
slightly gray-strigillose, especially toward the apex, angulate, 
sulcate in drying; bracts paired under the branches of the in- 
florescence, resembling the leaves in all respects but smaller; 
bractlets linear-subulate or lanceolate, one pair subtending the 
group of 3 spikes and one subtending each flower in the spike, 
the latter 1.5 — 2 mm. long, acute at the apex, very minutely and 
obscurely grayish-strigillose, closely appressed to the calyx; 
calyx tubular, 2,5 — 3 mm. long, 1 mm, wide, densely and very 



1961i Moldenke, Monograph of Verbena l5l 

conspicuously" white-strigose, the teeth scmewhat unequal; corolla 
hypocrateriform, lilac or rose to violet, its tube very slender, 
about h mm. long, glabrous, its limb about 2 mm, wide, strigil- 
lose beneath, glabrous above. 

The type of this interesting species was collected by Csqptain 
P. King, R. N. ( no. 78) at Montevideo, Uruguay, aind is deposited 
in the Delessert Herbarium at the Conservatoire et Jardin Botan- 
iques at Geneva. The type specimen was annotated " Verbena minu- 
tiflora Briq." by Briquet before his death and was photographed 
by Macbride as his Type Photograph no. 2U693, but a valid de- 
scription of the plant does not appear to have been published be- 
fore 1959. The species is related to V. montevidensis Spreng., 
but is easily distinguished in any series of specimens . It in- 
habits campos, hedges near streamlets, marshy or wet swampy cam- 
pos, marshes, swamps, swampy places, and secondary woods on var- 
zea land, at altitudes of 885 to 1550 meters, and has been col- 
lected in an thesis in October, December, January, and March, and 
in fruit in December and January. Herbaritun material has been 
misidentified and distributed under the names V. isabellei Briq., 
V. litoralis L., and V. montevidensis Spreng. In my I9UI and 
19lt2 publications I regarded V. minutiflora as a synonym of V. 
isabellei , while in my 1959 work I regarded it as a synonym of V. 
montevidensis . Rambo states that where V. minutiflora grows the 
temperature varies from 0° to 25° C, and there are two meters of 
rainfall per year, with rare snow. 

In all, 3$ herbari\im specimens and 2 mounted photographs have 
been examined by me. 

Citations: BRAZIL: Parand: Dusgn 6802 (S), 851;6 (N, S, W— 
Iii8l771), 9313 (S); Hatschbach k^h (N, N), 728U (Ca); JOnsson 
9l;2a (Ca— 50I69I, N, S, W~lli8l969)j Mattos & Moreira s.n. [5/58; 
Herb. Inst. Hist. Nat. 5261] (Z) . Rio Grande do Sul: Rambo 30979 
(N), 3U721 (N, N, S), h937k (N, S), 51616 (W— 2102102), 5U985 (B). 
Santa Catarina: Jj^ £. T_. Mttller s.n. (P); Rambo 5l6l6 (Le, S); 
Reitz 3260 (N, S, Sm, W— 2321355), 3l;03 (N, W— ?1lil669); Reitz & 
Klein 769U (Ita, S), 786U (Mm); Smith & Reitz 911^6 (Ok, W— 
225lli98) . URUGUAY: Capt. P^ Smith 78 [Macbride photos 2U693] 
(Kr — photo of type, N — photo of type). 

xVEKBMA MOECHINA Moldenke, Revist. Sudam. Bot. U: 19, hyponym 
(1937); Moldenke in Gleason, New Britton & Br. Illustr. Fl. 
pr. 1, 3: 126, 131, & 132. 1952. 
Synonymy: Verbena angustifolio-atricta Bigelm., Am. Joum, 
Sci. I|6: 101, hyponym. I8U4. Verbena atricto-angustifolia 
Engelm., Am, Journ. Sci. 1^6: 101, hyponym. l8Uli. Verbena angus- 
tifolia X stricta Palmer, Ann. Mo. Bot. Gard. 3: 292, hyponym. 
1916. Verbena simplex x stricta Palmer, Ann. Mo. Bot. Gard. 22: 
629, hyponym. 1935. Verbena angustifolia x stricta Kellogg ex 



152 PHYTOLOGIA Vol. 10, no. 2 

Moldenke, Revist. Sudam, Bot, U: 19, in syn. 1937. Verbena an- 
gustifolla X stricta Rydb. ex Moldenke, Revist. Sudam. Bot. lU 19, 
in syn. 1937. Verbena stricta x angustifolia Eggert (in part) ex 
Moldenke, Revist. Sudam. Bot. U: 19, in syn, 1937. Verbena sim- 
plex Lehm. x V. stricta Vent, ex Moldenke, Prelim. Alph. List In- 
valid Names 1;8, in syn. 19^0. Verbena simplex x stricta Gates, 
Fl. Kans. 191, nom. nud. I9U0. Verbena stricta x angustifolia 
Blankinship ex Moldenke, Suppl. List Invalid Names 10, in syn, 
19lil. Verbena angustifolia x stricta Bush ex Moldenke, Alph, List 
Invalid Names Suppl. 1: 22, in syn, 19li7. Verbena stricta x has- 
tata Gates (in part) ex Moldenke, Alph, List Invalid Names Suppl. 
1: 27, in syn. 19U7. Verbena stricta x angustifolia Gates ex 
Moldenke, Alph, List Invalid Names Suppl. 1: 27, in syn, 19li7. 
Veixmica angustifolia Michx. ex Moldenke, Am, Midi. Nat, $9: 355, 
in sjm. 1958. Verbena spicata Pammel ex Moldenke, Am, Midi. Nat. 
59: 355, in syn. 1958. Verbena stricta var. angustifolia Martens 
ex Moldenke, Am. Midi. Nat. 59; 355, in textu. 1958, Verbena has- 
tata X bract eosa Pammel ex Moldenke, Am. Midi. Nat. 59: 355, in 
syn. textu (1958), Rlsum6 365, in syn. 1959. Verbena simplex x 
stricta Hitchc, ex Moldenke, Am. Midi. Nat. 59: 355, in syn. tex- 
tu (1958), R^svmil 371^, in syn, 1959. Verbena stricta x angustifol- 
ia Pammel ex Moldenke, Am. Midi, Nat. 59: 355. in syn. textu 
(1958), R5sum6 375, in syn. 1959. Verbena stricta x angustifolia 
Popenoe ex Moldenke, Am. Midi, Nat. ^9: 355, in syn. textu (1958), 
R^sum6 375, in syn. 1959. Verbena stricta x urticaefolia Letter- 
man ex Moldenke, Am. Midi. Nat, 59: 355, in syn. textu (1958), 
R6sum5 375, in syn. 1959. Verbena stricta x urticaefolia Pammel 
ex Moldenke, Am. Midi. Nat, 59, 355, in syn. textu (1958), R6sum€ 
375, in syn. 1959. Verbena angustifolia x stricta Eggert ex Mol- 
denke, R6sum6 357, in syn. 1959. 

Bibliography: Engelm., Am. Journ. Sci. U6: 101. iSUh; E. J, 
Palmer, Ann, Mo, Bot, Gard, 3: 292. 1916; Kanda, Bot, Gaz, 69: 
5U-— 71, pi. 6, fig. 2, I92OJ E, J, Palmer, Ann, Mo. Bot, Gard, 
22: 629. 1935; Moldenke, Revist, Sudam. Bot, U: 19. 1937j Mol- 
denke, Prelim. Alph, List Invalid Names U; & U8, 19l|0j C. C, 
Deam, Fl. Ind. 797 & 1232. 19i|0j Moldenke, Suppl, List Invalid 
Names 10, 19ia} Moldenke, Alph, List Invalid Names U5 & 50, 19li2i 
Moldenke, Known Geogr, Distrib, Verbenac, [ed, 1], 6, 7, 9, 10, 
& 101, 19li2: Moldenke. Bot. Gaz. IO6: I60. 19U5; Moldenke, Cas- 
tanea 10: 38 & 39. 19U5; Moldenke. Phytologia 2: 7ii & 115. 19li5; 
Moldenke, Alph, List Cit, 1: 28, U5, 80, 81, m3, m8, lU9, 159, 
181, 193, 262, & 267. 19U6j Hill & Salisb,, Ind. Kew. Suppl. 10: 
2U2. 19li7; Moldenke, Alph. List Invalid Names Suppl. 1: 22 & 27. 
19U7; Moldenke, Castanea 13: 112. 19^8; Moldenke, Phytologia 2: 
U78. 19h8; Moldenke, Alph. List Cit. 2: 35U, 390, 391, 39h, 395, 
397, U05, U07, U38, U66, U8l, 5U7, & 606 (19li8), 3: 657, 720, 
783, 790—793, 826, 828, 869, 887, & 973 (19^9), and h: 1137, 
1255, & 1261. 19U9i Moldenke, Knoim Geogr. Distrib. Verbenac., 



196U Moldenke, Monograph of Vertena 1^3 

[ed. 2], 12— Hi, 16—18, & 198. 19U9i Moldenke, Phytologia 3: 131 
(19U9), 3: 28U (19^0), 3: U67 (1951), and k: 6?. 19^2; Moldenke 
in Gleason, New Britton & Br. Illustr. Fl., pr. 1, 3: 126, 131, Sc 
132. 19^2; Moldenke, Phytologia U: 18$ & l86. 1933; S. F. Thome, 
Proc. Iowa Acad. Sci. 62: 182. 1955; Moldenke, Am. Midi. Nat. 59: 
355 — 356. 1958; Moldenke in Gleason. New Britton & Br., Illustr, 
Fl., pr. 2, 3: 126, 131, & 132. 1958; Moldenke, Rfisumfi 15—18, 
21, 22, 26, 22li, 357, 365, 37U, 375, 379, & U72. 1959; Moldenke, 
R6sum5 Suppl. 2: 3. I960; Moldenke, Phytologia 8: 121 & lii6 (1961) 
and 8: li05. 1962; Moldenke, R6suml Suppl. 3: U & 5 (1962), U: Hi 
& 18 (I962), 6: 2 (I963), and 7: 2 & 10. 1963; Gleason & Cron- 
quist, Man. Vase. PI. 580. 1963; G. N. Jones, Fl. 111., ed. 3, 
[Am. Midi. Nat. Monog. 7:] 213. 1963; Moldenke, Phytologia 9: Hi?, 
220, 356, & 359. 1963. 

Ill\istrations : Kanda, Bot. Gaz. 69: pi. 6, fig. 2. 1920; Mol- 
denke in Gleason, New Britton & Br. Illustr. Fl., pr. 1, 3: 132 
(1952) and pr. 2, 3: 132. 1958. 

This is a natural hybrid (also produced artificially by Kanda) 
between V, simplex Lehn. and V. stricta Vent, It occvirs ab\an- 
dantly where the ranges of the two parental species overlap in 
the central portions of the United States. It usually resembles 
V. simplex in habit, but has broader leaves, which are often el- 
TTptic or elliptic-ovate, more or less densely short-pubescent or 
velutinous nebeath, the stems and branches usually densely pubes- 
cent, and the spikes poorly and irregularly fruited. From V, 
stricta it differs in its more slender and poorly fruited spikes, 
smaller flowers, and narrower leaves. 

The binomial name was proposed by me originally in 1937, based 
on V. angustifolio-atricta of Engelmann (l8Uli), which, however, 
was published by him vrithout description or reference to a spec- 
ific collection. I am therefore dssigr^ting as logotype the 
specimen collected by Charles Clemon Deam ( no, 20357) one mile 
west of Palmyra, Harrison County, Indiana, on June 22, 1916, and 
deposited in the Britton Herbariim at the New York Botanical 
Garden. Deam reports that the plant was "Plentiful along road- 
side very variable in width and pubescence of leaves, and I 

suspect some are hybrids with V, stricta Vent., varies also 
greatly according to quality of soil." 

Engelmann' s V. 3 trie to-angus tif olia was apparently based on 
his unnumbered collection from Saint Louis, Missouri, gathered 
in July, I8U2, and deposited in the Torrey Herbarium at the New 
York Botanical Garden. V. angustifolia x stricta Eggert is 
based on Eggert' s unnumbered collection of July U, I896, frcm 
Pacific, Franklin County, Missouri, deposited in the herbarium 
of IgWa State College at Ames. V. angustifolia x stricta Bush 
is based on Bush 15697, collected on barrens in Stone Coxrnty, 
Missouri, and deposited in the herbariimi of Kansas State College 
at Manhattan; V. angustifolia x stricta Rydb. is based on Rydbeirg 
& Imler h3h, from roadsides in Montgomery County, Kansas, also 
deposited in the Kansas State College herbartum; V. hastata x 



15U PHTTOLOGIA Vol. 10, no. 2 

bracteosa Pammel is based on Painmel s.n. from Steam Boat Rock, 
Hardin County, Iowa, collected in September, 1912, and deposited 
at Iowa State College, Ames; V. stricta x angustifolia Blankin- 
ship is based on Blankinship s.n. , collected in Greene County, 
Missouri, on July 17, 1919, and deposited at Pomona College, 
Claremont, California} V. stricta x angustifolia Popenoe is based 
on E. A. Popenoe s.n. from Topeka, Shawnee County, Kansas, col- 
lected on July 7, 1879, and deposited at Kansas State Collegej 
both V. stricta x angustifolia Pammel and V. stricta x urticaefo- 
lia Pammel are based on Pammel s.n. from Oxford Junction, Jones 
County, Iowa, collected on July 25, 1919, and deposited at Iowa 
State College. 

Verbena simplex x stricta Hitchc. is based on A. S. Hitchcock 
s.n. from Montgomery County, Kansas, collected in August, I896, 
and deposited at Kansas State College; V. stricta x urticaefolia 
Letterman is based on Letterman s.n, from Allenton, Saint Louis 
County, Missouri, collected on Jime 28, 1911, and deposited at 
Iowa State College; V. stidcta var. angustifolia is based on M. 
Martens s.n. ., collected at Saint Louis, Missouri, and deposited 
in the herbarium of the Jardin Botanique de I'Etat at Brussels; 
Veronica angustifolia Michx, is based on }[. N. Andrews , Jr. , s.n., 
collected at Hercxilaneum, Jefferson County, Missouri, on May 30, 
1936, and deposited at the University of Massachusetts, Amherst; 
and V. spicata Pammel is based on L. H. Pammel s.n. , collected on 
June 25, 1888, at 'Washington, Franklin County, Missoxiri, deposit- 
ed at Iowa State Collage. The type of V. angustifolia x stricta 
Palmer is E. J. Palmer 2932 from Jasper County, Missouri, depos- 
ited at the Missouri Botanical Garden in St. Louis. V. stricta x 
angustifolia Eggert is actually in part this hybrid and in part 
xV. engelmannii Moldenke, while V. stricta x hastata Gates is in 
part this, in part xV. rydbergii Moldenke, and in part V. stricta 
Vent. 

Engelmann (181U;) comments as follows: "I^brids of V. angusti - 
folia with any but V. stricta , and of V. bracteosa irith any but 
V. urticaefolia , or of V. aubletia, the only remaining species 
in this region, I have not yet found." The Bush s.n. [Oct. 9, 
1936] collection, cited below, was annotated by Miss Periy as nonr- 
typical V. simplex Lehm, 

Letterman s.n. [Allenton, June 188U & June 28, 1911] have very 
large leaves resembling those of V. hastata L., but much firmer in 
texture, with the upper ones narrow like those seen in V. simplex . 
L. H. Pammel s.n. [Sandusky, June & July 1912] is narrow-leaved 
and much like true V. simplex in appearance, but the same form is 
found with the broad-leaved hybrid by genetic segregation. His Sj. 
n. [Oxford Junction, Jvly 25, 1919] consists of two branches of 
the broad-leaved form and one dense branch of the narrow-leaved 



196U Moldenke, Monograph of Verbena 1^5 

form exactly similar to the Sandusky specimen. He apparently 
proposed two different hybrid designations for these parts of the 
same collection (see above). Somes 3301 is a mixture of xV. moe- 
china and V. simplex * 

E. J. Palmer records the hybrid frcm Jasper County, Missouri, 
but I have not as yet seen any substantiating specimens from that 
county. 

Collectors have found xV. moechina on high ground along road- 
sides, open dry ground, sandy prairies, lime uplands, rocky or 
clay ridges, bluffs, and sand ridges, in rather sandy soil, pas- 
tures, overgrazed prairie pastures, prairie pastures with lime- 
stone i?ock surfacing, creek bottoms, and barrens, on sandy hills, 
along riversides, and in the vicinity of stockyards; also in ui>- 
land sandy areas, open woods, rolling open pastured hillsides, 
and along roadsides in general, often closely associated with V. 
simplex and/or V. stricta . Fell actually says that it is "com- 
mon in pastures" in Illinois. It has been collected an anthesis 
and fruit fran May through September, at altitudes of 360 to 900 
feet. Herbarium material has been misidentified and distributed 
under the names V. angustifolia Michx., xV. blanchardi Moldenke, 
V. hastata L., xV. rydbergii Moldenke, V. simplex Lehm., and V. 
stricta Vent, Collectors record the common names "hoaiy vervsdn" 

and "la sacr^e verveine" . 

L. H^ Pammel s.n, [Cedar Falls, Sept. 28, 1920], identified 
as V. angustifolia , bears a notation by the collector "perhaps a 
hybrid with stricta ", while on his s.n. [Ames, 9-8-9U] he says 
categorically "hybrid between stricta and bracteosa" . C_. R. Ball 
s,n, [June 30, I898] was identified as V. angustifolia with the 
notation "hybrid possibly with stricta ". The label on F. E. Mc 
Donald s.n. [Peoria, Jul. I90U] says "evidently a hybrid". Shi- 
mek s.n. [Oct. lU, 1919] from an upland sandy area in Muscatine 
County, Iowa, was identified as V, stricta x angustifolia , pre- 
sumably by the collector, and is in part xV. moechina and in 
part typical V, stricta , while Ruth s.n. , collected at Knoxville, 
Knox County, Tennessee, in July, 1897, was determined by the col- 
lector as V. angustifolia x stricta and is indeed this hybrid. 
Schopf comments that of this taxon there were "scattered plants, 
over rolling pastured hillside". Thome (1955) cites two speci- 
mens from Johnson County, Iowa: "A presumable hybrid between this 
species [V. stricta] and V. simplex Lehm. was collected at Dwyer's 
Lake, Cedar Twp., R. P. Adams 1929, and in sandy prairie south of 
Iowa City, Shimek 1917 " . The R. Bebb U376, distributed as V. 
stricta x simplex , is typical V_. stricta Vent., while Danaree 
30963 is apparently a mixture of this hybrid and V. stricta . 

In all, 100 herbarium specimens, including the~types of almost, 
all the names involved, have been examined by me. 

Citations: OHIO: Erie Co.: L. H. Pammel s.n. [Sandusky, June & 



156 PHYTOLOGIA Vol. 10, no. 2 

July 1912] (lo—^hhOQ), ILLINOIS: Adams Co.: Evers 1)i)t)i (Ur). Pe- 
oria Co.: F. E. McDonald s.n. [Peoria, Jul. I9OJ1] (Ur) . Pope Co.: 
Schopf 6 (II — 16010). Saint Clair Co.: Eggert s.n. [111. opposite 
St. Louis, Aug. 12, 1875; Herb. Geete 5710] (Go, W--1323130) . 
Stephenson Co.: Eggert s.n. [stockyards; August 12, 1875] (I), £. 
n. [bluffs, 12 Aug. 1875] (I). Winnebago Co.: E. W. Fell 51307 
(Il--3890li), 51330 (11—38610), 51331 (I1--38611). INDIANA: Da- 
viess Co.: C_. C. Deam 25586 (In) . Harrison Co.: C. C^ Deam 20357 
(N— type, Pu— isotype, W— 769I68— isotype), 20357a (Dm). Marion 
Co.; £, £♦ Deam 6953 (On). Orange Co.: C^ C. Deam 26226 (Dm). 
Washington Co.: £• C_. Deam 20603 (Dm). IOWA: Black Hawk Co.: 
Burk 59U (Ur); L, H. Pammel s.n. [Cedar Falls, Sept. 28, 1920] 
(Io~98936); Pammel , Fisk , & Gilbert 272 (N) . Cerro Gordo Co.; 
A. Hayden 1, in part (N, N, N, N, N, N, N, N, N), 3 (N, N, N, N, 
N, N, N, N, N, N). Hardin Co.: L. H. Pammel s.n. [Steam Boat Rock, 
9-1912] (lo — 5197U). Jones Co.: L. H. Pamnel s.n. [Oxford Junc- 
tion, Jul. 25, 1919] (Io~97271, lo— 97273). Muscatine Co.: 
Shimek s.n. [Oct. lli, 1919] (N) . O'Brien Co.: Winge s.n. [Calu- 
met, Maj 1911] (Cp). Stoiy Co.: C. R. Ball s.n. [June 30, I898] 
(10—15309); L. H. Pammel s.n. [Ames, 9-8-9I1] (lo— 1532U). County 
undetemined: Semes 3301, in part [Blackstrap] (W~672170) . KEN- 
TUCKI: Warren Co.: S. F. Price s.n. [Bowling Green] (N) . TENN- 
ESSEE: Knox Co.: Ruth s.n. [Knoxville, July 1895] (Dt) . KANSAS: 
Greexnrood Co.: W. H. Horr s.n. [July 28, 1930] (Lw) . Leavenworth 
Co.; Jahns s.n. [June 27, 1929] (Lw). Miami Co.: Oyster s.n. 
[Aug. 1885; Herb. Prager I862U] (Gg— 3110-3) . Montgomery Co.: A. 
S. Hitchcock s.n. [Montgomery Co., Aug. I896] (Ka)j Rydberg & 
Imler li3U (Ka — 71^585, Lw, N). Shawnee Co.: A. S_. Hitchcock s.n. 
[Shawnee Co., July I896] (Ka); E. A. Popenoe s.n. [Topeka, July 
7, 1879] (Ka, W— 1U9628). Woodson Co.; E. W. Lathrop 369 (W— 
22350Uli), I188 (W— 2235073). MISSOURI: Barry Co.: Bush 15613 
(Ka— 8895077" Franklin Co.: Eggert s.n. [Pacific, h July I896] 
(Au, Cm, lo— 5Uli22, Mn— 6895, W— 75U958); L. H. Pammel s.n. 
[Washington, 6/25/88] (lo — 22913). Greene Co.: Blankinship s.n. 
[July 17, 1919] (Po— 638li8). Jackson Co.: C. A. Ripley s.n. 
[July 1898] (Ob— 507U9). Jefferson Co.: H. N. Andrews Jr. s.n. 
[Herculaneum, 5/30/36] (Ms), Phelps Co.: Kellogg s.n. [Jerome, 
June 20, 1912] (N), s.n. [Jerome, July 10, '12] (W— 1325386). 
Pike Co.: J. Davis 3379 (Ur) . Polk Co.: Steyermark 2U0J49 (Ky) . 
Saint Louis Co.: Letterman s.n. [Allenton, Jiine l881i] (lo — 
75363), s.n. [Allenton, June 28, I9II] (Au, lo— 76362, N, W— 
9861;30). Stone Co.: Bush 15697 (Ka— 88956), s.n. [Oct. 9, 1936] 
(Md). Saint Louis: Eggert s.n. [St. Loiiis, 12 Aug. 1875] (Al, 
N); Engelmann s.n. [St. Louis, Sept. I8UI] (W— 71991^), s.n. [St. 
Louis, July I81i2] (Pr, T), s.n. [St. Louis] (Br); M. Martens s.n. 



196U Moldenke, Monograph of Verbena 157 

[St. LoTiis] (Br, Br). ARKANSAS: Baxter Co.: Demaree 29320 (N) . 
Boone Co.: Demaree 3023 (We). Fulton Co.: Demaree 26322 (Au — 
1223lil, N) . Izard Co.: Demaree 227U5 (Bm) . Lawrence Co.; Rolfs 
s.n. [8/91] (Jo — Uli89). Netrton Co.: Danaree 222Ulia (N, Z) . Pope 
Co.: G_. Merrill 571 (Au — 1223UO) . Pulaski Co.: Hasse s.n. [Little 
Rock, May 30] (N) . Randolph Co.: Demaree 30963, in part (Au — 
122713). OKLAHOMA: Ottawa Co.: G. W. Stevens 2308 (Ok, Ok). CUL- 
TIVATED: Belgium: M. Martens s.n. [h. b. 1.] (Br). LOCALITY OF 
COLLECTION UNDESIGNATED: Herb. Bothe s.n. (B, B) . 

VERBENA MONACENSIS Moldenke, Phytologia 2: lU8~lii9. 19^6. 

Bibliography: Moldenke, Phytologia 2: lli8 — lU9. 19U6} Moldenke, 
Alph. List Cit. 2: Ul3. 19U8; Moldenke, Known Oeogr. Distrib, 
Verbenac, [ed. 2], 16U & I98. 19li9} Moldenlce in Chittenden, Roy. 
Hort. Soc. Diet. Bard. U: 2209 L 2211. 1951; E. J. Salisb., Ind. 
Kew. Suppl. 11: 263. 1953; Moldenke, R6sum« 22U & ii72. 1959; Mol- 
denke, R^sTflnl Suppl. 3: 29 (1962) and U: 10. I962. 

Herb; stems apparently prostrate, ascending toward the tips, 
branched, sharply tetragonal, brownish, lightly and irregularly 
pilose, less so in agej branches more sharply tetragonal (almost 
submargined) and more densely appressed-pubescentj nodes annulate; 
principal internodes 2 — 7 cm. long; leaves decussate-opposite, 
often bearing abbreviated and very leafy branches in their aixils; 
petioles to 1 cm. long, usually much shorter, winged and almost 
indistinguishable from the rachis of the lamina, strigose on both 
surfaces; leaf-blades uniformly green on both surfaces, chartace- 
ous, deeply trifid, the divisions again incised, the individual 
lobes mostly obtuse at the apex and elliptic or oblanceolate in 
outline rather than linear or oblong, strigose on both surfaces, 
the margins slightly revolute, the midrib and secondaries slender, 
obscure above, prominulous beneath; inflorescence solitary at the 
end of each stem and branch, at first condensed, later elongating 
to U cm. or more, densely many- flowered; peduncles slender, 1.5 — 
6.5 cm. long, densely strigose or appressed-pubescent, conspicu- 
ously tetrsLgonal like the branches; bractlets lanceolate, about 6 
mm. long, 1 mm. wide at the base, densely short-pubescent with 
subappressed whitish hairs, densely white-ciliate along the mar- 
gins, long-attenuate at the apexia calyx tubular, 8 — 9 mn. long 
(including the teeth), strigillose, 5-costate, its rim shortly 
5-toothed, the teeth triangular and usually less than 1 mm. long; 
corolla large, showy, its tube projecting about 5 nmi. beyond the 
calyx, glabrous outside, its limb about 1 cm. wide, the lobes 
shallowly bilobulate at the s^ex; anther-appendages not exserted. 

The type of this cvirious species in a specimen from the Uart- 
ius Herbarium deposited now in the herbarium of the Jardin Botan- 
ique de I'Etat at Brussels, said to have been collected from cul- 
tivated plants in the botanical garden at Munich, Germany, in 
1823 [not "18113" as erroneously stated by me previously] . The 
BlcHii specimen cited below, cultivated in Sweden, was apparently 
grown from seed obtained from Rouen, so presumably the species 
is/was cultivated also in France. I^ has in the past been misiden- 



1^8 PHYTOLOGIA Vol. 10, no. 2 

tified and distributed under the names V. erinoides Lam. and V. 
tenera Spreng. In all, 3 herbarium specimens, including the type, 
and 2 mounted photographs have been examined by me. 

Citations: CULTIVATED: Gennany: Herb. Martius s.n. [h. Monac. 
1823] (Br — type, N — isotype, N — photo of type, Z — photo of type). 
Sweden: Blom s.n. [from Rouen 1929] (Go). 

VERBENA MONTEVIDENSIS Spreng. in L., Syst. Veg., ed. 16, 2: 7li7. 
1825. 

Synonymy: Verbena isabellei Briq., Ann, Cohserv. & Jard. Bot. 
Genev. U: 23h. 1900. 

Bibliography: Spreng. in L., Syst. Veg., ed. l6, 2: 7U7. l825j 
Steud., Nom. Bot., ed. 2, 2: 750. iQhl} D. Dietr., Syn. PI. 3: 
600. I81i3; Walp., Repert. Bot. Syst. li: 20. lQh$i Schaai. in A. 
DC, Prodr. 11: 5U0. 18U7; Schau. in Mart., Fl. Bras. 9: 187. 
1851; Jacks, in Hook, f . & Jacks., Ind. Kew. 2: 1179. 1895} 
Kuntze, Rev. Gen, PI. 3 (2): 257. I898j Briq., Ann. Conserv. & 
Jarc. Bot, Genev, k: 23U. 1900; K. Schum. in Just, Bot. Jahresber. 
28 (1): li97. 1902; Thiselt.-Dyer, Ind. Kew. Suppl. 2; 191. 190U; 
Briq., Arkiv Bot, 2 (10): 12. 190ii; Briq., Aim. Conserv. & Jard. 
Bot, Genev, 10: 101 & 102. 1907; Moldenke, Suppl. List Invalid 
Names 9. 19la; Moldenke, Lilloa 6: 333 (19U) and 8: U32. 19^2; 
Moldenke, Alph. List Invalid Names U8. 19 U2; Moldenke, Known Ge- 
ogr. Distrib. Verbenac, [ed. 1], 39, Ul, & 101. 19U2; Rosengurtty 
Estud. Prad. Nat. Urug. 3$ 231; (19U3) and 5s 39U. 191^6; Schnack 
& Covas, Bol. Soc. Argent. Bot. 1: 28U. 19U6; Augusto, Fl. Rio 
Grande do Sul 211 & 233. 19U6; Troncoso & Burkart, Darwiniana 7: 
2lU & 215. 19U6; Moldenke, Alph. List Cit, 1: 171 & 26ii. 19U6; 
Schnack & Covas, Haumania 1: 5. [8], & 10, fig. 2h. 19li7; Molden- 
ke, Castanea 13: 117 & 119. 19U8; H, N, & A. L. Moldenke, PI. 
Life 2: 65. 19U8; Moldenke, Alph. List Cit. 2: U8I|, 577, 598, & 
612 (19l;8), 3: 665, 688, 693, 703, 705, 7li5, 8I4O, 8hS, 916, & 
923 (19li9), and h: 12U9, 1250, & 1257. 19U9; Moldenke, Known Ge- 
ogr. Distrib. Verbenac, [ed. 2], 9U, 99, 100, & I98. 19^9; Mol- 
denke, Phiytologia 3: 136 (191^9) and 3: 306, 1950; Stellfeld, 
Trib. Famac. 19 (10): I66. 1951} Moldenke, Inform. Mold. Set ii8 
Spec. [U] (195U) and U9 Spec. 3. 195U} Rambo, Sellowia 7: 260. 
1956; Angely Fl. Paran. 7: 13. 1957} Moldenke, Am, Midi. Nat. 
59: 33U. 1958; Reitz, Sellowia 11: 57 & 13li. 1959; Moldenke, r6- 
sum6 no, 118, 119, 127, 367, 370, & li72. 1959; Moldenke. R^sumS 
Suppl. 1: 23. 1959} Angely, Fl. Paran. 16: 78 & 79 (I960) and 
17: li6. I96I} Moldenke, RisumI Suppl. 3: 6 (1962) and U: 10. 
1962} Moldenke, Phytologia 8: 267, 3lU, 317, & 382 (1962) and 9: 
93. 1963} Moldenke, R6sum6 Suppl. 6: 7 (1963) and 7: 6. 1963} 
Moldenke, Phytologia 9: 367 & 380—382. I963. 

Illustrations: Schnack & Covas, Haumania 1: [8], fig. 2h. 
19U7. 

Tall perennial herb, subshrub, or suffrutescent shrub, 0.5 — 3 
m. tall, scoparious, very much branched; branches green, ascend- 
ing, acutely tetragonal, almost leafless, smooth and glabrous, 
slender; principal intemodes beneath the inflorescence to 17 cm. 



196U Moldenke, Monograph of Verbena 159 

long; leaves decussate-opposite} petioles short, 2 — 3 mm. longj 
lower leaf -blades rather small, lanceolate, green, membranous, 1 
to 3 cm. long and 7 mm. wide, acute at the apex, regularly convex 
and entire or subentire along the margins, cuneately attenuate 
into the petiole at the base, glabrous on both surfaces, the ven- 
ation simply pinnate, subprominulentj upper leaves much reduced, 
the blades lanceolate-linear, 0.5 — 1.5 can. long, the uppermost 
linear or setaceous, sessile; spikes short, 5 — 15 mm. long, 
slender-pedunculate, dense-flovrered, about U mm, wide; bractlets 
lanceolate-setaceous, about 1.5 mm. long, shorter than the calyx; 
calyx ovoid-campanulate, 2 — 2.5 mm. long, 5-costate, short-pubes- 
cent with antrorse hairs on the outside, especially on the ribs, 
its tube 1.5 — 2 mm, long, the rim shortly 5-dentate, the teeth o- 
vate at the base, short-mucronate at the apex, scarcely 0.5 mm. 
long; corolla varying from bluish, blue, or blue- violet to pale- 
violet, violet, pale-lilac [Saccardo U8X], dark-lilac, "hirhite- 
lilac", purple, clear-purple, rose, or red, [f^white to light- 
violet", according to Osten], about twice as long as the calyx or 
exceeding it by 2.5 mm., short-puberulent on the outside; stamens 
and pistil slightly exserted; cocci dark, oblong, about 1.5 mm. 
long; chromosome number: basic no. - 7, 2n - 21 (and h2) . 

The type of this species was collected by Friedidch Sellow at 
Montevideo, Uruguay, and was deposited in the herbarium of the 
Botanisches Museum at Berlin, where it was photographed by Mac- 
bride as his photograph no. 17U32, but is now destroyed. The 
type of V. isabellei was collected by Arsene Isabelle — in whose 
honor it was named — in Rio Grande do Sul, Brazil, and is depos- 
ited in the Delessert Herbarium at the Conservatoire et Jardin 
Botaniques at Geneva. 

The species was reduced to synonymy under V. stellarioides Cham. 
by Schauer (181|7, l85l) and by Jackson (l895)7"but is certainly 
not at all closely related to that species, as was pointed out by 
Kuntze on a label on the type sheet. In my l9Ul and 19U2 publica- 
tions V. minutiflora Briq. was regarded mistakenly as conspecific 
with V_. isabellei (which I now regard as a sjmonym of V. montevid- 
ensis), but I now regard this as distinct. 

Verbena montevidensis inhabits thickets, shrubby fields or 
eanpos, marshes and shrubby marshes, fields, open woods, pastures 
and dry sandy pastures, campos, meadows, parks, grassy plains, 
bogs, grassy places, dry stony meadows, cutover forest land, low 
wet places, and waste places in general. Collectors have found 
it along fencerows, roadsides, grassy roadsides, and rather wet 
roadsides, in woods, wet places, marshy ground, and sandy soil, 
at the edge of woods, along riversides and river banks, under 
trees at the edge of "monte" or in the interior of "monte", and 
in wet shrubby campos in a region of 2 meters rainfall and 0° — 25° 
temperature variation. Dus&i describes it as "ruderal" and Rambo 
reports it widespread in cultivated ground. Ewan found it in 
sandy soil of newly planted cane fields and in grassy pasture of 
bayou margin associated with Qnelista tora (L.) Britton & Rose, 
Ditremexa occidentalis (L.) Britton & Rose, and Vemonia, in full 



160 PHYTOLOGIA Vol. 10, no. 2 

STin, in Louisiana. 

It has been fovmd at altitudes of 200 to 16^0 meters, blooming 
in every month of the year, frxiiting fI^5m November to March, in 
June, and September. Legrand states that it flowers from Novanber 
to February in Uruguay. It is used medicinally in Brazil accord- 
ing to Reitz, and in Argentina according to Montes. Vernacular 
names reported for it are "anil", "camaradinha" , "formosa sem 
dote", "Jurupeba", and "quinanha", Osorio reports it as "rare in 
pradera" in Uruguay. 

Herbarium material has been misidentified and distributed under 
the names V. alata Cham., V. approximata Briq., V. bonariensis L., 
"V. cf . caracasana H.B.K.", xV. engelmannii Moldenke, V. interme- 
dia Gill, & Hook., V. litoralis H.B.K., V. litoralis Kunth, V. 
literal is L., V. littoralis H.B.K., V. littoralis Ktinth, V. litt- 
oralis var, brasiliensis Briq., and V. phlogiflora Cham. 

On the other hand, the JOnsson 9U2a, Rambo ^I6l6 & $U9Q$ , Reitg 
3260 & 3U03 , and Smith & Reitz 9ll;6 , distributed as V^ isabellei 
or V. montevidensis , are actually V» minutiflora Briq., while 
Smith & Klein 778? is V. reitzii Moldenke . Malme s.n. has extra 
large leaves, while MaLne 66I exhibits both large and small 
leaves on the same stem, 

Schnack & Covas (19U7) determined the basic chrcanosome number 
for this species as 7, 2n «= 21, fran a specimen cultivated at 
Capital, Mendoza, originally from Lavallol, Buenos Aires. They 
comment "En Verbena montevidensis hemos observado 21 crcanoscmas 
somdticas en seis individuos distintos y s6lo en uno 2n = U2. Es- 
to parece indicar que esta especie es apomictica, condici6n que 
estudiaremos oportunamente." Troncoso & Burkart (19li6) say "Ver- 
bena montevidensis Sprengel es vma especie hasta ahora dudosa, 

que Schauer coloca con interrogante entre los sin6nimos de 

V, stellarioides . Gracias a las notas de 0. Kiintze y a la 

fotografla del tipo (Montevideo, fotogr. 17U32 de la serie del 
Museo de Chicago) se llega a la seguridad que no tiene nada que 
ver con esa especie, Pso'ece que V, montevidensis es una buena 
especie, vecina de V, littoralis , pero distinta por se estatura 
menor, los -tallos menos escabrosos, las brdcteas pequeflas, etc, 
Es coratin en al Uruguay y las regiones vecinas de la Argentina," 
They describe the number of spikes as "infinitas" and say "V. 
intermedia y V, montevidensis tienen los tallos macizos." 

Rambo, in a letter to me dated August U;, 1955, states that 
in his opinion his no, 38055 is V. litoralis , not V. monteviden- 
sis , since it grew along a roadside (typical of that species) 
and not in a swamp (?rtiich he thinks is typical of V, monteviden- 
sis ) . I believe, however, that he momentarily confused V, monte- 
vidensis with V, alata when he made this statement, since the 
latter species is more typical of swamps. 

In all, 208 herbarium specimens and 7 mounted photographs, In- 
clmding phototypes of both the names involved, have been examined 



196U Moldenke, Monograph of Verbena l6l 

by me. 

Citations: LOUISIANA: Avoyelles Par.: Ewan I9O83 (Tl) . Evange- 
line Par.: Ewan 19367 (Tl) . BRAZIL: Minas Gerais: P. Clausen s. 
n. [Aug.— April I8U0] (Br, Br); Regnell 1. 326 (Ja~lll8U5). ?ara£&i 
Beetle 2025 (Ng~657U, W— 2Ui3908); Braga & Lange 8| (Gg, W— 
23693hh); Dusfn 2U9O (Ja—lli8U6, N, S, W— 1199U36), 7827 (Ca— 
5OI69O, N, N, S, S, W-II18I772), 10856 (N, S, W— li;8l77U), III69 
(S); Imaguire 18U7 (N, S); Mattos U710 (S), s^. [Herb. Mus. Par- 
an. 1|710] (N), s.n. [Herb. Mus. Paran. U751] (N); Nogiri k (Gg)j 
Reiss 56 (N); Stellfeld s.n. [h/9hh; Herb. Fac. Farm. Odont. 
1225] (N), s.n. [Herb. Mus. Paran. I63U] (N) . Rio Grande do Sul: 
Henz 353I46 (N); Isabelle s.n. [Macbride photos 2I169O] (Kr~photo, 
N— photo)} Llndman A.ti75 (N, N, S); Malme 166 (N, S), 661 (N, S)* 
3»n. [Cruz Alta, 20/1/1902] (S)j Moldenke & Moldenke I968I (Es, 
Lg, N, Sm); Rambo 937U (Rb), 38055 (N), U5339 (Au, Go), U6069 
(Au, W— 2026996), U9723 (Go), 51U;9 (S, W--2102035), 52083 ls) , 
5^Q7^ (B); Saldanha 6327 (Ja— 1^6579); Schwacke s.n. [ConceicSo 
do Uruguai, IV. I88O] (Ja— l|658l)j SchTrarzer s.n. [Colonia Sito 
Angelo, Oct. I9OO] (S)} Sehnem 3519 (B)j J. Vidal s.n. [Boca do 
Monte, March 1939] ( Ja--3U9U3) . Santa Catarina: Dus^n 17852 (S), 
17880 (S); Klein 2133 [Herb. Barb. Rodr. 1U209] (N, Sm, W— 
2321339); F. Mtlller s.n. [I876] (Ja— 31561), s^ru (Ja— U6535)} 
^^^^^ ^1358 (S), C^ (ja— 51358, N, N), H.939 (S); Reitz & Klein 
^-^0 [Herb. Barb. Rodr. 837U] (Cb, N, W— 2220170), 2772 (Sm), 
3758 (Sm), 3872 [Herb. Reitz lli211] (N, N, Sm), 5528 (W— 2252083), 
681;0 (Mm), 7l5U (Mm); Smith & Klein 750U (Ok, W— 2251285), II797 
(Ok, W— 2251815); Smith ^ Reitz 8988 (Ok, W— 225m78), 9730" (N~ 
Ok, w~225l595), 10175 (W— 22li9U67), IO303 (W— 22U9370)7^ith) 
Reitz, & Sufridini 9U63 (Ok, W—225l55il) , 962l£ (Ok, W— 2251575) . 
PARAGUAY: Grosse & Lindman 3651 (Ja— 28230); Hassler 6685 (Cb, 
N, N— photo, Z— photo), 706U (N); T. Rojas Uh8 [Herb. Osten 790U] 
(Ug); A. G. Schulz 7682 (Cb) . URUGUAI: J. Anderson 78 (Bm); An- 
derson & al. 78 [Herb. Reichenbach f . 13U25] (V); Arechavaletl 
39 (N, Ug— 1133, Ug), 3139 (Ug); H. H. Bartlett 20689 (Mi, W— 
2320Uil), 20751 (Mi), 21301 (Au~19h909, Mi, W-232028U); Berro 
5017 (N), QlkQ (N), 8232 (N)} Castellanos s.n. [Herb. InstTlBi- 
uel Ullo 150U6] (N), 8.n. [Herb. Inst. Miguel Lillo 15760] (N); 
Collector undesignated s.n. [Montevideo, Dec. 1885] (Ug), 3.n. 
(Ug, Ug); Qibert hh$ (Ug); Herter 269 [Herb. Herter 81713]~(n7 N, 
W— 13la8U9), s.n. [Sierra Acegui; Herb. Osten I81i99b] (Ug); Kun- 
tze s.n. [Montevideo, 7/XII/91] (N); Legrand 25U (Ug), 1992 "(Ug), 
2009 (Ug), 3809 (Ug), UOI9 (Ug), U188 (Ug); oigio 265 Jv^ 
13321), m (Ug— 13332), 662 (N, Ug-1323U, Ug), IOO9 (N, Ug, Ug- 
13212), s.n. [Valle Eden, Feb. 18, 19U7] (Ug— 13UH77r 



A NEGLECTED MEXICAN SPECIES OF ARUNDINARIA 

F. A. McClure 

Dept. of Botany, U.S. National Museum 

The studies on which this paper Is based were sup- 
ported by grant G-2314-2 from the National Science 
Foundation. 

Mumerous species of baraboo Indigenous to areas of 
the Western Hemisphere outside of North America have 
been placed at one time or another In the genus Arund- 
Inarla. Without apparent exception, these pertain to 
other genera. However, a misplaced Mexican species, 
represented by a single known collection, Llebmann no. 
132 , apparently does belong here. 

This collection consists of sterile leafy branch 
complements, each attached to a short section of the 
adjacent culm Internode. A specimen from this collec- 
tion, preserved at the U.S. National Herbarium, shows 
characters sufficiently critical In nature to warrant 
the allocation of this species In the genus Arundlnar - 
ia. 

ARUNDINARIA FLABELLATA (Fourn. ) McClure, comb. nov. 

G-uadua flabellata Fournler, Mexlcanas plantas 2: 
131. 1881. "Otate Mexlcanorura. . . In monte Orlzab- 
enal, Inter praerupta, 9OOO'." Type: Llebmann no. I32 
(HANX Fournler (I.e.) Inadvertently cited this specT^" 
men as "Llebmann I3I . " Llebmann I3I (L) Is actually 
the type of G-uadua aculeata Ruprecht, as verified by 
Hitchcock, who also examined Fournler' s type of G-uadua 
flabellata at Hannover. Hltchcockb notes (US) docu- 
ment the statement here published for the first time. 

Fournler' 8 brief diagnosis follows: "Ramls flab- 
ellatlm dlvergentlbus, follls llnearlbus longls angus- 
tls cum vaginls glabrls." The specimen examined by me 
(US 20^711 ; Llebmann I32 , ex C) shows additional fea- 
tures that are more dTsTinctly diagnostic than those 
recorded by Fournler. The following statements sup- 
plement the original description: 

fiulm sheatha not deciduous (eroded away by wea- 
thering — only a basal fragment remaining on the spe- 
cimen). Culm internodes cylindrical, not strongly 
sulcate but marked by a distinct transverse ridge at 
the level of (but external to) the locus of insertion 

162 



1961i McClure, Arundinaria 163 

of a branch complement; glabrous and lustrous; fistu- 
lous, the lumen filled (at least In part) with firm 
pith. Mid-culn branch complements pleloclade, the 
component axes unequal, the primary branch dominant. 
Leaf sheaths finely elevate-veined, minutely verrucose 
between the veins, otherwise glabrous dorsally, cill- 
ate on one margin near the apex; the inner llgule well 
exserted, fragile, apically concave, asymmetrical, in 
some sheaths densely clothed dorsally with fugaceoua 
antrorse hairs; the outer ligule linear, entire or e- 
rose on the margin; auricles and oral setae lacking. 
Leaf blades linear-lanceolate, attenuate-acuminate- 
caudate, gradually narrowed basally and decurrent on 
the slender, somevrtiat elongate, abaxially puberulent 
petiole; antrorsely scabrous near the outer edge, but 
elsewhere glabrous on the adaxlal surface, villous 
throughout on the abaxial surface; the midrib strongly 
salient and glabrous on the abaxial surface, and 
flanked on each aide by 4—3 secondary veins, these 
separated by 7"5 tertiary ones; the transverse vena- 
tion plainly visible on both surfaces of young leaves, 
less clearly so (especially on the adaxlal surface) in 
older ones. 

The classic station for the collection of this 
bamboo is given as "among precipices at 9OOO ft. on 
Mount Orizaba." This mountain is more popularly known 
today, and is shown on maps, as Citlaltepetl. The re- 
corded altitude of its highest peak is 18,696 feet 
(Hammond's Ambassador World Atlas, 195^ » P- 95)- Any- 
one who has the opportunity to botanize on its slopes 
will render a useful service to Agrostology if he can 
find and collect ample specimens of this inadequately 
known bamboo. — I shall be glad to supply direc- 
tions for the selection of material of critical inter- 
est. 

Flowering and fruiting material la essential to 
the completion of a description of the plant. Fuller 
representation of the vegetative structures is also 
needed, and sterile plants in good condition should 
not be by-passed. Most generally neglected are the 
rhizome (of importance for the confirmation of generic 
affinities) and the young culm shoot with persistent 
sheaths in good condition still attached. These lat- 
ter are useful for the recognition of specific enti- 
tiea. 



BOOK REVIEVfS 
Alma L. Moldenke 



"Families of Flowering Plants of Southern Africa", by Herbert P. 
Riley, 26? pp., Ikh colored illus.. University of Kentucky 
Press, Lexington, Kentucky. I963. $ll|..00 

This book covers quite accurately and very attractively a won- 
derfully and ihcreasingly interesting part of our world — Africa 
south of Tanganyika and the Republic of the Congo. The author's 
two decades of floral studies in this area, his access to the 
floristic works of other botanists, and his recent field experi- 
ence there give ample authority to his work. His camera loaded 
with color film, and those of Levyns and Jokl, give beauty allur- 
ing to the educated public the world over as well as to botanists. 
His writing style allows for easy reading of the pertinent data 
that the title proscribes. 

Plants endemic, indigenous, and introduced are described in 
their family groups. Chromosome nimbers, economic importance and 
special native usage are recorded. The maps, other introductory 
material, bibliography, and thorough index are very useful. The 
illustrations are the greatest asset of the book. 

Any work that covers so much in topic and area can seldom es- 
cape some errors. Considering just the Verbenaceae and its al- 
lied groups, the following items are noted: 

Avicennia officinalis does not occur in the region; A. mar - 
ina does, and the whole genus is better isolated in its own 
family, the Avicenniaceae . 

Lantana tri folia is probably not known frcan l^asaland be- 
cause it is aji American species j L. meamsii is the more cor- 
rect name for the African plant. 

Chascanum is not "equal to" Bouchea , but rather it is a 
segregate from that strictly American genus. 

These names are preferred: Lippia javanlca for L^ asperi- 
folia and Lantana galpiniana, Kalaharia uncinata for Clero- 
dendrrmi uncinatim , Vitex doniana for both V. cvmeata and V. 
cienkowskii , Duranta rep ens for D. plumieri , Lantana rugoaa 
for L. salvifolia , and Ptyla nodiflora var. reptans for Lip - 
pia reptans . 

Angola has 3, not 1, species of Premna; 2U, not 16, of 
Clerodendrum ; 3, not 1, of S tac hyt arphe ta . Duranta consists 
of 36, not 23, species, while Holmskioldia comprises U, not 
Just 3 or U, species. 

16U' 



196U Moldenke, Book revievrs 165 

"Selected Botanical Papers", edited by Irving W. Knobloch, 311 
pp., Prentice-Hall, Englewood Cliffs, New Jersey. I963. 
$3.95 paperback. 

What a wonderful service Dr. Knobloch achieved in making read- 
ily available these gems of botanicad literature I So many can 
profit from this work — thousands upon thousands of high school, 
college, and university students and teachers, more thousands of 
the general public, and thousands of botanists and other scien- 
tists who would be intrigued to see what is included or omitted 
and who would appreciate rereading these short selections because 
they know the writers and/or they are interested in the specific 
subject matter. 

How good it is to get to know sane of the great minds in this 
scientific discipline through important brief excerpts of their 
own writings] Included among the 59 are people of today (Leder- 
berg, Beadle, Sears), of yesterday (Humboldt, Ames, Camp), and of 
yester year (Ingen-Housz, Theophrastus, Aristotle whose author- 
ship of the included article is debatable) . There are people of 
many nations (Sinnott and Merrill of the United States, Erdtman 
of Sweden, Wallace of England), with the majority fron America. 
There are selections from many fields of botany (general impoi't- 
ance and development, ecology, phyt ©geography and exploration, 
systematic s and taxoncmy, anatomy and morphology, physiology, 
radiobiology, space biology, lower plants and pathology, horti- 
culture, forestry, conservation, paleobotany, cytology, genetics 
and plant breeding, evolution, the nature of scientific method) . 
The choices result in an emphasis on 20th century highlights. 

An impressive note in the preface reveals that, .according to 
a study by Dr. Barzan made in several of our midwestem colleges, 
75 percent of the students taking the first course in biology 
and 82 percent of those taking the first course in botany went 
no further in those disciplines. Perhaps making use of material 
like this book might lead to better education along these impor- 
tant lines. 



"The Plant Community", by Herbert C. Hanson and Ethan D. Church- 
ill, 218 pp., Reinhold Publishing Corp., New York 22. New 
York. 1961. $5.95 

The formation and nature of the plant community is the central 
theme of this ecological approach to the study of vegetation 
through discussion of the properties of the isolated species, of 
populations of uniform species (autecology), and of the ccmmunity 
of various organisms (synecology) . The scope of the book covers: 

1. the nature of the species in relation to the physical 
environment and in relation to other organisms^ 

2. the factors involved in the success of a species, 



166 PHYTOLOGIA Vol. 10, no. 2 

3. the formation of communities, their characteristics and 
their dynamics, 

U, the nature of the climax comzminity, 
5 . the classification of ccamminities . 

This is an excellent and simply "written text. It is also a 
fine book for general reading.. It is copiously illustrated in an 
effective manner with photographs provided mainly by the Soil 
Conservation and Forest Services of our government. There is a 
very good bibliography and a fairly good index. More cross- 
referencing would be helpful, as in the case of "kudzu" that does 
not appear under "K", but orJLy under "P" as Pueraria thunbergiana 
(kudzu). Larrea is listed for p. 79, but actually occurs on p. 
80. The authors did well to include scientific names as well as 
common ones, and well to avoid the fashion of using so many newly 
invented ecological terms that the basic concepts are muddled in 
all the verbiage. 



"Flora of Illinois", by George N. Jones, Ii02 pp.. University of 
Notre Dame Press, Notre Dame, Indiana, third edition. The 
American Midland Naturalist Monograph No. 7. 1963. $7.50 

This book review appears under my name, but I found it almost 
all written for me on one of my husband's memos after he checked 
the book for the Verbenaceae . "This book certainly should get a 
GOOD review. It is an example of how state floras should be 
Tfritten. I am delighted that synonyms are given under which the 
taxon may be found in other manuals. I am very happy to see 
synonyms given even for family names, but would have preferred 
seeing the standard " -aceae " tennination used for all family 
names. 

"Lippia is not a straight synorQTn for Phyla as implied on p. 
213. Since Phyla is a segregate from the still very large genus 
Lippia , it would have been better to write the synonym ' Lippia 
(Houst.) L., in part.' 

"The author claims that Verbena simplex hybridizes with V. 
bracteata . MacDougal made the same claim in 1907. I have ex- 
amined 36,000 specimens of this genus from 269 herbaria and have 
yet to see such a hybrid. Especially since neither author cites 
any substantiating specimens, I certainly would like to see one." 

I can add that there is considerable revision in this new 
edition, that the keys are relatively easy to follow, that there 
is a suitable conspectus, index and glossary, and that it would 
be wise to keep the second edition handy for its 1375 distribu- 
tional maps, fuller synonymy, and detailed bibliography. 



19^h Moldenke, Book reviews 167 

"Venomous and Poisonous Animals and Noxious Plants of the Pacific 
Region", edited by Hugh L. Keegan and W. V. Macfarlane, 1+56 
pp., Pergamon Press, New York 11, New Tork. I963. $21.00 

This is a collection of papers from the symposium of the Pub- 
lic Health and Medical Science Division of the Tenth Pacific Sci- 
ence Congress held in Hawaii in I962. The btilk of the papers 
deal with the deleterious effects of coelenterates, certain mol- 
lusks, leeches, scorpions, spiders, centipedes, those few milli- 
pedes with irritating secretions, those caterpillars with irri- 
tating hairs, certain fish and especially vencmous snakes. About 
liO pages are devoted to the poisonous and urticating plants of 
Australia and the poisonous plants of Guam. 

The book is well illustrated. The paper and print are very 
good. There is a useful index. Individual articles carry their 
own bibliographies. The price seems unusually high. 



"Chromoscsae Botany and the Origin of Cultivated Plants", by C. D. 
Darlington, 231 pp., revised second edition, Hafner Publish- 
ing Company, New York 3, New York. $5.50 

To the first edition of 1956 much new material has been added, 
the most valuable being the radiocarbon-^ studies facilitating 
chromosome counting in today's cultivated plants and those re- 
mains found with man's earliest evidences of civilization. Since 
the chromosomes have shown what the origins of cultivated plants 
were, it becomes possible to relate plant studies with the activ- 
ities of man in particular as well as with the processes of evo- 
lution in general. "Man's greatest biological esqjeriment has 
been the invention of agriculture, a process of understanding and 
controlling and improving certain flowering plants." So claims 
the author in his preface. 

The book deals with the nature and function of chromosomes, 
then with effects of chromosomes on plants in their systematic 
groupings, and on plants in their ecological and geographical re- 
lationships and on plants throughout time with its evolutionary 
changes. Almost half of the book is devoted to these problems 
as they effect cultivated plants and man. 

In an appendix Dr. E. B. Ford points out the bearing of the 
chromosome studies on the field of zoology. 

Forty diagrams and maps make the book excellently illustrated. 
There is a valuable bibliography and a good index. Both the pa- 
per and the binding are of poor queility, especially in view of 
the high price. 



"International Code of Nomenclature of Bacteria and Viruses", by 



168 PHYTOLOGIA Vol. 10, no. 2 

The Editorial Board of the International Committee on Bac- 
teriological Nomenclature, 186 pp., Iowa State University 
Press, Ames, Iowa, reprinting 1959. f!h*^0 

This is the ncanenclatural bible in this field and wiH remain 
so until possible changes of minor or drastic nature are made by 
the nomenclature committee of the next International Congress of 
Microbiology. Will Traub's new concept and terminology be follow- 
ed? The Rules and Recommendations seem to be clearly stated in 
heavy print and they are followed by explative annotations. An 
important feature is the compai*ison here between the botanical 
and the zoological codes with each other and with the microbio- 
logical one. 

This is an exceedingly valuable book and is well printed on 
good materials . 



"Form and Function in Plants", by John D. Dodd, 233 pp., Iowa 
State University Press, Ames, Iowa. 1962. $6.50 

This fine new introductory text on plant anatomy and physio- 
logy is the work of a seasoned and dedicated professor. Most 
basic information is introduced through laboratory stuc^ of 
freshwater algae and is then expanded to the other groups of 
plants. The main topics covered are: man in the world of 
plants, the green cell, reproduction, meiosis, inheritance, 
evolution, life on land, growth and differentiation, life with- 
out chlorophyll, and classification. An interesting and useful 
course could be taught frcm this text. 

The book is effectively illustrated with many of the author's 
excellent photographs and with more than three hundred very at- 
tractive drawings by Mrs. K. T. O'Siillivan. There is a good in- 
dex; the paper, printing, and binding are fine. A bibliography 
is missed. 



PHYTOLOGIA is financed entirely by its contributors, each one paying 
in advance for the entire cost of printing, binding, and distributing his con- 
tribution. All money received from siil)scribers, after the expenses of col- 
lections have l)een deducted, will be distributed among the contributors upon 
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Each contributor is therefore a shareholder in the magazine, assuming his 
part of the expenses and sharing in the profits, if any accrue. 

Each number consists of not less than 32 pages. .Ml manuscript accepted 
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PHYTOLOGIA 

Designed to expedite botanical publication 






\'ol. 10 June, 1964 No. 3 



CONTENTS 

REED, C. F., Stewartia malachodendron L. on Delmarva Virginia. .. 169 

MOLDENKE, H. N., Notes on new and noteworthy plants. XLI 170 

MOLDENKE, H. N,, Materials toward a monograph of the genus 

Verbena. XIX 173 

EITEN, L. T., A correction 237 

MOLDENKE, A. L., Book reviews 237 



Published by Harold N. Moldenke and Alma L. Moldenke 

15 Glenbrook Avenue 
Yonkers 5, New York, U.S.A. 

Price of this number, SI; per volume, S5.75, in advance 
Volume 10, No. 1, was issued March 19, 1964; Vol. 10, No. 2, was issued 

April 23, 1964 



STEWAF.TIA I1ALACH0DENEP.0N L. ON DELMARVA VIRGINIA 
Clyde F. Reed 



The Virginian stewartia, Stewartia malachodendron L, (Stewartia 
viTKinica Cav, ) , lives in moist woods on the coastal plain from 
eastern Virginia (Accomac Co\inty on the Delmarva Peninsula) south 
to western Florida and then westward to western Louisiana; thence 
northward to Tennessee and Arkansas. 

Tatnall (19U6, in Flora of Delaware and the Eastern Shore, p. 
175) states that there is "one station, head of Messango Creek, 
Accomac County, Virginia, Ellis Mears in 1886 (at GH and Acad, 
Nat. Sci. Phila.); not found in recent years, althou^ carefully 
searched for". 

Last year, while botanizing in Accomac County, Virginia, the 
author came upon a Mr. L. Floyd Nock, Jr. of Accomac who had found 
Stewartia malachodendron L, in a woods two miles south of Wacha- 
preague, Accomac County, about 19li5 or 19li6, At that time he trans- 
planted small plants of this species to slopes (above saline line) 
along Folly Creek at his home (The Folly), 2 miles east of Accomac. 
The plants have flourished to be shrubs 10-1^ feet tall. Except 
when the hurricanes forced salt water up on the land and killed 
those plants nearest the waterline, the rest of the plants have 
grown well. Some seedlings have come up and are growing fine. The 
plants in this area bloom about two weeks after dogwood has drop- 
ped its blossoms. 

Wachapreague is in the southeast corner of Accomac County, 
whereas Messango Creek is in the northwestern corner, some 25 to 
30 miles away. I have searched in the Messango Creek area and 
have not been able to find the plant, after several hunts over the 
years. The Gulf Stream Nursery, near Wachapreague, has been pro- 
pagating this species from seed obtained from Mr, Nock, 

Representative specimens from eastern Virginia: Accomac County : 
woods above saline line, at The Folly, on Folly Creek, 2 mi, east 
of Accomac. Sept, 11, 1963. Reed 65^h79-82 (Dupl. in US), Princess 
Anne County ; near Pungo, July"??, 1939, Fernald & Long 10726 (US), 
Norfolk County ; Yadkin, Great Dismal Swanp, June 11, 19liO, Fer- 
nald & Long 12131 (US). 



169 



NOTES ON NEW AND NOTEVfORTHT PLANTS . XLI 
Harold N. Moldenke 



ALOISIA SLEUMERI Moldenke, sp. nov. 

Frutex 2 m. altus; ramulis gracilibus stramineisj foliis ter- 
niis subsessilibus lanceolatis Ii — 7 cm. longis 1 — 1.5 era. latis 
supra minute piloaulis (pilis ad basin minute bulbosis), subtus 
minute resinoso-punctulatis; inflorescentiis axillaribus folia 
brevioribus laxe paucif loris . 

Shrub, 2 m. tallj branchlets and twigs slender, stramineous, 
glabrous; leaves temate, subsessile or very shortly petiolate; 
petioles, vdien present, filiform, 1 — 3 mm. long, very minutely 
appressed-strigillosej leaf -blades thinly chartaceous, dark-green 
above, lighter beneath, narrow-lanceolate, J4 — 7 cm. long, 1 — 1.5 
cm. -wide, acute at the apex, reg\ilarly sharp-serrate with appres- 
sed antrorse teeth except toward the base and apex, attenuate- 
acute at the base, minutely pilosulous and slightly scabrous a- 
bove with short broadly bulbous-based hairs, the hairs eventually 
wearing off and the bases collapsing to form numerous disk-like 
markings, minutely resinous-punctulate beneath, the venation ob- 
scure or slightly subimpressed above, rather conspicuous but not 
prominent beneath, the filiform secondaries short, straight, sub- 
parallel, each ending in the apex of a tooth; inflorescence ax- 
illary, abundant, temate, shorter than the subtending leaves, a- 
bout 3 cm. long in all; peduncles filiform, 1 — 1.2 cm. long, an- 
trorsely puberulent; spikes short, stout, rather few-flowered, a- 
bout 1.5 cm. long, 1 — 1.5 cm. wide, the flowers rather distant, 
wide-spreading J bractlets minute, triangular, densely puberulent; 
calyx cylindric, about 3 mm. long, densely puberulent; corolla 
hypocrateriform, white, its tube about 6 mm. long, densely puber- 
ulent on the outside, the limb about 2 mm. wide. 

The type of this distinctive species was collected by Dr. H. 
Sleumer — in -vrtiose honor it is named — and Federico B. Vervoorst 
( no. 2370 ) at Pozo de Piedra and vicinity, at I9OO meters alti- 
tude, department of Bel^n, Catanarca, Argentina, between January 
25 and 31, 1952, and is deposited in the United States National 
Herbarium at Washington. The species is obviously related to A. 
triphylla (L'H^r.) Britton, from which its toothed leaves and 
short congested spikes at once distinguish it. 

LIPPIA BELLATULA Moldenke, nom. nov. 

Lippia bicolor Mart, &: Schau. ex Schau, in A. DC., Prodr. 11: 
587. I8U7 [not L. bicolor Kxinth & BouchI, Ind. Sem. Hort. Berol, 
18U5: 10. 18U5]. 

LIPPIA DUARTEI Moldenke, sp. nov. 

Planta perennis ubique cinereo-pubescens; ramis gracilibus 
obtuse subtetragonis dense cinereo-pubescentibus, pilis brevibus 

170 



1961i Moldenke, New and notefworthy plants 171 

adpressis; intemodiis valde elongatis; foliis decuasatis; petio- 
lis maturis elongatis; laninis foliorum oblongis usque ad 7 cm. 
longis 2 cm. latis, supra rugosis breviter pubescentibus, subtus 
densiore pubescentibus; inflorescentiis axillaribus solitariis 
vel binis capitatis pedunculatis cinereo-pubescentibus . 

Apparently a perennial plsint with rather strict stems or 
branches, densely cinereous-pubescent throughout; stems or bran- 
ches slender, obtusely sub tetragonal, densely cinereous pubescent 
with short appressed antrorse hairs; nodes not annulate; princi- 
pal internodes conspicuously elongated, 3*5 — 9 cm. long; leaves 
decussate-opposite, the lower ones often with abbreviated leafy 
non-floriferous twigs in their axils; petioles slender, conspicu- 
ously elongated on upper mature leaves and there about 1,0 cm, 
long, very densely cinereous-pubescent, canaliculate above, the 
hairs straight and antrorse; leaf -blades chartaceous, dark-green 
above, lighter beneath, oblong, to 7 cm. long ?rtien mature and 2 
cm, wide, apparently only a few mature ones present during anthe- 
sis, attenuate-acute at the apex, rounded at the base, uniformly 
crenate along the margins, deeply rugose and scabridous-pubescent 
above, more densely cinereous-pubescent (especially on the larger 
venation) beneath; midrib and the numerous secondaries slender, 
deeply impressed above, prominent beneath; veinlet reticulation 
ab\mdant, deeply impressed above on mature leaves; inflorescence 
axillary, much shorter than the subtanding mature leaves, soli- 
tary or binary in each of the upper axils, 2 — 3 cm, long, capita- 
te; peduncles very slender, 1.5 — 2.5 cm, long, very densely white- 
pubescent vrith straight antrorse subappressed hairs; heads hemi- 
spheric, many- flowered, 5 — 10 mm. wide; bractlets ovate, the out- 
er ones about 2 mm. long and 1 ram, wide, acute at the apex, ap- 
pressed-puberulent . 

The t^-pe of this curious species was collected by Apparicio 
Pereira Duarte ( no, 7851) — in whose honor it is named — at 
Varzea da Palma, Faz . MSe d'Agua, IJinas Gerais, Brazil, on April 
28, 1963, and is deposited in the Brit ton Herbarium at the New 
York Botanical Garden, 

LIPPL\ FELIPPEI lloldenke, sp. nov. 

Fruticulus parvus; ramis gracilibus strictis dense glanduloso- 
hirtellis striatis; foliis decussatis; petiolis filiformibus 
brevibus; laminis foliorum cordato-ovatis par vis plerumque re- 
f lexis ad apicem acutis, marginibus regulariter grosso-serratis, 
ad basin cordatis vel subcordatis, utrinque glanduloso-pilosis; 
inflorescentiis axillaribus paucis folia superantibus ; capitulis 
globosis 2 cm. latis longisque; bracteolis dense imbricatis herb- 
aceis ovatis purpureis. 

Small shrublet with a woody xylopodium; branches few, erect, 
simple or subsimple, about 60 cm. long, slender, stramineous, 
striate, rather densely hirtellous with soft gland-tipped hairs, 
less so toward the base; leaves decussate-opposite, petiolate, 
mostly drooping-reflexed; petioles slender, 3 — 7 mm. long, dense- 
ly glandulose-hirtellous; leaf-blades thin-char taceous, bright- 
green above, lighter beneath, cordate-ovate, 1.7 — 3.2 cm. long, 



172 PHYTOLOGIA Vol. 10, no. 3 

l.U — 2.7 cm. wide, acute at the apex, regularly and coarsely ser- 
rate from the apex almost to the base with triangular somewhat 
antrorse but divergent teeth, cordate or subcordate at the base, 
glandulose-pilose with erect gland-tipped hairs on both surfaces, 
soft to touch, the larger venation slightly impressed above and 
prominulous beneath; inflorescence axillary in the uppermost 
leaf-axils, solitary, surpassing the subtending leaves, ascending; 
peduncles very slender, 1 — 2 cm. long, densely gl.^ndulose-hirtel- 
lous with soft spreading whitish hairs; heads globose, about 2 cm. 
long and wide, apparently rosy-purple, showy; bracts very nimier- 
ous, herbaceous, densely'' imbricate, ovate, to 12 mm. long and 6 
mm. Trvlde, venose, acute or short-acuminate at the apex, slightly 
surpassing the corolla-tube, glandulose-pubescent; corolla hypo~ 
crateriform, rose, with a yellow central "eye", the tube about 10 
1X1. long, glandulose-pubescent on the outside, the linb about 9 
ram . ivide . 

The type of this handsome species was collected by Gil M. 
Felippe (no. lli.O) — in whose honor it is named — in a cerrado 
along the road to the airport, municipality of Prata, Minas Ger- 
ais, Brazil, on IJarch 20, 1963, and is deposited in the United 
States National Herbarium at Washington. 

LIPPIA TURBINAIA f. MAGNIFOLIA Moldenke, f . nov. 

Haec forma a forma typica speciei laminis foliorum late ellip- 
ticis vel lanceolato-ovatis vel obovatis usque ad 7 cm. longis et 
3.3 cm. latis recedit. 

This form differs from the typical form of the species in hav- 
ing its leaf -blades regularly larger and wider when mature, more 
broadly elliptic to lanceolate-ovate or oblanceolate to obovate 
and to 7 cm. long and 3.3 cm. wide. 

The type of this form was collected by Santiago Venturi ( no. 
8120 ) at Carapo Quijaro, Rosario de Lerma, Salta, at an altitude 
of 1300 meters, Argentina, in January, 1929, and is deposited in 
the United States National Herbarium at Washington. The plant is 
described as a shmib 1.5 m. tall, -vrith white flowers. 

VERBENA SIMPLEX f . ALBIFLORA Moldenke, f . nov. 

Haec forma a forma typica speciei corollia albis recedit. 

This form differs from the typical form of the species in hav- 
ing white corollas . 

The type of the form was collected by Ronald Leighton McGregor 
(no. 13217) scattered in low places in shallow soil over limestone 
in a native bluestem prairie hay meadow tT/o miles south of Els- 
more, Allen County, Kansas, on J\ily 22, 1957, and is deposited in 
the herbarium of the University of Kansas at Lawrence. 



MATERIALS TOWARD A MONOGRAPH OF THE GENUS VERBENA . HX 
Harold N. Moldenke 



VERBSIA MONTE VIDENS IS Spreng. 

Pedersen found this species in grasslands on malezal (low fre- 
quently flooded ground) . 

Additional citations: URUGUAY: Osten 3165 (Ug), 3335 (Ug), 
368U (N, Ug, Ug)j Rosa-Mato 313 (Ug— 9808), 399 (Ug— 98IO), UOO 
(Ug— 98II); Seijo s.n, [Montevideo, 16 Nov. de I88U] (Ug); Sellow 
s.n. [Montevideo; Macbride photos I7I4.32] (Kr — photo of type, N — 
photo of type, N — photo of type); Teisseire l|l;8l (Ug) . ARGENTINA: 
Buenos Aires: A^ Alvarez i;53 (Bm, S); R. Alvarez i;27 (S); Cabrera 
861 (N), 1710 (N), 2108 (N, 7/~l57l|ii5U), 2336 (N, N, N); Cannelle 
s.n. [Rio Parana Mini] (Mv); H_, M. Gurran s.n. [Oct. 19, 1913] 
(N); Krapovickas 2678 (Gg— 353253, N), 27li2 (N); Nicora 377 (W— 
2196U6ii); Pastore 137 (N); Remiro s.n. [Mercedes, 20.IV.19U3] 
(Mv); Scala 10001 (N); A. G. Schulz 5690 (Z); Sparre 268 (S) . 
Gorrientes: Pedersen 77ii (W — 2122514;), 50 7U (S); Ruiz Huidobro 
2087 (S). Bntre Rfos: Cabrera 10756 (W~2370638) . Misiones: Ber- 
toni 3196 (N); Ekman 2018 (N, S), 2030 (N, S) ; Grltner 108 [Herb. 
Osten 23I8O] (Ug); Martinez Crovetto 3U1;3 (N); Medina 123 (Gg— 
353260, N)j T. Meyer 11297 (N); Montes 91 (Ca, Ca), 1237 (Es, Gg— 
35267U, N), 9553 (Vi, Vi); G. J_. Scfararz 62U (Ca), 821 (Ca), 1127 
(Ca), 1128 (S), ll£30 (N), 1572 (N), 1733 (N), 1976 (N), 3779 (N) . 
Santiago Island: Cabrera 2159 (N, W— 157UU95) . 

VERBENA MORICOLOR Moldenke, Phytologia 2: U2ii— U25. I9U8 . 

Bibliography: Moldeni:e, Phytologia 2: U2U — U25 & U32. 19U3; 
Moldenke, Castanea 13: 119 . 19ii8; Moldenke, Knovm Geogr. Distrib, 
Verbenac, [ed. 2], IO6 &: I98. I9l;9; Moldenke, Alph. List Cit. U: 
1090. 19l9; Moldenke, Phytologia 3: 290. 1950; Hcnyell, Vfasmann 
Journ. Biol. 10: 377. 1952; E. J. Salisb., Ind. Kew. Suppl. 11: 
263. 1953; Molden}:e, R^suml 127 & U72. 1959- 

Herb, to 1 ra. tall; ster.s and branches slender, more or less 
tetragonal, sparsely and irregularly pilose vrLth '.vhitish hairs 
of various lengths, the youngest parts rather densely liirsutulous 
with sharp-pointed hairs about 1 mn. long borne at right angles 
to the branch; nodes annulate; principal intemodes elongated, 
7.5 — 13 cm. long; leaves decussate-opposite; petioles slender, 
1.5 — 2 cm. long, irregularly pilose -.vith very short hairs and 
sparsely hirsutulous with long sliarp-pointed hairs like on the 
younger branches; leal-blades thin-chartaceous, somewhat lighter 
beneath, lanceolate, 1; — 3 cm. long, 1 — 2.3 en. v/ide, attenuate- 
acute at the apex, obtuse or abruptly acute at the base, irregu- 
larly dentate from tlie base to the apex vrith ratlier broad suba- 
cute teeth, the larger irregularly interspersed with smaller ones 

173 



17U PHYTOLOGIA Vol. 10, no. 3 

toward the base of the blade and often doubly dentate, rather 
densely strigose-pubescent above, densely pubescent beneath with 
hairs of various lengths; midrib slender, plane or slightly subim- 
pressed above, slightly proainulous beneath; secondaries about 7 
pairs, very slender, ascending, hardly arcuate, indiscernible a- 
bove, rather obscure beneath; veinlet reticulation indiscernible; 
inflorescence terminal, spicate, the flowering portion apparently 
elongating to about 7 cm.; peduncles similar to the upper branches 
or stems in texture, shape, and pubescence, 1 — 7.5 cm. long; ra- 
chis densely pilose-pubescent with hairs of various lengths; 
bractlets lanceolate, about h mm. long, attenuate to the apex, 
sparsely puberulent on the back, the margins regiilarly ciliate; 
calyx cylindric, 6 — 7 mm. long, about 1 mm. in diameter, rather 
sparsely puberulent-pilosulous, the teeth 0.5 and 0.9 mm. long, 
subulate; corolla hypocrateriform, varying from blue-violet or 
violet to purple, rose, or mulberry-colored, its narrow-cylindric 
tube 13 — lU mm. long, densely pilose-puberulent above the calyx, 
its limb 6 — 7 nim. wide, much darker in the throat, glabrous with- 
in, subglabrate on the outer surface. 

The type of this handsome species was collected by Santiago 
Ventu.'i (no. 5397) among spiny plants on a hillside at Sierra de 
Calilegua, at an altitude of 800 meters, in the department of 
Ladesma, Jujuy, Argentina, on October 11, 1927, and is deposited 
in the herbarium of the California Academy of Sciences at San 
Francisco. The species is obviously related to V. phlogiflora 
Cham, and V. incisa Hook., but differs conspicuously in its much 
smaller calyx and corolla. One collector describes the plant as 
a "shrub", but probably in error. It has bean found at altitudes 
of 360 to 1600 meters, flowering in January, February, September, 
and October. Herbarium material has been nisidentified and dis- 
tributed as V. megapotamica Spreng. and V. scrobiculata Griseb. 
In all, 15 herbarium specimens, including the type, and k mounted 
photographs have been examined by me. 

Citations: ARGENTINA: Catamarca: \'{all & Sparre s.n. [La Estan- 
cia, 27/II/U6] (Ew, N), s.n. [Alsaverade, 21/ll/k6] (Ew, Ew, N). 
Jujuy: Venturi $397 [Herb. Osten 20800] (Bm — isotype, F — photo of 
type, Gg — 157893 — type, N — isotype, N — photo of type, S — isotype, 
Si~photo of type, Ug~isotype, W— llUi0837~isotype, W— 1591U72— 
isotype, Z — photo of type). Salta: Lillo I8III [Herb. Osten 
3U69] (Ug); Parodi 9179 [Herb. Osten 2253U] (Ug); Schreiter 5037 
[Herb. Osten 2077U] (Ug). 

VERBENA MULTICAULIS Raf ., Herb. Raf . 65, ncan. nud. 1833. 

Bibliography: Raf., Herb. Raf. 65. 1833; E. D. Merr., Ind. 
Raf. 205 & 295. 19U9J Moldenke, R^sumS Suppl. 7: 3. 1963. 

Nothing is known about this plant except that, according to 
Merrill, the name was proposed by Rafinesque and published by 
him in 1833, without description, based on a type from the "Appa- 
lachian Mountains". It is very probably a synonym of V. simplex 
var. eggerti Moldenke, but of this I cannot be sure wTChout see- 
ing the type material. 



196U Moldenke, Monograph of Verbena 175 

VERBENA NANA Moldenke, phytologia 3: 119 . 19h9 - 

Bibliography: Moldenke, Known Geogr. Distrib, Verbenac, [ed. 
2], 105 & 198. 19U9; Moldenke, Phytologia 3: 119 & 136 (19U9) and 
3: 289 & 290. 1950j Stellfeld, Trib. Famac. 19 (10): 16?. 19^1} 
E. J. Salisb., Ind. Kew. Suppl. 11: 263. 1953 J Angely, Fl. Paran. 
7: 13. 1957; Moldenke, R^sumS 110 118, 127, & U72. 1959; Angely, 
Fl. Paran. l6: 79 (I960) and 17: U6. 1961; Moldenke, Phytologia 
9: 388. 1961i. 

Dwarf annual herb, 1 dm. or less in height, or sometimes suf- 
frutescent and to 6 cm. tall; stems seni-prostrate or erect, usu- 
ally with 2 ascending or erect branches at the base, obtusely 
tetragonal, deeply l;-sulcate in drying, densely hirsutulous with 
whitish mostly glanduliferous hairs of various lengths standing 
at right angles to the stem; principal intemodes 0.5 — 2 cm. long 
leaves decussate-opposite; petioles slender, 3 — 11 nim. long, 
densely hirsutulous like the stem and branches; leaf -blades 
chartaceous, rather uniformly green on both surfaces, ovate, 1.3 — 
2.5 cm. long, 8 — 12 mm. wide, obtuse or subacute at the apex, a- 
cute at the base, appressed-serrulate with rather wide, flat, 
rounded teeth from the ividest part to the apex, rather densely 
pubescent on both surfaces vd.th v;hitish, stiff, straight, more or 
less glanduliferous hairs; midrib slender, plane above, subpro- 
minulous beneath; secondaries slender, 2 — 5 per side, ascending, 
subprominulous beneath; veinlet reticulation rather obscure on 
both surfaces; inflorescence terminal, dense, rather few-flower- 
ed, sessile or subsessile; bractlets lanceolate, U — 5 nun. long, 
glandular-pubescent; caljoc tubular, about 11 mm. long, densely 
hirsutulous with short, whitish, often glanduliferous hairs 
standing at right angles to the surface, the rim irregularly 5- 
apiculate, the teeth about 1 mm. long; corolla hypocrateriform, 
varying from intensely red or rose to pink, rose-lilac, or lilac- 
tinged, its tube about 15 mm. long, glabrous on the outer sur- 
face, the limb 5 — 6 mm. wide, glabrous. 

The type of this very distinct species was collected by Ish- 
mael Vorel ( no. 117 ) at Piran6, in the department of P^ran^, For- 
mosa, Argentina, on October 23, 19U5, and is deposited in the 
Britton Herbarium at the New York Botanical Garden. The species 
has been found on carapos, in low wet soil, in Copemicia palm 
woods, and in Vachellia farinosa thorn-thickets, in flower in 
February, June, and October to December, and in fruit in October. 
It is said to be common in open ground in Paraguay, and rather 
abundant in the Argentine Chaco. A common name is "verbena de 
los palmares". 

Herbarium material has been misidentified sind distributed un- 
der the names "V. chamaedryfolia var. vel hybr." [by Osten] , "V. 
aff . kuntzeana I.;oldenke" [by Troncoso], V. phlogiflora Cham., 
and V. scrobiculata Griseb. In all, 29 herbarium specimens, in- 
cluding the t^'pe, and It mounted photographs have been examined 
by me. 

Citations: BRAZIL: Paran5: Stellfeld s.n. [Herb. Mus. Paran. 
311;0] (N). SSo Paulo: Brade 7002 [Herb. Inst. Bot. S. Pailo 672U] 



176 PHYTOLOGIA Vol. 10, no. 3 

(N); EdTOll s.n. [Herb, Comm. Geogr. & Geol. 2958] (N, Sp— 1^729, 
Sp_1^732) . PARAGUAY: Fiebrig 1218 (S), U371 (Bm, Cb, Cb)j Ped- 
ersen U069 (N, S, Ut— 90219b, W— 2283663); T. Rojas 2|U (Bm, S) , 
2g6 (S), 1895 [Herb. Hort. Parag. 10055; Kerb. Osten 13558] (N, 
N, Ug). ARGENTINA: Chaco: T. Meyer 68O [Herb. Osten 229Uii] (N, 
N, Ug), 29U3 (N); A. G. Schulz 286 [Herb. Osten 2311i7] (N, Ug), 
IU66 (N) . Formosa: I. Morel 117 (B— isotype, F— photo of type, 
fj — type. N — photo of type, Sg — photo of type, Z — photo of type), 
U96 (Bm). 

VERBENA NEOlIEnCANA (A. Gray) Small, Fl. Southeast. U. S., ed. 1, 
1010 & 1337 [as " neo-nexicana "] . 1903. 
Synonymy: Verbena officinalis var. hirsuta Torr. in Emory, 

Rep. U. S. fie Mex. Bound. Surv. 2: 123. 1859. Verbena c anescens 
var. neo-mexicana A. Gray, Syn. Fl. N. Am. 2 (1): 337. I878. Ver- 
bena neo-mexicana Briq. in Engl. & Prantl, Nat. Pflanzenfam. h 
( 3a) : liiS, hyponym. I89U. Verbena canescens x officinalis Briq. 
in Engl. & Prantl, Nat. Pflanzenfam. h (3a): 11^8, in syn. I89i;. 
Verbena neomexicana Small ex Prain, Ind. Kew. Suppl. 3: 187. 
1908 . Verbena canescens neomexicana A. Gray ex Standi., Contrib. 
U. S. Nat. Herb. 13: 171 & 211. 1910. Verbena officinalis hirsuta 
Torr. ex Standi., Contrib. U. S. Nat. Kerb. 13: 171 & 211, in syn. 
1910. Verbena canescens var. neomexicana A. Gray apud Perry, Ann. 
Mo. Bot. Gard. 20: 296. 1933. Verbena neomecicana (Gray) Small 
ex Moldenke, Suppl. List Invalid Names 9, in syn. I9UI. Verbena 
neo-mexicana Small ex Moldenke, Suppl. List Invalid Names 9, in 
syn. I9UI. Verbena neomexocana (A. Gray) Small ex Moldenke, 
Alph. List Git. 1: 200, sphalm. I9U6 . Berbena neomexicana 
(Gray) Small ex Moldenke, Resum6 237, in syn. 1959- Verbena 
canescens neo-mexicana Gray ex Moldenke, Rlsum6 36O, in syn. 
1959. Verbena neo-mexicana Gray ex Moldenlce, R6sum6 370, in syn. 

1959. 

Bibliography: Torr. in Emory, Rep. U. S. & Mex. Bound. Surv. 
2: 128. 1859; A. Gray, Syn. Fl. N. m. 2 (1): 337. 1878; Briq. in 
Engl. & Prantl, Nat. Pflanzenfam. U (3a): II18. I39ii; J. K. Small, 
Fl. Southeast. U. S., ed. 1, 1010 & 1337. 1903; Prain, Ind. Kew. 
Suppl. 3: 187. 1908; Standi., Contrib. U. S. Nat. Herb. 13: 171 & 
211. I9IO; J. K. Small, Fl. Southeast. U. S., ed. 2, 1010. 1913; 
Perry, Ann. Mo. Bot. Gard. 20: 2U6, 2U8, 261, 296—298, 301, 302, 
3UO, & 355. 1933; Dermen, Cytologia 7: I6I & 162. 1936; Cory, Texas 
Agr. Exp. Sta. Bull. 550: 38. 1937; Moldenice, Prelim. Alph. List 
Invalid Names U5 & U7. 19U0; Moldenke, Suppl. List Invalid Names 
8 & 9. 19ia; Schnack, Anal. Inst. Fitot6c. Sta. Catalina U: I8. 
19U2; Moldenke in Lundell, Fl. Texas 3 (1): 17 & 29—30. 19U2; 
Moldenke, Alph. List Invalid Names ii6 & U9. 19U2; Moldenke, Knovm 
Geogr. Distrib. Verbenac, [ed. 1], 13, lU, 19, 5c 101. 19U2; 
Schnack & Covas, Darwiniana 6: U70. I9UI1; Moldenke, Phytologia 2: 
128. 19U5; Darlington & Janaki Ammal, Chromosome Atlas 270. 19U5; 



196U Moldenke, Monograph of Verbena 177 

Moldenke, Alph. List Cit. 1: 3U, 116, 182, 199, & 200. 19U6; W. 
C. Leavenworth, Am. Midi. Nat. 36: 187. 19U6j L- C. Hinckley, Am. 
1/ridl. Nat. 37: 170. 19U7j Moldenke, Phytologia 2: 331 (19U7> and 
2: 163~16U. 19 U8; Moldenke, Wrightia 1: 225—227. 19liB; Moldenke, 
Alph. Ust Cit. 2: U71, 538, & 539 (19U3), 3: 708, 723, 729, 752, 
789, 939, 953, & 951 (19U9), and U: 990, llli2, 1150, 122U, 1230, 
& 1290. 19ii9; Moldenke, Knoiivn Geogr. Distrib. Verbenac, [ed. 2], 
2I1--26, 33, & 198. 19U9j McDougall & Sperry, Pl. Big Bend lii6. 
I95I; Moldenke, Phytologia 3: U67 £t ii68 (195l) and 5: 133. 1955i 
Moldenke, Biol. Abstr. 30: 1093. 1956; Moldenke, Am. Midi. Nat. 
59: 356 & 362. 1958; Moldenke, R^surn^ 29, 31, 32, 39, 22i;, 237, 
360, 361, 370, 371, & U72. 1959; Moldenke, Phytologia 8: 121 & 
I2U (I96I) and 8: 279. 1962; Moldenke, R^sum^ Suppl. 3: 10 (1962), 
h: h, Hi, 'St 17 (1962), 5: h (1962), and 6: U. 1963; Gleason & 
Cronquist, Man. Vase. Pl. 580. 1963; Moldenke, Phytologia 8; U71- 
U73 & U77 (I963) and 9: 3h, 36, 156, 165, & 166. I963. 

Short-hairy herb, to 1 m. tall, slender; stems upright, branch- 
ed from the base, hirsute; branches 10 — 35 cm. long; leaves de- 
cussate-opposite, the blades 1 — 5 (usually less than 5) cm. long, 
rather narrow, entire or coarsely toothed, pinnately cleft, or 
almost parted, with the segments again incised or coarsely toothed, 
rugose, somewhat scabrous and finely pustulate above, the veins 
more or less prominent beneath, hirsute on both surfaces; inflor- 
escence spicate, the spikes solitary or tending to be panicled, 
usually short-pedunculate, slender, open, hirsute; bractlets lan- 
ceolate-acuminate, commonly not longer than the calyx; flowers 
small or very small; calyx about 3 mm. long, hirsute-pubescent 
and very slightly glandular, its teeth short sind subulate; corol- 
la hypocrateriform, varying from blue, dark-blue, or blue-lavender 
to blue-purple, purple, or pink, its tube scarcely longer than 
the calyx, the limb approximately U mm. wide; cocci trigonous with 
a convex back, about 2 mm. long, very shallowly reticulate- 
scrobiculate on the upper half, longitudinally striate below, the 
commissural faces extending to the tip of the coccus; chromosome 
niimber: n (x) = 7; 2n = ll|. 

The type of this rather puzzling species was collected by 
Charles Wright ( no. ll;97 ) in the borders of thickets near Copper- 
mines, at 1900 meters altitude. Grant County, New Mexico, in 1351, 
and is deposited in the Gray Herbarium of Harvard University. 
Gray's original discussion says "Borders of thickets near the 
Coppermines, New Mexico, Wright, Bigelow. Appears as if a hybrid 
between V. canescens and V. of fi^cinalis . S. Arizona, similar in 
foliage but with long bracts, Rothrock." Torrey's V. officinalis 
var. hirsuta vras also described as from tie ar the copper mines" 
and was probably based on the same collection. 

The species in its typical form has been found in wet ground, 
pifion pine-oak forests, grasslands and mesquite grasslands, along 
streams, at the base of hills, on hills along creeks, rocky slopes, 
gravel and sandbars of creeks, in granitic soil on foothills, and 
amongst rubble along the beds of small streams, at altitudes from 
180 to 2665 meters, blooming in February and from April to Novem- 



178 PHTTOLOGIA Vol. 10, no. 3 

ber, in fruit in February and from July to November. Hinckley 
describes it as frequent at middle and upper elevations in can- 
yons. Warnock found it "sparse in deep igneous soil". Smith, 
Peterson, & Tejeda encountered it in gray to nhitish soils with 
occasional calcareous outcrops and thorn-scrub-cactus cover in 
Puebla. It is reported as cultivated in Massachusetts. 

Briquet (I89U) mentions, without description, a "V. neo- 
mexicana " which he says is a hybrid between V. canescens H.B.K. 
and V. officinalis L. I have no doubt that he was referring to 
the plant here under discussion, found rather abundantly in 
western Texas, New Mexico, and adjacent Arizona and Mexico. I do 
not believe that it is of hybrid origin; certainly one of the 
putative parents, V. officinalis , occurs only very sparingly in 
this region. Wooton says of his unnumbered collection from Rui- 
dosa Creek, July 3, 1895, "larger and of different habit frcam V. 
canescens . Probably a good sp." and with this statement I concur. 
Two rather ill-defined varieties are described hereinafter, but 
V. necmexicana var, grandiflora Moldenke is a synonym of V. 
clover ae Moldenke. " 

Verbena neomexicana has been crossed Ydth V. halei Small, 
producing xV. notha Moldenke, and with V. urticifolia L., produ- 
cing xV. torpa Moldenke. Herbarium specimens have been misiden- 
tified and distributed under thr names V. cainescens H.B.K., V. 
halei Small, V. neomexicana var. hirtella Perry, V. neomexicana 
var, ^lopoda Perry, and V. xutha Lshm. 

On the other hand, the G_. L. Fisher 32238, H_^ C^ Hanson 6h$, 
Herrick 71$ , LeSueur 53, E. G. Marsh 278 , C_. H^ IJueller 2375 <Sc 
8139, Parks & Cory 7396 , Sperry U78 & T.321 , B. H. Yfarnock 
U6615 & W.I68 , and Wooton 3852, distributed as V. neomexicana , 
are all var. hirtella Perry; Blumer l80ii , 2170 , &: H.159 , Goodding 
33U & 22U6 , Gould & Pultz 3I6U, E. H^ Graham 3231, Griffiths |jl22, 
G. J_. Harrison U778, Harrison & Kearney 5796 & 6689 , Knobloch 
I30U , Mearns 1013 & 1887 [and maybe 1913 ] , E. W. Nelson 616I & 
6271, Nelson & Nelson 2018 , Peebles , Harrison , & Kearney 3790 , W. 
W. Price s.n. [May 1, 139U] , Thomber & Thornber s.n. [Total 
Wreck Mts., May 7, 1902], Tourney s.n. [Copper Basin, Aug. 2U, 
189!^], ToTmsend & Barber 192, and Wooton s.n. [Santa Rita Range 
Reserve, May 12, 1912] are all var. xylopoda Perry; Kenoyer & 
Crum 2879 & 2951 and E. G_. Marsh 278 are V. canescens H.B.K.; E^ 
J . Palmer 10002 , Tharp 352 , I36I , s.n. [Austin, 5/27/20] , s.n. 
Tlangtry-Dryden, 6-lli-1931] , and s.n. [10/9/36], B_. U^ Warnock 
11238 , and Warnock & McBryde II1.9O8 are V. canescens var. roemeri - 
ana (Scheele) PerryJ E^ W^ Lathrop 20U6 is V. halei Small; H^ S. 
Gentry 1923 is V. pinetorum Moldenke; and Cory 53.U75 , V. L. Hams 
1752 , E. J_^ Palmer"T2"382 , Tharp U3-8OI; , and Warnock & Mullins 
])|)ing are V. plicata Greene; Kamraerer 33 is V. wrightii A. Gray; 



196U Moldenke, Monograph of Verbena 179 

0. B. Metcalfe 612 is a mixture with something not verbenaceous, 
while Warnock 21715 is definitely not verbenaceous . The Sixth 
Grade Brownsville 3U and s.n. [Nov. 19 3U] and Tharp 1201 , cited 
by me in Lundell's Flora of Texas as V. neomexicana , are actually 
V. runyoni Koldenke. Sperry $83 , distributed as "V. neomexicana 
var.?" is V. halei Small. 

Perry cites Ferris & Duncan 2607 (E) and E. J_. Palmer 30791 
(E) as var. hirtella , but I think that they are better placed in 
the typical form of the species. Similarly, E. A^ Phillips 753 
and S. S. White 2202 and 3836 have been annotated by me as var. 
xylopoda, but the University of Michigan sheets of these collec- 
tions show definitely very small flowers, and, I believe, repre- 
sent the typical form. Likewise, £. S. llhlte I678 and Yfhitehead 
1238 were annotated by me as var. hirtella in some herbaria, but 
the University of Michigan specimens show lobed or even very 
deeply lobed leaves and I feel that they had better be refeired 
to the typical form of the species. 

Perry (1933) cites the following 7 additional specimens not as 
yet seen by me: MEW MEXICO: Grant Co.: E, L^ Greene s.n. [Pinos 
Altos Mts., 1880] (E, F)j C. Wright ll;97 (E~isotype, G— type, W- 
isotype). Socorro Co.: 0. B. Metcalfe 612 (E, G) . She comments 
that "The collections of Young 1703 , Wooton 61;6 , and Wooton (col- 
lection of July 3, 1895) are by no means typical of the species, 
but perhaps are better placed here than elsewhere. This species 
appears to be very closely related to V. canescens and V. gracil- 
is. It differs from both, however, in the upright habit and the 
nutlets. In both V. canescens and V. gracilis the commissural 
face does not extend to the tip of the nutlet and the striae on 
the dorsal surface are less conspicuous." In all, 81 herbarium 
specimens, including the type collections of most of the names 
involved, have been examined by me. 

Citations: TEXAS: Brewster Co.: C_, H. Muller 8I38 (Au, Ca — 
88285U, Mi, N, N); Sperry U77 (Om)j B. H. Warnock 9677 (Rf ) . 
Jeff Davis Co.: Ferris S: Dime an 2606 (Du— 12U293), 2607 (Du— 
125U18, Gg~31385, N); Hinckley 170 [9/15/3U] (N), 170 [8/13/35] 
(N), s.n. [Mt. Livermore, 9/15/3U] (Au, Au) , s.n. [July 5, 
1936] (N); E^ J^ Palmer 30791 (Au), 31367 (Au) ; Sperry T.811 
(Om), T.312 (Om); Tharp U;93 (Au); M. S^ Young 1703 (Au, V^— 
lllii6U7). Travis Co.: A. M^ Ferguson s.n. [April 1901] (Au) . Val 
Verde Co.: Collector undesignated s.n. [Del Rio, Aug. 19, 1932] 
(Dp); Parry , Bigelow , Wright , ^ Schott s.n. [valley of the Rio 
Grande below Doflana] (V;~56l97) • IJEiY MEXICO: Catron Co.: Sggles- 
ton 20292 C.V~15228U6) . Grant Co.: Chapline 6O9 [Herb. U. S. 
Forest Serv. 2506U] (N); E^ L_^ Greene s.n. [Pinos Altos Mts., 
Aug. 23, 1880] (Vt); Holzinger s.n. [G. 0. S. Ranch, 2? Aug.— 12 
Sept. 1911] (W~660558), s.n. [East Canton] ('W~660559) ; Cj. 
'iVright IU97 (Ca — 2U83Uli — isotype) . Lincoln Co.: Goodman & Water- 



180 PHYTOLOGIA Vol. 10, no. 3 

fall U967 (Ok) J Wooton 6U6 (N, Ur), s.n. [Ruidosa Creek, July 3, 
1895] (C, N, W— 563952, W--736886). Sierra Co.: Beals s.n. [Lake 
Valley, July I90U] (Mi) ; 0. B^ Metcalfe 955 (W--U9785U) . Socorro 
Co.: Oj_ B^ Metcalfe 612 , in part (Ca— 882628, N, Po— 70886, W— 
U95596). ARIZONA: Cochise Co.: Gould & Pultz 3161; (W--1893186) j 
A. R. Moldenke 6I6 (B, S) . Gila Co.: Collom 355 (W~1683306) . 
Maricopa Co.: Whitehead 1288 (Mi, N) . Pima Co.: S. B. Parish s.n. 
[Herb. Osten I30II] (Ug); Skj^^t-Pedersen 9l|6 (Cp). Yavapai Co.: 
Gould & D arrow U-80 (W— 200873U) . MEXICO: Baja California: Meling 
3 (Du— 2255I;5, N, Po~209970, W— 1635283) . Chihuahua: S. S. 
White 2202 (Hi, Oa) . Coahuila: S^ S_. White 1678 (Mi, Tu— 35529) . 
Guanajuato: Yfaterfall 16553 (Ca) . Mexico: Vfaterfall I6I476 (Z) . 
Nuevo Le6n: Mueller & Mueller 157 (Au, Me); S^ S. "'.Thite 1577 (Ml, 
W— 1822811;). Sonora: E. A_^ Phillips 753 (Mi, Rf, Sm)j S_. S. 
White 2787 (Mi), 3099 (Mi), 3836 (Mi, N, Vf— 2132273), 3836a (Mi), 
3382 (Mi, W--213228U). LOCALITY OF COLLECTION UNDETERMINED: 
Herb. Torrey s.n. (T) . 

VERBENA NEQMEXICANA var. HIRTELLA Perry, Ann. Mo. Bot. Gard. 20: 
298—299. 1933. 

Synonymy: Verbena neomexicana var. hirstella Perry ex Molden- 
ke, Alph. List Invalid Names Suppl. 1: 25, in syn. 19U7. Verbena 
mexicsina Sperry ex Moldenke, R6suml Suppl. U: 17, in syn. I962 
[not V. mexicana L., 1753, nor Pers., I96O] . 

Bibliography: Perry, Ann. Mo. Bot. Gard. 20: 260, 261. 298 — 
299, & 355. 1933; Cory, Texas Agr. Exp. Sta. Bull. 550: 88. 1937; 
Sperry, Sul Ross State Teach. Coll. Bull. 19: 68. 1938; Moldenke, 
Annot. & Classif. List 108. 1939; Sperry, Sul Ross State Teach. 
Coll. Bull. 22: Ul. 19Ul; Moldenke in Lundell, Fl. Texas 3 (1): 
16, 17, & 30 — 31. 19U2; Moldenke, Knovm Geogr. Distrib. Verbenac ., 
[ed. 1], 3, 13, lU, 19, & 101. 19l;2; Moldenke, Phytologia 2: 128. 
19l;5; Moldenke, Alph. List Cit. 1: lOU, 176, & 22U. 19li6; Molden- 
ke, Phytologia 2: 331. 19U7; Moldenke, Alph. List Invalid Names 
Suppl. 1: 25. 1917} Moldenlce V^rightia 1: 227-— 228. 19l;8; Molden- 
ke, Alph. List Cit. 2: li67, U75, 1;76, U98, 5l7, 522, 527, 538, 
539, 596, & 6U0 (19U8), 3: 656, 666. 679, 68U. 698, 707, 7U8, 
752, 753, 756, 790, 795. 797, 798, 807. 833, 873, 905, 953, 9Sh, 
963, & 966 (I9li9), and U: 989~993, 1071, 1107—1110, 1121, 1122, 
1126, lll;2, llli9, 1150, 12U0, 12U1, I2U;, & I2U7. 19U9} Moldenke, 
Known Geogr. Distrib. Verbenac, [ed. 2], 6, 2U— 26, 33, & 198. 
19U9; McDougall & Sperry, Pi. Big Bend IU6, fig. li;5. 1951; Mol- 
denke in Gleason, New Brit ton & Br. Illustr. Fl., print. 1, 3: 
126, 131, & 132 (I952) and print. 2, 3: 126, 131, & 132. 1958; 
Moldenke, R6suml 8, 29, 31, 32, 39, 370, li2U, & ii72. 1959; Mol- 
denke, Resume Suppl. 1: 3. 1959; Lewis & Oliv., Am. Joum. Bot. 
Ii8: 639— 61;1. 1961 ; Moldenke, Phytologia 8: 12U. I96I; Moldenke, 
Resume Suppl. 3: 10 & UO (1962), U: 1; & 17 (1962), and 5: h. 
1962; Moldenke, Phytologia 8: U72, 1;77, & 1;78 (I963) and 9s 156, 
165, & 199. 1963. 



196U Moldenke, Monograph of Verbena l8l 

Illustrations: McDougall &: Sparry, PI. Big Bend fig. Iii5. 
1951 J Moldenke in Gleason, New Britton & Br. Illustr. Fl., print. 
1, 3: 132 (19^2) and print. 2, 3: 132. 1958; Lewis £t Oliv., Am. 
Journ. Bot. hQ: 6i;0. I96I. 

This variety differs from the typical form of the species in 
being densely canescent-hirtellous, the leaves more or less shal- 
lowly incised, the bractlets usually broadly ovate-acuminate, and 
the corolla-limb about 8 mm. wide. 

It is described as an erect, slender, perennial herb; stems up- 
right, branched, densely canescent-hirtellous; leaves 1 — $ cm. 
long, more or less shallowly incised or lobed, sometimes entire, 
canescent-hirtellous on both surfaces, the veins more or less 
prcsninent beneath; spikes solitary or sometimes tending to be 
panicled, usually short-pedunculate, canescent-hirtellous and 
glandular throughout; bractlets usually broadly ovate-acuminate, 
commonly not longer than the calyx; flowers large; calyx about 3 
mm. long, canescent-hirtellous and very slightly glandular, its 
teeth short and subulate; corolla varying frcm blue, bluish, or 
bluish-purple to blue-lavender, light bluish-lavender, lavender, 
purplish, purple, or deep-purple, its tube scarcely longer than 
the calyx, its limb about 8 mm. wide; nutlets trigonous, about 2 
mm. long, with a convex back, very shallowly reticulate-scrobicu- 
late on the upper half, longitudinally striate below; chromosome 
nxunber: n = 7. 

The type of this rather poorly defined variety was collected 
by Ernest Jesse Palmer (no. 3U065) among dry arroyos, plains, and 
foothills of the Chisos Mountains, Brewster County, Texas, on May 
22, 1928, and is deposited in the herbarium of the llissouri Bot- 
anical Garden at St. Louis. The variety has been collected in 
sandy, rockj'', limestone, or granitic soil, in dry arroyos, desert 
scmb and washes, crevices of rocks above vmterways, foothills, 
old laJcebeds, on gravel knolls, in rocky fields, grasslands, and 
rolling grasslands, on low mountainsides, flats, rocky hillsides, 
rocky open hilltops, plains, gravelly mesas, limestone hillsides, 
and sandstone mountainsides, along small sandy draws, and along 
the banks of dry streams, at altitudes of 1260 to 8000 feet, 
flowering fron January to November, fruiting from February to Oc- 
tober. Wamock found it "scattered" or "infrequent" in Brewster 
Coimty and "infrequent" or "frequent along highways" in Presidio 
County, Texas, and "frequent on limestone soil" in Presidio 
County; Young found it "common in open valley" in Brev^ster Coun- 
ty; Hinckley & Wamock describe it as "infrequent sind widespread 
along creek" in Presidio County; while Hanson reports it "common 
on mountainside". In Webb County it was encoiintered by litis, 
Koeppen, & litis "in fine sandy red clay soil on microphyll des- 
ert" with various species of Acacia and Opuntia, Prosopis , many 
spiny shrubs and weeds, Aristida, and Bouteloua . In Nu^vo Le6n 
it is said by Mueller to be "abundant in shaded situations" and 
"common in scrub oak and pinyon"; Mueller & Mueller describe it 
as "sparse"; while Stanford, Retherford, & Northcraft encountered 
it "in scrubby woodland association of pine, juniper, and oaks, 
heavily grazed by goats". 



182 PHYTOLOGIA Vol. 10, no. 3 

The name Verbena mexicana Sperry is based on Sperry li78, col- 
lected on flats in Brewster County, Texas, and deposited in the 
herbarium of Omer E. Sperry at Alpine, Texas. Verbena mexicana L. 
and V. mexicana Pers. are synonyms of Priva mexicana (L.) Pers. 

The variety is recorded from Gila and Maricopa Counties, Ari- 
zona, in my 19U9 and 1959 publications, apparently in error, tin- 
less the two collections cited hereinafter from that state prove 
to have originated in those counties. The Herb. Univ. Texas s.n. 
[Devine-Dilley, 3/1/30] cind Tharp s.n. [Devine-Dilley, March 1, 
1930] may have been collected in either Medina or Frio Counties, 
Texas, while Tharp 87i|ii may have come from either Duval or Jim 
Wells County. As to the LeRoy s.n. specimen cited below from Bal- 
timore County, Maryland — it bears a label inscribed "near Balti- 
more, Md., 1866", but Miss Perry says "surely not collected at 
Baltimore unless cultivated". There is also the possibility of 
it having been introduced accidentally as a weed there and the 
station now destroyed in the growt.h of the city. I am inclined 
to the latter theory. 

Herbarium specimens of this variety have been misidentified 
and dicjtributed under the names V. canescens H.B.K., V. gracilis 
Desf., V. halei Snail, V. nenthaefolia Benth., V. neomecicana 
(Gray) Small, V. neomexicana (A. Graj') Small, V. neomexicana var. 
xylopoda Perry, V. officinalis L., V. orcuttiana Perry, V. plica - 
ta Greene, V_. polystactya H.E.K., V. urticaefolia L., and V. 
xutha Lehm. 

On the other hand, the Meling 3 and S. S. liVhite 3836a, distrib- 
uted as V. neomexicana var. hirtella , at least in some herbaria, 
are typical V. neomexicana (A. Gray) Small (with the corollas only 
U, not 8, mm. wide); Mexican Biol. Exped. Univ. Ill . 992 and R. A. 
Schneider 992 are var. xylopoda Perry; and Tharp s.n. [Austin, 11- 
17-39] and E. W. Lathrop 2075 are V. canescens var. roemeriana 
(Scheele) Perry. M. E. Jones 2622U is a mixture with V. canescens 
var. roemeriana, irtiile B. H. Wamock and Wiggins i;360 are mixtures 
with V. plicata . S. S. White 2168 is annotated by me in some her- 
baria as var. xylopoda Perry, but the University of Michigan spec- 
imen shows leaves not at all lobed, so I think the collection had 
better be placed in var. hirtella. S. S. White 16 78 and Whitehead 
1288 Were annotated as var. hirtella or var. xylopoda by me in 
some herbaria, but the University of Michigan specimens show lobed 
or even deeply lobed leaves and I now feel that these collections 
are better placed in typical V. neomexicana. The Tharp s.n. [6/19/ 
31], R. H_- Painter 2U9 , and Ecology Class Uni v. Texas s.n. [2.28. 
30] cited by me as var. hirtella in Lixndell's "Flora of Texas" 
(19i;2) are now regarded by me as representing V- cane scens var. 
roemeriana instead. H. C. Hanson 6U5 bears a label reading "Flora 
of Texas", but was collected 17 miles north of Ruidosa, New Mexico. 

Perry (1933) cites the following 10 additional specimens not as 



1961i Moldenke, Monograph of Verbena 18 3 

yet seen by me: TEXAS: Brewster Co.: Moore & Steyermark 3277 (E); 
E. J. Palmer 3U065 (E — type); M^ S_^ Young 112 (E) . Culberson 
Co.: Parry, Bigelow, Wright, & Schott s.n. [between Van Horn 
Wells and Muerte, 2 July 18527 (N, W) . Presidio Co.: H. C. Hansen 
6U5 (G); McKelvey 20U6 (G) . MEXICO: Coahuila: Purpus 109U (E, F, 
G) . The Ferris ^ Dime an 2607 and E^ J_. Palmer 30791 , however, 
which she cites, I regard as typical V. neomexicana. She says 
"The pubescence of this variety is much finer and shorter than in 
the above [var. xylopoda ] . The leaves are not so deeply incised 
smd often tend to be elongated. The leaves of Purpus 109h are so 
narrow and shallowly incised that it appears superficially like 
V. perennis ; nevertheless the character of the pubescence allies 
it with this variety." I may add that Mueller & Mueller $06 also 
has the leaves so very narrow, only the basal ones with a pair of 
narrow lateral lobes, that it also greatly resembles V. perennis . 
The E^ Wilkinson s.n. [Sta, Eulalia hills, 30 July 188^] cited 
below, is cited by her as var. xylopoda . 

In all, 162 herbarium specimens, including type material of 
all the names involved, have been examined by me. 

Citations: MAJIYLAND: Baltimore Co.: LeRoy s.n. [near Baltimore, 
Md., 1366] (C). TEXAS: Bexar Co.: 0. M. Clark 7l4ia (Ok— 18719) . 
Brewster Co.: Lewis & Oliver 5U$7 (Nb); Moore & Steyermark 3277 
(Ca-~U 71230, Du— 22U1;76, Gg— I9U778, Mi, N); cT H. Mueller 8139 
(Au, N, W~1651901)} L^ T. Murray s.n. [May 22, 1928] (It)j E. J. 
Palmer 3U06$ (N—isotjT^e); Parks ?c Cory 703U (Tr), 7396 (Tr); 
Rose-Innes & Moon 1200 (Au) ; Rose-Innes & Wamock $31 (Au) ; 0_, E. 
Sperry 73 (W— 168U73), 9U (Om, W— I68UI83), U77 (W— 168U;23), 
U78 (Om, W--I68iil2l), 1536 (Om), I70U (Om), T.761 (Om, W— I8U8966), 
t7?21 (Om); Steiger 1373 (N); Tharp s.n. [Wilson Ranch, 6/19/31] 
(Au, Au, Au), s.n. [6-19-31] (Sm) , s .n ♦ [Wilson Ranch, June 1931] 
(Au, Au), s.n. [Wilson Ranch] (Au) ; B. H_. Wamock $37 (N) , 201; 36 , 
in part (Au) , C.8OI (N), W.I68 (N), s.n. [May 3, 1937] (Au, Mv) ; 
M. S_. Young 101 (Ur), s.n. [8/12/1$] (Au) . Dimmit Co.: M. E. 
Jones 28993 (Po— 137972) . " Frio Co.: C^ H. ttueller 2612 (Ar~ 
I7IOU2) ; Tharp s.n. [Devine-Dilley, March 1, 1930] (Au) . Pludspeth 
Co.: Tharp U>80U (N); V^aterfall $lli3 (N, Ok). Jeff Davis Co.: 
Hinckley s.n. [Limpia Canyon, July 1936] (Fs); D. C. Ingram 2$22 
[U. S. Dept. Agr. Forest Serv. 96I87] (W— 23l4ii7l47) ; 1^ E. Jones 
2622U, in part (Po— 1783U$); A. R. Moldenke 6U6 (S). Jim Hogg Co.: 
parks & Cory 1690$ (Tr) ; Tharp s.n. [June 1$, 1928] (Au) . Jim 
Wells Co.: Tharp Qlhh (Au) . La Salle Co.: £. Skottsberg s.n. 
[Millet, U/6/193$] (Go). Live Oak Co.: Tharp s.n. [March 21, 
1932] (Au) . Medina Co.: Herb. Univ. Texas s.n. [Devine-Dilley, 3/ 
1/30] (Au, Au). Presidio Co.: Correll & Rollins 2 36$ 2 (Ld); Gar- 
ner $0 (Au); H. C. Hanson 6U$ (N); Hinckley IO89 (Au, Au, N) , 



I81i PHYTOLOGIA Vol. 10, no. 3 

1971 (Au, N), 2U28 (N), 32ii6 (N), s.n. [Marfa, April 1938] (Fs); 
lanckley & Warnock U69U2 (Au, N, S); D. C Ingram 2607 [Herb. 
Forest . Serv. 96392] (S)j J. R. Lundell 5^(Ld)j Lundell & Lundell 
lli3U0 (N, Rf, W— 1927012); McKelvey 201^6 (Ca— It800U2, Tf— 1581336); 
Parks & Cory 26i;01 (Tr); B. H. Vfarnock 8 908 (Rf ) , 1U337 (Rf), 
Ii66l5 (Au, Ca— 7319li9, N, Ok, S); Warnock & McBryde lk$$$ (Rf ) . 
Travis Co.: C. L. York U6035 (Au, N) . Val Verde Co.: G. L. Fish- 
er 32238 (W— I62U2OI); M. E. J ones 28296 , in part (Du— 239763) J 
Parry , Big slow , Wright , & Schott 827 , in part (W— ^6198) ; White- 
house Illi79 (Sm). Webb Co.: H. J. Hamby 3U3 (Ar~206601), 792 
(Ar— 206289) J litis , Koeppen , & litis 19 (Vfs) ; Tharp 3687 (Au, W- 
1289911) • County undetermined: Havard s.n. [Texas] (W — 2203^9) . 
NEW MEXICO: Dona Ana Co.: Parry , Bigelow , Wright , & Schott 827, 
in part (N, N) . Lincoln Co.; H. C. Han3on~5Iir Tw— 982971) « So- 
corro Co.: C^ L. Herrick 715 (W~737262)} Woo ton 38^2 (Ur, W— 
73622U). ARIZONA: County undetermined: D. T. MacDougal 63 I1 (Au, 
N)} Purpus 8233 [Besner Creek] (Ca— 139759) . MEXICO: Baja Cali- 
fornia; T. Sj_ Brandegee s.n. [San Enrique, May 2, 1889] (Ca — 
10li853); G. Lindsay s.n. [July 11, 1937] (Du— 209288); Raven , 
Mathias , & Turner 12587 (Ca— 171723, Du— l4l8ll;9). Chihuahua: 
Correll & Johnston 21737 (Ld); Knobloch 252 (Z), 935 (Sm); Aj, Lee 
hS (Au, N); LeSueur 53 (Au, Ca— 712702, Gg— 319703, W— 1887827), 
231 (Au) ; F. W^ Pennell 13691 (W— 16U1250) ; Shreve 9O89 (W— 
179071a); S. S. White 2168 (Mi, Tu— 35536) , 2316 (Mi); E. Wilkin- 
son 37 (W— 1323066), s.n. [Sta. Eulalia hills, July 30, 1385] 
(Ob— 50870, Y^— 219la3). Coahuila: E. G. Marsh Ihl (Au, Au), 278 
(Au— 212U49), 681; (Au— 122335, Ok); Ed/f. Palmer 25 (Ca— 10U8U9); 
Purpus IO9U (Ca— 139758, II, Po— 63882), II3U (Ca— 139721); Stan- 
ford , Retherford, ^ Ilorthcraft 23U (Ca— 713322, Du— 289331, N, 
3e — 69590); Water fall 16623 (Ca) . Durango: Reko 5201 (',7— 
1318979); T/aterfail £; V/allis 133U2 (Ok). Nuevo Le6n: C_. H. Muel- 
ler 2375 (Au, Mi), 2391 (Au); !,:ueller & Mueller ii65 (Au) , 506 
(Au, Mi); R. A. Schneider 992 (Mi, N) . Oaxaca: Purpus 3U08 (Ca — 
138826) . Puebla: Purpus 3927 (Ca— 138825) . 

VERBENA NEOMEXICANA var. HLOPODA Perry, Ann. Mo. Bot. Gard. 20: 
297—298. 1933. 

Synonymy: Verbena canescens var. arizonica Gray ex Moldenke, 
Alph. List Invalid Names Suppl. 1: 23, in syn. 19U7. Verbena 
neomexicana xylopoda Perry ex Moldenke, RisumI 370, in syn. 1959. 
Verbenaceae neomexicana (Gray) Small ex Moldenke, R^sum^ Suppl. 
U: 21, in syn. I962. 

Bibliography: Perry, Ann. Mo. Bot. Gard. 20: 261, 285, 297~ 
298, & 355. 1933; Moldenke, Known Geogr. Distrib. Verbenac, [ed. 
1], 13—15, 19, & 101. 19li2; Moldenke in Lundell, Fl. Texas 3 (1) : 
17 & 30. I9U2; Moldenke, Phytologia 2: 128. 19ii5; Moldenke, Bot. 



1961; l!oldenke. Monograph of Verbena 185 

Gaz. 106: l6l. 19U5; lloldenke. Am, journ. Bot. 32: 610. 19U5; 
Moldenke, Alph. Ust Git. 1: i;6, 77, 182, 235, 2^2, 2hh, 260, & 
27li. 19U6; Moldenke, Alph. List Invalid Names Suppl. 1: 23. 19U7; 
Moldenke, Phytologia 2: l6U. 19 U8; Moldenke, Wrightia 1: 228. 
19ii8j Moldenke, Castanea 13: 113. 19U8j H. N. & A. L. Moldenke, 
PI. Ufe 2: Uh. 19l;8: Moldenke Alph. List Cit. 2: 370, li03, U09, 
U72, U73, U75, U76, ^78— hSO, h9h, U97, U98, 519, 523, 527, 597, 
& 598 (19l;8), 3: 656. 700, 723, 72U, 731, 752, 755, 769, 771, 783, 
803, 807, 809, 813, 828, 831, 881, 905, 933, 952, & 963 (19U9), 
and h: 996, 1003, IO68, 1071, 1121, 1125, 1126, 1129, 111^8, 1166, 
1171, 117li, 1192, 1199, 1210, 1217, 122U, 1225, 1229, 1231, I2U0, 
I2U2, 12li3, & 1252. 19U9; Moldenke, Known Geogr. Distrib. Verben- 
ac, [ed. 2], 10, 2U, 26, 27, 33, & 198. 19ii9; Moldenke, Phytolo- 
gia 3: 375. I95O; Moldenke, ResumS 13, 29, 32, 33, 39, 361, 370, 
& U72. 1959; Moldenke, R^sum4 Suppl. U: 21. 1962; Moldenke, Phy- 
tologia 8: U72 & 1;77 (1963) and 9: 36, 156, & 215. I963. 

This variety differs from the typical form of the species in 
having its stems somewhat coarser, the pubescence shorter, denser, 
and more glandular, the calyx about h ran. long and glandular- 
hirsute, with acuminate teeth, aid the corolla-limb 6 — 10 mm, wide. 

It is described as an herb to 30.5 cm. tall, the leaves deeply 
lobed, the bractlets 3 — 3-^ nm. long, the flowers large, the cal- 
yx 3.5 — h ram. long (including the teeth) and glandular or densely 
glandular, the corolla large or very large, varying from blue, 
blue-violet, or blue-lavender to dark-blue, lavender, lavender- 
pink, or purple, more or less 2- lipped, the tube i; — 6 mm. long 
(including the throat), the limb 9 — 10 mm. wide, and the nutlets 
scabrellous on the ccxamissure. 

The type of this rather ill-defined variety was collected by 
Herbert Christian Hanson (no. A. 1130) on rocky slopes in Sabino 
Canyon, Santa Catalina Mountains, Pima Coimty, Arizona, on April 
21, 1922, and is deposited in the herbarium of the Missouri Bot- 
anical Garden at St. Louis. Verbena canescens var. arizonica is 
apparently'- based on an unnumbered collection made by Cyrus Guern- 
sey Pringle in the foothills of the Santa Rita Mountains, pima or 
Santa Cruz County, Arizona, on May H, I88i|, and is deposited in 
the Gray Herbarium at Harvard University. 

The variety has been collected among rocks, in granitic soil 
on foothills, in sandy or limestone soil, on gravelly slopes, 
rocky hills, open dry rocky slopes in chaparral, dry plains and 
gentle slopes, gravelly brush-covered plains, rockj' limestone 
slopes, decomposed limestone, and stony slopes, along moist stream- 
sides, bordering streambeds, in pine woodlands, scxnibland on lime- 
stone mountainsides, canyons, pinelands or pine woodlands, grass- 
lands and oak-eras slands, dry rocky canyons, and calcareous des- 
erts, at altitudes of 100 to 8OOO feet, flowering and fruiting 
frcci Febmary to October. 

Maguire found the plant on "gravelly west slopes in raesquite and 
Opuntla cholla association", while Tucker found it in "pine-oak 
woodlands of Pinus chihuahuana, P. engelmannii, Quercus arizonica, 
Q. viminea, some shrubby Q. liypoleucoides, and Arbutus"; Wiegand, 



186 PHYTOLOGIA Vol. 10, no. 3 

Magiiire, Richards, & Moeller encountered it on "gravelly slopes, 
west exposure, under Quercus and Rhus "; Stanford, Retherford, & 
Northcraft found it in a "mountain valley with much bare lime- 
stone present, partially covered with Agave and Yucca". Mueller 
says that it is "common in meadows" and "common on rocky slopes 
in open pine-oak forests"; Wiegand, Maguire, Richards, & Moeller 
describe it as "uncommon on gravelly slopes under Quercus and 
Arctostaphylos"; iriiile Runyon avers that it is "widespread on 
hills in Starr Co, [Texas]". Benson collected it in the Lower So- 
noran Zone of Arizona. Blumer, in a note on his collection no. 
I80li, says "may be introduced". 

The Meams 1918 collection frcm the Niggerhead Moxintains, near 
Monument 82, is hereinafter cited from Sonora, Mexico, on the au- 
thority of Perry (1933) • 

T. H. Kearney comments in regard to Eastwood 18U;7 "approaches 
V. menthaefolia (nutlets scaberulous on the commissure)"; in re- 
gard to Harris on & Kearney 5796 he says "calyx .glandular, 3.5 mm. 
long, nearly as long or slightly surpassing the calyx [sic]"; and 
in regard to Kearney & Peebles 103^5 says "V, menthaefolia Benth. 
is very similar in appearance to V. neomexicana but has more 
slender spikes and more reduced bracts . It is questionable 
whether the two forms are specifically distinct." 

Wynd & Mueller 525 was originally distributed as " Verbena xu- 
tha Lehm. vel aff .", while Edw. palmer lOUl was annotated by Perry 
as~"Af f . V. Halei or V. xutha " . 

Material of V. neomexicana var. xylopoda has been misidentified 
and distributed in herbaria under the names V. brevibracteata (A. 
Gray) Eggert, V. canescens H.B.K., V. canescens var, neo-mexLcana 
A. Gray, V. canescens neo-raexicana Gray, V. halei Small, V, hasta- 
ta L., V, neomexicana (A. Gray) Small, V. neo-mexicana (A. Gray) 
Small, V. neo-mexicana Gray, V, neomexicana var. hirtella Perry, 
V. officinalis L., V. prostrata A. Br., V. remota Benth., V^ sor - 
oria D. Don, and V, xutha Lehm. 

On the other hand, the Mueller & Mueller 506, distributed as 
var. xylopoda , is actually var. hirtella . E. A. Phillips 753 and 
S. S. White 2202 and 3826 were annotated by me as var. xylopoda 
in several herbaria, but the University of Michigan sheets show 
definitely very small flowers (less than 6 mm. wide), so I now re- 
gard these three collections as representing typical V. neomexica- 
na, as I also do Gollom 355 and Gould & Darrow l^lSO . "~fhe corolla 
also seems very small on Peebles, Harrison, ft Kearney 1753, Tides - 
trom 872, and Peebles & Harrison 16U9 , but the pubescence on the 
stans in not that of typical V. neomexicana , so I think that in 
these cases the corollas were large when fresh and have merely 
shriveled up in the drying process. 

Helson & Nelson 5138 , cited hereinafter as from Terrell County, 
Texas, -was actually collected between Terrell and Webb Counties 



196U Moldenke, Monograph of Verbena 187 

("between Sanderson and Laredo") and is very ancsnalous in appear- 
ance, the fruiting-calyx and fruit being much larger than is nor- 
mal for this taxon. It may represent an undescribed species. 
Griffiths 3U31 is a mixture with something non-verbenaceous . 

Annette Carter, Principal Herbarium Botanist at the University 
of California, in a letter to me dated February 18, 1955, states 
that she has checked through the University of California herbar- 
ium and the Jepson herbarium and finds no California specimens of 
this variety. Miss Perry (1933) cites the following 33 addition- 
al specimens not as yet seen by me: ARIZONA: Cochise Co.: Blumer 
180U (E, F, G), 2170 (F); Goodding 33U (G) ; Lemmon 2857 (G) . Mar- 
icopa Co.: Coues & Palmer 571 (E) . Pima Co.: H^ 2ji Hanson A. 1130 
(E — type), Santa Cruz Co.: Pringle s.n. [Santa Rita Mts., Hay 11, 
I88ii] (D, F, G). Yavapai Co.: M. E. Jones s.n. [Skull Valley, A- 
pril 28, 1903] (E)j Yf. W. Jones 3kh (G), 3U6 (G) . CALIFORIIIA: 
County undetermined: Edw. Palmer s.n. [I876] (G) . LIEXICO: Baja 
California: Edw. Palmer 312 (E, F) , s.n. [Big Canyon of Tantillas 
Mts., 10 Sept. 1875] (G) . Chihuahua: Hartman 6O8 (G)j Nelson 
6161 (G), 6271 (G); Edw. Palmer 52 (E, F, G) ; Pringle 270 (D, F, 
G)j Towns end & Barber 192 (E, F, G); Wislizenus 150 (E) . Sonora: 
Thurber U;6 (G, N) . It is probable that the Thurber » Uh6 " which 
she cites from the New York Botanical Garden herbarium is the 
Thurber 336 cited by me hereinafter from the Torrey Herbarium. 
She comments that "This variety differs from the species in the 
denser and more glandular pubescence and the larger corolla. The 
nutlets also are slightly longer, with the reticulations somewhat 
deeper than in the species, and the commissursil faces hardly ex- 
tend to the tip of the nutlets. The specimens Nelson 6I6I, 6271, 
and Townsend & Barber 192 closely resemble the above variety [var. 
xylopoda ] in inflorescence, but are more like V. neomexicana in 
the long and somewhat sparsely hirsute indument on the lower part 
of the stem; Pringle 270 and Wilkinson approach variety hirtella ." 

in all, 181 herbaritmi specimens, including the type of one of 
the names involved, have been examined by me. 

Citations: FLORIDA: County xindetermined: Collector undesignated 
s.n. [Florida] (Vt) . TEXAS: Hidalgo Co.: R. Runyon 55 (Rr) . Ter- 
rell Co.: Nelson & Nelson 5138 (Au) . Victoria Co.: Owens 3125 
(Au). Webb Co.: Perkins I, Hal3^ 2627 (Po — 256897). ARIZONA: Catron 
Co.: Eggleston 20292 (N) . Cochise Co.: Bameby 5l5l (N); Benson 
10299 (Po— 26765U); Blumer I80U (Du— 2la79, Ka— 60la6, N, Tu, W~ 
S6l9^6), 2170 (^T— 563573), N.157 (Tu), N.159 (Tu)j J. I_. Carlson 
s.n. [Warren, May 20, 1915] (Gg— 31ii69, V^~873378); Darrow g.n. 
[July 26, 19U3] (Tu— 1U767); Goodding 33li (Fg— 8032, N, Tu, Tu), 
22l;6 (Ca— U66879, Ca— I3OO3O, S)j Gould t Pultz 3l61i (Ca— 705195, 
N); Harrison & Kearney 5796 (To, W— Ili35l78)j M^ E. Jones s.n. 
[Sept. 3, 1903] (Po— 2U8OO3)} Kearney & Peebles 138U7 (To)j Mearns 



188 PHYTOLOGIA Vol. 10, no. 3 

1013 (Du— 9555, N, W--229031); W^ W. Price s.n. [May 1, 189U] 
(Du— 97119, PO--88556); Wiegand, Maguire , Richards , & Moeller 
11065 (Gg— 295125, N, N, PI— 13055U, Ua—UTUOii); T_. E. Wilcox s_. 
n. [Ft. Huachuca, Aug. 1892] (C). Gila Co.: Nelson & Nelson 2018 
(N); Wiegand , Kaguire , Richards , & Moeller 11336 (Gg— 295126, N, 
N, PI— 130553, Ua— U7U03). Mohave Co.: Eastwood 18UV7 (Gg— 
I889U7). Pima Co.: Benson 1022U (Po— 267653); Bumham U79 (I)} 
N. C. Cooper C.725 (Ak— 12632) j Eastvfood 17921 (Gg— 17li989); Gil - 
man 1120 (Po— 172237); E, H. Graham 3231 (Cm, Du— 2U6092); Grif- 
fiths 3i;31 (W— li97009), ia22 (W— 660150); G. J_. Harrison ^778 (To, 
W— 1367707); M. E. Jones s.n. [Aug. 3I, 1903] (Du— I518OI, Po— 
183132); Kearney & Peebles 10355 (To), IO8U6 (To); King & Beckett 
lli;02 (S); Munz lll;9 (Po— 98OU3); Peebles 686O, in part (Gg— 
192656); Peebles , Harrison , & Kearney 3790 (To, V/— 1367U8ii) ; Spal- 
ding s.n. [March 21, I9O6] (Tu, Tu); J. J. Thomber 8III (Tu) ; 
Wooton s.n. [Santa Rita Range Reserve, May 12, 1912] (W— 66395U) . 
Pinal Co.: J, Arnold s.n. [April 16, 1937] (N); G. J. Harrison 
6689 (Fo— 173002); Harrison & Kearney IU88 (To), T6H9 (N); C. B. 
Maguire IO88U (N, N, Ua~U7U02); Nichol s.n. [May 6, 1925] (Tu); 
Peebles 686O , in part (Gg— 18U987) ; Peebles & Harrison I6U9 (To); 
Peebles , Harrison , & Kearney 1753 (To) ; Thomber & T hornber s.n. 
[Total Wreck Mts., May 7, 1902] (Ca— 882627, Tu) . Santa Cruz 
Co.: Kearney & Peebles I3863 (To); Pr ingle s.n. [Santa Rita Mts,, 
May 11, I88U] (Be, Br, Cm, Ll, Pa, Up— 17121, Vt, W— 56199); 
Tidestrom 872 (W— 5077it9) . Yavapai Co.: Crosswhite 718 (Hi— 
19U682); M. E. Jones s.n. [Skull Valley, April 28, 1903] (Po— 
2U8OOI, W~856988), s.n. [Skull Valley, May 1, I903] (Po— 
2U8OO2); Tourney s.n. [Copper Basin, Aug. 2li, I89U] (W— 6I9I8I) . 
Coimty undetermined: Herb. LeRoy s.n. [Ariz. 'SU] (C). CALIFOR- 
NIA: County undetermined: Edw. Palmer 339 1/2 [Cent. Cal. I876] 
(W— 56181). MEXICO: Baja California: D. R. Harvey 539 (W— 
1685982) . Chihuahua: Correll & Johnston 21528 (Ld) ; M^ E^ Jones 
s.n. [Colonia Juarez] (Po — 2li800U); Knobloch I30U (Mi); LeSueur 
875 (Au); C. Ej_ Mueller 3li27 (Ca— 720115, Mi, Rf); E. W. Nelson 
6161 (W— 35987ii), 6271 (W— 359988); EdiT. Palmer 52 (N, W— 57352 
Pringle 270 [Herb. Osten I30I8] (Br, Ll, Me, Me, Mi, N, Pa, Ug, 
Up— 17097, Vt, W— 56159); Shreve 9097 (Ca— 73l8Ii5, Fs); Tovmsenc 
& Barber 192 (Ca— I3883U, Me, Me, N, Po— 711148, Ur, Vt, V/— 38321^ 
J. M. Tucker 2508 (Ca— 2791, Gg— Ul5500, Z); V/aterfall I6IOO (Ca^ . 
Coahuila: Barkley , Webster , & Rowell 7189 (Mi, N); Herb. Inst. 
Biol. Univ. Nac. Mex. 7138 , in part (Me); Stanford , Retherford , & 
Northcraft i;26 (N, Se— 70UU5); %nd & Mueller 525 (Fs, I, N, Ok, 
S, Ur, W— 1639936) . Nuevo Le6n: Barkley , Webster , & Rowell 7l50 
(Au— 123253, N); Mexican Biol. Exped. Univ. 111. 992 (Fs, N); C. 
H. Mueller 2305 (Fs, Mi), 2391 (Mi); Ed»r. Palmer lOlq (Pa, W— 



196U Moldenke, Monograph of Verbena 189 

56200) ; F. W. Pennell I688O (Me) ; Schery 32 (Mi) j R. A_. Sc hneider 
992 (Ur). San Luis Potos£: F. W. PenneU 1753$ (D— 733971, Me, W 
— I6ii0627). Sonera: Mearns 1887 (W— 232U28), I9I8 (Du— 95U9, W— 
232676); Merton 201^2 (W— 232685) ; Thurber 336 (T); S. S_. Tfhite 
3310 (Ml), 3985 (Ml), UI76 (Mi, W— 2132336); Wiggins 6l61i (Du— 
253Ua). Tamaullpas: Stanford , Lauber, & Taylor 2331 (IJ, N) . 
State undetermined: E. "tTilkinson s.n. [1885] (Ui) . 

xVERBENA NBQUAM Moldenke, Phji^ologia 5: 133. 1955. 

Synonymy: Glandularia laciniata x dissecta Schnack L Covas, 
Darwiniana 7: 73, in textu. 19U5« Verbena laciniata x dissecta 
Dermen ex Moldenke, Phytologla 3: ii67. 1951 » Verbena dissecta 
Spreng. x V. teniiisecta Briq. ex Moldenke, Rlsiim^ 363, in syn, 
1959. Verbena tenui secta Briq. x V. dissecta Spreng. ex Moldenke, 
R6suin6 376, in syn. 1959. Verbena dissecta Willd. x V. tenuisec- 
ta Briq, ex Moldenke, RisumS Suppl. 2: 11, in syn, I96O. 

Bibliography: Schnack & Covas, Darwiniana 7: 73. 19U5; Schnack 
& Covas, Revist. Argent. Agronom. 12: 22li. 19U5; Moldenke, Phyto- 
logia 3: U67 (1951) and 5: 133. 1955; Moldenke, Biol. Abstr. 30: 
1093. 1956; Moldenke, Am. Midi. Nat. 59: 356. 1958; Moldenke, R^- 
svani 22li, 296, 363, 376, & ii73. 1959; Moldenke, R6sum6 Suppl. 2: 
11 & 12. I96O; Moldenke, Phytologia 8: 121. I96I; Moldenke, R6su- 
m4 Suppl. U: 19. 1962; Moldenke, Phytologia 9: 69. I963. 

The name, xV. nequam, was proposed by me in 1955 for the hy- 
brid between V. dissecta Willd. and V. tenuisecta Briq., produced 
artificially by Schnack & Covas from cultivated material in Ar- 
gentina and described by them in 19U5 under the designation 
" Glandularia laciniata x G. dissecta". Evidence seems to indi- 
cate that what they consistently identify as Glandularia lacini- 
ata is actually Verbena tenuisecta. 

Verbena dissecta and V. tenuisecta grow together in at least 
two states of Brazil, in Uruguay, and in at least six provinces 
of Argentina. It is possible that they also occur together in 
parts of Bolivia and Chile. One night expect to find this hy- 
brid in the wild where the ranges of the parent species overlap. 
It is very possible that some of the almost innumerable "forms", 
"races", "mutants", "ecotypes", "lusi", or "variants" now gener- 
ally regarded as Vj_ laciniata (L.) Briq., V. dissecta Willd., V. 
tenuisecta Briq., V. santiaguensis (Covas & Schnack) Moldenke, V. 
mendoclna R. A. Phil., or other species in this complex may actual- 
ly represent this hybrid. Much more intensive work is urgently 
needed in this group of the genus. xV. nequam should certainly 
have considerable horticultural merit"! 

XVERBENA NISA Moldenke, R6sum6 Suppl. h: 10, 11, 18, & 19 (1962), 
nom, nov. 
Synonymy: Glandularia perakii x laciniata Schnack & Covas, 
Darwiniana 7: 73, in textu. 19U5. Verbena perakii (Covas & 



190 PHYTOLOGIA Vol. 10, no. 3 

Schnack) Moldenke x V. tenuis ecta Briq. ex Moldenke, R6sum6 Suppl. 
1;; 18, in syn. I962. Verbena tenuisecta Briq. x V. perakii (Cov- 
as & Schnack) Moldenke, R6sum6 Suppl. Ij.: I9, in syn. I962. 

Bibliography: Schnack & Covas, Darwiniana 7: 73. 19U5; Molden- 
ke, R^suml Suppl. h: 10, 11, 18, & I9. 1962. 

The name, xV. nisa , was first proposed by me in I962 for the 
hybrid, artificial or natural, between Verbena perakii (Covas & 
Schnack) Moldenke and V. tenuisecta Briq. This hybrid was appar- 
ently first produced in 19U14. by Schnack and Covas from cultivated 
material in Argentina and was referred to by them in their 19U5 
publication cited above under the designation " Glandularia perak- 
ii X G. laciniata " . Evidence seems to indicate, however, that 
the plant which they consistently identify as Glandularia lacini- 
ata was actually Verbena tenuisecta . As far as known now, the 
two parental species do not occur together in the wild anywhere, 
so it is not probable that this hybrid will be found in the wild, 
unless the ranges of the parental species are much extended in 
the future either by natural distributional expansion or by more 
collecting on our part producing a more complete picture of the 
natural ranges of the species. The hybrid should have considerable 
horticultural merit. 

VERBENA NIVEA Moldenke, Phytologia 1: hS? - 19hO. 

Bibliograplior: Moldenke, Phytologia 1: U59. 19U0; Moldenke, 
Known Geogr. Distrib. Verbenac. [ed. 1], Ui & 101. 19li2j Hill & 
Salisb., Ind. Kew. Suppl, 10: 2li2. 19U7; Moldenke, Known Geogr. 
Distrib. Verbenac, [ed. 2], 98, I06, & I98. l9U9i Moldenke, R^- 
sum6 115, 127, & I4.73. 1959} Moldenke, R6sum6 Suppl. h: $. 1962. 

Low procumbent or trailing herb, branched frcm the basej 
branches spreading in radial fashion, very slender, sometimes al- 
most filiform, obtusely subtetragonal, rather densely puberulent 
iTith very short spreading cinereous hairs, more sparsely so or 
glabrescent in age, the youngest branchlets very densely short- 
pubescent with spreading cinereous hairs; nodes faintly annulai ; 
principal intemodes 0.5 — 2.5 cm. long, mostly greatly abbrev? 
ted; leaves decussate-opposite, numerous, usually TrLth several 
much reduced ones in their axils; petioles slender, 1 — k nun. 
long, flattened, deeply canaliculate above, narrowly winged- 
margined, rather densely or sparsely strigose with appressed 
whitish hairs; leaf-blades chartaceous, uniformly gray-green o, 
both surfaces, ovate in outline, U — 8 mm. long aind wide, densel 
strigose with appressed whitish antrorse hairs on both surfaces^ 
3-lobed or -parted almos^ to the base, each division often again 
2- or 3-lobed, the lobes all obtuse at the apex and revolute- 
margined; the very slender midrib and secondaries impressed a- 
bove, slightly proninulous beneath; veinlet reticulation indis- 
cernible on both surfaces; inflorescence tenninal, capitate; 
heads small, many-f lowered , dense, subsessile or very short- 
pedunculate; calyx tubular, 3 — h mm. long, often purplish, dense- 
ly short-pubescent with whitish rather spreading hairs; corolla 



196U Lloldenke, Monograph of Verbena I9I 

white, bluish-white, or slightly purplish, its tube about 5 mm. 
long, the limb 2.5 — 3 nun. in diameter. 

The type of this species was collected by Santiago Venturi (no. 
lOOllj ) along the highway to San Antonio, Rosario de Lerma, at an 
altitude of 3000 meters, Salta, Argentina, in December, 1929, and 
is deposited in the Britton Herbarium at the New York Botanical 
Garden. The species has been found at altitudes of 3000 to li200 
meters (according to labels, but see Dr. Asplund's letter quoted 
hereinafter), flowering frcxn December to March and in May, fruit- 
ing in March. Herbarium material has been misidentified and dis- 
tributed as V. microphylla Kunth and it is actually possible that 
these two taxa are not distinct. More intensive study is required 
here. 

In regard to the Asplund $871 collection cited below, whose 
label reads "General Campero, U200, 1921/6/III. 12366", Dr. As- 
plund, in a letter to me dated November 19, 1961, says "The rail- 
way station Jeneral Campero (that is the spelling on the station 
house) is in the province of Pacajes in the Department of La Paz, 
Bolivia. According to my annotations I collected there on March 
6, 1921, only one number of Verbena , viz. Nr. 2800 , which you de- 
termined in July 195U as Verbena microphylla H.B.K. From where 
the number 12366 cones I do not know at all, but the change of 
2800 to 5871 was most probably done by Dr. Otto Buchtien. He 
helped me in drying my plants in the rairiy season during the 
first months of 1921, and as a compensation for his help I al- 
lowed him to take fragments for his ovm herbariim. When he in- 
serted these fragments (which were scmetimes rather large) into 
his herbairium he seems to have regarded them as a continuation of 
his 'Herbarium Bolivianum' and gave them new numbers, continuing 
his own number series. In copying my field labels he seems to 
have been sometimes not very careful. For Nr. 2800 I have anno- 
tated an altitude of 3900 m («= the altitude of the railway station 
house in the bottom of the valley), not U200 m (= approximate al- 
titude of the plateau above the station) ." 

In all, 10 herbarium specimens, including the type, have been 
examined by me. 

Citations: BOLIVIA: La Paz: Asplund 5871 [12366] (W~113U873) • 
Potosi: Flebrig 26I3 (S, S) . ARGENTINA: Jujuy: TV. Meyer 3167 (N), 
s.n. [Herb. Inst. Miguel Lillo %h01] (N) ; Venturi 8777 (Du~ 
372^86, £—962052, N, W~l5911ilU). Salta: Venturi lOOll; (N— type) . 

VERBENA NIVFJl f . ROSEA Moldenke, Phytologia 8: 60. I96I. 

Bibliography: Moldenke, Phytologia 8: 60. 196lj Moldenke, R6- 
suml Suppl. 3: 15 . 1962. 

This form differs from the typical form of the species in hav- 
ing rose-colored corollas. 

The type of the form was collected by Santiago Venturi ( no. 
10111) in waste land between pastures at an altitude of 3300 meters, 
Cumbre del Cajon, dept. Tafl, Tucumin, Argentina, on April 20, 
1926, and is deposited in th : United States National Harbariian at 



192 PHYTOLOGIA Vol. 10, no. 3 

Washington. At the time of its publication, I was under the im- 
pression that the corollas of V. nivea are always white. However, 
some collectors describe them as "bluish-white" or "o ligeramente 
morada", possibly due to fading with age. The present form seems 
to have the flowers always pink in color. It is known thus far, 
however, only frc»n the type specimen. 

Citations: ARGENTINA: Tucumin: Venturi 10111 (W— I591ii28~type). 

xVERBENA NOAGKI Moldenke, Phytologia 2: 11|9. 19U6. 

Synonyiny: Verbena hispida x hastata Dermen, Cjrtologia 7: 170, 
in textu. 1936. Verbena noackii Moldenke in Chittenden, Roy. 
Hort. Soc. Diet. Gard. U: 2211. 19^1. Verbena hastata L. x V. 
hispida Ruiz & Pav. ex Moldenke, R^sum^ 366, in syn. 1959. Verbe- 
na hispida Ruiz & Pav. x V. hastata L. ex Moldenke, R^sumS 366, 
in syn. 1959. 

Bibliography: Dermen, Cytologia ?: 170. 1936; Moldenke, Phyto- 
logia 2: IU9. 19U6j Moldenke, Alph. List Invalid Names Suppl. 1: 
2U. 19i;7; Moldenke, Known Geogr. Distrib. Verbenac, [ed. 2], I6i(. 
& 198. I9ii9; Moldenke in Chittenden, Roy. Hort. Soc. Diet. Gard. 
h: 2211. 1951} Moldenke, Am. Midi. Nat. 59: 356—357. 1958: Mol- 
denke, R6sum6 22U, 366, & U73. 1959; Moldenke, Phytologia 8: 121 
(I96I) and 9: 219 & 296. 1963. 

The name, xV. noacki , was proposed by me in I9U6 for the hy- 
brid produced artificially by Dermen from cultivated material in 
Massachusetts between V. hastata L. and V. hispida RvlUzSc Pav. 
and discussed by him in 1936. The two parental species do not 
normally grow together anjrwhere in the wild — one being temper- 
ate North American and the other being temperate South American. 
It is not expected, therefore, that this hybrid will ever b'j 
found in the wild unless one or the other of the parents if in- 
troduced into the region of the other and becomes acclimat i 
there. The hybrid might, however, have horticultural pos 3il- 
ities. Its characters are intermediate between those of 3 two 
parental species. 

Adding a second "i" to the termination of the second b of 
this bincanial name, and of various other similar binomia in my 
article on Verbena in the Royal Horticultural Society's 1 tion- 
ary of Gardening (1951), seems to have been a policy of i edi- 
tor. It was definitely not due to ar^ intentional act on y 
part, since I do not approve of such so-called and wholly unnec- 
essary and ujidesirable "corrections" of original spellings'. 

xVEKBEWA NOTHA Moldenke, Phytologia 5: 133- 1955- 

Synonymy: Verbena halei x neomexicana Dermen, Cytologia 7: 170. 
1936. Verbena halei Small x V. neomexicana (A. Gray) Small ex 
Moldenke, Resimil 365, in syn. 1959* Verbena neomexicana (A. 
Gray) Small x V. halei Small ex Moldenlce, R6s\un6 370, in syn. 1959. 

Bibliography: Dermen, Cytologia 7: 170. 1936; Moldenke, Phyto- 
logia 3: U67 (I95I) and 5: 133. 1955; Moldenke Biol. Abstr. 30: 
1093. 1956; Moldenke, ResumS 22U, 365, 370, St U73. 1959; Moldenke, 



196U Moldenke, Monograph of Verbena 193 

Phortologia 8: 121 (I96I) and 9: 166. I963. 

The name, xV. notha, was proposed by me in 1955 for the hybrid 
between V. halei Small and V. neomexicana (A. Gray) Small pro- 
duced artificially by Dermen from cultivated material in Massa- 
chusetts in 1936 and discussed by him in that year. The two par- 
ental species grow together in the wild in at least four counties 
of Texas and one county of Arizona, as well as in at least four 
states of Mexico, so it is very possible that this hybrid occurs 
in the wild. It may account for some of the many puzzling "in- 
termediate" specimens hereinbefore cited under V. neomexicana and 
itb two varieties. 

VERBEia OCCULTA Moldenke, Phytologia 3: 280— 201. 1950. 

Bibliography: Moldenke, Phyi>ologia 3: 280—281 & 286 (1950) 
and 3: U85. 1951; E. J. Salisb., Ind. Kew. Suppl. 11: 263. 1953; 
Moldenke, R^sum6 85 & ii73. 1959 j Moldenke, Phytologia 9: 395* 
1963. 

Herb; stems and branches erect or ascending, brunnescent, ob- 
tusely tetragonal, appressed-pilosulous on the older parts, 
short-pubescent with spreading hairs on the younger parts; nodes 
annulate; principal internodes 0.5 — 3 cm. long; leaves decussate- 
opposite, sessile, more or less clasping-based; petioles obsolete; 
leaf -blades deeply 3-parted to about the middle, uniformly dark- 
green on both surfaces, brunnescent in drying, thin-chartaceous, 
very fragile in drying, densely pubescent on both surfaces with 
subappressed hairs of irregular length, the divisions often few- 
toothed or short-lobed, the lobes subacute, somewhat revolute- 
margined, the single vein in each segment flat or obscure above, 
prominulous beneath; inflorescence terminal, spicate, abbreviat- 
ed, 1.5 — k cm. long, densely many- flowered; peduncles obsolete 
or to 2 cm. long, spreading-pubescentj bracts few, foliaceous, 
oblong, to 12 mm. long and 2 mm. wide; bractlets large and con- 
spicuous, surpassing and mostly hiding the calyx, 8 — 10 mm. long, 
lanceolate, attenuate at the apex, rather densely appressed- 
pubescent; calyx about 5 ram. long, irregular, densely pubescent, 
its rim 5-lobed; corolla varying from blue or violet to clear- 
lilac, purple, or rose, hypocrateriform, its tube 8 — 10 mm. long, 
the limb U — 5 nim. wide. 

The type of this curious species was collected by Ram6n Ferrej'- 
ra (no. 1298 ) in a stony habitat, at an altitude of 3200 to 3500 
meters, near Nevado Cajamarquilla, in the province of Bolivar, La 
Libertad, Peru, on September 12, I9U6, and is deposited in the 
Britton Herbarium at the New York Botanical Garden. The species 
has been found in rocky places at altitudes of 2625 ^ UOOO meters, 
blooming in May, June, August, and November. The only common 
name recorded for it is "verbena". 

In all, 10 herbarium specimens, including the type, have been 
examined by me . 

Citations: PERU: Cajamarca: R. Ferreyra 8522 (Z); Scolnik 86? 
(S), I30U (Er, N). La Libertad; ~Angulo l675 (Z); R. Ferreyra 1298 
(N— type); L6pez Miranda IIO8 (W— 217371*5); L6pe2 Miranda & Sag- 



19U PHYTOLOGIA Vol. 10, no. 3 

£stegui 2718 (S)j Ochoa lhh3 (N), lU68 (N) . 

VERBENA OCCULTA f. ALBA Moldenke, Bxill. Torrey Bot. Club 77: U05. 
1950. 

Bibliography: Moldenke, Biill. Torrey Bot. Club 77: U05. 19^0; 
Moldenke, Revist. Sudam. Bot. 8: 167. 1950 j Moldenke, Phytologia 
3: 376 (1950) and U: l;5l. 1953j Moldenke, R^sum^ 85 & U73. 1959; 
Moldenke, Phytologia 9: 395. 1963. 

This form differs fron the typical form of the species in hav- 
ing white corollas. 

The type of the form was collected by Ram6n Ferre^rra ( no. 
6586 ) in a grassy steppe, at an altitude of 3800 to 3900 meters, 
between San Rafael and Cerro de Pasco, in the department of Pas- 
co, Peru, on January 31, 1950, and is deposited in the Britton 
Herbarium at the New York Botanical Garden. The plant has been 
collected at 3U00 to UOOO meters altitude, flowering in April and 
June. Sandeman describes it as a "low-growing herb, the leaves 
very viscous, groTring in full exposure on the mountain-side". His 
specimen is a perfect match for the illustration given by Ruiz & 
Pavon for their V. multifida , which, however, they state came 
from Concepcion, Chile. Their name, furthennore, seems to be on- 
ly a new name for Erinus laciniatus of Linnaeus and so must go 
into the synonymy of V. laciniata (L.) Briq. regardless of the i- 
dentity of the plant which they illustrated. 

In all, 7 herbarium specimens of this form, including the 
type, have been examined by me. 

Citations: PERU: Ancash: Cerrate 302 (N), 3l6 (N) . Junin: As- 
plund 12009 (S, S); Sandeman 103 (K) . Pasco: R. Ferreyra 6586 
(N~type)j Ochoa 333 (N) . 

VERBEJIA OFFICINALIS L., Sp. PI., ed, 1, 20—21. 1753 '' ^ Yj. 2L~ 
ficinalis Auct., 1957, nor Wats., 19U3] . 
Synonyny: Verbena Dorst., Botanicon 292. 15U0. C jnbina 
Dorst., Botanicon 292, in syn. 15U0. Crista gallir a Dorst., 
Botanicon 292, in syn. l5U0. Peristereon Dorst., F anicon 292, 
in syn. 15U0. Verbenaca Dorst., Botanicon 292, in /n. 15U0. 
Verber^aca recta Ruell., Ped. Dioscorid. Anazarb., ^ i. 1, 317. 
I5I49. Verbenaca Matth., Comment. U66 — ii67. 155U. Sacra herba 
Matth., Comment. U66 — U67. l55U. Communis Verbena sive sacra 
procerior recta L'Obel, Plant. Obs. 231. 1576. Verbena vulgo 
appellata Caesalp., De Plantis U50. 1583. Alterum genus, quod 
Mas apud~Plinium intelligltur Caesalp., De Plantis U5l. 1583. 
Verbena recta Dodon., Stirp. Hist. Pemptad. 150. 1583. Communis 
verbena & sacra recta L'Obel, Icon. Stirp. 53li. 1591. Verbena 
communis Gerarde, Herbal 580. 1597. Verbena sacra Gerarde, Her- 
bal 580. 1597. Verbena vulgaris Clus., Rar. Plant. Hist, h: xlv. 
1601. Verbena commiinis caeruleo flore C. Bauh., Pinax Theatr. 
Bot. 269. 1623. Verbena communis & sacra recta L'Obel ex C. 



196U Moldenke, Monograph of Verbena 19$ 

Bauh., Pinax Theatr. Bot., ed. 1, 269, in syn. 1623. Columbaris 
Herm. ex C. Bauh., Pinax Theatr. Bot., ed. 1, 269, in syn. 1623. 
Herba sacra Ang. Tiir. ex C. Bauh., Pinax Theatr. Bot., ed. 1, 269, 
in syn. 1623. Verbena sive verbenaca Guil. ex C. Bauh., Pinax 
Theatr. Bot., ed. 1, 269, in syn. 1623. Hierobotane mas Brunf. 
ex C. Bauh,, Pinax Theatr. Bot., ed. 1, 269, in syn. 1623. Verbe- 
na Gesn. ex C. Bauh., Pinax Theatr. Bot., ed. 1, 269, in syn. 
1^23. Verbena mascula Brunf els ex C. Bauh., Pinax Theatr. Bot., 
ed. 1, 269, in syn. 1623. Verbenaca vulgaris Lug. ex llatth.. 
Opera Oninia Comment. 7U2, in syn, 162)4.. Sacra herba Toum. ex 
Katth., Opera Omnia Comment. 7U2, in syn, l62li. Verbena recta 
Trag. ex Matth., Opera Omnia Comment. 7U2, in syn. l62lt.. Verbena- 
ca recta Dodon. ex tiatth,. Opera Omnia Comment. 7U2, in syn. I62U. 
Verbena communis C, Bauh, ex I'atth., Opera Ctania Comment. 7U2, in 
syn. l62ii. Verbena vulgaris J. Bauh., Hist. Plant. Univ. 3: Ui3 — 
)|)|)l . 16$0, Verbena Matth., Apolog, 21ii. 167U. Verbena communis, 
flore coeruleo C. Bauh. apud P. Herm., Hort. Acad. Lugd.-Bat, Cat. 
619. 1687. Verbena communis flore caeruleo C. Bauh. apud P. Herm,, 
Fl. Lugd.-Bat. $S » I69O. Verbena latifolia lusitanica procerior 
Tourn. apud P. Herm,, Fl. Lugd.-Bat. 55. I69O. Verbena off. 
Eysenkraut Rivin., Introd. Gen. Remherb, [2U] Icon. [56]. I69O, 
Verbena communis, caeruleo flore C. Bauh. apud Tourn., Hist. Pi. 
Paris 309. 1698. ? Verbena chalepensis major vulgaris facie R. 
Moris on. Plant. Hist. Univ. Oxon. 3: "I4O8" [=Ul8]. 1699^ Herba 
sacra R, Morison, Pi. Hist. Univ. Oxon. 3: "li08" [=Ul8], in syn. 
1699. Kierobotana Brunf els ex R, Morison, Plant, Hist, Univ. Ox- 
on. 3: "li08" [=Ul8], in syn. 1699. Verbena cominunis sive vulga- 
ris Clus. ex R. Llorlson, Plant, Hist, Univ. Oxon. 3: "U08»« 
1^13] 6: Ia9, sec. 11, pi. 25, fig. 1. 1699. ? Verbena chalepen - 
sis major , vulgaris facie IJoris . apud Ray, Hist. Plant. 3: Suppl. 
287. 170li. Verbena lusitanica latifolia procerior Tourn. ex Ray, 
Hist. Plant. 3: Suppl. 265. 170U. Verbena latifolia , lusitanica , 
procerior Tourn. apud Boerh., Ind. Plant. Hort. Acad. Lugd., ed. 

I, 75. 1710. Verbena; lusitanica; latifolia; procerior Tourn. 
apud Boerh,, Ind. Alt. Plant. Hort. Acad. Lugd., ed. 2, 1: 187. 
1720. Verbena; camnunis; flore caeruleo C, Bauh. apud Boerh., 
Ind. Alt. Plant, Hort, Acad, Lugd., ed. 2, 1: 187. 1720. Verbena 
foliis multifido-laciniatis, spicis filiformibus L,, Hort, Cliff. 

II. 1737. Verbena communis, flore caeruleo C. Bauh. ex L., Hort, 
Cliff. 11, in syn. 1737. Verbena officinarum Dal. ex L., Hort. 
Cliff. 11, in syn. 1737. Verbena Rivin. ex L., Hort. Cliff. 11, 
in syn. 1737. Verbena humilior foliis incisis Clayt. ex J. F. 
Gronov., Fl. Virg., ed. 1, 8. 1739. Verbena folio variegata 
Breyn ex Haller, Enum. Meth. Stirp. Helvet. 1: 661. 17li2. Verfae- 
na communis coeruleo flore C. Bauh. ex Haller, Enum. Meth. Stirp. 



196 PHYTOLOGIA Vol. 10, no. 3 

Helvet. 1: 661, in syn. 17U2. Verbena foliis laciniatis superi- 
oribus, tripartitis, spicis angustis longissimis Haller, Envua. 
Meth. Stirp. Helvet. 1: 661. 17U2. Verbena Ruell. ex Haller, E- 
num. Meth. Stirp. Helvet. 1: 661, in syn. 17i|2. Verbena Caesalp. 
ex Haller, Enum. Meth. Stirp. Helvet. 1: 661, in syn. 17ii.2. Ver - 
bena Rivin. t, 56 (sola summa planta) Blackw. t, I4I Haller, Enian. 
Meth. Stirp. Helvet. 1: 661, in syn. 17ii2. Verbena ccamniuiis 
caeruleo flore ( Foliis dissectis ) Tourn. apud Michel, Cat. Plant. 
Hort. Caes. Florent. 98. 17U8. Verbena ccmmunis caeruleo flore 
(foliis non , vel parmn dissectis ) Michel, Cat, Plant. Hort. Caes, 
Florent , 98 . 17li8 . Verbena vrticaefolia canadensis , foliis in- 
cisis, flore maiore Tourn, ex Michel, Cat. Plant, Hort. Caes, 
Florent, 98. 17U8. Verbena spuria L., Hort, Upsal. 8. 17li8j Sp. 
Pi., ed. 1, 20. 1753. Verbarta ( tetrandra ), spicis filiformibus , 
foliis multifido-laciniatis L . , Mat , Med . 6 . 1714-9 • Verbena su- 
pina Blackwell, Cur. Herbal 1; pi, Ul. 17^1 • Verbena foliis 
mnltifido laciniatis , spicis filiformibus L, apud Sauvages, Meth. 
Fol. p] . Fl. Monspel, 279, sphalm. 17^1 • Verbena foliis lacini- 
atis, suinmis tripartitis , spicis angustis longissimis Haller ex 
Sauvages, Meth. Fol. PI. Fl, Monspel. 279, in syn. 1751. Verveine 
Sauvages, Meth. Fol. Pi. Fl. Monspel. 279, in syn. 1751, Verbena 
tetrandra , spicis filiformibus paniculatis, foliis multifido- 
laciniatis, caule solitario L,, Sp, PI., ed. 1, 20. 1753, Verbena 
tetrandra , spicis filiformibus , foliis mult if ido-lac iniat is caul - 
ibus numerosis L,, Sp. Pi., ed. 1, 1: 20, 1753, Veronica humili - 
or, foliis incisls Clayt. ex L., Sp. Pi., ed. 1, 1: 20, in syn. 
1753, Verbena ( officinalis ) tetrandra spicis filiformibus panic- 
ulatis, foliis mult if ido-lac iniatis , caule solitario L., Fl. 
Svec, ed. 2, 10. 1755, Verbena tetrandra , spicis filiformibus , 
foliis multifido laciniatis, caulibus numerosis L. apud J. F. 
Gronov., Fl. Virg., ed. 2, U. 1762. Verbena urticae folio cana- 
densis , foliis incisis , flore ma j ore Tourn. apud J. F. Gronov., 
Fl. Virg., ed. 2, h, in syn. 1762. Verbena humilior foliis in- 
cisis J. F. Gronov., Fl. Virg,, ed, 2, h. 1762. Verbena foliis 
tripartitis , rugosis, spicis nudis gracillimis Haller, Hist. 
Stirp. Indig. Helvet. 1: [96]. 1768. Verbena tetrandra spiciis 
filiformibus paniculatis, foliis multifido laciniatis , caule 
solitario L. apud Haller, Hist. Stirp. Indig, Helvet, 1: [96], 
in syn, I768, Verbena foliis vix dissectis Haller, Hist, Stirp, 
Indig, Helvet, 1: [96], 1768, Verbena Rivin, t, 56, summa , ut 
fere solet , plsinta Blakwell t, Ul Haller, Hist, Stijrp, Indig, 
Helvet, 1: [96], in syn, I768, Verbena officinalis tetrandra , 
spicis filiformibus paniculatis , foliis multif ido-lac iniatis , 
caule solitario L, apud Pollich, Hist, Plant, Palat, 1: 22. 1776, 



196U Moldenke, Monograph of Verbena 197 

Verbena officinalis, tetrandra spicis filifonnibus paniculatis, 
foliis multifido-laciniatis, caule solitario L. apud 0. F. Mtlller, 
Icon, Plant, Fl. Dan. U: 5, in syn. 1777. Verbena spicata Gilib., 
Fl. Lithuan. 1: 92. 1735. Verbena foliis multifido-laciniatis ; 
spicis filiformibus L. apud Lour., Fl. Cochinch., ed. 1, 1: 27, 
in syn, 1790. Verbena tetrandra , spicis filiformibus ; foliis 
multifido-laciniatis. Caule solitario L., apud Lour., Fl. Cochin- 
ch., ed. 1, 1; 27, in syn. 1790. Verbena tetrandra; spicis fili- 
formibus, paniculatis J foliis multifido-laciniatis; caule soli- 
tario L. apud Desf ., Fl. Atlant, 1: 16. 1800. Verbena erecta 
sive mas Dodon. apud Desf., Fl. Atlant. 1: 16. 1800. Verbena 
foliis trif)artitis , rugosis; spicis nudis, gracillimis Haller a- 
pud Desf,, Fl. Atlant. 1: 16, in syn. 1800. Verbena tetrandra, 
spicis filiformibus, paniculatis, foliis multifido-laciniatis, 
caule solitario L. apud Poir. in Lam., Encycl, M^th. Bot. 8: 
Shh — 5U5. 1808. Verbena foliis rugosis , tripartitis; spicis nu- 
dis, strigosis Haller apud Poir. in Lam., Encycl. M^th. Bot. 3: 
$U5, in syn. 1808. Verbena erecta , sive mas Dodon. apud Poir. in 
Lam,, Encycl, M6th. Bot. 3: ^h$, in syn. 1808. Communis verbena 
& sacra, recta L'Obel apud Poir. ih Lam., Encj'-cl. M^th. Bot. 8: 
^U5, in syn. 1808. Verbena tetrandra , spicis filiformibus , fol- 
iis multifido-laciniatis , caulibus numerosis L. apud Poir. in 
Lam., Encycl. U^th. Bot. 8: 51i7, in syn, 1808. Verbena spuria , 
caule decumbent e , ramosissimo , divaricate ; foliis ovalibus, in- 
cisis ; spicis filiformibus, bracteis calices superantibus Kichx. 
apud Poir, in Lam., Encycl. M^th. Bot. 8: 5U7, in syn. 1808. 
Verbena humilior , foliis incisis Clayt . apud Poir. in Lam., 
Encycl, M^th. Bot. 3: 5U7, in syn. 1308, Verbena conmunis 
Thornton, Brit, Fl. 1: 27. 1812. Verbena tetrandra , spicis 
filiformibus paniculatis ; foliis multifidis , laciniatis; caule 
solitario L. apud Picot de Lapeyr., Hist. Abrig . Pi. Pyrin. 12, 
in syn. 1813. Verbena vulgaris Waller, New Brit. Domest. Herb. 
1: 30. 1822. Verbena sororia D, Don, Prod. Fl. Nep. lOU. 1825 . 
Verbena officinalis var. a Hook., Bot, ?iisc . 1: 160, 1829, 
Verbena fol. multif .-lacin. , spicis filif . L. apud H. E. Richter, 
Cod, Bot, Linn. 35. 1835. Verbena lusitanica latif . procerior 
To\irn. apud H. E. Richter, Cod. Bot. Linn. 35, in syn. 1835. 
Verbena lus. latif. proc . Tourn. apud H. E, Richter, Cod, Bot, 
Linn. 36, in syn. 1835, Verbena tetrandra , spicis filif ormib. , 
fol. multifido-laciniatis , caulibus numerosis L. apud H. E. Rich- 
ter, Cod. Bot. Linn. 35- 1835. Verbena tetrandra , spicis filif. 
paniculatis , fol. multifido-laciniatis, caule solitario L. apud 
H. E. Richter, Cod. Bot. Linn. 35- 1835. Verbena urticae fol. , 
canadens ., fol. incisis, f 1 . majore Tourn. apud H. E, Richter, 



198 P H I T L G I A Vol. 10, no. 3 

Cod. Bot. Linn. 35, in syn, 1835» Verbena officinsilis fl spuria 
(Willd.) Hook., Comp. Bot. Mag, 1: 176. I836. Verbena officina- 
lis var. spuria Hook., Comp. Bot. Mag. 1: 176. I836. Verbe na 
spuria Willd. ex Hook., Comp. Bot. Mag. 1; 176, in syn. I836, 
Verbena urticae fol. canadensis, fol. incis . Tourn. ex Peterm., 
Cod. Bot. Linn. Ind. Alph. I96, in syn. I8ii0. Verbena communis , 
fl. caerul. C. Bauh. apud Peterm., Cod. Bot. Linn. Ind, Alph. 
196, in syn, I8U0. Verbena lusitanica latif . proc. Tourn. apud 
Peterm., Cod. Bot. Linn. Ind, Alph. I96, in syn, I8U0, Verbena 
officinalis var. natalensis Hochst, ex F, Krauss, Flora 23: 68, 
hyponym, 18U5. Verbena conmxmis flore coeruleo C, Bauh. apud 
Walp., Repert. I4.: 2$, in syn. l8U5. Verbena officinalis Lo\ir. 
apud Walp., Repert. i;: 25, in syn. 18U5« Veronica humilior foliis 
incisus Clayt. apud V/alp., Repert, 1;: 26, in syn. 13U5. Verbena 
officinalis (X vulgaris Schau. in Mart., Fl. Bras. 9: I9I. 1851. 
Verbena officinalis et supina Lap. apud Zetterstedt, PI. Vase, 
Pyren. 220, in syn. 1857. Verbena rumelica Velen., Fl. Bulg. 
km. 1891, Verbena vulgaris Bub., Fl. Pyren, 1: 378, I897. Ver- 
bena officinalis var. genuina Briq., Ann, Conserv, & Jard, Bot, 
Genlv. 10: 105, in nota. I907. Verbena officinale L. ex Molden- 
ke, Suppl. List Invalid Names 9, in syn. I9UI. Verbena officio- 
nalis L. ex Moldenke, Suppl. List Invalid Names 9, in sjn, 19 Ul, 
Verbena officunalis L. ex Moldenke, Suppl. List Invalid Names 9, 
in syn. I9UI, Verbena officinalis var, ramosa L^vl. ex Moldenke, 
Alph. List Invalid Names U9, in syn. 19U2. Verbena dioscoridis 
L'Obel apud Moldenke, Alph. List Invalid Names Suppl. 1: 23, in 
syn. I9U7. Verbena officialis L. ex Moldenke, Alph. List Invalid 
Names Suppl. 1: 25, in syn. 19U7, Verbena officionallis L, ex 
Moldenke, Alph. List Invalid Names Suppl. 1: 26, in syn. 19U7. 
Verbena vulgaris L. ex Moldenke, Alph. List Invalid Names Suppl, 
1: 28, in syn. 19U7. Verbena spuria I*ursh ex Moldenke, R6sum^ 
37U, in syn. 1959. Verbena carolineana Hort. ex Moldenke, R§sum6 
361, in syn. 1959. Verbena spura L. ex Moldenke, R6sum6 Suppl. 
3: Ul, in syn. I962. Verbena oficinalis L., in herb. 

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Miq., Ann. Mus . Bot. Lugd.-Bat. 2: 97. 1865; A. W. Chapm., Fl. 
South. U. S., ed. 1, pr. 2, 307. 1865; Syme, Engl. Bot. 6: pi. 
1018. 1866; Reichenb. f ., Neuholl. Pfl. Am. Dietr. 9. I866; Gri- 
seb.. Cat. PI. Cub. 21h. I866; Ascherson in Schvfeinf., Beitr. Fl. 
Aethiop. 1: 120 & 278. 1867; Cesati, Passer., & Gibelli, Comp. 
Fl. Ital. pi. ii9. 1867—1886; LaMaout ^ Decne., Trait4 G6n. Bot. 
205. 1868: F. Muell., Fragm. 6: 153. I868; R^veil, Regne V5g. 3: 
pi. U2. I87O; Benth. & Muell., Fl. Austral. 5: 35 & 36. I87O; 
Dupuls & R^veil in R^veil, Dupuis, Gerard, S; H^rincq, Regne V§g- 
et. Flor. M^d. 3: li53— U55. 1870; Ulrich, Intemat. WOrterb. Pfl., 
ed. 1, 2U9. 1872} A. V/. Chapn., Fl. South. U. S., ed. 1, pr. 3, 
307. 1872; Pratt, Flow. PI. Sedges Ferns Gr. Brit., ed. 3, h: 
viii & 208—211. pi. 167, fig. 1. 1873; Somoku Dzusetsu, ed. 2, 
11: pi. i;0. 1871; Cusin, Herb. Fl. Fran?. 18 [Verb.]: pi. 1. 
1875; Ulrich, Internat. WOrterb. Pfl., ed. 2, 2^9. 1875; F. 
Muell., Fragm. 9: 5- 1875; Loret L Barrandon, Fl. Montpel. 532. 
1876; Benth. in Benth. l: Hook, f.. Gen. PI. 2 (2): llii6. I876; 
Lesacher & Mareschal, Nouv. Bot. M^d. 1: pi. [8]. I876; C. Berg, 
Anal. Soc. Cientif. Argent. 3: 199. 1877; C. Berg, Enun. PI. 
Europ. 17 & 2U. 1877; J. Ball, Journ. Linn. Soc. Lond. Bot. 16: 
607. 1878; A. Gray, Syn. Fl. N. Am. 2 (1): 335. I878; Pickering, 
Chron. Hist. PI. ii33 & 1220. 1879; RijputAna Gazeteer 30. 1879— 
1880; Karst., Deutsche Fl. Pharm.-medic. Bot. 989. I880~l883; 
C. Hoffm., Pflanzenatlas pi. 50, fig. 309. I88I; Strobl, Flora 
von Admont, Progranme des Melker Gymnasium. 1882; Mttller, Verb. 
Natur. Ver. Preuss. Rheinl. & Westfalens 39 (1): pl. 1. 1882; 
Wagner, 111. Deutsch. Fl., ed. 2, 617. 1882; J. Sm., Diet. Pop. 
Names Pl. U28. 1882; Franchet, Sert. Somal. 50. 1882; F. Muell., 
Syst. Cens. Austr. Pl. 102. 1882; F. M. Bailey, Sjti. Queensl. 
Fl. 376. 1883; H. Friend, Fls. & Flow. Lore 171, 313, 315, 529, 
532—5 3U, 5U5, 5ii7, 5U8, 591, 607, 609, 610, & 665. 1883; s. 
Wats., Proc. Am. Acad. Sci. 18: 135. 1883; Tristram, Surv. West. 
Palest., ed. 1, 378. I88U; F. M. Bailey, Proc. Roy. Soc. Queensl. 
1: 70. I88U; W. Mill., Diet. Eng. Names Pl. lla & 261. I88U: 
Schlecht., Lang., & Schenk, Fl . Deutschl., ed. 5, 19: pl- lo79. 
I88U; A. W. Chapm., Fl. South. U. S.. ed. 2, pr. 1, 307. I88I1; 
F. Muell., Key Victor. Pl. 2: U3. 1835; C. B. Clarke in Hook, f., 
Fl. Brit. Ind. h: 565. 1885; Heuz^ & al., Fl. Pittor. France 221, 
pl. Sh, fig. 321. 1885; Baill., Ic. Fl. Frang. U: pl. 376. 1885— 
I89U; Sterne, Herbs t- & Winterbl. pl. 16, fig. b. 1886; Moloney, 
Sketch Forest. W. Afr. 523. 1887; A. W. Chapm., Fl. South. U. S., 
ed. 2, pr. 2, 307. 1887; J. Ingram Lang, of Fls. 355. 1887; F. 
Muell., Key Victor. Pl. 1: 397. 1837—1889; Sterrart & Corrj', Fl. 
North-east Ireland 285—286. 1888; Tristram, Sxirv. West. Palest., 



202 PHYTOLOGIA Vol. 10, no. 3 

ed. 2, 378. 1888; Batt. & Trab., Fl. Alg. 1: 716—717. 1888; F. 
Muell., Sec. Census 171. 1889; A. W. Chapm.j Fl. South. U. S., 
ed. 2, pr. 3, 307. 1889; Dujard-Beaumetz & Egasse, PI. M^d. 753. 
1889; PotoniS, 111. Fl. Nord- & Mittel-Deutschl., ed. U, fig. 
53U. 1889; ThomS, Fl. Deutschl. U: pi. 531. 1889; Watt, Diet. &- 
conom. Prod. Ind. 6 (U): 229. 1889; Forbes & Hemsl., Fl. Sin. 2; 
252. 1890; F. M. Bailey, Cat. Indig. Nat. PI. Queensl. 35. I89O; 
R. Tate, Handb. Fl. Extratr. South Austr. 155 & 25U. I89O; Pao- 
lucci, Fl. Marchig. pi. 17. I89O; H. C. Hart, Some Acct. Fauna 
Fl. Sinai lOU. I89I; Leclerc du Sablon, Nos Fleurs pl. 3. 1892; 
Lubbock, Seedlings 2: 370. 1892; Coult., Contrib. U. S. Nat. 
Herb. 2: 327. 1892; Baill., Diet. Bot. U: 256. 1892; Engl., Uber 
Hoehgebirgsfl. Trop. Afr. 355—356. 1892; A. W, Chapm., Fl. 
South. U. S., ed. 2, pr. h, 307. 1892; G. Beck, Fl. Nied.- 
Oesterr. 2 (2): 975. 1893; Masclef, Atlas Pl. France pl. 270. 
1893; G. Watt, Diet. Eeonom. Prod. India 6 (U): 229. 1893; 
Moore, Handb. Fl. N. S. Wales 356. 1893; Sehimpfky, Uns. Heilpfl. 
2: pl. U2. 1893— I89U; Acloque, Fl. France 527. I89U; Briq. in 
Engl. & Prantl, Nat. Pflanzenfam. h (3a): lii7 t HiS. I89ii; 
Brehm, Merv. Nat. Monde des Pl. 2: 3U6. 189U~1896; Karst., Fl. 
Deutsch3. 2: 56U. 1895; Gareke, 111. Fl. Deutschl. U89. 1895; 
Mak., Bot. Mag. Tokyo 9: [321]. 1895; Jacks, in Hook, f, & 
Jacks., Ind. Kew. 2: 1179. 1895; Gtlrke in Engl., Pfl. Ost-Afr. 
C: 337. 1895; L. H. Bailey in A. Gray, Field Forest & Gard. 
Bot. 3UO — 3UI. 1895; Si^lain, Atlas Poehe Pl. Champs Prair. & 
Bois., sir. 2, pl. hi. I896; Bub., Fl. Pyren. 1: 378. 1897; A. 
W. Chapm., Fl. South. U. S., ed. 3, 369. 1897; Britton & Br., 
Illustr. Fl., ed. 1, 3: 70. I898; Lijst Volksnamen Genees. Utig. 
Ned. Mij. Phaimacie. I898; J. L. Stewart, Punjab Pl. 166. 1899; 
Journ. Agric. Soc. England, ser. 3, 11 [61]: 738. I9OO; Baker & 
Stapf in Thiselt.-Dyer, Fl. Trop. Afr. 5: 286. I9OO; H. H. W. 
Pearson in Thiselt.-Dyer, Fl. Cap. 5: 208 — 210. 1901; F. M. 
Bailey, Queensl. Fl. U: 1173. 1901; Fiori & Paoletti, Icon, Flor. 
Ital. 380, fig. 3218. 1902; C. T. Green, Fl. Liverpool Distr. 
fig. Ii87. 1902; Chod., Plant. Hassler. 9: 196. 1902; Murr, Deut- 
sch. Bot. Monatschr. 20: 52. 1902; Josef v. Oesterr. & Marg. v. 
Thurn und Taxis, Atlas Heilpfl. pl. I8I. 1903; Losch, KrSCuterb. 
pl. 61. 1903; Sturm, Fl. Deutschl., ed. 2, 11: pl. 22. 1903; 
Voss, Salomon's WOrterb. Deutsch. Pflanzennamen 37 & 251. 1903; 
Thiselt.-Dyer, Ind. Kew. Suppl. 2: 191. 190U; Maiden. Proc. 
Linn. Soc. N. S. Wales [Fl. Norf. Isl.] 29: 712. I90U; Masclef, 
Pl. Eur. pl. 63. 1905; Step, Wayside & Woodl. Bloss., new ed., 
2: pl. 6^. 1905; Burbidge, Book of Scented Gard. 7U & 95. 1905; 
Dtirand & Jacks., Ind. Kew. Suppl. 1: U5l. 1906: F. M. Bailey, 
Weeds & Poison. Pl. Queensl. 139 & lUl, fig. 2U3. 1906; Coste, 
Fl. France 3: Hil. I906; Kirby, Brit. Fl. Pl. pl. 7U. 1906; 
Merino, Fl. Galieia 2: 223. I906; Cheeseman, Man. N. Zeal. Fl. 
1083. 1906; Briq., Ann. Conserv. & Jard. Bot. Genev. 10: 105. 
1907; E. H. L. Krause in Sturm, Fl. Deutschl. 15: 305. 1907; 
Lazaro, Comp. Fl. Espafl. 2: 575. 1907; C. L E. Reid, K. Akad. 
Wetensch. Verb., sec. 2, 13 (6): 18, pl. 2, fig. hi. 1907; J. W. 
Blankinship in Engelm. & Gray, Pl. Lindheim. 3: I86. 1907; M. A. 



196U Moldenke, Monograph of Verbena 203 

Day, Check List 128. I908; Vxnr, Allg. Bot. Zeitschr. lU: 19. 
1908; Robinson & Fern, in A. Gray, New Han. Bot., ed. 7, 688 & 
92U, fig. 878. 1908; Hayek in Engl., Bot. Jahrb. Ii2: I63. 1908; 
Eeille, Bot. Pharm. 2: 1075 & IO76. 1909; Heukels, Fl. Nederl. 3: 
270. 1909; Backer, Ann. Jard. Bot. Buitenz. Suppl. 3 (1): UI9. 
I9IO; Dinand, Taschenb. Heilpfl. pi. Hi, fig. 1. 1910; Fedtsch- 
enko & Fler., Fl. Eur. Russia 796. 1910; Murr, Allg. Bot. Zeit- 
schr. 16: 187. I9IO; Hicken, Chlor. Plat. Argent. 197. 1910; Gor- 
revon, Fl. Champs & Bois pi. 83. 1911; Johns, Fls. of Field, ed. 
C. Elliott, pi. 7U. I9II; Gerth van iYijk, Diet. Plantnanes 1397— 
1398. 1911 ; Nakai, Fl. Korea 2: I3U. 1911; CM. Skinner, l^hs & 
Legends 262. 1911; l^atsum., Ind. 2 (2): 53l|. 1912; Creevey, Har- 
per's Guide 'Jild Fls. 331- 1912; Somoku Dzusetsu, ed. llakino [I- 
con. Pi. Nippon.] 11: pi. UO. 1912; Klein, Unsere Unkrauter pi, 
hO. 1913; Eritton Sc Br., Illustr. Fl., ed. 2, 3: 95, fig. 3552. 
1913: Hauman-Llerck, Anal. LIus. Argent, Hist. Nat. Buenos Aires 
2ii: UlU. 1913; F. M, Bailey, Compreh, Cat, Queensl. PI. 382. 
1913; Gadeceau, Fl. Moiss. 50, fig, 30 — 32, pi, U6. 191ii} Perrin 
L Boulger, Brit, Flow, PI, U: pi. 235 & 2U6 [A], fig, 1—7. 191ii; 
Horvrood, Plant Life Brit, Isles 3: 158, 1915; N. Taylor, Mem, N. 
Y. Bot, Card, 5: [Fl. Vic. N. Y,] 525. 1915; Blanchan, New Nature 
Lib. 6 [Vfild Fls,] UO — Iil, I9I6; Bonnier, Name This Flower pi. 
U5. 1917; C. L, Pollard in Webster's New Intemat. Diet, Eng. 
Lang, 2275 & 2278, 1917; L6v1,, Cat, PI, Yun-Nan 278. 1917; Basu, 
Ind. Medic. Pi. 3: 3, pl. 732b. I9I8; N. L. Britton, Fl. Bermuda 
310. 1918; Lindman, Svensk Fanerogamfl, U61i. I9I8; Holste, Zeit- 
schr, Exper. Patholog, 19: U83. 1918; Sanzin, Anal, Soc, Cient, 
Argent, 88: 96, 12U, & 13ii. 1919; H, J, Lam, Verbenac, Malay. 
Arch. 10 & 369, 1919; Fitch & Sm,, 111, Brit, Fl,, ed. rev. h, 
iss. 2, fig. 822. 1919; Quart. Journ. Geol. Soc. Lond. 76 (2): 157. 
I92O; Bose, Man. Ind, Bot. 252—253. 1920; Blanchan, Wild Fls. 
Woth Knowing 185—136. 1922; Dixon & Fitch, Personal. PI. 150~ 
151. 1923; K. V. 0. Dahlgren, Bot. Notiser Lund 1923: 12. 1923; 
Fitch & Sm., 111. Brit. Fl., ed. 5, fig. 822. 192ii; J. M, Black, 
Fl, South Austr, 3: U73, 1926; Hegi, Illustr. Fl. Mittel-Eur. 5 
(3): 2239— 22U3, pl- 222, fig. 5, 317U, & 3175, pl. 3U6. 1927; 
Bonnier, Fl. Corapl. France Suisse £c Belg. 9: pl. U97. 1927; Domii^ 
Bibl. Bot. 89 (6): IIO6. 1928; Molfino, Fl. Advent. 2: 20. 1923; 
H. F. Roberts, Pl. Hybr, Before Mendel 2U~28 pl. 17 & 18. 
1929; A. B. Seymour, Host Ind. Fungi N. Am. 587. 1929; Herter, 
Florula Urug. 105. 1930; Seckt, Rev, Univ, Nac, Cordoba 17: pl. 
18, fig, U86, I93O; Marzell in Fedde, Repert. Beih, 61: 176, 1930; 
Gier, Trans, Kans. Acad. Sci. 3U: 261. 1931; Eig in Fedde, Repert. 
Beih. 63: 172. 1931; Eig, Zohary, 2: Feinbrunn, Pl. Palest. 305— 
306. 19 31; Soest, Invl. Rivier. Fl. Gelderland 3 £: 6. 1931; Stapf, 
Ind. Lond. 6: U29— U31. 1931; Grieve, Modem Herb. 330. 1931; Ja- 
vorka & Csapody, Ic. Fl. Hiingar, Ul8, 1932; Hegi, Illustr, Fl, 
Mittel-Eur, 5 (3): fig. I319. 1933; Perry, Ann. Mo. Bot. Card. 20: 
2Ul, 2UU, 2li7, 259, 262—263, 265, 267, & 355. 1933; Post, Fl. 
Syria, ed. 2, 2: 321—322 & 925. 1933; Junell, Symb. Bot. Upsal. 
h: 11, 16U, 169—171, & 210. I93U; Crevost & P6telot, Bull. Eco- 
nom. Indo-chine 37: 1287, 193U; L. H. Bailey, Cat. Florists Handl. 



20U PHYTOLOGIA Vol. 10, no. 3 

Verbenac. mss. 1930; Bourdier, Arch. Phaim. 273: 3^7. 1935; 
Reichert, Arch. Pharm. 273: 357. 1935; Dop in Lecomte, Fl. Gin. 
Indo-chine U: 773 & 777—778, fig. 83, U— 7. 1935; L. H. & E. Z. 
Bailey, Hortus, new rev. ed., 631 & 632. 1935; Korsmo, UgressfrO 
pi. 17. 1935; Bedevian, Illustr. Polyglot. Diet. PI. Names 609 — 
610. 1936; Ltrtjeharms & Oostroom, Bltmea 2: 82. 1936; W. Young, 
Trans. & Proc. Bot. Soc. Edinb. 32: llU. 1936; Buchli, Beitr. 
Geobot. Landes. SchwBiz 19: 63. 1936; Dermen, Cytologia 7: I6I — 
I6U, fig. 15. 1936; Gheymol, Verbanaloside . 1937; Herter, Revist. 
Sudam. Bot. U: 186—187. 1937; Perez Arbelaez, Plant. Medic. Co- 
lomb. 2UO. 1937; A. A. & K. H. Rechinger in Fedde, Repert. Beih. 
C: 131. 1938; K. V. D. Dahlgren, Svensk Bot. Tidsk. 32: 231. 1938; 
Gallinal, Bergalli, Campal, Aragone, & Rosengurtt, Stud. Nat. 
Mead. Urug. 95. 1938; Gongalves da Cunha & Gon<jalves Sobrinho, 
Revist. Fac. Cienciaa 3: 5. 1938; Oppenheimer, Palest. Journ, 
Bot., ser. R, 2: 37. 1938; I. C. Verdoorn, Union S. Afr. Dept. 
Agr. & Forest. Bull. 185: h6 & 171, fig. 90. 1938; Moldenke, An- 
not. & Classif . List 108. 1939; F. C. Hoehne, Plant, Subst. T6x. 
& Med. 2U9 &: 250, fig. 201; II. 1939; Moldenke, Alph. List Coimon 
Names 2—);, 8—17, 19, 21, 23—26, & 28— 3U. 1939; Kuvrajima, To- 
hoku Joum. Exp. Med. 36: 28 — U3. 1939; Moldenke, Carnegie Inst. 
Wash. Publ. 522: lh9* 19iiO; C. C. Beam, Fl. Indiana 795, 108ii, & 
1232. 19l;0; Moldenke, Suppl. List Common Names 5, 7, 9, 10, 17, & 
20. 19 UO; Oppenheimer & Evenari, Bull. Soc. Bot. Genev. [Fl. 
Cisiord.], sir. 2, 31: 90 & 363. 19i;0; Ranson, Texas Wild Fl. 
Legends, ed. 2, 113. I9UO; Gonial ves da Cunha & Gonial ves Sobrin- 
ho. Bol. Soc. Brot., ser. 2, lit: 11. I9UO: Fern., Rhodora li2: 
U7o. I9UO; Robledo, Lecc. Bot. 2: U97. 19U0; Moldenke, Prelim, 
Alph. List Invalid Names U5 — U8. 19i4.0; Hutchinson & Bruce in 
Gillet, Kew Bvill. I9UI: 177. 19Ul; Pool, Fls. & Flow. PI., ed. 
2, 107, fig. 72c. I9UI; Moldenke, Suppl. List Invalid Names 9. 
I9UI; Telles Palhinha, Gon^alves da Cunha, gc Gon^alves Sobrinho, 
Contrib. Conhec. Fl. Agores 52. I9UI; Biswas, Ind, Forest Rec, 
Bot,, new ser., 3: ii2. 19iil; Worsdell, Ind. Lond. Suppl. 2: U86. 
I9UI; Schnack, Anal. Inst. Fitotlc. Sta. Catalina U: 13 — 20. 
19U2; H. N. Webster, Herbs 27, 31, & 32. 19U2j Moldenke in Lun- 
dell, Fl. Texas 3 (1): 21, 23, & 30. 19U2; Moldenke, Known Ge- 
ogr. Distrib. Verbenac, [ed. 1], 2—5, 7, 10, 15, 19, 21;, UO, 1;2, 
lUi— 116, U9, 50, ^3, $$, 57—61, 63, 7U, 75, & 101. 19U2; Moldenke, 
Alph. List Invalid Names U6, U9, & 50, I9I4.2; Breitwieser, Pharma- 
kogn. Untersuch. Verbenac. I9l;2; Webster's College Diet., ed. 5, 
1115. 19li3; Yuncker, Bishop Mus. Bull. 178: 101. 19U3; A. R. 
Sehultz. Introd. Estud. Bot, Sistem., ed. 2, U76, ii83, & 562, fig, 
175. 19U3; R. S. Woods, Naturalists' Lex. 271. 19Uli; E. L. D. 
Seymour, New Gard, Encyel, 1279. 19UU; Schnack & Covas, Darwinia- 
na 6: U70, U72, fig. 1 E— H, 1+73, fig. 2 A, & hlk, fig. 3 C. I9hhi 
Rozeira, Fl. Prov. Tras. & Alto Douro I6U. I9I1U; Darlington & 3aR- 
aki Aramal, Chromosome Atl. 270, 19U5; Mxirrill, Guide Fla, PI, UC 
19U5; Covas & Schnack, Darwiniana 7: 86. 19U5; Roig y Mesa, Plant, 
Medic. Cuba 670—671 & 811. 19U5; Moldenke, Phytologia 2: 7h, 76, 
78—80, 85, & 115—116. 19l;5; Moldenke, Bot. Gaz. IO6: I63. 19U5; 
Moldenke, Castanea 10: 36 & U2. 19U5; Moldenke, Torreya U5: 90. 



196U Moldenke, L'onocraph of Verbena 205 

Koldenke, Am. Journ.^Bot. 32: 611. 19h5; Schnacl: L Covas, Bol. 
Soc. Argent. 3ot. 1: 28U. 19U6j Rosangurtt, Ilstud, Prad. Hat. Ur- 
ug. 5 [Fl. J. Jackson]: 39U. 19ii6j Iloldenlre, Alph. List Cit. 1: 
10, 18, 19, 33, 31^, 38, 59, 71, 77, 30, 100, 105, 115, 120, 122, 
121;, liiO, I5ii, 169, 181, 196, 199, 205, 22U, 226, 236, 2hO, 2U5, 
2U9— 251, 262, 268—270, 27U, 231, 288, L 292—295. 19U6; R. R. 
Tatnall, Fl. Del. 217. 19U7; P'ei, Bot. Btil. Acad. Sin. 1: 2—3. 
19l;7j Kasapligil, Kuzey Anadol. Botanik Gezileri 7U, 122, Pz l55. 
19U7; Daniel, Verb. Gent. Antioq. 2. 19ii7j Lloldenke, Alph. List 
Invalid Names Suppl. 1: 22, 23, 25, 26, t 23. 19U7; Hill £i Sal- 
isb., Ind. Kew. Suppl. 10: 2U2. 19li7; Ik:oldonl:e, Phytologia 2: 330, 
339~3U3, 3U5, 3U8, & 332 (19U7) and 2: U78, U79, & U32. 19U3; 
Moldenke, Gastanea 13: 111, 112, 2.: 119—121. 19i;3; J. Hutchinson, 
Brit. Flow. PI. 129. 19U8; Hager in Reichert, Trat. Famac . 
Prict. 5: 1211i. 19U8; A. £; D. Love, Ghromos. Ilumb. North. Pi. 3p. 
88. 19ii8; Hara, Enum. Sperm. Jap. 1: 190. 1918; Hlosson, K. I. 
Herald Trib., sect. 5, P. lU, July 25. 19^3; Cabrera cz Fabris, 
Publ. Teen. 5 (2): [Pi. Acuat. Frov. Buenos Aires] 13- 19.'i3; 
Hausman, Begin. Guide Y/ild Fls . 262 "^ 375- 19U8; H. N. «c A. L. 
Moldenke, Pi. Life 2: 39, Ul, h2, F: UU. 19U8; Moldenke, Alph. 
List Cit. 2: 356, 357, 360, 362, 369, 391, 396, 1^02, U05, Uo6 
U09, UlO, hl3, UlU, h27, U55, U57, U62, U67, U71, h73, i;73, UoO, 
U90, U93, U95— )497, 506, 5il, 519, 520, 52Ji, 526, 523, 531, 550, 
555, 558, 560, 562, 572, 537, 600, 60U, 605, 617, 6l8, 621, 63O, 
631, 635, 6l;l, 6Ui, & 6h5 (19U3), 3: 653—660, 663, 667, 668, 671, 
698, 699, 708, 709, 712, 717—719, 72U, 727, 729, 733, 7Ul, 7U5, 
7h7, 750—752, 766, 769, 771, 772, 77U— 776, 773, 733, 787, SOO, 
302, 303, 807, 313, 823, 827, 332, 3Ul, 81;2, 8U8, 857, 365, 367, 
83U, 336, 837, 395, 900, OOh, 92U, 927, 932, 936, 939, 9h2, 9U3, 
9li5, 9U6, 96U, 968, 973, 977, ^. 978 (19U9), and h: 930, 932—936, 
993, 995, 993, 1002— lOOU, 1010, 1012, 1013, 1071, 1030, IO83— 
1085, 1093, 1095, 1097, 1103, noli, 1110, 1116, 1120, 1123, 1123, 
1129, 11 3h, 1137, 1139, 1116, 1152, II5I1, lluU, 1167— U69, 
1173—1130, 1192, 1200, 1202, 1203, 1205, 1207—1211, 1213, 12lU, 
1219—1221, 1225, 1227, 1223, 1230, 1232, 1236, 1233, 12U3, 12U7, 
12U3, 1251, 125U, 1260, 1261, 1238, I289, 129U, 1296, L 1297. 
I9U9; H. ::. ■: A. L. Moldenke, Anal. Inst. Biol. Mex. 20: lU. 19^9; 
Cabrera, Lilloa 20: cuadro VIII. 19U9; i.oldenke, Knovm Geogr. Dis- 
trib. Verbenac, [ed. 2], 3—3, 10, 11, lU, 19, 25, 27, 33, U, 
98, 101, 107—111, 115—119, 122, 121;, 125, 123, 129, 132—135, 
137, 133, lUO, Jli2, 152, 15U, 16U, 166, L 199. 19i;9; Moldenke, 
Phytologia 3: 136, 133, ^ 139 (19U9) and 3: 283, 291, 295, S: 380. 
I95O; Molden]-:e, Revist. Sudan. Bot. 8: 173. 1950 j Fern, in A. 
Gray, >an. bot., ed. 8, 1209, 1210, ■■ 1612, fir. II16I. 1950; Con- 
nor =-: Adams, Poison. PI. N. Zeal. 106 & 135. 1951 J Ahles, Bull. 
Torrey Bot. Club 73: 269. 1951; Acevedo de Vargas, Bol. Mus . Uac . 
Hist."^Nat. Chile 25: 70. 1951; Moldenke in Chittenden, Roy. Hort. 
Soc. Diet. Gard. U: 2208, 2209, & 2211. 1951; Moldenke, Inform. 
Mold. Set U6 Spec. 1;. 1951; Moldenke, Sp. Subsp. Cont. Mold. Set 
U5 [3]. I95I; Moldenke, Phytologia 3: li63 & U67 (1951) and U: 66 
& 71. 1952; Moldenke in Gleason, New Britton & Br. Illustr. Fl., 
pr. 1, 3: 126—128. 1952; Lid, Norsk Fl. 529, 530, & 771, fig. 



206 PHTTOLOGIA Vol. 10, no. 3 

303a. 1952 J Roig y Mesa, Dice. Bot. 1: 909 (19^3) and 2: llli;. 
19$3i Moldenke, Phytologia k: k^2 (1953) and 5: 97. 195U; Molden- 
ke, joum. Calif. Hort. Soc. 15: 82. 195U} Moldenke, Infoim. Mold. 
Set U8 Spec. [U]. 195U} Thome, Am. Midi. Nat. 52: 313. 195U} R. 
Mohr, Deutsch. Med. Wochenschr. 79: 1513. 195U; Gohn, Vis, & Kar- 
rer, Helv. Chim. Act. 37: 790. l95Ui Potztal, Drogist. Lex. 1: 
153. 1955; Hocking, Diet. Terms Pharmacog. 77, 117, 171, & 2l;0. 
1955j Univ. Hull List Seeds 7. 1955} Moldenke, Phytologia 5: 133. 
1955; Kara, Joum. Fac . Sci. Univ. Tokyo, ser. 3, 6: 366. 1956; 
Kasapligil, PI. Jordan 102. 1956; Moldenke, Inform. Mold. Set 51 
Spec. h. 1956; Moldenke, Biol. Abstr. 30: 1093. 1956; Alain in 
Le6n & Alain, Fl. Cuba U: 281. 1957: Braun-Blanquet & Bol6s, Anal. 
Estac. Exper. Aiila Dei 5: Ul, 62, 68, 109, 139, & 202, pi. 8 & 25. 
1957; Rothmaler, Exkursionsfl. Deutschl. 373 & 501, fig. 3731. 
1958; Univ. Hull List Seeds 7. 1958; Strausbaugh & Core, Fl. W. 
Va. 3: 77U & 775. 1958; Dandy, List Brit. Vase. PI. 101 & 176. 
1958; Moldenke, Am. Midi. Nat. 59: 3Ul, 3li2, 3kh, 3h7 , 35U, 356, 
358—361, 363, & 361i. 1958; Moldenke in Gleason, New Britton & 
Br. Illustr. Fl., pr. 2, 3: 126 — 128. 1958; Rothmaler, Exkursions- 
fl. Deutschl. Atl. Ii30 & 566, fig. 19U3. 1959; Gaster, Frazer's 
New Golden Bough 622. 1959; Reitz, Sellowia 11: 79. 1959; Molden- 
ke, ResuraS 6—11 13, Hi, 18, 23, 31—33, 39, ii9, 115, 122, 128— 
136, lii2, ll;3, lil5, 1U6, IU9, 15U, 158—160, 16U, 166, 167, 171— 
17U, 177, 178, 182, 135, 202, 207, 210, 22U, 275, 298, 357, 361— 
363, 369, 371, 37U, 378, 379, li21, 123, h2$—\x21, L U73. 1959; 
Moldenke, R^sun^ Suppl. 1: 7, 3, & 11 (1959) and 2: 8 & 10—13. 
I96OJ "lilnde L HtCnsel, Arch. Pham. 293: 556—567. I960; Winde, 
Echaust, L Hansel, Arch. Pham. 29V^^J 220—229. 1961; P. Jaeger, 
Wonderf . Life of Fls. 12U. 1961; Lev;is L Oliv., Am. Joum. Bot. 
U8: 6UI. 1961 ; Steam, Gat. Bot. Books Coll. Hunt 2: cxiv & 393. 
I96I: Moldenke, Phytologia 8: 119—122, 12i;, L m3— IU6 (I96I) 
and 8: I9U, 195, 206, 257, 279, 280, 310, 313, 316, 317, 373, & 
U35. 1962j Cuf., Bull. Jard. Bot. Brux. 32: Suppl. 787—788. 
1962; Zohary, Pi. Life Palest. I6U. 1962; Nair & Rehnan, Bull. 
Nat. Bot. Gard. Lucknow 76: 3 £: 5, fig. 3. 1962; Moldenke, R^sum4 
Suppl. 3: 1. 2, 16—19, ^.•. iil (I962), h: 1, 2, L U— 21 (1962), $: 
1, 2, & 6—3 (1962), and 6: 7, 3, ^t 11. I963; Gleason ?.• Cronquist, 
Man. Vase. PI. 579 & 580. I963: Coon, Using Pl. for Heal. 220. 
1963; Moldenke, Phytologia 8: U62, ii72, U77, ?: U33 (I963) and 9: 
3U, 36, 33, 39, lS—1^, 93, 1U8— 151, I60, 162, 165—167, 175, 
181, 190, 21>-215, 219—221, 293, & 296. 1963; R. Paris in T. 
Svjain^ Chem. Pl. Tax. 356. 1963; Moldenke, R^siun^ Suppl. 8: 6. 
196ii; Vilmorin-Andrieux & Cie. s'eed packet U9659. n.d. 

Illustrations: Dorsten, Botanicon 292. l5U0; Ruell., Ped. 
Dioscorid. Anazarb., ed. 1, 317. l5U9; Matth., Comment. Ii67. 155U; 
Tvlatth., Disc. Ped. Dioscor. Anazarb. 1107. 1568; Dodcn., Stirp. 
Hist. Pemptad. 150. 1583; L'Obel, Icon. Stirp. 53U. 1591; Gerarde, 
Herbal 580, fig. 1. 1597; L'F.cluse, Rar. Plant. Hist. XLV. I6OI; 
Matth., Opera Omnia Comment. 7U2. 1621;; J. Bauh., Hist. Plant. 
Univ. 3: UU3. 1650; Rivin., Introd. Gen. Remherb. Icon. [56]. I69O; 
R. Morison, Plant. Hist. Univ. Oxon. 3: sec. 11, pl. 25, fig. 1- 
1699; Zannich., Is tor. piante Venez. pl. 269. 1735; Blackvrell, Cur. 



I96I4 lloldcnke, I'onograph of Verbena 20? 

Herbal 1: pi. U. [in color]. 17^1; Kniphof, Herb. Viv. 2: pi. 
1156 [in color]. 176ii; Curtis, Fl. Lond., ed. 1, 1 (1): pi. Ul 
[in color]. 1772} Regnault, Eot. 1: 112 [in color]. 177U; 0. F. 
Ittller, Icon, plant. Fl. Dan. h: pl. 628 [in color]. 1775} Sabbat 
in L'artelli, Hort. Roman. 3: pl. 56 [in color]. 1775} Vfalcott, 
Fl. Erit. Indie, pl. [92]. 1775; Zorn, Ic. Pl. Med. 1: pl. 38 
[in color]. 1779; Bulliard, Herb. France 3 ^ U: pl. 215 [in col- 
or]. 1730—1793; Plenck, Ic. Pl. lied. 1: pl. 17 [in color]. 1788; 
Ho-^pe, Zct. Pl. P.atisb. pl. 332. 1789; Lam., Illustr. 1: pl. 17, 
fir. 1. 17^1; 0. F. Hoffm., Deutschl. Fl. pl. 2. 1791; C. K. 
Spren§:., Sntd. aeheimn. Natur pl. 1, fig. Ih — 16 & 21. 1793: 
.Voodville, Ted. Eot. Suppl., ed. 1, pl. 218 [in color]. 179U; 
Droves '■ V.ayne, Pot. Bilderbuch 2: pl. hS [in color]. 1795; Os- 
kar.p, Afb. Artscny— :"revnss. 1: pl. 38 [in color]. 1796; Sturm, 
Deutsclil. Fl. 1 (3): pl. [3U] [in color]. 1797; Kops, Fl. Eatava 
1: pl. 31 [in color]. 1300; Sowerby S, Sm., Engl. Dot. 11: pl. 767 
[in color]. 1300; Dreves ^ Ha:me, Choix de Pl. pl. U5 [in color]. 
1302; Svensk lot. 3: pl. 182 [in color]. ISOU; G. Savi, tot. Ued. 
Vog. Toscana pl. 52- i305; P.oques, Pl. Lsuel. Indig. f,-. Exot. 2: 
r-1. 59 [in color]. 1C03; Schlnihr, Dot. !!andb., ed. 2, pl. 3 [in 
color]. 1303; Jaune St.-':il., Pl. France h: pl. [39] [in color]. 
1309; A. &. F. ?l6e Ilerbor. P-j-is pl. [27] [in color]. l3ll; Pdll, 
r.inily Ilorbal pl. u9 [in color]. I3l?; Vicneux, Fl. Pitt. Envir. 
Paris pl. 1 [in color]. I3l2; Thornton, Erit. Fl. 1: 27. I8l2; 
Scl-iranlc, Fl. Lonac. 2: pl. 135 [in color]. I3ll+; Curtis, Fl. 
Lond., CO. 2, 1 (1): pl. Iil [in color]. I3l7; Ilayne, C-etreue Dar- 
st. Arzn. Ge-zfachse >: pl. u2 [in color]. 1317; Chauneton, Fl. Med. 
6: pl. 3U6 [in color]. I3l3; Vietz, Ic. Pl. L'ed.-oec.-techn. Suppl. 
pl. 93 [in color]. 1322; Waller, 'lev,' Brit. Domest. Herb. 1: 30. 
1322; sweet, Br::t. Flo.v. Gard. 3: pl. 202 [in color]. 1327; '.Vood- 
ville, Ked. Eot. Suppl., ed. 3, 2: pl. 133 [in color]. 1332; Bax- 
ter, Brit. Eot., ed. 2, 1: pl. 26 [in color]. 133U; Dietr., Fl. 
Boruss. 2: pl. 120 [in color]. I33li; Heger, Danske Cec. Pl. 3: pl. 
210 [in color]. 1335; Zenker, Fl. Thtfringen 3: ?!• 321 [in color]. 
1836 — 1855; Soenn. in Nees von Esenb., Gen. Plant. Gem. f asc . 21: 
20 (U20). 1339; r. E. Jacks., Pict. Fl. fig. 363. 13U0; Mees von 
i;senb.. Gen. Pl. Ganop. 2: pl. 50. IjUI; Pl6e, lypes Far.. Pl. 
France pl. 126 [in color]. l.iUIi — 136U; Deakin, Florigr. Brit. 2: 
fig. ICOli [in color]. 13U5; Duchesne, Pl. Util. pl. 56. I3ii6; J. 
'.". Loud., Brit. ".7ild Fls., ed. 2, pl. U9 [in color]. 13U6; Lorek, 
Fl. Pruss., ed. 3, pl. lU7, fig. 731 [in color]. I3h3; Petermann, 
Deutschl. Fl. pl. 72, fig. 565 [in color]. l8ii9; Honigberger, 
Thirty-five Years Fast 2: pl. 29. 1352; J. C. "Jeber, Fl. Bayern 5: 
pl. Ij83 [in color]. 1853; Reichenb., Icon. Fl. Germ. I8: pl. 1292, 
II [in color]. 1357; Irvine, Illustr. Handb. Brit. Pl. 21 & Ul5. 
1858; Johnson -!; So-;ferby, Brit. '.Yild Fls. fig. 98I. 1353—1360; 
Berg, Charact. pl. 36, fig. 232. I86I; T. L'oore, Field Dot. Conp. 
pl. 17 [in color]. 1862; Hogg Fc Johnson, 'wild Fls. Gr. Brit. 1: 
pl. 59 [in color]. 1863; Argenta, Alb. Fl. Led. -Farm. 3: pl. 238 
[in color]. I86U; Benth., Handb. Brit. Fl. fig. 8II;. 1865; Schu- 
bert, Naturg. Pflanzenreich, ed. 2, pl. 2, fig. 7 [in color]. 
1865; Syme, Eng. Bet, 6: pl. 1018 [in color]. 1866; Cesati, Passer., 



208 PHYTOLOGIA Vol. 10, no. 3 

& Gibelli, Comp. Fl. Ital. pi. U9. 1867—1886; LeMaout £c Decne., 
Traits Gin. Bot. 205 . 1868; R^veil, Regne Y6g. 3: pl. Ii2 [in col- 
or]. 1870} Pratt, Fl. Pl. Grasses Sedges & Ferns Gr. Brit., ed. 
3, ii: pl. 167, fig. 1 [in color]. 1873; Somoku Dzusetsu, ed. 2, 
11; pl. ho [in color]. 187U; Cusin, Herb. Fl. Fran?. 13 (Verb.): 
pl. 1 [in color]. I875j Lesacher & Mareschal, Nouv. Bot. M6d. 1; 
pl. [8] [in color]. 1876; Karst,, Deutsche Fl. Pharm. -medic . Bot. 
989. 1880—1883; C. Hoffm., Pflanzenatlas pl. $0, fig. 309. I88I; 
Mtlller, Verb. Natur. Ver. Preuss. Rheinl. & Westfalens 39 (1): 
pl. 1. 1882; Wagner, 111. Deutsch. Fl., ed. 2, 617. 1882; Schl- 
echt., Lang., & Schenk, Fl. Deutschl., ed. 5, 19: pl. 1879 [in 
color]. 1881;; HeuzS & al., Fl. Pittor. France 221, pl. 5U, fig. 
321 [in color]. 1885; Baill., Ic. Fl. Frang. U: pl. 376 [in col- 
or]. 1885~189U; Sterne, Herbst-/; Winterbl. pl. 16, fig. b [in 
color]. 1886; Dujard-Beaumetz & Egasse, Pl. Mid. 753. I889; Poto- 
nil. 111. Fl. Nord- 4 Mittel-Deutschl . , ed. h, fig. 53ii. 1889; 
Thoal, Fl. Deutschl. U: pl. 531 [in color]. 1889; Paolucci, Fl. 
Marchig. pl. 17. I89O; Leclerc du Sablon, Nos Fleurs pl. 3 [in 
color]. 1892; Lubbock, Seedlings 2: 370. 1892; Baill., Diet. Bot. 
h: 256. 1892; G. Beck, Fl. Nied.-Oesterr. 2 (2): 975. 1893; Mas- 
clef, Atlas Pl. France pl. 270 [in color]. 1893; Schimpfky, Uns. 
Heilpfl. 2: pl. U2 [in color]. 1893— I89U; Acloque, Fl. France 
527. I89h; Briq. in Engl. & Prantl, Nat. Pflanzenfam. h (3a): lii7. 
I89ii; Brehm, Merv. Nat. Monde des Pl. 2: 3U6. I89U— 1896; Garcke, 
111. Fl. Deutschl. U89. 1895; Karst., Fl. Deutschl. 2: 56U. I895j 
Sillain, Atlas Poche Pl. Champs Prair. & Bois., sir. 2, pl. ii7 
[in color]. I896; Britton & Br., Illustr. Fl., ed. 1, 3: 70, fig. 
3057. 1898; Joum. Agric. Soc. England, ser. 3, 11 [6J] : 738. 
I9OO; Fiori & Paoletti, Icon. Flor. Ital. fig. 3218. 1902; C. T. 
Green, Fl. Liverpool Distr, fig. U87. 1902; Josef v. Oesterreich 
& Marg. V. Thum & Taxis, Atlas Heilpfl. pl. I8I [in color]. 
1903; Losch, Krauterb. pl. 61 [in color]. 1903; St\mn, Fl. Deut- 
schl., ed. 2, 11: pl. 22 [in color]. 1903; Masclef, Pl. Eur. pl. 
63 [in color]. 1905; Step, Wayside & Woodl. Bloss., new ed., 2: 
pl. 65 [in color]. 1905; F. M. Bailey, Weeds & Poison. Pl. 
Queensl, 139. 1906; Coste, Fl. France 3: lUl. 1906; Kirby, Brit. 
Flow. Pl. pl. 7k [in color]. I906; Merino, Fl. Galicia 2: 223. 
1906; Lazaro, Comp. Fl. Espafl. 2: 575. 1907; C. & E. M. Reid, K. 
Akad. Wetensch. Verb., sec. 2, 13 (6): 18, pl. 2, fig. hi. 1907; 
Beille, Bot. Pharm. 2: 1075 & 1076. 1909; Heukels, Fl. Nederl. 
3: 270. 1909; Dinand, Taschenb. Heilpfl. pl. li;, fig. 1 [in col- 
or]. I9IO; Fedtschenko & Fler., Fl. Eur. Russia 796. 1910; Cor- 
revon, Fl. Champs & Bois pl. 83 [in color]. I9II; Johns, Fls. of 
Field, ed. C. Elliott, pl. 7ii [in color]. 1911; Creevey, Harper's 
Guide Wild Fls. 331 [in color]. 1912; Somoku Dzusetsu, ed. Makino 
[Icon. Pl. Nippon.] 11: pl. kO. 1912; Britton & Br., Illustr. Fl., 
ed. 2. 3: fig. 3552. 1913; Klein, Unsere UnlcrSCuter pl. i;6 [in 
color]. 1913; Gadeceau, Fl. Moiss. 50, fig. 30 — 32, pl. U6 [in 
color]. I9IU; Perrin & Boiager. Brit. Flow. Pl. U: pl. 235 & 21x6 
[A], fig. 1 — 7 [in color]. 19lli; Horwood, Plant Life Brit. Isles 
3: 158. 1915; Bonnier, Name This Flower pl. U5 [in color]. 1917; 
Basu, Ind. Med. Pl. pl. 732. 1918; Britton, Fl. Bermuda 310. 



196U Moldenke, Monograph of Verbena 209 

1918; Lindman, Svensk Fanerogainf 1 . U6U. 1918} Fitch & Sm., 111. 
Brit. Fl., ed. rev. U, iss. 2, fig. 822 (1919) and ed. 5, fig. 
822. 1921;; Hegi, Illustr. Fl. Mittel-Eur. 5 (3): pl. 222, fig. 5, 
317U, & 3175 [in color]. 1927; Bonnier, Fl. Compl. France, Suisse, 
& Belg. 9: pl. U97 [in color]. 1927; Javorka & Csapody. Ic . Fl. 
Iftingar. UI8. 1932; Hegi, Illustr. Fl. Mittel-Eur. 5 (3): fig. 
1319. 1933; Korsmo, Ugressfrt5 pl. 17 [in color]. 1935; Dop in Le- 
conte, Fl. G^n. Indo-chine U: 773, fig. 83, U— 7. 1935; Dermen, 
Cytologia 7: 163, fig. 15. 1936; Bedevian, Illustr. Polyglott. 
Diet. Pl. Name 6IO. 1936; F. C. Hoehne, Plant. Subst. Veg. Tox. 
fig. 20ii II. 1939; H. N. & A. L. Moldenke, Pl. Life 2: Ul, pl. 7. 
I9US; Blossom, N. Y. Herald Trib., sect. 5, p. li;, July 25. 19U8; 
Hausman, Begin. Guide Wild Fls. 262. 19U8; Moldenke in Gleason, 
New Britton h Br. Illustr. Fl., pr. 1, 3: 128. 1952; Le6n & Alain, 
Fl. Cuba I4: 281. 1957; Moldenke in Gleason, New Britton & Br. Ill- 
ustr. Fl., pr. 2, 3: 128. 1958; Strausbaugh & Core, Fl. W. Va. 3: 
775. 1958; Rothnaler, Exkursionsfl. Atlas fig. 19U3. 1959; Nair & 
Rehnan, Bull. Nat. Bot. Gard. Lucknovc 76: 3, fig. 3. 1962; Brit. 
Mus. (Nat. Hist.) Brit. Flovf. Pl. card F.80 [in color], n.d. 

Annual or perennial herb, 0.1 — 2 m, tall, varying from erect 
to spreading or even low sind sprawling, scented, often weedy, 
sometimes semi-woodj'^, diffusely much-branched; stans 1 or several, 
ascending or spreading to erect, slender, mostly herbaceous, 
somewhat tetragonal and margined, branched above, striate, glab- 
rous or almost so except on the muriculate or scab rid angles; 
leaves decussate-opposite, sessile or subsessile or sometimes 
neirrowed into a short winged petiole, sheathing at the base; 
leaf -blades 1.5 — 10 cm. long, 0.7 — 2 cm, wide, the basal and low- 
er ones more or less ovate, ovate-oblong, oblong, or oblong- 
lanceolate to lanceolate or oblanceolate or even rhomboid-ovate 
in general outline, varying from tripartite-lac iniate or deeply 
incised to more or less deeply trifid, 5-cleft, or even 1- or 2- 
pinnatifid, narrowed toward the base or cuneate into a petiole, 
the upper ones similar but smaller and less divided, the upper- 
most entire, the various segments, lobes, or laciniations acute 
or obtuse at the apex, often again incised or crenate-serrate, 
strigose-scabrous or roughly pubescent to appressed puberulent or 
even subglabrous or glabrescent on both surfaces, sometimes 
sparsely strigillose on the larger venation only beneath; larger 
veins impressed above, prominulous beneath; inflorescence compos- 
ed of long, slender, axillary and terminal spikes ("racemes" ac- 
cording to Verdoorn), 15 — 50 cm. long, forming a terminal much- 
branched panicle, wide and more or less leafy below; spikes few, 
very lax, slender or filiform, l5 — 23 cm. long, strict, open and 
remotely flowered or compact only during anthesis, greatly elon- 
gating after anthesis, paniculate or subpaniculate, bracteate; 
peduncles glabrous, smooth on the flat surfaces, roughened on 
the margins for their whole length; rachis hirtellous and glandu- 
lar, the branches slender; bractlets ovate or lanceolate, not o- 
ver 2 mm. long, shorter than (or the lovfer as long as) the calyx, 
acute or acuminate at the apex, pubescent or hirtellous beneath, 
glabrous above, glandular, with the midrib very prominent beneath 



210 PHYTOLOGIA Vol. 10, no, 3 

(i.e., on the back); flowers numerous, fragrant, small, loosely- 
arranged on the narrow spikes, distant below, crowded above, usu- 
ally only k OT S blooming simultaneously on one spike, about 12 
mm. long; calyx cylindric, about 2 mm. long, glabrous within, 
hirtellous or hispid on the ribs outside, glandular, the rim sub- 
truncate, minutely toothed; corolla hypocrateriform, varying from 
red, peach-red, red-violet, rose, rose-purple, rose-lavender, 
lavender-rose, pale rose-purple, or pink to purple, pale-purple, 
bluish-purple, mauve, lilac, violet, lavender, deep-lavender, 
lavender-blue, light-blue, bluish, or blue ("albo-rufescentes" 
according to J. A. Murray, 177U), its tube delicate, cylindric, a 
little longer to about twice as long as the calyx, usually less 
than 3 mm. long, the limb 1 — H mm. wide, the lobes more or less 
rounded; stamens inserted at about the middle of the corolla-tube; 
anthers not appendaged; pollen-grains oblate-spheroidal, 26 — 30 fj 
X 29 — 35 [i, usually 29 a x 32 u, the endocolpium lalongate, 3»$ux 
12 U ; ovaiy and style about 2 mm. long, glabrous; frui ting-calyx 
persistent, short-hairy, enclosing the fruit; fruit slightly lar- 
ger than in V. gracilescens , composed of U nutlets, easily separ- 
ating, the cocci broadly ellipsoid or shortly oblong, 1.^ — 2 mm. 
long, strongly striate, areolate-reticulate between the veins or 
striations toward the apex on the back; chromosome number: n - 7; 
2n = lU. 

Cufodontis (1962) states that this species, the type of the 
genus, is based on "syntypes: ex Europae mediterraneae ruderatisO 

It should be noted here that V. officinalis Auct., referred to 
in the synonymy above, is a synonym of V. g racilescens (Cham.) 
Herter; ?, officinalis Cuevas is Stachytarpheta jamaicensis (L.) 
Vahl; V. o fficinalis Wats, is V. menthaefolia Benth.; V. officin- 
alis var. gracibras Lehm. is V. litoralis H.B.K.; V. officinalis 
var. gracilescens Cham., var. gracilescens Graseb., and var. 
gracilescens Haumann-Merck are all V. g racilescens (Cham.) Hert- 
er; V. officinalis var. hirsuta Torr. is V. neomexicana (A. Gray) 
Small; V. officinalis var. major Osten is V. gracilescens (Cham.) 
Herter; V. officinalis var. mediterranea N^e is V. menthaefolia 
Benth.; 7. officinalis var. tenuis Cham, is V. g racilescens 
(Chan.) Herter; V. officinalis f . roseiflora Benke is V. halei f . 
roseiflora (Benke) Moldenke, as is also V. o fficinalis f . rosa 
Blackvell. 

The following hybrids are known: mth V. halei Small (=xV. 
conata Moldenke); with V. hastata L. (=xV. baileyana Moldenke); 
with Y. hispida Ruiz & Pav. (=xV. meretrix Moldenke); TrLth V. 
lasiostachys var. septentrionailis Moldenke (=xV. suksdorfi Mol- 
denke); with V. robusta Greene (=xV. c lemensorum Moldenke); with 
V. supina L. (=xV. adult erina Hausskn.); and with Veronica marit- 
ima L. (=xVeronicena haartmani Moldenke). The supposed " Verbena 
officinalis x bracteosa Barnes" is xV. perriana Moldenke (actual- 
ly V. bracteata Lag. & Rodr. x V. urticifolia L.), "Verbena offic- 



1961; Moldenke, Monograph of Verbena 211 

inalis x prostrata Dermen" is xV. suksdorfi Uoldenke (actually V. 
officinalis L. x V. lasiostachys var. septentrionalis Moldenke), 
and "Verbena officinalis x xutha Engelm." is V. halei Small. 

The "V. o fficinalis L." of C. Berg, Anal. Soc . Cientif. Argent. 
3: 199 (1877), which he found around Buenos Aires and describes 
as "Los ejemplares eran muy tiemos y t^nues", is V^ gracilescens 
(Cham.) Herter. The " Sacra herba Ruell." is a mint. 

Hooker (1829) describes a var, <X "caule erecto subhispido, 
foliis lanceolatis, inciso-serratis trifidisque segmentis incisis 
scabris, spicis filiformibus subpaniculatis , floribus remotiuscu- 

lis In Pampas ab urbe Bonaria usque ad Uendozam." He mentions 

also a var. ft "foliis grosse serratis vix profunde incisis ... .Apud 
Rio Saladi.llo ad limites occidentalis planitiei pampas dictae, et 

ad margines aquarum in Provinciae Mendozae Var. 5 differs fran 

the (J, and the common European state of the plant, solely in the 
less deeply cut leaves." His second variety is certainly V. grac - 
ilescens (Cham.) Hejrter. 

Linnaeus (1753) distinguishes his V. spuria by its "caulibus 
numerosis" from V. officinalis with its "caule solitario". He 
also uses the term "paniculatis" for the spikes of the latter and 
does not use it for those of the former. Poiret (1808) affirms 
that Michaux's "V. spuria, caule decumbente, ramisissimo , divari- 
cato; foliis ovalTbus, incisis; spicis fiirformibus , bracteis cal- 
ices superantibus" aind Clayton's "V. humilior, foliis incisis " 
are the V. spuria foxTii of the species, too. Hooker (I836) says 
"The V. spuria , Willd., which I possess from Pennsylvsuiia and New 
Jersey, I agree with Mr. Nuttall in considering a mere var. of 
officinalis ." 

The type of Verbena humilior foliis incisis Gronov. (and vari- 
ants of this pre-linnean name) is sometimes cited as " Clayton 8", 
but the figure here is the page number of Gronovius' work where 
the name appears . The " Verbena spuria inedito " of Ruiz & Pavon is 
V. menthaefolia Benth. 

Verbena recta Caesalp. is given under the genus Lye opus by Hal- 
ler in his Enum. Meth. Stirp. Helvet. 1: 66O (17U2) . This same au- 
thor places Verbenaca supina s. femina Fuchs and Verbenaca supina 
Cord, under what is no?/' known as Verbena officinalis on page 661 
of the sane work, but these names actually belong in the synonymy 
of V. supina L. On page 551 he places Verbena mas Fuchs and Verbena 
recta ^n mas Gesn. under what he calls Sisymbrium foliis pinnatis , 
extreme lobo triangulo, siliquis erectis, cauli adpressis , and 
which is what we now know as Erysimum officinale L. On page 62U he 
places Verbenaca recta Cord, in the synonymy of a species of Lyco- 
pus which he calls Alectorolophus calycibus glabris , foliis inter 
floris latioribus , 

Michel (17U8) regarded the Verbena of Caesalpinius as a "spe- 
cies" with dissected leaves, while the Verbenaca of Matthiolus he 



212 PHYTOLOGIA Vol. 10, no. 3 

regaixied as a "species" with not at all or only slightly dissec- 
ted leaves. Under the former he says "Haec, & sequens promiscue 
usiirpantur pro Verbena, Herba Sacra, Hierobotane, Herba S. loan- 
nis, & Berbena". C. Bauhin (1623) includes Verbenaca foemina 
Caesalp. and Verbenaca supina (& foemina) Fuchs in the synonymy 
of Verbena communis caeruleo flore (which is now V. officinalis), 
but these two names belong more properly in the synonymy of V. 
supina L. 

Steam (1961) notes that "although Linnaeus placed his genus 
Verbena in the class Diandria , having two stamens, he deliberate- 
ly included within it species with two stamens and species with 
four, among the latter being V. officinalis , the historic type of 
the genus." 

Of Verbena urticae folio canadensis, foliis incisis, flore 
ma j ore Tourn. Ray, in his Hist. Plant. 3: Suppl. 285 (170U), says 
"Hujus speciei folia pinguntur a Jacobo Zannoni in eadem tabiila 
cum Verbena con foglie d'ortica di Canada pag. 203- ubi plena ha- 
betur omnium partium descriptio." This appears to be V. urtici- 
folia L. 

Matthiolus (162U) regsirded Verbenaca supina Fuchs and Verbena 
foemina Caesalp. as synonyms of V. officinalis , but these names 
actually belong in the synonymy of V. supina L. 

Schauer (18U7) included V. setosa Mart. & Gal, in the synonymy 
of V. officinalis , but it really belongs with V. menthaefolia 
Benth. In my Prelim. Alph. List Invalid Names U6 (19^0) I regar- 
ded V. domingensis Urb. as a synonym of V. officinalis , but I now 
maintain it as distinct. Similarly, in my Alph. List Invalid 
Names Suppl. 1: 22 (19U7) I reduced xV. adult erina Hausskn. to 
synonjmiy under V. officinalis , but now regard it as probably dis- 
tinct. 

Lam (1919) places V. macrostachya F. Muell. and V. menthaefo- 
lia Benth. in the synonymy of V. officinalis, but I regard the 
former as a valid variety and the latter as a valid species. 

Verbena communis caervtleo flore C. Bauh. is accredited to 
"G. Bauh." by J. Bauhin (I689) and is given as a synonym of V. 
supina L. Actually, Caspar Bauhin" s given name was sometimes 
written "Kaspar" or "Gaspard". 

Dorsten (l^UO) says "Verbenae duo sunt genera. Altera floribus 
est albis, altera vero coeruleis". Haller (17U2) recognized a 
variety "6" with white flowers (now known as f. albiflora Strobl) 
and a variety "y"with variegated leaves ("Folio variegata Breyn 
Prodr. 11, p. 100"). I have not as yet seen any specimens with 
variegated leaves, but the form may well exist and may be worthy 
of nomenclatural recognition. On page [96] of his 1768 work, 
Haller regards the Hort. Florent. p. 98 plant as a variety » P> 
Foliis vix dissectis". Ray (1701;) and Linnaeus (1737) regarded 
Toumefort's "Verbena lusitanica latifolia procerior " as a vari- 
ety — designated as "(I"by Linnaeus — of V. officinalis. 



196U Moldenke, Monograph of Verbena 213 

No description is given of V. officinalis var. natalensis in 
the original place of publication, but the name is based on F. 
Krauss 1^1 "ad fluv. Umlaas, Natal, Dec." 

iiKoch & Almquist, Svenska Flora 1", "Hochst., PI. Schimp. Ab- 
yss. I. Iii5", and "Poeppig, Coll. PI. Chile 1^8" are sometimes 
cited as though they were literat\ire references, but appear to 
be exsiccatae. L'Ecluse, Rar, Plant. Hist. XLV (l601) is some- 
times cited as "2: US, fig. 1", as "xiv", or as ""Rar. Aliq. Stirp. 
Pannon. p. XLV.l.B.III.p.U;3. 1583" — the last-mentioned may be 
a reference to Bejthe, Stirp, Nomencl. [iJj.] . 1$Q3- The Haartman 
(17^6) reference is sometimes given as "pi. 11" [=11] . The date 
on the volume title-page is "17^6", on the second title-page is 
"August 20, 1756", and at the head of the article itself is "Nov- 
ember 23, 1751". On page 35 it is stated that "Floruit quidem 
haec planta canni anno felicissime, in annim que haec edimus 1755 
& vivi.*' This is sometimes misquoted as "1775"- 

It is perhaps worth mentioning here that J, F. Gronovius, Fl. 
Virg., ed, 1, 8 (1739) is sometimes cited as "Clayt., Fl. Virg. 
8. 17U3". 0. F. Mtlller, Icon. Plant. Fl. Dan. 1;: pi. 628 (1775) 
is also cited as "1777"; Briq. in Engl. &•. Prantl, Nat. Pflanzen- 
fam. 1; (3a): lii7 (I89U) is often cited as "1895"; and Fiori & 
Paoletti, Icon. Fl. Ital. fig. 3218 (1902) is also cited as 
"1895". The L., Fl. Suec, ed. 1, 9 (17h5) reference is some- 
times referred to as page "26", which figure actually is the 
species number on page 9; L., Mat, Med. 6 (17U9) is scmetimes 
erroneously cited as page "38"; J. Bauh., Hist. Plant. Univ. 3: 
hh3~hhh (l650) is sometimes cited as "l65l" or "1750"; Gerarde, 
Herbal 580 (1597) is sometimes erroneously cited as page "781"; 
Zetterstedt, Pi. Vase. Pyren. 220 (1857) is sometimes incorrect- 
ly cited as page "1050", that figure being the species nimiber on 
page 220. Roig y Mesa, in his Pi. Medic. Cuba 67I (19l;5), cites 
Robledo, Lecc. Bot. 2: 526 (19ii0), but the correct page is U97. 
Sabbat in Martelli, Hort. Roman. 3: H, pl. 55 (1775) is some- 
times cited as an illustration of V. officinalis , but it is pl. 
56 Tfhich has the illustration of this plant. Zannich., Istor. 
Piante Venez . pl. 269 (1735) is sometimes cited as "pl. 289", 
but the latter is an illustration of a composite, there designa- 
ted as Cyanus minor . Hofmeister, Jahrb. Wiss. Bot. 1: 82 — 190 
(1853) is sometimes cited in the bibliography of V. officinalis , 
but the species is mentioned only on page lUO, while the "M, 
Treub, Ann. Jard. Bot. Buitenz, 3: 77—87 (1883) reference is 
entirely incorrect, since the species is not mentioned anywhere 
on those pages. Hara (I9U8) cites fig. 1319 (1933) and fig, 563 
(19liO) in two books with Japanese titles not decipherable to me, 

Eig, Zohary, &: Feinbrunn (1931) state that the leaves of V, 
officinalis are "pinnate", but actually they are always simple I 
These authors probably meant to say "pinnately parted" . They 
note that in Palestine the species is "found almost all year 
round" (i.e., in anthesis?) . Zohary (1962) says that it com- 
prises part of the Rubion sancti Alliance in the Inuletum visco- 
sae Alliance among the ecologic formations there. 



2lU PHYTOLOGIA Vol. 10, no. 3 

Common and vernacular names for this plant are numerous, in- 
cluding "akkoraragg", "Altarblimie", "aristereon", "Aschlepius 
alceas", "ashthroat", "berbena", "berbina", "berbine", "bemena", 
"blue vervain", "camaradinha", "candelabra verbena", "chanarlycusy 
"chamaelygos", "chamelicos", "chamticos", "columbina", "columbine',' 
"common veruaine", "common vervain", "co roi ngua", "creisetta", 
"crista gallinacea", "crocetta", "curetis fersefomon", "cyparis- 
sos", "doaetrias", "diose lacete", "Druidenkraut", "Druivenkruid", 
"echtes Eisenkraut", "Eisen-bart", "Eisen-bart kraut", "eisencraut',' 
"Eisen-dek", "Eisenhart", "Eisen-hart", "Eisen-hart kraut", "eisen- 
hendrik", "Eisenherz", "Eisen-herz", "Eisen-herz kraut", 
"eisenhindrik", "Eisenkraut", "Eisen-kraut", "eisen krokt", 
"Eisenreich", "Eisen-reich", "Eisenweich", "Eisen-weich", 
"eisener Heinrich", "eiserich", "eiserich kraut", "eisem*, 
"eisemhart", "eisem-hart", "eisevri-ch", "eisewig", "eisick", 
"enchanter's plant", "enchanter' s-plant", "erba de berm^na", "erba 
de San Giovann", "erba trona", "erba turca", "erbo a touti li 
man", "European- verbena", "European vervain", "European vervein", 
"exupera", "eysencrut", "eysenhart", "eysenkruyd", "eyssen- 
chravrtt", "eyssen-kraut", "Feldcypresse", "ferraria", "gebr^ch- 
liches Eisenkraut", "gemeine Eisenkraut", "gemeines Eisenkraut", 
"geweihet kraut", "geweiht-kraut", "geweiht kraut", "gtfvercin 
otu", "Hahnenkampf", "Hahnen-kampf", "frog's-foot", "frossis- 
foot", "Hahnenkopf", "Hahnen-kopf " , "hardijzer", "heiliges Krauty 
"Heiligkraut", "Heilig-kraut", "herba coliombariae", "herba santa',' 
"herba S. lonnis", "herba veneris", "herba verbenae", "herba 
verbenae officinalis", "herba verbenae sanguinalis", "herbe a 
tous les maux", "herbe a tous maux", "herbe aux enchantements", 
"herbe aux sorciers", "herbe aux sorcieres", "herbe de foie", 
"herbe de sang", "herbe de vervaine", "herbe di San Giovani", 
"herbe du foie", "herbe sacr^e", "herbe-sacr^e", "herb-grace", 
"herb of the cross", "herb-of-the-cross", "herb o' grace", "herb 
of grace", "herculania", "herva de ferro", "hierabotane", 
"hierobotane", "holie herbe", "holy herb", "holy-herb", "holy 
veruane", "holy vei^ain", "holy vrort", "iezerkroet", "iherabotaney 
"iisercruyt", "ijsenkruyd", "ijserhard", "ijserhart", "ijserkruydy 
"ijzerhard", "ijzerkruid", "irenhard", "Iren Hendrek", "isarna", 
"isama chrUt", "isarnina", "isecruyt", "isekrut", "isena", 
"isenarre", "isen-bart", "isen-brut", "isen-hard", "isenharde", 
"is en-kraut", "fsenkrflt", "iserbart", "isercruit", "isercruyt", 
"isere", "iseren-bart", "iserene Hendrek", "iseren-hard", "iseren 
krut", "iserhark", "iserhart", "iserhert", "iserich", "iser-krauty 
"isem", "isernehart", "isem Kendreck", "isern krut", "isin- 
chlete", "isin-lna", "isin-un", "isim", "isimwurz", "isni", 
"issenkraut", "issemhar", "issinkraut", "lunos teares", "jagged- 
leaved vervain", "jernurt", "Juno's herb", "Juno's tears", "Juno'a- 
tears", "JunothrSCnen", "Juno-trSCnen", "jururuba", "kardita", 
"Katzenblut", "Katzenblutkraut", "kau kau yexik", "kau nga ts'o", 
"kerckkruyd" , "kerkkruid", "Krauskraut" , "kuma-tsuzura" , "la 
verveine", "la verveine officinale", "lecheri", "licinia", 
"lightning plant", "lung nga ts'o", "lustam", "lustrago", "ma pi6n 
tsao", "ma pin tso", "ma pin t'so", "ma tien thio", "matricalis", 



1961; lloldenke, Llonograph of Verbena 2l5 

"llercurie's noist blood", "IJircuries noist bloude", ''Wercury's 
moist blood", "Merkurblut" , "militarem", "mine ?iQ", "mine qxq o',' 
"native vervain", "ngoh sat na", "officinal vervain", "Opfer- 
braut", "Opfemblut", "pImTikh", "panchromos", "pancremon", 
wpempentar", "peristerian wort", "peristerion", "perstereona", 
wphiltrodotes", "pigeon's grass", "pigeon' s-grass", "pigeon's 
grasse", '♦pigeon weecJ", "pitagosas", "planta de sorte", "pushtu*, 
"reichhard", "reich-hard", "reich-hart", "Richard-kraut", "rigi 
el khamSm", "rigl el khamSm", "ri'I el khamSm", "sacra herba", 
"Sagenkraut", "S eg en-kraut", "seruftit", "serrufit", "shamuki", 
"shop vervain", "sideritis", "simpler' s joy", "simpler »s- joy", 
"sirpina", "sister vervain", "ssirugtiSCt", "Stahlkraut", "Stahl- 
kraut", "standing berbana", "Tauben-kraut", "Taubenlieb", 
"tialu", "tigrodion", *trygonivmi", "trygonum", "urgebSo", 
"varveino", "varveyn", "Venusader", "Venusblut", "verain", 
"verbena", "verbenaca", "verbena colombaria", "verbenam", 
"verbene colombaria", "verbene officinale", "verminacola", 
"verminacula", "vertiperdum", "veruaine", "veruayne", "veruana", 
"vervain", "vervains masse", "vervayne", "vervein", "verveine", 
"verveine batard", "verveine bttarde", "verveine ccsmnune", 
•nrerveine d' Europe", "verveine officinale", "verveino", "verven',' 
"vervena", "veirvene", "verveyn", "vervin", "vrgebacci", "".Veihs- 
prossen", "Wieselblut", "yrilder eiserrig", "wild hyssop", 
"worogobd", "yik mo tso", "ysen-hard", "ysen-krtlte", "ysercruit',' 
"ysere", "yseren", "yseren-hard", "yseren-hart", "yser-hard", 
"yser-hart", "yserkruyd", "ysem", "ysemhard", "ysinina", and 
"yzerkruid" . 

Verbena officinalis has been collected in anthesis and in 
fruit in ever;y- month of the year, growing at altitudes from sea 
level to 11,000 feet. It is found along roadsides and ditches, 
in fields, wheatfields, pastures, roadside meadows, waste places, 
grassy ground, damp or moist places, vacant lots, railroad 
grounds, and very dry pastures, on dry gentle slopes, grassy 
hillsides, village commons, and mountainsides, along village 
streets, rocky paths, sandy roadsides, and canal-banks, at the 
borders of rice fields and the edges of barnyards, in moist or 
sandy soil, in damp, grassy, or cultivated ground, on old ballast 
in shipyards, in d ry land under trees, lowlands, cleared areas, 
and the lawns of old houses . 

Stewart & Corry (1888) report that it is casual in northeast- 
ern Ireland, adding the comment that "Templeton notes this spe- 
cies as about gardens not indigenous, and not naturalized", 

but MacKay (I836) says "roadsides and waste ground in a limestone 
soil. Plentiful near Cork, and Killarney, and at Kilmacannick, 
County of V/icklow." L^rs . Clemens says that it is "gay in pad- 
docks when nothing else showed color except chenopods" and "pretty 
weed along road" in Queensland. Keogh found it "in densely tim- 
bered country" in New Guinea. Suvamakoses found it scattered in 
mixed deciduous forests in Thailand, while Kasapligil (1956) re- 
ports it growing along ditches and in moist places in Jordan. 

Domin (1928) gives its distribution as "temperierte und sub- 
tropische Gebiete der ganzen Erde. In Australien mit Ausschluss 



216 P H Y T L G I A Vol. 10, no. 3 

von Nord-Australien und West Australia, daftir aber aiif Tasmanien." 
He states that in Queensland it inhabits savanna woods, groTsing 
in sandj'- soil, and occurs as a weed on rsdnforest soil cleared for 
cultivation. Verdoom (1938) says "Widely distributed from the 
Cape Peninsula through the south-eastern and eastern districts 
[of the Republic of South Africa] to Natal; also recorded frcm the 
Orange Free State and the Transvaal; appears to be rare in the dry 
central and north-western districts as there is only one record 
from Prie3ka....A native of Europe and naturalized at the Cape 
over 100 years ago. In seme areas it is a troublesome weed in 
lands but plants may be destroyed by the usual methods of clean 
cultivation before the seeds ripen. In some parts of Griqualand 
East it is said to be a good winter feed for large and small 
stock." 

Franchet (1882) records it from French Scmaliland, -vriiile Lou- 
reiro (1790) records it as both wild and cultivated in China and 
Cochinchina. Hausman (19U8) describes the flowers as purple and 
states that the species is found in the United States from "Maine 
south to Florida; west to Tennessee, Texas."' All the Texan mat- 
erial so determined, hovrever, has proved to be V. halei Small. 
Berg (]877) records it from Buenos Aires, saying "Encontr^ esta 
especie en muy pocas partes cerca de Buenos Aires . Los e jempla- 
res eran muy tiernos & t^nues" . The material to which he refers 
here is actually V. gracilescens (Cham.) Herter. Bojer (1837) 
gives the temperate zones of Europe and Asia as the home of V. 
officinalis, and "Perou" as the original iiome of V. spiiria (1). 

Connor & Adams (19^1) aver that V. officinalis is "occasional 
and sometimes locally abundant in waste places on both islands" 
of New Zealand; Bose calls it a weed of waste places in India; 
Fahrenholtz found it abundant at the edge of ditches in Germain; 
Bailey encountered it as a weed in parks in China; Sampson found 
it to be a "common weed of vraste places, abundant on the city 
walls [of Canton]; flowers all the simmer"; Rodin reports it 
from among cultivated valleys and hills with sparse vegetation in 
Pakistan; Clemens calls it a weed near houses in Luzon; Ching 
calls it "common" in Anhwei, China, but Lei found it "rcire, among 
scattered shrubs in dry level land"; Collins reports it as com- 
mon in Bermuda. Puchtler encountered it growing with Carduus 
nutans , Lappa officinalis , Persicaria hydropiper , P . lapathifo - 
lia^ and Thymus serpyllum in Germany; Thedenius refers to it as 
"subspontaneous" in Sweden; Yuncker calls it a "roadside weed" 
on Okinawa and describes it as a "weed in waste areas and occas- 
ional along roadsides" and also as a "roadside weed" on Niue. 
Black (1926) describes it as rare or localized in temperate 
Australia; Ylatt (1893) reports it "common in the Himilaya from 
Kashmir to Bhutan, at altitudes of 1,000 to 6,000 feet, and in 
the Bengal plain to the Sunderabands." Radford & Stewart found 
it growing along a wagon road at the edge of a saltmarsh in 
North Carolina, while Mackenzie reports it as uncommon along the 
streets of Cape May, New Jersey. Murrill (19U5) calls it "rare 
in Alachua Co\mty, Florida", but the plant to which he refers 



196U Uoldenke, Monograph of Verbena 21? 

here is V. halei Small, 

Gonial ves da Cunha & Gongalves Sobrinho (1938) record V^ offio - 
inalis from the island of Faial, in the Azores, where they say- 
that it grows in association with Selaginella denticulata , Rubus 
rusticajms. Polygonum aviculare , Hypericum boeticum , Blechnum 
spicant, Asplenium hemionitis, and Scolopendrium vulgar e . In 
their I9U0 publication they aver that it is not very frequent and 
has about the same frequency as Ranunculus megaphyllus , Euphorbia 
stygiana, Rubus hochstetterorum, Ammi seubertianum, Vaccinium cy- 
lindraceum , Cymbalaria muralis ^ and Mentha piperita. Buchli 
(1936) calls it a field weed in northeastern Switzerland; W. Young 
(1936) records it frcm Scotland; Soest (1931) gives us a distribu- 
tion map of its occurrence in the Netherlands and Lid (19^2) does 
the sane for Norway; Bojer (1837) records it as cultivated on the 
island of Mauritius, \diere he says that it rarely flowers. Koch 
& Almquist record it from Finland, while Femald records it from 
Dinwiddle, Henrico, and Princess Anne Counties, Maryland. Kara 

(19^6) gives its distribution as "Europe Asia Minor, Caucasus, 

3,*. Asia, c. Asia, n. India, Siam, Indo-China, China, Korea, 
Honshu, south to Formosa, Australia, Polynesia. The plant is 
weedy, and the Japanese specimens agree well 7ri.th the European." 
Beam (I9U0), on the other hand, excludes it from the flora of 
Indiana. C. & E. Reid (1907) report it in the fossil form from 
the Pliocene of England ajid Belgium. 

The G. V. Nash s,n. [7 S. I898], cited below, was cultivated 
in New York from seeds secured from Geneva, Switzerland, in I898. 
The Herb. Calif. Acad. Sci. 31390, also cited below, does not 
bear any indication on its label that it originated from cultiva- 
ted material, but I assume that this was the case. The species 
has been in cultivation as early as 1799 in England, according to 
Murray (1799). 

Herbarium material of V. officinalis has been misidentified 

and distributed under the names V. littoralis H. & B., V. littor - 
alis H.B.K., Vj_ riparia Raf ., V. ~s"upina L«, V. supina Spreng., V_. 
urticifolia riparia (Raf.) Britton, V. xanthi Gray, y_, xutha 
Lehm., Bouchea sp., Buchnera sp., Stachytarpheta sp., and Veroni- 
ca officinalis L. The Bornmtlller s.n. [Gebirge Juda, Juli 1897] 
collection was originally distributed as V. tenuispicata Stapf, 
and no. $$16 as V. officinalis var. grandiflora Hausskn. Schimper 
lii^, at least insofar as the United States National Herbarium 
specimen is concerned, seems to be var. prostrata Gren. & Godr. 
Paez 112 is V. bipinnatifida Hutt. 

The E. F. Constable 11633 , Pedley 377 , and L. S. Smith 30U5, 
distributed as V. officinalis , are actually var. gaudichaudii 
Briq.j Schlieben 769I is V^ brasiliensis Veil.; Chardon 3U, J_.. T^ 
Curtis Jr. s.n. [July 27, 19hh] , Eggers 1823 , E^ L^ Leonard 3939 , 
Questel 1686, and C. Wright 36^8 are V^ domingensis Urb.; Jameson 
12^/26 and Shepard~2$U are V. gracilescens (Cham.) Herter; Beamer 



218 PHYTOLOGIA Vol. 10, no. 3 

3.n. [April 30, 1933], W. H. Dimcan 20211; , G. Een s.n. [29.3. 
19^1] , R. K^ Godfrey ^3168 , E. Hall U32 , R^ L^ McGregor 15^, E. 
J. Palmer 336oU, J, Skehan s.n. [Seymour & Earle 109], and B. L. 
Wagenknecht 2^99, as well as almost all the Texan specimens so i- 
dentified, are V. halei Small; Rodin 3917 and Tttrckheim 90li are 
y. litoralis H.B.K.j Abraps 3U06, Lyonnet 33U, C_. T^ Mohr s.n. 
[Huatusca, 18^7] , E. W. Nelson 19U3 & U577, C. R. Orcutt s.n. 
[Apr. 1889], J. G^ Ortega U21g , Edw. Palmer ll|l , 1^3, 268, & 10U2 , 
Pringle 853U, and Salazar s.n. [Nopala, Aug. 1, 1913] are V. 
menthaefolia Benth.j Havard s.n. [Texas], Parry , Bigelow, Wright , 
& Schott 827 , and Tharp s.n. [Wilson Ranch] & s.n. [V/ilson Ranch, 
[June 1931] are V. neomexicsina var. hirtella Perry; Meams IOI3 , 
Edvf. Palmer 339 V2 & lOUl , and F. W. Pennejl I688O are V. neo- 
mexicana var, xylopoda Perry; Gall egos 23l;2 is V. orcuttiana Per- 
ry; T. £. Hale s.n. [Baraboo, I86I] and T. A. Williams 119 are xV. 
perriana Moldenke; E. W. Nelson 6096 and Edw. Palmer 3^6 are V. 
pinetorum Moldenke; Sidey II96 is V. rigida Spreng.j M. E^ Jones 
s.n. [Crinnell, Aug. 1877] is V. striata Vent.; C. C_. Albers 
3UOO8 is V. tenuis ecta Briq.; Gauba I698 is V. tenuispicata Stapf; 
Parry & Palmer 718 is the type collection of V. teucriifolia var. 
corollulata Perry; Ripley s.n. [Sep. 10, 1902] is V. urticifolia 
L.; Clover 1695, Eggert s.n. [along railroads, 10 June I898], 
Harding 399 , and W. IL. Rhoades s.n. [Jackson, 1927] are V. xutha 
Lehm.; and Gorman hOO$ is Cochranea anchusaefolia (Poir.) Gttrke 
in the Heliotropiaceae, while Lap ham s.n. , distributed as »V. 
spura L.", is actually V. hastata L. 

The Behn s.n. [7 Decbr. 1920] collection has extra large 
leaves. The Small & Heller s.n. [July I6— -17, I89I] collection ap- 
pears to be exactly like some specimens of V. riparia Raf ., and 
may prove to be that species. Tanaka & Shiiaada 11032 and H. Smith 
1536 exhibit pubescent foliage like that seen in V. menthaefolia 
Benth. Meissner 572 possibly represents var. prostrata Gren, & 
Godr, Baldacci 1^2 has spikes that are rather long and stiff, 
while T. Tanaka 11032 has them rather dense and long. Phares 1733 
is described by the collector as "a form with deeply incised 
leaves". G. P. En 2022, from "a garden" in Fukien, may be from 
cultivated material, but the label does not plainly indicate this . 
Gowgill 1730 was cultivated in Maryland from seeds secured on Mt. 
Kaakerpu, at an altitude of 3OOO meters. The McCann s.n. [9-1$- 
36], cited below, was cultivated in Maryland from seeds seciired 
from Brussels, Belgium. Liu L.831 was cultivated in Peking from 
seeds secured from Copenhagen, Denmark, while L. H. Bailey s.n. 
[Sept. 21, 1923] was cultivated at Ithaca, New York, frcm seeds 
secured from Vilmorin-Andrieux & Cie . ( no. ii96$9) , sown on April 
27th. Bailey (1935) reports that the seeds of this species are 



196U Moldenke, Monograph of Verbena 219 

offered to the horticultural trade also by Floraire, Haage & 
Schmidt, Kew, and Taihoku, The snecies is known definitely to 
have been ciiltivated in Spain in iSlU and in France in I8I9. Cul- 
tural directions given by Vilmorin-Andrieux are "Semer de mars en 
Juillet, en p^pini^re; repiquer en p^piniSre; mettre en place en 
autonne, de pref^r^nce, ou au printemps, en terrain sain. — Fl. 
de juin en ao'flt." 

The Haussknecht s.n, specimens from Greece, cited below, may 
actually represent the type collection of xV. adulterina Hausskn., 
but I see no differences between them and small specimens of typ- 
ical V. officinalis. They are not at all hairy. Perhaps they 
represent one of the F2 segregates reverting to ancestral type. 

The " Verbena officinalis " of I. C, Verdoom, Union of S. Afr, 
Dept, Agr. & Forest. Bull. 185: 171, fig. 90 (1938) is actually 
V, litoralis H,B,K. The Kausch s.n. [Rottstock, I869] is a mix- 
ture with V. hastata L., while M. K. Clemens s.n, [Forest Hill, 
Jan. 19kh] is a mixture with two species of Fabaceae and A. H. 
Kahn s.n, [8.7.25] is also a mixture with something non-verbena- 
ceous. 

Urban, in his discussion of V. domingensis, cites Eggers 2175 
as V. officinalis . He maintains that it differs from V. domin- 
gensis in having subsimple stems, many times larger leaves, which 
are pinnatifid or the lower ones pinnately parted, and larger 
flowers. The length of pistil in relation to size of pollen 
grains is discussed by Covas & Schnack in Darwiniana f: 86 (19li.5) • 
According to Seymour (1929), V. officinalis is attacked by the 
fungus Erysiphe cichoracearum~DC"!i 

The tvro specimens of V. officinalis from Czechoslovakia pre- 
served in the Stockholn herbarium from Linnaeus' herbariimi are 
inscribed as follows: The first has on the obverse side in ink 
"■i " and " Verbena officinalis Linn. 13", the first and third 
words in a later hand, the second vrord in an early hand, the 
number probably in Linnaeus' hand; on the reverse side in pencil 
"LinnI herb." and in ink (in Linnaeus' handwriting) "Verbena 
communis Gerb. tamay 312" and in the same hand "Habitat in in- 
cultis Verania circa Lubnam", followed (in an unknown hand) by 
"von Linn6 sen. scripsit", and then "Herb. Alstroemerii", and 
finally (in Dahl's handwriting) "a Linn6 P." The second specimen 
has on its obverse side in pencil "Verb. 13 offic. L. sp. pi. 20" 
and in ink (perhaps in Linnaeus' hand) "13 officinalis"; on the 
reverse side in pencil "Linn6 herb." and in ink "Herb. Osbeckii" 
ajid "Verbena officinalis" and "Linn." (in a different hand), and 
finally "a Linn^ f." (in Dahl's handwriting). 

Gerarde (1597) calls this plant "Common veniaine" and "Holy 

veruane" . He says "Vervain is called in Latin Verbena, & 

Verbenaca, Herculania, Ferraria, & Exupera, or some Matricalis, & 
Hierobotane, of others Veruena, & Sacra herba. Verbenae are 
herbs that were taken from the altar, or from some holie place, 
which because the Consul or Praetor did cut up, they were like- 



220 PHYTOLOGIA Vol. 10, no. 3 

■wise called Sagmina, which oftentiines are mentioned in Liuie to 
be grassie herbs cut up in the capitol. Plinie also witnesseth 
in his 22. book, II. chap, that Verbenae & Sagmina be all one, L 
this is manifest by that which vre read in Andria in Terence: Ex 
ara verbenas hinc sumej Take herbs here from the altar, in -vriiich 
place Terence did not meane Veruaine to be taken from the altar, 
but some certain herbe: for in Menander, out of whom this Comedie 
was translated, is read MVP<5vi|, or Myrtle, as Donatus saith. In 
Spanish it is called Vrgebaomj in Italian Verminacula: in Dutch 
Isercruitj in French Veruayne: in English lunos teares. Mercuries 
moist bloude, Holie herbe, and of some Pigeons grasse, or Colvmi- 
bine, because Pigeons are delighted to be amongst it, as also to 
eate thereof, as Apuleius writeth." 

Mrs. William Starr Dana, in her "How to Know the Wild Flowers", 
says of the verbena: "It was believed to be the herba sacra of 
the ancients, until it was understood that the generic title Ver- 
bena was a word which was applied to branches of any description 
which were used in religious rites. It certainly seems however, 
to have been applied to seme special plant in the time of Pliny, 
for he writes that no plant was more honored among the Romans 
than the sacred Verbena. In more modem times it has been re- 
garded as an 'herb of grace' and has been gathered with various 
ceremonies and with the invocation of a blessing, which began as 
follows; 'HalloT/-ed be thou. Vervain, As thou growest on the 
ground. For in the Mount of Calvary There thou wast first f oundl ' 
Mrs. Alice Earle ^yde writes in a letter dated October 29, 19iiO, 
that it is said to have grown at the foot of the cross on Cal- 
vaiy." In the book entitled "Poety of the Flowers" the rl-^jrme 
quoted above is said to be an "old Cornish superstition". 

Friend (1883) records that the plant is called "Holy Herb", 
and according to Pliny was one of the sacred plants of the Druids 
and was gathered by them with all majmer of mystic ceremonies. 
An old rhyme is 

"Hail to thee. Holy Herb J 

Growing on the ground 

On the Mount of Olivet 

First wert thou found. 

Thou art good for many an ill. 
And healest many a wound; 
In the name of sweet Jesus, 
I lift thee from the grouiKl." 

William Coles, in his "Adam in Eden", says "It is known to 
such as have skill of nature, what wonderful care she hath of the 
smallest creatures, giving to them a knowledge of medicine to help 
themselves, if haply diseases are among them. The Swallow cureth 
her dim eyes with Celandine; the Wesell knoweth well the virtue of 
Herb Grace; the Dove the Veirven; the Dogge dischargeth his mame 
with a kind of grass; and too long it were to reckon up all the 
medicines which the beasts are known to use by nature's direction 
only." 



196ii Moldenke, Monograph of Verbena 221 

Friend states also that "vervain is said to have the power to 
open locks and unshoe horses" ... ."vervain leaf is called 'Frog's- 
foot' from fancied resemblauice to the foot of a frog. A cuplet 
reads 'Frossis-foot men call it. For his levys are like the 

frossys fet.' Witches a re said to be fond of vervain and to 

use its juice with that of henlock, nightshade, & St. Johnswort, 
to mix into their baleful draughts prepared for their enemies. 
But this is contradicted in St. Colne's charm, as sung by Meg 
Merrilies at the birth of Harry Bertram: 'Trefoil, Vervain, 
John's Wort, Dill, Hinder witches of their will'. It is also 
contradicted in the old rhyme given in the notes to the 'Demon 
Lover' in the 'Minstrelsy of the Scottish Border': 'Gin ye vmd 
be leman mine. Lay aside St, John's Wort and Verveine' ... .Conway 
remarks that in France the Mandrake superstition seems occasion- 
ally to invest some other root. Thus the author of 'Secrets du 
Petit Albert' (Lyon, 1718) says that a peasant had a Bryonia root 
of human shape, v/hich he received from a gipsy. He buried it 'at 
a lucky conjunction of the moon with Venus' in spring, and on a 
Monday, in a grave, and then sprinkled it with milk in T<hich three 
field mice had been drowned. In a month it became more human- 
like than ever. Then he placed it in an oven with Vervain, wrap- 
ped it afterwards in a dead man's shroud, and so long as he kept 

it, he never failed in luck at games or work' Conway says 'In 

reading accounts of the vdtch trials, especially those of the 
south of England, one can hardly help remarking that in the antics 
by which so-called vritches imposed upon their neighbours the 
plants used by them are almost always Rue and Vervain' ... .Brand 
quotes from Scot's 'Discover of V/itchcraft' , pp. l5l — 1^2, that 
'To be delivered from witches (in England), they hang in their 
entries verven' and 8 or 10 other plajits ... .Valerian and Ver- 
vain are famous plants, and are scarcely ever mentioned, especi- 
ally the latter, without reference being made to their mystic 

character I have named the Vervain already on more than one 

occasion, and shall therefore say as little about it here as 
possible. Several suggestions have been made respecting the or- 
igin of the word, Professor Max Mtfller being of the opinion that 
it is connected Td.th brahman, a word intimately associated with 
India. Although a favorite with witches, it was at the same time 
one of the plants which hindered them f rem carrying out their e- 
vil designs. Among other plants 'Vervain and Dill Hinder witches 

from their will' Among the ancients Vervain was sacred to the 

god of war, and was borne by ambassadors when they went to defy 
or challenge the enemy. It became associated with the god of 
war and thunder in Germany also, and was thus supposed to be cap- 
able of protecting houses from lightning and storm. Mr. Conway 
remarks that even yet, in some districts of England, children may 
be seen with Vervain tvaned about their necks, little laiov/ing how 
nearly it has been related in times of witchcraft to a halter. 
Pliny tells us that the Druids made use of it in casting lots, in 
drawing omens, and in othor pretended magical arts ... .But although 
it was once so famous, the plant has lost its glory. Among our- 
selves it has fallen into disuse, in spite of the fact that a 



222 PHYTOLOGIA Vol. 10, no. 3 

pamphlet was sane years ago written expressly to recommend it, 
directing the root to be tied with a yard of vihite satin ribbon 
round the neck, and to be allowed to remain there till the patient 

recovered The poet Drayton in his 'Muses' Elysium' says 

A wreath of Vervain heralds wear. 

Amongst our garlands named; 
Being sent that dreadful news to bear, 
Offensive war proclaim' d. 
The heads of Roman priests were scmetimes garlanded with Ver- 
vain, the sacred and magic herb .... .Vfreaths of flowers have scme- 
times been worn around the neck as an amulet or charm ... .Vervain 
7»a.s one of the flowers so employed. By the Greeks and the French 
it was known as a sacred herb, and was used not only as an amulet, 
but also to cure venomous bites and various diseases. Its reputa- 
tion was sufficient in the time of Ben Johnson for him to write: 
•Bring your garlands, and with reverence place The Vervain on the 
altar' . An English writer has also recommended that the root be 
tied with a yard of white satin-riband around the neck, there to 
remain till the patient recovers; and Mr. Conway says children 
may still be seen in some places with the plant twined about their 
neck. The ancient Persian magi made great use of this plant in 
their worship of the sun, al?ra,ys carrying branches of it in their 
hands as they approached the altar. It was one of the plants ded- 
icated to the goddess of beauty, while Venus vrore a crown of 
l/^tle interwoven with Vervain. Roman ambassadors or heralds-at- 
arms also wore cjrowns of Vervain when they went out to proclaim 
war, or bid their enemies defiance; a custom to wMch Drayton re- 
fers in the lines already quoted. Vervain 'was sacred to the god 
of war', says Mr, Conway; but we fail to see the force of this. 
The Germans are said to have until quite recent times presented a 
wreath of Vervain to the newly-^narried bride, as if to put her 
under the protection of Venus the Victorious. The history of the 
plant, and of its name as well (if it be from the same root as 
brahman, as Professor Max Mtlller suggests), is full of interest.. 
Besides being indebted to Brand's 'Popular Antiquities' for sun- 
dry notes, quotations, and references, I ovight to refer to the 
use I have made of 'Flora Historica' for Vervain and Mandrake." 
Ingram (188?) says that this plant is symbolic of magic and 
inspiration in the "language of flowers". Ranson (19U0) says 
■The name 'verbena' has come to us from the latin, and means 
sacred branch. Crowns of berbena, or vervain, the family name, 
were used in Roman public rites . Both the Greeks and Romans re- 
garded the verbena as a peacemaker, and diplomats wore it often 
in their efforts to make peace with others. I-t was used also in 
marriage ceremonies, just as the orange blosscmi is now.... In the 
Far East the verbena vras considered a symbol of enchantment, and 
was thought to be strong enough to open a locked door. 
"Verbena 

These dainty round bouqaets were made by fairies: 
Their tiny fingers, deft and very strong. 

Arranged the lovely fragrant purple blossoms 
In beautiful rosettes the whole day long. 



196Ii Moldenke, Monograph of Verbena 223 

I think the busy, sprightly little fairies 
Must know a thousand happy things to do — 

Perhaps if we were diligent and patient 

Our minds and fingers would be skillful, too, 

"Venrain 
This is a wayside altar — 

This quiet country place — 
The vervain candelabra 

Lends dignity and grace. 

To services of gratitude: 

Their blue-tipped candles flame. 
And all the flowers genuflect 

And praise their Maker's name. 

"The vervain was a sacred plant to the Druids, and of Thor, 
Scandinavian god of thunder. It was used by witches in their 
brews, and also was employed to hinder their spells. During the 
time of Shakespeare vervain was hung, with dill and a horseshoe, 
over the door to keep our evil spirits. It was considered by 
the ancient herb doctors, or 'simplers', as a cure for almost 
eveiy disease known. Anxious mothers hung bits of it about 
their children's necks as a charm against sickness and evil. 
Christians attributed its miraculous healing qualities to the 
fact that it was foiind growing on Mount Calvary, when Jesus was 
crucified. For hundreds of years it was woven into bridal 
wreathes and was thought to bring luck to the bride and to help 
in regaining lost affections . 

"With so many powers ascribed to it, it is no wonder that 
through the centuries it has merited such names as herb of the 
cross, holy herb, enchanter's plant, lightning plant, simpler' s 
Joy and many others. Even today this plant is gathered by 
French peasants, 'when the moon is right', for important cures. 
Bees delight in the dainty little lavender refreshment stands 
the vervain keeps in the sun for their entertainment, and feast 
on the nectar and pollen," Skinner (19II) says that boiling 
gun- flints with rue and vervain was supposed to ensure that the 
shot would reach the intended victim, no matter how poor the aim. 

Hegi (1927) says: " Verbena officinalis ist ein alter Kultur- 
begleiter, dessen urspmtnglic he Heimat unsicher ist, aber rait 
grosser V/ahrscheinlichkeit im Mittelmeergebiete liegt. Die 
pflanze genoss vermutlich bei den antiker VOlker eine grosse Ver- 
ehrung. Nach Plinivis war kein Kraut im Altertum berttmter. Im 
alten Aegypten hiess die Pflanze die 'TrJCne der Isis'. Sie wurde 
bei feierlichen Gelegenheiten verbrannt (H^rba sancta) und gait 
als das beste Wundkraut bei Venrandungen durch eiseme Waff en. 
Die Hippokratiker empfahlen sie gegen Unfruchtbarkeit. Auch wur- 
de sie gegen Blasensteine, sowie auch als allgemeines Volksmittel 
bentttzt. In Griechenland gilt sie noch gegenwartig als Gltfcks- 
pflanze. Wann sie nach Mitteleuropa gelangt ist, ist unbekannt. 
Sie soil aber schon bei den Kelten und Germanen in hohem Rufe ge- 
standen haben. Der heiligen Hildegard vrar sie als Heilpflanze 



22U PHYTOLOGIA Vol. 10, no. 3 

bekannt. Von Thai wird sie in der Hatrzflora (1577) aufgeftthrt, 
von Jungermann fttr Franken (l6l5) . In dem in L\md aufbewahrten 
Herbar von Rostius (1610) liegt sie unter dem Namen Sideritis. 
Frtfher wurde sie viel in GSCrten zu medizinischen Zwecken gebaut 
und die Herba Verb^nae s . H. Golumbiriae s . H. sanguina3J.s als 
schleimhaltiges, zusammenziehendes, bit teres Mittel, femer bei 
Wechselfi'eber, Steinleiden und AugenentztCndungen angewendet. 
Femer dienen sie mit Zimmt, Nelken, Vanille und bitteren Mandeln 
gemlscht als Ersatz ftlr chinesischen Tee, femer als Aufgruss (3 
g. auf 1/U Liter Wasser) gegen Kopfschmerzen, Migrane, allgeraeine 
Schwache tmsw. Kneipp empfiehlt diesen Tee auch gegen Keuchhus- 
ten, Nieren- und Leberleiden, Wasser- und Gelbsucht. Ein frtther 
viel bentttzter Haustee wurde aus 100 g Eisenkraut, 10 g Pfeffer- 
minze, 2 g Thymian, 2 g Majoran, 5 g Zimmt imd 1 g Muskatbltfte 
gemischt. Die HomOopathie bereitet auS der bltlhenden Pflanze 
eine Essenz. Gurken sollen beim Einmachen durch Zusatz von Eiser*- 
krautwurzel schmackhaf tiger werden. Chemisch wurde neben Gerb- 
stoffen, Bitterstoff, Invertin, Emulsin usw. das nicht giftige 
Gykosid Verbenalin nachgewiesen, das sich in der frischen Pflanze 
in Form von Kristallnadeln findet, beim Trocknen aber zum Teil 
verschwindet . — Die Verbena (verbenaca, hierobotane) spielte im 
antiken Aberglauben eine grosse RoUe. Nach Plinius (Nat. hist. 
2$, 105 ff .) behaupteten die Magier, dass man, wenn man sich mit 
der Pflanze salbe, alles erlange, was man wolle, dass sie das 
Fieber vertreibe, Freundschaft erwerbe und alle mOglichen Krsuik- 
heiten heile, dass man sie beim Auf gang des Sirius sammeln mtfsse 
und zwar so, dass es weder der Mond noch die Sonne sehe usw. Ob 
allerdings unter der antiken verbena unsere Verbena officinalis 
zu verstehen ist, bleibt recht zweifelhaft. Unter 'verbenae' 
verstanden die ROhier ganz allgemein die KrJtuter \ind Baumzweige, 
deren man sich bei Opfem und anderen Kulthandlungen bediente. 
Fussend auf den antiken Berichten Quellen erscheint die verbena 
auch im Aberglauben des deutschen Mittelalters . So sagt der 
Regensburger Domheer Konrad von Kegenberg in seinem 'Buch der 
Natur' (Mitte des lU. Jahrhunderts), dass das Eisenkraut 'den 
zaubraeren gar ntftz' sein. Auch Bock benerkt in seineiu KTctuter- 
buch (l55l): 'Das Kraut Verbena wtlrt noch heuttigs tags mehr zu 
der Zauberei dann zu der Artznei gesammelt.' In mittelalterlich- 
en Handschriften sind \ins verschiedene BeschwtJrungen der Verbena 
erhalten. Im heutigen Aberglauben ist das Eisenkraut so gut vie 
vergessen. Im Anhaltischen gehOrt es noch vor einigen Jahrzehn- 
ten zu den Pflanzen, die das Gewitter abwehren sollten. Man 
warf es zusamraen mit Hartenau ( I^ericum perforatm) bei einem 
aufziehenden Gewitter in das Feuer und sprach: 'Eisenhart tind 
Hartenau — Brennt an, dass sich das Wetter stau.' Im deutschen 
Volksaberglauben war das Eisenkraut wohl nie wirklich volksttfm^ 
lich. Durch die Kleriker \md Aerzte des Mittelalters war zwar 
Zeitweise manches tfber die geheimnisvollen Eigenschaften des 
Eisenkrauts ins Volk gedrungen, aber es blieb dort nicht haften. 

"Verbena officinalis ist zieailich warmebedUrftig und fehlt 
infolgedessen im Norden von Deutschland und in den subalpinen 
TSlem der Alpen fast ganz. Hingegen ist sie gegen BeschStdig- 



196U Moldenke, Monograph of Verbena 225 

ungen, vor allem gegen den Tritt von ITeidevieh und Menschen sehr 
unempfindlich xind findet sich meist gruppenweise in den BestSCnden 
von Lolium perenne vind polygonum aviculare, aug mageren, schwach 
gedrtlngten Weiden vom Festuca rubra — oder Brachypodium pinnatum 
— Sieglingia decumbena — Typus. Nicht selten erscheint sie auch 
an WegrJCndem mit Hordeum murinum und Sisymbrium officinale und 
S. Sophia . Auch aiif starker gedtfngten BOden pflegt sie nicht 
auszubleiben und ist dann mit Chenopodium- Arten, Lepidium draba, 
Potentilla supina, Ballota nigra , Aethusa cynapim, Dipsacus sil- 
vestris. Chrysanthemum parthenium usw, anzutreffen. In Rotklee- 
Stckem ist diese Pflanze in llitteleuropa selten zu finden, ob- 
gleich ihre Samen mit franzOsischen Rotkleesamen hSCufig einge- 
ftfhrt werden, — Die kleinen von Apiden, Tagfaltem, Fliegen usw. 
besuchten Bltften bergen im Grunde den von der Unterlage des 
Fruchtknotens abgeschiedenen Nektar und besitzen einen Ring von 
nach vom zusammenneigender Haare. Steckt das besuchende Insekt 
den Rtlssel in die KronenrOhre, so streift dieser zunachst zwisch- 
en den Staubbeuteln vorbei und trifft erst dann aiif die papillose 
FlStche der tiefer stehenden Narbe. Da aber die Risse der Staub- 
beutel nach abwarts gerichtet sind, so -nird er sich zunSchst 
nicht mit Pollen behaften. Dies tritt vielmehr in der Regel erst 
beim zurttckziehen des Rttssels ein, weil dann eine Drehimg der 
Staubbeutel bewirkt wird. In der Regel erfolgt also Fremdbe- 
staubung. Doch stehen die beiden unteren Staubbeutel der Narbe 
so nahe, dass auch mit einer selbsttatigen Selbstbestaubung zu 
rechnen ist, die anscheinend von vollem Erfolge begleitet ist. 
Gegentlich terten Bltlten auf, in denen nur 2 StaubblSttter vorhan- 
den sind. Zur Fruchtzeit ist der Kelch schrSg aufwarts gerichtet 
und gestattet dadurch ein Ausschtltteln durch den TTind. Auch vor- 
beistreichende Tiere kOnnen dadurch, das sich die etwas eiiiwarts 
gekrtlramten Kelchblattspitzen an ihrem Felle reiben, zur Ausstreu- 
ung beitragen. Als Schmarotzer treten Didymella effusa Niessl und 
D. rehmii (Kze,), Erysibe cichoriacearum DC, Mollisia verbenae 
(Opiz), Ophiobolus cesatianus (Mont.) usw. auf. 

"Der Name Eisenkraut (althochdeutsch Isama [erf^anze chrflt] 
mittelhochdeutsch Isenkrflt) ist zunachst eine Uebersetzung des 
griechischen sideritis (Pflanzenname bei Dioskuridis; grtech. 
sideros = Eisen) . Er ist in vielen Gegenden dem Volk wohlbe- 
kaiint und dttrfte hauptsSchlich durch Fachkundige (Aerzte, Apo- 
theker) bzw. durch Bttcher dahin gelangt sein. Iserhark [Eisen- 
hart] (MecklenbTirg), Eisenhindrik, iserene Hendrek, Iren Hendrek 
[=eisemer Heinrich, vgl. Pflanzennamen wie guter Heinrich, stol- 
zer Heinrich usw.] (GOttingen), Eisick (Nsihagebiet) ." 

Pulteney (1790) says: "In the meantime, in tracing the origin 
and progress of botanical science in Britain, a survey of its 
state in the druidical times, ought to claim first attention; but 
in fact, the little information transmitted to us fi*om the an- 
tients relating to this extraordinary sect, being almost yrtiolly 
confined to Caesar and Pliny, precludes any enlarged view respect- 
ing my particular object It is from Pliny we learn that the 



226 PHITOLOGIA Vol. 10, no. 3 

mistletoe , the vervain , the selago and the samolus , these antient 
fathers of druidism attributed efficacies almost divine; and or- 
dained the collection, and administration of them, with rites and 
ceremonies, not short of religious strictness, and such as coun- 
tenanced the grossest superstition The vervain , after previoxs 

libations of honey, •na.s to be gathered at the rising of the dog- 
star; when neither sun nor moon shone; with the left hand only; 
after describing a circle around the plant &c; and thus prepared, 
it banished fevers, and other distempers; was an antidote to the 
bite of serpents, and a charm to conciliate friendship (from 
Pliny, Lib. xxv. c. 9)..... With respect to this herb, the hiero - 
botane, the sacra herba of Dioscorides, although the modem bot- 
anists have not agreed to confine the term to verbena , which Pliny 
has described, as having narrower and smaller leaves than the oak, 
it may be remarked, that there has been a diversity of opinions 
among the commentators, relating to the plant and it is acknow- 
ledged that verbena or verbenacea, vra.s also applied, as a general 
term for all plants used about the altar in sacrifices. To this 
day the Tuscans apply the word vervena to slips, shoots, suckers 
or bunoles of plants of any kind." 

Webster (19l|2) says iTrtiy this pale-flowered weedy herb ever 
became so imbued with magical virtues is a mystery. The Dniids 
revered it. Perhaps the styptic tannic principle found in its 
stems and leaves bears out the legend, cited by Mrs. Grieves in 
her Herbal, that the herb was found on the Mount of Calvary and 
used to staunch the wounds of the Saviour. At any rate, as a 
very sacred and old-world herb, tuck it away in some corner of 
your mediaeval garden, the only place where it belongs." 

As to its supposed medicinal properties, Haller (1768) has 
this to say: "Inodora planta, & fere insipida: destillantem . , . . 
dat ad agnem spiritim acidum, oleum emKn^-eumaticum, & salem ali- 
quem colatilem: in cinere fixus & salsus sal est, non alcalinus. 
Febres intermittentes succi spissati drachmam bie de die sumtam 

sanare , aut certe pemviani corticis potestatem eo remedio 

adjuvari lego. Balsamum, cum oleo paratim, arcanum est antarth- 

riticum Fulvii Testi Decoctum cum spititu vini mistum glutem 

oculos connectens dissipare dicitur. Ad narium haemorrhagiam 

veteres adhibebant. In cataplasmate, cum rhodia radice tri- 

tam dolores capitis sanare, & solam, amuleti modo , eos 

dolores sedure, inque pleuritide fot\m prodesse Mihi non vi- 
de tur ferio in medicinam recipi Ad Verbenam n. 219. in convul- 

sionibus infantxim saepe appensum nihil praestitisse Rosen. Kinder 
Krankh. p. 79." 

Vilmorin-Andrieux report that "Les feuilles entrent dans cer- 
taines preparations." Watt (1893) states that "In the Pan jab the 
fresh leaves are considered febrifuge and tonic, and is said to 
be used as a rubefacient in rheimiatism and diseases of the joints. 
The root is believed to be a remedy for scrofula and snake-bite . 
At one time it was worn in Europe as a charm against evil, and 
for good luck. In Tuscany it is said to be still employed as a 
poultice for liver complaints, and taken internally for the same 



196U Moldenke, Monograph of Verbena 227 

disease, and for dropsy." 

UacKay (I836) speaks of the name. Verbena , with an origin in 
"ferfaen in Celtis, derived from fer to drive away and faen , a 
stone, from having been supposed to cure the complaint so called 
Theis." He continues: "The genus Verbena is placed by Sir James 
Edward Smith in the first order of the class Didynamia, but as 
Doctor Hooker remarks, it does not naturally rank there, being 
considerably different in the structure of its germen and fruit. 
This herb has scarcely any aromatic or other sensible quality. 
The root worn about the neck vrith a string, is an old superstit- 
ious remedy, or charm for the King's Evil [=scrofula] ." 

Ichekama tells us that the species is regarded as medicinal in 
Japan, too, while McClure says that on Hainan Island it is used 
in the treatment of dog bites. Roig y Mesa (19U5) avers that it 
is the leaves and floriferous tips of the stems that are used 
medicinally in Cuba. According to Pamo "La Verbena oficinal fu5 
considerada ccmo astringente, resolutiva, diafor^tica, antiespas- 
m6dica, vulneraria, etc. En la actualidad no se usa." In Eritrea 
"Las hojas se usan contra la inflamaci6n de los gldndulas del 
cuello." Paris (I963) states that it contains "verbenalin". Rob- 
ledo is quoted as stating that the plant contains "verbenalina", 
which "tiena sabor amargo y se eraplea como vulnerario." Caiflas 
reports that in Cuba "Las hojas y sumidades de sabor amargo y 
astringente, se aplican sobre la piel, machacadas y en cataplasmas 
como vulnerarias. La infusi6n de las flores (ii5 gramos por litro 
de agua), se emplea para los dolores de cabeza." 

Pliny, who died in Pompeii in 79 A.D., tells us that no plant 
was as highly esteemed by the Romans as this one; it was used to 
decorate Jupiter's altars, to cleanse houses, to guard against bad 
luck, and (according to Marzell [1930]) "mit der \insere Gesandten 
nach den Feinden gehen." P'ei (19U7) says that in China it is 
"used by natives as a remedy for inflammation of the intestines." 
He records it from Sikang. Jyh Ho Chen, in a letter to me dated 
August 16, I9U8, reports that Chinese doctors use V. officinalis 
in the treatment of malaria, and that the very bitter juice proves 
"very effective and successful." Perez Arbelaez (1937) states 
that the dosage is 10 — 20 grams in 200 grams of water in the 
treatment of fevers associated with amoebic dysentery and typhoid, 
administered as a decoction in enemas and tea, and that it is used 
with efficiency in this way at Bogota, Colombia. 

Loureiro (1790) says: "Virtus. Nervina, Deobstruens. Decoctum 
prodest in hydrope, cataplasma in tumore scroti." Connor & Adams 
(1951) report that "Vervain has been suspected of poisoning 
cattle in Australia on one occasion, but no poisonings have been 
reported in New Zealand.. . .Vervain contains a glycoside, verbenalin, 
vrtiich appears to be veiy mildly toxic." Kuwajima (1939) describes a 
new glycoside, verbenin, but, according to Winde, Echaust, & HfiCnsel 

(1961) "Das von Kuwajima als ein neue Glykosid der Verbena offio- 

inalis beschriebene Verbenin konnte in der Droge nicht nachgewiesen 
werden. Die Vermutung Breitwiesers (19i;2) wurde nunmehr experimen- 
tell belegt, dass Verbenin eine Kristallmodifikation des Verbenalins 



228 P H Y T L G I A Vol. 10, no. 3 

darstellt." Hocking (19$$) states that it contains a bitter glu- 
coside called verbenalin, a neutral bitter principle tannin, and 
is used medicinally as a bitter tonic, astringent, and vulnerary 
in asthma and kidney disturbances; the root being extremely use- 
ful for liver and gallbladder disturbances, and the leaves used 
as a tea substitute. 

Coon (1963) says "The constituent which brings Verbena into the 
medical field is a bitter glucoside and tannin, a simple infusion 
(2 teaspoons to 1 pint) being employed as a diaphoretic, tonic, 
and expectorant. There are, in herbal literature, no strong claims 
made for its efficacy.. ..JLn exploration of the story of vervain 
leads us dovm some ancient avenues and provides an explanation for 
belief in the efficacy of the plant in herbal medicine. This plant 
was first used by the Romans. They gave us the name 'verbena', 
which to them meant any one of a number of plants used in sacri- 
fices, purgation and supplications. Finally the name vras attached 
to one particular plant and the virtues ascribed to Verbena by the 
Romans were passed along through the centuries until, in the lliddle 
Ages, it was said to have been a plant v^iich, growing on the Mount 
of Cal"<ra.ry, staunched the wounds of the Savior. The transferral 
of virtues from pagan to Christian (it has happened in our Christ- 
mas celebrations) was not unusual and verbena early became one of 
the holy herbs associated with St. John. Pliny said 'if the din- 
ing chamber be sprinkled in water in which the herb Verbena has 
been steeped, the guests will be merrier.' Such a story led to 
the belief in its efficacy as a cure against the plague, and as a 
remedy for almost anything. It even had supernatural powers. Sev- 
eral Welsh names have meanings such as 'devils' hate' and 'enchant- 
ment herb . ' " 

Jaeger (I96I) reports that it produces pollen abundantly from 
7 to 11:30 a.m. and in small quantities to 2 pjn. only. 

Of very great interest in any discussion of Verbena officinalis 
is the problem of a possible hybrid between it and the scrophular- 
iaceous Veronica maritima L. This hybrid is now known as xVeroni- 
cena haartmani Moldenke. Roberts (1929) sums up the situation as 
follows: "On November 23, 1751, appeared a discussion, included in 
the Amoenitates Academicae (vol. 3, pp. 28 — $2, 176U) by another 
of Linnaeus' pupils, Johannes Haartman, entitled Plantae Hybridae. 
This discussion upon hybrid plants is to be noted, insofar as it 

reflects the views of Linnaeus and his school on the subject 

Veronica maritima (o) x Verbena officinalis (<?) is described in 
the greatest detail (p. 35), and is illustrated. (8b, pi. 11), 
This natural hybidd is reported as having been produced in the Bot- 
anical garden at Upsala in 1750. The statement is made 'neque 
longe ab his lecta est haec nostra planta [S] , quae antea nulli 
Botanica visa est' (p. 35). The vegetative and flower characters 
are described in some detail. The hybrid was perennial, bloomed 
annually, and was multiplied easily by the roots, but had no fruit 

('nullos vero fructus maturat') (p. 36) The description of 

the plant is as follows: In height, hoary color of the stem and 
leaves, smoothness of the stem, structure of the spike, and color 



1961; Moldenke, Monograph of Verbeiia 229 

of the corolla, the plant is stated to resemble the Veronica fe- 
male parent. If the flowers and their color and the roundness of 
the stem were emitted, 'the most vacuate botanist would have con- 
sidered it to be Verbena itself' (p. 35) J the leaves of the hy- 
brid are said to have had 'exactly the same singular division, 
with deeply furrowed lobes' (p. 35) • The flowers are stated to 
have been smaller than those of the female parent, and not larger 
than the flowers of Verbena; the leaves 'sometimes in threes, as 
in the Q but more often opposite, as in the J' . Although the 
plant flowered annually, it was sterile, and bore no fruit, but 
was perennial and multiplied by the roots. 'Floruit quidem haec 
planta cmni anno felicissime, in annum quo haec edimus, 1755, et 
vivis radicibus facillime immutata multiplicatiir, nullos vero 
fructus maturavit' (p. 35) • It thus appears that Linnaeus' hy- 
brid Veronica , originating in 1750, was still alive in 1755." 

Hooker (I836) cites T_^ Drummond 252 from New Orleans, Orleans 
Parish, Louisiana, for the typical form of V. officinalis and T. 
Drunimond s.n. [New Orleans, 1833] as his var. spuria, deposited 
in the herbariimi of the Royal Botanic Gardens at Kew. 

Schauer (18U7) cites Wallich 1825 and Edgeworth s.n. from Ne- 
pal; R, Brown s.n. and Lhotsky s.n. from eastern Australia; Bov6 
s.n. from "Africae ora septentr."; Herb. De Candolle s.n. frcan 
the Canary Islands; Bergius s.n. from the Cape of Good Hope; Raf - 
inesque s.n. frcan "America septentr."; Beyrich s.n. from Georgia; 
Humboldt & Bonpland s.n. , Schiede II67 , and Berlandier 159 from 
Mexico; Sagra s.n. from Havana, Cuba; Sellow s.n. from "Brasilia 
merid."; and Sellow s.n. from Buenos Aires, Argentina. He notes 
that Schimper lh$ differs in being "demissa, ramis procumbentibus, 
spica subglandulosa" . I suspect that the last^nentioned is var, 
prostrata Gren. &, Godr. 

Richard (l85l) cites Quartin-Dillon s.n. [ChirS, Aug.] and 
Schimper s.n. [Adoua, Mai] from Abyssinia. Ascherson (1867) 
cites cienkowsky s.n. [Roseres, Senaar, 9 Mai I8U8] and Schimper 
s.n. [Dehli-Dikeno, 23 Oct. 185U] , while Engler (1892) cites Hil - 
debrandt Ui5, Schimper I.lh5 , 7, '-c 28U , and Rohlfs & Stecker s.n. 
from Abyssinia. Bentham (I87O) records the species frcm Victoria, 
Australia. 

Baker & Stapf (1900) cite the following: ERITREA: Hildebrandt 
Ui5; Schweinfurth & Riva lll6 ;' Steudner I30U . ABYSSINIA; Schimp- 
er 7, ll;5, & 28U ; Quartin-Dillon s.n. [Shire] ; Rohlfs & Stecker 
s^nT [Godofelassi] . SUDAN: Senner: Cienkowsky s.n. KENIA: Scott - 
Elliot 7800 . BRITISH SOMALILAND: Revoil s.n. Pearson (I9OI) 
cites the following: REPUBLIC OF SOUTH AFRICA: Cape Province: 
Burchell 503 ; Pappe s.n. [near Rondebosch]; ¥olley Dod k92 . Natal: 
Dr^e s.n. [Bashee River]; Gerrard 12^9 ; Haygarth li73 [Herb. Wood 
I96U] ; Krauss l5l ; Sanderson 92; Tyson 2105 . Transvaal; Burke 59 
Holub s.n. [Linokana]; Wilms 1175 &^ 1175a. Province undetermined; 



230 PHYTOLOGIA Vol. 10, no. 3 

Harvey kO$ ; Miller s.n. ; Zeyher 136U. 

Domin (1928) cites A. Dietrich 1^00 and Domin s.n. [II .1910] & 
s.n. [III. 1910] from Queensland. Dop (1935) cites the following: 
INDO-CHINA: Annam: Chevalier s.n. [Lang-bian]j Couderc s.n. [HuS]; 
E^^^ard s.n. [Dalat] . Laos: Spire s.n. ; Thorel s.n. Tonkin: Bal- 
ansa s.n. [Haiphong]; Bois s.n. [Dong-dang]; Chevalier s.n. [Nam- 
dinh], s.n. [Phu-tho]; Clemens s.n. [Haiphong]; Couderc s.n. 
[Hanoi]; Demange s.n. [Hanoi]; Duport s.n. [Cho-ganh]; Eberhardt 
s.n. [Thuyen-quang] ; Hautefeuille s.n. [Chapa] ; Mouret s.n. [Nam- 
dinh] ; P^telot s.n. [Cho-ganh] ; Simond s .n. [vers Lcng-tcheou] . 

Rozeira (19iUl) cites the following: PORTUGAL: Coul. s.n. [Bra- 
gan(ja]; Ferreira s.n. [MurQa] ; Henriq. s.n. [Sedeilos]; M_. Lopes- 
s.n. [Vimioso]; Mariz s.n. [SantulhSo]; Rozeira s.n. [Amedo] , s. 
n. [Covas-do-Douro] . 

The pollen-grain description given by Nair & Rehman (1962) is 
on the basis of a specimen collected at Hastanapur, India — 
"N[ational] B[otanic] G[arden] Ui559; Sl[ide] 2725." 

Perry (1933) reduces V. doming ens is Urb. to synonymy iinder V. 
officinalis , noting that "On the whole, the specimens from Santo 
Domingo and Cuba differ from the typical V. officinalis in their 
slender and more elongate habit; the inflorescence is scarcely as 
glandular, the flowers are smaller, ana the nutlets often do not 
exceed 1.5 mm. in length. Nevertheless, the Cuban specimens vary 
greatly in size, and Curtiss 677 is hardly separable from typical 
V. officinalis . Since many of the specimens are rather poor, it 
appears probable that they may very well represent an impoverish- 
ed condition. Urban himself was somewhat uncertain of the status 
of his species as he appended the following note in a later pub- 
lication: 'An re vera a forma V. officinalis L. separanda? • " She 
cites, in addition to 15 specimens of Cuban and Hispaniolan col- 
lections cited hereinbefore by me under V. domingensis , the fol- 
lowing U2 specimens not as yet seen by me: MASSACHUSETTS: Essex 
Co.: Oakes s.n. [Rowley] (G) . RHODE ISLAND: Kent Co.: Thurber s. 
n. [T/arwick Neck, 18U8] (G) . NEV/ YORK: Kings Co.: Schrenk s.n. 
[Brooklyn, 13 Sept. 1879] (E) . NE\T JERSEY: Camden Co.: Parker 3_. 
n. [ballast, Camden, 30 Aug. 187U] (G), s.n. [Longacoming, 23 
July 1867] (G). Cape May Co.: Gershoy 583 (G) . PENNSYLVAinJL : 
Dauphin Co.: Porter s.n. [banks of the Susquehanna, Harrisburg, 
Oct. 1852] (G). Lancaster Co.: Porter s.n. [Lancaster, 21 Aug. 
1861] (G). Philadelphia Co.: Lea s.n. TPhlladelphia, 18U;] (E) . 
York Co.: MacSlwee 873 (E) . DELAWARE: New Castle Co.: Tatnall £. 
n. [Wilmington, 18U5] (G) . DISTRICT OF COLUMBIA: Yfard s.n. [vic- 
inity of Washington, 23 June 1878] (E) . VIRGINIA: Accomac Co.: 
Norton s.n. [Parksley, 11 Sept. 1902] (E) . Bedford Co.: Curtiss 
s.n. [30 June 1870] (E) . Sm^rth Co.: £. I£. Small s.n. [Marion, 
1892] (E). NORTH CAROLINA: Avery Co.: vaslizenus 121U (E) . Bladen 
Co.: Biltniore Herb. U762 (G, N) . Ciirrituck Co.; RsESolph 587 (G) . 



I96I1 Moldenke, Monograph of Verbena 231 

Washington Co.: Randolph 6It5 (G) . Counly undetermined: Curtis 3. 
ru (E). SOUTH CAROLINA.: Aiken Co.: Eggert s.n. [streets of 
Graniteville, 23 May 1899] (E) . GEORGIA: Floyd Co.: Chapman s.n. 
[Rome] (E). FLORIDA: County undetermined: Rugel 121 (E, F) . ALA- 
BAMA: DeKalb Co.: Eggert s.n. [Collinsville, 29 July 1897] (E) . 
Etowah Co.: Eggert s.n. [Attala, 9 July I898] (E) . TEMNESSEE: 
Carroll Co.: Eggert s.n. [Hollow Rock, $ Aug. 1897] (E, F) . Car- 
ter Co.: Small & Heller U8U (E, G) . Knox Co.: Ruth s.n. [Knox- 
ville, July 1893] (E) . LOUISIANA: Plaquemines Par.: Tracy h. Lloyd 
s.n. [Port Eads, 22 Aug. I9OG] (E, G, \\) , BERMUDA ISLAIIDS: Main: 
S. Brown U92 (D, G); Brown h Britton 28 (D, G); F. S. Collins 267 
(G); A.. H. Moore 2939a (G) . St. Georges: Robinson 113 (G) . 

In all, 111(2 herbarium specimens, including the types of sever- 
al of the names involved, and 27 mounted photographs and illustra- 
tions have been examined by me. 

Citations: MASSACHUSETTS: Martha's Vineyard: E. P. Bicknell 
7353 (N). CONNECTICUT: Fairfield Co.: H. C^ Beardslee s.n. [Hun- 
tington] (Ob — $08lii) . Hartford Co.: M. Brandegee s.n. [Berlin] 
(Ga — U68I23) . Middlesex Co.: M. Brandegee s.n. [Middletown] (Ca- 
U68I23) . New Haven Co.: W^ W. Denslow s.n. [Sept. I863] (Ms) . 
NEW YORK: Bronx Co.: A. Brown s.n. [Hunter's Point, Atig. 10, 
1879] (Tc); H. N. Moldenke 20^62 (B, F, Fy, Hw, Le, Lm, N, Rs, S, 
Ss, Ug); _J. Schrenk s.n. [Hunter's Point, Aug. 2, 1879] (C) . Hei>- 
kimer Co.: Collector undesignated s.n. [July 2, 1852] (N). Sche- 
nectady Co.: Tuckerman 7U8 (Al) . Suffolk Co.: Leggett s.n. [Mil- 
ler Place, Aug. I86I] (Tc, Tc); E. S. Miller s.n. [Mt. Sinai, Aug. 
29, 1873] (N). Ulster Co.: Halsey s.n. [Kingston, 1820] (C). 
County undetermined: Miller s.n. [Long Island] (Ka); C. H. Peck 
s.n. (Al)j £. Torrey s.n. (C, Pa) . NE7/ JERSEY: Atlantic~Co . : Rai 
3 (In — IOU25). Camden Co.: Beringer s.n. [Camden, Aug. I889] 
(Mi)j Martindale 3.n. [Camden, Sep. I876] (Pu); M. W. Twaddell £. 
n. [August 3, 1878] (Up). Cape May Co.: K^ K. Mackenzie s.n. 
[Cape May, Sept. 25, 1920] (H— 51908, N); W. Stone 15810 (Up), 
16317 (Up). Gloucester Co.: Brinton s.n. JWilliamstown, June 25, 
1889] (Ca~25l86). Hudson Co.: Billberg s.n. [Hoboken, 1826] (S); 
J. Schrenk s.n. [Jersey City, July 9, 1879] (Tc). Coxonty vmde- 
ternined: E. Pur and s.n. [Hew Jersey] (Ky)j R. E. Griffith s.n. 
[New Jersey, Julio] (Ky); Herb. Torrey s.n. (T); J_. Torrey 3 ji. 
[1835] (Br). PE^raSYLVANIA: Delaware Co.: Fogg 5695 (Up). Phila- 
delphia Co.: R. C. Alexander s.n. [Philadelphia] (Ca — 379997)j I_. 
Burk s.n. [Raighs Point] (Up— 17111); Meredith s.n. [Oct. 9, 
1920] (N); A. H^ Smith s.n. [Navy Yard, July 1866] (Up— 17112) . 
York Co.: MacElwee 753 (Um — 2li), s.n. [York Furnace, June I896] 
(Ka — 7II8U); W. Stone 2166 (Up). County undetermined: A. P_. Gar - 
ber s.n. [August I868] (S) . DELAWARE: New Castle Co.: W. M. Can- 



232 P H Y T L G I A Vol. 10, no. 3 

by i960 (Dt)j Morong 3.n« [Wilmington, Sept. 2, 1873] (Be). Sus- 
sex Co.: Commons 7 (N) . MARILA.ND: Baltimore Co.: Boldo 92 (Q)j 
LeRoy s.n. [Baltimore, I866] (C, Du— 123l|88, Gg— 31U03, Ms, N); 
K. A. Taylor s.n. [July 9, «91] (Ur) . Cecil Co.: Brinton s.n. 
[North East, July 20, I89O] (Up— 1711ii) . Dorchester Co.: £. P. 
Smith 293$ (I, I). Saint Mary's Co.: O'Neill s.n. [June 13, 
1930] (I). Somerset Co.: J. H. Holmes 76 (W) . Worcester Co.: C. 
E. Moldenke s.n. [H. N. Moldenke 66U8] (N); ^ ^ Rusby s.n. 
[Stockton, Aug. I889] (C, R) ; True 13 (Gu— 10U52, Up). County 
undetermined; Collector undetermined s.n. [W. D., Maryland] (Lu) . 
DISTRICT OF COLUMBIA: M. S. Bebb s.n. [1862] (N); J. W. Chicker - 
ing s.n. [6-2U-I873] (N); Kearney s.n. [J\me 20, 1897] (N, Z— 
drawing); B. R^ Patterson s.n. [July 9, '76] (Cm); Peck s.n. 
(Mi); E. S. Steele s.n. [July 21, I896] (Ka— 82^07, Ob~508l8, 
Ifin— 12). VIRGINIA: Accomac Co.: Wilkens $372' (Up). Montgomery 
Co.: Murrill s.n. [Blacksburg, 2k June 189$] (N) . Norfolk Co.: 
Meredith s.n. [Norfolk, June 2$, 19^] (If— 26197), s.n. [Ghent, 
June 28, I92U; Herb. Dreisbach 3003] (Mi, Mi). Princess Anne 
Co.: Ffemald & Long Ul$2 (Up); K. K. Mackexizie 1679 (Po— 267668). 
Smyth Co.: J. K. Small s.n. [Marion, June 29, 1892] (Ca— 10U82$), 
s.n. [Middle Fork, Holston River, Marion, July 6, 1892] (Ca — 
2018$, Fc, Ob— $0816, Up— 17116, W). Southampton Co.: A. A^ 
Heller 96^ (C, Up — 17117, W) . County undetermined: Collector 
undesignated s.n. [In Canada, Virginia] (Lu, S) ; Herb. U. S. 
Dept. Agr. s.n. (Fc). Chincoteague Island: H. A^ Gleason 8$^U8 
(N) . WEST VIRGINIA: Jefferson Co.: Guttenberg s.n. [Harper's 
Ferry, Aug. 2U, 18 78] (N) , s.n. [Harper's Ferry, Aug. 28, 18 78] 
(Vt), s.n. [Harper's Ferry, July 22nd] (Cm). NORTH CAROLINA: 
Bladen Co.: Biltmore Herb. 1;762 (N, W— 33210$) . Buncombe Co.: 
R. H. Ward s.n. [Asheville, Aug. 21, '77] (Ur) . Carteret Co.: 
Blomquist IO36O (H— 1|98$0). Currituck Co.: Randolph & Randolph 
$87 (Ba). H7de Co.: D. S. Correll 1771 (H~l;02l;2); Radford & 
Stewart 786 (Hi— 21$62). Iredell Co.: M. E. ^yams s.n. [states- 
ville, July I878] (W— lli7$83) . Jones Co.: A. E. Radford 37228 
(Hi— IOU887). Madison Co.: G. B. Grant 2926 (Po— 26763$); J. D. 
Smith s.n. [Warm Springs, July 27, 1830] (W— 132311U) . Mitchell 
Co.: Collector undesignated s.n. [Aug. 9] (Hi— $9U69). Polk Co.: 
E. £. Towns end 92 (Pi— 87I8O), s.n. [Columbus, May 31, 1897] (W- 
3la769). Stokes Co.: Small & Heller 1;8U , in part (Ca— 2$l87, Up- 
1711$, W), s.n. [near Hall's Store, Piedmont Springs, July 2, 
1891] (Fc). County undetermined: Ashe s.n. [Roan Mtn., July '93] 
(Hi— $9U78); McCarthy s.n. [No. Carolina, Aug. 188$] (Ba), s.n. 
[E. N. Carolina, Aug. 1887] (W— 218877) . SOUTH CAROLINA: Ander- 
son Co.: J, Davis 8U93 (W — I20U69U) . Darlington Co.: Collector 
Tindesignated s.n. [June lii, '11] (Hi — $9U6U) . County imdetermin- 



196U Moldenke, Monograph of Verbena 233 

ed: G. McCarthy s.n. [Sept. 1888] (Ka); Short s.n. (Pr) . GEORGIA: 
Rabun Co.: Cuthbert s.n. [Tallulah Falls, Aug. 5, 1899] (Fl— 
21115). Coiinty undetermined: B eyrie h s .n. (Br). FLORIDA: Escam- 
bia Co.: Curtiss s.n. [Pensaccla, summer 1885] (N) . County unde- 
termined :^^er~12ri'N, lff~5ll9U2). AUBAl'A: DeKalb Co.: Eggert 
s.n. [Collinsville, 29 July 1897] (Cm, 11, W— 75U256) . Mobile 
Co.: £. T_. Mohr s.n. [waste & cult, lands. Mobile, May 10, 1876] 
(VV~771863), s.n. [Mobile, July I883] (Mi), s.n. [July—Sept. 
I88U] (C), s.n. [ballast ground. Mobile, 5/30/1887] (W~77l862) . 
County undetermined: Buckley s.n. [Aug. 181|0] (Br, T). LOiSSISS- 
IPPI: Wilkinson Co.: Phares 1733 (VO . TEIIIJESSEE: Carroll Co.: 
Eggert s.n. [Hollow Rock, 5 August 1897] (N). Carter Co.: Small 
§L Heller IO6 (Ob~503l7), U8Ii, in part (Ka), s.n. [along banks of 
the Doe River, July 16~17, I89I] (Ba, Dt, lo— 2075U, Ok, W— 
298655). Knox Co.: Ruth s.n. [June 1893] (Se— 95716), s.n. [Knox- 
ville, July 1893] (Dt, Ob— 508l5) . LOUISIANA: Livingston Par.: 
Herb. Torrey s.n. [Albany] (T) . Orleans Par.; Gates s.n. [near 
New Orleans, 1829] (T). Plaquemines Par.: Tracy & Lloyd 17 (Cm, 
N, Tr, Up— 50898, W— 383530). NEW l^EXICO: Taos Co.: D. A. Johan - 
sen 600 (Gg— 19ii25l) . OREGON: Multnomah Co.: J. C_. Nelson 835 
(Du— 77653), USUI (Or— 17ii59); Suksdorf I89U (Pi— I38U02) . CALI- 
FORNIA: Amador Co.: Belshaw 2it5U (Ca~12UtOO); G. Hansen U77 (Du- 
2li213). San Diego Co.: H. P. Chandler 5122 (N); F. W. Johnson 
1665 (N); Orcutt & Bowne s.n. [San Diego, I88U] (C). MEXICO: 
Nuevo Le6n: F. W. Pennell 168 80 (D— 733979, Me, W— I6U0298). 
BERMUDA ISUNDS: Main: S. Brown U92 (N, W— 8U835il); Brown & Brit - 
ton 28 (N, Up— U5621, W— 52U78I); F. S_. Collins 267 (Cm, N, W— 
717561); 0. Degener s.n. [Hungry Bay, July 26, 1921] (Ba); A. H. 
Moore 2939a (Gg— 155395, Mi, N) . Island undetermined: Bailey , 
Bailey , Whetzel, Degener , ''. McCallan s.n. [July 26, 1921] (I); 0_. 
Degener I306 (N); Marble 791 (N) . BRAZIL: Rio de Janeiro: 0_. M. 
Barth 16 [Herb. Inst. 0. Cruz 233] (W— 23U2992) . BOLIVIA: Coch- 
abamba: Buchtien 2101; (N). CHILE: Arica: Buchtien 1;380 (W — 
1159367). Valdivia: Claude-Joseph 5U08 (W— lii70520) ; Hollermayer 
s.n. [Werdennann I9I6] (V/— 15U1196) ; Kuntcel 397 (Sm); Sparre 2262 
(S). Valparaiso: Behn s.n. [7 Decbr. 1920] (Ca— U986lli) . Teja 
Island: Hollermayer s.n. [V/erdormann I9I6] (E — 999029, N, N, S). 
TRISTAN DA CUNHA: Mejland 3UI (Bm, Go). AZORES ISLANDS: Faial: 
C. Sj, Brown 213 (W— 262279) . MADEIRA: N^ J. Anders son s.n. [Mad- 
eira, 1851] (S); GongalvBs da Costa s.n. [Alegria, Maio 1928] 
(Go); Hombeck s.n. [Madeira] (Cp) . CANARY ISLANDS: Gran Canaria: 
A. £. Cook 35 (N, W— 536157), 569 (Ca— 202072, Cm, Du— 81261, Gg— 
3IU07, Go, Mi, N, Ob— lli870. Or— lli239, Po— 220U39, Ur, W— 
5367U6); Kuntze s.n. [3I/XII/87] (N) . Gomera: W. M. A. Brooke U27 
(Bm) . Lanzarote: R. T. Lowe s.n. (Bm). Tenerife: Asplund lllt8 (S); 



23U PHYTOLOGIA Vol. 10, no. 3 

C_. B. Clarke 2018 (Br); Tullgren I9I; (S) . GREAT BRITAIN: England: 
J. Ball s.n. [South-Vifest part of Herefordshire, Aug. I81i8] (W — 
M2U55) ; Beeby s.n. [29.7.1888] (Go, S)j A^ Jj;_ Berggren s.n. [11. 
7.1920] (S); Britton L Britton 3.n. [Brighton, July 22, I888] (C), 
s.n. [Canterbury, Aug. 5, I888] (C)j R. Campbell s.n. [Hampton 
Courts, July 10, I89O] (Ito— 20ii79)i Collector undesignated s.n. 
[Freshwater Church, Sept. 1866] (Mi); M. Dehn 1 (N, N); Doubleday 
s.n. [Eppius] (C); Gentil s.n. [Teddington] (Br); P. J. Greemray 
31^0 (N); R. A^ Harper s.n. [Chisham, 9/21/1889] (Cm); Hairorth s_. 
n. (T)j Herb. Lemmon s.n. [Clifton, August 8th, I83I] (Ca— 329283, 
Ca--32928U); Herb. Lond. Univ. s.n. (W— 71960) ; Herb. Marie- 
Victorin s.n. [Croyden, 13 Oct, 18 31] (Vi); Hooker s.n. [Aug. 
1823] (Vi); Horvrood s.n. [23/8/36] (Go); J. H^ Lewis s.n. [22. 
VIII.I877] (U); 0. Lindblom s.n. [Aug. I929] (S); Linton s.n. 
[18.VII.1885] (S); Lomax s.n. [16 Augusti I89I] (Du— 90897, Ob— 
ll;871); R. M. Middleton Jr. s.n. [Oct. 18^9] (Mm— 15386); W. H. 
Painter s.n. [July I883] (S); Pease 8280 (Gg--3l501); Roekentz 
1217 (Br); C_. Skottsberg s.n. [11 Juli 1920] (Go); Tracke s.n. 
[Cornwall, 1832] (Vi), s.n. [near Helston, I836] (M); Turrill 3. 
n. [Keston, July 2, 1921] (Ba), s.n. [Oxford, 15 .VIII .1933] (S); 
T. Twining s.n. [near Kingston, 18U2] (Ca— 330213); L. F^ Ward 
s.n. [Corfe Castle, Aug. 18, I89U] (W— 229639); H. C. Watson s.n. 
(La) ; B. Welch 5217 (Go); £. W. White s.n. [Bank of Avon, JxlLj 
30, 1883] (Bl— li23U9), 3.n. [July 20, 1883] (Go); J. W. Wood s.n. 
[July I81a] (Ms). Scotland: B. James s.n. [1852] (Mm— 15387). 
Wales: A. E. Wade s.n. [July 1935] (Ms), s.n. [Aug. 1950] (Vi). 
EIRE: S^ A. Stewart 581|3 (Pr). SCILLI ISLANDS: Buller s.n. 
[Scilly Is.] (Wp). CHANNEL ISLANDS: Guernsey: R. Harvey s.n. 
[Uth July 1863] (Bl— i;235l). Jersey: L. Arsene li5U (Vi) . SWE- 
DEN: Agardh s.n. [Klagerup] (S); N. J^ Anders son s.n. [Skane, 
I8I16] (S); BjOrdlng s.n. [Lund, Augusti I88O] (N); C. Blom s.n. 
[lli/8/1936] (Go, S), s^n. [7/7/19U8] (S), s_^ [12/10/1952] (S); 
Cgster s.n. [I8/8/7I;] (S), s.n. [Aug. 18 7U] (Go, S), s.n. [1875] 
(S); Falck s.n. [Rrflunda, 7 Sept. I86I] (Go); Hallberg s.n. 
[GOteborg] (Go) ; Herb. Hort. Bot. Gothenburg s.n. [l8U0-talet] 
(Go); Herb. Linnaeus s.n. (F — photo, F — photo, N — photo, N — 
photo, S, Sg — photo, Sg — photo, Z — photo, Z — photo); Herb. Mas. 
Bot. Stockholm s.n. [I8UI] (S) ; Hjorth s.n. [August 1862] (Go); 
Kohler s.n. [1.IX.1921] (Go), s.n. [IX.1921] (Ew); Leche s.n. 
(S); Lilja s.n. [1855] (Go); Lindeberg s.n. [Aug. 1862] (Go);, 
Mortensen s.n. [11 Aug. I883] (Bl — li2350); Nordquist s.n. [Skane, 
1850] (S); Rasch s.n. [13/7/87] (Go); Ringirg s.n. (S); Ring- 
strand s.n. [Raflunda] (Go), s.n. (S); Sjoberg s.n. [1/7/1928] 
(S); SjPvall s.n. [September I88U] (Go); Sond6n s.n. [Tyckland, 
1908] (S); C. G. H. Thedenius 3.n. [Sept. 1898] (Ca— 206 308, Go); 



196U Moldenke, Monograph of Verbena 235 

K^ F. Thedenius s.n. [29/7/187U] (S); Visbig s.n. (Go); Wester- 
berg 3J1. [17/8/1872] (S); WinsloTT s.n. [GOteborg, 1872] (Go). 
BORNHODi ISLAND: M. Engstedt s.n. [U/8/1908] (S); KlPrker s.n. 
[25 Juli '83] (S)i Krok s.n. [Juli 1865] (S, S)j E. 0. F. I^rman 
s.n. [3/9/86] (S) . POLAND: Anderberg s.n. [m/7/1929] (Go); Dy- 
bowski 61i (S); Jungner s.n. [lii/10/1906] (S)j Nils son & Degelius 
s.n. [11^/7/1929] (S); Stendfinski s.n. [6.8.1872] (W— 2156791). 
DENMARK: Andersen s.n. [19/8/1895] (S, S); Christensen s.n. [2/ 
7/96] (S), 3.n. [28.VIII.1896] (Go)j poster s.n. [Juli 1868] (S)j 
M. Engstedt s.n. [U/8/1908] (Go, S) ; Ernsts en s.n. [1/7/68] (S, 
S); A. Hansen s.n. [31/8/98] (Go), s.n. (S); 0^ Hansen s.n. [10/ 
8/96] (S), 8.n. [10/10/97] (S); T. Holm s.n. [7/1881] (S); Johan - 
son s.n. [U Juli 1883] (S); Lorenzen s.n. [27/VII/1911] (S); Mor - 
tensen s.n. [21/7/1880] (S), s.n. [11 Aug. 1883] (S); H. F. Poul - 
sen s.n. [8.1919] (S); Rosenberg s.n. [KjOrup, 9/8/18U9] (S)j A. 
E. Thomsen s.n. [5/8/1873] (S), s.n. [Aug. 1873] (S), s.n. [27/7/ 
1908] (Hi--l8887U); Troiel s.n. [Haverup ved sorO, 2/8/1862] (S)j 
Virliz s.n, (N) j Visby s.n. [6/1853] (S); Vogel & J^rgensen s.n. 
[Fredensborg, 3/9/1876] (S). NETHERLANDS: Collector undesignated 
s.n. (I); De Mol s.n. (Bz — 2378U)j Goester s.n. [Schalkwijk] (Ba); 
Henrard & Tap s.n. [20 Juni 1917] (La); Jonker & Kramer 210 (We); 
Oudemans 501h (Br); Reclaire s.n. [Rotterdam, Juli I898] (Yf — 
1750319); Tap s.n. [Aug. 1913] (La). LUXEMBURG: Lundberg s.n. 
[23/9/1925] (Go). BELGIUM: Bamps s.n. [Hasselt, Aout I87I;] (Br); 
Bommer s.n. (Br); Busschodts s.n. [7 Juin I886] (Br); Coemans s. 
n. (Br); Collector undesignated 3787 (Ca — 9555iUi) ; Coemans & Coe - 
mans s.n. [31 Juillet I863] (Br), s.n. [lli-7-79] (Br), s.n. 
[AoUt 1896] (Br); Cr^pin 165 (Br), s.n. [22/7/66] (Br); De Wilde- 
man s.n. [Laeken, 1881] (Br); De Witte s.n. [26 Juillet 1923] 
(Br); Evrard s.n. [25 juillet I88U] (Br); Guns s.n. [Juillet 
1903] (Br), s.n. [16 juillet 192h] (Br); Hector 103 (Br); Henry 
s.n. [Namur, aoOt 1882] (Br); Herb. Hort. Brux. s.n. [16 juillet 
19 2U] (Ca~31l40U8), s.n. [2 aoflt 1927] (Ca— UO666U); Herb. Jard. 
Bot. Brux. s.n. [Blegny, 3 Juillet 1913] (Br), s.n. (Br, Br, Br); 
Lamabr^e 3787 (W— 2091277, W— 2319331); Laurent 109 (Br); Lejeune 
s.n. (Br, Br); Lejeune & Coart 165 (Br); Libertae s.n. (Br); Lcm - 
rabrge 3787 (Vi); Lundberg 3.n." T8/lX/1927] (Go); Martinis 3.n. 
[Juillet 1855] (Br); Mathieu s.n. [I8-8-I907] (Br, Br); Michiela 
s.n. [II/8/I938] (Vi); Nyst s.n. [Env. de Brux.] (Br), s.n. (Br); 
Pir^ 3.n. [AofJt 1873] (Br); Serclaes 9 (Br); Theux s.n. [Aotlt 
1908] (Br); Troch s.n. [Aoflt I883] (Br), s.n. [Juillet '90] (Br); 
Vandenbroeck s.n. [3I Juillet I878] (Br); Van Haesendonck s.n. 
(Br); Van Heurck h (Br); Vennoesen s.n. [Curango-Hasselt, 7-8- 
1920] (Br, Br), s.n. [27 Juillet 1922] (Br); Wathelet s.n. [15 



236 PHYTOLOGIA Vol. 10, no. 3 

Aoflt 1909] (Br). FRANCE: J. Ball s.n. [Paris, Oct. '39] (W— 
682Uii2); Beger s.n. [12 .VII .1916] (B); Billiet 1907 (Dm— 15) J 
Bonne s.n. [fin juiUet 1932] (It); Chamberet 25ll| (Vi)j S_. L. 
Clarke s.n. [Mt. Revard, Aug. 8, I898] (Lh)j Coomans & Coomans 
s.n. [Nancy] (Br); Copineau s.n. [Douellens, 7 Jul. '90] (W — 
202597)} Corbiere s.n. [Cherbourg, 17/8/1886] (W— 71990) ; Deize 
s.n. [Aofit 1896] (Du); L. Dempster 2i;02 (Ca— 179923); Dongg s.n. 
[Juillet 1927] (Br, Br); G. Een s.n. [Clermont, 2.7.1955] (S); L^ 
Engstedt s.n. [12/7/1937] (S); Erdmann s.n. [lU/7/10] (S, S), s.n. 
[I5/7/IO] (S); Francaville s.n. [Alnalit, Pyrenees] (S); Gautier 
s.n. [12 Juin I88O] (Du); Gombault s.n. [Pouan, 18 juillet 1917] 
(S); Grapengiesser s.n. [12/8/1926] (S); Hard av Seyerstad s.n. 
[IO/9/I9U7] (Go); Herb. W. H. Harvey s.n. [Liancourt] (Du--l66U63); 
Herb. Saldanha 2773 (Ja); Lassimonne 529 (W— 27U236); Lemie s.n. 
[Auch, 1922] (Bt— 33828); S. Lindman s.n. [23 Juli 1925] (S); 
Mougeau s.n. [Aofit i860] (Du) ; Nyman s.n. [7 Aug. i860] (S); gst- 
rand s.n. [25.6.1899] (S); Peyron s.n. [Juli I89I] (S, S); Puget 
s.n. [Annuy, Juillet 1353] (Vt) ; SegerstrOm s.n. [17/6/192U] (S); 
Tides trom 12985 (N, Y/— 155U891) , 13037 (I, W~l55U90U) , 13282 (I, 
N), 13736 (N); E. Wall 5, in part [29/725] (Ew) ; Zetterstedt s.n. 
[16/7/1856] (Go), s.n. [6/10/1856] (S). PORTUGAL: J. J. Barros 
s.n. [V.26] (B), s.n. [VI. 26] (B); Lemos 157 (Hi— 202055); Linde- 
roth s.n. [1858] (S), s.n. [Buzaco] (S); Rainha 2079 (W— 21899U3); 
Sjogren U53 (S) . SPAIN: Albo s.n. [Alrededores de Santa Elena, 
26.V.1933] (\7— 221338U) ; H. Deles sert s.n. [Valence, 1850] (Du— 
iSShhQ)} Knoche C.39 (Du); Kretzhmer 568 (S); Roivainen s.n. [17/ 
6/1950] (S), swi. [I950] (S), s£u [II/VI952] (Ca— 97085); Zet- 
terstedt s.n. [Barcugnas, 16/7/1856] (N, S). BALEARIC ISLANDS: 
Majorca: Knoche 2U96 (Du— 3983U3); SjOberg s.n. [Mallorca, 27/10/ 
1930] (S), s.n. [Mallorca, 7. XI. 1930] (S) . GERMANY: Benedicks 
s.n. [Gemrode] (S); C^ Billot 67 (S); Bommtlller s.n. [Leipzig, 
1878] (B), s.n. [Berka, IX.1897] (B), s.n. [12 .VIII .1911] (B), £. 
n. [Ettersberg] (B); Chalmot s.n. [GtJttingen] (W— 368114;) ; Col - 
lector undesignated I67 (Go), 172 (Ms), 226 (Du), s.n. (B); Deg- 
ener & Degener 23U05 (Ur); Dietrich 7^0 (Br); DtfU s.n. [I8i;3] 
(Br); Bmnert s.n. (Vi); Erdmsinn s.n. [Ik Juni I9IU] (S, S), s.n. 
[lii Juli I9II;] (Go); Erichsen s.n. [V7/86] (Go); Fahrenholtz s. 
n. [Anfang Septbr. I9OO] (Po— 63918, Vt, W— 9309U6); Felsmann s_. 
ru [2/8/86] (Go); Grapengiesser s.n. [I6/8/I929] (S, S); Gross 
s.n. [Brandenburg, 2U/6/1920] (Sp — 25790); Grossman s.n. [Leipzig, 
1896] (Mi); Guyot s.n. [Berlin, 7/33] (Pr); HafstrOm & HafstrOm 
s.n. [Wiesbaden, 1888] (S); Hase s.n. [ThUringen, I9O6] (B); He- 
arle 252 (Bt— Ii6379); Heiland 6 (La); Helledaij s.n. [2 Juli I886] 
(S, S); Herb. Calif. Acad. Scl. 31391 (Gg); Herb. Coll. Pharmacy 
s.n. (Pa); Herb. LandvrLrt . Hochsch. Berlin s.n. (B, B) . 



A CORRECTION 
L. T. Eiten 



In my article, "Egleria, a new genus of Cyperaceae from Brazil" 
(Phytologia, vol. 9, no. 8), I inadvBrtently used the words 
"holotype" and "isotype" when referring to the paratype collec- 
tion (page U82, below). These words, of course, should be used 
only for the type collection. What was meant was that duplicates 
of the paratype collection would be sent to the University of 
Brasilia and other herbaria throughout the world. 



BOOK REVIEWS 
Alma L. Moldenke 

"An Introduction to Plant Biology", by Dale C. Braungart and Ross 
H, Amett, Jr., Ull pp. C. V. Mosby Company, St. Louis, 
Missouri. 1962. $7.00 

The main advantages of this text are (a) an attractive and 
phylogenetic arrangement of 22 plant life cycles in a central 
separate section on green tinted paper and well illustrated by B. 
Melloni, (b) sound ecological and taxonomic approaches, (c) 
clear textual exposition, (d) effective use of photographs, espec- 
ially those frora the United States Department of Agriculture, and 
(e) useful bibliographer (which is given after each chapter), 
glossary and index. 

The main disadvantage is the conspicuous neglect of proper 
proofreading of galley and/or page proof resulting in the perpet- 
uation of authors' and/or printer's slips which might mislead the 
uninformed student and distress the critical one. On page UO we 
read that the "cell produces energy through respiration" instead 
of "releases"; page ii2 has "required a catalytic agent by a specif- 
ic enzyme" Triien the word "action" should have been substituted for 
"agent". On page U8 it is stated that "Fat is the most common 
plant food stored" and that maltose splits into 2 molecules of 
fructose only. Pages 229, 230 and part of 231 list taxonomic or- 
ders in ordinary type, while from there on they are given in bold- 
face type. In this whole section none of the family names are 
given in distinctive type as is to be expected in scientific lit- 
erature. The following terms are misspelled: nucleus on page UO, 
Chlamydomonadaceae , Euglenales, Euglenidae, Metazoa on page 79, 
Violales on page 226, Avicennia on pfiige 321, luxuriant on page 

237 



238 PHYTOLOGIA Vol. 10, no. 3 

337, and Vacciniiim on page 359. 

Other disadvantages are: insufficient explanation of anaerobic 
vs. aerobic respiration for many beginning students with poor 
secondary school chemistry backgrounds, absence of an electron 
microphotograph of a cell, no mention in late mitotic prophase of 
the replication of chromosomes, use of archaic mitosis diagrams, 
considering a reduction division a mitotic one on page 3U3 while 
giving its correct interpretation on psige 353, and preferring 
"desoxyribonucleic" for DNA. 



"Flowering Plants and Ferns of the Texas Coastal Bend Counties", 
by Fred B. Jones, Chester M. Rowell, Jr., and Marshall C. 
Johnston, 165 pp. Welder Series B-1, Welder Wildlife 
Foundation, Sinton, Texas. 1961. $2,35 

This fine botanical survey from this distinctive and little 
studied section of a huge state lists by scientific and common 
names almost all of the members of the rich floral composition 
to be found there. The flower colors, habitats and blooming 
times are added for each species. 

The covers are attractively illustrated, but the size of the 
book is awkwaixily and unnecessarily large and the spiral bind- 
ing is frail and also awkward. A map of the area covered would 
have been helpful. The specific epithet in Verbena cloverae is 
misspelled on page llxhi several specific and varietal names 
have been capitalized contrary to present day usage and the 
recommendations of the International Rules of Botanic Nomencla- 
ture. 

The avowed purpose of the Welder Wildlife Foundation for 
wildlife research and education is to be saluted enthusiastic- 
ally'. 



"Woody Flora of Taiwan", by Hui-Lin Li, 992 pp., 371 illustra- 
tions . A Morris Arboretiim Monograph, Livingston Publish- 
ing CcHnpany, Narberth, Pennsylvania. 1963. $18.75 

This is an unusually fine and comprehensive flora with keys, 
descriptions, literature citations, synonymy, and annotations 
on distribution, habitats and uses of 1030 species and many var- 
ieties in UH genera and 105 families. Original line drawings 
represent most of the genera with such artistic skill that the 
plants as a whole could easily be recognized from them, and with 
such scientific attention to detail that almost all the struc- 
tures of gross external anatomy are shown. This woik is monu- 
mental both in quantity and quality. 



196U Moldenke, Book reviews 239 

It is, of course, virtually impossible to produce such an ex- 
haustive work as this without some minor errors. The following 
comments on the treatment of the Verbenaceae are perhaps in order, 
but are not to be taken as any significant criticism of the work. 
The genus Avicennia is now usuaG-ly segregated as a separate fam- 
ily, Avicenniaceae, and contains 15 accepted species and ^ vari- 
eties instead of only "3 species". Callicarpa has li|.2 valid spe- 
cies and U? named forms and varieties instead of only "about UO"j 
C. kotoensis Hayata and C. japonica var. kotoensis (Hayata) Masa- 
mune belong in the synonymy of C. japonica var. luxurians Rehd. 
instead of in that of C. longifolia Lam. The name, C_. dichotoma , 
according to the International Rules must be credited to K. Koch 
rather than to Raeuschel. Caryopteris consists of 15 valid spe- 
cies, 1 hybrid, and 3 forms and varieties rather than "about 8 
species". Clerodendrum Burm., incorrectly written Clerodendron 
L., contains iil3 valid species and lU6 named varieties amd forms 
instead of only "about 100 species"; C_. canescens "HaJl. is a spe- 
cies distinct from C. viscosum Vent., and is not a synonym of it. 
The latter species is not known from Formosa or Japan. Premna 
has 198 valid species and 1|7 accepted varieties and forms; Vitex 
has 266 valid specific taxa and 105 forms and varieties, rather 
than "about l50 species". 

Nepeta is misspelled on page 82i4, Dr. P'ei's name on page 83O, 
and Gornutia on page 83I1. 

The following taxa have also been recorded from Formosa by H. 
N. Moldenke in his monographic studies, but are not accounted for 
by Li: Callicarpa pedunculata R, Br., Clerodendrum fragrans var. 
pleniflorum Schau., C. intermedium Cham., C. kaempferi (Jacq.) 
Sieb., C. ohwli Kanehira & Hatusima, C. trichotomum var. ferru- 
gineum Nakai, Duranta repens L., D. repens var, adba (Masters) L. 
H. Bailey, Lantana camara var, aculeata (L.) Moldenke, Premna 
foetida Reinw,, Sphenodesme involucrata (Presl) B. L. Robinson, 
Vitex negundo var. intermedia (P'ei) Moldenke, V. quinata var. 
puberula (H. J. Lam) Moldenke, and V. urceolata C. B. Clarke. 



"Biochemical Systematics", by Ralph E. Alston and B. L. Turner, 
UI6 pp., illustr. Prentice-Hall, Englewood Cliffs, New 
Jersey. I963. •'?13.25 

This welcome book culls those chemically natural relation- 
ships that exist among the orf^anisms studied from the many 
dissociated experimental works in this broad field, and the 
authors' e:ctensive study of the literature and their ovm ex- 
perimental work mainly on Baptisia by electrophoretic analysis 



2U0 PHYTOLOGIA Vol. 10, no. 3 

of certain non-essential and limitedly distributed metabolites 
found in the natural species, in natural and induced hybrids, and 
in taxonomically related genera. These compounds are many in 
number and from the following main groups: non-protein amino acid 
derivatives, certain organic acids, fatty acids, such phenols as 
anthocyanins and anthoxanthins and flavonoids, beta-cyanins, 
tannins, lignins, terpenoids, isothiocyanates, certain carotenoids, 
quinones, and sugars other than the ubiquitously essential glucose 
and fructose. After a few introductory chapters clearly and 
simply e:q5laining plant taxonrany, the bulk of the book is devoted 
to the nature, presence and work of these chemicals. The book 
continues with a survey of other chemicals as they have been 
traced from differing parents to hybrid offspring, and then sorted 
out, often in Kendelian pattern, to their offspring — as in the 
case of the inheritance of oil characteristics in the hybrids of 
Eucalyptus macarthuri x E. cinerea . The book closes with a gen- 
eral evaluation of physiological or chemical races, variations 
in development and with the mature plants, and metjiods of pre- 
senting comparative biochemical data for systematic purposes. As 
for evaluating specific biochemical data and for projecting the 
presont situation into the futiire, the authors "conclude that 
there is in the final analysis a much better chance of expressing 
specific biochemical differences in precise genetical terms (in- 
cluding characterization of the enzyme involved) . Therefore, 
although the art of assessing the phylogenetic value of morpho- 
logical data is farther advanced than the art of assessing the 
phylogenetic value of biochemical data, and we know far less at 
this time about variation in the chemistry of the plant, it is 
probable that in fifty years the situation will be reversed. 
Form is so subtly, delicately, and especially so indirectly 
regulated that its underlying genetics and biochemistry are 
likely to remain among the most intractable problems in biology 
for a long time." 

The text is lucid and comprehensive. The print is clear and 
clean, paper and binding serviceable, index useful, bibliography 
excellent for content but unfortunately arranged by chapter 
topics, and the illustrative material valuable. In the biblio- 
graphy many of J. B. McNair's works are included, but not his 
fine paper entitled "Energy and Evolution" published in volime 
2, Number 2, of the present journal in December I9UI. The 
generic name, Cycas , does not have its initial letter upper- 
cased, as it should, on page 366. 



Dn and 



.w 




Figure 12, Distribution of Verbena officinalis tn the United States 



Kerbariun curators vrtio have material of this species fron additional 
counties are asked to send it to the author for verification and 
record, so that future editions of this map may be more conplete 



appinc by coxmties done by Andrew R, Moldenl:e 




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PHYTOLOGIA^r,;; 

Designed to expedite botanical publication . ' ^^Al, 
Vol. 10 June, 1964 No. 4 



CONTENTS 



HARKNESS, B., A checklist of the cultivated woody plants of the 

Rochester parks. Part II 241 

MOLDENKE, H. N., Materials toward a monograph of the genus 

Verbena. XX 271 

MOLDENKE, A. L., Book review 319 



Published by Harold N. Moldenke and Alma L. Moldenke 

15 Glenbrook Avenue 
Yonkers 5, New York, U.S.A. 

Price of this number, SI; per volume, $5-75, in advance 



A CHECKLIST OF THE CULTIVAT2D WOODY PLAITTS OF THE 
H0CH2STEB PARKS. 
Fart II 

Berr3,rd Harkness, 

PTaiit Taxonomist, Monroe Co. Bept. of Parks 



PICEA, con. 






rut ens 


Biltmore Nur. 


1902 


Schreakiana 


Arnold Ar "bore tun 


1912 


Wilsonii 


Arnold Ar^boret-um 


1915 


PIERIS - Ericaceae 






florilranda 


J. Dawson 


1907 


japonica 






J. 'Variegata' 


nolden Ar"boretun 


1957 


PIIJUS - Pinaceae 






aristata 


Hicks llur. 


1917 


Amandii 


Arnold ArTDoretum 


1905 


Bi.ingeana 


Me e ban Nur. 


1900 


CemlDra 


Ellwanger & Barry 


1898 


cem'broides ed-olis 


H. Ullrich 


1953 


c. tionophylla (") 






contorta latifolia 


Biltmore Nur. 


1902 


denaiflora 


Arnold Ar"boret\m 


1902 


d. 'Globosa' 


Andorra llur. 


1916 


d. 'Oculls-draconis' 


Holden Ar"boretuin 


1957 


flexilia 


Veitch llur. 


1898 


f. 'Gleniaore Pyranid' 


E. More 


1956 


Jeffrey! 


Veitch llur. 


1898 


Hunnewellii 


V. MclTitt 


195^ 


koraiensis 


Veitch Nur. 


1899 


nonticola 


Arnold Artoretum 


1902 


Mugo 


Arnold Arhoretim 


1920 


M. 'Compacta' 


D, Hill Uur. 


1922 


M. MiA^hus 






M. ro strata 


Arnold Ar'boret-um 


1902 


M. rot-undata 


Ellwanger 3e "BarT-y 


1897 


M. 'Slavin' 


Rochester seedling 




nigra 


Arnold Arboretum 


1905 


n. austriaca 


Ellwanger & Barry 


1898 


n. carananica 


Ji-llwanger & Barry 


1897 


n. cebennensis 


Veitch llur. 


1903 


n. •Homi'brook' 


Rochester origin 




n. 'Monstrosa' 


Rochester seedling 




n. Poiretiana 


Veitch ITur, 


1898 


n. 'Pyranidalis' 


Rochester seedling 




parriflora 


Veitch Nur. 


1898 


p. pentaphylla 


Arnold Arhoretum 


1911 


Peuce 


Arnold Arhoretum 


1896 


ponderosa 


veitch Nur. 


1898 


p. ' Pendula' 


Arnold Arboretum 

2ia 


1913 



21;2 



PHITOLOGIA 



Vol, 10, no, U 



PIUUS, con. 






ponderosa scopulorum 


Arnold Arlioret-um 


1913 


resinosa 


Uoiiglas ITur. 


1898 


rigida 


e side, Irondequoit Bay 


1902 


Satini n.na, 


Schumacher 


1957 


StroMs 


Bllwanger & Barry 


1896 


S. 'Contorta' 


Hochester origin 




S. 'Densa' 






S. 'Fastigiata' 


Hicks Nur. 


1920 


S. 'Ontario' 


Kochester origin 




sylvestris 


Ellwanger & Barry 


1898 


s. 'Argentea' 


Veitch Nur. 


1903 


s. 'Fastigiata' 


Hicks Kur. 


1920 


s . ' Nana' 


Veitch Nur. 


1903 


s. 'Pumila' 






tatulaeformis 


Arnold Arboretum 


1911 


ThunlDergii 


Arnold Arboretum 


1902 


Wallichiana 


Little Nur. 


1898 


PLATAMJa - Platariftceae 






acerifolia 






occidentalis 






POFUIUS - Salicaceae 






alba 






a. 'Ei chard' 


Arnold Arboretum 


1916 


■berolinensis 


Arnold Arboretum 


1907 


caaadensis 'Eugene' 


Barbie r Nur. 


1907 


c. 'Regenerata' 






c. 'Serotina' 






canescens 






cathayana 






deltoides 


BHS, Eose Hill, Canada 




Fremont! i 


Spaeth Nur. 


1908 


gileadensis 


Horsey, I ronton, Ohio 




grandidentata 






Maximowiczii 


W. A. Smith 


1952 


nigra 


Spaeth Nur. 




u. 'Betulifolia' 


Arnold Arboretum 


1907 


n. 'Elegans' 


Arnold ArboretvBa 


1907 


. n. 'Italica' 






n. 'Volga' 


Teas Nur. 


1907 


rotusta 


Spaeth Nur. 




Sietoldii 


Arnold Arboretum 


1952 


Simbnii 


Arnold Arboretum 


1907 


S. 'Fastigiata' 


Arnold Arboretum 


1917 


tremuloides 






POTMTITiLA - Eosaceae 






artuscula albicans 


Arnold Arboretum 




fruticosa 


i4eehan Nur. 


1915 


f. 'Longacre' (K) 


U. S. D. A. 


1962 


f. 'Moonlight' 


Uolden Arboretum 


1956 



I961i 



Harkness, Cultivated woody plants 



2U3 



fruticosa 'Rigida' 


Mayfair ITur. 


1953 


f. 'Tenuilota' 


l<Iayfair ITrir. 


1958 


PEIITSEPIA - Eosaceae 






sinensis 


Arnold Artoretum 


1915 


uniflora serrata 


Morton Artoretum 


19^ 


PEUiruS - Eosaceae 






alle^eniensis 


Arnold Artoretiin 


1902 


americana 


Arnold Art ore turn 


1913 


an^ustifolia 


Arnold Ar "bore tun 


1912 


apetala 


Arnold Ar "bore turn 


1916 


Armeniaca 






avi-un 


Arnold Arboretum 


1912 


Besseyi 






cerasifera 


Arnold Artoret-un 


1916 


Davidiana 


Arnold Arboretum 


1919 


D. 'Allja' 






donestica 'Plantier' 






Dun"barii 


Eochester seedling 




Grave sii 


K. Y. Bot. Gard. 


1952 


Grayana 


Arnold Ar "bore tun 


1913 


"Hally Jolivette' 


Arnold Arl) ore turn 


1951 


incisa 


Arnold Artoretuxi 


1919 


i. Yaoadei 


Arnold ArTjoretun 


1917 


institia 


Spaeth. Kur. 


1902 


lanata (11.) 


Morton Artoretum 


1959 


La'orocerasus 'Mischeana,' 


(11.) Briiafield Gard. 


1958 


L. ' Scliipkaensis' 


Jiillwanger garden 


19^8 


I^aackii 


Spaeth l?ar. 


1902 


mandshurica 


Arnold Arboretum 


1931 


maritima 


Dunbar, Long Island 


1912 


I4axiiaovn.cz ii 


lliklco Bot. Gard. 


1956 


mexicana 


Arnold Arboretum 


1911 


nigra 


Arnold Arboretum 


1902 


nipponica 


ITikko Bot. Gard. 


1956 


Padus 


Arnold Arboretum 


1912 


P. commutata 


Arnold Arboretum 


1912 


Sargentii 


Arnold Arboretum 


1902 


serotina 


Arnold Arboretum 


1902 


serrulata 






s. 'Amonogawa' (IT.) 


Weston llur. 


1962 


s. 'Asagi' 


Arnold Arboretum 


1917 


s. 'Hata^zakuri' 


Arnold Arboretvim 


1917 


s . 'Ho s ofewa-odo ra ' 






s. 'Jo-llioi' 


Arnold Arboretum 


1917 


3. 'Kikushidare' 


Arnold Arboretum 


1917 


s . ' Kuramayama' 


Arnold Arboretum 


1917 


s. 'Kwanzan' 


Arnold Arboretvim 


1917 


s. 'Oklna' 


Arnold Arboretum 


1917 


8* ' Sakon' 


Arnold Arboretum 


1917 



2lll4 P H Y T 


L G I A 


Vol. 


10, no 


PEUHUS, con. 








serrulate ' Shirofu^eii' 


Arnold ArTDoretum 




1917 


s. 'Sliirotae' 


Arnold Artoret-uin 




1917 


sulshlrtella 


Arnold Artoretum 




1912 


s. ascendens (H.) 


Holden ArlDoretuin 




i960 


s. pendula 


Arnold Arboretum 




1907 


tomentosa 


Arnold Artoretum 




1892 


t. 'Geneva' 


U. Y. State Exp. 


Sta. 


1931 


t. ' Leucocarpa' 


N. Y. State Exp. 


Sta. 


1931 


Virginians 


Arnold ArlDoret-um 




1902 


V. demissa 


Spaeth Nur. 




1902 


V. melanocarpa 


Arnold Artoretum 




1915 


yedoensis 


Arnold Artoretum 




1917 


y. 'Perpendens' 


Arnold Artoretum 




1919 


PSETJDOLAEIX - Pinaceae 








amatilis 


Veitch Hur. 




1905 


PSEUTOSASA - Gramineae 








japonica 


Ellwanger garden 




195^ 


PSEiroOTSUGA - Pinaceae 








Menziesii glaiica 








M. g. 'Big Flats' 


Seedling in Big Tlats Nur. 


. 1955 


M. g. 'Densa' 


Arnold ArTDorettun 




1896 


M. g. 'Pastigiata' 


Rochester seedlii 


ig 


1913 



PTELEA - Eutaceae 
lutescens 
trifoliata 
t. mollis 



PTEROCARYA - Juglandaceae 



fraxinifolia 


Arnold Artoretuin 


1902 


rhoifolia (K.) 


Smith College 


i960 


stenocarpa 


Arnold Axtoretxim 


1919 


PTEROSTYRAX - Styracaceae 






hispida 


Arnold Artoretum 


1902 


PYRUIARIA - Santalaceae 






pu"bera 


BE, ZentTicky 


i960 


PYEUS - Rosaceae 






"betulaefolia 


Arnold Arboretum 


1912 


Bretschneideri 


Arnold Arlsoretum 


1918 


Calleryana 


Arnold Arboret-um 


1915 


C. dimorphophylla 


Arnold ArTDorettun 


1922 


C. Pauriei 


Arnold Arhoretum 


1920 


C. tomentella 


Arnold Arhoretum 


1918 


canescens 


Spaeth WuT. 


1902 


comia'unia 


Arnold Artoretiim 


191J^ 


elaegrifolia 


Spaeth Nur. 


1902 


Lindleyi 


Arnold Artorettun 


1919 



196U 



Harkneis, (Mltivated woody plants 



2kS 



PYEUS, con. 






Kichauxii 


Arnold Ar"boret-um 


1915 


nivalis 


Spaeth Nur. 


1902 


phaeocarpa 


Arnold Artoretum 


1918 


pyrifolia 


Arnold Artoretum 


I9I8 


p. 'Culta' 






salicifolia (E.) 


Vienna 


1952 


serrulata 


Arnold Artoretum 


1915 


ussuriensis 


Arnold ArtoretTim 


1919 


u. hondoensis 


Arnold Ar "bore turn 


1921 


u. ovoidea 


Arnold Artoret-um 


1915 


qUEECUS - Fagaceae 






acutissina 


Biltmore Nur. 


1902 


alba 






a. latilo"ba 






aliena 


Arnold ArlDoretum 


1909 


a. acute serrata 


Arnold Artoretum 


1907 


arkansana 


Arnold Artoretum 


1916 


Beblaiana 


Arnold Artoretum 


1902 


■bicolor 






castaneaefolia 






Cerris 


Biltmore llur. 


191^ 


coccinea 


Arnold Artoretum 


1921 


dentata 


Biltmore ITur. 


1902 


d. ' Pinnatif ida' 


A. J. Malefyt 


1953 


ellipsoides 


Arnold Artoretum 


1907 


Femowii 


Arnold Artoret-um 


1921 


Frainetto 


Veitch ITur. 


1907 


glandulifera 


Arnold ArtoretTm 


1907 


Haas (11.) 


U. S. D. A. 


i960 


ilicifolia 






inlDricaria 






Jackiana 






liaotungensis 


Arnold Artoretum 


1929 


macrocarpa 






marilandica 


Buntar, Bayvllle 


1908 


Mlchavucii 






Muehlenbergil 






palustris 


Horsey, Bowling Green 




petraea 






p. 'Mespilifolia' 


Jiillwnnger & Barry 




Phellos 


Teas ITur. 


1923 


prinoides 


Meehai llur. 


1992 


Prinus 


i-nintar, Baj'ville 


1908 


puliescens 


Arnold Artoretum 


1906 


Ro"bur 


Arnold Artoretum 


1906 


H. 'Fastigiata' 






E. 'Filicifolia' 


Ellwanger & Barry 




rutra 






r. ■boreal is 


Arnold Artoretum 




jrunclnata 


Arnold Artoretum 


1921 


Sargentii 


Arnold Artoretum 


1902 



2lt6 



PHTTOLOQIA 



Vol. 10. no. U 



qUEECUS, con. 






Saulii 






Schuettei 






Sliimaxdii 


Arnold Arl: ore turn 


1918 


S. Schneckii 


Arnold Artoretum 


1907 


Stella ta 






s. Margaret ta 


B.H.Slavin, Oklahoma City 




texana 


Arnold ArlDorettua 


1902 


Tariatilis 


Arnold Ar^b ore turn 


1907 


Telutina 


Arnold Art ore turn 


1907 


V. missouriensis 


Arnold Arboretum 


1907 


RHAMMUS - Ehajnnaceae 






alpina 


Spaeth. Nur. 


1892 


cathartica 


Spaeth Nur. 


1892 


davurica 


Arnold Arhoret-um 


1902 


d. nipponica 


Arnold Arhoret-um 


1916 


dolichophylla (N.) 


Alma-Ata 


i960 


Fra.ngula 


J. Dawson 


1892 


infectoria 


Spaeth Nur. 


1902 


japordca 


Arnold Artoretum 


1906 


koraiensis 


Kohankie Nur. 


19^5 


Pallasii (N.) 


Morton Arhoretum 


1956 


Parshlp„na 


Spaeth Hur. 


1902 


rutra (U.) 






Schneider! manshurica 


Arnold Artoretum 


1919 


utilis 


Spaeth Nur. 


1892 


Wilsonii (K.) 


Mt. Airy Artoretum 


i960 


EHODODEiroROlI - Ericaceae 






albicans 


A. Water er 


190/f 


ar'borescens 


J. Dawson 


1892 


atlanticTom 






August ini 


U. of Wash. Arboretum 


1952 


■braclij'carpuia 


Kegel & Kesselring 


1910 


calendijlaceum 


Kelsey Nur. 


1899 


calophytum 


Dr. Irving 


i960 


concinnum lepidanthum 


Wisley 


1953 


canadense alMflorum 


D. Leach 


1958 


carolinlanum 


Eobbins Nur. 


1922 


cuneat-um 


EdinlDurgh 


1955 


decorum 


U. of Wash. ArlDoret-um 


195^ 


Fauriei 


Edinburgh 


1952 


flavum 






haematodes 


Schumacher 


1955 


Mppophaeoides 


Bremen 


1953 


Houlstonii 


U. of Wash. Arboretum 


1952 


japonicTJm 


Arnold Arboretum 


191^ 


keleticum 






laetivirens 


Bobbink & Atkins 


1917 


leucapsis 


Schumacher 


1956 


Makinoi 


Bremen 


1952 


maxinrujn 


J. Dawson 


1892 



196h 



Harkness, Cultivated woody plants 



2U7 



EHDDOEEKDROU, con. 






nicranthxua 


Arnold Ar"borettim 


1922 


minus 


LnBar Nur. 


1921 


fflucronulatum 


Arnold Artoretum 


1902 


mucronatuin 






m. ripen.se 






ottusTia Eaenpferi 






praecox 


Dickson Nur. 


1892 


reticulattun 


Nikko Bot. Gard. 


1950 


rutignosuQ 


U. of Wash. Artoretum 


195^ 


rupicole 


Wisley 


1955 


russatum 


Bremen 


1953 


Smirnovd.i 


Edinturgh. 


1932 


sutchuenense Giraldii 


U. of VJash. ArlDoretum 


1951 


Vaseyi 


Eelsey Nur. 


1892 


viscosepaluiii 


A. Waterer 


190i; 


Weyricliii 






yakusimar.-um 


D. Leach 


1958 


yedoense 






RHODODEI^DROIT - (Cultivars) 






'AlTDun Elegans' 


Ellv/anger & Barry 


1892 


'Altjun G-randiflorun' 


illwanger & Barry 


1892 


'Altaclareuse' Az, 


A. Waterer 


190^ 


'imphion' 


A. Waterer 


1900 


'Appelles' Az. 


van Heiningen 


191^ 


'Ariadne' Ax. 


van Heiningen 


1914 


' As tr cans' Az. 


Veitch ITur. 


1907 


' Augusts Meclielynch' Az. 


van Heiningen 


1914 


'Aurore de Eooigliem' Az. 


Teitch i:-ar. 


1907 


'Beaufort' 


D. Leach 


1958 


'Beaute Celeste' Az. 






'Belle Vemeille' Az. 


Veitch ITur. 


1907 


'Bijou des Anateurs' Az. 


van Heiningen 


1914 


•Blue Tit' 


Brimfield Gard. 


1955 


'Boule de lleige' 


A. Vfeterer 


1900 


'Boule de Rose' 


D. Leach 


1958 


'Bouq.uet de Flore' Az. 


Veitch l^MT. 


1907 


'Caractacus' 


A. Waterer 


1904 


'Cardinal' Az. 


A. V/aterer 


1904 


'Caroline' 


D. Leach 


1958 


'Charles Barley' 


Cottage Gard, 


1937 


•Charles Dickens' 


A. V.'aterer 


1904 


'Charles Rogier' Az. 


van Heiningen 


1914 


'Coote de Goner' Az. 


A. Waterer 


1904 


'Conte de Fapadopoli' Az. 


van Heiningen 


1914 


• Comte de Q,uincy' Az. 


A. Waterer 


1904 


'Cunninghams VHiite' 






'Cynodocee' Az. 


A. V/aterer 


1904 


'Davies' Az. 


A. Waterer 


1900 


'Delicatissinum' 


Cottage Gard. 


1918 


'Domenico Scassi' Az. 


A. Waterer 


1904 


'Dr. M. Oosthoek' Az. 


Grootendorst 


1951 



>ll8 PHYTOLOG 


I A Vol. 


10, no 


RHOIX)DEKnEOH (cv), con. 






'Dr. Streiter' Az. 


Veitch Uur. 


1907 


'Doc de Provence' Az. 


Veitch iTur. 


1907 


'Edison' Az. 


A. Waterer 


190^ 


•ElizaTseth' Az. 


A. Waterer 


1904 


'Esmeralda' Az. 


van Heiningen 


1914 


'Everestlanxan' 


Ellwanger & Barry 


1892 


' Efijna' Az . 


A. V/aterer 


1904 


'Fanfare' 


D. Leach 


1958 


'Erere Ortan' Az. 


A. Vfeterer 


1904 


'General Brialmont' Az. 


A. Waterer 


1904 


'General Goffinet' Az. 


van Heiningen 


1914 


'Gloire de la Belgique' Az. 


van Heiningen 


1914 


'Gloria Mundi' Az. 


A. Vfeterer 


1900 


'Goldsworth Yellow' 




1958 


'Grandeur Triomphant' Az. 


A. Waterer 


1904 


'Henri Conscience' Az. 


van Heiningen 


1914 


'H. H. Hnnnewell' 


A. V7aterer 


1904 


'H. W. Sargent' 


Cottage Gard. 


1918 


'Ignatius Sargent' 


van Heiningen 


1914 


'11 Tasso' Az. 


van Heiningen 


1914 


« Imperator' Az. 


A. Waterer 


1904 


'Josephine KLinger' Az. 


Veitch Kur. 


1907 


'Julda Sclmpp' Az. 


A. Waterer 


1904 


' lady Armstrong' 


A. Waterer 


1904 


'Lady Grey Egerton' 


A. V/aterer 


1900 


'Liberty' Az. 


Grootendorst 


1951 


'Ma'bel Parsons' 






'Madame Carvallio' Az. 


A. Waterer 


1904 


'Mme. Gustave Guillemot' A2 


. 




'Itoe. Joseph Bavimann' Az. 


A. Waterer 


1904 


'Marie Ardent' Az. 


A. Waterer 


1904 


'Marie Verschaffelt' Az. 


A. Waterer 


1904 


'Mecene' Az. 


van Heiningen 


1914 


' Memoir' 


A. Waterer 


1909 


• Mlgnon' Az . 


A. Waterer 


1904 


'Monsieiir des Bois' Az. 


van Heiningen 


1914 


'Mrs. C. S. Sargent' 


A. Waterer 


1904 


'Nancy Waterer' Az. 


A. Waterer 


1904 


'Pallas' Az. 


A. Waterer 


1904 


. 'Parsons Gloriosum' 


Cottage Gard. 


1937 


'Parsons Grandiflorum' 


Cottage Gard. 


1918 


•Phoete' Az. 


van Heiningen 


1914 


'Phidias' Az. 


van Heiningen 


1914 


'Prince Albert' Az. 


A. Waterer 


1904 


'Pacelle' Az. 






'Porpureum Crispum' 






'Parpureian Elegans' 






'Parp\ireum Grandiflorum' 


Cottage Gard. 


1937 


'Q,ueen Emma' Az. 


Grootendorst 


1951 


'Eamapo' 


D. Leach 


1958 


'Eemhrant' Az. 


Veitch Nut. 


1907 


»Eoi des Peux' Az. 


van Heiningen 


1914 



196U 



Harkness, CultiTat«d woody plant* 



2li9 



EHODODEEDEON (cv. ), con. 
'Hosetud' Az. 



'Eosetta' Az. 




van Heiningen 


191^^ 


'Eoseum Blegans' 




Bllwanger & Barry 


1892 


'Eussel Harmon' 




D. Leach 


1958 


« Scintillation' 




D. Leach 


1958 


•Uniqiie' Az. 




A. Waterer 


190^ 


« WindlDeaia' 




Baldslef sen Nur. 


I960 


EHODO TYPOS _ Eosaceae 








Bcandens 




Parsons Nur. 


1892 


EHUS - Anacardaceae 








aromatica 




iillvwnger & Barry 


1892 


a. flatelliformis 




Arnold Artoretum 


1906 


chinensis 




W. A. Smith 


1958 


copallina 




Meehan Nur. 


1915 


glatra 'Laciniata' 




i^llwanger & Barry 


1892 


ponjatensis sinica 




Arnold Artoretum 




typhina 




Arnold Artoretum 




EIBES - Saxifragaceae 








alpintim 




Spaeth Nut. 


1892 


americantan 




Spaeth Nur. 


1892 


aureum 




Arnold Artoretum 




Carrierei 




Spaeth Nur. 


1892 


diacajithum 




Spaeth Nur. 


1892 


futurojn 








Gordonianum 




iillwanger & Barry 


1892 


holosericeum 








nivetun 




Arnold Arhoretum 


1907 


odoratum 




Slavln, Muskogee, Okla. 




tenue 




Arnold Arhoretum 


1914 


■uxceolatum 








Warscevriczii (ll.) 




Zornick 


1959 


EOBINIA - Leguminosae 








fertilis 








hispida 




J. Dawson 


1852 


Holdtii 'Britzensis' 








Kelseyi 




Kelsey Nur. 


1919 


luxurians 




Arnold Arhoretum 


1920 


Pseiidoacacia 








P. 'Decaisne' 








P. 'Dependens' 








P. 'Eelider* 








Slavinii 




Eoehester Seedling 




viscosa 




Eohhins Nur 


1922 


BOSA - Eosaceae 








acicxilaris BourgeauianA (N.)Andrews llur. 


1915 


adenosepala (N.) 




Morton Artoretum 


1926 


Albertii (N.) 




T. Smith Nur. 


1916 


arvensia ayreshirea 


(N.) 


Arnold Artoretum 


1916 



>50 P H Y T L 


G 


\ I A Vol. 


10, no, k 


ROSA, con. 

tella (K.) 




Griefswald 


i960 


iDlanda, alta (N.) 




Arnold Arhoretum 


1916 


centif olia muscosa (U. ] 


) 






cixuiafflomea plena (N.) 




Morton ArlDoretum 


1926 


consan^oinea (K.) 




Arnold Ar'boretuin 


1917 


corym"bifera (H.) 




Arnold Ar "bore turn 


1907 


danascena (N. ) 




Morton Ar'ooretum 


1926 


d. tringintipetala (ll. '. 


) 


Arnold Arlsoretum 


1917 


davurica (N. ) 




Arnold Arhoretum 


1907 


Fedtsclienkoanfl. (N.) 




T. Smith ITur. 


1899 


f ilipes (N. ) 




U. of Wash. ArTDoretum 


195^ 


fujisanensis (K.) 




Schumacher 


1955 


Helenae (H.) 




Arnold Ar "bore turn 


1911 


Hemsleyana (N. ) 




U. of Wash. Arhoretum 


195^ 


Lheritieriana 'Madis' 


(N.) Morton ArlDo re turn 


1926 


macrophylla (U.) 




Griefswald 


i960 


Malyi (N.) 




T. Smith ITur. 


1899 


manca (N.) 




Auderghem 


1962 


margiaata (K.) 




Arnold Arhoretum 


1917 


Maxino\id.cziana (IT.) 




U. of Wash. Arhoretum 


1957 


montana (IT.) 




Kew 


i960 


Moyesii ill.) 








m-ultiflora catheyensis 


(N.) Arnold Arlsoretum 


1915 


m. 'Mrs. F. W. Flight' 


(IT.) V. aibhs 


191^ 


m. ' Platyphylla' (N.) 




Arnold Ar'boretum 


1920 


Onoei (N.) 




U. S. D. A. 


1957 


palustris Hut; tal liana 


(IT.) 


T. Smith Nut. 


1899 


pendullna (N. ) 




Backhouse Uur. 


1892 


Primula (U.) 




Arnold Ar'boretum 


1917 


rugosa 'Alta* (H.) 




Toronto 


i960 


sertata (U.) 




Edinburgh 


i960 


spinossissima altaica 


(N.) 


Arnold Ar'boretum 


1917 


WetMflTia, (U.) 




T. Smith Hur. 


1899 


WichrdTiana X Sotilieana 


(VanFleet #18) (N.) 








U. S. D. A. 


1920 


Woodsii Fendleri (K.) 




J. Dawson 


1892 


EOSA, C\J.tivars 








«MySo"b3art' (U.) 




BllwH,nger & Barry 


1901 


'Apple Blossom' (H.) 




Arnold Ar'boretum 


1918 


'Bradwardine' (U.) 








'Burnet Brightness' (N.) 


T. Smith Nut. 


1916 


'Heart of fiold' (VanFl^ 


eet 


#5) (K.) 




'Jeannie Deans' (N.) 




Ellwanger & Barry 


1901 


'William C. Egan> (N. ) 




J. Dawson 


1896 


' Zuccariniana' (N. ) 




T. Git'bs 


191^ 



EUBTJS - Eosaceae 

Cockhurnianus 
phoenicolasius 



Willowwood Ar'boretum 



1950 



196U 



Harkness, Cxxltl-mt«d wooc^jr plants 



251 



SAGEEETIA - Ehaaoaceae 
pycnophylla 

SALIX - Salicaceae 



acutifolia 


Arnold Arboretum 


1917 


alTDa 'Calva' 


Arnold Artioretuia 


1911 


a. 'Ch.ermesina' 


Ellwanger & Barry 


1918 


a. 'Sericea' 


Arnold Arboretim 


1917 


a, 'Vitellina' 


Arnold ArTDoretum 


1919 


aaygdalina (N.) 


Mort6n Artoretura 


i960 


"batylonica 'Crispa' (U.) 


Holden Arboretum 


i960 


"blanda 


Arnold Arboretum 


1916 


cinerea (ll.) 


Morton Arboretvua 


i960 


ElaeaeD-os 






holosericea 


Arnold Ar"boret\im 


1906 


Hooteriana 


Arnold Arboretum 


1917 


jessoensis 


Arnold Arboretum 


1917 


icoreensis (IT.) 


Morton Arboretum 


i960 


Hatsudana 


U. S. D. A. 




M. 'Pendula' 


Morton Arboretum 


1933 


Meyeriftna (U.) 


Morton Arboretum 


i960 


Miyateana (N.) 


Morton Arboretum 


i960 


nigra 


native at Hendon Park 




Pierotii 


Spaeth Nut. 




purpurea 'Waucescens' 


Arnold Arboretum 


1906 


p. 'Pendula' (ll.) 


Holden Arboretum 


i960 


nilDens 


Ellwarger & Barry 


1892 


sepulcralis 


Arnold Arboretum 


1917 


•JiiiJi, U1.^CU.C^ 


Not cut t IJur. 


1959 


stlpularis 


Spaeth Hut. 


1892 


SAI-CBUCUS - Caprifoliaceae 






canadensis acutilota 


W. A. Smith 


1955 


nigra 


Spaeth ITur. 


1892 


n. 'lllJo-variegata' 


Horton Nur. 


1958 


SAPniDUS - Sapindaceae 






Drummondii 


Slavin, Okla. 


191^ 


SARCOCOCCA - Busaceae 






Hookeriana humilla 


Hicks llior. 





SASSAFRAS - Lauraceae 



albidum 


Arnold Arboretum 


1913 


SCHIZOPHKAGMA - Saxlfragaceae 
hydrangeoides 


Arnold Arboretum 


1916 


SCIADOPITYS _ Pinaceae 
verticillata 


Yokahajna Hur. 


1922 


SECURINEGA - Buphorbiaceae 
suffruticosa 


Morton Arboretum 


1950 



2^2 PHY 


T L G I A Vol. 10, no. 


SHEFHEHDIA - Blaea^naceae 






argent ea 




Zohankie Hur. 


19^5 


SMILAX - Liliaceae 








glauca 




Slavin, Asheville 


1911 


rotundifolia 




J. Dawson 


1892 


SOPHOEA - Leguminosae 








japonica 




Meelvi,n Hur. 


1899 


SOHBARIA - Rosaceae 








arlDorea 




Arnold ArTDoretiam 


1913 


a. gla"brata 




Arnold Ar "bore turn 


1913 


a. suTstomentosa 








assurgens 




Bartiier Nur. 


1907 


sortifolla 




Sllwanger & Barry 


1892 


s. stelliplla 




Arnold Artoretrum 


1911 



SOEBOPTRUS - Eosaceae 

auricularis 'bultiformis 



Arnold ArlDorettim 



1912 



SOEBUS - Eosaceae 






alnifolia 


Arnold ArTDoretum 


1920 


amerioana 


Arnold Artoretum 


1913 


Aria 


Dickson Nur. 


1907 


A. longifolia 


Dickson Nur. 


1907 


A. 'Latescens' 


Grrootendorst llur. 


1957 


arnoldiana 






Aucuparia 


Arnold ArlDoretum 


1907 


A. lanuginosa 






'Carpet of CJold' (H.) 


Weston Nur. 


1962 


commixta 


U. S. D. A. 


i960 


decora 






discolor 


Spaeth Nur. 


1908 


Folgneri 


Arnold Artoretum 


1919 


hylDrida 


Arnold ArlDoretum 


1906 


h. 'Meinich' 






intermedia minima 






japonica calocarpa 


Arnold Artoretum 


1920 


latlfolia 




1932 


Mougeotii 






randaiensis (N.) 


U. of Wash. Arhoretum 


1952 


rufo-ferruginea 


Arnold Ar^boretum 


1918 


serotina 


Eoyal Bot. Gard.. Hamilt 


iOn 
1961 


SPIEAEA - Eosaceae 






alta 


Ellwanger & Barry 


1892 


Billiardii 


Spaeth Nur. 


1892 


"blanda 


Arnold Ar "bore turn 


1902 


Blumei 


Arnold ArlDoretum 


1919 


"brachytotrys 


Ellv/ajiger & Barry 


1892 


Buinalda 






B. 'Anthony VJaterer' 







196U 



Hso-kness, Cultivated woody plants 



253 



SPIBASA, con. 

Bumalda 'Norman' 

3. 'Pnihonlca' 

chamaedryfolia 

c. u] mi folia 

cinerea 

conspicua 

DoToglasii 

Henryi 

hypericifolia 

latlfolia 

lon^igemmis (Nt) 

Margaritae 

media 

m. sericea (N») 

Menziesii 

Miyabei glatrata 

mollifolia (N.) 

nipponica 

n. rotixadifolia 

nudiflora 

oxyodon 

prunifolia 

p. simplicifolia 

pyramidata 

salicifolia 

ThunlDergii 

trichocarpa 

trilo'bata 

Van Houttel 

Watsoniaaa 



Willowwood Artioretua 1952 

Arnold ArlDoretuin 1906 

Arnold Artoretum 1919 

Spaeth ITur. 1892 

iillwanger & Barry 1892 

Arnold Ar"boretum 1909 

Bllwanger & Barry 1892 

Kew 1958 

Spaeth Nur. 1892 

LeEoy Nut. 1892 

Leningrad 1958 

V. aiT3-bs 191^ 

Lemoine Niir. 191^ 

Kew 195^^ 

Arnold Ar"boretujn 1909 

Illvranger & Barry 1892 

V. GitlDS 191^ 

Ellwanger & Barry 1892 

Spaeth llur. 1892 

Arnold Artoretum 1919 

Nikko Bot. Sard. 1950 

Arnold Artoret-um 1919 

iillwanger & Barry 1892 

Dickson N\ir, 1892 



STACHYUHUS - Stachyuraceae 



chinensis 


Willo^vwood ArTDoretiim 


1957 


praecoz 


&. Landis Ar "bore turn 


1954 


STAPHYLEA - Staphyleaceae 






colchica 


Ellwanger & Barry 


1892 


pimiata 


J. Dawson 


1892 


trifolia 






STEPHAIIAIIDEA - Eosaceae 






Incisa 


Meelian Nur. 


1919 


i. 'Crispa' 


Gulf Stream ITur. 


1958 


sp. aff. chinensis 






Tanakae 


T. Smith Nur. 


1899 


STEWARTIA - Theaceae 






koreana 


Arnold Ar"boretum 


1952 


Malacodendron 






Pseudo-Camellia 


Parson Nur, 


1892 



25U 


P H Y T L 


G I A Vol. 


10, no. U 


STYEAI . 


- Styracaceae 






dasyantha 


T. H. Everett 


1952 


japonica 


Veitch Nur. 


1899 


Olsassia 


Saul Nut. 


1892 


SIMPHORICAEPOS - Caprif oliaceae 




al'bus 


Spaeth llur. 


1892 


a. laevlgatus 


Slavin, Milwa\ikee 


1918 


Chenaultii 


E. 0. Orpet 


191^ 


orbiciilatus 


LeEoy Nur. 


1892 


STMPLOCOS _ Symplocaceae 






panloiilata 


Parsons liTor. 


1892 


SYEINGA 


- Oleaceae 






araurensis 


Arnold ArlDoretum 


1919 


&' . 


japonica 


EllvffUTger & Barry 


1892 


chinensis 


Ellvreinger & Barry 


1892 


c. 


Al-ba« 


Ellwanger & Barry 


1892 


c. 


Blcolor' 


Lemoine ITur. 


1905 


c. 


Le Troyes' 






0. 


Metensis' 


Kohankie Ntot. 


1940 


c. 


Orchid Beauty' (N.) 


Arnold ArTjoretuin 


1956 


c. 


President Hayes' 


BarMer Nur. 


1902 


c. 


Saxigeana' 


Transon Nur. 


1892 


c. 


Steencruysii' 


Parsons Hur. 


1892 


diversifolia (ll.) 


Elan Mem. Park 


1958 


emodi 


Spaeth Nur. 


1892 


e. 


Aurea' 






Henryi 'AlTia' (ll.) 


Arnold Arhoretum 


1956 


H. 


Charles Heptum' 


BoTDtink & Atkins 


19'^2 


H. X tomentella - 'Prairial' Morton ArTDoretum 


19^ 


hyacinthiflora 'Alice East 


wood' 








Swarthmore 


1947 


h. 


Assessippi' 


Swarthmore 


1941 


h. 


Berryer' 


Lemoine Nur. 


1914 


h. 


Blue Hyacinth' 


Swarthmore 


1947 


h. 


Bountiful' (H.) 


Swarthmore 


1957 


h. 


Buffon' 


Col. Plum 


1925 


h. 


Catinat' 


Morton Arboretum 


1927 


h. 


Charles llordine' (IT.) 


S'rfartl-Miore 


1957 


h. 


Clarkes Giant' (U. ) 


R\iliffson 


1953 


h. 


Claude Bernard' 


Lemoine Nur. 


1916 


. h. 


'Descartes' 


Lemoine Nur. 


1917 


h. 


Dr. Chadwick' (N.) 


Morton Artoretum 


1962 


h. 


•Estlier Staley' 


V/ayside Gard. 


1949 


h. 


Evangeline' 


Arnold Arhoretum 


1942 


h. 


•Excel' 


Swarthmore 


1941 


h. 


Fantasy' (U. ) 


Weston Nur. 


1962 


h. 


Penelon' 


Morton Arhoretum 


1944 


h. 


Gertrude Leslie' (N.) 


Sv/arthmore 


1957 


h. 


Grace' (H. ) 


Sira.rthmore 


1957 


h. 


]fe.zel Opper' (N.) 


Dr. Rankin 


1955 



196U 



Harkness, Cultirated woody plants 



255 



SYBIUGA. con. 

nyacinthiflora 'Jewel' (N.) 

h. ' Eate Sessions' 

h. 'Lainartine' 

h. 'Louvois' 

h. ' Minnehaha' 

h. 'Montesq-oieu' 

h. 'Necl^r' (IT.) 

h. 'Nokomis' 

h. 'Uorah' (IT.) 

h. 'Pascal' 

h. 'Peggy' 

'Pink Cloud' 

'Pink Spray' (H.) 

'Plena' 
h. 'Pocahontas (N.) 
h. 'Purple Glory' (K.) 
h. 'Scotia' 

' Splendor' 

' Suiraer Skies' 

'Sunset' (IT.) 

' Swartluaore' (IT.) 

'Turgot' 

' Yautan' 
josifleza 'Anna Amhoff (N.) 
j. 'Bellicent' 

'Guinevere' 

'ITcllie Boan' (N.) 

'Eoyalty' 

•Eutra' (N.) 
Joeikea 
J. 'Eximia' 
J. 'Pallida' 
J. 'Rubra' 
J. 'Zatel' 

J. X ? - 'Kim' (IT.) 
Julianae 

J. 'Hers Var.' (N.) 
Koiiiaro\irii 
laciniata 
1. X pinnatifida 
Meyeri 
microphylla 
m. ' Superle' 
nanceieina 'Floreal' 
n. 'Rutilant' 
otlata 
0. dilitata 
0. Giraldii (N.) 
0. G. 'Nana' (U.) 
pekinensis 
p. 'Pendula' 
persica 
p. 'Al"ba' 



h. 

h. 

h. 



h. 
h. 
h. 
h. 
h. 
h. 



Upton Nur. 1958 

Wayside Gard. 19^ 

Lemoine Nur. 1912 

Cottage Gard. 1933 

Arnold Arboretum 19^2 

Swarthmore 19^1 

Arnold ArTjoretum 19^ 

Arnold Artoretua 19^ 

Morton Arboretum 1961 

Lemoine ITur. 1917 

Morton Arboretum 19^ 

Swarthmore 1957 

Swarthmore 1957 

dc liessemaekcr 191^ 

Strawberry Hill 1963 

Swartlmore 1957 

Swarthmore 19^7 

C. V;. Burr ITur. 1952 

Swartl^more 1957 

Weston Nur. 1962 

Strawberry Hill 1963 

Arnold Arboretum 19^ 

Lemoine ITur. 191^ 

Colprit ITur. 1959 

Bobbink & Atkins 19^-2 

Arnold Arboretiaa 19^2 

Colprit ITur. 1959 
Zelsey IT-orsery Serv-. 1955 

Elan Mem. Park 1958 

Ellwanger & Barry 1892 

Barbie r Nux. 1902 

Arnold Arboretum 1915 

Transon Nur. 1892 

Sllwanger & Barry 1917 

Boerner Bot. Gard. 1958 

Upton lT\ix. 19^5 

Ji:ian Mem. Park 1959 

Arnold Arboretum 1912 

Spaeth ITur. 1908 

Upton ITvir. 1958 

Arnold Arboret'um 191^ 

Arnold Arboretum 1915 

Wayside Gard. 19^ 
1933 

Arnold Arboretum 19^3 

Upton IT\ir. 19^3 

Arnold Arboretum 1919 

Arnold Arboretum 19^^ 

Upton Nur. 195^ 

Arnold Arboretum 1913 

Elan Mem. Park 195^ 

Lemoine Nur. 1902 

iillwanger & Barry 1892- 



256 



PHTTOLOGIA 



Vol. 10, no. k 



pimiatifolia 


Arnold Arboretum 


1911 


PotaJiinii (H.) 


Eart & Vick 


1943 


Prestoniae 'Alice' 


Brand Nur. 


19^7 


P. 'Calparnia' 


Morton ArTjoret-um 


19^^ 


P. 'Charmian' 


Morton Arboretum 


19^ 


P. 'Coral' (N.) 


Upton ITur. 


1955 


P. 'Desdemona' 


S\vartlxnore 


19^7 


P. 'Doiiald Vfyman' 


Gardner llur. 


1950 


P. 'Elinor' (N.) 


Morton Arboretuiji 


1961 


P. 'Enid' (IT.) 


iilaji Kern. Park 


1958 


P. 'Ethel M. Wetster' 


(N.) Swartlimore 


1957 


P. Trancisca' 


Morton Arboretum 


19^ 


P. 'Hfl-ndel' 


Arnold Arborettun 


19^2 


P. 'Hecla' 


Arnold Arboretum 


19^2 


P. 'Horace' (IT.) 


Elan Mem. Park 


1958 


P. 'Isabella' 


Bobbink & Atkins 


19^2 


P. 'Jessica' 


Bobbink cS: Atkins 


19i^2 


P. 'Lucetta' 


Brand Nur. 


19^1 


P. 'Miranda' 


Snyder Bros. ITur. 


19ii7 


P. 'ITerissa' 


Arnold Arboretum 


19i^2 


P. 'Nocturne' 


Zelsey ITur. Service 


1955 


P. 'Oteron' 


Bobbink & Atkins 


19^2 


P. 'Paulina' 


Arnold Arboretum 


19'4-2 


P. 'Pack' 


Morton Arboretum 


19^ 


P. 'Red Wine' (11.) 


±jlan Mem. Park 


1959 


P. 'Romeo' 


Arnold Arboretum 


19^ 


P. 'Ursula' 


Arnold Arboretiom 


1948 


P. 'Valeria' (K.) 


Morton Arboretum 


1961 


P. 'Viola' 


Morton Arboretum 


l9iJ« 


putescens 


Lemoine ITur. 


1902 


reflexa 


Arnold Arboretum 


1912 


r. 'AllDa' 


Upton Nur, 


1943 


rhodopea 


Arnold Arboretum 


19^ 


SMraieri (U.) 


Svra,rthnore 


1957 


swegiflesa 


Arnold Arboretum 


19^+2 


Svreginzowil 


Arnold Arboretum 


1912 


S. 'AlMcLa' 


Jackson & Perkins 


1939 


S. 'Densiflora' (N.) 


Morton Arboretum 


1961 


S . ' Superta' 


Lemoine Hur. 


1916 


Tigerstedtii (IT.) 


V.'ageningen 


1956 


tomentella 


Arnold Arboretum 


1913 


velutina 


Arnold Arboretum 


1919 


villosa 


Jiillwanger & Barry 


1892 


V. 'Aurea' 


Morton Arboretum 


19^ 


V. Bretschneideri 


Col. Plum 


1925 


V. 'Rosea' 


Morton Arboretum 


19^*4 


V. X Sweginzo\d.i - 'Hedin' Swartlanore 


19i^l 


V. X S. 'Hunting Tower' 


(N.) Arnold Arboretum 


1956 



196U 



Harkness, Cultivated woody plants 



257 



SYRIIIGA 


con. 






vulgari 9 


Arnold Arboretum 


1909 


V. 


At el Carriere* 


Lemoine Kur. 


1900 


V. 


A. B. lamberton' 


Dunbar seedling 


1917 


V. 


Adelaide Dunbar' 


Dunbar seedling 


1916 


V. 


Admiral Farragut' 


Dunbar seedling 


1923 


■7. 


AlTsa' 






V. 


Al'ba &randi flora' 


Jiillwanger & Barry 


1892 


V. 


AllDa Virginalis' 


Ellwanger & Barry 


1892 


V. 


Altert The Good' 


lillwnnger & Barry 


1892 


V. 


Alexander Hamilton' 


Dunbar seedling 


1923 


V. 


Alice Harding' 


Swarthmore 


19^7 


V. 


Aline Mocq.ueri3' 


Spaeth ITur. 


1892 


V. 


Allison Gray' (K.) 


Morton Arboretum 


1958 


V. 


Alphonse Lavallee' 


Jiillwanger & Barry 


1892 


V. 


Am'bassade-ur' 


V/ayside Gard, 


19^ 


V. 


Amtroise Verscbaffclt' 


Morton Arboretum 


1925 


V. 


Amethyst' 


Spaeth Uur. 


1892 


T. 


Ami Schott' 


Clarke Nur. 


1938 


V. 


Amoena' 


Spaeth IT-ur. 


1892 


V. 


Anna Elizabeth Jacq.uet 


' Morton Arboretum 


I9ii8 


V. 


Anne Schiach' (K.) 


Gardner Imr. 


1950 


T. 


Anne Tighe' (N.) 


S\vrarthmore 


1957 


V. 


Archeveque' 


Arnold Arboretum 


19^ 


V. 


Arthur William Paul' 


Lemoine llur. 


1902 


V. 


Astra' 


Svrarthmore 


1947 


V. 


Aucuhaefolia' 


Col. Plum 


1925 


V. 


Aurea' 


Spaeth Nur. 


1892 


V. 


Azurea Plena' 


Spaeth Nur. 


1892 


V. 


Bfi.nciuise' 


Lemoine Nur. 


1907 


V. 


Belle de Nancy' 


Lemoine Nur. 


1908 


V. 


Be ranger' 


lllv;anger & Barry 


1892 


V. 


'Bicolor' 






V. 


Bleuatre' 


de Messemaeker 


191^ 


V. 


Blue Angel' (N.) 


M. Eaton 


i960 


V. 


'Boule Azure e' 


Col. Plum 


1925 


T. 


'Boussingault' 


Lemoine Nur. 


1902 


V. 


'Calvin C. Laney' 


Dunbar seedling 


1923 


V. 


•Candeur' 


Lemoine Nur. 


1950 


V. 


Capitaine Baltet' 


Col. Plum 


1925 


V. 


'Capitaine Perrault" 


Cottage Gard, 


1938 


V. 


Carmen' 


Lemoine Nur. 


1919 


V. 


Carmine' (H.) 


iilan Mem. Park 


1959 


V. 


'Caroli' 


Lemoine Nur. 


1900 


V. 


'Cavour' 


de Messemaeker 


191^ 


V. 


'C. B. Van Nes' 


Lemoine Nur. 


1905 


V. 


Champlain' 


Swarthjnore 


19^7 


V. 


'Charlemagne' 


Jiillwanger & Barry 


1892 


V. 


'Charles Baltet' 


Morton Arboretum 


195^ 


V. 


'Cliarles Joly' 


Lemoine Nur. 


1900 


V. 


'Charles Sargent' 


Lemoine Nur. 


1907 


V. 


•Charles X' 


Spaeth Nur. 


1902 


V. 


'Charlotte Morgan' (N.) Elan Mem. Park 


1958 



258 



PHITOLOGIA 



Vol. 10, BO. U 



SYEINaA 


con. 






V. ' 


Charm' 


Swarthmore 


1947 


V. " 


Christophe Colom"b' 


Lemoine Kiir. 


1907 


V. " 


City of Greshflpi' 


Arnold ArTjoretum 


19^ 


V. ' 


City of Longview' 


Arnold Arho return 


19^ 


V. ' 


Clara' 


Morton Arhoretum 


1944 


V. 


Clara Co chef 


de Messemaeker 


1914 


V. 


Clarence D. VanZandt' 


Duntar seedling 


1923 


V. 


Claude Le Lorrain' 


de Messemaeker 


1914 


V. 


Coerulea Super'ba' 


BllwR.nger & Barry 


1892 


V. 


Colbert' 


Lemoine Nur. 


1900 


V. 


Colmariensig' 


Ellwanger & Barry 


1892 


V. 


Col. Wm. R. Plum' 


Brand Hur. 


1947 


V. 


Comte Adrian de Monte"bello' Lemoine Nur. 


1912 


V. 


Comte de Kerchove' 


Lemoine Nur. 


1902 


V. 


Comte Horace de Choiseul 


' Lemoine Nur. 


1900 


V. 


Comtesse Horace de Choiseul' Dickson Hur. 


1892 


V. 


Condorcet' 


Lemoine llur. 


1902 


V. 


Congo' 


Lemoine Nur. 


1900 


V. 


Corinne' 


Lemoine Nur. 


1900 


V. 


Crampel' 


Lemoine Nux. 


1900 


V. 


Crepuscule' (N.) 


Swarthmore 


1957 


V. 


Croix de Brahy' 


Jiillwanger & Barry 


1892 


V. 


Dane Blanche' 


Lemoine Kur, 


1903 


V. 


Davm' 


Gardner Nur. 


1950 


V. 


Decaisne' 


Lemoine ITur. 


1914 


V. 


De Croncels' 


Farquhar Kur. 


1917 


V. 


De Humtoldt' 


Lemoine Nur. 


1900 


V. 


De Jussieu' 


Lemoine Nur. 


1900 


V. 


De Louvaln' 


de Messemaeker 


1914 


V. 


De Miriljel' 


Lemoine Nur. 


1905 


V. 


De Saussure' 


Lemoine Nur. 


1905 


V. 


Desfontaines' 


Morton Artoretum 


1954 


V. 


Deuil d'Emile Gall^' 


Lemoine Nur. 


1905 


V. 


Diane' (ll.) 


A. McKean 


1958 


V. 


Diderot' 


Lemoine Nur. 


1916 


V. 


Dillia' 


Morton Arhoretum 


1948 


V. 


Dipl ornate' 


Euliffson 


1953 


V. 


Dovmfield' (N.) 


M. Baton 


I960 


V. 


Doyen Keteleer' 


Lemoine Nur. 


1900 


V. 


Dr. Charles Jaco"bs' 


de Messemaeker 


1914 


T. 


Dr. Lindley' 


Ellwanger & Barry 


1892 


V. 


Dr. Maillot' 


Lemoine Nur. 


1902 


•7. 


Dr. Mastets' 


Lemoine Nur. 


1902 


V. 


Dr. Notte' 


Dickson Nur. 


1892 


V. 


Dr. Troyanowsky' 


Lemoine Nur. 


1904 


V. 


'Dr. von Regel' 


Transon Nur. 


1892 


V. 


Dresden Clilna' 


Morton Arhoretum 


1954 


V. 


Due de Massa' 


Lemoine Nur. 


1907 


V. 


Eaton Eed' (N.) 


M. Eaton 


I960 


V. 


Eden' 


Clarke Nur. 


1940 


V. 


Edith Cavell' 


Lemoine Nur. 


1917 


V. 


Edmond Ahout' 


Lemoine Nur. 


1900 



196Ii 



Harkness, Cultivated woody plants 



259 



SYRimk, 


con. 






V. ' 


EcLiond Boissier' 


Lemoine Nur. 


1907 


V, ' 


Edna Dimbaai' (IJ. ) 


Dr. Eankin 


1955 


V. ' 


Bdoiiard Andre' 


Lemoine ITur. 


1902 


V. ' 


Ed;vard J. G-ardner' 


Gardner ITur. 


1950 


V. ' 


Elceiiholm ' 


Spaeth ITur, 


1892 


V. ' 


Elihii Root' 


Buntar seedling 


1923 


V. ' 


Ellen Willmott' 


Lemoine Nur. 


1905 


V. 


Ellie44arie' (N.) 


Svrarthmore 


1957 


T. ' 


Enil Lie-big' 


Spaeth ITur. 


1892 


V. 


Bnile Gentile' 


Lemoine Nur. 


1916 


V. ' 


Emile Lenoine' 


Lemoine ITur. 


1902 


V. 


Erzlierzog Johann' 


Spaeth ITur. 


1892 


V. ' 


Etna' 


Swarthmore 


19^1 


V. 


Etoile de Mai' 


Lemoine llur. 


1907 


V. 


Farrionensis' 


Spaeth ITur. 


1892 


V. 


Fimament' 


Clarke Hur. 


1938 


V. 


Jraicheur' 


Lemoine ITur. 


1950 


V. 


Francisq.ue Morel' 


Lemoine Kur. 


1900 


V. 


Frank KLager' 


Arnold Ar "bore turn 


19^ 


V. 


Frank Patterson' (IT.) 


Ellesmere Nur. 


1962 


V. 


Frau. Bertha Dasimann' 


Ellwanger & Barry 


1892 


V. 


Frau Wilhela Pfitzer" 


de Messemaeker 


1912 


V. 


Fred Payne' (N.) 


Dr. Eankin 


1958 


V. 


Fritz' 


Morton ArTsoretun 


195^ 


V. 


Fuerst Bleucher' (N.) 


Timm ITur. 


1951 


V. 


Fuerst B-aelow' (U. ) 


Morton Arhoretum 


1952 


V. 


Faerst Liechtenstein' 


Spaeth ITur. 


1908 


V. 


Gaudichatid' 


Lemoine Kur. 


1905 


V. 


Geant des Batailles' 


Spaeth Nut. 


1902 


V. 


Gelieimrat Heyder' 


Trail son ITur. 


1892 


V. 


Geheimrat Singelmann' 


Spaeth ITur. 


1892 


V. 


General Drouot' 


de Messemaeker 


191^ 


V. 


General Elwell S. Otis 


Dunhar seedling 


1923 


V. 


General OraJit' 


Dunhar Seedling 


1917 


V. 


G-eneral Jolm Pershing* 


Ihinhar Seedling 


1917 


V. 


'G-enerail Kitchener' 


Ihinhar seedling 


1917 


V. 


'G-eneral Pershing' 


Clarke Nur. 


19^ 


V. 


General Sheridan' 


Danhar seedling 


1917 


V. 


'General Sherman' 


Dunhar seedling 


1917 


■7. 


'George W. Aldridge' 


Dunhar seedling 


1923 


V. 


'Georges Bellair' 


Lemoine ITur. 


1902 


V. 


•Georges Claude' 


Swarthmore 


19^7 


V. 


'Seminal' (IT.) 


Elan Hem. Park 


1958 


V. 


' Gigantea 


Bll\iranger & Barry 


1892 


V. 


'Gil-bert' 


Lemoine ITur. 


1912 


V. 


'Gismonda' (N.) 


Lemoine Nur. 


1950 


V. 


'Gloire de Lorraine' 


iJllwanger & Barry 


1892 


V. 


•Gloire de la Eochelle' 


Transon Nur. 


1892 


V. 


'Gloire de Moulins' 


Ellwanger & Barry 


1892 


V. 


'Glory' 


Ruliffson 


1950 


V. 


'&odron' 


Lemoine Nur. 


1910 


V. 


'Goliath' 


Spaeth Nur. 


1902 


V. 


'Grace Orthwaite' 


Brand Nur. 


19^ 



260 



PHYTOLOGIA 



Vol. 10, no, U 



SYRINGA, 


con. 


V. ' 


Grand Due Constajitin' 


V. ' 


Guizot' 


V. ' 


Hallelujah' (K.) 


V. ' 


Hazel Opper' (N.) 


V. ' 


Heather" (W.) 


V. ' 


Helen Schoen' (N.) 


V. ' 


Henri Martin' 


V. ' 


Henri HolDert' (H.) 


V. ' 


Henry Clay' 


V. ' 


Henry Ivadsworth Longfellov;' 


V. ' 


Henry V/ard Beecher' 


V. ' 


Heroan Bilers' 


V. ' 


Hippolyte Haringer' 


V. ' 


Hiram H. Edgerton' 


V. 


Hugo d© Yries' 


V. ' 


Bigo iioster' 


V. ' 


Hyazinthenfleider' 


V. ' 


J. de Messeniaeker' 


V. 


Jacq.ues Callot' 


V. 


James Booth' 


V, 


James Stuart' 


V. 


Jan3 Day' (H. ) 


V. 


Jan van Tol' 


V. 


Jean Bart' 


V. 


Jean Mace' 


V. 


Jeanne d'Arc' 


V. 


Jessie Gardner' (K.) 


V. 


Joan DunTsar' 


V. 


Jules Ferry' 


V. 


Jules Simon' 


v.«, 


Julien Gerardin' 


V. 


Justi' 


V. 


Kate Harlin' 


V. 


Katharine Haveneyer' 


V. 


'Konigin lAiise' (ll.) 


V. 


Lady Lindsay' (N. ) 


V. 


' Lamarck' 


V. 


'La Mauve' 


V. 


'Languis' 


V. 


'Laplace' 


V. 


'La Tour d'Auvergne' 


T. 


'Laura L. Barnes' (lI.) 


V. 


'Lavoisier' 


V. 


'Le Gaulois' 


V. 


'Lemoinei' 


V. 


'Le Notre' 


V. 


'Leon Gamtetta' 


V. 


'Leon Mathieu' 


V. 


'Leon Simon' 


V. 


'Leopold II' 


V. 


'Le Printemps' 


V. 


'Lilarosa' 



Lemoine Nur. 1900 

Lemoine Hur. 1900 

M. Eaton I960 

Dr. Ranldin 1955 

M. Eaton I960 

Ellesmere Hur. 1962 

Lemoine Nur. 191^ 

Swarthaore 19^7 

Dun^bar seedling 19§3 

DunlDar seedling 19§0 

Dunl)ar seedling 1923 

Kobankie Nur. 1938 

Lenoine ITur. 1910 

DunTjar seedling 1919 

Clarke llur. 19^ 

Kallen & Lunneman 191^ 

Baroier Kur. 1908 

de Messemaeker 191^ 

Ellv/anger & Barry 1892 

Col. Plum 1925 

Arnold Ar'Doretum 19^ 

Dr. Rankin 1958 

Cole llur. 193^ 

de Messemaeker 1908 

Lemoine llur. I916 

Lemoine Kur. 190^1- 

Boerner Bot. Gard. 1958 

Don^bar seedling 1923 

Ellwanger & Barry 1917 

de Messemaeker 191^ 

Lemoine Kur. 1917 

Arnold Artoretum 1919 

de Messemaeker 191^ 

Col. Pl-om 1925 

Morton ArlDoretura I96I 

Arnold Artoretum 1956 

Ellv;anger £: Berry 1892 

Lemoine Hur. 1900 

Jiillwanger & Barry 1892 

Lemoine Uur. 191^ 

Transon Nur. 1892 

Boerner Bot. Gard. 1958 

Lemoine llur. 191^ 

iiillwanger & Barry 1892 

iiillv;anger & Barry 1892 

Col. Plum 1925 

Lemoine Hur. 1908 

de Messemaeker 191^ 

Transon ITur. 1892 

cie Messemaeker 191^ 

Lemoine llur. 1902 

Spaeth Uur. 1892 



196U 



Harkness, Cultivated wocxfy plants 



261 



NGA 


con. 


V. 


Llcne' 


■7' 


L'Oncle Tom" 


V. 


Louis Henry' 


V. 


Lucie Baltet' 


V. 


Ludwig Spaeth' 


V. 


Macro stachya' 


V. 


Madeleine Lemalre' 


V. 


1-Iagellan' 


V. 


Marceau' 


T. 


Marc Micheli' 


V, 


Marechal de Bassompierre' 


T. 


Mareclvi.1 Toch' 


V. 


Marechal Lanneg' 


V. 


Marengo' (N.) 


V. 


Marie Finon' 


V. 


Marie Legraye' 


V. 


Marleyensis' 


V. 


Marleyensis Pallida,' 


V. 


Martha Kounze' (N. ) 


V. 


l-Iassena' 


V. 


Mathieu de Doa'basle' 


V. 


Ifeud llotcutt' (N.) 


V. 


Maurice Barres' 


V. 


Maurice de Vilmorin' 


V. 


Mauve Mis.t' (N. ) 


V. 


1-Iaxime Comu' 


V. 


Maximowicz' 


V. 


Michel Buchner' 


V. 


Midwest Gem' 


V. 


Milton' 


V. 


Miraheau' 


V. 


Mireille' 


V. 


Missrao' 


T. 


Mile. Ferr^nde Viger' 


V. 


Mile. Melide Laurent' 


V. 


Mme. Ahel Chatenay' 


V. 


Mme. A. J. Zlettenherg' 


V. 


Mne. Amelie Duprat' 


V. 


Mme. Antoine Buchner' 


V. 


Mme. Aiieraste G-ouche.ult' ( 


V. 


Mne Briot' 


V. 


Mme. Casinir Perier' 


V. 


Mme. Catherine Bruchet' 


V. 


Mme. Charles Souchet' (S 


V. 


Mme. de Miller' 


V. 


Mine. Fallierea' 


V. 


Mme Felix' 


V. 


Mme. Florent Stepraan' 


V. 


Mme. F. Morel' 


V. 


Mme. Henri G-aillard' 


V. 


Mme. Jules Finger' 


V. 


line. Krouter' 


V. 


I-Jme. Lemoine' 



Lemoine llur. 


1902 


Lemoine ITur. 


190ilf 


Lemoine ITur. 


1900 


de l-Iessemaeker 


191^ 


Heards Nux. 


1955 


Lemoine Nur. 


1900 


Jackson $ Jerklna 


1939 


Lemoine Nur. 


1919 


Lemoine Umx, 


1914 


Lemoine Hur. 


1902 


e' Lenoine Hue. 


1900 


Cottage Gard, 


1938 


de Messemaeker 


1914 


Upton llur. 


1958 


Cottage Gard. 


1938 


iillwa.nger & Barry 


1892 


Ellwanger & Barry 


1892 


Spaeth ITur. 


1902 


M. Baton 


i960 


Morton Ar "bore turn 


1927 


Bar tier Hur. 


1902 


ITotcuti Nur. 


1958 


Lemoine Nur. 


1919 


Lemoine llur. 


1902 


M. Eaton 


i960 


Arnold Arhoretum 


19^ 


Lemoine ITur. 


1907 


Kllv/anger & Barry 


1892 


Morton Arhoretum 


1954 


de Messemaeker 


1914 


Lenoine l"ur. 


1912 


Lenoine llur. 


1905 


Wayside Gard. 


19^ 


Bar "bier Uur. 


1902 


Lemoine ITur. 


1900 


Lemoine llur. 


1900 


Morton Arhoretum 


1954 


Goucliault & Turhat 


1907 


Lemoine ITur. 


1910 


(N.) M. Eaton 


i960 


Lenoine ITur. 


1900 


Lemoine ITur. 


1910 


de Messemaeker 


1914 


(N.) Lenoine ITur. 


1950 


Lemoine ITur. 


1905 


Morton Arhoretaan 


19^46 


Cottage Gard. 


1938 


de Messemaeker 


1914 


Lenoine ITur. 


1902 


Koliankie ITur. 


1938 


Dickson ITur, 


1892 


Spaeth ITur. 


1892 


Lemoine Hur. 


1900 



262 



PHYTOLOGIA 



Vol. 10, no, U 



SYHINGA, con. 



Mme. Leon Simon' 


Lemoine Nur. 


1905 


«Mme. Moser' 


Ellwanger & Barry 


1892 


'Mme. E. Foyer' 


Maarse 


1953 


' Monge ' 


Lemoine Nut. 


191'^' 


'Monique Lemoine' 


Swarthmore 


19^7 


'Mons. LePage' 


Barrier Nur. 


1902 


'Montaigne' 


Lemoine Nur. 


1908 


'Mont Blanc' 


Lemoine Nux. 


1916 


'Montgolfier' 


Morton Arhoretun 


195^ 


' Monument ' 


Clarke Nur. 


1938 


'Monument Camot' (JS.) 


Brooklyn Bot. Gard. 


19^^-3 


'Mood Indigo' (N.) 


Swarthmore 


1957 


'Moonlight' (N.) 


Swarihmore 


19i^7 


'Motmtain Haze' (N.) 


Morton Arhoretum 


1958 


'Mrs. Calvin Coolidge' 


M. Franklin 


1950 


'Mrs. Edward Harding' 


Clarke Nur. 


19^ 


'Mrs. Flanders' (ll.) 


Arnold Arh ore turn 


1956 


'Mrs. John S. Willians' 


S;i;arthmore 


19i^7 


'Mrs. Watson Weh"b' 


Swarthmore 


19^7 


'Mrs. W. E. Marshall' 


Col. Plum 


1927 


'Murillo' 


Lemoine Nur. 


1902 


'My Favorite' 


Arnold Arhoretum 


19U8 


'Nana' 


Ellwanger & Barry 


1892 


'Hancy Frick' 


Swarthmore 


19^7 


'Uaudin' 


Lemoine Nur. 


1914 


' Negro ' 


Lemoine Nur. 


1902 


'Night' 


Swarthmore 


19^1 


'Nigricans' 


Transon Nur. 


1892 


'Noisettiana Alba' 


iiillwRjiger & Barry 


1892 


'Ohelisque' 


Lemoine Nur. 


1900 


'Olivier de Serres' 


Lemoine Nur. 


1910 


'Ostrander' 


Merrill 


1930 


'Othello' 


Lemoine Nur. 


1905 


'Pasteur' 


Lemoine Nur. 


1905 


'Patrick Henry' 


Dunhar seedling 


1923 


'Paul Deschanel' 


Cottage Gard. 


1938 


'Paul Hariot' 


Lemoine Nur. 


1905 


'Paul Thirion' 


Lemoine Nur. 


1916 


'Peau de Chamois' 


bwarthmore 


1947 


'Perle von Stuttgart' 


de Messemaeker 


1914 


'Perle von Teltow' 


KH,llen & Lunneman 


1914 


' Philemon' 


Lemoine Nur. 


1900 


'Pierre Joigneux' 


Lemoine Nur. 


1902 


'PinMe' (N.) 


Dr. Rankin 




'Pink Mist' (N.) 


M. Eaton 


i960 


• Planchon' 


Lemoine Nur. 


1910 


'President Camot' 


Lemoine Nur. 


1900 


•President Fallieres' 


Lemoine Nur. 


1912 


' President G-revy' 


Ellwanger & Barry 


1892 


'President Harding' 


Dunhar seedling 


1922 


'President John Adams' 


(N.) Bunhar seedling 


1923 


'President Lamheau' 


de Messemaeker 


1914 



196U 



Harioiess, Cultivated woody plants 



263 



SYRIUGA 


con. 






V. 


President LeBrun' (ll.) 


Arnold AtTdo return 


1956 


V. 


President Lincoln' 


IXm'bar seedling 


1916 


V. ' 


President Loutet' 


Lemoine llur. 


1905 


V. ' 


President Massart' 


iillwanger & Barry 


1892 


V. 


President Monroe' 


D-anbar seedling 


1923 


V. 


President Poincare' 


Lemoine ITur. 


191^^ 


V. " 


President Roosevelt' 


Dun'bar seedling 


1919 


V. 


President Viger' 


Lemoine Nur. 


1902 


V. 


Primrose' (i:.) 


Wayside Gard. * 


1957 


V. 


Prince de Beauveau' 


Lemoine Kur. 


1900 


V. 


Prince Iraperial' 


Morton Arhoretum 


192i* 


V. 


Prince Notger' 


Jiillwanger & Barry 


1892 


V. 


Prince of Wales' 


Ellwflnger & Barry 


1892 


V. 


Princess Ale::andra' 


i!;llwR.nger & Barry 


1892 


V. 


Princesse Camilla de Rohan' Parsons Kur. 


1892 


V. ' 


Princesse Clementine' 


de Messemaeker 


1914 


V. 


Princesse Marie' 


Lemoine Nur. 


1900 


V. ' 


Prinzessin Klotilde' {ll.) Timrn l^ur. 


1951 


V. 


Priscilla' 


Swartlinore 


19^7 


V. 


Prodige' 


Clarke ilur. 


1938 


V. 


Prof. 3. H. Wilson' 


Arnold Arhoretum 


19^ 


V. 


Prof. E. Stoelchardt' 


Ellwanger & Barry 


1892 


V. 


Prof. Sargent' 


Spaeth Hut. 


1902 


V. 


Pyramidalis' 






V. 


Pyramidalis Al"ba' 


iillwanger & Barry 


1892 


V. 


Q,uadricolor' 


Spaeth ICur. 


1892 


V. 


Ratelais' (N. ) 


Morton Ar'boretum 


1958 


V. 


Reaumur ' 


Lemoine IJur. 


1905 


V. 


Red Feather' 


Ruliffson 


1953 


V. 


Reine Elizabeth' 


ae Messemaeker 


1914 


V. 


Reine Margnierite' 


Arnold Arhoretum 


19^ 


T. 


Rene Jarry Desloges' 


Lemoine Nur. 


1907 


V. 


Renoncule' 


Moon Hut. 


1913 


V. 


Bochamheau' (H. ) 


Morton AtTd ore turn 


1927 


V. 


Roi Albert' 


de Messemaeker 


1914 


V. 


Romance' (H.) 


M. Baton 


i960 


V. 


Ronsard 


Lemoine Kur. 


1914 


v.. 


Rosea Grcmdiflora' 


Trans on ITur. 


1892 


V. 


Rose a Grand Pleur' 


ae Messemaeker 


1914 


V. 


Ro-uge de Trianon' 


ae Messemaeker 


1914 


V. 


RuTDella Plena' 


Ellwanger & Barry 


1392 


V. 


Ru^ora Insignis' 


Elluanger & Barry 


1892 


V. 


Ruhm von Eorstensteln' 


Ruliffson 


1937 


V. 


Rustica' 


Lemoine llur. 


1950 


V. 


Sarali Sands' 


Swarthmore 


1941 


V. 


Sat-ornf.le' 


Lemoine ITur. 


1917 


V. 


Schermerhom' 


Parsons Nur. 


1892 


V. 


'Scipion Cochet' 


Ellwanger & Barry 


1892 


V. 


'Senateur Volland' 


Dickson ilur. 


1892 


V. 


' Sensation' 


Kluis Nur. 


1952 


V. 


'Serene' (N.) 


M. Eaton 


i960 


V. 


' Sieljold' 


lemoine Nur. 


1907 



26U 

SIEIIIGA, con. 



PHYTOLOGIA 



'Silver Kir^' (U.) 

'Snowflake' (U.) 

'Souv. de Claudius G-raindorge 

' Souv. de Georges Traffaut' 

' Soutr, de Henri Sinon 

'Souv. de Louis Chasset' (N. 

'Souv. de Louis Thitaut' 

'Souv. de Simono' 

'SpecWtilis' 

' Stadtgartner Eothpletz' 

'Susan B. Anthony' 

'Sv;eetheart' (IT.) 

'Taglioni' 

'Thomas A. Edison' 

'Tliomas Jefferson' 

' Thunherg' 

' Todmorden' 

•Tournefort' 

'Toussaint L'Ouverture' 

'Triomphe de Moulins' 

'Triomphe d' Orleans' 

'Triste Barharo' (H.) 

'Turenne' 

'Yaletteana' 

'Van Aerscliott' 

' Yergissmeinnicht' 

'Tersalicnsis' 

•Verschaffelti' 

'Vestale' 

' Vesuve' 

'Victor Lemoine' 

'Ville de Limoges' 

'Ville de Troyes' 

'Violacea' 

'Violetta' 

'Virginia Becker' 

'Virginite' (IT.) 

'Vi viand Morel' 

'Vivian Evans' 

' Volcan' 

' Waldeclo-Eousseau' 

'Weddle' 

'VJhite Svran' 

'Vfilliara G. Barry' 

' William Eohinson' 

'William S. Riley' 

'W. T. Lee' (IT.) 

•Zukunft' (n.) 

'Zulu' (N.) 
Wolfii 
W. hirsuta 
yunnanensis 



L Vol. 


10, no. 


Heards ITur. 


1955 


M. Eaton 


i960 


' Maarse 


1953 


(iT. ) V/eston Uur. 


1962 


Col. Plum 


1925 


) Weston ITur. 


1962 


-Wllv.'anger & Barry 


1899 


Morton Arhoretum 


19^ 


Parsons ITur. 


1892 


Barhier ITur. 


1908 


Dunhar seedling 


1923 


V/eston llur. 


1962 


Lemoine Uur. 


1907 


Bunhar seedling 


1922 


Dunhar seedling 


1922 


Lemoine Nur. 


1914 


Sv/arthmore 


1947 


Lemoine ITur. 


1900 


Transon llur. 


1892 


Spaeth Nur. 


1908 


Morton Ar^boretum 


1958 


Lemoine Nur. 


1917 


de MessemaekEr 


1914 


Spaeth ITur. 


1892 


Spaeth ITur. 


1892 


Ellvio.nger & Barry 


1892 


Lemoine ITur. 


1914 


Lemoine Nur. 


1917 


Lemoine Uur. 


1907 


Parsons ITiir. 


1892 


Ellv,'anger & Barry 


1892 


Transon Nur. 


1892 


Lemoine Nur. 


1917 


Upton Nur. 


1954 


Morton ArTioretum 


1958 


Lemoine Nur. 


1905 


Heard Nux. 


1955 


Lemoine Nur. 


1900 


Lemoine Nur. 


1905 


Morton ArlDoretum 


19^- 


Arnold Artoretum 


19^ 


Duntar seedling 


1917 


Lemoine Nur. 


1900 


Dunhar seedliiig 


1922 


Ellsmere Niir. 


1962 


Boerner Bot. Gard. 


1958 


M. Eaton 


i960 


Arnold Ar "bore turn 


1913 


Arnold Arhoretum 


1920 


Upton Nur. 


1943 



196U 


Harkness, Cultivated woody plants 


265 


SmilTGA 


, con. 






yuiuaanensis 'Rosea' 


Morton Ar"boretum 


19i^7 


M-IARIX 


- Tanaricaceae 






Jmiiperina 


iJll\iranger & Boxry 


1892 


parvi flora 






pen 


bandra 


■BJllwanger & Barry 


1892 


TAXOEIUll - Hnaceae 






dis 


bichun 


Ellwflnger & Barry 


1897 


d. 


Pendens' 


Ellwanger & Barry 


1898 


TAXUS - 


Ta^aceae 






"baccata 


Ellv/anger & Barry 


1897 


t. 


Adpressa Stricta' 


Arnold Arboretum 


1906 


•b. 


Aurea' 


Ellwflneer & Barry 


1897 


I. 


Balkans' (IT.) 


Missouri Bot. &ard. 


195^^ 


•b. 


Ciieslrontensis* 


Dickson ITur. 


1907 


b. 


Conpe.cta' (U.) 


Secrest Arhoretum 


1958 


b. 


Dovastoniana' 


Dickson llur. 


1907 


t. 


Erecta' 


i!illvm,nger & Barry 


1897 


t. 


Ericoides' 


BarlDier ITur. 


1911 


Td. 


Pastigiata T&riegata' 






Id. 


Glauca' 


iillwanger & Barry 


1897 


Id. 


Jackson' 


Barhier ITur. 


1911 


t. 


Nedpath Castle' 


BarlDier IT-ur. 


1911 


t. 


Fend-Jla' 


Arnold Artoretum 


1905 


t. 


Repfl.n dens' 


Bartier ITur. 


1911 


t. 


Variegata' 


Barrier ITur. 


1911 


t. 


Washington' 


JiJllwanger & Barry 


1897 


"bre^ 


trifolia 


fi. Fillmore 


1958 


C£LU< 


idensis 


Arnold Arhoretun 


1901 


CUSj 


pic^ita 


Ellv/anger & Barry 


1897 


c. 


Aflpins' (N.) 


Secrest Artcretum 


1958 


c. 


Densa' (IT. ) 


Secrest ArTsoretum 


1958 


c. 


Minima" 


Rochester seedling 




c. 


liana' 


Arnold Ar^boretum 


1896 


c. 


Prostrata' (N.) 


Secrest Arhoretum 


1958 


c. 


Thayer' 


Arnold Arh ore turn 


1930 


Hiarj 


'.ev;ellianA 


Arnold Arhoretum 


1930 


laed 


La 


Arnold Artoretum 


1930 


m. 


Amherst' (N.) 


Mitiska ITur. 


1957 


n. 


Andorra' (IT. ) 


Secrest Arhoretun 


1958 


m. 


Bottink' (N.) 


Secrest ArTsoretum 


1958 


m. 


Cole' (11.) 


Secrest Arhoretum 


1958 


m. 


'Dutweiler' (IT.) 


Secrest Arhoretum 


1958 


m. 


•Flener' (IT.) 


Secrest Artoretum 


1958 


m. 


'Hatfield' 


Arnold Arhoretum 


1930 


m. 


Hatfield #23' (IT.) 


Secrest Arhoretum 


1958 


m. 


'Hicks' 


Hicks Kur. 


1920 


m. 


'Setian' (11.) 


Secrest Artoretum 


1958 


m. 


' Slavin' 


B. H. Slavin sel. 




m. 


'Ward' (U.) 


Secrest Arhoretum 


1958 



266 



PHYTOLOQIA 



Vol. 10, no. U 



THUJA - Pinaceae 






koraiensis 


Arnold Arhoretum 


1930 


occidental is 


D. Hill N\ir. 


1904 


0. 'Bodmer' (N.) 


Morton Artoretum 


1951 


0. 'Booth' 


Barhier IJur. 


1911 


0. 'Buchanan.' 


Arnold Arhoretum 


1921 


0. 'Columbia' 


Bohhink & Atkins 


1927 


0. 'Compacta' 






0. 'Cristata' 


Barhier Eur. 


1911 


0. 'Douglas Aurea' 


D. Hill Nur. 


1927 


0. 'Douglas Pyramidnlis' 


iillwanger & Barry 


1897 


0. 'Ellwanger' 


Ellwanger & Barry 


1897 


0. 'Fastigiata' 


Ellvffijiger & Barry 


1897 


0. 'G-lohosa' 






0. 'Gracilis' (N.) 


U. S. D. A. 


1962 


0. 'Eetz Midget' 


Arnold Arhoretum 


1958 


0. 'Ho opes' 


Biltaore Nur. 


1902 


0. 'Hovey' 


Little Kur. 


1897 


0. 'Hudsonica' 


Biltmore Hur. 


1902 


0. 'Leptoclada' (N.) 


Morton Arhoretum 


1951 


0. 'Lutea' 


Arnold Arboretum 


1896 


0. 'Masters' 


Arnold Arboretum 




e. 'Nigra' (N.) 






0. 'Ohlendorff (U.) 


Secrest Arboretum 


1958 


0. 'Pendula' 


Arnold Arboretum 


1907 


0. 'Pumila' (11.) 






0. ' Recur va lilana' (H.) 


H. Y. Bot. Gard. 


1951 


0. 'Reeves' 






0. 'Eeid' 






0. 'EiTers' 


Bobbink & Atkins 


1923 


0. 'RolDusta' 


Arnold Arboretum 


1896 


0. 'Rosenthal' 






0. 'Spiralis' 


Bobbink & Atkins 


1923 


0. 'Uiibraculifera' (H. ) 


Morton Arboretum 


1951 


. ' Vervaeneana' 


Ellwfinger & Barry 


1897 


0. ' Vfagneriana' 


Bar bier Nur. 


1911 


0. 'V^oodward' 


D. Hill Nur. 


1927 


orientalis 


Arnold Arboretum 


1902 


0. 'Aurea' 


D. Hill Nur. 


1927 


0. 'Aurea Nana' 


Princeton Nur. 


1927 


0. 'Bonita' 


D. Hill Nur. 


1927 


plica ta 


Arnold Arboretum 


1396 


p. 'Atrovirens' (ll.) 


11. Y. Bot. Gard. 


1951 


p. 'Hillier' (l^I.) 






Standi shii 


Arnold Arboretum 


1902 


THUJOPSIS - Pinaceae 






dolohrata 


Veitch Nut. 


1903 



TILIA - Tiliaceae 
americaiia 
a. 'Fastigiata' 
car©iiniana rhoophila 



Horsey, Metamora, Ohio 
Slavin selection 
Arnold Arboretum 



1919 



196h 



Harkness, Cultivated woody plants 



267 



TILU, con. 
cordata 
euchlora 
europea 

e. 'Ebnii^gslinde' 
floridana 
heterophylla 



h. Micbauxii 

japonica 

mandshurica 

MaxiIno^^d c z iana 

Miqueliana 

Moltkei 

mongolica 

Qonticola 

neglecta 

Oliver! 

or"bicularis 

petiolaris 

platyphyllos 

P 



(IT.) 



'Fastigiata' 
'Laciniata' 
'Rubra' 
' Tortuosa' 
'Vitifolia' 



Tuan 

TOREEIA - Taxaceae 
nucifera 

TEIEETALEIA - Ericaceae 
"bracteata 

TSUGA - Pinaceae 
canadensis 
c. 'AtroTirens' 
c. 'Gracilis' 
c. 'Pendula' 
c. 'Pumila' 
caroliniana 
diveraifolia 
Sieboldii 

ULI-IUS - Ulmaceae 
anericana 
a. 'Ascendens' 
caxpinifolia 
c. comutiensis 
' Suterosa' 
umbraculifera 
'Variegata' 
'Wetliiaiia' 
'Wrede' 



Veitch llTir. 


1907 


Ellv;anger & Barry 


1906 


Veitch "ur. 


1907 


Grootendorst Kur. 


1952 


Arnold Arboretion 


1921 


Horsey, Steubenville, 


Ohio 




1916 


Arnold Ar'ooretujn 


1920 


Ellwfinger & Barry 


1906 


Zomik 


1959 



Jfillv/anger & Barry 



1906 



Arnold Arho return 
Bartier llur. 


1913 
1907 


Arnold Artoretum 
Ellwanger & Barry 


1911 
1919 


Grootendorst Nur. 
Arnold Arhoretum 
iJllwanger & Barry 
Arnold Arhoretum 


1956 
1915 
1906 
191^ 


Veitch Kur. 


1907 


Edin"burgh 


1952 


Rothins Nur. 
Hicks ITur. 
BohlDink & Atkins 
Arnold Arboretum 


1922 
1917 

1923 
1901 


Zelsey Nur. 
BarMer ITur. 
Veitch ITur. 


1899 
1903 
1899 


Brown Bros. ITur. 
B. H. Slavin sel. 


1919 
1919 


Spaeth Nur. 
Arnold Arhoretum 
ELehms Nur. 


1908 

191^ 


Spaeth Nur. 
J^illwanger & Barry 


1908 

192'* 



268 



PHITOLOQIA 



Vol. 10, no. (^ 



UIMJS, con. 






gla"bra 






g. 'Dovae' 


Ellwacger & Barry 


1907 


g. 'Ezoniensis' 






g . ' Nana' 


Arnold Artoretua 


1907 


g. 'Pendula' 


Arnold Artoretum 


1923 


hollandica raajor 


Spaeth Nur. 


1908 


h. 'Pitteurs' 






h. 'Saperlsa' 


Arnold ArTDoretum 


1923 


h. 'Vegeta' 


iillvv-anger & Barry 


1907 


japonlca 


Arnold Ar'boretvun 


1907 


laevls 


Spaeth ITur. 


1908 


macrocarpa 


Arnold ArTsoretum 


191^ 


parvifolia 


Arnold Artoret-um 


1906 


procera 


Mr. Ellwai'i^er planted 


l8i^3 


p. 'Viniinelis' 


Spaeth Nur. 


1908 


pujLlla 


U. S. D.-A. 


1918 


p. ar"borea 


Arnold ArtoretuQ 


1905 


ru'ora 






serotina 


Riverview Kur. 


1915 


Thonasii 


Forest I'ur. 


1919 


Wilsoniana 


Arnold Ar "bore tun 


191'^' 



VaCCIHEUI.I _ Ericaceae 
staminerun 



Kelsey Nur. 



1892 



VIBURlTUl-l - Caprifoliaceae 
acerifolium 
a. ovatiun 
"betulifoliuia (ll.) 
"brae tea turn (N.) 
"buddleifoliun 
■burejaticiun (IT.) 
Burlcwoodii 
B. 'Cheimult' 
carlcephalmi 
Carlesii 
cassinoides 
cotinifolia (H.) 
dentatum 
d. puuescens 
dilitatur.1 

d. 'Xanthocarpura' (lI.) 
eru"bescens gracilipes 
fra^'rans 
f. 'llanum' 
Juddii 
Lantana 
L. 'R-ugosum' 
Lentago 

L. sphaero carp-am 
lotophyllun 
macroceplial'oiu 'Sterile' 



Morton ArTDoretuQi 
Veitch llur. 
Morton Artoretim 
Clarke ITur. 
Grootendorst "ur. 
Wayside Gard. 
Lemoine ITur. 

Dr. Egolf 

J. I)av;son 
Spaeth llur. 
Lemoine llur. 
Arnold ArTrjoretujn 
Arnold ATtoretuni 
Arnold Artoret-oin 

ffrootendorst ITur. 
iillviranger & Barry 

J. Daxifson 
Arnold Arhorct-um 
Arnold Artorctuni 
Trans on Imr. 



1951 
1911* 
19^9 
19^^*0 
1951 
1952 
1907 

1959 
1892 
1892 
1902 
195^ 
1915 
1913 

1951 
1893 

1892 
1921 
1920 
1892 



196U 



Harkneis, Cultivated woody plants 



269 



VIBURiroi'i , con. 



molle (IT.) 


Dr. Egolf 


1959 


iuongolicum (N.) 


Morton Artoretutt 


1951 


Opulus 


Ellvffinger & Barry 


1892 


0. ' Compact-um' 


Wayside Gard. 


1957 


0. 'Notcutts Var.' (IT. ) 


Dr. Egolf 


i960 


0. 'Roseua* 


iillwFinger & Barry 


1892 


0. 'Xaathocarpum' 


Arnold Artoretuin 


1915 


orientale (IT.) 


Dr. Egolf 


i960 


plicat'jn 'Plicatum' 


JiJllWringer & Barry 


1892 


p. Mferies' (IT.) 


Dr. Egolf 


1959 


p. 'Pink Surprise' (IT.) 


Holden Ar'boretum 


1961 


p. tomentosiun 


Arnold Artoreixun 


1913 


prunifolium 


Spaeth ITur. 


1892 


Eaf inc squian-uai 






rhytidophylloides 


Rochester seedling 




rhytidophyllun 


Veitch ITur. 


1907 


Sargentii 


Arnold Ar'boretum 


1902 


S. 'Flavum' 


Arnold Artorettm 


1907 


schensianiam (N.) 


Dr. Egolf 


1959 


setigerum 


Arnold Ar'boretum 


1915 


Sietoldii 


"Teitch ITux. 


1907 


trilotun 


iillwanger & Barry 


1918 


t. ' Compc.ctum' (IT.) 


Dr. Egolf 


1959 


urceolat-um (IT.) 


Dr. Egolf 


1959 


VJrightii 


Arnold Ar'boretum 


1902 


VITSX - Verbenaceae 






Agnus-castus 


Meehan Ntir. 


1893 


A. 'AlTDa' 






A. latifolia 


Brown Eros. ITur. 


19^3 


Negundo 






IT. heterophi'lla 


Spaeth ITur. 


1892 


N. h. 'I-iiiltifida' 






VITIS - Vitaceae 






acerifolia (IT.) 


Arnold Ar'borct-om 


190i; 


axmrensis (IT. ; 


Arnold Ar'boretum 


1907 


Anderson! i (H.) 


Arnold Ar'boretum 


1920 


argentifolia (K.) 






arizonica (IT.) 


Arnold Ar'boretum 


1906 


Baileyana (IT.) 


Arnold Ar'boretum 


1915 


calif omica (IT.) 


Arnold Ar'boretum 


1914 


cinerea (iT.) 


Arnold Ar'boretum 


1915 



Doaniana 


(IT.) 


La'brusca 


(IT.) 


monticola 


(IT.) 


Romanetii 


(N.) 


Slav»nii 


(IT.) 


VffilGELA - Cap: 


.'ifoliaceae 


'Biformia' 





Arnold Ar'boretum 



Lemoine Nur. 
Rochester native 



Spaeth Uur. 



1907 



1905 
1919 



1892 



270 PHITOLOGIA 


Vol. ID 


, no. 


WEIGEIA, COri, 






coraeensis 'Lavalle' 


LeRoy Hur. 


1892 


c. 'Styriaca' 


Lemoine llur. 


1917 


decora (U.) 


U. S. D. A. 


1961 


•neux de Mai' 


Lemoine Hiir. 


1905 


flori'bimdA 






f. 'Abel Carrlere' 


iill\ffljiger & Barry 


1892 


florida 


Ellwanger & Barry 


1892 


f. 'Stelzner' 


Bllwanger & Barry 


1892 


f. 'Van Houtte' 


Dickson Niir. 


1907 


f . 'Variegata ' 


Bllwanger & Barry 


1892 


f. venusta 


Arnold ArlDoretum 


1915 


hortensis 'Des^bols' 


Bllvranger & Barry 


1892 


japonica 


Arnold ArTDoretum 


1915 


'Messager' 


Lenoine Uur. 


1912 


Middeudorffiana 


Grootendorst KTur. 


1951 


'Mme. Tellier' 


Ellwanger & Barry 


1892 


praecox 'Floreal' 


Lemoine Kur. 


1905 


VnSTERIA - Leguminosae 






flori"bunda 




190I*' 


f. 'AllDa' 


Lemoine Nur. 


1902 


f. 'MacroTDotrys' 


Ellwanger & Barry 


1892 


f . 'ITaga ITocla,' 


Brown Bros. Hur. 


19^3 


f. 'Rosea' 


Brown Bros. Uur. 


19^3 


f. 'Shlro IToda' 


Brown Bros. Nur. 


1943 


f . ' Yi olac ea-pl ena ' 


Bllwpjiger & Barry 


1892 


macrostachys (ll.) 


D. J. O'Connor 


i960 


sinensis 'Al^ba' 


Ellwanger & Barry 


1892 


venusta 


Arnold Artoretum 


1919 


XAIITHOCERAS - SapindAneae 






sortifolia 


Transon Hiir. 


1892 


XANTHOEHIZA - Eammculaceae 






sLuplicissiiaa 


Smith College 


1951 


?UGQA - Liliaceae 






filamentosa 


Transon Uur. 


1892 


flaccida 






glauca 


Arnold Arljoretum 


1922 


ZANTHDXICLUM - Eutaceae 






americamam 


Zblianlsie Nur. 


19^ 


scMnifoliiam 


Arnold Ar"boretum 


1906 


simTolans 


Arnold ArTjoretum 


1906 


ZELKOVIA - Ulmaceae 






carpini folia 


Arnold Ar "bore turn 


1906 


serrata 


Meelian llur. 


1899 


ZEITOBIA - Ericaceae 






pulvejrulenta 


G. Landis Arljoretum 


1962 


ZlZ'iPrlJS - Rhiiinnaceae 






JujuTaa 


Arnold Ar"boret-ujn 


1916 



lUTERIAI^ TOmKD A MONOGRAPH OF THE QZNUS VERBEMA , U. 
Harold N. Moldenke 



VERBHIA OFFICIHALIS L. 

Additional citations : GERMANY : Herb. Martius s.n, (Br) | Herb. 
Monac. a.n. [Mtochen, VII .1901] (Ga— UOU33U) ; Herb. Mua, Bot. 
Stockholji s.n. [29/7A889] (S), s.n. [Berlin] (S), s.n. [Breslau] 
(S), s.n. [Gennania] (S); Herrenkohl s.n. (La); Hisinger s.n. 
[Chariot tenburg] (S); Holzinger s.n. [Kunzelau, 188U] (W— 2U5962), 
s.n. [Wurtemberg] (N)j Q. Jensen s.n, [30/8/1870] (S)j Katisch s. 
n, [Rottstock, 1869] (Lajj Kelleiman s.n« [GOttingen, k Aug. l^T?] 
(Ka); E, H. L. Kraoise 1682L (B)j Kretzer s.n. [Braunschweig, 23/ 
7/83] (lo— 92222), s^n. [23/7/83] (Go, Pl--769U9)j Kretzhmer s.n. 
[7.7111.25] (S); Lademann s.n. [Brandenburg, 25.8.1933] (N); 
LJungstrOn s.n. [21/7/1879] (Go, S)| Lundberg s.n, [8/7/195U] 
(Go) J Martius s.n. (Br)| Meister s.n, [kug, 'O8] (Go, S), s.n, 
[Ponmelsbrunn] (Gg)j Mettzbach s.n. [August I87O] (Gg--31392)j 
Mohlins s.n. [Lindau, 13/7A937] (S); Moll s.n. [Marlof stein, 
1781] (Br) J Ohl s,n, [Kiel, 11,07] (Go)j Olin 3,n. [Juli 1892] 
(Gg, Go, S)j Ploser s,n, [Fa l keriburg, 6,72] (S)j A. Prager s.n, 
(Cm) J Prechtelsbauer s.n. [Aug, I9OI] (S); Pnchtler 12lih (Ca— 
180756, Ca— 988783, Gg, Go)j Relneck s.n, [Amstadt, VII.l91]i] 
(Oa)| Scheaaann 8.n. [22/8/1907] (Go)j Scheppig s.n, [I889] (Go)| 
Schiede s.n. [Mttnchen, Aug. 20] (Br)} R. Schulz s.n, [August 
1913] (B)} Schulz s,n, [Jimiienhausen, 7 August 1907] (B)j Schulz & 
Schulz Bji, [Chorin, 1892] (B), 8.n. [Chorin, 25.7.1896] (B); 
SchwPder s.n, [MShren, Juli 1882] (Io--92221)j Simmons s.n. [29 
Juli 1893] (Go, S)j Stares 7565 (S); Vestergren s.n. [29/7 A907] 
(S); H. Wagner 172 (Ca — 1M775) ; Wahlberg s.n, [Schttoeberg] (S)| 
Wibbe 3,n. [Stromberg, 1862] (I), s.n. [Stromberg, Juli I863] 
(I) J T. Wolf s.n. [28 Aug, 1893] (La)} Zetterstedt s.n. [Hamburg, 
1852] (S) . RdGEN ISLAND: Herb, Mus. Bot. Stockholm s.n. [Insula 
Rugia, 1819] (S). AUSTRIA: Battes s.n. [Tjt^I, 1876] (Mi); Blan- 
hy s.n. [1876] (Ca— 25l8U)i Gander s,n, [25/8/1870] (W— m7579)j 
Gibbe s.n. [Tirol, 26.6.19UO] (B); Hayek 701 (Go), U260 (Go), s_, 
n. [20 Juli 1902] (Go), s.n. [22 Juni I903] (Go), s.n. [17 .VII. 
1921] (S)j Helmerl s.n. [Aug. 21, 1923] (Se— lii62); Herb. Mus. 
Bot. Stockholm s.n. [15 Atig. 1853] (S); M, Johnson s.n. [20/8/ 
1931] (S); K. Keck s.n. (Pa, Po— 7067I); Leffler s.n. [Sept. 1867] 
(Go); D, K. Rechinger s.n. [21 Septemb. 1922] (Go); SOderlund s. 
n^ [l6Tt.l922] (S); M^ F. Spencer s.n. [Aachensee, 9A887] (Ob— 
6729); Steam s.n. [Moldenke & Moldenke 9159] (N) . CZECHOSLOVAKIA: 

271 



272 PHYTOLOGIA Vol. 10, no. k 

Bohemia: Anderberg s.n. [28/6/1936] (S); Herb. Linnaeus 8.n. (S)} 
Lindell 8.n. [Prag, 1906] (E»r)j Sommer s.n. [15.VII.1910] (Vi). 
Moraid-a: Suza 260 (Br, Du— rl8l627, Gg~-267617, Go, I, S, S, Ih— 
1373931), s.n. [VII. 1925] (Ote — 13). State undeterminedi Enander 
s.n. [20/8/1925] (S)j Frisendahl 3.n. [28/6/36] (Go)j H. Lam s.n. 
(N); Petrak 665 (S, S); SchwOder s.n. [Juli 1882] (Go), s.n. 
[Axigust 1887] (S)j Sommer s.n. [22.VII.1913] (S); Sterner s.n. 
[9/7A923] (S); Vestergren s.n. [25.VII.1922] (S). HUHGARI: Hein- 
Itz s.n. (S)j Herb, F. Phllippi s.n. [Hangaria] (Sg--2556U); JAaos 
8.n. [VIII .1906] (Go) J Porutln s.n. [7AV187U] (Pu); Stelnltg 
1278 (Cm) J Sztehlo 16892 (Bz— 23785), s.n. [3/9/187U] (N). STOT- 
ZERLAHDj Ayasse U995 (Du, Uni — 16)} Beger s.n. [Ztirich, 28,6.12] 
(B); Bemet s.n. [Geneve] (N)j R. Campbell s.n. (Ito — 20U80); Dahl- 
sted s.n. [12/8/1893] (S)j Ekman s.n« [26,7.1912] (S)j Frymann s. 
n. [VII .99] (Go, S)j Herb. Mas. Bot. Stockholm s.n. [8/8/72] (S); 
Httlphers s.n. [3.VIII.07] (S); Undmaa s.n. [17/10/188U] (S)j Mac- 
farlane , Bechtel , & Harvey s.n. [Chur, July 23rd 1906] (Hp)} Mels- 
sner 572 (Qo)| Raas s.n. [Tessin, Sept. 1929] (S); Scbneck s.n. 
[July 28, 1903] (Ur); SegerstrOa s.n. [30/10/l92ii] (S)} SOderlund 
8.n. [Juli 1906] (S)} Tiselino 8.n. [U.8.1883] (S), s.n. [7.8. 
1883] (Hi~19U850)j Wijk 8.n. [8/1892] (Go); Wulff 8.n. [Lugano, 
Jtini 1895] (Oo, S). GREECE: Haussknecht S4i« [Kardiza, 1885] (B), 
s.n. [Thessalia superior] (Lu), s.n. (Br, F — photo, M — photo, Si- 
photo, Z— photo), IONIAN ISLANDS: Cephalonia: Bormtaier 1328 
(B). Corfu: D. Hummel 8.n. [22/9/1957] (S). ITALY: Ahlfvengren 
s.n. [26/7 A921] (S); Alstroemer s.n. [Calabria] (S)} J. Ball s.n. 
[Udo, Oct. 18U7] (W~682U56); Cacciato s.n. [10/7/1956] (S); Col- 
lector undesignated 765 (S), 1217 bis (Br)} Commons s.n. (Florence] 
(Cm)} De Toni 8.n. [Modena, Auglio 1903] (N); Engelhard t s.n. [2/ 
6/193U] (B)} Errera & Errera s.n. [12 Sept. 1875] (Br); Gavioli Sj, 
n, [VII.1938] (N)} Gresino s.n. [29.Vn.1938] (N)} Hayek 8,n. [Is- 
turien, 19 Juli 1900] (Go)} Herb. Mas. Bot. Stockholm s.n. [Bagno- 
li] (S), s.n, [Syracuse] (S)} Herb, Itus. Florentini s.n. [Sept, 
1812] (Pa)} Herb. W. H. Harvey s.n. [Bellovaco, Julio 1817] (Du— 
l661Ui9)} Landerer & Sartori s.n. [Nauplia, 183U] (Br)} Lenander a, 
n. [July 21, 1931]~(S)} Savi s.n. [9bre 181;5] (S)} S»derlund s.n. 
(S)} Vignolo-Lutati s.n. [VII.1938] (N, N)} E. Wall 5, in part 
[18/621] (Ew)} L. F. ITard s.n. [Leaning Toirer, Pisa, Aug. I89U] 
(W--229752) } Zoolola s.n. [Ceppo, Augusto 1952] (Hi--1772U2) . SIC- 
ILY: Babington s.n. [Messina, 18U5 — 1851] (C)} Eric Hultfci s.n. [U. 
lt.1960] (S)} Todaro 697 (Du, S), sja. [Palermo, 1850] (S), s.n. 
[1000 m. alt.] (S). CRETE: R. Lauche 2337 (B). JTIGOSUVIA: Bos- 
nia: Lenander s.n. [Juni 30, 1938] (S) . Dalmatia: Lenander s.n. 
[June 19, 1938] (S); Novak s.n, [Lesina] (Lu), Istria: Fruada 8.n. 



196U Moldenke, Monograph of Yertena 273 

[10,11.1913] (S). Macedonia: BornnrtOler I6OI (B)j Seheen a«n. 
[jLlsar, 25.6.I8] (B). Montenegro: Lenainder s,n, [Jxini 10, 1938] 
(S)} Pejovlc 8,n. [11.935] (S). Serbia: Nicic s.n. [27.VII.I896] 
(Qo) . Slovenia: Lenander s.n. [Bled, Juni 3, I93I4] (S). Trieste: 
Collector undesignated s.n. [Sta. Anne] (Lu) j Lenander 8.n« [Maj 
2U, 19 3U] (S); Verouvi s.n. [26,5.1886] (B). Province undetermin- 
ed: E. Wall 5, in part [25/123] (Eir). ALBANIA: Alston St. Sandwith 
2786 (S)i Baldacci 152 (Br). ROMANIA: Lohler s.n. [Transsilv.] 
(S). BULGARU: Bommtfller 3.n. [Vni.l886] (B) . UNION OF SOCIAL- 
IST SOVIET REPUBLICS: Adzharia: Massaloky s.n, [Batum, 3/TL] (Br), 
Azerbaijan: Heideman & Heidetaan 8.n. [13 .VIII. 19 3U] (S). Tuxianan- 
skaya: Sin tenia 932 (B) , Uzbek: Hert?, Hort. Bot. Univ. Asiae Med, 
s.n, [Taschkent] (Se — U;023)j Wedenroy s.n. [Taschkent, 1919] 
(S). Republic undetermined: Kikodse bm, [Kutars, Ozvirgety] (S)> 
Pavlov 517 [Karatau Mts. & Talaa Alatau] (B), 1218 [Karatau Mts. 
& Talas Alatau] (B). MOROCCO: Boitel s.n. [17 juillet I9I8] (Ca— 
882671) J Cos son 8.n. [Mardochee, 1875] (Pa); Favire s.n. [30.5. 
1931] (B)} Quer 5oU ( C a— 370001 ) . ALGERIA: Bov6 s.n. [Aofit 1837] 
(B). TUNISIA: Murbeck s.n. [16/6/1896] (S) . TRIPOLITANIA : Dick- 
son s.n. [circa Tripolium, 1827] (Du — 166UU7) . EGYPT: Hedenborg 
s.n, [Bujukdene] (S, S), s,n. (S, S)} Sabet s.n. [22AA927] (Ka- 
77987); Shabetad U6O.II36 (Ka— 72820); Tackholm s.n. [2UA0A926] 
(S). MAURITANIA: Bov6 s.n. [Aottt 1837] (Br, Br). E31ITREA: Pappi 
U331 (Ca— 902362, N, S), Uljlli (S), ABYSSINIA: CurU I8U (Bn); 
Hildebrandt Uli5 (B); Schiaper 7 (W~9U5770) . BRITISH SOMALIUND: 
Collennatte 1^09 (B) ; Glover & Gilliland 920 (Bm) . CAPE VERDE IS- 
LANDS: Santiago: Brunner s.n. (C). CONGO LEOPOLDVILIE : Bequaert 
5537 (Br), 5976 (Br); De Witte 156 (Br, Br); Hendrickx 201 (Br, 
Br); Humbert 7566 bis (Br)j Quarrg U622 (Br), UGANDA: Edel 2 
(N), 32U (N). TANGANYIKA: J. W. Gregory s.n« [E. of falls, Han- 
nington River, 18 Apl.] (BmY^ s.n. [West of Inkuyuni] (Bm); A. 
Peter U22b [0.1.17] (B), I78O [0. I.U3] (N), 1852 [O.LliUb] (B), 
h2li60 [Y,287] (B), U3726 [V,318] (B). KENYA: P. Chandler 2ZLk 
(B); Meama 82 (N, ¥—630082), 1165 (W— 631208), 197U (W— 
631933), 199|~(W— 631958). SOUTHERN RHODESIA: R. B. Drummond 
U858 (S); C. E. Oodman 221 (Ba)j Rattray 397 (Rh); Wild 82 (N). 
REPUBLIC OF SOUTH AFRICA: Cape Province: Brueckner lt67 (N); H^>- 
trCto 1006 (S, S)t W. H. Harvey Uo5 (Bn); Zeyher 1365 (S) . Natalt 
I. C. Verdoom s.n. [87l2/20] (Eir) . Transvaal: E. E. Oalpin 9061 
(Br) I Obermeyer 30315 (Gg); Rep ton 1298 (Z); E. Wall s^. [2/10/ 
1938] (S); Ri Oi N. Young s.n. [22/5/27] (Hk). Province undeter- 
mined: W. H. Harvey U05 (T) . MASCARENE ISIANDS: Mauritius: F. L. 
Hill M.1 (a»). ARABUt Yemen: Kuntz s.n. [11 Feb. «5l] (W— 
199U907y. TURKEI: BommtQler 812 (B)7 T^6 (B); Fidao s.n. [En- 



27U PHITOLOGIA Vol. 10, no. U 

viron de Snyme, Aottt I90U] (Ca — SkBZOh, N) } Frlvoldaaky s.n. (W- 
26U613)| Kasapllgll s.n. [8-7-19U5] (Ca— 938375); Tengwall 37 (S); 
Wledemajm s.n. [Anatolia] (Ob~m872) . ISRAEL: R. Aar-onaobn 359 
(S); Jouannet-Marle 509 (Du)j Eaeucker U59 (B), 5U6 (B)j Mgyers & 
Dinamore B.)|1)|1 [J.1588] (S), B.8lia [Cat. no. 1588] (S). JOHEU«J 
Bormnttller 3.n. [Gebirge Juda, Jul! 1897] (B)} Kasapllgil 1679 
(Ca--85086). SYRIA J Blanche l51a (B, Br, Da, S, S), srfi. (Du)| 
M. Martens 8.n« (Br)j Post 200 (W--805056)j E. Wall a.n. [2/lU/ 
1932] (S). IRAQ: BomatOler 165U (B), s.n. [Basra, 23.III.1893] 
(B)j Field & Lazar |16 (N), 806 (N), 967 (M)j Lazar U86 (N), 3^1*2 
(S). IRAN: BommtOler 5127 (B); Brans sui. [Teheran, I909] (B)j 
Field & Lazar 1125 (N); Pravitz 77 (S), 100 (S)j K. H. Rechlnger 
1805 (W--206lli37) . PAKISTAN: North»rest Provinces: Dathle 3.n. 
[Thinkiari, 29-8-99] (Qg— 127009), s.n. [Hazara, 1.9.99] (Ca~ 
29U805)} A. H. Kahn s.n. [Hazara district, 8.7.25] (W— 1239955) . 
Swat: Rodin ^2U (Ca— 366OO, W— 22U2319) . NEPAL: WalHeh s.n. 
(Cp). INDIA: Assam; Chand 3259 (Mi); Chatter Jee s.n. [April 
1902] (Br, Po— 63870); T« L, Jenkins 3.n. [Assam] (Bz— 23770); 
Koelz 22817 (Mi) . Chamba: Koelz 8806 (N). East Punjab: J_. R, 
Dnnrunond 26707 (Ca— 2Uli6U5); Koelz 1623 (Mi, N), 30li6 (N), UI9O 
(Mi, W— 1608020), U769 (Mi, N, N, W— I6O8363), jW (Mi, W— 
16097U8); Parmanand 337 (Mi); R^ 17U0 (N); Schlagintrreit 10233 
(S); R. R. Stewart 1937Ua (N); T, Thopson s.n. [Lucknow, April 
1858] (Br), 3.n. [Panjab] (Br, M, sfl Jammr & Kasimir: C. B. 
Clarke 22896 [603a] (W— 802856) ; Collector undesignated 2227 
(Xa)j Ganmie s.n. [Sunagar, U. 7.1891] (Ca — 26979U); Schlagint - 
weit lil;96 (S, W — 80li521). Uttar Pradesh: Blinckworth s.n, [In 
Kamoan] (M); Duthie U282 (W— 80U935); N. Gill 77 (B); U. Singh 
212 (Ca — 361108, La, N); Strachey & Winterbottom s.n, [Kumaon] 
(Br); Vaid s.n. [30,6.U9] (N); Wallich 1825A (L^, S). State 
undeteimined: Duthie s.n. (Ca — 26979U) ; Voigt s.n. (Cp, Cp, Cp) . 
BURMA: Southern Shan States: Malaise 282 (S). Upper Burma: R. 
E. Cooper 6OI9 (Ca— 170238). TIBET: Moribeig s.n. [I908] (S); Sou- 
lig 1075 (Bz— 7281t6), s.n. [Tsekou] (B). CHINA: Anhwei: R. C. 
ChiniTa23 [Herb. Univ. Nanking 85lU] (Ca— 261521, ¥—137011331. 
Chekiang: Barchet 553 (W— 59611U). Fokien: T. C. Chang Ullt7 (La); 
Chang & Metcalf 90 (Du— 250191); £. H. Chen s.n. [Aug. 16, I9U8] 
(N); Cheng 1821 (Ba— 23783); Chung jSEl (Bz— 23782); C, P. ^ 
2022 (Du— 250193), 2689 (Du— 200936); Ging 7229 (Mi); Metcalf 997 
(Urn— 6llli2); Metcalf & Chang 90 (Vi— 128); Pi 6002 (Du— 200935, 
Uto— lli)| Po 12311* (UrT, I2U9I (Ur); L. I. Tai tTIiT"? (Ur) . Honan: 
L. H. Bailey sjn. [Sin-Tien, June 1^1917] (Ba). Hupeh: L. H. 
Bailey s.n. [Hankow, June 10, 1917] (Ba); Gheo 83 [Herb, Univ. 
Nanking I8U16] (Bz— 23780); Chow 598 (N). Kiangsi: S. K. Lau 



196Ii Moldenke, Monograph of Verbena 275 

U280 (S, W--1752972), U758 (S)j W. T. Tslang 9873 (N). Kiangsuj 
H. T_. Chang 369 (Du — 20093U); E, Deschamps sji, [Shanghai, Sept. 
m, 1907] (¥^95698) i Tsu 658 (Ca— 230126, Vi, W--13U6009), 658a 
(Bl). Kirangai: R, C. Chlng 3272 (W— 1508U02), 5272 (Ca— 1+09675, 
N)i Steward & Cheo 513 (N, S), 929 (N); W. T. Tslang 27635 (W— 
1757080) . Kwangtung; N_. J^ Anders son s.n, [Whampoa, Dec, 1852] 
(S); DahlstrOte 25h (S); 0. W^ Groff 163 [Herb. Canton Chr. Coll, 
11001+] (Ca— 2880S9); M^ 112 (Bz— 23751) J P\ A. McClure 2595 
[Herb. Canton Chr. Coll. 9153] (Oa); Pent , Tak , & Kin 68O [Herb. 
Canton Chr. Coll. 12679] (Ca--275057, S, ¥—121+7932), 9U2 [Herb. 
Canton Chr. Coll. 129U1] (Ca— 271+511, ¥•— 121+811+9)} Ping 10622 
(Bz— 23779); 2i Sampson s.n. [Canton, April 1831+] (an); Tsang 
20228 (N); S. W. Wllllaas s.n. [Canton] (N); Wong 21+ (Ba) . Kwei- 
chow: Stewart , Chlao , & Cheo 52 (N); Tslang 5059 (N), 5399 (N, 
S, ¥—1575158). Slkang: Chiao 1126 (S) . Szechuan: C. L. Chorw 
7208 (¥—1990615); Fang 3661 (N, ¥—1525366), 5201 (¥— l6717Ui) , 
5588 (¥—1525367), 10222 (Du— 289053), 12327 (Bm), 121+38 (Bm) . 
Yttanan: Enander s.n. [21/9A926] (S), s.n. [2i+/9/1926] (S); For- 
rest 8036 (S); A. Henry s.n. [Szemao] (N); Malre 1561 (Ca— 
22289!+), 6181 (Ca— 386801, S); Schoch 75 (¥—1235269); H. Smith 
1536 (Go, S); Tsal 53593 (S); Ytt 10518 (Bm) . Province undeter- 
mined; Dunn s.n. (Cp); F. B. Forbes s.n. [lawn, June 1, '79] 
(Bm); Hugh s.n. [Mt. Mias-Wang-san, north central China] (Bm); 
Osbeck 3.n. [ca. 1750] (Lu), s.n. [China] (S) . CHINESE COASTAL 
ISLANDS: Amoy: H. H. Chung 1+70 (Ca— 1+20275) ; A. N. Steward 3059 
(Ca— 1+1+778) . Hainan: Chun & Tso 1+3U71 (N); C. Ford s.n. [31.3. 
93] (¥-1+56231); Lau 1685 (N); C. I_. Lei 231 (B, Ba, Bz— 23777, 
Ca— 61161+8, N), 9112 (B, Ba, Bz— 23778, Ca— 61221+2, N, ¥— 175U*28); 
Liang 61+870 (N); F. A. McClure 7590 [Herb. Canton Chr. Coll. 
9153] (VI), 9153 [Herb. Canton Chr. CoU. 9153] (Ca— 21+8680, Gg— 
1279U5), 3.n. [Herb. Canton Chr. Coll. 9153] (Bi); Tak 13ii [Herb. 
Lingnan Univ. 15633] (Ca— 31573U), 753 [Herb. Canton Chr. Coll. 
17502] (Du— 250192); Tsang I3I+ [Herb. Lingnan Univ. 15633] (N), 
753 [Herb. Lingnan Univ. 17^02] (Ml); Tsang , Tang , i Fung 200 
[Herb. Lingnan Univ. 17731] (Bl), s.n. [Herb. Lingnan Univ. 
17831] (Bi); Ifn 1088 (Du^250190) . Honam: C. 0. Levlne 1+88 (¥— 
778811); E. D. Merrill 9906 (Ca— 29I698, Gg— 311i06) . HOICKONG: 
£. ¥rlght s.n. (¥—71987) . THAILAND: Snitijiand I689 (Ss); Suvar - 
nakoses II8I+ [Royal Forest Dept. 15131] (Sm) . INDOCHINA: Annam: 
Eberhardt l+Ol+O (B) . Tonkin: Clemens & Clenens 3591 (Ca— 339823); 
P^telot I37U (N, W— 1525838). KOREAN COASTAL ISLANDS: Quelpart: 
Kitamura s.n. [16 Jul. 1930] (Mi). ¥ESTHiN PACIFIC ISLANDS: JA- 
PAN: Honshius Baker & Baker s.n. [Kobe, May 18, 1915] (Gg— 31398); 
Collector xindetermlned 36O (¥—71938); Dorsett & Morse 886 (N, ¥- 



276 PHYTOLOGIA Vol. 10, no. h 

I553I469); Herb. Mus. Bot. Stocldiolin s.n. [Tokyo, 30/lO/lh] (S)j 
Ichlkawa 200678 (Ca— 382672, S, W~13li7U59) J £. Matsumura s.n. 
[Tokio, June 11, 1879] (W— 1U75;80) . Ikishina: Hlroe 7336 (Ca— 
96156I). Kiushiu! Ichikaira 200131 [Tanaka U7] (Ca— 2U3938)j Old- 
ham 619 (Br, S, T); Takenouchl 172U (W— 127167U) . ShUcoku: Col- 
lector undesignated s.n. [Shimura, June 19, 1893] (W — 206019), s. 
n. [Mosashi, 2U Juni I910] (W— 117828U) j Herb. Scl. Coll. Imp. 
Univ. s.n. [June, Musashi] (Vt). Island undetermined: Burzes s. 
n« (S)7" ^rb. Ames s.n. [June 19, 1893] (Oa)j Herb. Lugd.-Bat. s_. 
n. [1865] (M), s.n. (T); Noda s.n. (Po— 128708) j J. Petersen s.n. 
[Japan] (S)j Thuiiberg s.n. [1777] (S) . RYUKIU ISLAND ARCHIPELAGO: 
OEDJAWAN ISLft-NDS: Okinawa: Boehmer 117 (N); Field & Loew 21t (Ca- 
7li52li2, Hi), 96e (Bi)i R. V. Moran 50^6 (Bi, Ca— 7^U20)j Walker , 
Taiiada , & Amano 5720 (N) . FORMOSA: Baker & Baker s.n. [Keelung, 
Dec. U, 19lli] (Gg— 31396)} Faurie 372 (V— 8193); Kuntz 33 (W— 
2336836); Sasaki 21397 (Ca— 3Uii389, La); Simada 885 (Ca— 3U5l23)| 
T, Tanaka 11032 (Br, La, S)j Tanaka & Shimada 11032 (B, Go, Mi, 
H, ¥—1577527) . PHILIPPINE ISLANDS: Luzon: J. Clemens 17532 (N); 
M. S. Clemens I9OO3 (Bz— 23767, Ca— 2957U7)j E. D. Merrill 305 
(Bz— 23768); M, Ramos s.n. [Herb. Philip. Bur, Sci. 7789] (N, W— 
629315), 8.n. [Herb. Philip. Forest Bur. 7976] (Bz— 23769); Ramos 
& Edaflo s.n. [Herb. Philip. Bur. Sci. U6562] (Br, Bz— 23766, Ca— 
50897U) ; Ramos & Ramos s.n. [Herb. Philip. Bur. Sci. U6562] (N) . 
INDONESU: GREATER SUNDA ISUNDS: Java: C. A. Baker 1909 (Bz— 
23765) ; Herb. Hort. Bot. Bogor. Ph. 19 (Bz— 26505) . MELANESU: 
HEff GUINEA: Northeastern New Guinea: Keogh s.n. [Mt, Wilhelm] 
(Ng— 6576). AUSTRALIAN RBSION: AUSTRALIA: New South Wales: Boor - 
man s.n. [Cambewarra, 2.1910] (Bi); R. Brown s.n. [Port Jackson] 
(Bm); Collector xmdesignated 62 [Herb. Prager 18628] (Gg— 31393); 
Hagman s»n. [1887 — 89] (Go); A. A^ Hamilton s.n. [Menangle, 8-U- 
1912] (Ew57 S. Helms ^hh (W— 1271320); B. Llnder s.n. [15/10/1936] 
(S); F. Mueller s.n. [Darling River] (Bz— 23775), a.n. [Jvinction 
of Murray & Darling Rivers] (Bz— 23776, W— lli758l) . Queensland: 
M, K. Clemens s.n. [Forest Hill, Jan. I9U4] (Or— 507U5, Or— 507U6), 
8.n. [Oct. -Nov. 19U5] (Or— 53U9li, Or— 53U95, Or— 53U96); Michael 
688 (Bz— 23773); F. Mueller 3.n. [Barcoo] (Bz— 2377U) . South 
Australia: F. Mueller s.n. (Br) , State undetermined: Herb. Mart- 
ius s.n. [Austral, felix] (Br). NEW ZEAIAND: North Island: Wilkes 
3.n, [Bay of Islands] (W— 71985) . POLTNESIA: NIUE: H. F, Moore 
393 (W— 653393); Yuncker 961t5 (Bi, Bi, Ca— 7U*06l, Ca— 918727, Pp- 
290li6, W— 1967969), 1011t5 (Bi, Dp— 290U7, W— 2156U78). CULTIVATED: 
Belgium: M. Martens s.n. (Br). California: Walther s.n. [June 
1925] (Gg— 128725) . Canada: A. Blaln s.n. [L. H. Bailey access. 
617U-39] (Ba); Jard. Bot. Montreal Cult. PI. 617U-39 [seed YO 
10518] (Mv). Chile: Looser 3996 (N). China: Liu L.83I (Ba), 



196U Moldenke, Monograph of Verbena 277 

L«2063 (Ba) , Denmark: Herb. Llebmann 8.n» (Cp); Herb. Mu8» Bot, 
Stockholm s.n. [ex horto Havn.] (S); Herb. Schumacher s.n. (Cp); 
Herb. Vahl s.n. (Cp); Herb. Hort. Hot. Hafn. s.n. [Sept. i860] 
(Cp). England: Herb. Linnaeus Q.3g , S.15 (N — photo of type, Z — 
photo of type), 3.n. [Hort. Cliff. G,83U, S.6; Hort. Bailey neg. 
61|07] (N—photo of type) . France: Herb. W. H. Harvey s.n, [1819] 
(DU--166U3U) . Germany: Herb. Calif. Acad. Scl. 31390 (Gg); Herb. 
Lemmon 8.n. [22.8.l83li] (Ca — 36687U) . Italy: Herb. Harvey s^n. 
[ex semlnib. h. R. P. I81a] (Du— 166U73). Java: Herb. Hort. Bot. 
Bogor. IV.K.A.XLIV,U (Bz— 26U2U), IV.K>A.3iXV.9 (Bz, BZ--26U36) . 
Maryland: W. H. Cowglll 1730 CU. S. Plant Introd. I30I4I6J seed Ytt 
10518] (Oa— 9235); McCann s.n. [9-15-36] (Md, Md, Md) . New loiicj 
L. H. Bailey 8.n« [Sept. 21, 1923] (Ba); G. V. Nash s.n. [17 S. 
1898] (N). Spain: Herb, Hort. Matrlt. 31 (Q), U7 (Q). LXALITI 
OF COLLECTION UNDETERMINED: Bower s.n. [Sept.] (Ms); Capus s.n, 
(Ca— 38875); Collector undesignated 376 (Br), K.ll (W— 36958U), 
s.n. [fin Aoflt l83li] (Pr), s.n. (S); Freed s.n. [N. Am.] (S)j 
Garrigues s.n. (Ml); Herb. J. Angstrttei s.n. [Drageron, P. Sever] 
(S); Herb. H, R. Bassler s.n. [Graigns Point, Aug, 29, '79] (Ka); 
Herb, Bot. Bogor. 23771 (Bz) ; Herb. Coll. Pharmacy s.n. [9-U7] 
(Pa), s.n. (Pa); Herb. Columbia Univ, s.n. [I8UI] (C); Herb. 
Linfaaeus s.n. (S); Herb. Marie-Victorin s.n. [Monde] (Vi); Herb. 
Mu3. Bot. Stockholm 13 (S), s.n. [Majo 18U9] (S), s.n. [Niulam] 
(S); Herb. Rafinesqae s.n. [13.1.1tl] (N); Herb. Saldanha 2766 
[Herb. Mus. Nac. Rio Jan. ref. V] (Ja); Herb. Schleicher s.n. 
(Ca— 73651i); Herb. U. S. Dept. Agr. s.n. (W); Knudsen nv.2 
[Schwirsen] (B); Matveeva 211; [Iter Lencoranlcum] (N); C. T. Mohr 
228 (W— 771861); N^ 80, in part (Q); Poepplg s.n. [N. Am.] (S); 
L. Schaeffer s.n. [Waldhof, 1886] (N); Slmoda s.n. (W); A. Smith 
s.n. [Canavus] (Cp); Stewart s.n. [Campellpore] (Bz — 23772); 
Trolander s.n. [5/7A920] (Ew), s.n. [15 Aug. 1922] (Ew). MOUN- 
TED ILLUSTRATIONS: Anon., fig. 625 (N); F. Bauer, Icon. Nov. 
Holl. 963 (V), 963a (V), s.n. (V); Brut. Mus. (Nat. Hist.) Brit, 
Flow, PI. card F.80 (N); Bulliard, Plant. M6dlc. France pi. 215. 
1780—1793 (N); H. N. & A. L. Moldenke, PI. Ufe 2: pi. $, 19U8 
(Z — negative), pi. 7. 19U8 (Z — negative); Schubert, Naturg. 
Pflanzenreich, ed. 2, fig. 7. 1865 (N); Thornton, Brit. Fl. li 
27. 1812 (N); Zannich., Istor. Piante Venez. pi. 269. 1735 (N) , 

VERBENil OFFICINALIS var, ALBIFLORA Strobl, Flora van Adnont, Pro- 

graamie das Melker Gymnasium. 1882; Strobl, Oesterr. Bot, 

Zeltschr. 33* U06. I883. 

Synonynyj Verbena floribus albidis Vaill, apud Haller, Bmun. 

Meth. Stirp, Helvet. It 661. 17U2. Verbena communis , floribua 

albldla C. Bauh. apud Toum., Inst. Rei Herb. 200. 1700. Verbena 

commmis floribus albidis C. Bauh. apud Sabbat in Martelli, Hort. 



278 PHYTOLOQIA Vol. 10, no, U 

Reman, 3: 11. 1775. Verbena officinalis f . albiflora Strobl ex 
Hegi, Illustr. Fl. Mittel-Eur. 5 (3): 22hl, 1927. Vertena offlcte - 
alis f . albiflora Kraiise ex Moldenke, Rlsum6 371, in syn, 19^9. 

Bibliography: Dorsten, Botanicon 292. l^UO; C. Bauh,, Pinax 
Theatr, Bot., ed. 1, 269 (1623) and ed. 2, 269. I671j Toum,, Inst, 
Rei Herb, 200. 1700; Haller, Emm. Meth. Stirp. Helvet, 1: 661. 
17U2; Dalibard, Fl. Paris. 9* 17U9i Sabbat in Martelli, Hort, Ro- 
man. 3: 11. 1775; Strobl, Flora van Admont, Programme das Melker 
Gymnasium. 1882; Strobl, Oesterr, Bot. Zeitschr, 33: U06, 1883; 
Hegi, Illustr, Fl, Mittel-Bur, 5 (3): 22U.. 1927; Moldenke, Rfisu- 
m6 130, 371, & U73. 1959; Moldenke, R6sum6 Suppl. Ux 15 (1962) 
and 6: 7 & 11. 1963. 

This variety differs from the typical fonn of the species in 
having white corollas. 

Strobl' 8 original publication of this variety has not as yet 
been seen by me, nor do I know whether Krause ever validly pub- 
lished his f , albiflora . The type of Krausefs trinomial, how- 
ever, appears to have been collected by Ez*nst Hans Ludwig Krause 
( no. 28270 ) at Herrenstein, Germany, on August 11, 1907, and is 
deposited in the heirbarium of the Botanisches Museum at Berlin, 
In his 1883 work Strobl says "TM Mascalucia [Etna] (Herb. Gass.). 
Blttht fast das ganze Jahr.^j-," 

This color form is first mentioned by Dorsten (15U0), who 
says **Verbenae duo sunt genera, altera floribus est albls, altera 
vero coeruleis". Later it is referred to again by G, Bauhin 
(1623 & 1671), Toumefort (1700), Haller (17U2), Sabbat (1775), 
and Dalibard (17U9). Bauhin (1623) says "Genera ejus duo sunt 
Plinio 1, c. foliosa, quam foeminam putant: mas rarioribus foliis^ 
&c. folia minora qudm quercus colos glaucus, &c, quidam xaxm om- 
nino genus faciunt, quoniam utraq eosdem effectus habeat, ftc," 
Under his first species, "I, VeAena communis caeiruleo flore", he 
makes the comment "Floribus est caeruleis, qaand6que albidis," 

Only a single specimen has been seen by me. 

Citations: GERMANY: E. H. E, Krause 28270 (B). 

VHIBIKA OFFICINALIS var, ANARRHINOIDES Murr, Allg. Bot, Zeitschr, 
Hit 19. 1908. 

Bibliography: Murr, Allg, Bot. Zeitschr, lUt 19. 1908; Hegi, 
Illustr, Fl. Mittel-Eur, 5 (3)t 22U.. 1927; Moldenke, Known Ge- 
ogr, Distrib, Varbenac, ed. 2, 108 & 199. 19U9; Moldenke, r6su- 
m6 130 & U73. 1959. 

This variety differs from the typical form of the species in 
having 15 — 25 flowers blocnaing simxiltaneously on a single spike, 
the corollas somewhat paler and somewhat larger than in the 
typical fora. 

Murr»s original description of this variety is "Verbena offic- 
inalis L. «nov. var. anarrhinoides Mh. Corollis pallidloribus et 
paulo maioribus, floribus 15 — 25 (in typo U — 5) in una spica si- 
mul evolutis; foma lepida et habitu insignis. In mehreren Exem- 
plaren an Ardetzenberg neben dem Typus." It comprises part of an 
article on the flora of the Tirol, Vorarlberg, and Liechtenstein. 



196U Moldenke, Monograph of Verbena 279 

The type locality, according to Dr, Karl Rechinger, is in Levis, 
Vorarlberg, Aiistria. Hegi describes the plant as "Blttten zu 15 
bis 2$ in eineoi einzigen Bltttenstaiid vereinigt, etwas bleicher 
und etwas grOser als beixi Typus." 

It is knoim to me only from the literature, 

VERBQIA OFFICINALIS var. BRACHYAIITHA Murr, Allg. Bot. Zeitschr. 
16: 187. 1910. 

Bibliography: Murr, Allg. Bot, Zeitschr. 16: 187. 19lOj Hegi, 
Illustr. Fl. Mittel-Eor. $ (3): 22U.. 1927j Moldenke, Known Ge- 
ogr. Distrib. Verbenac, [od. 2], 108 & 199. l9U9j Moldenke, r6- 
sum« 130 & U73. 19^9. 

This variety differs from the typical fonn of the species in 
having all its flowers much smaller and the corolla- lip shorter. 

Murr's original description of the variety is "Floribus omni- 
bus multo minoribus limbo breviore, Ziemlich zahlreich unter dan 
T^us am Tubs des Ardetzenberges in Levis," This locality, ac- 
cording to Dr, Karl Rechinger, is in Vorarlberg, Austria. Hegi 
says "Blttten alle viel kleiner als belm Typus, Lippe ktlrzer," 

It is known to me only from the literature, 

VERBESA OFFICINALIS var. GADDICHAUDII Briq., Ann, Conserv, & 
Jard, Bot, GenSv. 10: 105. 1907. 

Synongrny: Verbena officinalis gaudichaudii Briq, ex Moldenke 
in Chittenden, Roy, Hort. Soc. Diet, Gard. 6: 2209, 1951. 

Bibliography: Briq., Ann, Conserv, & Jard, Bot, GenSv, 10: 
105. 1907; Moldenke, Known Geogr, Distrib, Verbenac,. [ed, 1], 
70 & 101, I9U2; Moldenke, Alph, Ust Cit, 1: 226 (I9U6) and 2: 
li8l, 19 U8; Moldenke, Castanea 13: 121. 19U8j H, N. 5c A. L. Mol- 
denke, PI, Life 2: 60, I9U8: Moldenke, Known Geogr, Distrib, 
Verbenac., [ed, 2], 15U, I6ii, & 199. 19U9} Moldenke, Alph, Ust 
Cit. 3: 750. 19li9i Moldenke in Chittenden, Roy. Hort. Soc. Diet. 
Gard. U: 2209 & 2211. 1951j Moldenke, Rfisumfi 210, 22U, & U73. 
1959; Moldenke, RisumI Suppl. 3: 26 & UO (1962) and 5: 6. 1962. 

This variety differs from the typical form of the species in 
having the boc^ of even the lower leaves linear-oblong and nar- 
row. 

An erect euinaal or "apparently perennial" herb, to 1 m. tallj 
stems several, erect, branched, from a "woolly" [-wDody?] root- 
stockj leaves dull-green, paler beneath, the stem leaves elon- 
gate-lanceolate, narrow, to 7 cm, long and 8 mm. wide, deeply 
incised-dentate, the segments 2—5 mm. long, the upper leaves 
linear-lanceolate, sparsely and superficially dentate, the upper- 
most linear and subentirej corolla deep-blue, bluish, or lilac. 

The type of this variety was collected by Charles Gaudichaud- 
Beaupr6 ( no, lldi ) at Port Jackson, New South Wales, Australia, and 
is deposited in the herbarium of the Conservatoire et Jardin Bot- 
aniques at Geneva. Briquet (1907) says of it "Planta habitu a 
typo valde aliena, tamen vix segreganda." 

The plant has been found on gray-brown clay in cleaired brigalow 
scrub, on riverbanks, on granite formations, and growing as a weed 
of cultivated and fallow land, at 2U0 meters altitude, flowering 



280 PHTTOLOOIA Vol. 10, no. h 

in January, April, Seprtesaber, and October, and fruiting in Octo- 
ber. It has been misiflentified and distributed under the names 
V, caroliniana L. and V. officinalis L. A common name for it in 
New South Wales is said to be "native tobacco" . Everist found it 
to be a "weed around shed in pale-brown loam". 

m all, 9 herbarium specimens and 2 mounted photographs have 
been examined hy me. 

Citations: AUSTRALIA t New South Wales; E. F. Constable 11633 
(W— 199U38U) J Goode 92 (an); A. B. Oldfield s.n. (W— 20686U) . 
Queensland: Everist 6101 (N); Pedlgy 377 (N)j L. S. Smith 30U$ 
(N). Victoria: A. Morrison s.n. [banks of Tarra River, Kew, 1/ 
16/85] (Mi, N — photo, Z — photo) . State undetennined: Herb. Coll. 
Phaiaacy s.n. [Austral, felix] (Pa). CULTIVATED: Germany: Herb. 
Prager 18633 (Gg— 311i60) . 

VERBENA OFFICINALIS var. GRACILIS 0. Cta. ex Moldenke, R6aum6 
Suppl, 3: 15, nom. nud. 1962. 

Bibliography: Moldenke, R6sum6 Suppl. 3: 15. 1962. 

As 7et I do not know where or when — or if ever — Gon^alves 
da Costa described this variety. Possibly the specimen cited be- 
low may represent the type collection, because on its label he 
has written "Sera-t-il une espSce novelle?" The variety is known 
to me only from this single specimen. 

Citations: MADEIRA: Gonial ves da Costa s.n. [Porto Santo, 
Maio 1939] (Go) . 

VERBENA OFFICINALIS var, GRANDIFLORA Hausskn., Mittheil. Thttring. 
Bot. Ver., new ser., 10: 65. 1897. 

Bibliography: Hauaskn., Mittheil. ThtXring. Bot. Ver., new 
aar., 10: 65. I897j Moldenke, R6sumS 131 & U73. 1959. 

This variety differs from the typical foim of the species in 
having its flowers almost twice as large, very remote on greatly 
elongated spikes, the calyx longer, and the corolla deep-blue. 

The type of the variety was collected by Heinrich Carl Hauss- 
kneoht along roadsides near Mount Korona, Greece. The plant is 
known to me only froa the original description. 

VHiBHIA OFFICINALIS var. MACRDSTACHTA (F. Muell.) Benth. in 
Benth. & Muell., Fl, Austr. 5: 36. 1870. 
Synonymy: Verbena macrostachya F. Muell., Fragn. 1: 60. 1858. 
Verbena officinalis var. macrostachya Benth. ex F. M. Bailey, 
Canpreh. Cat. Queensl. Pi. 382. 1913. Verbena officinalis macro- 
staotya Benth. ex Moldenke in Chittenden, Roy. Hort. Soc. Diet. 

Gard. U: 2209. 1951. 

Bibliography? F, Muell,, Fragm. 1: 60. 1858 j Benth. & Muell., 
Fl. Austr. 5: 36. 1870; F. Muell., Syst. Cens. Austr. Pi. 102. 
1882; F. M. Bailey, Cat. Indig. Nat. Pi. Queensl. 35. 1890; F. M. 
Bailey, Compreh. Cat, Queensl, Pi. 382. 1913; H. J. Lam, Verbena 
ao. Malay, Arch. 10, 1919; Moldenke, Known Geogr. Distrib. Ver- 
benac., [ed. 1], 7U & 101. 19U2} Moldenke, Phytologia 2: 339. 



196U Uoldenke, Uonograph of Verbena 281 

l9U7j Moldenke, Kncmn Geogr, Diatrib, Verbenac, [ed, 2], 109, 
16U. & 199. 19U9| Moldenke, Alph. Ust Git. 3x 667 (19U9) and U> 
12Uo, I9U9; Moldenke in Chittenden, Roy. Hort. Soc, Diet, Ga:d. 
Ut 2209 & 2211. 1951 J Moldenke, RfieumS 132, 22U, 369, & U73. 
19^9; Moldenke, Rfisum^ SMppl, 3 J 1*0 (1962) and 5« 6 & 7. 1962. 

This variety differs from the typical form of the species in 
having its spikes unlfonnly elongated to h$»S cm, and very 
coarsely glandular-puberulent or hirsute and the flowers rather 
larger. It is said to be a spreading herb, to 66 cm. tall, with 
pink corollas. 

The type of the variety was collected by Ferdinand Jacob 
Heinrich von Mueller at Peak Downs, Queensland, Australia, in or 
before 1858, Bentham (1870) cites also a Bownan s.n. from Rock- 
hampton, Queensland. 

The Liu collection cited below was cultivated in Chihli, Chi- 
na, from seeds secured from Copenhagen, Denmark, where one may 
assume that the plant was also cviltivated — the original source 
of the seeds is not known. It has been collected in flower and 
fruit in October. Herbarium material has been misidentified and 
distributed under the name V. officinalis L. 

In all, 2 herbarium specimens and 2 movmted photographs have 
been examined by me. 

Citations: ALGERIA: Dukerly s,n, [Setif] (Br). CULTIVATED: 
China: Liu L.2063 (Ba, N — photo, Z — photo), 

VERBENA OFFICINALIS f , MONTANA Goiran ex Moldenke, Known Geogr, 
Distrib. Verbenac, [ed. 1], U5 & 101, nca«, nud, 19U2. 

Bibliography: Moldenke, Known Geogr, Distrib, Verbenac, [ed, 
1], U5 & 101, 19U2j Moldenke, Alph. Ust Cit. 1: 229. l9U6j Mol- 
denke, Known Geogr. Distrib, Verbenac, [ed, 2], 108 & 199, 
19U9J Moldenke, R6sum6 131 & U73. 19^9. 

Where or irtien — if ever — Goiran described this form is not 
as yet known to me, nor do I know where, when, and by whom the 
type collection was made. It is known to me only from the singls 
specimen cited below. 

Citations: ITALY: Gavioli s,n. [19,VI,1927] (N) , 

VERBENA OFFICINALIS var, PROSTRATA Gren, & Godr., Fl. Fran?. 2: 
718. 1852. 

Bibliographer: Gren, & Godr., Fl, Fran?. 2: 718. I852j Hegi, 
niustr. Fl, Mittel-Bur, $ (3): 22U1. 1927; Moldenke, Known Ge- 
ogr. Distrib, Verbenac, [ed. 1], lili & 101. 19li2; Moldenke, 
Phiytologia 2: 339. 19^7} Moldenke. Known Geogr. Distrib. Verben- 
ac, [ed. 2], 107, 108, & 199. 19U9; Moldenke Alph. Ust Cit. 3: 
832 (I9U9) and U: 1210, l9U9i Moldenke, R^suml 129, 130, & U73. 
1959 I Moldenke, R6sum5 Suppl, 6: 7. 19o3. 

This variety differs from the typical form of the species In 
having prostrate stems. 

The original description of the variety is " Verbena officina- 
lis var. pro strata . Tige 6tal6e-couch6e sables des environs 

de Bayonne [France] ." It is not known to me when or by whom the 
type was collected. It is possible that the Meissner 572 cited 



282 PHTTOLOGIA Vol. 10, no. U 

by me hereinbefore tinder V. officinalis may actually represent 
this variety. Hegl (1927T'de8cribe8 it as "Stengel nlederlieg- 
end." A common name is said to be "seruftit*. Herbaritm mater- 
ial has been misidentified and distributed under the names V. of- 
ficinalis L. and V. supina L. The variety is known to me only 
from the folloiiing 7 specimens. 

Citations: FRANCE t Collector undesignated s.n. (Du, Du). 
SfflTZERUND: Probst s.n. [13.9.36] (Pb). ERITREA: Pappi 2981 
(W— 1969120). ABTSSmiA: Schimper JhS (S, S, W— 9U5092). 

VERBaiA OFFICINALIS var. RESEDIFOLIA Uurr, Deutsch, Bot« Monat- 
schr. 20: $2, 1902. 

Bibliography: llurr, Deutsch. Bot. Monatschr. 20: 52. 1902; 
Hegi, Illustr. Fl. Mlttel-Eur. $ (3): 22U1— 22U2. 1927| Moldenke, 
Known Qeogr. Distrib. Verbenac,, [ed. 2], IO8 & 199. 19^9 J Mol- 
denke, Alph. List Git. U: 1099. 19U9} Moldenke, R6snm6 I30, 132, 
& U73. 1959 J Moldenke, Rfisumfi Suppl. 6: 7. 1963. 

This variety differs from the typical fom of the species In 
having its stem-leaves twice pinnately parted, with blunt round- 
ed tips. 

Murr's original descripticm of the variety is "StengelbllCtter 
doppelt flederspaltig mit stumpfen gerundeten Zipfeln. Am Damme 
der Valsuganabahn bel S. Christoforo. Die Form, welche mlr 
einen fremdartigen Eindruk machte, dtlrfte aus den vielbesproch- 
enen griechischen sabiereln stammen." Dr. Rechlnger states that 
this locality is now in the province of Alto Adlge, in northern 
Italy (before World War I it was in Bozen, Bezirkshauptmann- 
schaft Trient, StCdtirol, Austria) . 

It is known to me only from the description and the single 
following specimen. 

Citations: MOROCCO: E. Wall 7$38 (Go). 

xVERBENA OKLAHOMENSIS Moldenke, Known Geogr, Distrib. Verbenac, 
[ed. 1], 78—79. 19U2. 

Synonymy: Verbena bipinnatiflda Nutt. x V. canadensis (L.) 
Brit ton ex Moldenke, Alph, List Invalid Names Suppl. 1: 22, in 
syn. I9U7. Verbena canadensis (L.) Britton x V. bipinnatiflda 
Nutt. ex Moldenke, Alph. List Invalid Names Suppl. 1: 22, in 
syn. 19lt7. 

Bibliography: Moldenke, Known Geogr. Distrib. Verbenac., [ed. 
1], 11, 76, 78—79, & 101. 19U2| Moldenke, Known Geogr. Distrib. 
Verbenac. Suppl. 1: 2. I9l*3i Moldenke, Alph. List Clt. 1: 1$7. 
19U6) Moldenke, Alph. List Invalid Nanes Suppl. 1: 22. 19U7} Mol- 
denke, Alph. List Clt. 2: $1$ & $17 (19U8), 3* 822 (19U9), and kt 
1085. 19U9} Moldenke, Known Geogr. Distrib. Verbenac., [ed. 2], 
21 & 199. 19U9J E. J. Salisb., Ind. Kew. Suppl. 11: 263. 1953; 
Moldenke, Am. Midi. Nat, 59: 357. 1958; Moldenke, RSsumfi 22, 26, 
358, 360, & U73. 1959; Moldenke, Plytologia 8t 121 (I96I) and 8: 
210, 213, 21U, U37, & UUO. 1962; Moldenke, Rfisumfi Suppl. 3j 6 4 
7. 1962. 

This hybrid has characters more or less Intermediate between 



196U Uoldenke, Monograph of Verbena 283 

those of 7. canadwisls (L.) Britton and those of V. bipinnatlfida 
Nutt.y with those of the latter predominating. The leaves are 
deeply blplnnatlfld and the corollas are small and densely pubes- 
cent outside as in V. bipinnatlfida , but the calyx and bractlets 
are finely appressed-pubescent or puberulent as in V* canadensis . 
The loirer bractlets have their lower margins more or less strong- 
ly ciliate, but these marginal cilia are only dense and conspic- 
uous on the lower portions of the lower bractlets and are very 
sparse or absent from the upp>er ones* 

The type of the hybrid was collected by Dona Pitts sixteen 
miles east of Norman, Cleveland County, Oklahona, on April 19, 
1915, and is deposited in the herbariim of the University of Ok- 
Isthana. It was distributed as V* bipinnatlfida Nutt. by the col- 
lector. Duplicate specimens of the type collection and other 
collections of the same taxon are mounted on the same sheets with 
V» canadensis and V. bipinnatlfida under one label, indicating 
that the plant apparently grows in close association with the two 
parental species. This fact leads me to believe that it is an 
interspecific h7l>rid, rather than a mere form or variety of V, 
bipinnatlfida . The two parent species grow together in at least 9 
counties of Kansas, 2 of Arkansas, I9 of Oklahoma, 19 of Texas, 
one county each of Missouri, Indiana, and Nebraska, and 3 parishes 
of Louisiana, so it is very possible that this hybrid will be 
found more often. It may even account for same of the tremendous 
"variability" noted in herbarium material now usuadly regarded as 
one or the other of its parental species. The Pitts collection 
is a mixture of the hybrid with specimens of V. blpinnablfjda , 
while the Van Vleet collection consists of a mixture with stems 
of V. canadensis , indicating, apparently, the very close proxim- 
ity of at least one of the parental species in each case. The 
Demaree collection was originally identified as V. ciliata var, 
longldentata Perry, and was collected on sandy hills. 

The hybrid has also been found in open or scrubby prairies and 
glades, on outcroppings of Cretaceous Annona Chalk, along road- 
sides and gravelly roadsides, in rocky railroad fill, on the top 
of limestone hills, in gravelly limestone, on rolling limestone 
prairies, along streams, in rich cork elm-hickory-oak woods on 
the Woodford Chert foimatlon, in rocky xeric limestone gullies, 
and on outcrops of Arbuckle Lime formation, flowering and fruit- 
ing from April to June and in September. 

The Stratton U302, cited by me under V. bipinnatlfida , looks a 
bit like xV. oklahomensls . Verberta damareel Moldenke, regarded 
by me hereinbefore as a synonym of V» blpl nnat if Ida , seems to 
combine the ordinary leaf -characters of V. bipinnatlfida with the 
large flowers of V» canadensis , and may possibly also prove to be 
a natural hybrid between these two species, perhaps in reverse 
form. Its flower-characters, however, are not the same as those 
seen in ±V. oklahomensls . Even if it should prove to be the same 
l^rbrid, perhaps in reverse form, its name would not, in my opinion, 



28U F H Y T L G I A Vol. 10, no. k 

replace xV. oklahonensls , because it was originally proposed by 
me as a true species — not as a hybrid. My understanding of the 
International Rules of Botanic Nomenclature is that the oldest 
valid epithet must be used only if proposed in the same category 
of classification. My good friend, Conrad V. Morton, however, in 
a letter to me dated March 5» 1962, disagrees. He maintains that 
"There is nothing in the Code that would make this true. On the 
contrary, if V. demareei is prior then it would displace V. x 
oklahomensis , if the two are considered the same; whether one, or 
both, or neither was fii^t proposed as a hybrid or a species is 
irrelevant.'* The plant should have considerable horticultTiral 
merit. 

In all, lU herbarium specimens and 2 mounted photographs have 
been examined by me. 

Citations: ARKANSAS: Little River Co.: Moore & litis s.n. [A- 
pril 5, 1953] (Ok). OKLAHOMA: Carter Co.: M. Hopkins 609U (Ca— 
882575). Cleveland Co.: Pitts s.n. [UA9A5] (N— photo of type. 
Ok — 20U77 — isotype. Ok — 20U79 — type, Z — photo of type). Coman- 
che Co. J Van Vleet s.n. [Mt. Sheridan, 7/h/03] (Ok— 10269). Mur- 
ray Co.: Hopkins , Nelson , & Nelson 667 (St). Payne Co.: E. W. 
Michael Th (St). Pontotoc~Co . : Duffer 28 (St); D. McCoy 58U 
(St), 2U79 (St), 2522 (St), 2539 (St). Tillman Co.: Demaree 
12188 (Ok— 20U70). TEXAS: Lampasas Co.: Mahler 12la (St), 

VERBENA ORCUTTIANA Perry, Ann, Mo, Bot. Card. 20: 28U— 285, 1933. 

Bibliograpiy: Periy, Ann, Mo, Bot. Card, 20: 2^7, 2li9, 250, 
260. 28U— 285, & 355. 1933; A, W, Hill, Ind. Kew. Suppl. 9s 295. 
19 38 J Moldenke, Known Geogr. Distrib. Verbenac, [ed. 1], 19 & 
101, l9U2j H. N, & A, L. Moldenke, Pi. Ufe 2: 7U. 19U8j Molden- 
ke, Alph. Ust Cit, 2: 519 (19l;8), 3: 779 (19U9), and U: 1126, 
1127, 12U3, 12hh, & 1295. 19U9; Moldenke, Known Geogr. Distrib, 
Verbenac, [ed, 2], 33 & 199. 19li9; H. N. & A, L. Moldenke, Anal. 
Inst. Biol, Mex. 20: lU, l9U9j Moldenke, R6sum6 39 & U73. 19 59 J 
Moldenke, Phytologia 10: lUO. I96U. 

Stems several from a common base, tetragonal in cross-section, 
branching, glabrous or veiy sparsely hirtellous, the hairs short; 
leaves decussate-opposite; petioles 1—2 cm. long, margined; 
leaf -blades lanceolate-elliptic to spatulate, U — 6 cm. long, ta- 
pering at the base into the petiole, coarsely serrate, rugose and 
appressed-pubescent above, more densely spreading-pubescent or 
hirtellous beneath and prominently veined; spikes pedvinoulate, 
solitary or somewhat panicled, strict, elongate, mostly dense- 
flowered, finely glandular, closely appressed-pubescent, the ra- 
chls more or less angulate; bractlets lanceolate-acuminate, 
shorter than the calyx, sparsely ciliate, the midrib and margins 
more or less decurrent along the rachis; calyx U mm. long, ap- 
pressed-pubescent and finely glandular, the teeth short-subulate 
or acuminate, more or less connivent above the schizocarp after 
anthesis; corolla iypocrateriform, blue, its tube about as long 
as the calyx, the limb 3 — k mm, wide; cocci trigonous, 2 mm. long, 



196U Moldenke, Monograph of Verbena 285 

raised-reticulate at the apex, striate toward the base, the com- 
missural faces extending to the tip of the nutlet, miiricxilate- 
scabrous . 

The typ>e of this endemic species was collected by Charles Rus- 
sell Orcutt ( no. 909 ) — in whose honor it is named — on table- 
lands at Hanson's Ranch, Baja California, Ifexico, on Jiily 30, 
1883, and is deposited in the Gray Herbarium of Harvard Univer- 
sity at Cambridge, The species has been collected on dry flats, 
tablelands, and mountains, at the margins of dry pools, and in 
mountain meadows, at altitudes of U650 to 6000 feet, in flower 
and fruit in April, Jtily, Septanber, and October. The common 
name "verbena azul" is recorded for it. It has been misidenti- 
fied and distributed in herbaria under the names V, littoralis 
H.B.K,, V, officinalis L., and V. scabra Vahl, On the other hand, 
the Wiggins U360, distributed as V. orcuttiana , is actually V. 
neocexicana var. hirtella Perry (or V. plicata Greene), and Carter, 
Alexander , & Kellogg 2135 is V, menthaefolia Benth. 

The Gallegos 23U2 specimens cited below have a photograph 
moiinted on the same sheet with the specimen. A note on the United 
States National Herbarium specimen of Wiggins 5506 states that the 
inflorescence is not glandular. 

In Plant Life (19U8) it was stated that this species was named 
in honor of Heman Chandler Orcutt (1825 — 1892), but apparently 
this is not the case,' There is no evidence that Perry did not in- 
tend the name to honor the collector of the type specimen, Charles 
Russell Orcutt (186U— 1929) . 

Perry (1933) cites the following 3 additional specimens not as 
yet seen by me: MEXICO: Baja California: C. R. Orcutt 909 (G — 
type), s.n. [Hanson's Ranch, 29 July 1883T"(G» ^T* She notes that 
•This species, which has been passing aa V. lit oralis , is much 
like 7. neomexicana var. xylopoda in the finely glandular spike 
and the angle of insertion of the flowers. It differs, however, 
in the shorter nutlets, the smaller corollas, and the type of pub- 
escence. In V. orcuttiana , the trichomes are shoirt and somewhat 
hirtellous, and the pubescence of the inflorescence is closely 
appressed. In gross habit, it is scarcely to be distinguished 
from V. simplex , but the latter has somewhat harsher pubescence 
and larger non-glandular flowers." The C. R. Orcutt 118 cited 
hereinafter is what Perry cites as " Orcutt s.n. [Hanson's Ranch, 
29 July 1883]" and his 5 _ 21 is her s.n. from "fountains, northern 
Lower California, 8 July 1885". 

In all, 20 herbartan specimens, including material of the type 
collection, and one mounted photograph have been examined by me. 

Citations: MEXICO: Baja California: Gallegos 23U2 (Me, ¥— 
1209871) J C. R. Orcutt 118 (W--132310U) , 521 (W— 1323105), 909 
(Cm— isotype, Pa~isotypey, sji. [Pinery, 7-27-1883] (Mi, W— 
56176), s.n. [Santa Catalina Mts,, JuJy 27, 1883] (Ca— 10li81i0), s^ 
n. [7-31-1883] (Vt), s.n, [July 1883] (C), s^n. [7-8-1885] (I, Up- 



286 PHTTOLOQIA Vol. 10, no. k 

17110, Vt); Wiggins 5^08 (Du— 265866, ¥—182101^3), 9157 (Du— 
25851i5, W— 17U7513), 11258 (Du--321768, W--1976620):; 

VESBEM ORIGENES R. A. Phil., Linnaea 29t 20. 1857. 

SynonynQr: Verbena deserticola R. A. Phil,, Fl. Atac. I4.O. i860. 
Verbena palmata Relche, Fl. Chile 5> 285 tt 287* 1910. Glandularla 
origenea (Phil.) Schnack & Covas, Darwiniana 6t U75» l9Ui* 

Bibliography: R. A. Phil., Linnaea 29: 20, l857i R. A, Phil,, 
Fl, Atac. UO. i860; F. Phil., Cat. PI. Vase. Chil. 220 & 221. 
1881; Hook, f . & Jacks., Ind. Kew. 1: II78 & 1179. 1895; Reiche, 
Fl. Chile 5: 285, 287, & 289—291, 1910; Prain, Ind. Kew, Suppl, 
U: 2U5. 1913; Sanzin, Anal, Soc, Cient. Argent, 88: 98, 127—129, 
& I3U. 1919; Baeza, Nomb, VUlg. Pi. Silv. Chile, ed. 2, 100, 206, 

6 269. l930;Stapf, Ind. Lond, 6: U30. 1931; R. Espinosa, Okolog, 
Stud. Kordillerenpfl, 36 & 38. 1932; M. R. Espinosa Bustos, Ri- 
vadavla 327. 1938; Moldenke, Suppl. List Common Names 9 & 18, 
I9UO; Uoldenke, Enoim Qeogr, Distrib, Verbenac,, [ed. 1], k2 !c 
102. I9lt2; Moldenke, Alph. List Invalid Names U6 & U9. I9l*2; 
Schnack & Covas, Darwiniana 6: li75 {19hh) and 7: 72. 19U5; Mol- 
denke, Phgrtologia 2: 116. 19U5; Cabrera, Bol. Arg. Soc. Bot, 1: 
67. 19U5; Moldenke, Alph, List Cit. 1: 8U. 19U6; Moldenke, Alph, 
List Invalid Names Suppl, 1: 10, 19^7; Moldenke, Alph. List Cit, 
3: 688 & 813 (19li9) and U: 1115 & 1116, 191*9; Moldenke, Known 
Geogr, Distrib. Verbenac, [ed, 2], 102 St 199. 19U9; Acevedo de 
Vargas, Bol, Mus, Nac, Hist. Nat, Chile 25: 58—59. 1951; E, J, 
Salisb., Ind, Kew, Suppl, 11: 101, 1953; Moldenke, Rfisum^ 122, 
296, 363, 371, & U73. 1959; Moldenke, Phytologia 8: 20U (1962) 
and 10: U3. 196ii. 

Illustrations: Sanzin, Anal, Soc, Cient, Argent. 88: 128, 
1919. 

Perennial herb, to UO cm, tall, robust, glandular-hispid or 
densely short-haizy, sometimes almost tomentose, yellowish- 
green; stems several, loosely tufted, from a matted caudex, up- 
right, 15— 60 cm, long, stout, trichotOTiously branched, leafless 
at the apex; leaves sessile, the uppermost smaller and forming a 
many-parted involucre beneath the inflorescence; leaf -blades 
trapezoid in outline, 3 — h cm. long and wide at the middle of the 
stems, usually broader than long, cuneate and somewhat clasping 
at the base, 3-parted to the middle or trifid, the segments o- 
vate'-triangular, acute at the apex, entire or 1 — 3-dentate, the 
margins sinuate; venation impressed above, prcminent beneath; in- 
florescence terminal, consisting of a cluster of visually 3 many- 
flowered heads; flowers numerous; bract lets linear to awl- 
shaped, almost as long as the calyx, ciliate; calyx prismatic, a- 
bout 8 mm. long, hispid, membranous between the 5 green ribs 
which terminate in short triangular teeth; corolla white or 
white-lilac to pink, lavender, or lilac, its tube about 10 mm. 
long, hairy on both surfaces, the limb often rose or blue, about 

7 mm. wide, the 5 lobes obovate-cuneate, enarginate; cocci iabout 
3.5 mm. long, red-bz'own, smooth on the back, the lateral angles 
narz*owly alate. 



1961i Moldenke, Monograph of Verbena 28? 

The type of this distinctive species was collected bj Claude 
Gay ( no. 1906) in Coqulmbo, Chile, a clastotype being preserved 
in the herbarium of the Uuseo Nacional de Historia Natural at 
Santiago, Chile, The type of V. deserticola was collected by Ru- 
dolph Amandus Philippi ( no. " 1293 ") at Pogonal, Atacama, Chile, 
in February, 185U, and is no, ShiSh in the herbarium of the same 
institution — Macbride's type photogr^h 17li3U being of an iso- 
type in the herbarium of the Botanisches Museum at Berlin, now 
destroyed, Vgrbena palmata is based, ^parently, on Peralta s,n, 
from Dofla Ana, Reiche s.n, from Baflos del Toro, collected in Jan- 
uaiy, I90U, and on Volckmann s.n, frcm Rio Turbio, collected in 
the summer of i860 — 1861, all these localities being in Coquimbo, 
Chile, and all the specimens deposited at Santiago. 

Morrison reports the species as "very common on rocky screes 
above the baflos, Bafios del Toro", Coquimbo. Johnston found is on 
dry benches at the foot of talus slopes in quebradas and "on ta- 
lus slopes in gorge above bafios" in Atacama. He distributed his 
no. 6102 as "Y. deserticola var," Common names reported for the 
plant are "hierba del incordio", "rica-rica", "ricanrica", and 
"yerba del incordio", the firfet of which is also applied to V» 
laciniata (L.) Briq, The species has been found at altitudes of 
3200 to 3800 meters, flowering from Decanber to February, fruit- 
ing in Febnjary, 

Reiche says of his V. palmata ; "Difiere de V, cuneifolia R, & 
P. por el tallo mas corto, las hojas mas anchas que largas, las 
espigas mas costas e involucradas , La V* cimeifolia es del Perfi, 
pero segun Qay V p&J* 23 se observ6 tambien en las Cordilleras 
entre Santiago i Mendoza," For the type of his species he seems 
to cite only one collection: "Cordilleras de Coquimbo (Dona Ana, 
valle superior del Rio Turbio)", but Acevedo de Vargas regards 
the three collections cited above as cotypes. She cites (1951) 
the following 12 specimens not as yet seen by me: CHILE: Atacama: 
Bore hers 3,n. [Baflos de Inca, 1. 1886] (Sg— 51i762); F. Philippi 
8,n, [Quebr. de Paipote, U-1-1885] (Sg— U2U65, Sg~5li763, 3g— 
68387); R. A. Philippi s.n. [Pajonal, II.l85Uj Hacbride photos 
I7U3U] (Sg--5U761i, Sg — photo) . Coquimbo: Alamos s.n, [Cordille- 
ra de Los Patos, Aestate l881t] (Sg— U2iA6l)j C, Gay I9O6 (Sg— 
5U769 — clastotype); Peralta s.n. [Dofla Ana] (Sg — 5U768); Reiche 
s.n. [Baflos del Toro, I.I90U] (Sg — 5U765, Sg — 5U767)} Volckmann 
s.n. [Rio Turbio, Aestate i860— 61] (Sg— 5U766) . 

In all, 22 herbarium sp>eclmens, including photographs of the 
type collection, and U mounted photographs have been examined by 
me. 

Citations: CHILE: Atacama: I. M. Johnston hSUh (W— lli97721) , 
$95Q (W— Ui96088), 61^ (¥—11*9^935) j R. A. Philippi 1293 (W— 
1323079), s.n. [Pogonal, Feb. 185U| Herb. Mas. Nac. Hist. Nat. 
Chile 5U76UJ Macbride photos 17U3U] (Kr— photo, N— photo, N— 
photo, N— photo) J ¥erdenaann 959 (Ca— 311i836, Qg— 114737U, N, S). 



288 PHYTOLOQIA Vol, 10, no. U 

Coqulnbo: Cabrera 3^28 (N, N, S)} M. R. Esplnosa 2U (N); £• L. 
Morrison 17271 (Ca — 63020^, S) j Wagenknecht s,n. [Bafloa del Toro, 
IX.19U7] (Ew); Werdermann 225 (Ca~2383lU, Qg~3U5lli, S, S, W— 
1233138), Ovalle: Tribarren 8,n« [T, Meyer UOOl] (N) . Province 
undetermined: Herb, Mus, Nac, Hist. Nat. Santiago 6 (N) . 

VERBENA ORIGENES var. SMPERI Moldenke, Phytologia 3i hh — h$» 
I9U8. 

Bibliography: Moldenke, Pt^rtologia 3: hh—h$ (19U8) and 3: 76. 
I9U9; Moldenke, Known Geogr, Distrib. Verbenac., [ed. 2], IO6 & 
199. I9U9J Moldenke, Inform. Mold. Set $1 Spec. U. 1956; Moldenke, 
R6sum6 127 & U73. 1959. 

This variety differs from the typical foim of the species in 
having broadly ovate 3-parted leaves about 3 cm. long and 2.5 om. 
wide, each division deeply lobed with rounded incised lobes, 
densely spz^ading-hirsutulous or scabrid-hirsute on both surfaces, 
the margins conspicuously revolute. The leaves are very similar 
to those of V» crithmi folia Gill. & Hook., but the inflorescence 
is that of V. origenes R. A. Phil. 

The type of the variety -was collected by Juan Semper — in 
whose honor it is named — at Quebrada de la Vacas, at an alti- 
tude of 2UOO meters, in the department of Las Her as, Mendoza, Ar- 
gentina, between March 2 and 20, 1938, and is deposited in the 
Brit ton Heitarium at the New York Botanical Garden. It was dis- 
tributed "by Ruiz Leal, iriio describes the plant as "ccmmon", as 
his no. U937. The plant has been collected in flower and fruit 
in February and March. The Miers UiS, cited below, bears a no- 
tation "G* macrocephala ", but to what genus the initial refers ia 
not clear. 

in all, 5 herbarium specimens, including the type, have been 
examined by me. 

CiUtions: ARGENTINA: Mendoza: Miers IM (Bm); Ruiz Leal 11^629 
(Z); Semper s.n. [Ruiz Leal U937] (N — type). San Juan: Castella- 
nos 15207 (W— 21982U5) J F. A. Roig s.n. [Ruiz Leal I3OIO] (Ss) . 

xVERBENA OSTENI Moldenke, Phytologia 2: 323— 32U. 19ii7. 

Synonymy: Veii?ena peruviana (L.) Britton x V^ plat ens is Spreng, 
ex Moldenke, Alph. List Invalid Names Suppl. 1: 26, in syn. 19U7. 
Verbena platens is Spreng. x V, peruviana (L.) Britton ex Moldenke, 
Alph. List Invalid Names Suppl. 1: 26, in syn. 19U7. Verbena 
teucrioides x charaaedryfolia Osten ex Moldenke, Alph. List Invalid 
Names Suppl. 1: 27, in syn, 19U7. Verbena melindres x teucrioides 
Osten ex Moldenke, R^sum^ 369, in syn. 1959. 

Bibliography: Moldenke, Phytologia 2: 32>-32U & 337. 19U7j 
Moldenke, Alph. List Invalid Names Suppl. 1: 26 & 27. I9li7; Molden- 
ke, Known Geogr, Distrib. Verbenac, [ed. 2], 100 &. 199. I9h9i 
Moldenke, Alph. List Cit. 3: 780. 19U9} E. J. Salisb,, Ind. Kew. 
Suppl. H: 263. 1953; Moldenke, Am. Midi. Nat. $9: 357. 1958; Mol- 
denke, R5sum6 120, 369, 372, 376, & U73. 1959; Moldenke, Phytolo- 



196U Moldenke, Monograph of Verbena 289 

gia 8x 121 (1961) and 10: 128. I96U. 

A natiiral hytrid between V. peruviana (L.) Britton and V, 
platensis Spreng,, with intermediate characters j stems slender, 
more or less densely short-pubescent, the younger parts spreading- 
pubescent or hiirtellousj petioles about 1 mm, long, hirtellous; 
leaf -blades small, ovate, 1 — 1.5 cm, long, h — 9 mm. wide, coarse- 
ly dentate, pustulate-scabrous and very sparsely or more densely 
strigose-hirsut ulcus above, scattered-pubescent or hirsutuloxis 
beneath, especially on the larger venation; peduncles about 1.5 
cm, long or almost obsolete, densely spreading-hirtellous with 
hair of several lengths j heads densely rather few-flowered; 
bractlets lanceolate, S^—6 mm. long, long-attenuate, densely 
short-pubescent, long-ciliate on the margins; calyx about 1 cm, 
long, densely hirsutulous, irregtilarly apiculate; corolla light- 
red, orange, or violet, its tube glabrous, about 15 mm, long, its 
limb about 15 mm. wide. 

This natural hybrid apparently occurs sporadically idiere the 
ranges of the two parents overlap in Uruguay, The type was col- 
lected by Cornelius Osten ( no, 3177 , in part) — in whose honor 
it is named — at Coquimbo, growing vrith the two parental species, 
in the department of Soriano, Uiniguay, on November 16, I89U, and 
is deposited in the herbaiT.um of the Museo de Historia Natural at 
Montevideo, Osten, in a note written at Montevideo in January, 
1931, says that "Der letztere [xV. uruguayensis Moldenke] ist 
tfbrigens, wie der spontane Bastard chamaedryf . ( Melindres ) x teu^ 
crioides Gill, sehr selten, was eigentlich Wander nimmt bei der 
Haufigkeit der Eltem, die Leichtigkeit mit der sich die Verbenen 
ktlnstlich kreuzen lassen." Osten describes its corollsis as or- 
ange, but Herter cauLls them violet. 

The plant inhabits dry svinny sandy fields, dry sandy soil, 
carapos, seiTanias, and arroyos, at altitudes of 100 to 200 metejrs, 
blosscHaing in September, November, and January. Herbarium mater- 
ial has been misidentified and distributed under the names V. 
humifusa Cham., V. incisa Hook., and V. marrubioides Chan, 

In all, II4 herbarium sp)ecimens, including the type, have been 
examined by me 

Citations: URUGUAY: Castellanos s.n. [Herb. Inst. Miguel Lillo 
I50U8I (N); Herter 1000 [Herb. Herter 82763; Herb. Osten 22625] 
(B, Ca— 3U8972, N, S, Ug, W— 11^22050), 1000a [Herb. Herter 83913] 
(Ca— 360220, N), s.n. [Valle Eden, IX.1928; Herb. Osten 20Ulli] 
(Ug); Legrand 3U91 (Ug); Osten 2977 (Ug), 3177, in part (N— iso- 
type, Ug— type). 

VERBENA OVATA Cham., Linnaea 7: 263~26U. 1832. 

Bibliography: Chan.. Linnaea 7t 263 — 26U. 1832; Steud., Nom, 
Bot., ed. 2, 2: 750. l8la; D. Dietr., Syn. PI. 3: 602. 18U3; 
Walp., Repert. h: 19. 18U5; Schau. in A. DC., Prodr. 11: 51il. 
18U7; Schau. in Mart., Fl. Bras. 9: 187. 1851; Jacks, in Hook, f, 
& Jacks., Ind. Kew. 2: 1179. 1895; Briq. in Chod. & Hassler, Bull. 
Herb. Boiss., s6r. 2, Us 1058. I90U} Briq. in Chod. & Hassler, 



290 PHYTOLOGIA Vol. 10, no. U 

Plant. Hassler. 10: U79. l90Uj Molfino, Physis 7: 103. 1923; Her- 
ter, Florula 105. 1930} Moldenke, KnoTin Geogr. Distrib. Verbenac., 
[ed. 1], 39, khf & 102. 19U2; Schnack, Anal. Inst. Fitotfic. Sta, 
Catalina U: 19. I9li2| Schnack & Covas, Darwiniana 6; U70. 19Uaj 
Darlington & Janaki Ammal, Chromoa. Atl. 270. 19U5} Augusto, Fl. 
Rio Grande do Sul 232. 19U6; Moldenke, Alph. List Cit. 2: 375 & 
IM (I9I18), 3: 688, 863, 865, 921, & 922 (19U9), and U: 1257. 
19U9j Moldenke, ISncnm. Geogr. Distrib, Verbenac., [ed. 2], 9U, I06, 
& 199. 19U9i Moldenke, Phcrtologia 3: 377. 1950; Moldenke, Am. 
Midi. Nat. 59: 353 & 35U. 1958; Moldenke, R^sumS 110. 118, 127, k 
li73. 1959; Lewis & Oliv., Am. Journ, Bot. U8; 61a— 6l2, 1961; 
Moldenke, Phytologia 8: 121, 126, & 127 (1961) and 9: 37U & 375. 
1963. 

Perennial herb, 1 — 1.6 m. tall; stems strict, mostly simple, 1 
m. or more tall, tetragonal, scabrous on the angles, densely 
leaiy to about the middle, subaphyllous above, issuing £rom per- 
ennial nodose roots, deeply canaliculate-stilcate, especially on 
the side alternate Tri-th the plane of the leaves, the margins ob- 
tuse or roTinded; nodes annulate; lower intemodes 3.5—8.5 cm. 
long, upper ones greatly elongate, to 28 cm. long; leaves decus- 
sate-opposite or approximate, sessile, coriaceous, very stiff, 
reticulate- veined and very riigose, shiny above and much roughen- 
ed by bulbous-based hairs, subcanescent-hirtellous beneath and 
hispidulous on the venation, the lower ones broad for the genus, 
often 1/3 or 1/2 longer than the succeeding intemode, broadly 
ovate or subrotund-ovate, 5«5 — 9 cm. long, 3 — 5*5 cm. wide, cor- 
date at the base, semiamplexicaul and adnate, acute at the apex, 
unequally and coarsely dentate-serrate except at the very base, 
the teeth irregular, large and small ones interspersed, the lar- 
ger ones often incised, acuminate; upper leaves (between the 
greatly elongate intemodes) much reduced or even scale-like, 
triangulai>-acuminate or elliptic-lanceolate, 1— It cm, long, 2— 
12 mm. wide, similar to the lower ones in texture and pubescence; 
venation deeply impressed above and very shaiply prominent be- 
neath, even to the teirtiaiy veinlet reticulation; panicle ter- 
minal, cymose, compact, spreading-hirtous on its branches and 
peduncles, the spikes cylindric, 2 — 2.5 cm. long, usually ter- 
nate at the apex of the short branches of the panicle, the cen- 
tral one sessile, the lateral ones short-peduncvQate; bractlets 
membranous, lanceolate, about 6 mm. long, 1-nerved, concave, 
slender, attenuate-acute, glabrous except for the ciliate margins^ 
imbricate, about twice as long as the calyx, shiny, tinted ■with 
lilac when fresh, hiding the calyx; calyx during anthesis mem- 
branous, tubTolar, very slightly incxirved, lightly 5-nQrved, pu- 
berulent, the teeth very short, rounded, apiculate, ciliate, in 
fruit oblong, split, 3 times as long as the fruit, contracted a- 
bove; corolla clear-blue or violet, twice as long as the calyx, 
its tube slightly surpassing the bractlets, subvilloua-lanuginoua 
outside, its limb exiguous; style slightly longer than the calyx, 
with a little terminal horn adjacent to the stigma; fruit regu- 
larly 6-seeded; cocci 2 mm, long, bright-fascous on the back, 
striate-ribbed, rather shloy, the commissural surface obtusely 



196U Moldenke, Monograph of Verbena 291 

angled, white-leathery j chromosome number: 2n • 72, 

The type of this very distinct and unmistakable species was 
collected by Friedrich Sellow ( no. 3671 ) somewhere in "Brasiliae 
meridional!" and was deposited in the herbarium of the Botanisch- 
es Uusexm in Berlin, where it was photographed by Macbride as his 
type photograph no,17U35, but is now destroj'-ed. August© (19U6) 
states that Herter collected the species somewhere in southern 
Uruguay, but I have seen no Uruguayan material of it thus far. 
Noack (1937) reports that the chranosooe number is 2n - 72, the 
highest nomber known for the genus. Walpers (18U5) places it in 
his Section Verbenaca , Subsection Inermes , Group Foliosae , Sub- 
group HolophyHae , along with 22 other species. A natural hybrid 
of 7. ovata with V. bonarienais L. is knowa as xV. intercedens 
Brlq. 

Verbena ovata has been found in bogs, swamps, and shrubby 
marshes, at altitudes of 850 meters, blooming in September and 
from November to February, fruiting in February. 

Lewis Sc Oliver (I96I), in dlsciissing the probable genetic his- 
tory of the genus, say that '•It seems logical that the 2 sections 
are monophyletic and consequently that their basic chromosone 
numbers of 5 and 7 are derived from a common number of x • 6. 
Such a base number is known in the modem Verbenaceae , e.g., 
Priva . Not yet considered because of its debatable origin is V. 
ovata Cham, with its 2n - 72 chromosomes (Noack, 1937)* This 
species, found in east-central South Amez*ica, might have arisen 
as an amphidiploid in a cross between an n « 5 &nd n > 7 species 
followed by genomic doubling to reach this polyploid level. Der- 
men's (1936) inability to produce an intersectional hybrid, the 
rare occurrence of 1 parent in South America, and the unquestion- 
ed classification of V. ovata in the section Verbenaca fthe taxon 
is not morphologically intermediate) are all evidences against 
such an origin. Alternately, V. ovata may be a terminal dodeca- 
ploid of an extant x - 6 series which was morphologically more 
closely related to Verbenaca than to Glandularis. Additional 
cytological studies might reveal other 'relic' taxa in Soirth A- 
merica, but the very existence of V. ovata supports the hypothe- 
sis of an ancestral x ■ 6 stock," 

Herbarium material of V. ovata has been mis identified and 
distributed as V. litoralis H.B.K. 

In all, 17 herbarium specimens, including the type collection, 
and 7 mounted photographs have been examined by me. 

Citations! BRAZIL: Rio Grande do Sul: Jttrgens UU2 (B) ; Raabo 
967U (Rb). Santa Catarina: Smith & Klein 11115 (N, W— 2251752, 
Z). State undetermined: Sellow 3^71 "[Macbride photos 17U35] (Br — 
isotype, F — photo of isotype, Er — photo of type, N — photo of 
type, N — photo of type, N — photo of isotype. Si — photo of iso- 
type, Z— photo of isotype). PARAQUAT: Fiebrig 56U5 (¥—1159391) J 
Haasler U695 (N) . ARGENTINA: Misiones: Ekaan 2032 (N, S)> D. 
RodriguerT96 [Herb. Inst. Miguel Lillo 32315] (Ca— 3I188, nTT s. 



292 PHTTOLOQIA Vol. 10, no. U 

n, [Herb, Mus. Argent. Cienc. Nat. 23779] (N)| A. G, Schalz 6986 
(Sz); Q. J. Schwarz lgl3 (N, S), 3673 (N). 

VERBEHA PARAGUARimsIS Moldenke, Phytologia 1: li83--ii8U. 19liO. 

Bibliography: Moldenke, Phytologla 1: li83 — 1;8U (19Uo) and 1: 
511. I9UIS Moldenke, Knoim Geogr, Distrib, Verbenac, [ed. 1], I4I 
&. 102. 19U2; Moldenke, Alph. List Cit, 1: 26U (I9I46) and 3: 869. 
19U9} Moldenke, Knonn Geogr. Distrib. Verbenac., [ed. 2], 99 & 199* 
19i;9j E. J. Salisb., Ind. Kew. Suppl. 11: 263. 1953; Moldenke, R6- 
sun^ 118 & U73. 1959. 

Herb, to 60 cm. tall; steas usually simple and erect, strigose 
with closely appressed iihitish antrorse hairs; leaves decussate^ 
opposite, numerous, sessile, appressed to the stem; leaf-blades 
lanceolate, 6—23 om» long, 1—3 inm. wide, attenuate above the 
rather sharply acute apex, rounded or obtuse at the base, entire 
and usually subrevolute along the margins, strigose on both sur- 
faces like the stems; inflorescence terminal, spicate, terminating 
the stcBis, solitary, 10—20 cm, long, about 1 cm, wide thtoughout, 
many-flowered, the flowers alternate, imbricate, somewhat spread- 
ing; rachis slender, more or less flexuous, strigose like the 
stems; propl^lla 2—3 mm. long, closely appressed to the calyx, 
strigose . 

The type of this most distinctive species was collected hy 
Teodoro Rojas ( Hassler 9701) on a high plateau and in declivities 
in the Sierra de Amambay, Paraguay, in December, I907, and is de- 
posited in the Delessert Herbarium at the Conservatolire et Jardin 
Botaniques at Geneva. It is known only from the type collection, 
of which 7 herbarium specimens and ^ mounted photographs have been 
examined by me. 

Citations: PAR/LGUAI: T. Rojas s.n, [Hassler 97$1] (B — isotype, 
Bm — isotype, Cb— type, N — isotype, N — isotype, N — photo of type, 
N — photo of isotype, S — isotype, S — photo of type, V — isotype, Z— 
photo of type, Z — photo of isotype). 

VERBENA PARANEMSIS Moldenke, Phytologia 6: 331. 1958. 

Bibliography: Moldenke, Phytologia 6: 331. 1958; Moldenke, r6- 
sum6 U73 & U9U. 1959; Moldenke, R6sum6 Suppl. 1: 7. 1959; Angely, 
Fl, Paran, 16: 79 (I960) and 17: I46. I96I, 

Herb; stem apparently procumbent or dectmibent and rooting, tet- 
ragonal, densely hirsute with fulvous-brown hairs; branches tet- 
ragonal, densely hirsute with fulvous-brown hairs; principal in- 
temodes 1—7 cm, long, elongated on the side branches; leaves de- 
cussate-opposite; petioles 1—3 mm. long, obscure, very densely 
hirsute like the branches; leaf -blades chairtaceous, rounded-ovate, 
scanewhat lighter beneath, 1 — 2.3 cm. long, 8 — 25 mm. wide, rounded 
in outline at the apex, more or less subcimeate or acute at the 
base, coarsely but regularly dentate along the margins with acute 
or subacute teeth, densely villous on both surfaces with long, 
brownish, more or less appressed hairs, the pubescence scanewhat 
more grayish beneath; midrib very slender, impressed above, pro- 
minulous beneath; secondaries slender, 3 — 5 per side, ascending, 



196U Moldenke, Monograph of Verbena 293 

hardly arcuate, impressed above, promlnulous beneath; veinlet re- 
ticulation sparse, obscure on both surfaces) inflorescence ter- 
minal, subcapitate, densely villosulous throughout, many-flowered, 
the heads about 1.5 cm, long and widej peduncles very slender, 
1,5—3 cm. long, densely hirsutvilous with fulvous-brown hair; 
calyx tubular, straight, about 3 nun« long, densely villosulous or 
hirsutuloxis on the outside, its ria 5-toothed, the teeth about 
0,7 am. long; corolla hypocrateriform, violet, its tube about 5 
mm, long, densely barbate-tomentose in the throat, the limb 5- 
parted, about 2.5 — 3 am. wide, the lobes obovate; stamens U, in- 
cluded, didynamous, 2 inserted at about the middle and the other 
2 above the middle of the corolla- tube; style glabrous, about U 
mm. long, included; stigma 2-lobed, only one lobe papillose. 

The type of this remarkable species was collected by my good 
friend, Gert Hatsohbach ( no. U211i ) in the campo along the road to 
Palmeirinho, in the municipality of CJuarapuava, ParanA, Brazil, 
on November 15, 1957, and is deposited in the H, N, Moldenke her- 
barium at Yonkers, New York. The species is very distinct be- 
cause of its capitate inflorescences and very small flowers. 

Vy good friend, Dr. Angely, is of the opinion that the specif- 
ic name of this plant should be written ' ♦paranaensis ". However, 
I see no valid reason for adding another syllable to names like 
this or " canadensis or " virginiensis " simply because the state 
name from which they are derived terminates in an "-a" (Parani, 
Canada, Virginia) . 

The species is known to me only from the type specimen. 

Citations: BRAZIL: Parand: Hatschbach li211; (Z — type), 

VHIBENA PARODII (Covas & Schnack) Moldenke, Phytologia 2: lh9 * 
19U6. 

Synonyny: Glandularia p>arodii Covas & Schnack, Revist, Argent, 
Agron, 11 J 9U— 97, fig. 3. 19hh. 

Bibliography: Covas & Schnack, Revist, Argent. Agron, 11: 9U — 
97, fig, 3. 19Ui; Covas & Schnack, Darwiniana 7: 86. 19li5j Schnack 
& Covas, Darwiniana 7: 71, 72, 7U, & 75, pi. 2 A & D, 19U5j Mol- 
denke, Phytologia 2: m9. l9Uo; Moldenke, Alph, List Invalid Names 
Suppl, 1: 10, 19U7; H. N. & A. L, Moldenke, PI, Ufe 2: 75. 19U8; 
Moldenke, Phytologia 2: U82. 19U8: Moldenke, Known Geogr. Distrib. 
Verbenac., [ed. 2], 106 & 199. I9ii9; E. J. Salisb,, Ind. Kew, 
Suppl. 11: 101 & 263. 1953; Moldenke, Inform, Mold. Set 51 Spec. 
U. 1956; Moldenke, Rfisum^ 128, 296, & ii73. 1959; Moldenke, Phorto- 
logia 8: 123 (1961) and 8: 396. 1962; Moldenke, R6sum6 Suppl. 3: 
15 (1962), 5: 7 & 8 (1962), and 6: 7. 1963; Moldenke, Phytologia 
9: UOO (1963) and 10: 136. 196U. 

Illustrations: Covas & Schnack, Revist. Argent. Agron. 11: 95, 
fig. 3. 19U;; Schnack & Covas, Darwiniana 7: pi. 2 A & D. 19li5. 

Annual or perennial prostrate herb, 5 — 15 ca. tall, forming 
small mats; stems creeping, rooting below, ascending at the apex, 
cylindric-quadrangular, densely pubescent, the hairs simple, 
rather rigid, erect or oblique, not appressed; leaves decussate- 
opposite; petioles 3 — 8 mm. long; leaf -blades triangular in out- 



29U PHITOLOGIA Vol. 10, no, k 

line, 1,5 — 2,8 cm, long, 1.2 — 2,5 cm, iiide, laciniate, attenuate 
into the petiole at the base, the segiaents linear^lanceolate, 
0,5 — 1.5 Jnm. wide, obtuse or subacute at the apex, not revolute 
along the margins, pubescent above and on the margins and on the 
venation beneath, the hairs simple, rigid, to 0,6 om, long, ob- 
lique to subappressedj inflorescence terminal, spicate, often 
solitary, abbreviated during anthesis but elongating after an- 
thesis; bractlets linear^ lanceolate, more than half as long as 
the calyx, ciliate along the margins, subappressed-pilose on 
both surfaces; flowers very fragrant; calyx tubular, 7—8 lan, 
long, densely pubescent, the hairs simple, erect or oblique, not 
appressed; corolla hypocrateriform, varying frcan lilac, pale- 
lilac, rose, violet, or blue-violet to bluish, cream, or white, 
sometimes described as •'yellow and violet", "rose and white", 
"white and bluish", "white and pink", or "white with patches of 
lilac", completely pubescent on the outer surface except for the 
portion covered by the calyx, pubescent also at the base of the 
lobes within; stamens typical for Gland\ilaria , the two upper 
ones with a glandular appendage about 1 mm. long and 0,5 mm. 
wide, flattened, rounded at the apex, longer than the theca, ex- 
serted; gynoecium typical; ovary about 1 mm. long; style about 9 
mm. long; cocci subcylindric, about 3.5 nmi. long, obtuse at the 
apex, truncate at the base, the upper third reticulate on the 
dorsal svirface; chromosome number: n ■> 5* 

The type of this species was collected by Guillerme Covas and 
Benno Julian Christian Schnack ( no. 2112 ) between Anchoris and 
Zapata, in the department of Tupungato, Mendoza, Argentina, on 
March 5, 19Ui. It is named in honor of Lorenzo Raimimdo Parodi, 
distinguished Argentine educator and botanist. 

The species has been collected along roadsides, on plains, in 
arroyos, at the edges of ditches, and on "paramillas", at alti- 
tudes of 850 to 3500 meters, flowering from October to May and in 
August, in fruit in December, Eyerdam, Beetle, & Grondona report 
it as "not common, in sandy loam, full sun, in a small garden, 
associated with Chenopodium ," It has been misidentified and 
distributed in herbaria under the name V, erinoides Lam, The 
length of pistil related to size of pollen-grains is discussed by 
Covas & Schnack (19l45) , These authors state that the species is 
related to V, laciniata (L,) Briq, They give the following inter- 
esting notes: "Hemos hallado en la localidad del tlpo una pobla- 
ci6n fomada, nuy probablemente, por hfbridos (y formas derivadas 
de ^stos) entre esta especie y Glandularia mendocina. . . ,La pobla- 
cl6n hlbzdda presenta una amplia gama de variaci6n que comprende 
formas inteirmedlas y foraas vecinas a ambos padres; en algunas de 
estas formas hemos podldo obsei*var f lores con pequeflos l6bulos 
petaloldes en la base del limbo de la corola, car^cter que ntinca 
hemos observado anteriormente en el gSnero Glandularia . . .EL polen 
de esta especie (observado en el ejemplar tipo) presenta cierto 
porcentaje de granos estSrlles. Ademds hemos observado, en indi- 
viduos de la poblaci6n hibrlda mencionada, Irregularidades en la 
melosls (mieBibi>os de un par de cromos<»iia8 separados en diacinesj.s. 



196U Moldenke, Monograph of Verbena 295 

lo cual Indlca falta de homologla en parte del material cromos&n- 
ico) Da el misma poblaci6n h£brida hemos encontrado una for- 
ma con flores rosadas, color aparentemente debido a un derivado 
de cianidina," This hybrid is discussed b7 me hereinafter under 
xV. perturbata Moldenke. 

^Schnack & Coras (I9lili) give the following key for the differ- 
entiation of V. parodii from its immediate allies: 

1, Stems erect or suberect, not creeping, 

2. Entire plant covered with a dense pubescence of mixed simple 

and glandulose haii^; leaves tripartite-pinnatilobed 

V. perakii . 
2a. Entire plant covered with a sparse pubescence of only- 
simple hairs; leaves pinnatisect V. mendocina . 

la. Stems procumbent, rooting at the base, ascending at the tips. 

3. Cocci 2 mm, long. 

U, Bractlets one-third as long as the calyx; spikes not elon- 
gating after anthesis V. dissecta . 

Ua, Bractlets more than half the length of the calyx; spikes 

elOTigating after anthesis .V. santiaguensis . 

3a. Cocci more than 3 mm. long. 
5. Corolla externally glabrous; glandular appendages of the 
anther connective subcylindric , scarcely visible froo 
outside or included; pubescence appressed,.V« laciniata . 
$a. Corolla extemaljy pubescent; glandular appendages of the 
anther connective much compressed, clearly exserted; 

pubescence composed or erect or oblique hairs.. .,, 

V, parodii . 
Herbarium material of V, parodii has also been misidentified 
and distributed as _V. laciniata (L.) Briq. In all, U5 herbariim 
specimens have been examined by me. 

Citations: ARQENTHIA: Cataoarcaj Peirano s.n. [Cerillos; Heit>. 
Inst. Miguel Ullo 328Ii9] (N, Ug— U9U7) . Chubutt Eyerdam , Beetle, 
i Orondona 2h^0 (Ca— 6^120) , Mendoza: Araqae Molina & Barkley 
l9Ar762 (N); Araque Molina & Paci 261 (N, S); CAceres h (N); Car - 
et te 8,n. [Ruiz Leal 2^6$] (N); Caret te & Ruiz Leal s.n, [Ruiz 
Leal 7838] (N); Covas , Schnack, & Ruiz Leal s,n. [Ruiz Leal 9U21] 
(N); Lourteig 772 [Herb, Inat, Miguel Ullo 113935] (Ca— I66OOO, 
N)j Melis & Barkley 2(3Mz012 (N); G'Donell 11^2 (N); Ruiz Leal 
1167 (N), 1208 (N), 1507 (N), 2235 (N), 1^373 (N), Ii390 (Rl), 1^825 
W; 6165 (n5T 6650 W^ 6888 Wi 5108 (N), 8U65 (N), 8556 (N), 
8700 (N), 9102 (N), IOU88 (N), llU^f lz), 13U05T sa); Sanzin 632 
[Herb. Oaten 1281U] (N, Ug), 800 [Herb. Osten 12809] (Ug), 3099 
[HeA. Osten II4638] (Ug); Semper s.n. [Ruiz Leal U3U5] (N), s.n, 
[Ruiz Leal 10212] (N), B.n. [Ruiz Leal 10302] (N, Ss) , Salt*: 
Yenturi 6937 (»— 1591503). Tucxaulnj Wall 4 Sparre s.n. [29/11/ 
U6] (Ew), B.n. [10/l2/li6] (Ew, Ew, N) . 



296 PHTTOLOGIA Vol. 10, no. 1* 

VERBENA PARVUIA Hayek in Engl., Bot. Jahrb. U2: 162--163. 1908. 

Synonymy: Verbena hirsuta Ruiz & Pav. ex Moldenke, Rfisume' 
Suppl. 6: 10, in syn, 1963 [not V. hirsuta Mart. & Gal., l8Ui]. 

Bibliographer: Hayek in Engl., Bot, Jahrb. U2: 162—163. 1908 1 
Praln, Ind. Kew, Suppl. Uj 2U5. 1913; Moldenke, Known Geogr. Dis- 
trib. Verbenac., [ed. 1], UO & 102. 19li2: Moldenke, Alph. List 
Cit. 1: 201 (19U6), 2: 602 (19U8), 3: 688 & 968 (19U9), and U: 
1079. 19U9j Moldenke, Known Geogr. Distrib, Verbenac, [ed. 2], 
73, 98, & 199. 19U9; Moldenke, Phytologia 3s 7U & 75 (19U9) and 
5: 96. 195Ui Moldenke, RfisumS 81, 85, & U73. 1959 j Moldenke, r6- 
sumS Suppl. 3: 13. 1962; Moldenke, Kjytologia 8: 317, 321, & U12 
(1962) and 9: 51, l5l, & 296. 1963; Moldenke, RSsumS Suppl. 6: 
10. 1963. 

Perennial herb to 38 cm. tall or decumbent, with straggling 
spreading habit to U5 cm* wide; roots thick, man^-branched; stems 
woody, wiry, erect or arcuate-ascending, decumbent at the base, 
6 — ^10 cm. tall, sulcate-tetragonal, subsimple or branched, scab- 
rous with appressed hairs; branches prostrate or appressed to the 
ground} leaves decussate-opposite, short-petiolate; leaf-blades 
oval or obovate, 1 — l.li cm. long, rather obtuse at the apex, con- 
tracted into the petiole at the base, irregularly incised-crenate 
or -serrate to almost pinnately lobed, appressed-setulose on both 
surfaces, linear-nervose on the lower surfaces; spikes teminal, 
simple or few-branched, at first narrowly conic, pointed, and 
head-like, later elongating as the seed forms ; flowers small, 
sessile; bractlets ovate, subequaling the calyx, acute at the a- 
pex, setose-ciliate on the margins; calyx about 1,5 mm. long, 
very minutely setulosej corolla hypocratertfom, varying from 
blue, pale-blue, light-blue, or deep-blue to lilac, pale-purple, 
or pTirple, or even "pinkish-blue, white at base", 3 nun, long, 
subpuberulent on the outer surface. 

The type of this species was collected by Karl Fiebrig-Gertz 
( no. 3Ul5 ) in Tarija, Bolivia, in 1903 or 190U, and is probably 
deposited in the heitarium of the Naturhistorisches Museum in 
Vienna, Hayek says of it "Ein zierliches kleines PflShchen vom 
Habitus einer zwerghaften V. officinalis , aber schon durch die 
ganz anders gestalteten Blatter verschieden." 

The species has been found on open or open rocky hillsides, 
in usually very dry waste or cultivated ground, and creeping in 
grass steppes. West found it near shrubs on the floor of rocky 
canyons, Asplund encountered it among weedy vegetation and in 
hard gravelly ground, and Ellenberg collected it in moist meadows 
with Festuca dolichophylla , The only common name recorded for it 
is "verbena". It has been found at altitudes of 600 to UOOO me- 
ters, flowering from February to May and in September & November, 
fruiting in February and March, It has been misidentified in 
herbaria and distidbuted under the names V. brasiliensis Veil., 
V. braziliensis Veil., V. cuneifolia Ruiz & Pav,, V. hispida Riilz 
& Pav,, and V. lit oralis H.B.K. West 8281 is a mixture with V. 
graeilescens (Cham.) Herter, A. S. Kalenbom 160 is erroneously 



196U Moldenke, Monograph of Verbena 297 

cited by me in my Alph. List Cit. 2: 602 (19U8) as V. brasilien- 
8 is, as was Kalenbom & Kalenbom 160 in Phytologia~5: 321 (19^2). 

In all, 39 herbarixmi specimens, as well as 2 mounted photo- 
graphs of the type, have been examined by me. 

Citations: ECUADOR: Az\iay: Asplund 17801; (S) . Chimborazo: As- 
plund 20l4l;3 (S). PERU: Cuzco: Balls 6781; (Ca— 683072, W— 17778U6); 
Cook & Qilbert 373 (W— 603603)j F, L. Herrera s.n, [Cuzco, July 
1923 ]""(¥— 1190015); F. W. Pennell llqSl; (N) . Junfn: A. S^ Kalen- 
bom 160 (W— 10l4ii398)j Kalenbom & Kalenbom 160 (N)} Killip & 
Smith 21855 (N, W— 1356976), 2211|2 (N, W— 1357187) J Kunkel 38| 
(Z), 386 (Mi), 387 (Mi), 390 (Mi); Ledlg 33 (W— 11M172) . Lima: 
Diera 982 (Ko); Killip & Smith 215U1 (N, W— 1356719) . Puno: El- 
lenberg 250 (Ut— U539Ub) . Tacna: H. H. Rusby 912 (C, Pa) . De- 
partment undetermined: Hrdlicka s.n« [vicinity of Huarochlri, 
February 1913] (W— 602735) . BOLIVIA: Cochabamba: Steinbach 8729 
(N, S), 8729a (W— l857l;la) . El Beni: Buchtien 5887 (W— 113U883); 
H_. H, Rusby 908 (C, Pa) . La Paz: Asplund 2303 (S, Us) j Buchtien 
135 (W~117798l)j Haamarlund 167 (N), 196 (N). Tarija: J. West 
8281 , in part (Ca— 565121;); Fiebrig 3la5 [Macbride photos 17l;36] 
(Kr — photo of type, N — ^photo of type). State undetermined: 
Kuntze s.n. [Bolivien, 600 m., l/U April 1892] (N, W— 702210) . 

VERBENA PARVULA var. GIQAS Moldenke, Phytologia 7: 85. 1959. 

Bibliography: Moldenke, Phytologia 7: 85. 1959; Moldenke, Rl- 
sum6 1;25 & 1;73. 1959; Moldenke, R^siim^ Suppl, 1: 6. 1959; Soukup, 
Biota 3: 30, I960; Moldenke, R^sum^ Suppl. 3: 13, 1962; Moldenke, 
Phytologia 8: 321 & la2. 1962. 

This variety differs frcxa the typical form of the species in 
having its stems erect, to 90 cm. tall, the intemodes greatly 
elongated, and the leaves elliptic-obianceolate, to 7.5 cm. long 
and 2.5 cm. wide. The corolla is described as pinkish-white. 

The type of the variety was collected by Albert Charles Smith 
and Ellsworth Paine Killip ( no. 21925) on an open hillside at an 
altitude of 3000 to 3200 meters, at Tarma, Junln, Peru, between 
April 20 and 22, 1929, and is deposited in the Britton Herbarium 
at the New York Botanical Garden, The plsint has much of the as- 
pect of V. litoralis H.B.K., but the inflorescence characters are 
those of V. parvula Hayek. It has been found in anthesis in A- 
pril and Jvine, and his been misidentified in herbaria as V. lit - 
oralis , Hammarlund 631 was incorrectly cited in Phytologia 8: 
321 (1962) as V. brasiliensis Veil, 

In all, 3 herbarium specimens, including the type, have been 
examined by me. 

Citations: PERU: Cuzco: Hammarlund 631 (S) . Junln: Killip & 
Smith 21925 (N— type); Kunkel 389 (Z) , 

VERBENA PAirLENSIS Moldenke, Phytologia 3: 1;26~1;27. 1951. 



298 PHYTOLOGIA Vol. 10, no. k 

Bibliography: Moldenke, Phytoiogia 3: U26— U27 & U5U. 195lj 
Moldenke, Biol. Abstr. 25: 3051. 195lj G. Taylor, Ind. Kew. Suppl, 
12: 1U9. 1959 i Moldenke, RfisumS 110 & U73. 1959. 

Herbj stems slender, obtusely tetragonal, densely hirsute- 
pubescent with sordid-grayish hairsj nodes not annulate; princi- 
pal intemodes 0.8 — 2.5 cm. long; leaves deciis sate-opposite, ses- 
sile or practically so, ovate, 1.5 — 2.5 cm. long, 1.2 — 2.2 cm. 
wide, subacute at the apex, rounded or truncate at the base, 
coarsely and irregularly dentate along the margins, the lowest 
teeth simost lobe-like, rather densely hirsutulous-pubescent on 
both surfaces, especially beneath; midrib slander, impressed a- 
bove, prorainulous beneath; secondaries filiform, h — 7 per side, 
the lower ones issuing palmately frcxn the base of the blade, im- 
pressed above, prominulous beneath, ascending, hardly arcuate; 
veinlet reticulation rather abundant, impressed above, prominu- 
lous beneath; inflorescence terminal, the spikes subcapitate, 
densely many- flowered, about 2 cm. long and wide during anthesis, 
sometimes ■with 2 or a few flowers slightly separate frcm the main 
head; peduncles abbreviated, mostly 1 — 1.5 cm, long, densely hir- 
sutulous-pubescent; bractlets narrowly lanceolate, 7 — 8 mm. long, 
1 — 1.5 mm. wide at the base, densely pubescent, attenuate at the 
apex; calyx cylindric, about 9 mm. long, strongly 5-costate, 
densely hirsutulous-pubescent or hirsutulous on the outside, its 
rim irregularly 5-subulate-toothed; corolla hypocrateriform, 
showy, its tube about 1 cm. long, very sparsely pilosulous on 
the outside, the limb almost 1 cm. wide. 

The type of this species was collected by Edwin Friedrichs 
( no. 27901 ) in thickets at Campo do JordSo, SSo Paulo, Brazil, in 
January, I9hh, and is deposited in the herbarium of the Colegio 
Anchieta at Porto Alegre, Brazil. In all, 3 herbarium specimens, 
including the type, and 2 mounted photographs have been examined 
by me. 

Citations: BRAZIL: SSo Paulo: Friedrichs 27901 (N— isotype, N- 
photo of type, Kb— type, Z — photo of type); Lanstyack s.n. [Herb. 
Rio de Janeiro 33107] (B) . 

VERB0IA PADLSENI R. A. Phil., Anal. Univ. Chile 90; 607. I896. 

Synonymy: Verbena porrigens var. paulseni (R. A. Phil.) Aceve- 
do de Vargas, Bol, Mus, Nac. Hist. Nat, Chile 25 J 59. 1951. 

Bibliography: R. A. Phil., Anal. Univ. Chile 90s 607 . I896; 
Durand & Jacks., Ind. Kew. Suppl. 1: U5l. 1906; Reiche, Fl. Chile 
5: 291. 19 10: Moldenke, Known Geogr. Distrib, Verbenac., [ed. 1], 
U2 & 102 (19U2) and [ed. 2], 102 & 199. 19ii9; Moldenke, Phytoio- 
gia 3: 75. I9I49; Acevedo de Vargas. Bol. Mus. Nac. Hist. Nat. 
Chile 25: 59. 1951; Biol. Abstr. 28: 90li. I951i; Moldenke, R6sum6 
122, 372, & li73. 1959; Moldenke, R6sum6 Suppl. kt 5 (1962) and 5: 
6. 1962. 

Suffruticose, procumbent, hirtous; stems ascending or erect; 
leaves 3-parted, rarely 5-pairted-pinnatifid, about 2 cm. long and 
1 cm. wide, the segnents narrowly linear, revolute-margined, or 
almost filiform like the rachis of the blade, hardly 1 mm. wide. 



196U Moldenke, Monograph of Verbena 299 

obtuse at the apex, gtrigose; peduncles about 12 cm, long; spikes 
long-pedunculate, many-flowered, capitate; bractlets about U mm. 
long, half as long as the calyx, lanceolate, long-ciliate at the 
base; calyx hispid, 8 mm. long; corolla-tube glabrous, 11 mm. 
long, about 1 1/2 times the length of the calyx, dark-violet, 
pilose in the throat, becoming brownish in drying; anther appen- 
dages exserted, black. 

The type of this little-known species was collected by the or- 
nithologist, Ferdinand Paulsen — in whose honor it is named — 
near Quillota, Valparadso, Chile, in September, 188$, and is de- 
posited in the herbarium of the Museo Nacional de Historia Natur- 
al at Santiago, Chile. Philippi states that the plant has the 
habit of V. sulphurea D, Don, but Acevedo de Vargas (1951) is of 
the opinion that it is a mere variety of V. porrigens R. A. 
Phil., in which opinion she may be coirrect. She says "Difiere 
del tipo por sus hojas mis angostas y largas y por sus flores 
violSceas." She follows Durand & Jackson (1906) in dating Phil- 
ippi' s originaO. work as "1895". She cites M6dano3 s.n. [Concan, 
2.X.l881i] (Sg— 68378), not as yet seen by me. The photograph of 
the type specimen, in the Britton Herbarium, seems to have "1835" 
written on the label as the year of collection, rather than 
"1885" as stated by Acevedo de Vargas. Judging frcxn this photo- 
graph, it is possible that the species is conspedific with V. 
cimiingii Moldenke. 

The species is said to have been collected by Grandjot at 1200 
meters altitude near Santiago, Chile, where it had dark-red 
flowers and bloomed in November. It is known to me thus far only 
from the type photograph. 

Citations: CHILE: Valparaiso: F. Paulsen s.n. [Quillota, Sept, 
1885; Herb. Mus, Nac. Hist. Nat, Chile 5U728] (N— photo of type). 

VHIBENA PERAKII (Covaa & Schnack) Moldenke, Phytologia 2: lli9— 
150. 19U6. 

Synonymy: V erbena erinoldes var, glandulifera Sanzin, Anal, 
Soc. Cientif ."Argent. 88: 129—133, fig. 3Ub. 1919. Glandular ia 
perakii Covas & Schnack, Revist, Argent, Agron, 11: 89 — 91, fig. 
1. I9UU, Verbena dissecta f . glandulifera (Sanzin) Moldenke, 
Phytologia 2: HxQ . 19ii6, 

Bibliography: Sanzin, Anal. Soc. Cientif. Argent, 88: 129 — 
131, fig. 3Ub. 1919; Stapf, Ind. Lond. 6: U29. 1931; Covas & 
Schnack, Revist. Argent. Agron. 11: 89—91, 96, L 97, fig. 1. 
19hh; Schnack & Cowas, Darwiniana 7: 71—75, pi. 1 C & 2 B. 19U5; 
Covas i Schnack, Darwiniana 7: 86. 19ii5; Moldenke, Phytologia 2: 
1U8~150. 19U6i Moldenke, Alph. List Invalid Names Suppl. 1: 10 & 
2li. 19U7; H. N. & A. L. Moldenke, PI. Life 2: 75. 19U8; Moldenke, 
Known Geogr, Distrib. Verbenac., [ed. 2], 106, 197, &; 199. l9U9j 
E. J, Salisb., Ind. Kew. Suppl. 11: 101 & 263. 1953; Moldenke, 
R6sum6 127, 128, 296, 36U, U71, & U73. 1959; Moldenke, Phytologia 
8: 123 (I96I) and 8: 378, 1962; Moldenke, Rlsum^ Suppl. i;: 17—19 
(1962) and 5: 6 & 7. 1962; Moldenke, Phiytologla 9: 70, 131, 39U, 



300 PHTTOLOGIA Vol. 10, no. U 

& UOO (1963) and 10: 136. I96U. 

Illustrations: Sanzin, Anal. Soc, Cientif. Argent, 88: 170, 
fig, 3Ub, 1919; Covas & Schnack, Revist, Argent. Agron. Ut 90, 
fig. 1. 19l4tj Schnack & Covas, Danriniana 7: pi. 1 C & 2 B. 19U5. 

Perennial herb; stems erect or suberect, cylindric-quadrangu- 
lar, pubescent with two kinds of hairs, the one type simple, to 
1,5 mm, long, and erect, the other type to 0.5 mm, long and 
glandulif erous ; leaves decus sate-opposite j petioles k — 6 mm, long; 
leaf -blades ovate, tripartite-pinnatilobed, 2,U — 3.8 cm, long, 
1,5 — 2,2 cm. wide, attenuate into the petiole at the base, with 
obtuse lobes, pubescent especially above and on the venation be- 
neath, the pubescemse composed of simple hairs 0,2 — 1 mm, long 
and glandulif erous ones almost 0,3 mm, long; inflorescence spi- 
cate, often solitary, abbreviated during anthesis, elongate after 
an thesis; bractlets lanceolate, more than half the length of the 
calyx, slightly pubescent on the veins, loosely ciliate on the 
margins; calyx tubular, almost 6 mm, long, composed of both sub- 
erect and glsmdulif erous hairs; corolla hypocrateriform, varying 
from purple, lilac, pinkish-lilac, lilac-rose, or rose, to violet 
or blue, sometimes described as "rose, drying violet", pubescent 
on the outer surface at and near the apex of the tube, glabrous on 
the upper part of the limb; stamens typical of Glandulaida , the 
two upper ones with subcylindric glandular appendages, the thecae 
small, included; pistil typical; ovary 1 mm, long; style 8 mm, 
long; cocci subcylindric, almost 3 mm, long, obtuse at the apex, 
truncate at the base, reticulate on the upper central portion of 
the back; chromosome number: n « 5* 

The type of this species was collected by Guillermo Covas 
( no, 2110 ) at Dique Papagallos, in the department of Las Heras, 
Mendoza, Argentina, on December 3» 19U3, and is deposited in the 
herbarium of the Instituto de Botlnica DarwLnion at San Isidro, 
Argentina. The species is named in honor of Juan Tomis Perak 
(1916— I9li3) , ill-starred Argentinian geneticist, who did note- 
irorthy experimentation on the effects of colchicine on diploid 
species of cultivated plants and on the duplication of chromo- 
somes, who obtained tetraploid maize, experimented on mutations 
induced by short-wave radiation, x-rays, and ultraviolet rays, and 
who died of radiation poisoning at the age of 27. The relation of 
the pistil length to pollen-grain size is discussed ty Covas & 
Schnack il9k$) . 

The species has been found in dry riverbeds, xmder pine trees 
on arid hillsides, and growing as a weed in cultivated soil, at 
altitudes of 820 to 1200 meters, blooming fran August to February 
and in May, fruiting in December, 

Verbena erinoides var, glandulif era was apparently based by 
Sanzin on his nos, 139, 1700 , 3099, 3129 , and 3130 , collected a- 
bout the city of Mendoza and exactly at the edge of the Cordil- 
lera at altitudes of 1000 to 1200 meters. He says of it: "Cerca 
de la ciudad de Mendoza, y precisamente del lado de la Cordillera 
a una altura de 1000 y 1200 metres, abunda una variedad ( Herb « 
Sanzin 139, 1700, 3099, 3129, 3130), que lleva gldndulas en el 



196U Moldenke, Monograph of Verbena 301 

cdliz y que tiene las hojas anchas, triangulares, de base cuneada 
y trlfidas o tripartidas con los segmentos casi enteros o con 
unos lobulitos laterales: A typo differt caule, foliis, calici- 
busque hirsutis, pilis glanduliferis mixtis. Tube calice subduplo 
longiore, appendicibus antherarum subexsertis clavatis violaceis. 
Laciniis foliarum lanceolatis (Osten, in 11 tt,). M5s al sur y a 
las mismas alturas indicadas exLste otre variedad que se acerca 
mis al tipo por sus hojas tripartido-pinatifidas con segmentos 
angostos, pero que se diferencia esencialmente por sus glSndulas 
estaminales apenas salientes de la garganta del tubo corolar en 
vez deser inclusas . La V. mendocina Phil, es intermediaria entre 
estas dos variedades, pues el examen de ejemplares aut^nticos de 
Philippi, del museo de Santiago, me permiti6 constatar que tiene 
hojas de dos clases, idfoticas en la forma a las hojas de lais dos 
variedades citadas. El cardcter de los tallos erguidos de la V« 
mendocina , no es const ante, pues en la variedad glandulifera h\iy 
individuos erguidos y otros semirastreros . Por todo esto ma par- 
ece convenient e unir en xina sola las dos especies, V. erinoides y 
V« mendocina." The Gray Herbarium's Card Index to New Species 
states erroneously that Sanzin' s plant is from Peru and Brazil, 

Covas & Schnack (19Ui) say "Esta especie fu6 descripta por 
Osten (ex Sanzin, en Anal, Soc. Cient, Argent. 88: 131, 1919) co- 
mo Verbena erinoides var, glandulifera , pero evidentemente se 
trata de una buena especie que nada tiene que ver con Glandularia 
laciniata (L.) Schnack et Covas (- Verbena erinoides Lam.) . En la 
clava que figura al final de esta trabajo se podrAn apreciar los 
principales caractei^es diferenciales .El polen de esta espec- 
ie es normal, Consignamos esta observaci6n porque en varias es- 
pecies de Glandularia el polen es irregular, con granos normales 
y abortivos en porcentaje variable, pudi^ndose tambi6n encontrar 
granos de polen con m&s de tres poros germinativos , . .G^ Perakii 
posee flores relativemente grandes y de un atractivo color viold- 
ceo, por lo que merece ser introducida al cultivo." 

Hybrids are known between V. perakii and V, peiniviana (L.) 
Britton ^xV, tentamenta Moldenke), with V^ santiaguensis (Covas & 
Schnack) Moldenke (=xV. gonzalezi Moldenke), and with V. tenuisec- 
ta Briq, (-xV, nisa Moldenke). 

In all, 20 herbaritun specimens have been examined by me. 

Citations: ARGENTINA: La Rioja: Ruiz Leal 16293 (Z) . Mendoza: 
H. H, Bartlett 19351i (Mi); D. 0. King 139 (Bm); Mexia U372 (Ca— 
?60S23)i Ruiz Leal 922 (N), 33^ W, 3390 (N), WH (N), 631U (N), 
8U53 (N), 9378 (N), 9503 (Rl), 9^0h (N); Sanzin s.n. [Ruiz Leal 
152^] (N); Semper s.n, [Ruiz Leal Ul$8] (N), s.n. [Ruiz Leal 9$38] 
(N), 3.n. [Ruiz Leal 98U9] (N). San Juan: Ruiz Leal 16 388 (Rl)| 
Ruiz Leal & Roig 189$6 (Ok) , CULTIVATED: New York: H. N. Molden - 
ke I8238 (N). 

VERBENA PKRE3INIS Wooton, Bull, Torr. Bot. Club 25: 262, I898. 



302 PHYTOLOGIA Vol. 10, no. k 

Synonymy s Verbena perenna Wooton ex Moldenke, Suppl. List In- 
valid Names 9, in syn. 19ill« Verbena perrenis Wooton ex Moldenke, 
R^STimS Suppl. 3: UO, in syn. 1962. Bouchea perennls Wooton ex 
Moldenke, RfisumI Suppl. $: 6, in syn. 1962. Verbena perennis 
Woot. Sc. Standi, ex Moldenke, RSsumfi Suppl. ^: 8, in syn. 1962. 

Bibliographer: Wooton, Bull. Torr. Bot. Club 25: 262. I898} 
Thiselton-Dyer, ind. Kew. Suppl. 2: 191. 190U} P. C. Standi., 
Contrib. U, S. Nat. Herb. 13: I6I, 173, & 211. 1910 j Glaz., M&i. 
Soc. Bot, France 3: 5UU. 1911} Perry, Ann. Mo. Bot. Qard. 20; 21*8, 
260, 299 — 300, & 356. 1933; Steyermsirk & Moore, Ann. Mo. Bot, 
Gard. 20: 805. 1933} Cory, Texas Agr. E^. Sta. Bull. 550: 89. 
1937; Sperry, Sul Ross State Teach. Coll, Bull, 22: Ul. I9I4I} 
Moldenke, Suppl. List Invalid Names 9* I9I4I} Moldenke in Lundell, 
Fl, Texas 3 (l)t 16 & 31, 19li2; Moldenke, Known Geogr, Distrib. 
Verbenac,, [ed. 1], 1>-15, 19, & 102, 19U2} Moldenke, Alph, Ust 
Invalid Names k9* 19U2} Moldenke, Bot. Gaz. I06; I6I, 19U5} Mol- 
denke, Alph. List Cit. 1: 126, 127, 15U, 175, 182, 203, & 283. 
19li6; Moldenke, Phytologia 2: 328. 19U7; Moldenke, Wrightia 1: 
228, I9U8} H. N. & A. L, Moldenke, Pi. Life 2: 65. 19U8} Moldenke, 
Alph, List Cit. 2: UOl, U67, U69, U71, U77, U93, 506, 523, 525, 
526, 532, 5U9, & 595 (19U8), 3: 708, 729, 752, 768, 797, 8U3, 
9lli, 939, 95U, 965, 966, & 990 (19li9), and U: 1107—1110, 1121, 
1122, llljl, 1150, I2U0, & 12U3. 19U9} Moldenke, Known Geogr. Dis- 
trib. Verbenac, [ed, 2], 2U— 26, 33, & 199. 19U9; Moldenke, r6- 
sumS 29, 31, 32, 39, 372, & U73. 1959} Moldenke, Phytologia 8: 
I2U. I96I; Lewis & Oliv., Am. Journ. Bot. Ii8: 639— 6UI. 1961; 
Moldenke, RSsumS Suppl. 3: 7, 8, & UO (1962) and 5: U, 6, & 8. 
1962} Moldenke, Phytologia 8: U72 (I963) and 9: 113* 1963} Mol- 
denke in Shreve & Wiggins, Veg. & Fl. Son, Des, 2: 1239— 12U0. 
I96U. 

Illustrations: Lewis &. Oliv., Am. Journ, Bot, 1*8: 6U0. I96I. 

Suffruticose perennial herb, 3 — k dm, tallj roots perennial; 
rootstock woo^y, heavy, perennial} stems several or numerous, 
from a woody base, varying from erect to ascending or divari- 
cately ascending-erect, more or less strictly branched, glabrate 
or often finely glandular and slightly hiapidulous with short 
stiff antrorse hairs, strlate-angled; leaves predominately line- 
ar, 1— it cm. long, entire or the lower ones 3-lobed to pinnately 
few-lobed, erect-ascending, sparsely hispidulous with short sharp 
stiff antrorse hairs, the margins revolutej spikes terminal, ter- 
minating the stems and branches, pedunculate, slender-filiform, 
elongate, loosely max^y-flowered; bractlets ovate, persistent, 
1.5— '3 mm. long, acute at the apex, hispidulous (or glabrate a- 
bove), ciliate; flowers small, sessile, 7—8 mm. long; calyx tu- 
bular, h — 5 ram. long, herbaceous-ribbed, more abundantly pubes- 
cent along the ribs, hyaline and glabrous between the ribs, the 
teeth short- triangular, equal or subequal, acute; corolla hypo- 
crateriform, varying fran blue, deep-blue, or blue-lavender to 
purplish-blue, purplish-lavender, lavender, pink-lavender, or 
purple, its tube slightly longer than the calyx, pubescent, ex- 
panded just below the throat, forming a ring in which the almost 



196Ii Moldenke, Monograph of Verbena 303 

sessile anthers lie, the thjroat filled -with hairs, the limb 5- 
parted, bilabiate, 5 — 7 nm. wide, the lobes elliptic, repsuid, the 
lower lip particiilarly undulate-margined; style short, included; 
stigma unequally 2-lobed, clavate; fruits more or less remote, 5- 
7 mm, apart on the rachis; schizocarp about 3 mm. long, strongly 
constricted along the lines of cleavage; cocci U, cylindric or 
subcylindric, r^ticulate-scrobiculate except at the base, glab- 
rous, brownish, minutely retrorsely scabrous on the conmissure, 
enclosed in the persistent frui ting-calyx, the commissural faces 
smooth or slightly scabrous and not extending to the tip of the 
nutlet; chromosome number: n ■ 7. 

The type of this species was collected by Elmer Ottis Wooton 
(no. 187 ), growing in crevices of rocks at 1800 meters altitude 
along the road about two miles west of the Mescalero Agency in 
the White Mountains, Lincoln County, New Mexico, on July 21, 
1897, and is deposited in the Britton Herbarium at the New York 
Botanical Gar^len. Standley (1910) states that the type locality 
is in Otero County, but Wooton" s labels definitely are inscribed 
•♦Lincoln County", Wooton (I898) says that •This species is most 
nearly related to V. canescens H«B.K, var. neo-mexicana Gray, but 
may be easily separated from that variety (?) by the linear, gen- 
erally simple revolute leaves and the pec\iliar pubescence." 

The species has been found in limestone or calcareous stony 
soil, on rough grassy slopes and the slopes of canyons, on hills, 
flats, gravelly hills and hillsides, rocky slopes and ledges, low 
rocky ridges, banks, rocky hillsides, rocky limestone hills in 
the thorn-shrub-grassland conmunity, calcareo\is ledges, and road- 
side prairies, on arid rocky hills, foothills, limestone hills, 
in juniper woodland and the juniper belt, and on calcareous grav- 
el among scrub oaks, at altitudes from 3200 to 8000 feet, flower- 
ing frcra March to November, and fruiting frcsa April to August and 
in October. Waterfall found it growing with Que reus , Juniperus , 
and Pinus in Culberson County, Texas; Janszen found it "in buff 
silt, alluvial cover, hilly topograqshy"; and Mueller describes it 
as "sparse on grassy limestone slopes" in the same county and "in- 
frequent" in Pecos County, Wamock avers that it is "infrequent" 
or "scattered" and also "abundant on hills" in Brewster County, 
and "frequent along highway in lower limestone slopes". Mueller 
found it "scattered on limestone on brow of canyon" in Hudspeth 
County, while McVaugh encountered it in "rocky limestone hills, 
abundant in thorn-shrub-grassland community" in Pecos County. 
Mueller found it "sparse on rocky arroyo banks" in Coahuila. In 
ShrevB & Wiggins (I96I1) it is said to be foxind "On rough grassy 
slopes, gravelly or rocky hillsides, flats, canyons, and low 
rocky ridges, Sonoran to Transition Zones, western Texas to Pima 
County, Arizona, and to Coahuila." 

The M.S. Young collection cited hereinafter appears to repre- 
sent a form with very large corollas, and is so noted also by the 
collector. It is perhaps worthy of nomanclatural recognition, 
but evidence from many other specimens indicates that the flowers 
vary in size and apparently shrivel quickly after being picked and 



30U PHYTOLOaiA Vol. 10, no. k 

before being pressed. Tharp 3682 exhibits corollas that are inr- 
termediate in size. 

Glaziou (1911) cites his no, 17716a from SSo Paulo, Brazil, as 
"V. perennis Woot." with a question, and describes it as an herb 
-with white flowers, blooming in August sind September. What his 
plant really is, I cannot as yet say, since I have not as yet 
seen any material of the coUectionj his 17716 is cited by him as 
V. ephedroides Cham, 

Herbarium material of V, perennis has been mlsidentified and 
distributed in herbaria as V. oauiescens H,B,K., V. halei Small, V, 
menthaefolia Benth., V. neomexicana (A. Gray) Small, V, neomexi" 
cana var, hlrtella Perry, V. xutha Lehm,, Boiichea linifolia A. 
Gray, Buchnera elongata Sw., and Lobelia sp. 

Perry (1933) cites the following 17 additional specimens not 
as yet seen by me: TEXAS: Culberson Co.: Clarke U2$0 (E)j Havard 
197 (0); Moore & Steyermark 36II (E)j M, S. Young s.n. [Gixadalupe 
Mts., 8/13A6] (E), Martin Co.: G. E. Seler s.n. [Loyola, 5 Nov. 
1902] (G). NEW MEXICO: Eddy Co.: Wooton s.n. [Queen, Aug. I909] 
(E). Uncoln Co.: F. Sj, Earle 387 (E)j J. Skehan 20 (E, F, G); 
Wooton 187 (E — isotype, G — Isotype), s.n. [Ruidosa Creek, Aug, 5, 
1901] (E) . Otero Co.: P. C^ Standley 3.n. [17 Aiig. I899] (W) . 
Sierra Co.: £. B. Metcalfe 1^68 (E, F, G) . She makes the follow- 
ing comments: "The relationship of this species is some?rtiat anom- 
alous. The loblng of the leaves and the character of the nutlets 
seem to ally it with V. canescens and its relatives; whereas the 
pubescence and the predominauice of practically entire linear- 
oblong leaves recall V. simplex . It could scarcely be confused 
with either, since the character ccmibination of an open spike, 
very narrow leaves, and sparsely short-strigillose hairs is not 
fo\md elsewhere in the group under consideration." 

In all, Ihh herbarium specimens and 1 mounted clipping have 
been examined by me, including the types of all the nanes involved. 

Citations: TEXAS: Brewster Co.: Cory 30082 (N)} G. L. Fisher 
s.n. [Alpine, Aug. 2U, 1932] (Hp, Wi); D. C. Ingram 2736 (Ar~ 
1U79U, Mi, W— II489829)} Parks & Cory 18$0U (Tr~l6l73)i 0. E. 
Speriy T.12g (Om, W— 16792U8), T.563 (Fs, Om, W— 1766U37), T,$6U 
(Om), T,813 (On); Steiger 20 (N), IO66 (N), 12U8 (N)j Tharp 3682 
(Au, W^^3I59910); B. H. Wamock 287 (Au), li65o7^982 (Au— 12239$), 
21090 (Au), 21205 (Au), 21279 (Au), 21827 (Al, Ca— 882787, Du— 
327912, N, N, Ok, S, Ur), ff,l69 (N), W.282 (N), W.287 (N)i Var- 
nock & Hinckley 3926 (N); Whitehouse 18633 (Ml) . Culberson Co.: 
Correli & Johnston 22006 (Ld)j Correll & Rollins 23897 (Ld); 
Hinckley )|I|)|1 (N, W— 2005U60)j Janszen h2h (Au— 122^01); C. H. 
Muller 8253 (Rf, Sm)j Ripley & Barneby 111^1 (Gg~3826l7, N)} 
Smith & Robertson 217 (Ar— 305860); Waterfall 3795 (N, St, T\i~ 
128^oy, 1glO (Au,"Ny. 5209 (Gg— 316101, N, Ok, Sm, St, St); 



196U Moldanke, Monograph of Verbena 305 

Whltehouse &7k2 (Au), lg968 (Sm), 17019 (Mi, N), 3,n. [Signal 
Peak, 7.5.31] (Au); M. S. Young s.n. [Guadalupe Mts., 8/13A6] 
(Au, Au), Hudspeth Co.: Cj^ H. Muller 621ii (Rf, Sm, St)j Turner , 
Tharp , & Wamock 3233 (Au~122397) . Jeff Davis Co.: E. D. Schnlt 
s.n. [Davis Mts., 8/2^928] (Wi) . Pecos Co.: Hinckle7 U839 (W— 
2095622); R. MeVaugh 7935 (Ar— 233930, Au— 178252, Du--355386, Mi); 
B. H. Wamock 13hhSa. (Au--123222, Rf ) . NEW MEXICO: Chaves Co.: 
G. £, Ikenberry 376 (St) . Eddy Co.: V. Bailey s.n. [Carlsbad 
Cave, April 193U] (W--1220Ui5) J Coty s.n. [Carlsbad, U-2U-192U] 
(Tr), s.n. [Carlsbad, li-2>1925] (Tr); O. Degener 5038 (Ms, N)j 
Havard s.n. [Guadalupe Mts., Oct. l88l] (W~lli7569, W— 218869); 
Hers hey s.n. [Guadalupe Mts., 5/12/39] (Bt— 59779); A. Nelson 
llii06 (Ca— 500718, Du— 218829, S, Um— 17, Up); P. C. Standley 
li0686 (W--1222072) ; Wilkens 1568 (En); Wooton s.n. [Queen, Aug. 2, 
1909] (W— 56U65ii), sji. [Queen, Aug. 3, 1909] (W— 56U653). Guada- 
lupe Co.: Arsene & Benedict 16656 (W-— 1033519) . Lincoln Co.: F. 
S. Earle 619 (N);~Earle & Earle 387 (N, Po— 63866, W— 382530), s^ 
n, [Lincoln, 7/31A900] (N); Eggleston I89IO (N); Goodman & Water - 
fall U967 (Ok); Hitchcock , Rethke , & Raadshooven U202 (Ca— 603911, 
Du— 256977, En, Gg— 3O8I38, 10—1146632, La, Pi— 90851, Se— 182U1, 
Se— U1887, Ua— 28658); E. L, Reed 3655 (Bl— li2331, I); £. Skehan 
20 (Ca— 25153, Cm, Ka, N, N, Po— 6I16U6, W— 350138), 1316 (Ca— 
882788); Wooton 187 (Ca— I2U666— isotype, Ka— isotype, N— type, 
Po— 70679 — isotype, W — 736221 — isotype), s.n. [Ruidoso Creek, Aug. 
5, 1901] (S, W— 73687U). Otero Co.: W. V. Fisher s.n. [August 20, 
1950] (St); Wooton s.n. [Tularosa Creek, Aug. 19, 1899] (W— 
563953, W^736875). Sierra Co.: 0^ B_. Metcalfe 1568 (Gg— 31386, 
N, n, W— 890290). ARIZONA: Pima Co.: M. E. Jones 2ii99lf (Po— 
192935). MEHCO: Coahuila: E^ G. Marsh 859 (Au), 1378 (Au— 
213159, St); Cj^ H, Muller 30li5 (Ca— 7201^7, Mi, Rf ) . MOUNTED 
CLIPPINGS: Bull. Torr. Bot . Club 25: 262. I898 (W) . 

VERBENA PERENNIS var. JOIINSTONI Moldenke, Phytologla 2: 150. 19U6. 
Synonymy: Verbena shrevei Johnston ex Moldenke, Phytologia 2: 

150, in syn. I9U6, 

Bibliography: Moldenke, Phytologia 2: 150 (19U6) and 2: 331 & 
38U. 19U7; H. N. & A. L. Moldenke, PI. Life 2: 65 & 83. 19U8; Mol- 
denke, Alph. List Cit. 2: 370 & U97 (19U8), 3: 963 (19U9), and h: 
I2U6. I9U9; Moldenke, Known Geogr. Distrib. Verbenac, {ed. 2], 
33 & 199. I9U9; Moldenke, R6sum5 39, 37U, & U73. 1959; Moldenke, 
Phytologia 8: U73. 1963. 

This variety differs from the typical form of the species in 
having its stems, leaves, and rachis densely spreading hirtellous, 
the leaves to 5 cm. long, the lowermost often irith several linear 
lobes. The corolla is described sis blue-purple. 

The type of the variety was collected by L. R. Stanford, Ken- 



306 PHYTOLOGIA Vol. 10, no. h 

neth lynn Retherford, and R. D. Northcraft ( no. 91^ ) among varied 
vegetation of large shrubs, small trees, and herbs, in a broad 
damp riverbed, at an altitude of 19^0 meters, 12 kilometers 
northwest of Palmillas on the road to Miquihuana, Tamaulipas, 
Mexico, on August lU, 19U1, and is deposited in the Britton Her- 
barium at the New York Botanical Garden. It was originally dis- 
tributed as V. shrevei by Ivan Mxirray Johnston (I898--I96O), ex- 
pert on the Boraginaceae and temperate South American plants, in 
whose honor it is named. By his specific epithet. Dr. Johnston 
meant to honor Forrest Shreve (18 78 — 19^0), distinguished Ameri- 
can ecologist and plant collector in northern Mexico and south- 
western United States. 

The variety has been found at altitudes of 19^0 to 2^00 meters, 
flowering in August. It has been misidentified and distributed 
in herbaria as V. neomexicana (A. Gray) Small, V. neomexicana var. 
hlrtella Perry, and V. neomexicana var. ::ylopoda Perry. Stanford, 
Retherford , & Northcraft I4B6 is a mixture with V. canescens H.B.K. 
and (perhaps) with V. neomexicana var, hirtella Perry. Their no, 
$07 may actually have been collected in Coahuila, since the label 
is not definite . Shreve & Tinkham 9728 was distributed as "Ver - 
bena n, sp," and was collected "in the pinyons" at 69OO feet al- 
titude, flowering and fruiting in August, 

In all, 8 herbariian specimens, including the types of both 
names involved, have been examined by me. 

Citations: MEXICO: Coahuila: Stanford , Retherford , & North- 
craft U86, in part (Du— 288703, N) . Nuevo Le6n: Herb,"'ln8t. Biol. 
Univ. Nac. Mex, 2375? (Me) , 7138 (Me) j Shreve & Tinkham 9728 (Tu— 
1257liO). Tamaulipas: Stanford , Retherford , & Northcraft 915 (N— 
type, Tu — 15150— isotype) . Zacatecas: Stanford , Retherford , ft 
Northcraft 507 (Tu— 10909). " 

xVERBENA. PERPLEXA Moldenke, RfisumS Suppl. U: U, Ik, & 16, hyponym 
(1962); hybr. nov. 

Synonymy: Verbena gooddingii Briq, x V, bipinnatiflda Nutt, ex 
Moldenke, R6sum6 Suppl. U: 16, in syn. I962. Verbena bipinnati- 
fida Nutt. X V. gooddingii Briq. ex Moldenke, R^sum^ Suppl. U: lU, 
in syn. I962. 

Bibliography: Moldenke, R6sum6 Suppl. U: U, 111, & 16, 1962 j Mol- 
denke, Phytologia 8: 378 & 381. I962. 

Planta hybrida aspectu V. bipinnatiflda et V. gooddingii inter- 
media; foliis inciso-pinnatifidis utrinque hispidisj inflorescen- 
tiis terminalibus multifloris conges tis dense albo-hirsutisj 
bracteolis calycem excedentibus albo-hirsutls . 

Herb, apparently a natural hybrid between V. bipinnatiflda Nutt, 

and V, gooddingii Briq., with intermediate characters; stems rather 
slender, tetragonal, white-hirsute throughout, less densely so in 
age; principal intemodea 1.5~U,5 cni, long; leaves decussate- 
opposite, usually with abbreviated leafy shoots in their axils; pet- 



196U Moldenke, Monograph of Verbena 307 

loles about 1 cm. long, winged, white-hirsute; leaif -blades in- 
cised-pinnatifid, rather densely white-villous on both svirfaces; 
inflorescence terminating the branches, erect, many- flowered, 
congested, at least during anthesis, conspicuously and densely 
white-villous; peduncles 2 — 5 cm. long, densely white- and 
spreading- villous; bractlets elongate, narrow, lanceolate, equal- 
ing or surpassing the calyx, mostly 10—12 mm. long, densely 
white-villous on both surfaces; calyx tubular, 3 — 10 mm. long, 
densely white-villous, the rim subulate-toothed, the teeth iu>- 
equally elongate; corolla-tube equaling or somewhat exserted frcm 
the calyx, densely white-pubescent outside above the calyx- tube, 
the Hmh about 5 nun. wide. 

The type of this hybrid was collected by Robert A. Darrow five 
miles south of Patagonia, Santa Cruz County, Arizona, on March 20, 
1938, and is deposited in the Brit ton Herbaidum at the New York 
Botanical Garden. The plant has been collected at an altitude of 
5200 feet, flowering in March and April. It has been misidenti- 
fied and distributed in herbaria as V. bipinnatifida Nutt., V» 
gooddl ngii Briq,, and V. macdougalii Heller. Only 2 herbarium 
specimens, including the type, have been exsonined by me. 

Citations: ARIZONA: Cochise Co.: Q. Martin s.n, [April 12, 
i960] (Hi— 19li9^). Santa Cruz Co.: Darrow s .n. [March 20, 1938] 
(N— type) . 

xVHlBENA PERRIANA Moldenke, Revist. Sudam. Bot. U: 19. 1937. 

Synonymy: ? Verbena laciniata Raf ., Herb. Raf . 61, nom. nud, 
I833 [not V. laciniata Briq., I96O, nor Kuntze, I9UI, nor (L.) 
Briq., I90II7 nor (Lam.) Briq., 1939, nor Sessfi & Moc, 19liO] . 
Verbena br a ct e os o-urtic aef olia Engelm., Am. Joum. Sci. U6: 101. 
I8UI4.. Verbena urticaefolio-bracteosa Engelm,, Am, Joum, Sci, 
US: 101. l81ju Verbena bracteosa x hastata Webber, Trans. Acad. 
St, Louis 6: UO, 1892. Verbena bracteosa x hastata Rydb,, Fl, 
Rocky Mts. 7U0. 1917. Verbena bracteosa x urticifolia Rydb., Fl. 
Cent, N. Am, 678. 1932. Verbena bracteosa x hastata Mackenzie ex 
Moldenke, Revist. Sudam. Bot. U: 19, in syn, 1937, Verbena 
bracteosa x stricta Britton ex Moldenke, Revist, Sudam, Bot, hi 
19, in syn, 1937. Verbena bracteosa x urticaefolia Mackenzie ex 
Moldenke, Revist. Sudam, Bot. Ii: 19, in syn. 1937, Verbena 
bracteosa x urticifolia Eggert ex Moldenke, Revist, Sudam, Bot. h: 
19, in syn, 1937. Verbena bracteoso-stricta Engelm. ex Moldenke, 
Revist. Sudam, Bot. k: 19, in syn, 1937, Verbena hastata x 
bracteosa Rydb, ex Moldenke, Revist. Sudam. Bot. U: 19, in syTi, 
1937. Verbena officinalis x bracteosa Barnes ex Moldenke, Revist. 
Sudan, Bot. k: 19, in syn. 1937. Verbena stricta x bracteosa A. 
S. Hitchc. ex Moldenke, Revist. Sudam. Bot. ht 19, in syn. 1937. 
Verbena stricta x urticifolia Stevens (in part) ex Moldenke, Re- 
vist. Sudam. Bot. U: 19, in syn, 1937. Verbena stricto-bracteosa 



308 PHYTOLOGIA Vol. 10, no. U 

Engelm. ex Moldenke, Revist, Sudam. Bot. U: 19, in sjm. 1937* 
Verbena urticlfolia x bracteoaa Eggert ex Moldenke, Revist . Sudam. 
Bot. U: 19, in syn. 1937. Verbena bracteata Lag, & Rodr. x V. 
xirticifolia L. ex Moldenke, Prelim. Alph, List Invalid Names kS, 
in syn. 19liO, Verbena bracteata x hastata Gates, Fl. Kans. 190. 
I9U0. Verbena bracteata x urticifolia Gates, Fl. Kans. I90, I9U0. 
Verbena bracteata x stricta Schneck ex Moldenke, Suppl, List In- 
valid Names 8, in syn. I9UI. Verbena bracteosa x stricta Palmer 
ex Moldenke, Suppl. List Invalid Names 8, in syn. 19iil. Verbena 
bracteosa x stricta Schneck ex Moldenke, Suppl. List Invalid Names 
8, in syn. 19 1*1. Verbena hastata x bracteosa Schneck ex Moldenke, 
Suppl. List Invalid Names 8, in syn. I9UI. Verbena bracteosa x 
stricta Clothier ex Moldenke, Alph. List Invalid Names Suppl. 1: 
22, in syn, 19li7. Verbena bracteosa x urticaefolia Carleton ex 
Moldenke, Alph. List Invalid Names Suppl, 1: 22, in syn. 19U7. 
Verbena bracteosa x urticaefolia Deam ex Moldenke, Alph, List In- 
valid Names Suppl. It 22, in syn. 19U7. Verbena bracteosa x ur- 
ticifolia Stevens ex Moldenke, Alph, List Invalid Names Suppl. It 
22, in syn, 19li7. Verbena urticifolia x bracteata Gates ex Mol- 
denke, Alph, List Invalid Names Suppl. 1: 27, in syn. 19U7, Ver - 
bena urticifolia L, x V, bracteata Lag, & Rodr, ex Moldenke, 
Alph. List Invalid Names Suppl. 1: 27, in syn. 19U7. Verbena 
urticifolia x bracteosa Patterson ex Moldenke, Alph. List Invalid 
Names Suppl, 1: 27, in syn. 19U7, Verbena bracteo-hastata Hall 
ex Moldenke, RfesumI 359, in syn, 1959"! Verbena urticifolio- 
bracteosa Engelm, ex Moldenke, R6sum6 378, in syn, 1959. Verbena 
brachlata Nieuwl. ax Moldenke, R^sum^ Suppl, 6; 10, in syn, I963. 

Bibliograpl^r: Raf,, Herb, Raf, 61, 1833; Engelm,, Am, Joum. 
Sci. U6: 101. l8Uij Webber, Trans. Acad. St, Louis 6: UO, 1892; 
E. J, Palmer, Ann, Mo. Bot. Gard, 3: 292, 1916; Rydb,, Fl, Rocky 
Mts, 7U0. 1917; Wolden, Proc. lo. Acad. Sci. 39: 123. 1932x Rydb., 
n. Cent. N. Am. 678. 1932; Moldenke, Revist. Sudam. Bot. U: I9. 
1937; Gates, Fl. Kans. 190. I9UO; Moldenke, Prelim, Alph. List 
Invalid Names U5— li8. I9UO; C. C. Deam, Fl. Indiana 798 & 1232. 
I9UO; Moldenke, Suppl, List Invalid Names 8, 19U1; Moldenke, Known 
Geogr, Distrib, Verbenac, [ed, 1], 1, 6—11, & 102. 19li2; Molden- 
ke, Alph, List Invalid Names U5, U7, & U9~5l. 19U2; G, N, Jones, 
Fl, 111, [Am. Midi. Nat, Monog, 2:] 216, l9hSi Moldenke, Castanea 
10: 37, 19li5; Moldenke, Am. Journ. Bot. 32: 610. 19li5; Deam, Krie- 
bel, Yuncker, & Friesner, Proc, Ind. Acad. Sci. S^: 56. 19U6; Mol- 
denke, Alph. List Cit. 1: 9, 31, 9h, 110, 113, 1U9--151, 163, I8I, 
193, & 23I. 19146; Hill & Salisb., Ind, Kew, Suppl, 10: 2U2. 19U7| 
Moldenke, Alph, List Invalid Names Suppl, 1; 22 & 27. 19U7} Mol- 
denke, Phytologia 2: 327 (19U7) and 2: U78. I9U8; H. N. & A. L. 
Moldenke, Pl. Life 2: 75. I9I48; Moldenke, Alph. List Cit. 2: 390— 
392, 39li, 396— UOO, li5l, U72, 5Uli, 5ii9, & 596 (19li8), 3: 653, 699, 
706, 723, 788, 790, 793, 800, 822, 852, 887, 90U, 926, 927, 932, & 



196Ii Moldenke, Monograph of Verbena 309 

970 (19U9), and li: 98I, U38, 1139, 1199, 1208, 1221, 1226, 1227, 
1255, 1261, & 1298. 19U9; Moldenke, Knowi Geogr. Distrib. Verben- 
ac., [ed. 2], 2, 12—15, 17, 18, 20, 21, 26, & 199. l9U9j E. D. 
Merr., Ind. Raf , 205 & 295. 19U9j Moldenke in Gleason, New Britton 
& Br. Illustr. Fl., pr. 1, 3: 127, 131, & I3U. 1952; Moldenke, 
Phytologia I^: 185. 1953; Moldenke in Gleason. New Britton & Br. 
Illustr. Fl., pr. 2, 3: 127, 131, & 13U. 1958; Moldenke, Am. Midi. 
Nat. 59: 3U5 & 357—359. 1958; Moldenke, R^sum^ 5. 16— 19, 21, 22, 
25, 26, 32, 359, 360, 365, 371, 375, 377, 378, & U73. 1959; Mol- 
denke, Rfisun^ Suppl. 2: 2. I96O; Moldenke, Phytologia 8: 121 (1961) 
and 8: 268, 272, 279, 280, & U35. 1962; Moldenke, R^sumfi Suppl. 3: 
6 (1962), U: 3 (I962), 5: 7 (1962), 6: 1, 2, & 10 (I963), and 7: 
9. 1963; Steyenn., Fl. Mo. 1260 & 1261, map iBtih. 1963; G. N. 
Jones, Fl. 111., ed. 3 [Am. Midi. Nat. Monog. 7:] 213. 1963; Glea- 
son & Cronquist, Man. Vase. PI. 581. 1963; Moldenke, Phytologia 8; 
U6li (1963), 9: 53, 5U, 215, & 220 (I963), and 9: 356 & 359. 1963; 
Moldenke, Rfisum^ Suppl. 10: 1. I96U. 

Illustrations: Moldenke in Gleaaon, New Britton & Br. Illustr , 
Fl., pr. 1, 3: 13U (1952) and pr. 2, 3: 13U. 1958. 

This is the natural hybrid between V. bracteata Lag. & Rodr, 
and V. urticifolia L. with more or less intermediate characters. 
It resembles V, bracteata in habit, being a large diffuse plant, 
stouter and more erect when young, but sprawling, procumbent, or 
half-prostrate when mature, the stems assurgent or ascending, the 
branches many, large, and diffuse, the leaves broad and laciniate, 
rather than dissected, the spikes long and slender, and the 
bractlets smaller, usually only slightly siurpassing the calyx, u- 
sually not conspicuous nor foliaceous, and mostly only 3 — U (rare- 
ly 7) mm. long. The flowers are blue, pinkish-blue, or white 
with a lavender tint. It is found rather commonly where the 
ranges of the two parental species overlap in central North Amer- 
ica. It is based on the Verbena urticaefolio-bracteosa of Engel- 
mann, of which cotypes were collected by Carl Andreas Geyer at 
Beardtown, Cass County, Illinois, in July and August, 18U2, and 
by George Engelmann — in whose honor it is named — in Saint 
Louis, Missouri, in July (and perhaps at other times), I81i2, the 
originals being deposited in the Torrey Herbarium at the New York 
Botanical Garden. 

Synonymous designations are typified as follows » V. brachiata 
Nieuwl. is based on Nieuwland s.n. , collected at Saint Mary's, 
Saint Joseph County, Indiana, in October 1926, and deposited in 
the United States National Herbarium at Washington; V. bracteata 
X stricta Schneck is based on J. Schneck s.n. , collected in dry 
clayey soil along streets of Mount Carmel, Wabash County, Illinois, 
on June 25, 1879, deposited in the herbarium of the University of 
Illinois; V. bracteata x urticifolia Gates is based on J. L. Shel- 
don s.n. , collected at Lincoln, Lancaster County, Nebraska, on 
July 16, 1898, deposited in the herbarium of West Virginia Univer- 
sity; V. bracteosa x hastata Mackenzie is baaed on K. K. Mackenzie 
s.n., collected in a barnyard at Little Blue Tank, Jackson County, 



310 PHTTOLOGIA Vol. 10, no, U 

Uissouid, on August 2, I896, deposited in the herbarium of the 
University of Illinois; V. bract eoaa x striata Clothier is based 
on G. L. Clothier s.n. , collected at Saint George, Pottawataaie 
County, Kansas, on July U, I896, and deposited in the herbarium 
of Kansas State College; V* bracteoaa x striata Palmer is based 
on E. J. Palmer U02£, collected in waste places at Webb City, 
Jasper Co\mty, Missouri, on August 7, 1913, and deposited in the 
herbarium of the University of Illinois; V« bracteosa x stricta 
Schneck is based on J. Schneck s»n. , collected on G. U. Kneippe's 
place. Mount Carmel, Wabash County, Illinois, on July 8, 1880, 
and deposited in the herbarium of the University of Illinois; V« 
bracteosa x urticaefolia Deam is based on C . C . Deam 39228 , col- 
lected in Fulton County, Indiana, and deposited in the Deam Her- 
barium; V, bracteosa x urticifolia Eggert is based on H. K. D. 
Eggert s.n. , collected at Glencoe on the Meraaer River, Saint 
Louis County, Missouri, on July 28, 1879, and deposited in the 
herbarium of the Carnegie Museum; Verbena bracteoso- stricta 
Engelm. is based on C. A. Geyer s.n. , collected at Beardstoim, 
Cass County, Illinois, in August, 18U2, and deposited in the 
herbarium of Dartmouth College; V. hastata x bracteoaa Schneck 
is based on J. Schneck s.n. , collected at Mount Carmel, Wabash 
County, Illinois, on June 15, 1888, and deposited in the herbar- 
ium of the University of Illinois; V. stricta x bracteosa Hitchc. 
is based on A. S. Hitchcock 972, collected in Pottawatomie 
County, Kansas, and deposited in the herbarium of Kansas State 
College; V. stricta x urticifolia Stevens is based on G. W, Ste- 
vens 1703 , collected in woods near Alva, Woods County, Oklahoma, 
on July lU, 1913, and deposited in the herbairLum of the Univer- 
sity of Illiaois; V« strict o-bracteosa Engelm. is based on G. En- 
glemann s.n. , collected at Saint Louis, Missouri, in August, 
i8U5, and deposited in the herbarium of Dartmouth College; V. ur- 
ticifolia X bracteata Gates is based on G. L. Clothier s.n. , col- 
lected at Saint George, Pottawatcmie County, Kansas, on July k, 
1896, and deposited in the herbarium of Kansas State College; V. 
urticifolia x bracteosa Eggert is based on H, K^^ D. Eggert s.n. , 
collected at Glencoe, Saint Louis County, Missouri, on July 28, 
1879, and deposited in the herbarium of the New York State Muse- 
um; and V. urticifolia x bracteosa Patterson is based on H. N. 
Patterson 8»n. , collected in Henderson Coiinty, Illinois, in July, 
and deposited in the herbarium of the New York State Museum. It 
should be noted here that the designation, V. stricta x urticifo- 
lia Stevens applies also, in part, to xV. illicita Moldenke. 

The name. Verbena laciniata Raf ,, is placed here provisionally, 
Merrill (I9U9) states that it was published by Rafinesque without 
description, with the type from "Kentucky or Illinois". Whether 
or not this disposition of the name is correct depends on an ulti- 
mate examination of the type, not available as yet to me. The V. 



196U Moldenke, Monograph of Verbena 311 

laclnlata of Briquet ["(L.) Briq." and "(Lam.) Briq,"] is a valid 
species from South Ameidca, that of Kuntze is V. dissecta Willd., 
while that of Sess^ & Mocino is Bouchea prismatica var. laciniata 
Grenz . 

Collectors have forrnd xV. perriana in very sandy soil, clayey 
or dry clayey soil, dry or open ground, sandy open ground, waste 
places, and barnyards, along sandy i*oad3ides, roadsides, and 
streets, near houses, in woods and grassy woods, in bottomlands, 
and on or near riverbanks, to 1200 feet altitude, flowering and 
fruiting from June to September. Schneck reports that it blocsns 
when V. hastata is not yet in flower sind that "the flowers when 
fresh are very much like [those of V.] hastata in shape and color, 
[but the] whole plant [is] procunbent." A specimen in the Colum- 
bia University herbarium, with no collector designated, bears the 

note "Erect or half prostrate I find it assuming the erect 

position while young only. My best specimens are from those which 
are half prostrate. This species is plentiful here. Brendel 
thinks it is only one of the many hybrids." Dodge reports it as 
"plentiful" at Point Edwards, Ontario, Patterson says that it is 
"the most common hybrid Verbena here". Shacklette calls it "a 
weed of meadows and pastures" in Kentucky. Deam describes it as 
a large diffuse plant or sprawling, with assurgent stems and many 
diffuse branches. Hitchcock, on the label of his Iowa City col- 
lection, says "grassy woods near river bankj same as found at 
Hamburg which Watson calls a hybrid." The W. H. Rhoades sai. 
from Edinberg, Johnson County, Indiana, is the finest specimen I 
have as yet seen, Nieuwland descidbes it eis a "plant perfectly 
prostrate in mats a yard wide". Popenoe says "flowers larger 
than in V. bract eosa and irtiite with a lavender tint." 

Gates (I9U0) reports the hybrid from McPherson County, Kansas. 
Herbarium material has been mis identified and distidbuted in her- 
baria under the names V. bracteata Lag, & Rodr,, V, bi^cteosa 
Michx,, V. bracteosa brevibracteata Gray, V, bracteosa var, brev- 
ibracteata Gray, V. canadensis (L.) Britton, V. hastata L,, V. 
hybrida Bicknell, V^ officinalis L., xV, deamii Moldenke, V. spu- 
ria L., V. haatata x urticae folia Pammel, and " Verbena hybrid". 
Rhoades records the common names "pidgeon grass", "holy-herb", 
"enchanter's plant", "European vervain", aad "Juno 's-t ears" for 
this plant, but these are all names applied to the European V, 
officinalis with which he mis identified his collection. Palmer 
(I9I6) cites his no. 601. The E. Hall s.n, [1361], distributed 
as "V. bracteosa x urticifolia " in the United States National 
Herbarium, is actually V. canadensis (L.) Britton, 

In all, 112 herbarium specimens, including the types of most 
of the names involved, and k movinted photographs have been exam- 
ined by me. 

Citations: ONTARIO: Lambton Co,: £. K. Dodge s,n, [Point Ed- 
ward, July 27, 1902] (Mi, Mi, Mi), GBOflGIA: County undetermined: 



312 PHYTOLOGIA Vol. 10, no. k 

A. W^ Chapman 69 [Mts. of Georgia] (W~1323060), s.n. [Northern 
Georgia] (W— 1323061) . ILLINOIS: Adams Co.: Seymour s.n. [Fall 
Creek, July 29, 1879] (Ur) . Cass Co.: Geyer s.n. [Beardstown, 
July I81i2] (Pr — cotype, T — cotype, T — cotype), s.n. [Beardstown, 
Aug. I81i2] (Dt— cotype). Coles Co.: Ahles & Gilpin 7U5$ (Ur) . 
Fulton Co.: J. Wolf s.n. [Canton, 1881] (W~$6221)| L^ Wolf s.n. 
[Canton, 187U] (Al) . Hancock Co.: S. B. Mead s.n. [Augusta, 181^2] 
(Pr), s.n. [A^^gusta, June iQhh] (C). Henderson Co.: H. N. Patter - 
son s.n. [vicinity of Oquavrica] (W— I32306U, W—132309li, W— 1323- 
137), 3.n. [July] (Al). Menard Co.: E. Hall s.n. [Athens, Aug. 
1866] (Ms). Monroe Co.: Winterringer 38lli (11—29272). Wabash 
Co.: Schneck s.n. [June 25, 1879] (Ur), s.n. [Ju3y 8, I88O] (Ur), 
s.n. [Mt. Carmel, June 15/88] (Ur) . Woodford Co.: V. H. Chase 
11)|73 (Ur) . County undetermined: F. Brendel s.n. [Illinois, 
1873] (W~71968l). INDIANA: Fulton Co.: £. C. Deam 39228 (Al, 
Dm). Jennings Co.: C_. R. Barnes 28 (N) . Johnson Co.: W. H^ 
Rhoad es s.n. [Edinberg] (Hs). Kosciusko Co.: C. C. Deam 55323 
(Dm) . Lagrange Co.; C. £. Deam 36661 (Dm). Lawrence Co.: C. C. 
Deam 17287 (Dm). Saint Joseph Co.: Nieuwland s.n. [St. Mary's, 
Oct. 192li] (W— 12JiU5lU) . IOWA: Black Hawk Co.: Carver s.n. [Ce- 
dar Falls, July 5, 1895] (lo— 22919). Clarke Co.: Pammel & Pam- 
mel s.n. [Osceola, Sept. 27, 192U] (lo— -1111682) . Decatur Co.: 
£. P. Anderson s.n. [July 23, 1903] (lo— 52109) . Emmet Co.: Wol- 
den 3.n. [EsthervLlle, Aug. 5, 1922] (lo— 105155), s.n. [Esther- 
ville, Jul. 12, 1927] (lo— I30U6I). Fremont Co.: A. S. Hitchcock 
s.n. [Hambxirg] (lo — 15300, Ka) , Hardin Co.: M. E. Jones s.n. 
[Iowa Falls, Aug. I876] (Po— 71002). Johnson Co.: A. S. Hitch- 
cock s.n. [Iowa City] (lo — 15301). Story Co.: Carver s.n. [Ames, 
July Hi, 1896] (lo— 15326)} L. Leonard s.n. [Collins, Aug. 8, 
1929] (lo— I3U217). KENTUCKY: Union Co.; Shacklette h99 (Ky) . 
County undetermined: Short s.n. [Barrens of Ky., I81i0] (Pr). WIS- 
CONSIN: Dane Co.: T. J. Hale s.n. [Madison] (Ws)j Lapham s.n. [M. 
Spear, 1858] (Ws). LaCrosse Co.: L. H. Pammel 3. n« [LaCrosse, 7- 
20-1887] (lo — 95llli). Lafayette Co.: Manning s.n. [Shulesbxirg, 
July 19, 1883] (N). Milwaukee Co.: £, S. Douglas s.n. [Wis.] 
(Je). Sauk Co.: T. £. Hale s.n. [Baraboo, 1861] (Ws, Ws) . KANSAS: 
Doniphan Co.: Agrelius , Hall , Love joy , & Maroney s.n. [8-18-19] 
(Lw). Pottawatomie Co.: Clothier s.n. [St. George, 7-li-96] (Ka)j 
A. S. Hitchcock 972 (Ka, N, W— 353769). Sedgewick Co.: M. A. 
Carlton 276 (Du— 9089O, W— 71932). Shawnee Co.: E. A. Popenoe 23 
(W— 56202), s.n. [Topeka, July 17, '79] (W— III9629) . MISSOURI: 
Jackson Co.: £. ^ Mackenzie s.n. [July 19, I896] (Dt, N), s.n. 
[Uttle Blue Tank, Aug. 2, I896] (Dt, N, Ur) . Jasper Co.: E, J. 
Palmer U025 (Ur) . Saint Louis Co.: Eggert s.n. [1| Aug. l87^(Cm), 



196U Moldenke, Monograph of Verbena 313 

s.n. [Eureka, Ju2y 28, 1879] (Vt), 3»n. [Glencoe, 28 July 1879] 
(Al, Cm, Cm, N), Saint Louis: Eggert 5318 (N); Engelmann s.n. 
[St. Louis, Sept. 18U1] (W— 71928), s.n. [St. Louis, July I81i2] 
(Au — 1228lli~ cotype, Dt—cotype, T — cotype), s.n. [St. Louis, 
I81i2] (Br—cotype), s.n. [St. Louis, Aug. 18U3] (T), s.n. [St. 
Louis, Aug. 18U5] (Dt), 3»n. [St. Louis, Aug. 18^9] (Br, T), s.n. 
[St. Louis] (F — photo of cotype, N — photo of cotype, S — cotype, 
S~cotype, Si — photo of cotype, Z — photo of cotype); Geyer s.n. 
[St. Louis, Sept. I81il] (Dt). ARKANSAS: Benton Co.: Plank s.n. 
(N) . NEBRASKA: Cass Co.: £. L. Sheldon s.n. [Weeping Water, Aug. 
17, 1898] (We); T. A. Williams 119 (W— 71931) . Lancaster Co.: J. 
L. Sheldon 3.n. [Lincoln, July 16, I898] (We). Otoe Co.: Thorn- 
ber s.n. [Nebr. City, Aug. I900] (Tu— 98878). Saunders Co.: Ryd- 
berg 153 (N), s.n. [Mead, June I89O] (C, ¥— 5l8l2]i) . OKLAHOMA: 
Lincoln Co.: J. W. Blankinship s.n. [Baker, 25 Aug. 1895] (Tf— 
313602). Woods Co.: G. W. Stevens 1703 (Du— 65779, N, Ok, Ok, 
St— 9270, St— 9279, Ur, W— 589635). WASHINGTON: Klickitat Co.: 
Sukadorf s.n. [July 9, I898] (N) . LOCALITY OF COLLECTION UNDE- 
TERMINED: Collector undesignated 27 (C), 103 (Q); Engelmann s.n. 
[Amer. bor.] (Br, S, Sg— 16096) . 

xVERBHIA PERTURBATA Moldenke, R6sum6 Suppl. $t 6—8, hyponym 
(1962), nom. nov. 

Synonymy: Glandularia parodil x mendocina Covas & Schnack, 
Reviat. Argent. Agron. 11: 96. 19Uii. 

Bibliography: Covas & Schnack, Reviat. Argent. Agron. U: 96. 
l9Ui; Moldenke, R6sum6 Suppl. 5: 6 — 8. 1962; Moldenke, Phytologia 
8: 396 (I962) and 10: 137. 196U. 

This is the natural hybrid between Verbena parodii (Covas & 
Schnack) Moldenke and V. mendocina R. A. Phil., with more or less 
intermediate characters. It is known only from the area in Men- 
doza where the ranges of the two parental species overlap. It is 
described by Covas & Schnack (19Ui) as follows: "Hemos hall ado en 
la localidad del tipo [i.e., the type locality of V. parodii ] v>na 
poblaci6n formada, iiiuy probablemente, por hfbridos (y formas der- 
ivadas de fistos) entre esta especie y Glandularia mendocina . . . .La 
poblaci6n hlbrida presenta una amplia gama de variaw;i6n que com- 
prende formas intermedias y formas vecinas a ambos padres; en al- 
gunas de estas fonnas hemos podido observar flcres con pequeflos 
l6bulos petaloides en la base del limbo de la corola, car&cter 
que nunca hemos observado anteriormente en el g6nero Glandularia ." 

Planta hybrida aspectu Verbena p)arodii et V. mendocina inter- 
media; limbo corollae interdum lobulis parvis petaloideis ad 
basin ornato. 

VERBENA PERUVIANA (L.) Britton in Morong, Brit ton, & Vail, Ann. 
N. T. Acad. Soi. 7j 197. I892. 
Synonymy: Lychnidaea, veronioae folio, flore coccineo Feuill,, 



31U PHYTOLOGIA Vol. 10, no. h 

Jotim. Obs. Phyg. CStes Orient. [3]: 36 — 37. 1725. Ijrchnldaea , 
veronlcae folio , flor6 coccineo Feuill., Journ, Obs, Phys, COtes 
Orient. [3]: pi. 25, [fig. 3]. 1725. Erinus penivianvis L., Sp. 
Pi., ed. 1, 1: 630. 1753. Lychnidea veronicae folio , flore coc - 
cineo Feuill. apud L., Sp. PI., ed. 1, 1: 63O, in syn. 1753. 
Verbena chamaedryfolia A. L. Juss,, Ann. Mus. Nat. Hist. Paris 7: 
73. 1806, Verbena veronicifolia J. Sm. in Rees, Cycl, 36: no, 28. 
I8I7. Verbena chamaedrifolia A. L. Juss. ex Stexid., Ncai. Bot,, 
ed. 1, 873. 1821 [not V. chamaedrifolia Briq., 190li], Verbena 
melindres Gill, ex Lindl. in Edirards, Bot. Reg. ll;j pi. 118U» 
1828 . Lychnidaea veronlcae folio flore coccineo Feuill . apud 
sweet, Brit. Fl. Gard., ser. 2, 1: pi. 9, in syn. 1829. Verbena 
coccinea Hort, ex Reider, Ann, Blumisterei 7: 288 — 289 & 298, 
pi. [2U]. 1831. Verbena melindroides Cham., Linnaea 7: 270 — 271. 
1832. Verbena melissoides Sweet ex Cham., Linnaea 7: 270, in 
syn. 1832, Verbena chamaedrifolia var. elfordiana Benth. in 
Maund & Henslow, Botanist 3: pi. 129, in part. 1839. Verbena 
picta Mamock, Floricult. Mag. $: 87. 181^0, Verbena veronicae- 
folia J, Sm. apud Steud., Nom. Bot., ed. 2, 2: 750, in syn. I8UI 
[not V. veronicaefolia Humb,, 1825, nor Humb. & Bonpl,, I8UI, nor 
Humb, & Kunth, I8U6, nor H.B.K., I8I8, nor Kunth, 18U7] . Verbena 
chamaedryoides Hofflngg., Preis. Verz. Orchid. 28, 18U2; Linnaea 
16: Litt, 282. I81i2, Verbena chamaedryfolia var, latifolia Aut. 
ex Hofftagg,, Preis. Verz. Orchid. 28, in syn. l8U2j Linnaea l6s 
Litt. 282, in syn. 18U2. Verbena chamaedrifolia <X melindres 
(Gill.) Schau. in A. DC., Prodr. 11: 537. I81i7. Verbena chamae - 
drifolia P melindroides (Cham.) Schau. in A. DC, Prodr. 11: 537. 
I8U7. Verbena chamaedryfolia CL melindres (Gill.) Schau. in 
Mart,, Fl. Bras. 9: 182. 1851. Verbena chamaedryfolia f melin- 
droides (Cham,) Schau. in Mart., Fl. Bras. 9: 182. 1851, Verbena 
chamaedrifolia L. ex Lorentz, Veg. Nordeste Proc. Entre Rlos, ed. 
1, 150. 1878. Verbena peruviana Kuntze ex Briq., Ann. Conserv. & 
Jard, Bot. Qenev. 7-8: 290, in syn. I90U. Verbena peruviana Druce 
ex Lindman, Vi och Vara Blomm. pi. U2. 1911 — 1913. Verbena peruv- 
iana (L.) Druce, Rep. Bot. Exch. Club Brit, Isles 1913 (3): U25. 
I91II. Verbena sanguinea Larrafl,, Escritos D. A, Larraflaga 2: 9. 
1923. Verbena chamaedryfolia elfordiana Maund ex Stapf, Ind, 
Lond. 6: U29» 1931. Glandularia peruviana (L.) Small, Man. South- 
east, Fl. 1139 & 1508. 1933. Verbena peruviana Juss. ex Gallinal, 
Bergalli, Campal, Aragone, & Rosengurtt, Stud. Nat. Mead. Urug. 
81. 1938. Verbena chamaedrifolia var. melindres (Gill.) Schau. 
ex Moldenke, Prelim. Alph. List Invalid Names 56, in syn. 19 UO. 
Verbena chamaedrifolia var. melindroides (Cham.) Schau. ex Molden- 
ke, Prelim. Alph. List Invalid Names 56, in syn. I9U0. Verbena 



1961i Moldenke, Monograph of Verbena 31$ 

penivlana R. & P. ex Moldenke, Lilloa 10: 385, in syn. 19hht 
Verbena chamoedrlfolla var. mellndres Schau. ex Augusto, Fl. Rio 
Grande do Sul 232, in syn, I9li6. Verbena chamaedrifolia var, 
mellndres Jiiss. ex Daniel, Verb, Cent. Antioq, 2, 19U7. Verbena 
chamaedrifolia var. melindroides Schau. ex Moldenke, Alph, List 
Invalid Names Suppl, 1: 23, in syn, 191; 7. Verbena c hamaedryf olia 
var. melindres (Gill.) Schau. ex Moldenke, Alph. List Invalid 
Names Suppl. 1: 23, in syn. 19U7. Verbena coccinea Waterer ex 
Moldenke, Alph. List Invalid Names Suppl. 1: 23, in syn. 19U7. 
Verbena peruviana (L,) Brlq, ex Moldenke, Alph. List Invalid 
Names Suppl. 1: 26, in syn, 19U7. Grandularia peruviana (L,) 
Small apud Martinez Crovetto & Piccinini, Revist. Invest, Agxdc. 
U: 77 & 225. 1951. Lychnidea veronicae folio Feuill, ex Molden- 
ke, R6sum6 318, in syn, 1959 • Verbena chamaedrifolia var, 
melindre Schau, ex Reitz, Sellowia 11: 13U, sphalm, 1959. Verbena 
chamaedryfolia var. elfordiana Maund ex Moldenke, R4sum6 362, in 
syn. 1959. Verbena melinderi Prager ex Moldenke, R6s\aii6 369, in 
syn, 1959. Verbena melindri Prager ex Moldenke, R6sum6 369, in 
syn. 1959. Verbena chamaedryfolia L. ex Moldenke, Resume Suppl. 
5: 7, in syn. I962. Verbena melindres var, major Martens ex Mol- 
denke, R6s\m6 Suppl. 5» 7, in syn. I962. Verbena melindri s Gill. 
ex Moldenke, R^sumfi Suppl. S'- 7, in syn^ 1962. Verbena chamae - 
drifolia Or ill. ex Maria, PI. Vail. Cochab. 38. 1962. Verbena 
chamaedryfolia f . camporum Osten ex Moldenke, R6sum6 Svtppl. 5s 7, 
in syn. 1962. Verbena peruviana L. ex Moldenke, RSsumfi Suppl. 6: 
11, in syn. I963. Verbena peruviana Michx. ex Moldenke, R^sum^ 
Suppl. 6: 11, in syn. I963. 

Bibliographer: Feuill,, Joum, Obs, Phys, C6tes Orient. [3]: 
36—37, pl. 25, [fig, 3]. 1725; L., Sp, PI., ed, 1: 630, 1753; L., 
Suppl. PI. 87. I78I; A. L. Juss., Ann. Mus. Nat, Hist, Paris 7: 
73. 1806; J, Sm. in Rees, Cycl. 36: no, 28, I8l7j Steud,, Nom, 
Hot,, ed. 1, 873. 1821; Spreng, in L,, Syat, Veg,, ed. 16, 2: 7U8. 
1825; Lindl. in Edwards, Hot. Reg. Hi: pl. II8U. 1828; Sweet, 
Brit, Flow. Gard. h [ser, 2, 1]: pl, 9. 1829; C Mcintosh, Flora 
& Pomona pl. 11, 1829; Hook., Bot. Misc. 1: I67. 1829; Lodd,, Bot, 
Cab, 16: pl, I51U. 1829; M. Roscoe, Fl. Illustr. Seasons pl. 31. 
1829— I83I; Sweet Hort. Brit., ed. 2. Ul8. 1830; Reider, Ann. 
Blumisterei 7: 288—289 & 298, pl. [2li]. I83I; Maund, Bot. Gard, 
U: 7U. 1831—1832; Joum. & Fl. Jard, 1832: 21, 1832; Cham,, Lin- 
naea 7: 270—271. 1832; Curtis, Bot. Mag. 51: pl. 3333. I83I1; 
Lindl. in Edwards, Bot. Reg. 21: pl. 17U8. 1835; Florists' Mag. 
1: U5. 1836; Meisn., Pl. Vase. Gen, 2: I98. 1839 J Maund & Hens low. 
Botanist 3: pl. 129. 1839; Mamock, Floricult. Mag, 5: 87, I8UO; 
Steud., Nom. Bot., ed. 2, 1: 58U (I8U0) and 2: 750 & 751. I8ia; 
Paxt,, Mag. Bot. 1: 173. iSUl; Hoffmgg., Preis. Vera. Orchid. 28. 
I81i2; Hoffingg. Linnaea 16: Litt. 282. 18U2; D. Dietr., Syn. Pl. 
3: 601—603. loU3; J. W. Loud., Ladies Fl.-Qard. Omam. Perenn. 



33.6 PHITOLOGIA Vol. 10, no. U 

2: pi. 91. iQhhi Bohn, Florist's Joiim. 5: Ul. ISUU; Walp,- Rep- 
ert, U: 28, l8U5j Spach, Hist, Nat. V6g. Phan. Atlas pi. 8U. 18U6; 
Schau. in A. DC., Prodr. 11: 537. 18U7; Schau. in Mart., Fl. Bras. 
9: 181—132. 1851| Bocq., Adansonia 2: 10$ & 107, pl. 11. 1861— 
1862i Bocq., R6v. Groupe Verb&iac. 27 & 105, pl. H. I862j Rdreil, 
Regno V^g. lU: pl. 16. 1870; Lorentz, Veg. Nordeste Prov. Entre 
Rlos, ed. 1, 1$0. 1878; Griseb., Abh, K. Gesell. Wisa. QOtting, 
2U: [Symb. Fl. Argent.] 275. 1879| Regel, Oartenfl. 28: 369—372. 
1879; Pickering, Chronolog. Hist. Pl. 1002 & 1220. 1879; Lorentz 
& Niederlein, Exped. R£o Negro 2 (Bot.): 263. 1881; J. Sa., Diet. 
Pop. Names Pl. U28. 1882; W. Robinson, Engl. Fl. Qard. pl. 262. 
1883; Lillo, Fl. TucunUbi 9U. 1888; Baill., Hist. Pl. 11: 78. 1891^ 
Baill., Diet, Bot. U: 256. 1892; Morong, Britton, & Vail, Ann. N. 
T. Acad. Sci. 7: 197. 1893; Jacks, in Hook, f , & Jacks., Ind, Kew. 
1 (2): 877. 1893; Bois, Diet, Hort. II89. 1893—1899; Briq. in 
Engl. St Prantl, Nat. Pflanzenfam. h (3a): lli7, fig. 57 A— D & G. 
I89U; L. H. Bailey in A. Gray, Field Forest & Gard. Bot. 3Ul. 1895; 
Jacks, in Hook. f. & Jacks., Ind. Kew. 2s 1178 & 1179. 1395; 
Kuntze, Rev. Gen. 3 (2): 257. I898; L. H. Bailey, Cycl, A»er. 
Hort., ed. 1, 6: 1912, pl. 26U9. 1902; Chod., Bull. Herb. Boiss., 
s6r. 2, 2: 8I8. 1902; Chod., Plant. Hassler. 9: 197. 1902; Uaclos- 
kie in ¥. B. Scott, Rep. Princeton Univ. Exped. Patag. 8 (2): 68I4. 
1905; Rev. G6n. Bot. 17: 299. 1905; L. H. Bailey, Cycl. Amer. 
Hort., ed. U, 6: 1912, pl. 26U9. 1906; Hassler, Flor. Pile. 101. 
1909; Hieken, Chlor. Plat. Argent. I96. 1910; P. Henderson, Handb. 
PL, new ed., U75. 1910; Wettstein, Handb. Syst. Bot., ed. 2, 739. 
1911 ; Lindman, Vi oeh Vara Blonna. pl. U2. 1911—1913; Druce, Rep. 
Bot. Exch. Club Brit. Isles 1913 (3): 1*25. 1911i; Garden 79: 510 
(1915) and 80: 355. 1916; Herzog, Meded. Rijksherb. Leiden 29: U3. 
1916; L. H. Bailey, Stand. Cycl. Hort. 6: 3hk$, pl. 3910. 1917; 
C. L, Pollard in Webster's New Intemat. Diet. Eng. Lang. 2275. 
1917; Alvarez, Fl. Santiago del Est. I06. 1919 J Prain, Ind. Kew. 
Suppl. 5: 269. I92I; Molfino, Physis Si 22 & 285. 1921; Larrafi., 
Escritos D, A. Larraflega 2: 9. 1923; Britton & P. Wils., Scient. 
Surv. Porto Rico & Virg. Isls. 6: I38. 1925; Hegi, Illustr. Fl, 
Mittel-Eur. 5 (3): 2239--22UO. 1927; Larrafi., Atlas Bot, pl. U2, 
1927; Herter, Anal. Mus. Hist. Nat. Montevid., ser. 2, 2: la8— 
Ia9. 1928; Herter, Florula 105 & cover plate. 1930; Seckt, Rev, 
Univ. Nac. Cordoba 17: 90—91. 1930; Stapf Ind. Lend. 6: U29— 
U30. 19 31; Herter, Estud. Bot. Reg. Urug. 8b: 170 & 201. 1933; J. 
K. Small, Man. Southeast. Fl. 1138, 1139, & 1508. 1933; Junell, 
Symb. Bot. Upsal. U: 11, 19, 171, & 210, fig. 16. 193U; Parodi, 
Rev. Argent. Agron, 1: 202. 193ii; T. Meyer, Memor. Comis. Invest. 
Lang. Corresp. 193U: 15U. 193Uj L. H. Bailey, Cat. Florists 
Handl. Verbenac., mss. 1935; L. H. & E. Z. Bailey, Hortus, new 
rev. ed., 632. 1935; Herter, Revlst. Sudam. Bot. U: 186—187. 
1937; Florists Exch. 89 (3): U5. 1937; Gard. Chron., ser. 3, 101: 
U23. 1937; Troncoso, Darwiniana 3: 53 & 5U. 1937; Moldenke, Cvdt, 
Pl. 35. 1938; G. Grimm, N. Y. Herald Trib,, April 2k. 1938; Gal- 
linal, Bergalli, Campal, Aragone, & Rosengurtt, Stud. Nat. Mead. 
Urug. 81. 1938; A. W. Hill, Ind. Kew. Suppl. 9: 12U. 1938; Mol- 
denke, Alph. List Common Names 6, 8, 12, 21, 26, & 32. 1939; Mol- 



1961i Moldenke, Monograph of Verbena 317 

denke, Annot. k Claasif . List 108. I939j Fedde, Bot. Jahresber. 
59 (2): Ia7. 1939; N. Y. Times, March 12, 1939 1 Herter, Beih. Bot. 
Centralbl. 59s 275. 1939: Beale, Chron, Bot. 5: 517. 1939| Beale, 
Journ. Genet. UO: 338, 3U0, 3li8, & 35U~356. l9U0j Moldenke, 
Suppl, List Common names 8 & Hi. 19 UO; Parodi, Darwiniana It: 55 • 
19hO', Moldenke, Prelim. Alph. List Invalid Names 26, U5, & 55~57. 
19U0; Moldenke, PhTtologia 1: U80 (19U0) and 1: 511. 19U1; Mol- 
denke, Suppl. List Invalid Names 9. 19U1} Balls, Gard. Chron. 
Amer. U5t 36. 19U1; Ragonese, Darwiniana $: U13. 19U1; Questel, 
Fl. Isl, St.-Barth. 179. 19Ul: Moldenke, Known Geogr. Distrib, 
Verbenac, [ed. 1], 39— Ul, hh, 7U, & 102. 19U2i Moldenke, Alph. 
List Invalid Names 2U, 25, I46. Ii8, U9, & 51. 19ll2; Schnack, Anal. 
Inst. FitotSc. Sta. Catalina U: 17, 20, & 21. 19U2; G. Grijnm, N. 
Y. Herald Trib,, January 31. 19U3j Rosengurtt, Estud. Prad. Nat, 
Drug. 3: 236. 19U3; A. R. Schultz, Introd. Estud. Bot. Sistam., 
ed. 2. U83 & 562, fig. 178. 19U3j Schnack & Covas, Darwiniana 6: 
U70, Li72, U73, & U76. 19hh; Descole, Gen. & Sp. PI. Argent. 2: 
Icon. Pi. Argent, pi. 165. 19hh; Moldenke, Lilloa 10: 385. 19UUj 
RosengiiTtt, Estud. Prad. Nat. Urug. U: 8. 19kh; Moldenke, Phyto- 
logia 2: 116. 19U5j P. I. Acufla, Cat. Fl. Catamarq. 30, l9U5j 
Cabrera, DAGI Pub. Tec. 3t 27, 38, U5, & 75. 19U5j Schnack & Co- 
vas, Revist. Argent. Agron. 12: 222—228, fig. 1 A, 2, & 3 A— C, 
pi. 11 & 12 B, C, & F. 19U5; Moldenke, Bot. Gaz. IO6: I63. l9U5i 
Moldenke, Torreya k$i 90. 19U5; Schnack & Covas, Darwiniana 7: 
71—75, pl. 3 B & C, & pi. h A. 19U5} Covas & Schnack, Darwinia- 
na 7s 86, 19U5} Darlington & Janaki Ammal, Chromosome Atlas 270, 
19U5; Rosengurtt, Estud. Prad. Nat. Urug. 5: 395. 19U6j Augusto, 
Fl. Rio Grande do Svd 209 & 211, fig. 97. I9I46J Schnack & Covas, 
Bol. Soc. Argent. Bot. 1: 282, fig. lb. I9U6: Moldenke, Alph. 
Ust Cit. 1: 5, 19, 28, 30, U7, 52, 62, 72, 83, 8U, 95, 120 lli5, 
197, 200, 202, 20U, 218, 219, 230, 2U7, 251, 263, & 285. 19u6j 
Moldenke, Alph. List Invalid Names Suppl. 1; 23 & 26. 19U7j 
Hodge, Revist. Fac. Nat. Agron, 7: 313. 19U7i Lorentz, Veg. Nord- 
este Prov. Entre Rios, ed. 2, 150. 19U7; Daniel, Verb. Cent, An- 
tioq. 2. 19U7i Moldenke, Phytologia 2: 3U8 (19li7) and 2: li78 & 
U82. I9U8; Moldenke, Castanea 13: 118 & 121. I9U8; Moldenke. 
Alph. List Cit. 2: 356—358, 360, 36U, 365, 367—369, 371, hh2, 
U58, U81, 532, 533, 551, 561, 580, 585, & 6Ui (19U8), 3: 672, 
673, 687, 696, 697, 702, 703, 7U5, 765, 767, 768, 770, 771, 780, 
781, 826, 837, 8U8, 859, 861, 866, 868, 87li, 889, 89O, 905, 913. 
920, 922, 923, & 963 (19U9), and h: 1010, 1075, I08I, 1082, II5I4, 
1162, 1163, 1178, II9U, 1208, 1210, 1231, 1233, 12U9, & 1302. 
19U9; Cabrera, Lilloa 20: 175 & 315, cuadro I. 19U9j Moldenke, 
Known Geogr. Distrib, Verbenac, [ed. 2], 7ii, 9U. 98—100, IO6, 
16U, & 199. 19U9; Moldenke, Am. Wild Fls. 292 & U50. 19U9; E. L. 
Palmer, Handb. Nat. Hist. 297 & 663. 19U9; H. N. & A. L. Molden- 
ke, Anal. Inst. Biol. Mex. 20: Hi, 19ii9} Moldenke, Phytologia 3: 
136. 19ii9; Martinez Crovetto & Piccinini, Revist, Invest. Agric. 
U: 180 & 225. I95O; Politi, The Garden 2 (h): 22. 1950; N. Y. 
Bot. Gard. Seed Exch. List 1951: U. 1950; Moldenke, Phytologia 3: 
305, 306, & 378 (I950) and 3: U67. 1951; Moldenke in Chittenden, 
Roy. Hort. Soc. Diet. Gard. U: 2208, 2209, 2211, & 2212. 1951; 



318 PHYTOLOGIA Vol. 10, no. U 

Moldenke, Phytologia Us 67 (19^2) and U: U5l. 19^3; Schnack & 
Solbrig, Revist, Fac. Agron. La Plata 29: 2^5 — 266, fig. 1 C — E, 
fig. 3 A & B, & fig. U C & H. 1953; Roig, Dice. Bot. 1; 909 
(1953) and 2: lllU. 1953; Rambo, Sellowia 6: 60, Qh, & 153. 195Ui 
Lombardo, Invent. Pi. Cult. Montev. 222, 232, & 262. 195U; Molden- 
ke, Inform. Mold. Set h& Spec. [U] . 195U} Moldenke, Journ. Calif. 
Hort. Soc. 15: 80. 195U; Moldenke, Phytologia 5: 132. 1955; Mol- 
denke, Biol. Abstr, 30: 1093. 1956: Rambo, Sellowia 7: 260. 1956; 
Moldenke in Humbert, Fl. Madag. 17U: 8. 1956; Moldenke, Am. Midi. 
Nat. 59: 3hh, 350, 357, 361, & 370. 1958; Mattoon, PI. Buyers 
Guide, ed. 6, 287 & 288. 1958; Reitz, Sellowia 11: 57 & 13U. 1959; 
Moldenke, Pl^ologia 7: 85. 1959; Moldenke, R^sumS 16, 85, HO, 
115, 118, 120, 128, 22U, 285, 296, 318. 361, 362, 369, 370, 372, 
373, 378, & U73. 1959; Moldenke, R^suml Suppl. 1: 23 (1959) and 
2: 9, 12, & 13. I96O; T, H. EvBrett, New Illustr. Encycl. Gard. 
13: 2U0I4 & 2I1O5, pi. 13-11. I960; Angely, Fl. Paran. 16: 79 
(i960) and 17: u6. I96I; Reitz, Sellowia 13: HO. I96I; Moldenke, 
Phytologia 8: 120, 121, & 123 (1961) and 8: 257, 280, 316, 378, 
Ul8, la9, & U35. 1962; Maria, PI. Vail. Cochab. 38. 1962; Mol- 
denke, R5sum6 Suppl. 3: 15 & 33 (1962), U: 17 (1962), S'- 3 Sc 6—8 
(I96?), 6: 7, 8, & 11 (I963), and 7: 9. 1963; Moldenke, Phytolo- 
gia 8: U89 (1963), 9: UO, 67, 69, 72, 130, 178, 215, 303—305, 
307—309, 315, 330, 333, 33U, 336, 352, 362, 365—367, 395, 397, 
& 399 (I963), and 10: 113, 127, 128, 132, & 133. 196U; J. A. 
Clark, Index New Spp. Gray Herb, issue 117 & 123. n.d. 

Illustrations: Feuill., Journ. Obs. Phys. COtes Orient. [3]s 
pi. 25, [fig. 3]. 1725; lindl. in Edwards, Bot. Rag. 11^: pi, 
118U (in color). 1828; Sweet, Brit, Fl. Gard, k [ser, 2, 1]: pi, 
9 (in color), 1829; Lodd,, Bot, Cab, 16: pi. I5lli (in color). 
1829; C. Mcintosh, Flora & Pomona pi, 11 (in color), 1829; M, 
Roscoe, Fl. Illustr. Seasons pi. 31 (in color). 1829 — 1831; Rei- 
der, Ann. Blumisterei 7: 298, pi, [2U] (in color). 1831; Maund, 
Bot. Gard. U: 7U (in color). I831~l832; Journ. & Fl. Jard. 1832: 
21 (in color). 1832; Curtis, Bot, Mag. 61: pi. 3333 (in color). 
18 3U; Florists' Mag. 1: 1;5 (in color). 1836; Maund & Henslow, 
Botanist 3: pl. 129, in part (in color). 1839; Mamock, Floricult. 
Mag. 5: 87. I8I4O; Paxb., Mag. Bot. 1: 173 (in color). iSUl: J. W, 
Loud., Ladies Fl.-Gard. Omam. Perenn. 2: pl. 91 (in color). 
18hh; Spach, Hist. Nat. V5g. Phan. Atlas pl. 8U (in color). 181*6; 
Bocq.. Adansonia 2: pl. 11. 1862; Bocq., r5v. Groupe Verbfinac. pl, 
11, 1863; R^veil, Regne V6g. 11*: pl. 16 (in color). I87O; Regel, 
Gartenfl. 28: 372. 1879; W. Robinson, Engl. Fl. Gard. pl. 262, 
1883; Bail!., Hist. Pl. 11: 78. I89I; Bois, Diet. Hort. II89 (in 
color), I893— 1899; Briq. in Engl. & Prantl, Nat, Pflanzenfam, h 
(3a): 11*7, fig. 57 A— D & G, I89U; L. H. Bailey, Cycl. Amer, Hort., 
ed, 1, 6: 1912, pl. 261*9. 1902; Rev. G6n, Bot, 17: 299. 1905; L. 
H. Bailey, Cycl. Amer. Hort., ed, 1*, 6: 1912, pl. 261*9. 1906; 
Wettstein, Handb. Syst, Bot,, ed. 2, 739. 1911; Lindman, Vi och 
Vara Blomm. pl. 1*2 (in color). 1911—1913; Garden 79: 5l0 (1915) 
and 80: 355. 1916; L. H. Bailey, Stand. Cycl. Hort, 6: 3UU5, pl. 
3910. 1917; Larrafi., Atlas Bot. pl. 1*2 (in color). 1927; Herter, 
Florula cover plate (in color), 1930; Junell, Symb. Bot, Upsal. 



196U Moldenke, Monograph of Verbena 319 

1;: fig, 16. 193U; A. R. Schultz, Introd« Estud. Bot. Sistem., ed, 
2, fig. 178. 19li3i Descole, Gen. & Sp. PL. Argent. 2: Icon. Pi. 
Argent, pi. 16^ (in color) [as V, scrobiculata ] . 19kh} Schnack & 
Covas, Revist. Argent. Agron. 1^: 225, fig. 1 A, 226, fig. 2, 227, 
fig. 3 A— C, pi. 11 & 12 B, C, & F. 19hSi Augusto, Fl. Rio Grande 
do Sill 211, fig. 97. 19U6; Schnack & Covas, Bol. Soc. Argent. Bot. 
1: 282, fig. lb. 19li6; Moldenke in Chittenden, Roy. Hort. Soc. 
Diet. Gard. 6: 2212. 195lj T. H. Everett, New niustr. Encycl. 
Gard. 13: 2li05, pi. 13-11 (in color). I960. 

Low procumbent half-hardy perennial herb, with loose spreading 
habit} stems prostrate or creeping, rooting at the nodes, very 
slender or filiform, forked, to k$ cm. long, hiirboixs, the tips 
and branches ascending; leaves small, decussate-opposite, gray or 
slightly grayish, oblong or oblong-lanceolate to ovate, broadly 
cuneate at the base and narrowed into the extremely short petiole, 
crenate or doubly crenate-serrate to unequally incised-serrate or 
almost incised-dentate, strigose above, more or less short- 
hirtous especiEilly on the venation beneath; inflorescence spicate; 
spikes solitary, long- pedunculate, ascending, densely corymbose- 
flowered, flattened-capitate when young, to 6.5 cm. wide dxiring 
anthesis; bractlets less than half the length of the calyx, cili- 
ate-^nargined; flowers odorless, very showy; calyx canescent- 
hixtellous; corolla hypocrateriform, very showy, varying from i^d, 
brillismt-red, cinnabar-red, fire-red, spectrum-red, or dark-red 
to scarlet, vivid-scarlet, brilliant-scarlet, red-scarlet, or 
purple-cinnabar, its tube scmewhat surpassing the calyx, the limb 
to 1 cm. wide; chromoscme number: 2n ■ 10, 

Linnaeus (1753) based his Erinus peruvianus on Feuill6e*s 
Lychnidaea , veronicae folio , flore coccineo, of which the type 
was collected by Louis Feuill6e in Paraguay (not in Pei-u as im- 
plied by Linnaeus and as stated by Schauer in 1851), who says of 
it "Je trouvai cette plante dans les campagnes qui sont sur le 
bord septentrional de la riviere de la Plata, dans le Paraguay." 
There seems to be no actual specimen in the Linneaji Herbarium, 
and Linnaeus seems simply to have supplied the brief descriptive 
sentence "ERINUS foliis lanceolato-ovatis" . I therefore regard 
FeuiUfie's collection as the type of the species. 



BOOK REVIEIf 
Alma L. Moldenke 



"Flora of Missouri", by Julian A. Steyennark, 1808 pp., 2300 
illustr. Iowa State University Press, Ames, Iowa. 1963* 
|18,50 

This work is superb and monimiental. Seldom can an active sci- 



320 PHYTOLOGIA Vol. 10, no. k 

entist with considerable other interests and responsibilities — 
as has the author — manage to publish so accurate and thorough a 
work as this. Dr. Steyermark deserves kudos of congratulations j 
so do the fellow scientists who collaborated on some of this 
work; so does the National Science Foundation for wisely choosing 
to assist in the publication. 

Taxa to the total of 3351, comprising 799 genera, 2li38 species, 
517 subspecies and varieties, and 297 forms of wild Pteridophyta 
and Spermatophyta are treated, After a historical introduction 
and a general description of the flora and vegetation, there is a 
50-page key to families which is followed by family descriptions 
and keys to their genera and species with enough details so as 
not to require additional species descriptions . The first key 
has a built-in glossary which is an excellent time-saver. The 
following data are given for each species: scientific name with 
authorities, some ccanraonly encountered synonyms, common name or 
names, period of flowering and fruiting, habitat within the 
state, pertinent ecological data, nature of local variations and 
intergradations . The following data are added where signifi- 
cant: medicinal usage, food and forage value, industrial ^plica- 
tions here and abroad. This compendium is a culmination of 30 
years of field work and associated taxonomic studies. 

There are many other special features: county distribution 
maps with about 3 for each page of text, over 2000 clear line 
drawings conveniently placed near the telated text and often sen- 
sibly taken from fine earlier works, a keyed map that makes the 
locating of counties easy, an alternate key for the tracing of 
completely sterile ferns, a list with location and dimensions of 
the ••champion" trees of the state, two-columned format in the 
text and three-columned format in the index to allow for most ef- 
ficient use of printing space on the paper, abbreviations to in- 
dicate differences in the author's choice of accepted names over 
those in the widely used Gray's Manual, the new Bntton & Brown 
and the Annotated Catalogue of the Flowering Plants of Missouri. 

The author claims that he uses the system of repeating the 
specific epithet for the typical variety of a species to achieve 
precision as do an increasing number of modem taxonomic botan- 
ists and zoologists. But no precision need be lost if the concept 
is inferred and the nonenclature left uncluttered; on page 1599 
we find Achillea Millefolium L. subsp, Mniefollum var. Millefo- . 
lium f . Millef oli\mi . 

The fact that Dr. Steyermark has been the guiding impetus in 
the founding of several nature preserves for education in and 
appreciation of natural beauty indicates that he has the heart of 
the true naturalist as well as the fine mind of the skilled sci- 
entist. Many more states in our country need to have svch 
SteyeiraarkLan work done for them. 



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i^^: 



PHYTOLOGIA 

Designed to expedite botanical publication 



^^. 



^^. 



*fi7V 



Vol. 10 July, 1964 No. 5 



CONTENTS 



REED, C. ¥., A flora of the chrome and manganese ore piles at 
Canton, in the Port of Baltimore, Maryland and at Newport 
News, Virginia, with descriptions of genera and species 
new to the flora of the eastern United States 321 

MOLDENKE, H. N., Materials toward a monograph of the genus 

Verbena. XXI 406 



Published by Harold N. Moldenke and Alma L. Moldenke 

15 Glenbrook Avenue 
Yonkers 5, New York, U.S.A. 

Price of this number, $1.25; per volume, $5.75, in advance 



A FLCRA OF THE CHROME AND MANGANESE CRE PILES AT CANTON 
IN THE PCRT OF BALTIMORE, MARYLAND AND AT * 

NE^VPQRT NEWS, VIRGINIA, WITH DESCRIPTIONS 
OF GENERA AND SPECIES NEW TO THE FLORA OF 
EASTERN UNITED STATES 

Clyde F. Reed 

Seaports and industrial areas along our seacoasts have always 
been welcon« havens for the plant waifs that reach our shores by 

yards and just docks all afforded good conditions for foreign 

lllT^^ f\li^^..l^^ ^^""^ ^" ^ "^" ^°"«- ^^ orHj need! to 
check the Eighth Edition of Gray's Manual for the species natu- 
ralized to be well aware of the large number of weeds which are 
coirmon with us today, which were introduced usually innocently 
50 to 100 years ago, '' 

M«J?® I'^^'i °^ Newport News, Virginia and the Port of Baltimore, 
Maryland, have been the sites for ships for a long time. Recent^ 
ly both ports have been the unloading stations for ores of various 
kinds, as well as for coal, oyster-shells, gypsum, and oil. With 
the huge steel mills at Sparrows Point not far from the Port of 
Baltimore, Canton soon became the unloading ground for chrome ore 
manganese ore and even iron ore. For at least twenty years Canton* 
has been a temporary unloading ground for ores on the move. 

As early as the 1890- s ballast was duirped at Canton and other 
areas along the Port of Baltimore. Dr. C.C.Plitt of Baltimore 
r no!^^ collections of the adventive weeds in Canton from 1900 
to 1906. Some of these plants have been recorded in the "Extracts 
from the Journal of C.C.Plitt".* Most of the herbarium specimens 
mentioned here and many others collected ly Dr. Plitt at Canton 
are in the Reed Herbarium. 

Other areas in the Port of Baltimore where ballast has been 
dmiped in the past are Port Covington, Dundalk, Curtis Pay. 
Brooklyn, Westport, Locust Point, and Fairfield. The present study- 
is minly concerned with the plants that have been found growing 
on the ore piles in the Canton area, although some species from 
the other areas will be included. 

The areas studied in Virginia have been in the Port of Newport 
News, the coal piles and wastes immediately adjacent to the docks 
and the chrome, manganese, and iron ore piles about seven miles 
inland along the C.& 0. Railroad, where an ore refining companv 
has stocked huge piles of these ores. 

* Fessenden G.R. Plants of Baltimore Ballast Grounds. Wild 
Flower. 32(3): Uh-ii7. 19^6. 

321 



322 



PHITOLOGIA 



Vol. 10, no. 5 





Views of chrome ore piles at Canton, Maryland 



I96h Reed, Flora of chrome & manganese ore piles 323 

Nearly ?^0 species of flowering plants have been collected on 
these mineral ore piles in Maryland and Virginia, about 80 per cent 
of >^ich are new to either the Ei^th Edition of (i-ay's Manual or 
Gleason's Illustrated Flora. Mai^ of these species have been col- 
lected in these areas year after year since 1953 when this study 
started. As these ores lie in piles awaiting to be shipped else- 
>rfiere, the seeds germinate and the plants most often reach matur- 
ity in our region and set seed. Some of these plants have continued 
to seed themselves here and may be found every year on undisturbed 
piles. Some have been carried various distances and already have 
become pests. 

Those genera and species which are entirely new to our flora will 
be listed and described below. Other species which are extensions 
of known ranges will be mentioned. Finally, other Icnown species which 
are found in these regions will be merely mentioned. A few species 
found in other areas in Maryland and Virginia will mentioned also. 
All specimens are in the Reed Herbarium, unless otherwise noted. 
Some species have been deposited in the herbarium of the United 
States National Museum in Washington, D.C. 

ACANTHACEAE 

1. Andrographis eehtaoides Nees. fropical India, in the drier dis- 
tricts from the Punjab and Chota Nagpore to Ceylon. Virginia: New- 
port News, on chrome ore piles. Nov. 1959. Reed 15902 (US). Det. "by 
E.C.Leonard. Description of genus and species, see Reed, Castanea, 
26: 128. 1961. 

AIZOACEAE 

2. Mollueo gracillima Ands. Maryland: Canton. Oct. 195U. Reed 35232; 
Oct. 19587 Reed U13I49. Native of Galapagos Isl. (Many sheits"in US); 
also from Uganda, Africa, marked (on US sheets), "Introduced, road- 
sides, Mmmer 2712". This species resembles M. verticil lata , to 
which it has been reduced by Hook. fil. end Jackson (Index Kew, 2: 
253), from which it differs hy having smaller flowers, a shorter 
capsule relative to the blunter sepals, and in its minute black, 
tessellated, not ribbed seeds. 

3. Mollugo nudicaulis Lam. Maryland: Canton. July 1958. Reed h0892. 
Native of the Old V/orld Tropics; also in the West Indie s.~irglabr- 
ous herb, 7-30 cm. high; leaves all basal, in an erect-spreading 
tuft, spatulate or obovate, 1.5-5 cm. long, 6-16 mm. broad, rounded 
at the apex, tapering at the base into the petiole; cymes long- 
peduncled, the peduncles slender, somewhat angled and grooved; 
flowers slender-pedicelled; calyx-lobes elliptic, about 2 mm. long, 
0.8-1 rem. broad; capsule ellipsoid, 2-2.5 mm. long; seeds reni- 
form, 0.5-0.6 mm. long, black, granulate. 

I4. milago verticillata L. Maryland; Canton, a common weed on the 
chrome ore piles. 

5. Trlanthema portulacastrum L. Maryland: Canton, on chrome ore 
piles. Sept. 1933. Reed3?7?2. 



32U PHYTOLOGIA Vol. 10, no. 5 

ALISMTACEAE 
6, Sagittaria latifolia Willd. Maryland: Canton, in ditches near 
the ojrster shell piles. July 19^8. Reed li0879. 

AMMIANTHACEAE 
The following new species and new genera of this family which 
have been found in these regions have been discussed by the author 
in Castanea 26: 123-127. 19'6l. Records and specimens are cited there 

7. Achyranthes aspera L. 

8. Alternanthera ficoidea (L.) R.Br. 

9. Alternanthera repens (L.) Kuntze 

10. Alternanthera sessilis R.Br. 

11. Guilleminea densa (Vfilld.) Moq. 

12. Hermbstaedtia elegans Moq, 

13. Amaranthus palmeri Wats, 
lli. Amaranthus cruentus L. 

15. Celosia argentea L. 

16. Froelichia gracilis (Hook.) Moq. Also see Reed, Castanea , 

27: 59-6l7l962. Additional records: Maryland, Canton, 
on manganese ore pUes, Newkirk Street. Aug. 9, 1961. 
Reed 521L5; June 10, 1963. Reed 62h07j Aug. 20, 1963. 
Reed 65U10 (US). 

17. Gomphrena celosioides Mart, 

18. Gomphrena dispersa Standi. From ore piles, Newport News, 

19. Gomphrena globosa L. /also, 

20. Alteranthera achyrantha R.Br. Newport News only, 

21. Amaranthus spinosus L, 

22. Amaranthus albus L. 

23. Amaranthus graecizans L. 
2U. Amaranthus retroflexus L. 

25. Amaranthus deflexus L. 

26. Amaranthus lividua L. Adventive from the Tropics; 

from Mass. and New Tork south to Maryland and the Dis- 
trict of Columbia, In Maryland this has become a genu- 
ine pesty weed, especially in spinach fields. Maryland: 
Baltimore Co., from Sparrows Point to Essex, Chase, 
Loreley and White Marsh, and then to the Harford Co, 
line. In the District of Columbia the author collected 
it in the outdoor flower beds near the foot of the 
Capitol, Sept. 2?, I96I. Reed 53271 (US). 

ANACARDIACEAE 

27. Rhus copalllP<> L. Maryland: Canton, common in thickets and 
alongTailroad tracks, Oct, I, 19^7, Reed 39^58, 

28, Rhus radicans L, Maryland: Canton, Oct. h, 1957. (obs.); 
AprirrU, 1958. Reed 10703, Common in thickets and in wastes on 
ground. 



196U Reed, Flora of chrome & manganese ore piles 32^ 

29. Rhus ^hlna L. Maryland: Canton, along Newklrk St.. several 
very large trees, about 25 ft. tall, with trunks h-6 in! in 
diameter . 

APOCTNACEAE 

30. Apocynum cannabinum L. Maryland: Canton, a common weed through- 
out the area. Sept. 2?, 1953. Reed 32725. «irougn 

ARISTOLOCHIACEAE 

31. Aristolochia clematitis L. Introduced from Europe; local from 
S^^'Tr n^,^^.°n^? Maryland. Maryland: Canton. May 22, 1900, 
May 16 1903; July 6, 1906. All collected by C.C.Plitt. Colon? 
entirely destroyed after the last date). ^ 

ASCLEPIDACEAE 

32. AscljE^ iricarnata L. var. pulchra (Ehrh.) Pers. Common on 
wastes between the chrome ore piles. July 30, 1959. Reed h38h2. 
Canton, Maryland. 

33. Asclepias syriaca L. Common throughout the area. Canton, 
wastes along Newklrk St. Aug. 18, 1959. Reed 11^388. 

BERBER IDA CEAE 
31i. Podophyllum peltatum L. In several patches along ditches in 
i^anton near the gypsum piles along Newklrk St. April 2li 1958 
Reed It0707. ^ * -^7^^, 

BCRAGIKACEAE 
Several new species in this family have been found in the 
Canton area. Most of these have survived the winters and have been 
found every year since 1953. Some have persisted since 1901, when 
they were collected by C.C.Plitt. 

35. Anchusa italica Retz. Native of the Mediterranean region and 
the Near East; N. Africa, France, Corsica, Europe meridioSle et 
So^S!* n^^^^^^"^; Canton, Newklrk St., chrome ore piles. June 
29, 1956. Reed 38821. Plant 30-80 cm. tilll, branched; lea;esX 
long, lanceolate to ova to-lanceolate, entire, the lower ones 
petiolate, the upper ones sessile; raceme paniculate; flowers 

fK»^ Z "Tv' ^^v °^ "^y^ ^^'^^^ "®^ ^« ^se, acute, shorter 
than the tube of the corolla, spreading in fruit; corolla IO-I3 mm. 
long, in a tube; nuUets oblong, wrinkled, granulate. 

36. Echium italicum L. Native of Turkestan, Asia Minor, and the 
Mediterranean region. Maryland: Canton, Newklrk St.. on chrome 
ore piles. June 29, 1955. Reed 388I8. Plant biennial, 3O-8O cm. 
tall stem robust, branched to form a pyramidal panicle; leaves 
hispid, the radical ones farming a rosette, elliptical-lanceolate, 
one-nerved, the cauline leaves sessile; flowers red or rose, ouit^ 
small; calyx very hispid, the lobes lanceolate; corolla lO-U "mm. 
long, the tube long, the limbs spreading and quite regular; the 



326 PHTTOLOGIA Vol. 10, no. 5 

stamens as long as the nearly regular corolla; flowers in lateral 
spikes; nutlets strongly tuberculate, sli^tly ridged. 

37. Echium vulgare L. Maryland: Canton, along Clinton St. in 
wastes. Oct. l8, 19^7. Reed 39iil6. Common in between the chrome ore 
piles. 

38. Heliotr opium cT3rassavicum L. Washington south to southern 
California and Mexico, east to Delaware and Florida; casually on 
ballast northward; widely distributed on all continents, Maryland: 
Canton, Newkirk St., chromB ore piles, June 22, 1951. Reed 33151. 

39. Heliotr opium europeum L, Naturalized from Europe; from Florida 
and Alabama north to New Jersey, and locally to Massachusetts. 
Maryland: common on ballast and on the chrome ore piles of Canton 
since 1901. Ballast bounds. June 19, 1901. C.C.Plitt (Not mention- 
ed in Plitt's Journal for this date, ace. to Fessenden, see Wild 
Flower 32: U5. 1956), in Reed Herb. No. 26612; same locality. June 
18, 1905. Plitt, in Reed Herb. No. 26611; Canton, on chrome ore 
piles near Newkirk St. Sept. 27, 1953. Reed 32673. 32812 and 32668; 
Oct. 12, 1953. Reed 32883; June 22, 195irrReed 33Uh9; July 22,195U. 
Reed 3h032; SeptTT, 1951. Reed 3^393; Sept. 30, 1955. Reed 382hO; 
jSne 29, 1956. Reed 38825; Oct. 17, 1956. Reed 382U3; Sept. 20, 
1957. Reed 39317; Clinton St. on wastes. Oct. U, 1957. Reed 39592; 
on manganese piles, Newkirk St. July 15, I960. Reed 1673 «; on 
pumice piles (from Italy). Aug. 20, I963. Reed 651i2U (US). 

UO, Heliotropium indicum L. Widely distributed in the Tropics of 
both hemispheres, probably of American origin. Maryland: Canton, 
Ne^ricirk St., on chrome ore piles. July 17, 1958. Reed la008 and 
li0889; Sept. 21^, 1958. Reed la238; July 9, 1959 (frequent). Reed 
U36I8. 

la. Heliotropium procumbens Mill. Native of tropical and sub- 
tropical America, Costa Rica. Maryland: Canton, Sept. 19, 1958. 
Reed 11327. Annual, sparsely or densely appressed-pilose, usually 
grayish, the stems erect or decumbent, 10-50 cm. long, much- 
branched; leaves elliptic, obovate, or broadly oblanceolate, 1-U en 
long, obtuse or rounded at the apex, petiolate; flowers small, 
white, in slender, scorpoid, mostly geminate or ternate, bractless 
racemes; corolla 5 mm. long or less; fruit depressed- globose, 
li-lobate, strigose, the nutlets 0.5-1 mm. long. 

U2. Lithospermum arvense L. Naturalized from Europe; a common weed 
from Nova Scotia to British Columbia, south to Florida, Louisiana 
and California. Maryland: Canton, on wastes. April 2U, 1958. 
Reed U0699. Throughout Maryland in waste places and fields. 



196U Reed, Flora of chrome & manganese ore piles 327 

CANNABINACEAE 
h3» Cannabis satlva L, Adventive from Asiaj Quebec to British 
Columbia and southward, Maryland: Canton, May 1903, Plitt in Reed 
Herb,; Sept, 1905, Plitt , in Reed Herb, 

hh. Huimilus Japonica Sieb. & Zucc, Introduced and naturalized from 
Asia; New England to Michigan, south to Virginia and Missouri, 
Maryland: Canton, just south of Hi^landtown, Sept, 19^7, Reed 
39387; Canton, near chrome ore piles. Oct, 19?7, Reed 39hlT, — 

h^. Humulus lupulus L. Native from New Brunswick to Montana, south 
to New England, northern Pennsylvania, West Virginia, eastern Ken- 
tucky, Ohio, Indiana, Illinois, Missouri, Kansas and New Mexico; 
also introduced from Europe; Maryland and Delaware. Maryland: Can- 
ton, plentiful, April 1958. Reed h0692; Sept, 1958, Reed II316 (fr.), 

CAPPARIDACEAE 
I46, Cleome viscosa L, Naturalized from the Old World Tropics; Ber- 
muda and West Indies, Maryland: Canton, Newkirk St,, on chrome ore 
piles. July 17. 1958. Reed hlOlO; July 19, 1958. Reed I1O887; Sept. 
6, 1958. Reed Ia303; Sept. 19, 1958. Reed ia319; JS^ 30, 1959. 
Reed U383lu ^Virginia: Newport News, on chrome ore piles. Aug. 7. 
1959. Reed iUjOh9; Nov. 15, 1959. Reed U5887. An erect viscid- 
glandular annual, 3-9 dm. tall; leaves digitately 3- or 5-f olio- 
late; leaflets obovate, oblanceolate or elliptic, 10-17 cm. long, 
0.7-3 cm. broad, acute to acuminate, rarely obtuse at the apex, 
often inequilateral at the base, the margin gLandular-ciliate; 
flowers solitary in the axils of the upper leaves; sepals otlong- 
lanceolate or lanceolate, 6*5-8 mm. long; petals yellow, obovate, 
about 1 cm. long; stamens 12-20, free; capsule cylindric, 5.5-10 
cm. long, li-J4.5 mm. thick, tapering to the apex, striate, viscid- 
glandular; seeds about 1.8 mm, in diameter, flattened, trans- 
versely ridged. 



328 



PHTTOLOQIA Vol. 10, no. 5 



CAFRIFOLIACEAE 
h7« Sambucvis canadensis L, Rather common in wet ditches in Canton, 
forming thickets. Oct. h, 19^7. Reed 39^79. 

US. Viburnum prunifoliim L, Forming fencerows in wastes along 
Clinton St., Canton. Oct. h, 19^7. Reed 395^3. 

h9. Lonicera japonica Thunb. Naturalized from Asia. Canton, common 
throughout the wastes, forming thickets over trees and bushes. 
Some patches appear to be var. chinensis (P.W.Wats.) Baker, with 
the branchlets and leaves purple and glabrous, and with a carmine 
corolla. 



CARYOPHYLLACEAE 



Newkirk St., on chrome 
s lavender). 



$0, Agrostemma githago L, Maryland: Canton, 
ore piles. May 2U, 195U. Reed 333U8 (flowers 

HERNIARIA L. Burstwort. Sepals 5; staminoides small or none; 
stamens 2-5 j style short, deeply bifid; capsule enclosed in the 
calyx and containing one seed. — Small procumbent herbs, with 
small green flowers; stipules minute, scarious. 

51. Herniaria cinerea DC. Introduced from southern Europe; also 
known from southern Arizona and California. Maryland: Canton, on 
chrome ore piles. June 10, 1957. Reed 38308. Small erect annual 
herbs, 2.5-7 cm. tall, sometimes forming mats 7-30 cm. in diameterj 
branches bearing 2-ranked branchlets; leaves opposite with minute 
papery stipules, the blades oblong and about 0.5 cm. long; flowers 
very small, green, crowned in clusters in all the leaf axils; h-5 
sepals, united at the base, usually no petals; 2-5 stamens, in- 
serted on the calyx base, and a short 2-cleft or 2-parted style; 
fruit an achene, enclosed hy the calyx; seeds shining black, minute 
double-convex lens-shaped, rimmed "by a thin margin. 

52. Herniaria glabra L. Native of central and southern Europe. 
Maryland: Canton, on chrome ore piles. Oct. 28, 1958. Reed Ul3h8. 
Stems very branching, slender, prostrate, 5-10 cm. long, forming 
mats on the ground, glabrous or clothed with short and somewhat 
deflexed hairs; root perennial; leaves very glabrous or ciliate at 
the base, the inferior ones opposite, the upper one alternate and 
opposite the floral branches, oblong to elliptic, obtuse, entire, 
attenuate at the base; stipules ciliate; flowers in axillary 
clusters, coalescing on the lateral branches into a somewhat leafy 
spike; calyx nearly hairless, the divisions obtuse; seeds black, 
lustrous. 

53. Saponaria vaccaria L. ( Vaccaria sege talis (Neck.) Garcke). 
Adventive from Europe. Maryland: Canton, Newkirk St., on jJhrome 
ore piles. June 22, 195U. Reed 33371; Oct. 28, 1958. Reed U13U6; 
Nov. 23, 1958. Reed Ul728;"j3l> 9, 1959. Reed U3629; on manganese 
ore piles. July~9n.959. Reed U36535 JulyTFT I960. Reed U6732. 



196U Reed, Flora of chrome & manganese ore piles 329 

5U. Saponarla porrigens L. Native of Amienia and The Levant. 
Maryland: Canton, on chrome ore piles. June 19Sh. Reed 333^7 (US). 
Stem erectj branches divaricating, hairy, viscid; flowers axil- 
lary, on long stalks; peduncles filiform; calyx terete; fruit 
egg-shaped, drooping; leaves lanceolate, connate; petals flesh- 
colored; stamens irtiite. 

55. Silene cserei Baurab, Native of Europe, Maryland: Canton, 
on chrome ore piles. May 21, I960. Reed U6327, Leaves broader 
2-U cm. wide) than in S, cucubalus Wibel (1-3 cm. wide); calyx 
but little inflated and the veins not conspicuously reticulate. 

Also found on the chrome ore piles in the Canton area are the 
following species vti±ch are fairly common. 

56. Cerastium viscosum L. 

57. Cerastium vol ga turn L, 
58» Silene antirrhina L. 

59. Silene noctiflora L. 

60. Stellaria media (L.) Cyrillo 

CEIASTRACEAE 

61. Celastrus scandens L. Maryland: Canton, frequent climbing over 
thickets. Oct. U, 1957. Reed 39559. 

CHENOPODIACEAE 
A triplex patxila L. is represented ty the following varieties in 
the Canton area, on chrome ore piles, in wastes and along the rail- 
roads. 

62. A triplex pa tula L. var. hastata (L.) Gray. Native of Eurasia; 
nat. from Newfoundland to British Columbia, and south to New Eng- 
land, South Carolina, Ohio, Indiana, Illinois and Missouri. Maiy 
collections from Canton, Dundalk and Newport News areas. 

63. A triplex pat^lla L. var. littoralis (L.) Gray, Prince Edwards 
Island to Ontario, south to New Jersey, Pennsylvania, Indiana and 
Wisconsin, Maryland: Canton. Oct. 1957. Reed 39568, 

61. A triplex pa tula L. var. Japoniea Levi. Native of Japan and the 
Orient. Maryland: Canton, on chrome ore piles, forming huge mats 
up to 10 ft. in diameter; Oct. 28, 1958. Reed ia3hh; Nov. U, 1958. 
Reed la36l. This variety has linear to broadly linear leaves, 
similar to those of var. littoralis ; but the fruits are bracteate, 
the bracts being from 1-1.5 cm. broad and long, similar to var. 
brae tea ta Westlund. Plants very large, woody and very branched, 
the bark reddisk, striated with green; the leaves up strai^t, 
glabrous, entire, without widening at the base, obtuse, petiolate, 
the petioles winged; fruits with valve-like bracts, 1-1.5 cm. 
long, disposed in glomerules of 3-1, forming terminal spikes and 
axillary ones as well. 



330 PHYTOLOGIA Vol. 10, no, 5 

65. A triplex nitens Schkuhr. Native of central Europe. Maryland: 
Canton, Newkirk St., on chrome ore piles. Sept, 19, 19^8, Reed 
U1322; Baltimore County, Cub Hill in fields. May 28, 19^9. Reed 
U3122j Aug. 3, 1959. Reed U5?91. Stems herbaceous to nearly woody, 
up to 1,5 ni, tall, erect branched; the leaves triangular, acuminate, 
greenish shining above, glaucous beneath, up to 12 cm. wide and 10 
cm, long, margins dentate, the lower ones cordate-hastate, the 
upper ones deltoid and somewhat auricled, the uppermost ones lan- 
ceolate; perianth of firuit ovate-acuminate, entire, smooth on the 
back, thin and finely reticulate, 

66. A triplex rosea L, Adventive from Eurasia; New York to Wisconsin 
and southward; California, Maryland: Canton, Sept. 1907. Plitt, in 
Reed Herb. (No. 17628). 

67. A triplex thornberi (Jones) Standi. Native of southern Arizona, 
Maryland: Canton, collected many times on the chrome ore piles. 
Sept. 1953. Reed 327514, 32812, 32770, 32715 and 32661; Oct. 12, 
1953. Reed 3?^5T; June 22, 195h. Reed 33li37. (A. elegans var. 
thornberi Jones), Annual or perennial, 2-5 dm7 tall, much-branched, 
the branches slender or stout, obtusely angled, erect or ascending, 
coasrely furfuraceous or gLabrate in age; the leaves numerous, al- 
ternate, sessile or subsessile, the blades oblong to narrowly ob- 
long or lanceolate, 7-20 mm. long, 2,5-6 mm, wide, obtuse or acute 
at the apex, the upper ones mucronate, crenate or attenuate at 
base, the lower ones remotely repand-dentate, the upper entire, 
rather thin and whitish-furfuraceous below, furfuraceous or glabrate 
above; flowers monoecious, disposed in small axillary clusters; the 
fruiting bracts short-pedicellate, orbicular, 3 mn, long, the mar- 
gins deeply laciniate-dentate nearly to the base, the sides each 
having 2 laciniate crests; the seeds 1 mm, long, pale brown; the 
radicle superior, 

Chenopodium L, is represented by several species which have been 
collected maiiy times since 1953 on the chrome ore piles and in the 
nearly wastes. Most common are the following three species, 

68, Chenopodium ambrosioides L» 

69, Chenopodium botrys L, 

70, Chenopoditun album L , 

71. Chenopodium viride L, Native of Russia, northern and western 
Asia (Kashmir to Nepal); and Europe, Maryland: Canton, on manganese 
ore piles, July 17, 1958. Reed lilOOh and Ia006; Sept. 6, 1958. Reed 
la300; Sept. 19, 1958. Reed TIT3U2; July 9, 1959. Reed li3650 and 
U3656, Virginia: Newport News, on chrome ore piles. Aug, 1959, Reed 
liU063 and U140U2, Entire plant usually mealy; leaves long stalked, 
broadly triangular, upper ones lobed, about 3 cm, each way, mar- 
gins sinuate or irregularly lobed; utricle only partially covered 
by the persistent teeth; seeds minutely dotted. 



I961i Reed, Flora of chrane & manganese ore piles 331 

72. Chenopodium vulvar la L. Adventive from Eur ope j local from 
Quebec, Ontario and Wisconsin, south to Maryland and Indiana, Mary- 
land: Canton, on chrome ore piles, collected many times from 19^3 
to 1958. The plant possesses a fetid odor which is distinctive. 
Sept. 27, 1953. Reed 32817; Oct. 2, 1953. Reed 32681; Sept. 30. 
1955. Reed 39239 and 39227; Aug. 23, 1956. Reed 37967; Sept. 19, 
1958. Reed I132U. 

73. Chenopodium paganum Reichenb, Naturalized from Europe; Quebec 
to Alaska and southward. Maryland: Canton, on chrome ore piles, 
Newkirk St. Sept. 27, 1953. Reed 328l5; Sept. 30, 1955. Reed 38235. 

7h. Chenopodium carnosulum DC, Native of Mexico, Maryland: Can- 
ton, on chrome ore piles. Sept. 1955. Reed 38211i, 3 8216. 

75. Chenopodium farinosum (S.Wats, ) Standi. (Ch, macrospermum Hook, 
fil, var. halophilum (Phil.) Standi.; Ch. macrospermum ssp. halo- 
philum (Phil.) Aellen forma farinosum IWats.) Aellen), Native of 
South America (Argentina north to Bolivia, Mexico); California and 
North Carolina. Maryland: Canton, on chrome ore oiles. Sept. 30, 
1955. Reed 3821^2 and 38217; Nov. 10, 1953. Reed 33036; Oct. 25, 
1951. Reed 35189. North Carolina: Wilmington, on ballast. (US- 
h39h$). See Aellen & Just, Amer , Midland Nat,, 30: 58, 19li3, 

76. Chenopodium glaucum L, (incl, Ch, salinum Standi,), Adventive 
from Europe; Alaska to North Dakota, south to Missouri, New Mexico, 
Arizona and Oregon, Maryland: Canton, on chrome ore piles, July 

9, 1959. Reed 1|3626; on manganese ore piles. July 30, 1959. Reed 
U3839. Annual, succulent and glabrous, or nearly so, except the 
densely white-mealy under-surface of the leaves; stems freely 
branching, decumbent or prostrate, h-20 cm. long; leaves usually 
oblong, varying fl-om lanceolate to narrowly ovate, 2-5 cm. long, 
sinuate-dentate, or the uppermost sometimes entire, narrowed to a 
short petiole; flowers in short axillary, often branched spikes; 
calyx lobes narrowly oblong, thin, not keeled, concealing only a 
small portion of the fruit; utricle dark brown; pericarp free; 
seed vertical in the lateral flowers, often horizontal in the ter- 
minal ones, sharp-edged, 0.6 ram, broad, 

77. Polycnemum majus A.Br, Adventive from Europe; also known in 
Ontario. Maryland: Canton, on manganese ore piles. Aug, 1959. 
Reed Ul;376. 

78. Salsola kali L, Newfoundland to Louisiana; also in Eurasia, 
Maryland: Canton, frequently collected on the chroite and manganese 
ore piles since 1953. 

79. Salsola kali var. tenuifolia G. F.W.Meyer, Native of central 
Asia; a troublesome weed, Michigan to Missouri and westward, 
southward to Indiana, West Virginia and North Carolina, Maryland: 
Canton, on chrome ore piles, frequent. June 22, 1951. Reed 33hhh 
and 33Uii6; June 29, 1956. Reed 38827; Sept. 19, 1958. Wee^ U1326. 



332 P H T T L G I A Vol. 10, no. 5 

COMMELINACEAE 
80 » Cojmnellna commcmls L, Common in the wastes throu^out the 
Canton area. 

81. CoimaBllna diffusa Burm. f 11. Native in pantropic and warm- 
ten^jerate regions in American tropicsj northward to eastern Vir- 
ginia and Maryland; a casual weed north to Massachusetts, Ohio, 
Indiana, Illinois and eastern Kansas. Maryland: Canton, on chrome 
ore piles, Nev*:irk St. Sept. 2U, 19^8. Reed 11231; Oct. Hi. 1958. 
Reed 10156; Oct. 28. 1958. Reed 10352; Nov. U, 1958. Reed la36U; 
Octr 6, 1959. Reed U5716 (seems to be spreading to maiy other piles' 

COMPOSITAE 

Mary new genera and species of -this family have been found on th< 
various ore piles in the Canton area. I am indebted to the late Dr. 
S.F.Blake for identifying or verifying the identification of most 
of the species listed below. 

82. Acbi'l 1 ftfl millefolium L. Naturalized from Europe, A common weed 
throu^out the wastes of Canton. Aug. 18, 1959. Reed Uh352. 

83. Ageratum conyzoides L, Old World tropics and subtropics; West 
Indies and tropical continental America. Maryland: Canton, on 
chrome ore piles. Sept. and Oct, 1958. Reed U1226 and lai6u. Annua: 
more or less pubescent, branched, 9 dm. hi^ or less, the stem 
terete, the branches widely ascending; leaves thin, ovate, 2-8 cm. 
long, mostly obtuse at the apex, cuneate to subcordate at the base, 
crenate or crenate-dentate, the slender hirsute petioles 3 cm. long 
or less; corymbs conpound, convex, the head several to numerous, 
about 6 mm. broad, mat^r-flowered; involucre can^janulate, its bracts 
oblong, glabrous or sli^tly pubescent, green with scarious margins 
acuminate; receptacle naked; corolla blue or white; achenes black, 
shining; pappus of 1-5 lanceolate scales, attenuate at the apex, 
sometimes very unequal in length and the shorter ones bluntish, 

8h. Ambrosia artemisilfolia L. A common weed throughout the wastes 
of Canton. 

85. Ambrosia trifida L, A common weed -throughout the wastes of 
Canton. 

86. Anaphalis margaritacea (L.) C.B.Clarke var, intercedens Kara 
Native of Japan; also from Newfoundland to Alaska, south to North 
Carolina, West Virginia, Ohio, Michigan, Wisconsin, Missouri, South 
Dakota and New Mexico, Maryland: Canton, common in waste areas, 

87. Antherais arvensis L. Naturalized from Europe; Maine to western 
New York and south to Georgia. Maryland: Canton, frequent on 
chrome and manganese ore piles. 

88. Anthemis cotula L. Naturalized from Europe; Newfoundland to 
Alaska and southward. Maryland: Canton, Oct. 1900. FOitt. 



196U Reed, Flora of chrcme & manganese ore piles 333 

89. Arctium minus (HiH) Bernh. Naturalized from Europe; Newfound- 
land to British Columbia, south to Virginia, West Virginia, Mis- 
souri, Kansas and California. Maryland: Canton, on chrome ore 
piles; Port Covington on wastes. 

90. Artemisia annua L, Naturalized from Eurasia; Prince Edwards 
Island to Ontario, south to Long Island, Virginia, Alabama, Tennes- 
see and Arkansas. Maryland: Locust Point, along RE tracks, rather 
frequent, 

91. Ajtemisia ludoviciana Nutt. var. gnaphalodes (Nutt.) Torr. & 
Gray Native on the prairies from southern Ontario and Michigan 

to southern British Columbia, south to Arkansas, Oklahoma and Texas; 
naturalized eastward to Quebec, New England, New Jersey, Delaware 
and Maryland, Maryland; Canton, in large patches along RR., col- 
lected from 1958 to I963. Virginia: Newport News, RR wastes at 
port. Aug. 19^9. Reed lii:082, 

92. Artemisia scoparia W, & K. Native of central Europe (Bohemia, 
Austria). Maryland: Canton, on wastes. Aug. 1956. Reed 37971, 
Stem sirple, erect; leaf bi-tripinnatifid, ovate in outline; seg- 
ments of the lowest leaves linear-lanceolate, of the rest, linear; 
upper leaves sessile, singly pinnatifid; uppermost undivided or 
with a tooth at the base; outer florets with anthers; receptacle 
naked; auricle at base of leaf-stalk about the middle of the 
flowering- stem; flower broadly ovoid or subgLobose, hairless, 

93. Artemisia abrotamun L, Introduced from southern Europe. 
Virginia: Newport News, on chrome ore piles. Nov. 1959. Reed U5888. 

9h* Aster sin^lex Willd, var. ramosissimus (T.& G.) Cronq, Fre- 
quent on the wastes at Canton, Oct, kt 1957, Reed 39565 and 39572. 

95. Aster subulatus Michx. Natural range from southern Maine to 
Delaware and Maryland, and southern Michigan, Maryland: Canton, 
on chrome ore piles. Sept, 27, 1953. Reed 32662, 

96, Baccharis halimifolia L, Native from Mexico and Texas to 
Florida, north along the coast to Massachusetts, Maryland: Canton, 
on wastes on chrome ore piles and gypsum piles, Oct. 2^^ 195U. 
Reed 35222; Sept. 20, 1957, Reed 39320; Oct, l, 1957. Reed 3956h 
and 39586, 

BAHIA Lag, Ray flowers pistillate and fertile, rays from in- 
conspicuous to definite, or in 1 species lacking, yellow; disk 
flowers perfect and fertile, vellow; anthers united, not caudate 
at the base; achenes narrow, U-angled; pappus of several paleae, 
these sometimes with liie thickened midrib excurrent as an awn, or 
without pappus; annual or perennial herbaceous plants, with alter- 
nate or opposite leaves, these entire to variously divided or dis- 
sected; heads corymbose; involucres campanulate, hemispheric, tur- 
binate or obconic, bracts in 2 or 3 series, herbaceous to scarious 



33U PHTTOLOQIA Vol. 10, no. 5 

or colc«red in partj receptacles mostly flat, naked or alveolate. 

97. Bahia schaffnerl Wats. Native of Mexico, Maryland: Canton, 
on chrome ore piles. Sept, 19^3. Reed 3278?. Annual; stem decum- 
bent and branched at base, strigose; leaves mostly opposite, 2-3 
cm. long, ternately 3 or h times dissected, with narrow linear 
divisions, strigose; heads terminating ttie branches; peduncles 
2-lt cm. long; involucre hemispheric, h-5 mm. hi^, about 8 mm. 
broad; bracts strigose, oblanceolate or obovate, acute, somewhat 
yellowish towards the tip; ligule yellow, oval, 3 ran. long; tube 
of disk-corollas slender, glandular, 2 mm. long, throat campanula te, 
1.? mm, long; achenes narrowly obpyramidal, 3 inm, long, hispidulous 
on the angles; pappus of 8 obovate squamellae, 1 mm. long, with a 
short ovate thickening at the base. 

98. Bidens frondosa L. A common weed from Newfoundland to Wash- 
ington and southward. Maryland: Canton, on wastes between the 
chrome ore piles. Oct. U, 19?7. Reed 39^88; Nov. 28, 19?8. Reed 
Iil7h8. 

99. Bidens pilosa L. Adventive from tropical America, northward to 
Massachusetts. Maryland: Canton, on chrome ore piles. Sept. 27, 
1953. Reed 32780; Oct. 12, 19^3. Reed 32873, 32878 and 32882; Nov, 
2, 1953, Reed 33001, Weecfy square-stemmed annual, with siii?)le 
ovate or 3-5'-parted leaves, white to pale yellow or purplish 
ligules and linear-tetragonal achenes, 5-9 nmi, long, 

100. Bidens polylepis Blake. Native in the Mid-west from Illinois 
to Iowa, Kansas and Colorado, south to Tennessee, Missouri, Okla- 
homa and Texas; natiiralized east to the Atlantic States, Maryland: 
Canton, on chrome ore piles, Sept. 6, 1958. Reed ljl289. 

101. Bidens tenuisecta Gray. Native from Idaho to Colorado, Texas 
and Arizona, south to Mexico, Maryland: Canton, on chrome ore 
piles. Oct. 1959. Reed U5691. Annuals, with stems up to 6 dm. 
tall, branched from the base, glabrous; leaves twice or thrice 
ternately or pinna tely divided into linear lobes; heads about 1 cm, 
high and broad; bracts linear, hirsute, the outer usually slightly 
longer than the inner; achenes glabrous, the outer 6-8 mm, long, 
the inner 10-12 mm, long; awns fully 3 vm, long, those of the outer 
achenes shorter. 

102. Carduus nutans L. Adventive from Europe; naturalized from 
St. P. et Miq. to Iowa, south to Nova Scotia, New England, ^te^y- 
land. District of Columbia and Missouri. Maryland: Canton, on 
wastes, 1901 and 190U, Plitt , 

103. Cnicus benedictus L, (Carthamus tinctorius L.), Native of 
southern Europe and Asia Minor; sparingly established from New 
Brunswick and Illinois and southward, Wbryland: Canton, plentiful 
on wastes. June 1955. Reed 36552 (US), 



196U Reed, Flora of chrome & manganese ore piles 335 

lOli, Centaurea depressa Bieb. Native of Persia and Caucasus, 
Maryland: Canton, on chrome ore piles, June 22, 19^h. Reed 33372. 
Perennial, almost prostrate, white-pubescent; lower leaves ob- 
long, entire or Ijrrate, the upper ones linear- lanceolate; heads 
blue, up to 3 cm, across, the marginal flowers enlarged; bracts 
of involucre with black or white margins. 

105, Centaurea calcitrapa L. Naturalized from the Mediterranean 
region in the southeastern states, north to New York and southern 
Ontario, Maryland: Canton, on chrome ore piles. May 2ii, 195U. Reed 
333^7; Aug, 23, 1956. Reed 37971; and 37966; June 29, 1956. Reed 
38826. 

106, Cichorium intybus L. Common as a weed throu^out the Canton 
area. Flowers varying from blue to pink or white. 

107, Cirsium arvense (L.) Scop. Natxiralized from Europe. Mary- 
land: Canton, July 6, 190li. Plitt . 

108, Cirsium arvense var, horridum Wimmer & Grab, Naturalized 
from Europe; the more commion variety in eastern United States. 
Maryland: Canton, on wastes near the chrome ore piles. June 12, 
1959. Reed U2977. Plant very spiiy; leaves deeply pinna tif id. 

109, Cirsium discolor (Muhl.) Spreng, Frequent on the chrome ore 
piles and wastes in Canton. Sept. 2h, 1957. Reed 395U9. 

110, Cosmos sulphureus Cav. Introduced from Ifexico; New Jersey 
and Pennsylvania, southward; spread fi'om cultivation. Maryland: 
Canton, on chrome ore piles, Nov, 5, 1953. Reed 32976 (US). 

111, Cosmos parviflorus (Jacq.) HBK, Native from Colorado south 
to Texas, Arizona and Mexico; adventive on wool wastes in Maine 
and Massachusetts, Maryland: Canton, on chrome ore piles. Oct, 
12, 1953. Reed 32916 (US); Nov. 2, 1953. Reed 32988, Stems 3-8 dm. 
tall, glabrous; leaves bi- or tripinnatifid intx) filiform divisions; 
involucres 6-7 mm, long; rays 7-12 mm, long; achenes 7-10 mm, 

long including the beak which is about one-third to one-half as 
long as the bo<^; pappus awns about 2 mm, long, 

112, Coryza bonariensis (L.) Cronq, Native of Venequela and Para- 
guay, Virginia: Newport News, in RR yard at port. Aug, 1959, 
Reed hh038. 

113, Dyssodia papposa (Vent.) Hitchc, Native from Louisiana to 
Mexico and Arizona, north to southern Ontario, Minnesota, North 
Dakota and Montana; adventive east to New England. Maryland: 
Canton, on chrome ore piles. Nov. 2, 1953. Reed 32998; Nov, 10, 
1953. Reed 33028. 



336 PHYTOLOGIA Vol. 10, no. $ 

nil. Eclipta alba (L.) Hassk. A common weed vhich grows luxuriant- 
ly on the chrome ore piles, as well as in wastes areas of Canton, 
Sept. 27, 1953. Reed 32737; Oct. U, 1957. Reed 39589; Nov. 23, 1958. 
Reed hl73U. 

115. Eclipta erecta L. Native of Lower California and Mexico, south 
to Guatemala, Honduras, Colombia and the Galapagos Islands. Maryland 
Canton, on chrome ore piles. Aug. and Oct. 1959. Reed 11356 and 
U5703. Ihis species is sometimes considered as a variety of E. alba, 
having long-peduncled heads. 

Eupatorium is represented in the Canton, DundaUc and other 
wastes areas fay three species, the last species forming huge plants 
on the chrome ore piles. 

116. Eupatorium hyssopifolium L, July 6, 190li. Plltt . 

117. Eupatorium purpureum L. 

118. Eiroatorium serotinmn Michx. Maryland: Canton, on chrome ore 

piles. Aug. ^3, 1956. Reed 37972; Clinton St. Oct. U. 
1957. Reed 39581;; Dundalk. Sept. 2U, 1957. Reed 3953U. 

119. Flaveria trinervia (Spreng.) Mohr. Florida and Alabama to 
soutbern Arizona, southward to South America; Cuba; introduced at 
Westford, Massachusetts, Maryland: Canton, on chrome and manga- 
nese ore piles. Sept. 27, 1953. Reed 328UO; Nov. 2, 1953. Reed 
3298li and 32997; Nov. 23, 1958. |eed U1731; on manganese ore 
piles. Oct. 6, 1959. Reed li5699. Virginia: Newport News, on man- 
ganese ore piles. NovTTF, 1959. Reed U 5866. Annual, dichoto- 
mously and divaricately branched; stem erect, 3-12 dm. high, some- 
times tinged with red, glabrate; leaves connate, lanceolate, 3- 
nerved, serrate, 3-9 cm. long, the lower short-petioled; cymes 
congested and head-like, 2-2.5 cm. broad, axillary, subtended ly 
3 pairs of leaves; heads small, usually 1-flowered, the flower 
either suKLigulate and pistillate or regular and hermaphroditic; 
involucral bracts concave, 1 c* 2, if the latter, the outer 
shorter; corolla of the pistillate flowers 1.5 mm. long, the 
ligule oblique, 3-dentate; corolla of the hermaphroditic flower 
2 ram. long; tube villous; throat can^sanulate; achene 2 ram. long, 
someviiat clavate; pappus wanting. 

120. G uizotia abyssinica (L. fil.) Cass. Adventive ffom tropical 
Africa, from Connecticut to Michigan and Pennsylvania. Maryland: 
Canton, on chrome ore piles. Nov. li, 1958. Reed 11371. 

IZL. Helianthus annuus L. Native from Manitoba and Minnesota 
to Texad and westward; cultivated and spreading eastward to 
Quebec, New Brunswick, Nova Scotia and the Atlantic States. 
Maryland: Canton, whole areas have been taken over by this sun- 
nower. The flowers range from 2 to 6 inches in diameter, and 
most of the plants have many-headed stalks, up to 10 ft. ^±. 
Some plants have been collected on the chrome ore piles which 
are only 2 ft. tall and are similar to Helianthus lentlcula ris 
Dougl. <July 1959. Reed U3657). Oct. 2, 1953. Reed 32680; Sept. 



196U Reed, Flora of chrome & manganese ore piles 337 

20, 1957. Reed 39386j Nov. U, 19^8. Reed la358; Nov. 23, 19^8. 
Reed lil738; Port Covington. Nov. 8, 19^7. Reed 381485^, 

122, Helianthus tuberosrus L. Common in the wastes Just south of 
Highlandtovm. Sept. 20, 1957. Reed 39389. 

123. Helianthus laetiflorus Pers. Common in the wastes about 
Port Covington. Nov. 8, 1957. Reed 39h83. 

HETEROSPERMA. Cav. (Icon. Ill, p. 31, t. 267. 179U (1795), pre- 
face on p. V says 10 Jan. 1795j also in 1802). (Willdenow in 180U, 
without due cause changed it to Heterospermum — vide KLake, Jo\irn, 
Bot. 53: 322. 1915). 

12U, Heterosperma pinnatum Cav. Native of Mexico. Jferyiand: Can- 
ton, on chrome ore piles, Oct, 12, 1953. Reed 329l8j Nov. 2, 1953. 
Reed 32989; Nov. 5, 1953. Reed 32978. Stems herbaceous, branches 
oppositej leaves opposite, very glabrous, connate, pinnate with 
uneqxial acute-linear pinnulesj flowers terminal, peduncled, sin^e- 
floweredj calyx very deeply laciniate, acute-linear, longer than 
the flower, ciliate at the base; corolla yellow j anthers fuscous; 
paleae ovate, striate; florets bearly 16, distinguished by the 
violaceous-fuscous of the rays and the yellow of the hermaphroditic 
ones. 

125. Helenium tenuif olixim Nutt, (H, amarum (Raf,) Rock), Common 
in RR yards in port of Newport News, 7a, Aug, 1959. Reed lih036, 

126. Hsrpochoeris radicata L, Naturalized from Europe; Newfound- 
land to Ontario, south to North Carolina, West Virginia, Ohio, 
Indiana and Illinois, Virginia: Newport News, on chrome ore piles, 
Nov. 1959. Reed 15890. 

127. Iva frutescens L, Frequent in the wastes in Canton. Sept. 
1957. Reed 39533. 

128. Inula helenium L, Naturalized from Europe; from eastern On- 
tario to southern Quebec, and southward, Maryland: Canton. Sept, 
12, 1900 and Oct. 18, 1900, C.C.HLitt . 

>EIAMPODIUM L, Ray flowers pistillate and fertile, rays white, 
yellow to sometimes pink, spreading, conspicuous; disk flowers per- 
fect but sterile, with undivided style; anthers united, entire at 
base; achenes broadening upward and more or less incurved; pappus 
wanting, — Perennial herbaceous plants with taproots and caudex; 
stems leafy, the leaves opposite, entire to sinuately lobed; heads 
terminal on the stem and branches; involucres canpanulate, bracts 
in 2 sets, the outer U or 5 herbaceous, broad, flat and partially 
united, the inner hooded, each embracing a ray achene and de- 
ciduous with it; receptacle convex or conical, chaffy, 

129. Melanpodium hispidum HBK, Native of S, Arizona and Mexico. 
Maryland: Canton, on chrome ore piles. Sept, 1953. Reed 32739 and 
327li6, Rays yellow, inconspicuous; usually not more than 2 mm. 



338 PHYTOLOGIA Vol. 10, no. $ 

longj plants annual, strictly herbaceous; heads 2-6 mm. wide, 
usually sessile or subsessile; fruit not hooded or beaked at apex, 
Kearney, Peebles et al, Arizona Flora, p. 890. I960, 

130, Onopordum acanthium L, Naturalized from Europe (and eastern 
Asia) J from New Bminswick to Ontario, south to Alabama and south- 
eastern Missouri, Scotch thistle. Maryland: Canton, forming 
large patches. 1901-1 90U, G.C.Plitt . 

131, Parthenium hysterophorus L, Adventive from tropical America, 
from Florida to Texas, locally north to Massachusetts, Pennsylvania, 
Dhio, Michigan, Illinois, Missouri and Kansas, Maryland: Canton, 
on chrome ore piles, Sept. 2?, 1953. Reed 32806; Oct. 2^, 195U. 
Reed 33363. 

132, Pulicaria dysenterica (L.) Gaertn. Naturalized from Europe; 
otherwise only recorded from the shores of the Potomac River, 
Charles Co., Maryland. Also Maryland: Canton, on chrome ore piles. 
Oct. 17, 1956. Reed 382U9, 

SANVITALIA Lam, Heads heterogamous, radiate; rajrs pistillate, 
fertile, ligulate, persistent on the achene; disk fl-owers perfect, 
fertile, tubular; involucre depressed hemispheric, of herbaceous 
bracts imbricated in 2-3 series; receptacle convex to conic, 
chaffy; scales enfolding the disk flowers, persistent on the re- 
ceptacle; achenes heteromorphous, ray achenes triangular, dorsally 
conpressed, 3-a'wned, the disk achenes laterally conpressed, winged 
or wingless, 2-awned or awnless; annual herbs with rather showy 
heads of yellow rays and dark purple disks, 

133, Sanvltalia angustifolia Engelm, Native of Mexico. Maryland: 
Canton, on chrome ore piles. Sept, 2?, 1953. Reed 32775; June 22, 
195U. Reed 33ltl8. Stem diffused; leaves lanceolate, strigose-hispid 
ligules small, aristate, \d.th shorter acute setulae, 3-5 barbate- 
apiculate; achenes or outer disk exalate subclavate, the inner 
short winged, bi-ari-stellate, 

13h, Sanvitalia procumbens Lam. Native of Mexico, Yucatan, Mary- 
land: Canton, on chrome ore piles, Sept. 27, 1953. Reed 32735 and 
32785; June 22, 195U. Reed 33Ul6. Annual, diffuse and procumbent, 
wi-th subangles stems and branches; leaves opposite, petioled, lan- 
ceolate-ovate, strigose; inflorescence of solitary heads terminat- 
ing the slender hirsute branches, the heads 7-8 mm, high, 12-15 
mm, broad, subtended by 2-3 foliaceous, hirsute bracts; rays 8, 
disk flowers numerous; involucral bracts orbicular, appressed,with 
foliaceous spreading ciliate tips, pubescent, the inner little ex- 
ceeding the outer; rays yellow, turning white in fruit, ovate, ^-6 
ram, long, minutely bidentate; disk corollas dark purple, 5-lobed; 
receptacle convex to conic; scales lanceolate, conduplicate, ex- 
ceeding and partly enfolding the disk flowers; achene of ray flowers 
cineraceous black when mature, 1,2 by 3 mm,, oblong cuneate, in 
section triangular, tuberculate, hirsute at the base; awns 3, di- 
vergent, 2 mm, long; achenes of disk flowers dull brown, ,85' by 2 
mm,, obovoid, in section plano-convex to rhomboidal biconvex. 



196U Reed, Flora of chrome & manganese ore piles 339 

with one or a pair of loosely cellular, cHiate-fimhriate wings, 
5-7 ranu wide, or winglessj the winged with 1 or 2 awns, 1,2 ram, 
long or awnless, faintly tuberciilatej the wingless achenes awn- 
less and covered with tubercules each tipped by a short stiff 
hair or its stub. 

SCHKUHRIA Roth. Annuals, rarely perennials, with slender, bran- 
ching, erect or decumbent stems; stems gLabrate to hispidj lower- 
most leaves opposite, upper alternate, pinnately or bipinnately 
divided into linear-filiform lobes, rarely siii5)le, often irpressed- 
punctatej heads discoid or radiate with one to few rays j involucre 
obconic to turbinate j bracts of the involucre U-18, obovate to ob- 
lanceolate, rather narrow, scarious and frequently colored on the 
margins, occasionally one or more smaller bracts present; ray 
flowers 1-3, rarely more, yellow to white, minute; disk flowers few 
to numerous, yellow, rarely red-tipped, lobes 5, extending about 
half the length of the throat, glandular tube equal to or less than 
the length of the throat; style branched with short acute appen- 
dages; achenes elongate; obpyramidal, generally U-angled, villous 
or hispid on the angles, particularly at the base; pappus of 8, 
rarely more, scarious squamellae, calloused at the base or with 
prominent mid-rib becoming an awn in some of the species. 

135, Schkuhria wisllzeni A.Gray, (= S. anthemoides var wislizenii 
(Gray) Heiser, Ann, Missouri Bot. Gard., 32: 273, 19li5). Native 
of Arizona and Mexico, Iferyland: Canton, on chrome ore piles, 
Oct. 25, 195ii, Reed 35198. Erect annual; stems glabrate, striate, 
20-50 cm. in height; leaves glabrate, pinnately or bipinnately 
dissected into linear-filiform segments, lO-UO mm, long, 0,5-2 
mm, wide, or the upper and lower ones entire, conspicuously glan- 
dular-punctate; involucre bracts yellow to purple at the apex; 
ligules seldom over 1 mm. long, almost as wide; squamellae lanceo- 
late to ovate-lanceolate, those of the an^es awn-tipped, the 
intermediate ones shorter, rauticous. 

SI>BIA Pers. Coarse annual or perennial herbs; leaves alter- 
nate or the lower ones usually opposite; heads rather large, radi- 
ate or discoid, the flowers yellow or purple; phyllaries 3-U-seri- 
ate, subequal or slightly graduated, narrow; receptacle slightly 
convex, paleaceous; rays usually yellow; disk achenes flat, ob- 
ovate or oblong, glabrous or seriaceous, the edges thin, not mar- 
ginate; pappus none or of 2 awns, 

136, Simsia lagascaeformis DC. ( Encelia lagascaeformis A.Gr, in 
litt."] Native of I4exico. Maryland: Canton, on chrome ore piles. 
Oct, 12, 1953, Reed 3^915; Nov, 2, 1953. Reed 32985 and 33000; 
Nov, 5, 1953, Reed 32980. 

137, Solidago altissima L, Common on wastes along Clinton Street, 
Canton, Oct. h, 1957. Reed 39585; along creek near Dundalk, Sept, 
2U, 1957. Reed 39532. 



3U0 PHITOLOGIA Vol. 10, no, 5 

138. SolldagQ senpervlrens L, Coramon along creek near Dundalk, 
Sept . 2U, 1957. Reed 39h^, 

139, Sonchua arvenals L. var. glabrescens Guenth., (jraebn. & Wiwnu 
Natttralized from Europe; from eastern Quebec to Minnesota, south 
to Nova Scotia, Connecticut and Indiana. Maryland; Canton, on 
oyster shell piles. Oct. 31, 19^7. Reed 39hli5; Nov. 23, 19?8. Reed 
UI729. 

lliO. Sonchus oleraceus L. A frequent weed on the oyster shell piles 
and the chrome ore piles in Canton. On chrome ore piles, Sept. 27, 
1953. Reed 32790: June 22, 19^. Reed 33U27; on cyster shell piles. 
Oct. 3I7T957. Reed 39Ul;6j July 177^958. Reed 10995? Oct. U, 19^7. 
Reed 39?69j Nov7T5, 1958. Reed U1737. 

Ha. Tagetes erecta L. Native of Mexico; cultivated and escaped in 
various areas of the West Indies and North America. Maryland: Can- 
ton, on dirorae ore piles. Nov. 2, 1953. Reed 32983. Glabrous, 
often much-branched, up to 8 dm. tall, or less; leaves pinnately 
divided, somewhat petioled, the lower 10-15 cm. long; leaf-segisents 
narrowly oblong to oblong-lanceolate, gland-dotted, sharply ser- 
rate, acute, 1.5-2.5 cm. long, or the lower ones much smaller; 
heads solitary at the ends of the branches, 2.5-b.5 cm. broad; 
peduncles lI-lO cm. long, swollen below the heads; involucre 1.5-2 
cm. high, its bracts united to near the summit; rays 10-20 cm. long 
or longer, yellow. 

Ili2. Tagetes minuta L. Introduced fi-om South America; South Caro- 
lina to eastern Virginia, ncjrth to Massachusetts. Maryland: Canton 
on chrone ore piles. Nov. 23, 1958. Reed U 1730. 

IU3. Tagetes pusilla HBK. Native from Guatemala and Costa Rica to 
Ecuador and Bolivia. Maryland: Canton, on chrome ore piles. Oct. 
12, 1953. Reed 32890 (US); Oct. 25, 195li. Reed 35200. Annual, 
diffuse; stem rarely more than 1 dm. talL, branched at the base, 
angled, striate, sometimes minutely scabrous above; leaves opposite 
or the upper alternate, pinna tif id, or bipinnatifid with linear- 
filiform divisions; heads subsessile or short-peduncled; peduncles 
rarely 1 cm. long, angled, sulcate; involucre turbinate-clavate, 
6-7 mm. long, 2-2.5 mm. broad, decidedly an&Led at the base; bracts 
about 5, with rounded, mucronate, membraneous-margined tips, and 
with 3 rows of mar^r minute glands; ray-flowers 1-3, ligulae glab- 
rous, elliptic, white, sometimes wanting; disk-flowers 8-10, corol- 
las glabrous, 3 mm. long; tube shorter than the trunpet-shaped 
throat; achenes 5-6 mm. long, strongly striate; 2 or 3 of the 
squamellae bristle-like, hispidulous, about h mm. long, fully twlf4 
as long as the other 3 or h which are linear or oblong, truncate^ 
often unequal in length, 

IRinAX L. Disk-flowers perfect and fertile, the ray-flowers pis- 
tillate, the rays often 3-lobed; involucre ovoid to hemispheric, 
its nearly equal bracts in few series, or the outer smaller than 
the inner; receptacle flat or convex, the chaff subtending the 



1961i Reed, Flora of chrcme & manganese ore piles 3Ul 

disk-flowers; anthers auricled at the base or sagittate; style-bran- 
ches of the disk-flowers subulate-appendaged; achenes silky-villous; 
pappus of many aristate plumose scales, perennial herbs with oppo- 
site, dentate or incised leaves, and long-peduncled heads of tubu- 
lar and radiate flowers. 

Ihh, Tridax accendens Blake. Native of ^fexico, Maryland: Canton, 
on chrome ore piles. Oct. 12, 1953. Reed 32899. Stems 5 dm, tall, 
with numerous erectish branches above, green, purplish-tinged, sub- 
terete, somewhat sulcate below, sparsely spreading pilose with 
several-celled hairs 1-1.5 inm, long, mostly tipped with brownish 
glands, the branches and pedicels puberulent with minute several- 
celled mostly incurved glandless hairs; leaves opposite, shorter thar 
the internodes, petioles 2-8 cm, long, pilose with gland-tipped hairs 
blades of larger leaves 5 cm, long, h cm, wide, subtruncate at the 
base and then shortly cuneate-de current with the petiole, remotely 
repand- serrate with 5-8 pairs of low bluntish teeth, herbaceous, 
green on both sides, above sparsely hirsute-pilose with subtubercu- 
lated -based hairs, beneath more sparsely hirsute-pilose, chiefly 
along the veins, triplinerved about h mm, above the base; panicle 
about 30 cm, long, 19 era, wide, nearly naked, its lowest branches 
subtended by reduced leaves, the remaining bracts linear- lanceolate 
or narrowly triangular, 3-10 mm, long, pedicels mostly 13-15 cm, 
long; heads canpanulate, about 22-flowered, about 5 mm, high, 7 mm, 
thick; phyllaries 5, 1-seriate, all subtending flowers, greenish 
with narrow whitish scarious margin, usually browning at apex, 
6-8-vittate, obscurely ciliolate above, glabrous on back; recep- 
tacle low-conical, the palaes readily deciduous; corollas white, 
densely hirsutulous on tube, sparsely so on some of the nerves and 
on teeth, 3.5-3.8 mm, long (tube 1 mm., throat cylindric-oblong, 
about 1,5 ram, in outer corollas, 1.8 mm. long in central flowers; 
teeth 5, broadly triangular, 1.2 mm. long in outer corollas, 0.8 
mm. in central flowers); achenes of outer flowers obovoid, some- 
what conpressed, densely and shortly silky-pilose, 5-ribbed, 2 mm, 
long, their pappus persistent, of about 20 lanceolate lacerate- 
fimbriate squamellae, united at the base, about O.U ram, long; cen- 
tral achenes obpyramidal with 5 black glabrous ribs, densely 
short-pilose, 1,8 ram, long, their pappus of about 20 alternately, 
somewhat uneqiial oblong obtuse fimbriate squamellae, 0,8-1 mm, 
long, iinited at the base in a thick ring; intermediate fruits with 
intermediate characters. 

Ilt5« Verbesina encelioides (Cav,) B, & H, Native from Montana to 
Arizona, east to Kansas and Texas; adventive to Missouri and east- 
ward to New England, Maryland: Canton, on chrome ore piles. Sept, 
27, 1953. Reed 32660; Oct, 12, 1953. Reed 32892; June 22, 195U. 
Reed 33la5j~Oct« 5* 195li. Reed 35207. 



3U2 



PHITOLOGIA 



Vol. 10, no. $ 




¥eeds on chrome ore at Canton, Tferyland 




Araaranthus, grasses on chrome ore at Canton, Maryland 



196U Reed, Flora of chrome & manganese ore piles 3U3 

lU6, Xanthlum chlnense Mill, Maryland: Canton, common in wastes, 
Sept. 27, 1953. Reed 32829. 

lLi7« Xanthlum spinosa L. Maryland: Canton, common in several waste 
places and on chrome ore piles. Sept. 27, 1953. Reed 32708, 32781, 
328U9J Oct. 12, 1953. Reed 32875? Nov. 2, 1953. Reed 32987; Oct. 
25, 19^. Reed 3523U; Sept. 30, 19^5. Reed 3821a; Oct. 17, 1956. 
Reed 382U6. 

CONVOLVULACEAE 

lli8. Col volvulus arvensis L. Natiaralized from Eurasia, from south- 
ern Quebec, southward and westward. Maryland: Canton, on chrome 
ore piles. Sept. 27, 1953. Reed 32658; July 17, 1958. Reed ' li09935 
wastes, Clinton and Holabird Streets, Canton. May 22, 1958. Reed 
h0lil5. 

1149. Cuscuta gronovii Willd. Not uncommon on plants in wastes in 
Canton. July 6, 190U. Plitt. 

150. Evolvulus filipes Mart. Central and tropical and subtropi- 
cal South America. Maryland: Canton, on chrome ore piles. Sept, 
27, 1953. Reed 3281;6, Herb with slender, erect or ascending 
stems, appressed hairy; leaves oblong- or linear-lanceolate, acute 
or acuminate, li-lO mm, long, 1-2 mm, broad, appressed hairy or 
nearly glabrous, very shortly petioled; Inflorescences axillary, 
1- or rarely 2-3-flowered; the pedicels 1-U mm, long; bracts 
minute, persistent, linear-lanceolate; sepals lanceolate, acute, 

2 mm. long; corolla rotate with short tube, white or lilac; cap- 
sule globose, 2-3 mm, in diameter, Ij-valved, U-1-deeded; seeds 
glabrous, 

151. Ipomaea aff . angustifolia Jacq, Native of Africa; also known 
from Australia and Puerto Rico, Virginia: Newport News, on chrome 
ore piles, marked from India, Nov. 1959. Reed U5901. 

152. Ipomaea nil (L.) Roth, Old World Tropics j Hawaii} continental 
tropical America; West Indies; Florida, rferyland: Canton, on 
chrome ore piles, Oct, 1959. Reed U58m, 

CRUCIFERAE 

153. Arabidopsis thaliana (L. ) Henyh. Naturalized from Europe; 
Massachusetts to Michigan and Illinois, and southward. Frequent 
on the wastes in Canton. April 2ii, 1958. Reed U07OI. 

15I1. Brassica kaber (DC.) L.C.Wheeler, var. pinnatifida (Stokes) 
L.C.Wheeler. (B. arvensis (L.) Rabenh., non L.; Slnapls arvensis L.) 
Natxiralized from Eurasia. Maryland: Canton, on wastes. May 16, 
1903. Plitt . 

155. Brassica hirta Moench, (B. alba of Amer. auth., non Gilib.| 
Sinapis alba L7T^ Maryland: Canton, on wastes. May 1903. Plitt. 
(Reed Herb.). 



3141 PHTTOLOGIA Vol. 10, no. $ 

1^6. Brassica juncea (L.) Coss. Maryland: Canton, on wastes and on 
chrome ore piles. June 10, 1957. Reed 388OO. 

1^7. Brassica nigra (L. ) Koch. Maryland: Canton, on chrome ore 
piles. Oct. 25, 195U. Reed,Herb. No. 9l66. 

158. Brassica rapa L. (B. campestris L.) Naturalized from Eurasia; 
throughout eastern North^America as a weed. Maryland: Canton, May 
1903. Plittj on chroma ore piles, June 22, 195li. Reed 33359j Oct. 
25, 195It. Reed 352U3j also fields near Fork, Baltimore Co. May 10, 
1959. Reed"35H'79. 

159. Cakile maritima Scop, Adventive from Europe; sporadic on 
rubbish piles along coast, eastern United States. Maryland: Canton, 
along shore near Newklrk Street. Sept. 27, 1953. Reed 3270Uj May 
2h, I95I4. Reed 333U6j on chrome ore piles. July 17, 1958. Reed 
U099U. 

160. Capsella bursa-pastoris (L.) Medic. Naturalized ffom Europe, 
throufihout North America and beyond. Common in the wastes in 
Canton. April 2U, 1958. Reed U0706. 

161. Cardaria draba (L.) Desv. ( Lepidium draba L.) Naturalized 
from Europe; from Nova Scotia to District of Columbia and westward. 
Maryland: Canton in wastes. May 1903. Plitt. 

162. Coronopus didymus (L.) Sm. Naturalized from Europe, from 
Florida to Texas, north to Newfoundland. Maryland: Canton, on 
chrome ore piles. July 17, 1958. Reed laOOO. 

163. Diplotaxis tenuifolia (L.) DC. Naturalized from Europe, from 
New Brunswick to Ontario, south to Nova Scotia, New England, Vir- 
ginia and Michigan. Maryland: Canton, on ballast lot. Oct. 17, 
1900. C.C. Plitt . (Reed Herb.); May I6, 1903. C.C. Plitt . 

I6H. Diplotaxis siifolia Kunze. Maryland: Canton, on chrome ore 
piles. Oct. 2, 1953. Reed 32682. 

l65. Efcyj5i™i«i repandum L. Adventive from Eurasia, from Massachusetts 
to Ctt^on, south to Alabama, Arkansas and Texas. Maryland: Canton, 
on chrome ore piles. June 23, 1955. Reed 36550. 

166. Lepidium campestre (L.) R.Br. Common in wastes areas throu^- 
out the Canton area. Canton, on the chrome ore piles. April 2k, 
1958. Reed U0697. 

167. Lepidium yirginicum L . Common in wastes areas throu^«ut the 
Canton area. June 23, 19^5. Reed 36551. 

168. Lepidium graminifolium L. Native of Europe and the Medi- 
terranean region; waif in New Jersey and Pennsylvania. Maryland: 
Canton, on chrome ore piles. June 22, 19514. Reed 33U2U, 3336U; 
May 21,; 195U. Reed 333U6; June 23, 1955. ReeT335U9; May 2U, I960. 
Reed U6326. 



196ii. Reed, Flora of chrome &. manganese ore piles 3hS 

169. iJiSpldlmn hyssop ifollum Desv, Native of New Holland. Mary- 
land: Canton, on chrome ore piles. Sept, 2h, 19^8, Reed Ul2l8, 
(vel L. sativtun ?) Anmial herb, winged, branched, the branches 
divaricate, pubescent to glabrous; leaves linear-lanceolate, 
acute, minutely dentate toward the apex, glabrous; silicle orval, 
apex obtuse, eraarginatej stigma sessile, capitate, 

170. Raphanus raphanistrum L, Maryland: Canton, on chrome ore 
piles. June 22, 195h, Reed 33ii29. 

171. Raphanus sativus L, Maryland: Canton, on chrome ore piles, 
June 22, 195U. Reed 33360. 

172. Rorippa islandica (Oeder) Borbas, Native of a*eenland and 
Eurasia; adventive from Europe; Anticosti Isl., Ouebec to Michigan, 
south to Nova Scotia, New England, New Jersey and Pennsylvania, 
Maryland: Canton, on ballast. July 6, 190U, C.C.Plitt , 

173. Rorippa sinuata (Nutt,) A.S.Hitchc, Native from western On- 
tario to Washington, south to Michigan, Illinois, Missouri, Okla- 
homa, Texas, New Mexico, Arixona and California, Maryland: Canton, 
on chrome ore piles, June 10, 19^7, Reed 38809. 

17h. Rorippa sylvestris (L.) Bess. Naturalized from Europe; New- 
foundland to Ontario, south to New Brunswick, New England, Virginia, 
Kentucky and Missouri. Maryland: Canton, Jvily 6, 190li, C.C.Plitt . 

175« Sissntibrium altissimum L, ( Norta Britt.) Naturalized from 
Europe, throu^out eastern North America and beyond, Maryland: 
Canton, common on the oyster shell piles. May 22, 19^8, Reed 
UoltLl; on chrome ore piles. July 22, 19$h, Reed 3U03hj wastes, 
Clinton and Holabird Streets. May 22, 19?8. Reed Ii0lil8, 

176. Descurainia sophia (L. ) Webb, ( Sisymbrium L,). Naturalized 
fl-om Europe; Quebec to Washington, south to Delaware, Pennsylvania, 
Illinois, Kansas, Colorado, Utah and California, Maryland: Canton. 
June 1900. C.C.Plitt . 

177. Thlaspi arvense L, Naturalized from Europe; Labrador to 
Alaska, and southward, Maryland: Canton, on chrome ore piles, 
frequent, Oct. 2^, 19514, Reed 3^195, 

CUCURBITACEAE 

BLASTANIA Kotschy & Peyr, Slender twiner; stems sli^tly 
scabrid, ribbed; leaves digitately 3-5-partite, up to 7 cm. long, 
thin, lobes sharply toothed, scabrid-pubescent; petiole pubescent; 
bract foliaceous and resembling a stipule at base of inflorescence, 
ovate-orbicular, narrowed to the base, fimbriate, about 1 cm. longj 
male flowers few and very small on a common axillary peduncle up 
to 2 cm. long, female solitary, subsessile or short pedicellate; 
fruit about 2-seeded, globose, smooth, 1-1,5 cm, in diameter. 



3^6 PHYTOLOGIA Vol. 10, no. 5 

178, Blastania fimbristipula Kotschy & Peyr. Native of Togoland, 
West Africa to eastern Sudan, Abyssinia, Angola, east and south 
Africa; Arabia and India. Maryland: Canton, on chrome ore piles. 
Sept. 2h, 1958. Reed U1236. Flowers creary-irtiite; fruit scarlet, 
cherry-like . 

179, Citrullus vulgaris Schrad. (Watermelon). Maixr varieties of 
these have been collected on various chrome ore piles, in Canton, 
between 1953 and I960. 

CTPEEIACEAE 

180, Cyperus fuscus L, Adventive from Europe; Massachusetts to 
western New York and Virginia. Maryland: Canton, June 19, 1900; 
July 20, 1901; Aug. 19, 1902. Oct. 1, 1902. All collected by Plitt. 

181, Cyperus globulosus Aubl. Native of tropical America; Florida 
to Texas, north to Virginia, New Jersey, Pennsylvania, Missouri and 
Oklahoma. Maryland: Canton, on chrome ore piles. Sept, 19, 1958, 
Reed la325. 

182, Cyperus iria L, Native of the warmer parts of Eurasia; Florida 
to Texas, north to southeastern Virginia and Maryland, Maryland; 
Canton, on chrome ore piles, Aug. 1959, Reed UU366; Oct, 1959, 
Reed l;5705, Virginia: Newport News, on chrome ore piles, Nov, 1959, 
Reed 15882. 

183, C3rperus rotundus L, Naturalized from Eurasia; Florida to 
Texas and Mexico, north to Virginia and southern New York. Mary- 
land: Canton, Sept. 27, 1953. Reed 32723 and 32758; July 17, 1958. 
Reed U1019; Sept. 6, 1958. ReeTTT292; Sept. 19, 1958. Reed U1339, 
jfenganese ore piles, Aug. 1F7T959. Reed UU381; July 15, I960. 
Reed 16737. Virginia: C&O RR at Port of Newport News. Aug, 7, 1959. 
Reed UliOUO and liU077. 

18U, Cyperus strigosus L, Florida to Texas and New Mexico, Cali- 
fornia, nor-tti to New England, southwestern Quebec, Michigan, Wis- 
consin, Minnesota, Nebraska and Washington. Maryland: Canton, on 
chrome ore piles. Sept. 1, 195U. Reed 31^382; Sept. 19, 1958. Reed 
la323 and la333. 

185. Cyperus polys tachyos Rottb. var. texensls (Torr.) Fern, 
Tropical Airerica to Argentina; Florida to Texas and Mexico, north 
on the coastal plain to Cape Cod, Massachusetts and inland north 
to se Mtssouric Maryland: Canton, on manganese ore piles, Aug, 
1959, Reed lUi380; Oct. 1959. Reed li5698, 

186, Cyperus compressus L. Widely disytbuted in Old World; throu^ 
tropical America to Brazil; Florida to Texas, near the coast, north 
to Delaware and Maryland, inland to Missouri. Maryland: Canton, 

on manganese ore piles. Aug. 1959. Reed ll;377; Oct. 1959. Reed 
U5695. 



196U Reed, Flora of chrome & manganese ore piles 3U7 

KTLLINGIA ROTTB, Annual or perennial sedges, with slender tri- 
angular culms, leafy below and with 2 or more leaves at the summit 
forming an involucre to the strictly sessile, sinple or conpound 
dense head of spikelets; spikelets numerous, corrpressed, falling 
away from the axis of the head at natixrity, consisting of only 
3 or U scales, the 1 or 2 lower ones small and eit^jty, the middle 
one fertile, the upper empty cr staminatej joints of the rachis 
wingless or narrowly winged; scales 2-ranked, keeled; perianth none; 
stamens 1-3; style 2-cleft, deciduous from the summit of the achene. 

187. Kyllingia brevifolia Rottb« West Indies, southern United 
States, tropical continental America and Old World Tropics; common 
on Pacific Islands; this similar to specimens from Pitcairn Isl. 
in IE. Maryland: Canton, on chrome ore piles. Oct. ih, 19^8, Reed 
lail^ Nov. 23, 1958. Reed la7U3. 

188. Sclrpus americanus Pers. Native of Eurasia; Florida to Texas, 
south to South America, west to Pacific, north to Newfoundland. 
Maryland: Canton, in ditches near oyster shell piles. July 19, 
1958. Reed U0877. 

189. Scirpus hystrix Thunb, Native of South Africa, Natal. Vir- 
ginia, Newport News, on chrome and iron ore piles, Aug. 1959. Reed 
UU059 (US); Nov. 1959. Reed U586h. Tufted annual, 8 cm. high or 
less; leaves as long as the flowering stems, flat above, roumded 
below; spikelets few to several in a cluster, the bract at base of 
cluster leaf -like; spikelets bristly with the green sonswhat re- 
flexed tips of the bracts; bracts widely ovate, green, sometimes 
with a reddish tinge, cuspidate; stamens 2, the anthers small, at 
maturity many times shorter than the filament; style branches 3, 
divided almost to the base; fruit sharply 3-angled, greenish- 
black, the posterior face largest. 

DICHONIRACEAE 

DICHONIRA Forst, Small prostrate or creeping herbs, rooting at 
the nodes, with stalked roxinded or reniform leaves, and axillary 
5-partite flowers; carpels 2, free, each 1- or 2-celled, with 1 or 
2 ovoles to each cell, the 2 styles arising between the carpels; 
fruits of fruitlets, usually 1-seeded. Ihis genus is sometimes 
placed in the Colvolvulaceae . 

190. Dichondra repens Forst. Native of Australasia and almost cos- 
mopolitan to the warm regions of the world, Maryland: Canton, on 
chrome ore piles. Nov. 1958. Reed U1726, A small perennial pro- 
strate, slender creeping herb, with small roundish, silky leaves, 
1-2 cm, broad; flowers small, pale white or yellow, with 5 calj^ 
and 5 corolla parts; sepals oblong, 2-U mm. long, the corolla 
hardly longer, with 5 oblong lobes twice as long as the tube; 
fruitlets 2, each with 1 obovoid seed, or only 1 developing; fruit 
a capsule. 



3U8 PHYTOLOGIA Vol. 10, no. $ 

nrSPHANIACEAE 

DTSPHANIA R.Br. Herbs, low, glabrons, with small alternate 
leaves without stipiiles; flowers small, in axillary or terminal 
clusters, with the inner ones often female; perianth of 1-3 
■vrfiite segments, stalked, with a broad limb; stamens 1-3 J styles 
1-2, short and thin; fruit 1-deeded, the pericarp closely ad- 
herent to the seed. TMs genus is related to Scleranthus, 
•vdiich has opposite leaves and typical sepals and is placed by 
Pax next to that genus in the Caryophyllaceae; it is more 
closely related to Chenopodium and thus links the families 
together. The genus has U or 5 species in Australia. 

191 • Dysaphania plantagineHa. F. v. M. Native of western 
Australia, Maryland: Canton, on manganese cxre piles. July 
1959. Reed U381i3. (Det. by L.B.Smith). 

EQUISETACEAE 

192. E quisetum arvense L, Eurasia; North America, south to 
Virginia, Missouri, Nebraska, Colorado and northern California. 
In Canton, between the chrome ore piles. Jiine 1957. Reed 38797; 
in wastes in Canton. Oct. 19^7. Reed 39lil8 and May 1958 » Reed 

uom6. 

EDPHCRBIACEAE 

193. Acalypha virginica L. Georgia to Texas, north to Massachu- 
setts, etc. Weeds throughout the Canton area. Some have green 
bracts (Reed 35208), others have red bracts ( Reed 35210), 

CHROZOPHCEIA Neck. Plants monoecious; MEiIe flowers: calyx 
5-partite, densely stellately hairy, lobes ovate, petals 5, 
raerabraneous, equalling or shorter than the sepals; disc in- 
conspicuous, of 5 glands or connate into a ring; stamens 5-15, 
more or less in 1-3 series; filaments connate into a minutely 
pubescent tube; anthers of outer series subsessile; pistil 
none. Female flowers: calyx segments narrow linear; petals 
narrow, sometimes small or setaceous, or none; disc glandular, 
alternating with the petals or disc somewhat tumid; ovai-y 
3-chambered, with a single ovule in each chamber, stellately 
hairy; styles 2-fid, covered with distinct papillae; capsule 
someidiat fleshy, splitting into 3 cocci, stellately haiiy; 
seed subglobose, with a woody testa, sometimes rough; endo- 
sperm fleshy, copious; enihryo straight; cotyledons subqua- 
drate. Branching herbs, often spreading, covered with stellate 
hairs or hispid; leaves alternate, petioled, ovate, often with 
undulate margins; stipules small, setaceous; inflorescences a 
short dense sessile axillary raceme. 



196U Reed, Flora of chrome & manganese ore piles 3ii9 

I9I;, Chrozophora tinctoria (L.) Juss, Native of the Mediter- 
ranean region, Spain, Italy, Greece, France, Syria, Egypt and 
Algeria. Maryland: Canton, on chrome ore piles. July 19^9. 
Reed 13620; Aug. 19^9. Reed hh3S9. 

19^, Croton glandulosus L. var, septentrionalis Muell, Florida 
to Texas, north to Delaware; adventive to New Jersey and Penn- 
sylvania. Virginia: Port of Newport News, weedy. Aug. 7, 1959. 
Reed UI1O37 and l4li080. Also known as far north in Maryland as 
Anne Arundel Co. July 22, 1956. Reed 38U25 and July 18, I960. 
Reed UTOUTj and Caroline Co., on wastes near Denton. Sept. 9, 
1956, R _ eed 381;96, Also collected in railroad yards at Ports- 
mouth, Scioto Co., Ohio ^ere some of the ores from the east 
have been taken. 

196. Croton bonplandianus Baill. Native of Argentina. Mary- 
land: Canton, on chrome ore piles, Sept. 2?, 1953. Reed 327li2; 
July 19, 1958. Reed 10888; July 17, 1958. Reed la003; July 1959. 
Reed ii3628; Aug7T959. Reed Uli36l;; Oct. 19^r 'Reed li5702 and 
Iif5?2; Sept. 1959. Reed" T3^77; on pile from Turkey. Oct. Hi, 

1958. Reed hlllS; Sept. 6, 1958. Reed hl302. Some of these piles 
miles apart. Virginia: Newport News, on chrome ore piles, fi'om 
India. Nov. 1959. Reed 15897. 

197. Euphorbia chamaesyce L. (E. prostrata Ait.). Nattiralized 
from tropical America; Florida to Texas and rfexico, north to 
southeastern Virginia, Maryland and Missouri. Maryland: Canton, 
on chroms ore piles. Sept. 27, 1953. Reed 32729 and 328ii8; Oct, 
1953. Reed 32896; June 195U. Reed 33li33; Sept. 1957. Reed 
3932U; Oct. 1959. Reed U58l6. 

198. Euphorbia dentata Michx. New York to Virginia, west to 
Minnesota, South Dakota and V^oming, south to Louisiana, Texas 
and Mexico. Maryland: Canton, on chrome ore piles. Sept, 27, 
1953. Reed 32811; Locust Point, wastes along Railroad. July 

1959. Reed U3828 (US) and U3829 (leaves linear). Virginia: 
Newport News, common in the railroad yards in the port. Aug. 
1959. Reed Ui4069 and UU035. 

199. Euphorbia esula L. Naturalized from Europe; Quebec to 
Alberta, south to Nova Scotia, New England, Pennsylvania, Indi- 
ana, Illinois, Iowa. Nebraska, etc. Maryland: Canton. May 1903, 
C.C.Plitt ; July 190li. C.C.Plitt . (Reed Herb.). 

200. Euphorbia falcata L, Naturalized from Europe; Pennsylvania 
and Chio, south to Virginia and West Virginia. Maryland: Can- 
ton, on chrome ore piles. June 1956. Reed 388I6. 

201. Euphorbia glomerifera Mlllsp, (E. hypericif olia L. ) . Flor- 
ida to Texas and Mexico, West Indies, Bermuda. Maryland: Canton, 
on chrome ore piles. June 19Sh» Reed 33hhO, 



350 PHYTOLOGIA Vol. 10, no. $ 

202, Euphorbia hernarlaefolia Willd, Native of Crete, Mt. 
Olynpus, Asia Mnor (US), Iferyland: Canton, on chrome ore 
piles, Sept. 2?, 1953. Reed 32669. 

203. Euphorbia hirta L , Widely distributed in tropical and sub- 
tropical lands, including southeastern United States. Maryland: 
Canton, on chrome ore piles, Nov. 2, 1953. Reed 3 2990; July 
1959. Reed li3631; Oct. 1959. Reed U570U, 

201;. Euphorbia hyssopifolia (L, ) Small. Native of Florida, the 
West Indies and tropical continental America, Maryland: Canton, 
on chrome ore piles, Sept. 195U. Reed 3U392. 

205. Euphorbia macula ta L, (E, preslii Guss,) Florida to Texas 
smd Mexico, north to New EngTand, New York, etc. Frequent on 
the chrome ore piles in Canton. Sept. 1, 195ii. Reed 3U379 and 
3li38l. 

206. Euphorbia marginata Pursh. Minnesota to Colorado and 
Texas; introduced in wastes in the central and Atlantic States. 
Virginia; Newport News, common in wastes about the port in 
railroad yards. Aug. 1959. Reed 1;U072, 

207. E\3phorbia supina Raf , Southern Canada to North Dakota, and 
southward. A common weed in railroad wastes, on chrome ore 
piles, roadsides, ill both Canton and Newport News, 

208. Euphorbia vermiculata Raf, Gaspe Peninsula to northern 
Michigan, south to Maryland, Ohio, Indiana and southern Wis- 
consin; New ^fexico and Arizona. Maryland: Canton, on dirorae 
ore piles. Sept. 1953. Reed 327lilt; Oct. 1959. Reed U579h. 

209. Mercurialis annua L, Local on wastes and ballast-ground, 
Quebec to Ohio, and southward, Maryland: Canton, on ballast, 
July I90U. C.C.Plitt . (Reed Herb.). 

FAGACEAE 

210. Quercus phellos L, Northern Florida to eastern Texas, 
north to Long Island, New Jersey, southeastern Pennsylvania, etc. 
In Canton this oak has been collected, Aug, I88O, Geo, L , 
Smith . (Reed Herb. No. I83O8), 

GMANIACEAE 

211. Er odium cicutariu m (L.) L'Her, Naturalized from Europe; 
Quebec to Mchigan and Illinois, south to Nova Scotia, New 
England, Virginia, Tennessee, Arkansas, Texas and ^fexico. Mary- 
land: Canton, on chrome ore piles. June 22, 1951^. Reed 33362. 



1961i Reed, Flora of chrome & manganese ore piles 351 

212, Geranium rotimdlfolium L. Native of ETnrope, Russia, Asia, 
Britain and Ireland. Maryland: Canton, on chrome ore piles, 
June 22, 195U. Reed 33366, Annual with wide spreading "branches, 
with small orbicular or reniform leaves, with soft hairs; -Uie 
peduncles rather short, the blades shortly divided into broad 
lobesj the flowers small with entire obovate petals, scarcely 
extending the slightly pointed sepals; carpels hairy, without 
ijrinkles; the seeds dotted, 

(51AMINEAE 

I wish to thank Dr, Jason Swallen and am indebted to Mrs, 
Agnes Chase of the United States National Herbarium for verify- 
ing or identifying the grasses collected in the Canton and New- 
port News areas. Many new genera and species have been found in 
these areas, some of which have already been published in 
Rhodora (Reed, 1951, 56: I78-I8I), 

213, Aegilops triuncialis L, Introduced from Europe; a weed 
in California; adventive in Pennsylvania, Maryland: Canton, 
on chrome ore piles, June 22, 195U. Reed 33379; June 1956, 
Reed 38819; June 1957. Reed 38805. Culms branching and spread- 
ing at the base, 20-liO cm, tall: blades rather rigid, sharp- 
pointed, spreading; spike 3 to u cm. long, 2 or 3 of the lower 
spikelets often reduced, the fertile spikelets 3 to 5; glumes 
with 3 strong scabrous somewhat spreading awns, ij to 8 cm, 
long; lemmas with three rigid unequal awns, 

21I1, Agrostis alba L, Native of Eiirasia; Newfoxmdland to Yukon, 
south to Georgia, etc, Maryland: Canton, on chrome ore piles, 

June 1957. Reed 38802; om manganese ore piles, Aug. 9, 1961. 
Reed 52l50. 

215. Agrostis capillaris L, Native of Europe, (Related to A, 
tenuis Sibth,). Maryland: Canton, on chrome ore piles. Aug, 
1959. Reed Uh362, 

216. Agrostis palustris Huds. Eurasia; Maine to Virginia, 
west to Wisconsin and Illinois, British Colximbia, Vfeshington 
and Idaho, California and Texas. Maryland: Canton, on pumice 
piles, from Islands off Italy, Aug. 20, I963. Reed 65li21. 

217. Alopecurus nyosiiroides Huds, Adventive from Europe; 
Massachusetts to Michigan, south to North Carolina and Kansas, 
Maryland: Canton, on chrome ore piles, June 19%. Reed 3 3381 
and 33397. 

218. Andropogon pertusus (L.) Willd. Introduced from the Old 
World; West Indies; Mississippi. Maryland: Canton, on chrome 
ore piles, Nov. 1958. Reed UI889 (US). 



352 PHTTOLOGIA Vol. 10, no. $ 

219, Andropogon virKinicus L, A coiranon weed from Massachusetts 
to Florida; common between the chrome ore piles at Canton, Oct. 
25, 195U. Reed 35220; Oct. U, 1957. Reed 39563. 

APLIEIA L, Tall slender perennial leafy grasses; stems geni- 
culate, base creeping or decumbent, branched above; leaves 
narrow petioled, vtpper ones spathe-like, raucronate; spikes 
numerous, very small, solitary, simple, on the ends of the 
branches, each enclosed in a membranous peduncled spathe, of 
one joint; spikelets 3, one sessile 2-flowered, bisexual, one 
pedicelled inperfect and one terminal male; sessile spikelets 
on the bulbous base of the rachis; glumes U; stamens 3; anthers 
short; stigmas plumed, free, 

220, Apluda mutica L. (A, arista ta L.) Native of China, East 
Indies, and South Africa, India and Malaya to Hawaii, Mary- 
land: Canton, on chrome ore piles, Oct. iLi, 1958. Reed UII38; 
Oct. 28, 1958. Reed 11350 (US), (Description in CoHett, 
Flora Siralensis, p. 598 sub Apluda arista ta Hack. 1902; in 
Hook., Fl. Brit, India, 7: 150, sub A, varia, 1897). 

221, Aristida adscensionis L, Originally described from 
Ascension Island; a common weed in the American tropics from 
Argentina northward to southern California and Nevada, east- 
ward to Texas, southern Kansas and Missoxiri. Maryland: Can- 
ton, on chroite ore piles. Sept. 1953. Reed 32752 and 32826. 

222, Arthraxon hispidus (Thunb.) Makino, Introduced ffom the 
Orient; Pennsylvania and Maryland to Florida, west to Missouri 
and Louisiana, Maryland: Canton, on iihroine ore piles, Oct, Ih, 
1958. Reed U1128; Cub Hill, Baltimore, in fields. Oct. 1953, 
Reed 33513; Nov. I6, I96I. Reed 53750. 

223, A vena fatua L, Introduced from Europe; rare in easlsern 
United States, Maine to Pennsylvania, Maryland, Missoxrri and 
westward; a common weed on the Pacific Coast, Maryland: Canton, 
on chrome ore piles, June 1951j. Reed 33h26. 

22li. Avena sativa L . Commonly cultivated and escaped. Maryland, 
Canton, on chrome ore piles, June 195U, Reed 33li25 and Nov, 
1958. Reed 11719. 

225, Bouteloua aristidoides (HBK) Griseb, Native of Texas to 
NeTteda, southern California and northern Mexico; Argentina, 
Maryland! Canton, on chrome ore piles. Sept. 27, 1953, Reed 
32727, Annual, erect or spreading, branching; culms slender, 
10-30 cm, tall; blades small and few, in vigorous plants as 
much as 15 cm. long; spikes mostly 8-II1 on a slender axis, re- 
flexed, readily falling, the base of the rachis farming a sharp, 
bearded point; spikelets 2-I4, narrow, appressed; rudiment of 3 
scabrous awns about 5 nim, long, exceeding the fertile floret. 



196U Reed, Flora of chrome & manganese ore piles 353 

226, Bouteloua barbata Lag. Native from Texas and Colorado to 
Nevada and southeastern California and Mexico, Maryland: Canton, 
on chrome ore piles, Sept. 1953. Reed 3282U, Annual, tufted, 
branching, erect to prostrate, often forming mats with ascending 
ends, the culms as much as 30 cm. longj foliage scantj blades 
l-li cm, long, 1-1.5 mm. wide; spikes U-7* 1-2 cm. long; spike- 
lets 25-UO, 2.U-I4 mm. long, nearly as broad j fertile lemma dense- 
ly pilose at least along the sides, usually throu^out, the awns 
from minute to as long as the body, the intermediate lobes sub- 
acute to obtusej rudiment from obscurely to conspicuously bearded 
at summit of rachllla joint, cleft nearly to the base, the inter- 
mediate lobes brbad, eubouctillate, the awns of rudiment and 
fertile lemma reaching about the same hei^t, a second nidiment, 
hroad and a>mless, often developed, 

227, Brachlaria distichophylla (Trin,) Stapf. Native of West 
Tropical Africa, where it is a common wayside grass, loosely tuft- 
ed, occasionally half -creeping and forming a carpet, Virginia: 
Newport News, on chrome ore piles. Aug. 1959. Reed hli062 (JB); on 
iron ore piles, Nov. 1959. Reed li5863. Annual, 12-30 cm. tail, 
culms very slender, terete, geniculate, usually ascending from a 
decumbent or prostrate stem, rooting at base, often much branched 
below, 5- to many-times noded, the uppermost internode usually 
very long, pubescent, rarely glabrous; leaf-sheaths more or less 
herbaceous, rather tight, those supporting a branch slipping off 
the internode and rolling round the branch, finely striate, more 
or less pubescent, sometimes densely so, or finely hirsute, 
rarely glabrous; ligules reduced to a narrow cilioate rim; blades 
lanceolate or linear -lanceolate from a suddenly contracted 
rounded base, gradually tapering from very low down to an acute 
point, 2.5-U cm. long, rarely longer, rather stiff and somewhat 
succulent, green, glabrous or more or less pubescent, margins 
markedly cartilaginous and usually cuspid downwards, spinulosely 
ciliolate to serrulate, primary lateral nerves 3-U on each side, 
very fine and differentiated from the numerous and close second- 
ary nerves only below, midrib very slender; panicle at length 
long-exserted, secund, 3.5-7 cm. long, of U-8 obliquely spreading 
or almost horizontal distant solitary slender spike-like mostly 
single racemes; common axis very slender, terete, terminating 
with a spikelet; racemes straight or slightly curved, the lowest 
1-3 cm. long, gradually decussing upwards, sinple or very rarely 
slightly conpoimded at base with very short and scanty secondary 
racemes; lower florets neuter; upper floret male and female, 
slightly shorter than the lower, or equalling it; glumes ver^ 
unequal. 

228, Brachiaria erucaeformis (J.E.Smith) Griseb. in Ledeb, Na- 
tive of Old World, occasionally cultivated in grass gardens. 
Maryland: Canton, on chrome ore piles. Sept. 1953. Reed 3^763; 
Sept. 1958. Reed I4I216; Oct. U, 1958. Reed lUlhl (US). Spread- 
ing annual, with rather delicate erect racemes and pubescent 
spikelet s 2.5 mm. long. 



3^U PHYTOLOGIA Vol. 10, no. $ 

229. Brachiaria subguadripara (Trin.) Hitchc. Asiaj Mariana 
Islands; occasionally planted in southern Florida. Maryland: 
Canton, on chrose ore piles, Sept. 1958. Reed 41296 j Sept. 194S. 
Reed 41318 (US); Nov. 1958. Reed 41900; on nany chroae ore piles 
iirAug. 1959. Reed 44367. Creeping leafy perennial; culas 25-60 
CBU long; blades flat, 5-10 ca. long, 4-S anu iride; racemes 
mostly 3-5, spreading, rather distinct; spikelets 3.5 to 4 ««• 
long, elliptic, glabrous. 

230. B roams comutatus Schrad. Maryland: Canton, on chrome ore 
piles. June 22, 1954. Reed 33423. 

231. Bronms rigidus Roth. Maryland: Canton, on chroiae ore piles. 
June 22, 1954. Reed 33376 and 33378. 

232. Broaus tectona L. Maiylatrl: Canton. May 22, 1958. Reed 
40407. 

233. Cenchrus echiaatus L. CoimiDn weed in tropical America} 
South Carolina to southern California; sparingly introduced to 
Hawaii and Malaysia . Maryland: Canton, on chrome ore piles, 
Oct, 1954. Reed 35205. Annual; cuIjk compressed usuallj- geni- 
culate, branching at base, 25-60 cm. long; blades 3-8 am. wide, 
pilose on the upper surface near the base; raceme 3-10 cm. long, 
the burs larger, fewer and less crowded than in C. brownii; 
bur 4-7 vau high, as broad or broader, pubescent, the lobes of 
the involucre erect or bent irorard but not interlocking; spike- 
lets usually 4 in each bur. 

234. Cenchrus. incertus. M.A. Curtis. Native on the Coastal Plain, 
from southeastern Virginia and North Carolina, west to Cali- 
fornia, and south to Mexico, Central and South America, and in 
West Indies; South Africa (weed), Maryland: Canton, on chrome 
ore piles. Sept. 19, 1958. Reed 41332. Det. D.G.DeLisle (See 
Iowa State Joum. Sci. 37(3): 308-316, fig. 15. 1963). 

235. Geachrua longispinus (Hack.) Fern. Ontario, southward through- 
out United States to Mexico, Central America and West Indies; 
Bermuda; locally naturalized ijri western Europe, South Africa and 
Australia. Common in the wastes of Port of Newport News, in 
RR yards, Virginia. Reed 44079 and 44083. Det. D.G.DeLisle (See 
Iowa Stat© Joum. Sci. 37(3) J 294-301, fig. 10. 1963), 

236. Chloris virgata Swartz. Native from Nebraska to Louisiana, 
Texas and s outhem California; also Ohio, Indiana, Maryland to 
Florida; tropical America. Maryland: Canton, on chrome ora piles. 
Sept. 27, 1953. Reed 327U; Oct. 25, 1954. Reed 35239; Oct. 31, 
1957. Reed 394^. 



1961i Reed, Flora of chrome & manganese ore piles 355 

237, Clnna arundlnacea L, Maine and Ontario to Minnesota and 
South Dakota, south to Georgia and Texas, Marylarri : Canton, 
on chrome ore piles, July 15, I960, Reed I16730. 

CCRIDOCHLQA Nees Spikelets flattened, ovate, in 2's or 3's, 
subsessile along a slender rachisj glumes and sterile lemma 
papery, the second gliiine stiffly ciliate; fruit stlpitate, con- 
cavo-convex, awned; annual, with several digitate racemes, naked 
at the base, 

238, Coridochloa cimicina (L.) Nees ex Jacks, Native of southern 
Asia J sparingly introduced in Florida, Maryland: Canton, on 
chrome ore piles, Oct. lU, 1958, Reed U1139, Culms 20-60 cm, 
tallj sheaths hispid; blades 3-8 cm, long, 1,5-2,5 cm, wide, sub- 
cordatej racemes mostly U-8, digitate, sometimes a second whorl 
belowj spikelets about 3 nm, long, the awn of ;the fruit curved, 
about 1 ram, long, 

239, Cynodon dactylon (L.) Pers. Naturalized from Europe; St. P. 
et Miq,, Massachusetts to Michigan, Iowa and eastern Kansas, 
southward, where abundant, Maryland: occasional in Canton on 
chrome ore piles, Sept. 1953. Reed 3 27 8U; Oct, 1953. Reed 3291iij 
Sept. 19, 1958. Reed U1328 and ia321; Nov. 23, 1958, Reed Iil7li2j 
on manganese ore piles, Aug. 9, I96I, Reed 52151. 

2li0, Dactyloctenium aegyptiiun (L.) Beauv, (1812 j Richter made 
same combination in 1890, using the same basii^on). Introduced 
from the Old World Tropics, from North Carolina to Florida and 
Texas; occasional north to Maine, New Jersey, Illinois,; Colo- 
rado, Arizona, California; tropical America, Maryland: Canton, 
common on chrome ore piles since 1953, collected every year, 
becoming weedy. Sept. 1953. Reed 32718 and 32720; Sept. 1958. 
Reed U1335 (US) and Ul3lj3 (USTPJuly 1959. Reed h363U and 
li3837; on manganese ore piles. Oct. 1959. Reed , many collections; 
Sept. i960. Reed U8058; Oct. 1961. Reed 53^5^7 Virginia: Newport 
News, on chrome ore piles. Nov. 1959. Reed U5913, U3892 and 
U5883. 

2UI, Dactyloctenium geminatu m Hook, Native in sandy soils on the 
coast of Zululand, in brackish soils in eastern Transvaal, (Chip- 
pindall. The Grasses and Pastures of South Africa, p. 131, 1955). 
Maryland: Canton, on chrome ore piles. Sept, 19, 1958, Reed 
U336 and 11337; Nov. 23, 1958, Reed 11901 (IB). A stoloniferous 
perennial with culms up to 70 cm. high; spikes 2-3, more rarely 1, 
2.5-6 cm. long; spikelets 3-U mm. long, 3-5-flowered; keel of 
the lemma and the rachis smooth; lemmas usually awnless; leaves 
glabrous or with a few tubercle-based hairs. 



356 PHYTOLOGIA Vol. 10, no, 5 

2U2, pigltarla longlfolla (Retz.) Pers. Tropical regions of 
the Old World J introduced from Tropical America into southern 
Florida, Maryland; Canton, on c|jrome ore piles. Oct, 19^8, 
Reed Ull3il. 

2li3, Dlgitaria ischaemum (Schreb.) Schreb. ex Muhl, Virginia: 
Newport News, Common ob chrome ore piles. Aug, 19^9. Reed hhO$9i 
on iron ore piles, Nov, 19?9, Reed U?858, 

2Ui, Digitaria sanguinalis (L.) Scop, Ifeiyland: Canton, common 
on chrome ore piles, many collections between 19?3 and 1960j 
on manganese ore piles, Oct, 19^9. Reed U5693, 

2U5» Digitaria adscendens Henr, Native of tropical regions of 
the World, ffajryland: Canton, on manganese ore from the Tamatogo 
Maru from India, Clinton Street, Aug. 20, 1963, Reed 651t09 (US), 

DINEERA Jacq, Inflorescence of 2 to many one-sided spikes, or 
spike-like racemes, much reduced so that the spikelets are in 
small cludters on the central axisj glume acuminate or tapering 
into awns, as long as or longer than the rest of the spikeletj 
rachis of all or of the spikes short and greatly reduced; ligule 
a membrane, 

2li6. Dinebra retroflexa (Vahl) Panzer. Native of Bechuanaland 
and the Transvaal. (Chippindall, I.e., pp. 18^-186, f. 160, 1955). 
Maryland: Canton, on chrome ore piles. Oct, lii, 1958, Reed Ull55» 
Annual, with culms up to 80 cm, tallj leaf-blades expended, 
glabrous or loosely hairyj ligule a membrane; inflorescence 8-30 
cm, long, of nuirerous short spikes on a central axis; the flat- 
tened rachis of ftie spikes often short or greatly reduced, especi- 
ally vrpwards, so that the spikelets are more or less clustered 
on the central axis and the inflorescence is narrow, dense and 
spike-like; spikelets 5-8 mm, long, 2-3-f lowered, glabrous; 
glumes about equal, usually much longer than the r est of the 
spikelet, acuminate or tapering into short awns; lemmas 3-nerved, 
awnless, 

2U7, Echinochloa colonum (L.) Link, Adventive and naturalized 
from the Old World; Florida to Texas and Mexico; locally north to 
New England, Ohio, Pennsylvania and Illinois, Maryland: Canton, 
rather common on chrome ore piles, collected many times up to 
I96O; on manganese ore piles, Aug. 1959. Reed hU351» Virginia: 
Newport News, on chrome ore piles, Aug, 1959, Reed hh055 and 
Nov, 1959. Reed U5896; on iron ore piles, Nov." 1'959, Reed h5860. 

2I18, Echinochloa crus-galli (L.) Beauv, New Brunswick to Wash- 
ington, south to Florida and California; Eastern Hemisphere, 
Maryland: Canton, common on wastes Ibetween chrome ore piles. 
Aug, 1959, Reed Uli385, 



196U Reed, Flora of chrome & manganese ore piles 357 

2k9» Echlnochloa crus-galll var, ndtis (Pursh) Peterm. Distri- 
buted over the same area as the typical form, and nearly as coin- 
raon, Maryland: Canton, on chrome ore piles, Jiily 19, 1958. Reed 
l;0886j Sept. 6, 1958. Reed lil290j Aug. 1959. Reed Uli386; Locust 
Point. July 1959. Reed h3827. Racemes dense, mostly somewhat 
spreading-fleruousj spikelets awnless or nearly, the awns less 
tiian 3 mm. long; basal sheaths occasionally hirsute, 

250. Echinochloa cms- gal li var, aelayensis (HBK) Hitchc, Mexico 
to Argentina J Oklahoma to Oregon, south to Texas and California, 
Maryland? Canton, on chrome ore piles, Oct. 25, 195h. Reed 35237. 
Differs from var. mitis in having less succulent culms, mostly 
sinple, mere or less appressed racemes, the spikelets less strong- 
ly hispid but papillose, usually green, 

251. Eleusine indica (L.) Gaertn, Naturalized from the Old 
Wcrldj Quebec to Minnesota and South Dakota, and southward, Mary- 
land: Canton, on chrome ore piles. Sept 27, 1953. Reed 32718 and 
3272OJ Oct. 1951i. Reed 35192 and 35l95j Sept, 1958. Reed hl293j 
Nov, 1958. Reed lil72U (depauperate form)j on pumice piles, from 
Island off Italy, Aug. 20, 1963, Reed 651^5, 

ENNEAPOGON Desv, ex Beauv, Spikelets 3-flowered, the first 
floret fertile, the second smaller, sterile, the third rudi- 
mentary; glumes strongly 7-nervedj lemmas rounded on the back, 
firm, the truncate summit bearing 9 plumose equal awnsj palea 
a little longer than the body of the lemma, the keels near the 
margin; slender tufted perennials, with narrow feathery panicles, 

252. Enneapogon desvauxii Beauv. Utah and Texas to Arizona, 
south to Mexico (Oaxaca}7 Peru, Bolivia and Argentina, Maryland: 
Canton, on chrome ore piles, Dct, 1953, Reed 32907 j Nov, 1953, 
Reed 33002, Culms numerous, slender, decumbent- spreading, 20-^0 
cm, tall, the nodes pubescent; blades flat to subinvolute, about 
1 ram, wide; panicle spike-like, gray green or drab, mostly- 2-5 
cm, long, sometimes interrupted below; glumes longer than the 
bodj of flie lemmas, 7-nerved, acuminate, pubescent; lemma of 
first floret (incl, awns) i;-5 iwn, long, the body about 1,5 ™n, 
long, villous, 9-nerved, the awns plumose, except at the apex, 

253. Eragrostis atherstonli Stapf , Native of South Africa and 
south Tropical Africa (n. Cape, Grange Free State, Transvaal, 
SW Africa), (Chippindall, l,c., p. 150, fig. II6. 1955), Mary- 
land: Canton, on manganese ore piles. Sept. 1958. Reed hl33ii ; 
Aug, 1959, Reed Ui383; Oct, 1959, Reed b569U, Perennial, ttifted, 
erect or stoloniferous, forming dense stands; culms up to 70 

cm, tall, simple or branched, straight or repeatedly geniculate 
or prostrate and rooting at nodes, few to many noded, usually 
glabrous, very occasionally with a few hairs; leaf-sheaths 
pallid or purplish, more or less papery, nerves with small 



358 PHYTOLOGIA Vol. 10, no. $ 

glandular dots above the node, and below the collar j collar 
pallid, smooth, or often with a few purplish glandular dots; 
leaf -blade loosely rolled or expanded, some nerves much finer 
than others, seen from below, usually with glandular dots especi- 
ally above the collarj inflorescence 5-20 cm, long sind 2-7 cm. 
wide, usually narrowly ovate in outline, lower branches, and 
often branches higher up, whorled, whorls with a tuft of spread- 
ing hairs; spikelets about 5 mm. long, 1-1.5 mm, wide, usually 
3-5-flowered, silvery-gray to greenish-gray in color; lower 
glume equal or subequal to the lower floret, silvery-gray, 
thinly membranous, acute to acuminate, often wrinkled on the 
back; lemmas acute, dark gray with white, thin, membranous tips; 
anthers about 0.7 mm. long; grains about 0,7 mm. long, oblong, 
slightly grooved on the back, embryo very dark green to blackish, 

255, Eragrostis barrelieri Daveau, Introduced from southern 
Europe; Colorado and Kansas to Texas and California and Mexico, 
Maryland! Canton, on manganese ore piles. July 1959. Reed li3639 
and li36Ui (US); July I960. Reed U6736; Aug. 9, 196l, Reed 5211j7. 
Annual; culms erect or decumbent at base, 20-50 cm, tall, 
branching at base, sometimes with a glandular band below the 
nodes; sheaths pilose at the sumnit; blades flat, rather short, 
2-U mm, wide; panicle erect, open but narrow, 8-15 cm, long, 
the branches ascending or stiffly spreading, few-flowered, 
spikelet-bearing nearly to base, the axils glabrous; spikelets 
linear, usually 12- to l5-flowered, mostly about 1 cm, long and 
1,5 mm, wide; lemmas 2 mm, long or slightly longer, 

256, Eragrostis horizontalis Peter, Native of South Tropical 
Africa, to N, Transvaal, SW Africa and Bechuanaland, ffaryland: 
Canton, Nev^irk Street, on manganese ore piles, Aug, 9, 1961, 
Reed 521It9* Agreeing in most respects with E, atherstonei 
Stapf, but not a distinct perennial and with slender culms; 
the sheaths usually somewhat keeled and conspicuously glandular 
(usually more so than in E, atherstonei ), while the basal 
branches of rVhe inflorescence have a tuft of hair in the axils 
of, cr between the branches, but not in a ring right round, 
(Chippindall, l,c., p, 150, fig. 117. 1955). 

257, Eragrostis ciliflnensis (All,) Lutati, Introduced from the 
Old World; Maine to Washington, south throughout United States 
and West Indies, south to Argentina, (E, megastachya (Koel. )Link, 
in Fernald, 8th Ed., p. 12h), Maryland: Canton, frequent on 
chrome ore piles. Sept, 1953. Reed 32711; Oct. 1953. Reed 32893; 
Oct. 195U. Reed 35201 and 352lII7~Nov, 1958, Reed lil725TTocust 
Point. July"T9^9. Reed U3821, 

258, Eragrostis curvula (Schrad,) Nees, Widespread in South 
Africa; introduced by cultivation, spontaneous in Florida, Texas 
and Arizona; useful in erosion control and revegetation of grass- 
lands in SE United States, Maryland: Canton, on chrome ore piles, 
Sept. 1958. Reed Ul3hl. Culms 60-120 cm. tall, densely tufted. 



196U Reed, Flora of chrone & manganese ore piles 359 

erect, simple or soitietiines branching at the lower nodes; sheaths 
narrow, keeled, glabrous or sparsely hispid, the lower densely 
hairy toxrard the base; blades elongate, involute, attenuate to 
a fine point, arcuate spreading, scabrous; panicles 20-30 cm, 
long, the branches solitary or in pairs, ascending, naked at the 
base, at least the lower densely pilose in the axils; spikelets 
7- to 11-flowered, 8-10 mm. long, gray green; lemmas about 2,5 
mm, long, obtuse or subacute, the nerves prominent. 

259. Eragrostis diffusa Buckl, looming, Idaho, Oklahoma and 
Texas to Nevada, California and Mexico; introduced occasionally 
in eastern United States, Maryland: Canton, on chrome ore piles. 
Sept. 1953. Reed 32750, 32770 and 32785; Oct. 1953. Reed 32908 
(panicle f ewer-flowe-red than typical plants) ; Oct. 195U. Reed 
35206, 3522li, 352li5 A and B, and 35236. More robust than"T7"pec- 
tinacea, usually 30-50 cm. tall, sometimes taller; panicle larger, 
the primary branches bearing appressed secondary branchlets 

with few to several spikelets, the main panicle branches thus 
more densely flowered. 

260. Eragrogtis pectinacea (Michx. ) Nees, Maryland: Canton, 

on chrome ore piles, on manganese ore piles; in wastes at Locust 
Point; common and frequently collected in this area, Virginia: 
Newport News, on chrome ore piles. 

261. Eragrostis pilosa (L.) Beauv. Maryland: Canton, on chrome 
ore piles. Nov, 1958. ^e^d I1I36O; on pumice piles, from Island 
off Italy. Aug. 20, 1953. Reed 651il6, Virginia: Newport News, 
on ore piles, Aug. 1959. Reed kh06l and Ul057. 

262. Eragrostis poaeoides Beauv. ex Roem. & Schult, Maryland; 
Canton, on chrome ore piles, Sept. 1953. Reed 32713 and 32792. 
Virginia: Newport News, on chrome ore pHes, Nov, 1959. Reed 
15916. 

263. Eragrostis unioloides (Retz.) Nees in Steud. Introduced 
from southern Asia; Georgia and Florida, Maryland: Canton, on 
chrome ore piles. Oct. 1958. Reed U1130; Nov. 1958. Reed Ia7h6. 
Annual; culms erect or ascending, 20-UO cm. tall; blades flat, 
2-h mm. wide; panicle elliptic, open, 10-50 cm. long, about 
half as wide, the branches ascending; spikelets ovate-oblong, 
strongly compressed, truncate at base, obtuse, 15- to 30- 
flowered, 5-10 mm. long, 3 nm. wide, often pink or purplish; 
lemmas closely imbricate, nearly horizontally spreading, strong- 
ly keeled, acute, 2 mm. long, the lateral nerves prominent; 
palea falling with the lemma or soon thereafter; grain about 
0.7 ram. long. 

26L. Erlochloa gracilis (Fourn.) Hitchc. Oklahoma and western 
Texas to southern California, south through the highlands to 
Mexico, Maryland: Canton, on chrome ore piles. Sept, 27, 1953. 



360 PHITOLOGIA Vol. 10, no. S 

Reed 32728, Annual; claims erect or decumbent at base, LO-lOO 
cm, tall; blades flat, glabrous, mostly "^-lO mm, wide; racemes 
several to numerous, approximate, ascending to slightly spread- 
ing, 2-U cm, long, the axis and rachis softly pubescent, the 
pedicels short-pilose; spikelets U-^ mm, long, rather sparsely 
appressed-pubescent, acuminate, or the glume sometimes tapering 
into an awn-point as much as 1 mm, long; sterile lemma eii^)^; 
fruit about 3 mm, long, apiculate, 

26?, Eriochloa punctata (L.) Desv, Southwestern Louisiana to 
southern Texas; American Tropics, Maryland: Canton, on chrome 
ore piles, Aug, 19?9, Reed hh3^9 and liU368; on manganese ore 
piles, Oct. 1959. Reed Ii5701. Perennial; culms in tufts, usually 
^0-100 cm. tall; blades flat, mostly 5-10 mm, wide, glabPOUBj 
raoenes several, ascending, overlapping, 3-5 cm, long, the axis 
rachises and pedicels scabrous only; spikelets h-5 mn, long, 
lanceolate, rather sparsely appressed-pilose; glume tapering to 
an avm-point about 1 mm, long; sterile lemma a little shorter 
than the glume, enpty; fruit about half as long as the glume, 
with an avm 1 mm, long ot more, 

HAGKELOCHLQA Kuntze Spikelets awnless, in pairs, the rachis 
joint and pedicel grown together, the two clasped between the 
edges of the globose alveolate first glume of the sessile spike- 
let; pedicellate spikelet conspicuous, staminate; freely branch- 
ing anniial with flat blades, the numerous racemes solitary and 
more or less enclosed in the spathes, these usually fascicled 
in the axils of the leaves, 

266, Hackelochloa granular is (L.) Kuntze, Native of East Indies 
and West Africa; in tropics of both hemispheres; introduced in 
North America, Georgia and Florida to Louisiana; New Mexico to 
Arizona, Maryland: Canton, on chrome ore piles, Oct, 1958. 
Reed lill22. Culms 30-100 cm, tall; sheaths papiUose-hispid; 
blades flat, 5-15 cm. long, 3-l5 mm. wide, papillose-hispid, 
cHiate; racemes 1-2 cm. long; sessile spikelets about 1 mm. 
thick; pedicellate spikelets about 2 mm, long, 

26?, Holcus lanatus L, Introduced from Europe, In Canton, on 
chrome ore piles, June 195^4, Reed 33393. 

268, Hordeum hystrix Roth, Introduced from Europe; Utah to Brit- 
ish Columbia, Arizona and California; adventive in Massachusetts, 
New Jersey and Pennsylvania, Maryland: Canton, on chrome ore 
piles, June 195h. Reed 33396; June 1957. Reed 38806. Annual; 
culms freely branching and spreading or geniculate at base, 15- 
UO cm, tall; sheaths and blades, especially the lower, more or 
less pubescent, the auricle wanting; spike erect, 1,5-3 cm, long, 
10-15 mm, wide, the axis usually not readily breaking; glumes 
setaceous, rigid, nearly glabrous to scabrous, about 12 ram, long; 
lemma of central spikelet 5 nun. long, the awn somewhat longer 
than the glumes; floret of lateral spikelets reduced, short-awned. 



196U Reed, Flora of chrome & manganese ore piles 361 

269, Hordeum leporlnmn Link. Introduced ft-om southern Ehirope; 
Massachusetts to Georgia; Vancouver Island and Washington to 
California, Utah and Texas. Maryland: Canton, on chrome ore 
piles. June 19%. Reed 33380, 33ii03, 33iiOU and 33h09. Annual; 
Taranching at base, spreading; sheaths glabrous, blades pilose 
to glabrous; auricle at base of blade well-developed; spUce 5-9 
cm, long, often partly enclosed by the inflated uppermost sheath, 
the rachis internodes mostly 3 mm. long; glumes of the central 
spikelet lanceolate, 3-nerved, long-ciliate on both margins, the 
nei^es scabrous, the awn 2-2,5 cm, long; floret 1-1,2 cm, long, 
raised on a rachilla segment 1 mm, long, the awn 3-li cm. long; 
lateral spikelets usually staminate, the glumes much shorter, 
unlike, the inner similar to those of the central one, the outer 
setaceous, not ciliate, the lemma broad, 10-20 mm. long, the 

awn 2-h cm. long. 

270, Hordeum vulgare L, Barley, adventive from grain fields, 
Maryland: Canton, on chrome ore piles, Sept. 27, 1953. Reed 32825. 

ISCHAEMUM L. Sessile spikelets perfect, awned; pedicellate 
spikelets perfect but not always fruitf\il; rachis disjointing; 
raceme 2 to several, digitate or aggregate on a short axis; culms 
branching with flat blades and digitate or flabellate inflores- 
cences with prominent awns, 

271, Ischaeraum ciliare Retg. Introduced ftrom Old World; Panama; 
British Guiana. Maryland: Canton, on chrome ore piles. Nov, 23, 
1958, Reed li3130. Slender much-branched perrenial, with creep- 
ing rooting bases; fertile culms 3O-6O cm, tall; blades flat, 
3-10 cm, long, I4-8 mm, wide; racemes usually 2, 3-5 cm, long, 
green, finally spreading; spikelets about h mm. long; flrit glume 
broadly winged at the summit, smooth across the back, longitudi- 
nally striate above; awn 5-8 mm. long. 

272, Ischaemum rugosum Salis. Introduced firom Old World; Panama, 
Cuba and Jamaica. Maryland; Canton, on chrome ore piles, Oct, lit, 
1958. Reed U1133, 10126^ Ull62; Nov. h, 1958, Reed 11363; Nov, 23, 
1958. Reed li3131 and U3132, Branching annual; culms 0,5-1 m, 
tall; geniculate below; nodes bearded; blades flat, 8-12 mm. wide, 
sparsely pilose; racemes 5-10 cm, long, erect, so closely ap- 
pressed to each other as often to appear like a single spike; 
spikelets 3-U mm, long, obtuse, the awn about 1,5 cm, long; first 
gliime strongly rugose across the back, 

273, Koeleria p hleoides (Vill.) Pers, Introduced from Europe; 
Pensacola, Florida; Mobile, Alabama; Cameron Co,, Texas; Portland, 
Oregon; several places in California; oiltivated in nurseiy plots 
in Belts\'ille, Maryland; and Tucson, Arizona, Maryland; Canton, 
on chrome ore piles. June 10, 1957. Reed 38798. Annual; culms 
15-30 cm. tall, smooth throughout; sheaths and blades sparsely 
pilose; panicle dense, spikelike, 2-7 cm, long, obtuse; spikelets 



362 



PHTTOLOGIA Vol. 10, no. 5 



2-1; mm, longj glunres acute; lemmas short-awned from a bifid apex; 
glumes and lemmas in the typical form papillose-hirsute on the 
back, but commonly papillose only. 

27U, Leptochloa dubia (H.B.K. ) Nees. Southern Florida; CSclahoma 
and Texas to Arizona, south through Mexico j Argentina, Maryland; 
Canton, on chrome ore piles, Sept. 27, 19?3. Reed 32?^. Perennial; 
culms wiry, erect, ^0-100 cm. tall; sheaths glabrous; blades flat 
or sometimes folded or loosely involute, scabrous, as much as 1 cm, 
wide, usually narrower; panicle of few to many spreading or ascend- 
ing racemes, 3-12 cm. long, approximate or somewhat distant on an 
axis as much as l5 cm. long; spikelets 5- to 80-flowered (or in 
reduced srpecimens only 2-f lowered), 5-10 ram. long; lemmas broad, 
glabrous on the internerves, obtuse or emarginate, the midnerve 
sometimes extending into a short point, the florets at maturity 
widely spreading, very different in appearance from their early 
phase, 

275, Leptochloa uninervia (Presl) Hitchc. & Chase, Mississippi 
to Texas; Colorado and New Mexico to Oregon and California, south 
to Mexico; Peru to Argentina; North Carolina and introduced in 
Maine, Massachusetts and New Jersey, IT-irginia; Newport News, on 
chrome ore piles. Aug. 1959. Reed hh0^2 (US); om manganese ore 
piles, Nov, 1959, Reed 15867, 

276, Lolium niultiflorum Lam, Natioralized from Europe; Newfotmd- 
land to Alaska, south to Virginia and California and southward. 
Italian rye-grass, fferyland: Canton, on chrome ore piles, Oct. 
1958 • Reed l4ll7li. Probably the same, though identified and 
labelled as L, multiflorum var. italicum (A.Br.) Beck, June 195li, 
Reed 33lJ-2-iir, from same area, 

277, Lolium perenne L. Newfoundland to Alaska, south to Virginia 
and California, and southward, Maryland: Canton, on chrome ore 
piles, June 1951. Reed 33377, 33385 and 33102; Oct. 1958, Reed 
11176, 

278, Lolium temulentum L, Adventive from Europe; Quebec and 
New England to Minnesota, Missouri and Kansas, south to Gulf; 
Pacific States, Maryland, Canton, on chrome ore piles, June 195U, 
Reed 33383. 

279, Microstegium vimineiim (Trin.) A, Camus. Native of Asia, 
Maryland: Cub Hill, Baltimore Co., in rock garden and fields, 
persistent since I96I. Oct. 19, 1961. Reed 53670 (IB); Nov. I6, 
1961. Reed 537U9; Nov. 1962. Reed 5991?rOct. 20, I963, Reed 
6li721 (SABC), Although Hitchcock and Chase (1951, Man, Grasses 
of U.S., p. 7U8) list this species from Virginia, North Caro- 
lina, Kentucky, Ohio, Tennessee and Alabama, and the var, imberbe 
(Nees) Honda from Berks Co., Pennsylvania and Greenville, 
Virginia, neither Fernald, in the 8th Edition of Gray's Manual, nor 
Gleason, in the New Britton ft. Brown, Illustrated i^.ora mention 
this genus or species, A description and illustration are pro- 
vided in Hitchcock and Chase, fig, 1137, p, 718. 1951. 



196U Reed, Flora of chrome & manganese ore piles 363 

280, Muhlenbergia asperlfolla (Nees Sr Mey.) Parodi. Indiana and 
Alberta to British Colmnbia, south to Texas, California and Mex- 
icoj southern South Americaj New York, Maryland: Frederick, in 
RR yards, Frederick Co, Sept. 12, 1959. Reed U56U2 and h?6h5. 

281. Qryzopsis miliacea (L.) Benth, Native of Mediterranean 
region; introduction in Califomiaj ballast, Camden, New Jersey; 
Philadelphia, Pennsylvania, Maryland: Canton, on pumice piles, 
from Islands off Italy, Aug. 20, 1963, Reed 65U22. 

282. Panicum adspersum Trin. Native of West Indies; introduced 
on ballast at Philadelphia, Pennsylvania; Camden, New Jersey; 
Mobile, Alabama. (Hitchc. & Chase, I.e., p. 682. 19^1), Virginia: 
Newport News, on chrome ore piles. Aug. 1959. Reed Ul;0li6 (US); 
Nov. 1959. Reed 15871. 

283, Panicum di cho tomifl orum Michx, Nova Scotia and Maine to 
Minnesota, south to Florida, Texas, California and Mexico; West 
Indies. Maryland: Csinton, on chrome ore piles. Sept. 27, 1953. 
Reed 327U8; Oct. 195U. Reed 35219; Sept. 1958. Reed I1I3UO; 
Oct. 1958. Reed UII78. 

28U. Panicum capillar e L. Maine to Montana, south to Florida 
and Texas, and westward. Witch-grass. Maryland: Canton, on chrome 
ore piles. Oct. 195U. Reed 35216; Sept. 1957. Reed 39333; Oct. 
1958. Reed mi82; Locust Point, wastes. July 15^97 Reed li3B2h, 

285. Panicum capillare var. occidentale Rydb. P.E.I, and Quebec 
to British Columbia, south to New Jersey, Maryland, Missouri, 
Texas and California. Maryland: Canton, on chrome ore piles. 
Oct. 1957. Reed 35216. 

286. Panicum paludosum Roxb, (As Panicum proliferum Lam, in 
Hooker, Fl. Brit. India, 7: 50. 1897). Native of Pacific Islands; 
Okinawa; India, Ceylon, Assam. Maryland: Canton, on chrome ore 
piles. Nov. 1958. Reed U3133. Perennial; stem 6-9 dm. or more; 
lower nodes spongy, about 1 cm. in diameter; leaves 15-30 cm. long, 
1-2 cm. wide, base broad but hardly cordate; sheaths loose; 
ligule a ridge of hairs; panicle 15-25 cm. long, often nearly as 
broad, lower branches whorled and fascicles, trigonous, scaberu- 
lous; spikelets green, variable in size, terete, palea of glume 
III absent, or minute, or linear, neuter or male, rarely bisexual. 

287. Panicum psilopodium Trin. Native of China, Sikkim, Macao, 
India and Ceylon. Maryland: Canton, on chrome ore piles. Nov. 
1958. Reed h6012. Annual; stems rather slender, simple or branch- 
ed, 3O-6O cm. tall, leafy to the panicle; leaves narrow, acuminate. 



36U PHYTOLOGIA Vol. 10, no. 5 

glabrous, 7-20 cm, long. 0,5-0,8 cm, widej sheaths glabrous or 
hairyj panicle 5-10 (20) cm,, rather conpact or loose, pedicels 
slender, as long as the spikeletsj spikelets about 3 mm, long, 
green or purpulishj glume I ovate, broader than long, one-half 
length of III or shorter, 3-5-nerved, II ovate acuminate, 9-11- 
nerved. III as long, IV oblong or rounded, obtuse, shining dark 
brown, 

288, Panicxim reptans L, Tropical regions of both hemispheres; 
Florida to Texas, Maryland; Canton, on chrome ore piles. Sept, 
1959. Reed h5680j Oct. 1959. Reed h5805 and 15811, Culms ascend- 
ing 10-30 cm, above the creeping basej blades 1.5-6 cm. long, 
U-12 mm, wide, cordate, usually glabrous, clliate on the undulate 
margin at basej panicle 2-6 cm, long, the 3-12 ascending or spread- 
ing racemes 2-3 cm, long, aggregate, the rachis usually pilose with 
long weak hairs j spikelets secund, about 2 mm, long, glabrous, on 
pubescent or pilose pedicels about 1 mm, longj first glume very 
short, truncate or rounded, 

289, Pf-nicum purpurascens Raddi, Throughout tropical America at 
low altitudes J probably in Brazil at an early date from Africa; 
Florida, Alabama, Texas; Oregon, Para Grass, Maryland: Canton, 
on chrome ore piles, Nov, 1958, Reed Ul7lil, Culms decumbent 
and rooting at base, 2-5 m, long, the nodes densely villous; 
sheaths villous or the upper glabrous, densely pubescent on -Uie 
collar; blades 10-30 cm, long, 10-15 mm, wide, flat, glabrous; 
panicle 12-20 cm, long, the rather distant subracemose densely 
flowered branches ascending or spreading; spikelets subsesslle, 
3 mm, long, elliptic, 5-nerved, glabrous; fruit minutely trans- 
versely rugose. 

290, Panicum ramosum L, Tropical Asia; cultivated as a bird 
feed; North Carolina to Florida, Arkansas and Lotiisiana, Mary- 
land: Canton, on c hrome ore piles. Sept, 1958, Reed lil223. Pedi- 
cels bristly; spikelets glabrous to finely pubescent, about 3 mm, 
long, tawiy or dull brown; otheirwise resembling P, fasciculatum 
var, reticulatum (Torr,) Beal, (New Mexico and Arizona; Mexico), 

291, Panicum scoparium l,.am. Native from Massachusetts to Florida,