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Project Name Data (Last Modified On 11/19/2012)
 

Flora Data (Last Modified On 11/20/2012)
Genus PIPER L
PlaceOfPublication Gen. P1. ed. 1. 333. 1737.
Description Shrubs or trees, occasionally subherbaceous, rarely scandent, often nodose; leaves alternate, simple, with rare exceptions entire, a few species peltate, frequently with one side attached to the petiole at the base several mm. lower than the other, smooth to rugose or bullate above and lacunose beneath, glabrous or sparsely to densely hairy, often scabrous, frequently sparsely to densely glandular-dotted, palmately nerved or more commonly pinnately nerved from the lower half or two-thirds, or throughout, usually somewhat venulose with finer cross-connecting and anasta- mosing nervules; petiole of various lengths, rarely lacking, commonly more or less vaginate-grooved at least at the base and often to the blade, the margins of the groove wing-margined, the wing very narrow or apparently lacking, or broad and conspicuous; spikes leaf-opposed, cylindrical or rarely globose or subglobose, 1-10 mm. or more thick and up to 50 cm. or more long; peduncle from less than 5 mm. up to several centimeters long; rachis commonly somewhat ridged between the flowers, the ridges low and smooth or papillate to fimbriate; flowers sessile and perfect (American species) or imperfect (Old World species) ; usually densely congested, in the axis of variously shaped bracts, the bracts sometimes glabrous but more commonly pilose or fimbriate; stamens 2-5; ovary 1, glabrous or sometimes papillate or pubescent; stigmas 2-4, rounded to filiform, sessile or on a short and thick to elongated and slender style; fruit variously shaped, one-seeded, drupe-like, with a thin pericarp and somewhat hardened endocarp.
Habit Shrubs trees
Key

a. Ovary and fruit stylose ..................-. I. Spp. 1-18 (pp. 5-14)

aa. Ovary and fruit not stylose. b. Leaves scarcely peltate. c. Leaves palmately nerved, or plinerved within the lowermost 5 mm .................... II. Spp. 19-25 (pp. 15-20)

cc. Leaves pinnately nerved. d. The principal branches of the midrib arising from somewhat below the middle to the upper third of the blade. e. Leaves mostly 20-25 cm. or more long X 10-20 cm. or more wide, base equi- or unequilaterally lobed (shallowly so in a few species) .-............................................... III. Spp. 26-34 (pp. 20-26)

ee. Leaves mostly smaller or, if large, not lobed at the base ...... IV. Spp. 35-82 (pp. 30-60) dd. The midrib more or less equally branched to the upper fourth or throughout .V.............................................. V. Spp. 83-93 (pp. 61-69)

bb. Leaves distinctly peltate .-............................................... VI. Spp. 94-96 (pp. 70-71)

I. KEY TO SPECIES HAVING STYLOSE OVARIES AND FRUIT (In a few species included here the styles may be very short or easily deciduous. Such question- ably stylose species have been also included in the non-stylose section of the key.) a. Leaves rugose-bullate ...................- ........ 1. P. PESARESANUM

aa. Leaves not rugose-bullate. b. Leaves peltate -----------------------------------------------------....95. P. MAXONII

bb. Leaves not peltate.

c. Spikes 15-20 cm. or more long; leaves 10 X 20 cm. or more; styles short.

d. Petioles, twigs, etc. more or less warty-verrucose 3 0. P. IMPERIALE

dd. Petioles, twigs, etc. not warty-verrucose.

e. Twigs glabrate; nerves 6-8 on each side f. Leaves elliptic, base oblique, cordate-auriculate .2. P. ALBOPUNCTU- LATISSIMUM

ff. Leaves ovate, base not oblique or auriculate 34. P. PITTIERI

ee. Twigs densely brown-hairy; nerves 5-6 on each side 31. P. TARDENS cc. Spikes and leaves much smaller.

d. Spikes mostly 3 cm. or more long when mature. e. Leaves pinnately nerved throughout; plants entirely glabrous; fruit tetragonous; bracts cupulate, glabrous 3. P. DARIENENSE

ee. Leaves pinnately nerved below the upper third (or sometimes throughout in P. phytolaccaefolium).

f. Young twigs and/or leaves hairy.

g. Spikes 6-8 cm. long; peduncle 1.5-2 cm. long .................. 4. P. LINCOLNENSE

gg. Spikes 3-4 cm. or less long; peduncle 0.5-1 cm. long.

h. Leaves ovate, 7-12.5 cm. wide; twigs retrorsely hirtellous .......................... 5. P. PERVENOSUM

hh. Leaves lance-elliptic, 3.5-5 cm. wide; twigs villous ...... 6. P. PERULIGERUM

hhh. Leaves elliptic-obovate or obovate, 7-12 cm. wide . 7. P. EXIGUICAULE

ff. Entirely glabrous.

g. Leaves ovate, base rounded; spikes 5 cm. or more long 8. P. CRASSINERVIUM var. TOCOTANUM

gg. Leaves not ovate, base acute to acuminate.

h. Leaves lance-elliptic; spikes 2-4 cm. long 9. P. PHYTOLACCAE- FOLIUM

hh. Leaves obovate, strongly revolute 10. P. MACROPUNCTATUM

dd. Spikes mostly 2 cm. or less long when mature.

e. Leaves with 6-8 main lateral nerves.

f. Leaves 10 cm. or more wide, nerves beneath puberulent 11. P. LATIBRACTEUM

ff. Leaves less than 5 cm. wide, nerves beneath glabrous 12. P. CANDELARIANUM

ee. Leaves with 3-5 main lateral nerves.

f. Leaf base 5-10 mm. shorter on one side, glabrous 13. P. MAGNANTHERUM

ff. Leaf base essentially equilateral, or if unequal more or less pubescent.

g. Young twigs and/or leaves hairy.

h. Leaves more or less long-ciliate, base acute 14. P. GARAGARANUM

hh. Leaves not ciliate, base more or less cordulate.

i. Twigs, petioles, etc. minutely hirtellous 5. P. PERVENOSUM

ii. Twigs, etc. densely villous.

j. Leaves about three times longer than wide, oblong- lanceolate 15. P. PUBISTIPULUM

jj. Leaves about 2 times longer than wide, elliptic . 16. P. COLON-INSULAE 

 gg. Young growth glabrous. h. Leaves mostly 6 cm. or more wide.

i. Leaves broadly elliptic, abruptly short-acuminate, subrugose ............................................. 17. P. DISTIGMATUM

ii. Leaves ovate, gradually acuminate, not rugose ............ 18. P. DAVIDSONII

hh. Leaves mostly narrower ............................................. 9. P. PHYTOLACCAE- FOLIUM

II. KEY TO SPECIES WITH PALMATELY NERVED LEAVES
(The nerves may coalesce within the lowermost 5 mm.)
a. Fruit papillate, with a prominent, glabrous, apical disc .--..... 19. P. RETICULATUM

aa. Fruit lacking any evident apical disc.

b. Leaf base cordate.

c. Leaves 9- to 11-nerved -------- --- --- 20. P. MARGINATUM

cc. Leaves 5-nerved --------------21. P. ARISTOLOCHIAE-
FOLIUM

bb. Leaf base rounded or narrowed, scarcely cordate, leaves more or less elliptic.

c. Leaf base inequilateral with one side about 2 mm. shorter; flowers approximated.

d. Petiole vagina margin glabrous; leaves lanceolate, 3-nerved .......... 22. P. HUMORIGAUDENS

dd. Petiole vagina margin ciliate; leaves lance-elliptic, 5-nerved - 23. P. CONVERSUM

cc. Leaf base essentially equilateral, 5- to 7-nerved.

d. Flowers distant; bracts glabrous ......... 24. P. PINOGANENSE

dd. Flowers more compact; bracts papillate-pilose. 25. P. MEDIUM

III. KEY TO SPECIES WITH LEAVES MOSTLY 20 CM. OR MORE LONG AND MORE OR LESS LOBED AT THE BASE
(The base may be only slightly lobed but none are rounded or acute; the petiole mostly vaginate-
winged to the middle or above.)
a. Leaves rugose or bullate.
b. Leaves scabrid ................-.. 41. P. PERACUMINATUM
bb. Leaves not scabrid.
c. Leaves loosely long-villous; peduncle 1-2 cm. long ............................ 26. P. PARTITICUSPE
cc. Leaves hirtellous to pubescent; peduncle 1-2.5 cm. or more long-- 27. P. COOPERI
aa. Leaves scarcely rugose or bullate.
b. Leaves and stems glabrous ........................................................................ 28. P. HASTULARUM
bb. Leaves and/or stems pubescent.
c. Leaves scabrous . 41. P. PERACUMINATUM
cc. Leaves not scabrous.
d. Leaf margin densely ciliolate, drying thin; spikes scarcely 5 mm.
thick, yellow when dry ...........................................-. 29. P. AURITUM
dd. Leaf margin glabrous or essentially so.
e. Young twigs, nodes and/or petioles and midrib beneath more
or less fleshy-warty .........................................-........ 3 0. P. IMPERIALE
ee. Not fleshy-warty.
f. Stems matted brown-hairy, leaves 5- to 7-nerved.
g. Leaves acute, basal sinus wide, essentially glabrous above... 31. P. TARDENS
gg. Leaves acuminate; basal sinus narrow, nerves pubescent
above 32. P. CAMPANUM
ff. Stems glabrous; leaves mostly 7- to 9-nerved.
g. Leaves glabrous above; petiole vaginate below the middle;
bracts glabrate .............-....-............................ 3 3. P. CARRILLOANUM
gg. Leaves more or less hairy above; petiole vaginate to the
blade; bracts fringed .............-...........-..... .... 34. P. PITTIERI

 IV. KEY TO SPECIES PINNATELY NERVED FROM SOMEWHAT BELOW THE MIDDLE
TO THE UPPER THIRD OF THE BLADE AND WITH THE LEAVES MOSTLY
LESS THAN 20 CM. LONG; IF LONGER, THE BASE IS NOT LOBED.
a. Leaves scabrous.
b. Twigs glabrous.
c. Leaves glabrous on both sides, scabrous beneath, smooth above - 35. P. DAVIDANUM
cc. Leaves hairy beneath, at least along the nerves.
d. Leaves mostly 3-4.5 cm. wide, strongly granular-dotted beneath.. 36. P. MINUTE-
SCABIOSUM
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FLORA OF PANAMA (Piperaceae) 27
dd. Leaves 5-6 cm. wide, scarcely granular-dotted beneath ................- 52. P. STORKII
bb. Twigs more or less hairy, at least when young.
c. Spikes strongly and regularly curved; peduncle longer than the
petiole .. .......................... ............................................................. .... 37. P. ADUNCUM
cc. Spikes not strongly or regularly curved.
d. Stem hairs stiff-hispid, comparatively short; leaves harshly
scabrous; fruit papillate-puberulent at apex ---------------------------------- 38. P. HISPIDUM
dd. Stem hairs retrorse; leaves finely scabrous.
e. Plant scandent; leaf base obtuse or acutish -------------------------------- 39. P. VITABUNDUM
ee. Plant shrubby; leaf base rounded, cordulate .............................. 78a. P. WEDELII var.
RETROHIRSUTUM
ddd. Stem more softly pubescent or villous.
e. Hairs on the midrib beneath erect or at least not upcurved-
strigose.
f. Hairs on midrib beneath mostly up to 1 mm. or more long.
g. Leaves mostly 18-20 cm. or more long.
h. Leaves 6-8 cm. wide, very slenderly acuminate, base
obtuse or cordulate .......................-...- .......-... ...... 40. P. PERSUBULATUM
hh. Leaves mostly 10 cm. or more wide, not slenderly
acuminate, base rounded-cordate ........................... . 41. P. PERACUMINATUM
gg. Leaves scarcely up to 15 cm. long.
h. Leaves broadly elliptic, less than two times longer than
wide .......-...-.................. ...... 73. P. CATIVALENSE
hh. Leaves 2-3 times longer than wide.
i. Leaves rhomboid-elliptic, strongly inequilateral; bracts
glabrous ...-....................... .......... 42. P. ERUBESCENTI-
SPICUM
ii. Leaves lance-elliptic, elliptic, or elliptic-obovate.
j. Petioles scarcely 5 mm. long; stem hairs scarcely
1 mm. long; blades slightly scabrous ........................ 72. P. PSEUDO-
GARAGARANUM
jj. Petioles up to 2 cm. long downward.
k. Blades mostly 5-8 cm. wide, scarcely scabrous
above ...................................-............. 43. P. CULEBRANUM
kk. Blades 3-4.5 cm. wide, harshly scabrous above_. 44. P. VIRIDICAULE
ff. Hairs on midrib mostly scarcely 0.5 mm. long, dense.
g. Leaves less than 5 cm. wide; veins above impressed in age.. 45. P. PERVELUTINUM
gg. Leaves 6-8 cm. wide; veins not impressed ........................ 46. P. VARIITRICHUM
ee. Hairs on the midrib beneath strongly upcurved and appressed
or strigose.
f. Leaves less than 4 cm. wide --------------------------------...-....-......--47. P. CHANGUINOLANUM
ff. Leaves mostly wider.
g. Blades moderately to densely hairy on the upper surface.
h. Peduncle scarcely 5 mm. long; blades villous above and
harshly scabrous; bracts of young spikes not conspic-
uously bristly ...................... 48. P. VILLIRAMULUM
hh. Peduncle 5-10 mm. or more long; blades crisp-hirtellous
or subhispid above, slightly scabrous; young spikes very
short, conspicuously bristly from erect marginal hairs.
i. Leaves mostly narrowed downward, usually only the
longer side rounded at the base ........ ............. 49. P. TABOGANUM
ii. Leaves rounded-cordulate on both sides .................... 50. P. PSEUDO-
CATIVALENSE
gg. Blades sparsely hairy or glabrous, hairy mostly only along
the nerves above.
h. Young spikes not conspicuously bristly because of the
erect bract hairs.
i. Bracts prominently densely yellow-fringed .................. 70. P. LEPTOCLADUM
ii. Bracts very narrowly and sparsely fringed.
j. Nerves beneath spreading-hirsute ..........................-.... 51. P. LUCIGAUDENS
jj. Nerves beneath obscurely strigose .......................... 52. P. STORKII
iii. Bracts glabrous, with a dorsal callus ............................ 78. P. WEDELII
hh. Young spikes about 1 cm. long, conspicuously bristly
because of erect bract hairs ..................-. 49. P. TABOGANUM
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28 ANNALS OF THE MISSOURI BOTANICAL GARDEN
aa. Leaves not noticeably scabrous.
b. Spikes short, scarcely exceeding 2 cm. in length.
c. Stems and leaves glabrous.
d. Spikes less than 1 cm. long; bracts glabrous ..................................- 53. P. WAGNERI
dd. Spikes about 2-3 cm. long; bracts densely ciliate .......................... 54. P. DUNLAPI
cc. Stems and/or leaves hairy.
d. Stems glabrous or sparsely pubescent; nerves beneath appressed-
puberulent ........................................................ 55. P. PALMASANUM
dd. Stems rather densely hairy, at least when young ........................... 16. P. COLON-INSULAE
bb. Spikes longer, mostly 3-5 cm. or more long (or unknown).
c. Stems and leaves essentially glabrous (nerves hirtellous or puberu-
lous in P. carrilloanum and P. aequale var. laurifolium).
d. Peduncle 4-7 cm. long ......................................................... 56. P. REPTABUNDUM
dd. Peduncle scarcely 2 cm. long at most.
e. Leaves ovate.
f. Base mostly acute at the petiole; petiole 1-2.5 cm. long ...... 57. P. CHIRIQUINUM
ff. Base obtuse to cordate.
g. Leaves 10-14 cm. long; petiole 5 mm. long ...................... 65. P. TECUMENSE
gg. Leaves 15 cm. or more long; petiole 1-2 cm. or more long.
h. Nerves beneath puberulous; peduncle 1-2 cm. long ......- 33. P. CARRILLOANUM
hh. Nerves beneath glabrous; peduncle 0.5-1 cm. long ......- 93. P. GRANDE
ee. Leaves elliptic, lance-elliptic, subobovate or obovate.
f. Leaves mostly 2.5-4.5 cm. wide (rarely wider); apex sharp-
to strongly mucronate-acuminate.
g. Leaves scarcely more than 2-2 /2 times longer than wide.
h. Base acutely decurrent, apex strongly mucronate ..........- 58. P. TENUIMUCRO-
NATUM
hh. Base rounded, obtuse ......................................................... 59. P. SEPTUPLINERVIUM
gg. Leaves mostly 3-4 times longer than wide, or base not
decurrent.
h. Bracts smooth; leaves not glandular.
i. Leaf base rounded, obtuse; nerves 4-5 on each side.... 59. P. SEPTUPLINERVIUM
ii. Leaf base narrowed, acute to cuneate; nerves 3-4 on
each side ......................................................... 60. P. SUBQUINQUENERVE
hh. Bracts conspicuously yellow-fringed; leaves more or
less glandular-dotted.
i. Leaves drying dull, yellow ............................................ 61. P. INFRALUTEUM
ii. Leaves drying glossy, green .......................................... 80b. P. HIRTELLIPETIOLUN
var. HARVEYANUM
ff. Leaves larger, mostly 5 cm. or more wide.
g. Leaves nerved from about the lower third, sharply ab-
ruptly acuminate ...................... 62. P. CRICAMOLENSE
gg. Leaves nerved to about or above the middle.
h. Leaves mostly 4- to 5-nerved on each side of the mid-
rib (or sometimes 6 in P. aequale).
i. Leaves obovate, revolute, apex mostly obtuse .............. 10. P. MACROPUNCTATUM
ii. Leaves oblong-elliptic-subobovate, apex a b r u pt 1 y
blunt-acuminate ............................................. 63. P. AEQUALE
iii. Leaves elliptic, apex gradually acuminate.
j. Leaves, petioles, etc. strongly glandular-dotted 64. P. SAMBUANUM
jj. Leaves, etc. not glandular-dotted ............... ............... 65. P. TECUMENSE
hh. Leaves 7-nerved on each side of the midrib, oblong-
ovate, 10-13 cm. wide ..................................................... 66. P. CRASSISPICATUM
cc. Stems and/or leaves more or less hairy (minutely and obscurely so
in P. caflibracteum).
d. Leaves nerved from the lowermost I/4 or 1/3 67. P. BOGOTENSE
dd. Leaves nerved mostly to about the middle or above.
e. Leaf base with one side strongly auriculate and covering the
petiole (auricle about 1 cm. or more long).
f. Pubescence strongly dimorphic; leaves 6 cm. or more wide... 68. P. SPERDINUM
ff. Pubescence not dimorphic; leaves less than 5 cm. wide.. 69. P. PERBREVICAULE
ee. Leaf base may be cordulate but scarcely auriculate.
f. Leaves hairy on the upper surface, at least along the mid-
rib above the very base.
g. Leaves 10 X 25 cm. or more ............................................... 85. P. COLONENSE
gg. Leaves mostly smaller.
h. Pubescence on the upper leaf surface restricted to the
nerves.
i. Nerves beneath glabrous; spike 2-3 cm. long ............ 59a. P. SEPTUPLINERVIUM
var. SUBPUBINERVIUM
ii. Nerves beneath obviously hairy .................................... 70. P. LEPTOCLADUM
hh. Pubescence not so restricted.
i. Leaves 6-7 cm. wide, scarcely 3 times longer than
wide; spikes straight.
j. Crisp-hirtellous or subhispid above; young spikes
short, bristly4............................................................... 49. P. TABOGANUM
jj. Sparsely long-villous above, glabrescent; young
spikes not noticeably bristly.
k. Peduncle up to 6 cm. long; stem and leaf hairs
strongly dimorphic-................................................ 71. P. DIMORPHOTRICHUM
kk. Peduncle much shorter; hairs not obviously
dimorphic.
1. Apex gradually acuminate, blades 15-20 cm.
long ............................................... 72. P. PSEUDO-
GARAGARANUM
11. Apex abruptly acuminate, blades 12-14 cm.
long ............................................... 73. P. CATIVALENSE
ii. Leaves mostly narrower, 3-4 times or more longer
than wide, rugescent, appressed-pubescent above;
spikes commonly curved.
I. Nerves mostly 7-8 on each side; leaves mostly 3-5
cm. or more wide ............................................... 74. P. PSEUDO-
LANCEAEFOLIUM
jj. Nerves mostly 4-5 on each side; leaves mostly
2-3.5 cm. wide ............................................... 75. P. LINEARIFOLIUM
ff. Leaves glabrous above.
g. Fruit and ovary papillate-pubescent.
h. Petioles scarcely 10 mm. long.
i. Spikes 2-3 mm. X 7-8 cm.; plant vine-like .............. 39. P. VITABUNDUM
ii. Spikes 3-5 mm.
X
4-5 cm.; plant shrubby .............. 76. P. PANAMENSE
hh. Petioles up to 3 cm. long; spikes 8-10 cm. long .......... 77. P. CALLIBRACTEUM
gg. Fruit glabrous, or unknown.
h. Stems glabrous or essentially so.
i. Leaf nerves beneath thinly strigose; bracts glabrous .... 78. P. WEDELII
ii. Leaf nerves beneath obscurely puberulent; bracts
fringed.
I. Leaves 7-10 cm. wide ............................................... 79. P. VERRUCULAE-
PETIOLUM
jj. Leaves less than 6 cm. wide ...................................... 65. P. TECUMENSE
hh. Stems obviously pubescent, at least when young.
i. Leaf base narrowed, acute or obtusish, the sides
nearly equal.
j. Nerves beneath glabrous; spikes about 4-5 cm.
long ................... 80. P. HIRTELLIPETIOLUM
jj. Nerves beneath pubescent; spikes 5-6 cm. long.... 80a. P. HIRTELLIPETIOLUM
var. TAPIANUM
ii. Leaf base with one or both sides rounded-obtuse, or
the longer side mostly cordulate.
j. Bracts conspicuously fringed; spikes 3-4 mm.
thick.
k. Lateral nerves loop-connected upward, not con-
tinuing toward the apex ................. 81. P. SUBCAUDATUM
kk. Lateral nerves not strongly loop-connected up-
ward, continuing unbroken toward the apex ...... 43. P. CULEBRANUM
jj. Bracts essentially glabrous; spikes 7-8 mm. thick .... 82. P. GONOCARPUM

 

VI. KEY TO SPECIES WITH PELTATE LEAVES
a. Petiole 7 cm. or less long, vaginate to the middle or above; mostly with
3 nerves on each side of the midrib above the base.
b. Leaves strongly bullate-rugose; young growth more or less rusty-
tomentose -94. P. HARTWEGIANUM
bb. Leaves not bullate-rugose, not hairy -95. P. MAXONII
aa. Petiole 12 cm. long, vaginate at base only; with 2 nerves on each side
of the midrib above the base -96. P. VERAGUENSE

 
 
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