, 
THE PIPERACEAE OF PANAMA 
By WILLU.K Taa .lt A 8E 
l1i1'RODUCTIOli 
No botanist who visits Central America can fail to take an interest 
in the great number of species of Piper and Pepe'fVJ'l'nia which he 
finds everywhere in this region. The family is most abundantly 
developed in the wet forests of the Atlantic watershed, but many 
spedes of Piper and some Peperomias are common in the drier 
forests of the Pacific slope. It is Costa Rica, doubtless, which of 
Central American countries possesses the richest representation of 
the Pepper family, chiefly in the wet forests at middle and upper 
elevations, but in Panama also the family is well reprerented, as 
may be seen by the 138 species here listed. . 
There is no doubt that the real number of species existing in 
Panama is much greater, since no part of Panama except the 
region of . the Canal Zone has been well explored botanically. The 
species of Poth01MrpM are common and widely distributed plants, 
and this is true also of a few species of Piper and Pepero""a. 
Most of the species of the latter genera, however, seem to be very 
limited in their distribution, and not until every secluded valley 01 
Panama has been explored will it be possible to make a complete 
census of the Piperaceae of the country. 
The species of Piper ate mostly low shrubs, and it is easy to 
collect and preserve specimens of them. Most of the Peperomias 
are fleshy herbaceous epiphytes. It is difficult to collect them, and 
even harder to dry specimens, since there is no other group of 
Central American plants which dry so slowly in the press. For this 
reason the species of PepM'Omia are neglected by most collectors and 
inadequately represented in herbaria. 
This paper is based largely upon the collections existing in the 
U. S. National Herbarium. These consist chiefly of material col­
lected by Mr. H. Pittier and Dr. William R. Maxon in 1911 and 
1912, in the course of the Smithsonian Biological Survey of the 
Canal Zone; by Mr. Ellsworth P. Killip in 1918 and 1922; by Dt. 
Maxon in 1923; by Dr. C. V. Piper in 1923; and by the present writer 
inl92$-24. 
1ft 
• 
• 
16 CONl'BIBUTIONS FROM THE NATIONAL HEBBAR!UK 
The collections of 1911-12 were identified by Casimir de Candolle, 
who based numerous new species upon them. The recent collections, 
as well as the earlier ones, were submitted to Doctor Trelease, who has 
kindly prepared the report here presented, which lists all the 
Piperaceae now known from Panama.-PAUL C. STANDLEY. 
SYSTEMATIC 'tREATMENT 
KEY TO Glunz' 
Stigmas 2 to 5 (mostly 3 or 4). 
Spikes axWary. 
Spikes several on a common peduDcle. Plants sutrruUcose. 
I. POTHOKOBPHE. 
Spikes solitary in the leat axii. Plants sulfrutlcose __ 2. SABCOBHACHlS. 
Spikes opposite the leaves, solitary. Plants woody _______________ S. PIPER, 
Stigma 1, often peniclliate. Piants herhaceous _____________ A. PEPEBOlllIA. 
1. POTHOlllOl!.PHE Miquei 
Lea.e distinctly peltste _______________________________________ l. P. peltsta. 
Leaves cordate, not peltate __ n_nnunn ____ nn _____ n ___ 2. P. nmbellata. 
1. Pothomorphe peltata (L.) Miquei, Comm. Phyt. 37. 1840. 
Piper peltatum (" pelaturn ") L. Sp. PI. 30. 1753. 
Peperomw peltata Dietr. Syn. PI. 1: 142. 1831. 
LepI<J"tke. peltatum Rat. Syl •. Teliur. 85. 1838. 
Heckeria peltata Kunth, Ltnnaea 13: 565. 1839. 
'I'tH LOCAUTY: U America calldiore." 
RANGE: Cuba, through the West Indies (the prototype figured by Plumier) 
Jnd northern South America to Mexico on the mainland. 
CHDlIQul: Cerro de la Plata, San Felix, Pitti6r 5160. CAUL ZoNE: With· 
out locality, E,pe181r.e1mer. Balboa, KUlip 3064; StandJetl 25602. Fort Lorenzo, 
Piper 5929. Fort Sherman, 8ta"dle~ 30969. Frijoles, 8taKdle~ 27619. Frijoles 
to Monte Lirlo, Kim", 12117. Gamboa, Standle1l 2B:i2ti. Gamboa to Cruces, 
Plttler 3782. Gat~n, 8taftd/e~ 27257. Juan Mina, Piper 5689. Las Cascadas 
Plantation, Statt4lell 25797. Monte Lido. Mazon 6858. Summit, Stooena 31-4, 
313. RIo Slrr~ Pltti6r 4016. PAKUlA: Rio Tapia, Sta"d/e~ 28062. Rio Tecu­
men. StancUe" 26735. DARIEN: Boca de Cupe, William8 697. 
II. Pothomorphe umbellata (L.) Miquel, Cornm. Phyt. 36. 1840. 
Piper umbellalum L. Sp. PI. 30. 1753. 
Peperomia tlmbeUata Knoth, Syn. PI. Aequln. 1: 124. 1822. 
Lepluflt1l.e, umooUatum Rat. Sylv. Tellur. M. 1838. 
H eckeria umbellala Kunth, Llnnaea 18: 569. 1839. 
TUm LOCALITY: Santo Domingo, Hispaniola. 
RANGE: Through the West Indies; on the continent reaching Brazil Bnd 
Peru to the south and Mexico to the north, in a number of often segregated 
fOrIDs, also (? naturalized) in tropical Asia BDd Africa. 
CHIRIQui: El Boquete, Pi-ttier 3133. 
2. SABCORRACHIS Trel., gen. nov. 
Trailing or climbing, rather succulent, glabrous shrubs. Leaves alterIlJite. 
Spiktc>s solitary in the a::dls. Flowers splcate, perfect, Daked, se8slle. suh­
tended ·by minute concave bracts. Stamens usually 4. distinct, essentially 
, 
• TIlEIJ!ASE PIPIIIIACEAE OP PAN AVA 17 
hJ>pogynoo8, ohort, wllb rounded antbers. Pistil 1, 1"",Ued, wltb a Blogle 
erect ovule i stigmas mostly 4, sessile. Fruit a small 1 seeded drupe, pertly 
tmmeUJCd In and coalescent with the somewhat ~(>8hy rachis at maturity. 
1iype species, 8. mc..:rva. Tret. (Piper lnou'VIf .. Sieber). 
1. S' rcorhaehtl naranJoana (C. DC.) '1 reI. 
Piper .... ra .. Joa .... '" C. DC. Llnoae& 37: 363. 1872. 
TYPE LOCAl,iTt : Naranjo, Costa Rica. 
R.£.KGII:: Costa Rica and Panama. 
DAlIltN: SambO River, Piltier 5562. 
3. PIPER L. 
KEY TO SPECIES 
1. Leaves palmately nerved _____ _______________ ______ ____________ _________ 2. 
!Jeavee pinnately nerved. _____ ______________ • ____ ______________________ 4. 
2. Inner Denes confluent below ___ ________________ ____ ____ 3. P. piDogaDense. 
~erves separate to the base __ __ ___________ ___ ___ _______________________ S. 
8. Petiole short, wingless; leaves greeo ______________ ___ l. P. amllacifoUum. 
Petiole elongate, winged ; leaves brown _ when dry _______ 2 . P. aau·jose,num. 
4. Midrib snbequaUy brancbed IhrougbouL ___________ ________ • ______ _____ o. 
Midrib without etrong ascending brancbes upwards ____________ ____ ____ __ 23. 
n. Leaf base appreciably acute ___ __________ _________________________ _____ 6. 
Leat base appreciably obtuse. at least OD one 81de _______________________ 16. 
6. I..eaves glabrous above ___ ________________ _________ __________________ ___ 7. 
Leaves pubescent above, large _______________ _______ ____ 24. P. gatunenH. 
7. Leaves glabrous beneath ____________________________ ___________________ 8. 
~erve8 (at least) pubescent beneath _________ ______ ____________________ 14. 
8. Leaves about balt 88 broad 8S long ___________ __________________________ 9. 
Leaves Dlore elongate ________________________________________________ _ 10. 
9. Leaves 10 10 14 by 20 to 30 CID ______________ n n_n_nnl!2. P. calocoma. 
Leaves scarcely bait as large _______________________ ____ 14. P. darienenae. 
10. Leat base nearly equlIateral _____________________________________ ____ ll. 
Leaves distinctly obUque ___ __________________ __ ____ ___ I8. P. falcltollum 
11. Spfkes !)Careety 2 em. long __ ________________ ______ 3~. P. amphoricarpnM. 
Spikes Dlore elongate ___________________________ _____________________ 12. 
12. !.eaves drying green __ _____ ______ ________ ____ __ ____ _______________ ___ 13. 
!.eave8 pale or bronzed beneath ______________ _________ 17. P. cordu1,tnm. 
18. Berries depressed·globose ___ _________________ _____ ___ 18. P. aeutlsaJmum. 
Berries elUpeoJd.mucronate ___________________________ 18. P. ]avsptcum , 
14. Spikes scarcely 8 COl. long ; leaves large ____________ ~27. P. latibracteum, 
SpikeR more eloDgate __________ __________ ____________________________ 15 
15. Leave8 scarcely. by 14 cm ____ __________ ________ ______ 15. P. trlcb0pU8. 
Leaves much larger _________ _________________________ 23. P . pol711eurum. 
16. Leaves glabrous above ___________________________ ________________ ____ 17, 
Leaves BOmewhat vlllou8 on both face9 ______ _____ ______ 24. P. gatuna •. 
17. Leaves large, granular or pilted _____________ ___ ______________________ 18. 
Leav .. 8ca ... ly 6 by 15 CDl _______________________ _____ ._._ •• _. ______ _ 20. 
18. Leaves firm and rugoge _____________________________ 27. P. lattbraeteum. 
Leaves thin aDd not rugose _____________________________________ ______ 19. 
19. Leaves coarsely glandular·granular ______________ __ ___ 2S. p, polTQeurum. 
Leaves finely glandular; pubescence whlte _____________ 25. p, ladrillense. 
20. Leat base nearly equllateral _______________________ __________ _________ 12. 
Leaves distinctly oblique or shorter OD one alde _____ _______________ ____ 21. 
• 
18 CONTRIBUTIONS FROM THE NATIONAL HEBBABIUM • 
21. Leayes about baIt 8S broad as IODg ___________________________________ -22. 
Leaves more eloocate ______________________ ___ _____ 19. P. laevibn.cteum. 
22. Leaves acumlDate, 6 by 15 cm ____________________ 20. P. aubnucliopioum. 
Taves rather blunt and 8JJ1aUer ____________________ 21. P. tuberculatnm , 
28. Leaf ba .. appreciably acute __________________________________________ 24. 
Leat baBe appreclably obtuse, at least OD one Ilde ______________________ 47. 
24. Mature leaves glabrous above between the Derve8 ______________________ 2fi. 
T.eave& scabrous or pubeac-ent on both slde& ______ n ____ n _________ h __ 76. 
25. Leaves quite glabrous beneath ________________________________________ 26. 
Nerves, at least, puberulent or pubeBCeDt beneath ______________________ 41: 
26. Leaves about halt as broad as long __________________________________ 27. 
Leaves more elongate _________ __________ • ____________________________ 31. 
~. Spikes scarcely 1 em. long _________________________________ ____ __ ____ 28. 
Spikes more elongate _____________ ___________________________________ 29. 
28. Petiole winged; flowers perfect __________________________ 84. P. wagner!. 
Petiole wingless ; dloec1ous _____ n _ _______ 84a.. P. wagneri m1nutiapicum. 
29. I.eoves drying green, 5 by 11 cm _______________ 38. P. tenuimucronatom . 
Leaves llrylng papery, rather large. _________________________________ 30. 
30. Stigmas sessile; berry subglobose ___________________ 44. P. ca.llibraeteum. 
Stigmas borne on a short style ______________________ 47. 1'. papyraceum . 
81. Spikes 2 em. long; leaves dark-gtanular ___________ 33. P. magnantherum. 
Spikes more eiongs te ________________________________________________ 32. 
32. Petiole glabrous _________________________ ___ _________________________ 33. 
Petiole sparsely hirtellous or villous ___ n ____ u __ 40. P. hirtellipetlolum. 
33. Leaves scarcely 6 by 14 cm. ________________________________________ 34. 
In yes appreciably larger _______ n _____________________ nnn un ____ 39. 
M. Some petioles winged or some leaves subcordste ______ lO. P. chiriQ"inum. 
Petiole not winged above the base ________ ____ ________________________ mii. 
35. Leaves green ___________________________ ______ ____________ ___________ 36. 
Leaves drying yellow beneatb ________________ '-_______________________ 38. 
86. Leaves 3 or 5--nerved near the base ____________________________________ 37. 
Leaves not trl ple~ nerved ________ _____________________________________ 12. 
87. Midrib without ascending branches upward __ n __ .88. P. I'n-lorenunum. 
Midrib with a lew such branche8 _____________________ 39. 1'. aubUlnerve. 
38. Leaves lance-oblong: plaDt glabrous ________ nn ______ 63. P. infralutel1m . 
89. Petiole 5 mm. long; spikes ftl1torm _____ n_n _________ 51. P. 8·m bU&nl1m. 
Petiole 20 mm. long; splltes stouter ___________ _______ 45. P. nitiditolinm. 
40. Nerves glabrescent benenth ____________________________ 87. P. tecumen8e. 
Nerves persistently pnberulent or pubescent beneath _______ __________ A1. 
41. Leaves about halt 8S broad as long ____________________ 48. P. amphloxY8. 
Leaves more elo~te _________________________________________________ 42. 
42. Twigs glabrous, pe.le-granular _______________________________________ AS. 
Twigs at least locally or transiently pubescenL _________________________ 46. 
43. Leaves granular·()unctulate beneath ____________________________ _______ 44. 
Lenves not dark·punctulate; petiole sborL _________________ ti2. P. atorkli. 
44. Petiole scnrcely 10 mm. long ____________ n ______ 50. P. subnudibracteum. 
Petiole 15 to 80 mm. long ____________________ _______ 45. P. nitidifolium. 
46. Leaves gI088Y, equUateral at base _______________________ 41. P. tapianum. 
lJCaves dull, obUque ________________________________________________ 46. 
46. Leaves lanceoJate _________________________________________ 61. P. kllllpl. 
Leaves laDco eUlptlc _________________________________ 55. P. luelgaudens. 
Leaves lanee-oblong _____________________________________ 42. P. turbense. 
4-7. Mature leaves glabrous above between the nerves ______________________ 4S. 
I..eaves rough or pubescent on both Rhles ____ nn ________ _ __ unn ______ 76. 
TREI.J!ASE PlPE&AOEAE OJ!' PANAVA 19 
4& tesTes ,labron. bentath _________________________ ____________________ 49. 
Ne ...... at Ieut, puberulent or pu-..ot be"""th ______ ~ ____________ 81. 
49. Leav .. about balf as broad as 10011 ____________ ______________________ 50. 
[eaTes 1D00e eloD'abe-_______________________________________________ 67, 
GO. ~ves peltate _____ ~ _______________________ ________________ ~--_____ Gl, 
I~ves not peltate ___________________________________________________ ft2. 
• 
~ scarcely Itano1ar ___ __ ~ _______________________________________ eo. 
M. Petioles not wlnged _________________________________ 51. P. ambnenum. 
Lower petlol .. wlnlrOO _______ ______________________________ 9. P. dunlapl. 
60. T eaves scarcely 12 em. long __ ____________________ 60. P. aeptupUnerrium, 
Leaves much larcer _____________________________ l1. P. lIuudo-varlabile. 
61. Older leaves rather deeply cordate or auricled, or pelt.ate ______________ 62. 
Leaves at most cordulate or shallowly subcordate ______________________ 67. 
62. Leaf base equilateral; petiole not winged ___________ 7. P. pauloWDUol1um. 
r .... t base InequlJateraL _____________________________________________ 63. 
63. Petiole tleshy-wo.rt1 ___________________________________ 92. P. Imperiale. 
Petiole not warty ___________________ ..;. ________________________________ 64. 
64. :Leaves Bubacute ________ ________________________ ___ ______________ -+-_ _ 6li. 
Lea ve8 long-Bcum l Da te ___________________ ___ ___ _______ -+-_____________ 66. 
65. Leaves neither rugose Dor peltate _________________ _______ 91. P. aurttum. 
leaves rugose and peltate ________ .. __________ ___________ 88. EI. u7..t1. 
66. Leayes eurieled OD ODe 81de ___________________________ 87. P. 
laves InequUaterally 8uboordate ________________ 71. P. II nacmm'natum, 
61. Petiole clabrous ____________________________________ -t-_____________ __ 68. 
Petiole subpuberulent ________ __________________ _______ !liS. P. eolonenae. 
Petiole more or less halry ___ _____________________ __ __________________ ott 
68. Leaves acute on one e1de at base ________________ tliO. P. aUbnudlbracteom. 
Leaves rounded at ba8e _____________________________ ___ S7. P. tecumen8e. 
69. 'twIgs sparsely subvelvety; leaves oblanceolate _______ 69. P. n.bd1latabtm. 
TwIgs tranalently appressed-batry ______________ _____ ___ 68. P. dtespDum. 
Twigs crlsp-tomentuI08e __ h _ ____ h ____ h ____________ __ 54. P. wt]Uam.ti. 
~~ cri~halr1 or hirsute _____________________ ___________ + ________ 70. 
Twlga vlllou8 _______________ ________________________ _ 29. P. culebranurn. 
70. TwIga glabreoceot. creen _____________________________________________ 71. 
Pubescence perelstent ________ __________________________ ;... __ -+-__________ 73. 
11. LoWer petioles wlDged ____ ___________________________ 8. P. pe,lm".anum. 
Petioles not wlnged _____ _________________________________________ ___ _ 72 . 
.. 
20 CONTRIBUTIONS FOOM THl! NATIONAL HERBARIUM 
72. Leaves 8 em. wide, not caudate ____________________ 46. P. erectamentum. 
Leaves 7 em. wIde. caudate _________________________ 49. P. subcaudatum , 
Leaves 5 to 6 em. wlde _________________________________ AlI. P. turbense. 
Leaves 4 em. wlde ______________ _____________________ 82. P. taboganum. 
73. Leaves cordulate, rather glossy __ + ___________________________________ 7 •. 
Leaves not cordulate, firm, dulL. _____________________________________ 7:i. 
14. Leaves glabresceut between the nerves. ______________ .28. P. I'Duocarpum. 
Leaves loosely bah")" beneath; spikes very shorL _____ 30. P. pubbUpulum, 
7tS. Leaves smooth ______________________________________ .59. P. pall_meuM. 
Leaves lepldote-roughened j nerves imprel!8e<L ______________ 61. P. killlpl. 
76. Leaves about halt as broad as 100g __________________________________ 77. 
~ves Inore elongate ________________________________________________ 87. 
77. Leaves glandular·grRDular above _________________ 71. P. peracuminatum. 
Leaves scabrous above _______________________________________________ 78. 
Leaves scarcely scabrous ____ ____ ____ ______ ___________________________ 83. 
78. Twigs erIBP-pubescenL ________ ___ _________ ____________________ ___ _____ 79. 
Twig. acabrld-blspld or b1rsute _______________________________________ 80. 
'I wigs villous __ uu ____ n _____________ u ___________ 76. P. villiramulnm. 
79. Leayes &earcely 6 by 12 cffi ______ ______________ 66. P. pseudo-cativalenae. 
Leaves becoming 8 by 16 cm _____ _____________________ 73. P. barbinerve. 
SO. Pubescence Bbort-Bcabrld ______________________________________ _______ 81. 
Pubescence more upcurved and hirsnte ________________________________ 82. 
B1. LeaYes scarcely 6 by 12 cm _____ _______________ 66. P. pseudo-catlva!ense. 
Leaves becoming 9 by 15 CIDu ______ nnn ____ n __ 64. P. tracbydermum• 
82. £.aves pale-granular beneoth ______________ ____ n __ 74. P. nOD-retronum . 
Leaves not markedly pale-granular beneathn __ ______ 72. P. laDctl-felicia. 
83. Plants climblngj leaves very lepldote ______ ___ ________ 83. P. vitabunduIn. 
Plants not climbing ____________ _________ _____ • _______ ___ __ ____ ___ ____ 84. 
84. Leaves 10 by 20 e01 ____________ __ ______________________________ __ ____ 85. 
Leaves distinctly swaller _______ ________ ___ _____________ _______ ___ ___ 86. 
85. Leaves Dot corduJate _____________ _______________________ 26. P. stevens!. 
Leaves unequally cordulate _______________________ 71. P. peracuminatnrn • 
86. Pubescence velvety-tomentulose _________ _____ ____________ 6. P. bigelovii. 
Pubescence sott, crlsp-appressed. _____ n _ _ ____ h _ _ _____ 70. P. obaldtannm. 
Pubescence sparsely vUlous ____________ ___ ____________ 65. P. caUvalense. 
87. Leaves scabrous above ________ ________________________ ___________ ____ 88. 
-Leaves scarcely scabrous _______ ___ ___________________________________ 96. 
88. Spikes hooked _____ ____ ______ ____ _____ . ________________ 77. P. elongatum. 
Spikes nearly or quite stralghL ________________________________ ______ 89. 
89. Twigs densely roo-velvety ____________ ____ ____ ______ 86. P. pervelutinum. 
Twigs crisp-pubescent or appressed-pubescent __________________________ 91. 
Twigs blspld·hlraute __________ __ __________________________ 61. P. ldlllpl. 
Twigs toment()$' or scabrous-hlspid __________ __ ______ ___ ___________ ___ 90. 
Twigs villous ___________________ _____________________________________ 94. 
90. Leaves oblong. 4 by 16 cm ______ n _______ n ____ __ 79_ P. changulnolanum. 
Leaves obliquely elliptlc-ovate, 8 by 18 Cln ____ _______ 75. P. IIcabrilimbum. 
91. Twigs densely canescent upward~ ___________________________ 85. P. breve. 
Pubescence Olore sparse ______ ______ ______________________ ___ ___ __ ____ 92 
92. Leaves rough, oblong ______________________________________ 85. :P. breve. 
Leaves sIDootber __ ____________ __ _______________________ _____________ .93. 
93. Leaves broadly 18nce-el11ptic ______ ____ _____ ____ _____ 78_ P. chagresianum. 
Leaves broadl)' oblanceolnte __ ____ n_nn __ n _______ 69. P. aubdUatat1Jm. 
94. Leaves subequllateral, elllpUc-oblouS ____ ___ _ • _________________________ 96. 
Leaves obliquely ovate-e!Uptic ____ ___________ _______ 76. P. vilUramulum. 
... 
TRIII,EASE PIPEBACEAE OF PANAMA 21 
9Ci, r ... _ ocar<ely f by 12 cm ________ h ___ h ___ h_
h
_
h
li6. p, viridlcaule, 
Leaves twice 88 IODg, gnmulaf __ _____ nn ________ __ 57. P. penubulatum, 
96. Leaves ocarcely 2 by 10 cm ___ h_h __ h ____ _________ _ 81, p, Uuearl!oll1!m, 
Leaves dIstinctly larger _____________ ______ _____ __ __ _________________ 97. 
fY1. Twigs glabrous, gr8nular __________________________________ SlI, P. otorldl. 
'1wlga 8pt1lCSJed·hlBpidj plants cllmblug _______ ___ _ u_83. P. vitabundnm. 
'1 wigs sparingly crls~pubescent or appressed-pube8cent or bJrsute _______ 98. 
Twigs almost cane8Cent above ______________________________ 85. P. breve. 
~lgs villo08 __________ ____________________________________________ 100. 
98. Leaves scarcely Ij by 16 em., the base obtuse ___________________________ OO. 
Leaves 8 by 20 em., the base Bubacute _____________ 53. P. palUdlbractellm. 
99. Leaves lanceolate ____ __________________________________ 80. P. fatoanl1m. 
Leav .. obianceoiate ________________________________ 69. P. aubdllatatum, 
100. Spikes short ______________________________________________________ 101. 
Spikes slender and elongate __________________ _____ 87. P. pll0alU8cul11m. 
101. Leaves granular beneath ________________________ __ 32. P. garagarlLDum. 
laves not grnnular ____ __ ___ ______________ 31. P. p.eudo~garagara.nllm , 
1. Piper omilaclfollum H. B. K. Nov. Gon. & Sp. 1: 56. 181~. 
'rtPE LOCAI.ITt: Caripe, Venezuela. 
RANOI:: Venezuela to Panama and Guatemala. 
CANll Zoo: Barro Colorado leland, 810""1611 31267. cano Quebrado. Pit· 
t4er 6986. Cerro Gordo, Culebra, 8tafldwg 26008. Fort Sherman. BtGtwileu 
30959; 8t."."., 1066. France Field to catlval, 81andler 30338. Oamboa to 
Cruces, Pill"'" 3718. OatOn, 8tandley 27201. Gatuncfilo .. Pi_ 5603, 5620. 
Gorgona to Galan, Pill"'" 2279. La. C88cadas Plantation, 81afldlev 25728. 
Mamel Hfil. PltI"" 3811. Matachln, Kuntze 1918. Obispo, 8tandlev 31720. 
Rro Agull 8alud, FriJoles, P'rJer ~9, 5860. Rio Pedro Miguel, East Paratso, 
8tandl6JI 29969. CoWl<: Chagres Mouth, Htev.,.. 278. Fat6, Pili"'" 381 2. 
Rio Fat6. PltI"'" 3872, 8882. P .. H ........ : RIo Tecumen, 8tandlev 29363. Rio 
Tapia, Matton d Baf'1Jeg 6600, 6708; Standley 26160, ~,28"41. Juan Dfuz. 
BtandleJi 30548. DUliN: SambQ River, Pittier 5570. BOCAB DEL TORO: Lower 
ChaDgulDola River, 8t",." 00. 
2. Piper san~joseanum C. DC. Llnnaea 37: 351. 1872. 
Piper margiMtum Auct. as to Panama. 
TYPE LOCALITY: Mount Aguacate. Costa Rica. 
• 
RANGE: Costa Rica and Panama.---One of the continental eQulvaleots of P. 
. . 
marginatvm of the Antilles, which Is quite 18I!kinc trom the.: ~r1b~ees. . 
PRoVINCE (?): Santa RUa Trail, CO'IOeU 122. VSUOUA..s: I sla de Uva, 
Pltl4er 5112. CANAL Zorn,,: A«U8 Clara, Btew,,,, M6. Anc6n Hill, KUUp :ro,u, 
12087; 8tarull.eJl 26326. Balboa, Stan4re_ 26600. Cerro Gordo, Culebra, PUtler 
3737: BtandleJi 25964. Culebra, PUtier 2146 j 8teven' 927. DariED. Statioll, 
Sta"dleJ/ 31M7. Fort Clayton to Corozal, 8tan.dZey 29214. Fort Randolph, 
Standley ~~. Lns Cruces TralI, 81antUeJi 29214. Fort Sherman, Stand­
lev 80942; 81.-. 267. FrIJoles, 8tandlev 27627. Frijoles to Monte 
Llrlo, Kmip 12116. Gamboa, 8tandley 28334, 28536. Gatc1D, 8tandlev 21247, 
27288. Corozal, 8te-v6rt1 63, 118. Monkey Hill above Coi6o, LehmaM 768. 
Juan Mlna, Piper 5600, 5698. Las Cascadas Plantation, 8tandlev 25751. 
Summit, StandleJl 26938. New Um6n, Btevens 1015. CoWN: FatO to Playa de 
Damas, Pitt"", 3934. Rio Chagres, Piper 5898, 5917, 11962. SaJlta Isabel, 
Pillier 4176. PAN .. M .. : Bella Vlota, KilUp 12012; St .... dley 25382. Chepo, 
Pilt4er 4708. Corozal Road, 8tand'etI 26834. CamIno de la Granja, Bro. 
Benberto 166. Junn Diaz, Btarsdlell 80621. JAlS Sabano.& to MaUas HernAndez. 
8la"dley 31835. Punta Paltllla, 8tandley 26274. RIo Tapia, Standley 28207 . 
• 
22 CONTRIBUTIONS FROM nU: NATIONAL HERBARIUM 
Rlo Tecumen. 8tcnuneJl 26590, 
3M3: SloAdI6v 27044, 270Ci7, 27847. 
Tabcga Ialand, Ki/lip 3165, 8195; Piltier 
DuliN : Garaehln~, Plllier 5612. 
iL Piper aan-Joaeanum aristolochlaefolium Trel., var. nov. 
A form. with more narrowly ovate. dull dark green, bullate leaves 8 to S.ri by 
15 em., deeply cordate, with parallel margin. to the sinus. 
T1P6 In the U. S. National Herbarium, DO. 1,217,625. collected on bUlB north 
ot Frijoles, Canal Zone, Panama, by Paul C. Standley (no. 27524). 
3. Piper pinol'anense Tret, sp. nov. 
A shrub; flowering internodes greeD, much 11uted, minutely and very 
transiently puberulent; leaves elliptic, caudate, rounded at base but suJcateJy 
narrowed to the petiole, moderate (4.5-5.5X9-12 em.), 5--nerved, the nerves 
aaUent on both sides, glabrous, very thick and firm, drying green; petiole very 
abort (scarcely 5 mm. ), puberulent. winged and deeply channeled; spikes op­
posite the young leaves, 1llltorm, 4 em. long, loosely flowered; peduncle scarcely 
1 em. long, slender: bracts minute, concave j :ft.owers sessile, perfect; atigmas 
3, minute. 
Type in the Delessert Herbarium, Geneva (duplicate in the U. S. National 
Herbarium>. collected at Pinogana, BOuthern Dari~n. Panama, by B. Pltt1er 
(DO. 6566). 
BANGE: Southeastern Panama. 
4. Piper pervenosum C. DC. (Candonea 1: 268, name only. 1923) Schroeder, 
Condollea 3: 137. 1926. 
A sbrub: o.owerlng Internodes moderately slender but sbort, gray~tomentuloae; 
leaves broadly ovate, subacute, equally rounded or sUghtly cordulate at base, 
rather large (8XI4-11XI8 em.), pinnately nerved from below the upper third, 
the main nerves 1 3 or 4X2 but with intermediates and spaced shorter veins 
nearly to the end. glabrous, gray-gleen, paler beneath; petiole very short 
(G mm.). tomentulose-puberulent, winged to the end; spikes terminal or oppo­
,Ite the leaves. small (4X2ti mm.); peduncle sbort (scarcely 10 mm.). gray~ 
tomentulose; bracts lunulate; flowers sessile, perfect i ovary round, with a 
Tery Bhort style. 
T!n LOCAI.tTY: Puerto Obaldfs, Sao BIas Coost, Province of Col6n (P4ttler 
4302, the type). 
RANGE: Northeastern Panama. 
5. I'iper obUtum Trel., sp. nov. 
A small BCaDdent (1) shrub; flowering internodes slender and short, from 
sllgbtly velvety quickly glabrescent i leaves broadly ovate, very bluntly acumi­
nate, Bubequally rounded at base, moderately large (a--aXIO-13 COl.), sub­
mUltiple-nerved from the lower fourth. the nerves 4X2, the lower eoncested. 
the nerves aboTe sparsely hairy, rusty-tomentuloBe beneath; petiole ratber 
short (13+2 oc 3 mm.) fteecy, wIni"d at ba .. ; splkee opposite the lea.es. 
moderate (3 X30-40 mm.); peduncle short (6-7 mm.), sparingly balry; bracts 
lunulateIy subpe1tate, velvety; flowers sessile, pertect. 
Type in the berbarium of the New York. Botanical Garden, collected in 
.. Panama" by Sutton Bayes (no. 803) . 
RANom: Northeastern (1) Panama . 
• 'lwo forms of expression used In this paper in the description of leaves 
perhaps require explanation. The phrase .. main nerves 3 or 4 X 2" indicates 
that there are S or 4- chief lateral nenes on each side of the costa. The phrase 
.. petiole rather abort (13+2 mm.)" Indlcates that In a leaf with unequal base 
the blade extends on one side 2 mm. below the apex of tbe petiole. 
TBI!I.EASE PIPERAOEAE OF PANAMA 23 
6. Piper blgelovU 'Ii e1. sp. DO •. 
A.. Rbrub, nodose, rnsty-tomentulose or velvety througbout; llowerlnc lDter­
nodes rather slender and short; leaves elliptic to ovate, 8ubacumlnate, 
rounded or cordulate at bftBe with one side commonly shorter, moderately 
I."" (5-7XI2-lB em.). plDnately Derved from below the middle, the De"eo 
5 or 6X2; petiole -rather short (10-10+4 mm.). winged at base; spIltes oppo­
site the Ie ..... moderately elong.te (3X~ rom.); peduncle ahort (5-10 
mm.); bracts triangular-BUbpeltate. ciliate; dowers BCSIUe, perfect; berrlea 
minute. obpyramfdal, 3-aided i sUpes 3, minute, tlrear, Berlle. 
iyve in the Torrey Herbarium. New York Botanical Garden. eollected on 
the II Isthmus of Panama" by 1. N. Bigelow. 
RANGE: Panama. 
7. Piper paulowDlfolium C. DC. Aoal. lost. Fl •. Googr. Costa Rica 9: 173. 1393. 
Pl"er oa.rimiriant"rJ Auet. as to Panama. 
TiP.: I..OCALITY: La Palma, Costa Rica. 
BANDE: Costa Rica and Panama. 
CBmIQui: H.to de Jobo. San Felix, Plllw 1l427. CelTO de la Plaia. San 
Felix. Plllw 5153. CANAL Zon: Borro Colorado IalaDd, AlOOf.", Ha"'"1l 
" VllJen.t"'e 6809; 8tan4lell 81282, 31893; StuEm r 645. Catlo Quebrado, 
Plllw 6666. Frljol ... 81 .... <1161/ 216l2. GatllD, 8tGfI<IIell 27226. GorgoDa to 
Gat1lD. Pitlier 2261. Mamel HlI~ Pitner 3312. CoL6N: PalO, PUller 33<14. 
S862. DAartN: Boca de Pauarand6, Sambti River, NIUe,. 5687. 
8. Piper palmaaan1!m C. 00. Smith •. Mlac. Coli. 71': 3. 1920. 
'1 iPJ: LOCA..LITY: Cuesta de las Palmas, Cerro de la Horqueta, Chiriquf 
(PiIlIer 3225. the type). 
IU.l'I'O.: Southwestern panAma. 
9. Piper dunlapl TreL, sp. no •. 
A shrub, glabrous, nodose; 1I0wering internodes rather slender and short, 
Il'anular, dark In drying; leaves ovate or lsncc-ovate, more or leBS sharply 
acuminate, rather unequally acute to equllater.llly rounded at ba8EI, moderately 
.mall (5-5.GX13 em.). 8ubmultlple·Derved. the midrib OD eaeh side with about 
I5i branches from below the middle or upper third, the lowermost busl, some­
what glandular-granular; peUole short (lO-lG mm.), wlnged at base only or 
on the larger leaves throughout; spikes opposite the leave, rather Bman 
(3X2S mm.) before flowering, mucroDate j peduncle slender and short (lc'ree17 
10 mm.) j bracts roundlsh-subpeltate, broWD-umbonat:e, with bz-oed yellow cillo­
Jate margin j flowers sessile. 
Type in the U. S. National Herbarium, DO. l,2OlS,Uil, eolJected in Ohangulnow 
Valley. Boca. del Taro, Panama. by V. C. Dnnllp (no. 281). 
RANGI!: Northwestern Panama. 
10. Piper chlrlqulnum C. DC. Smiths. Misc. Call. 71': 2. 1920. 
TTPB LOOAIJTY: EI Boquete, Chlriqul (PiUIer 2930, the type). 
RA.NGE: Southwestern Panama. 
11. Piper pseudo-varlablle 'nel., gp. DOl'. 
Pir>er grafl4tfoltum Aucl .&.8 . to Panama. 
PlJ)fIf" -variabfls Anct. as to Panama. 
A shrub, glabrous; flowering internodes rather long and slender; leaves 
e1l1ptlc to broadly ovate, acuminate, equllateraUy rounded or the larger sub­
truncate at base, large (1-8X17-13X20 em.). ptnnately to 8ubmnltlple-ne"ed 
trom below the upper fourth, tb"e ne"es :') to 7 X 2 with the lower congeate<l 
toward the bue; petiole relatively Bbort (lO-U~ liim.), winced toward the baae; 
• 
24 CONTRIBUTIONS FROM 'rnE NATIONAL HERBARIUM 
spikes opposite tbe leaves, ratber slender and long (3XOO--OO mm.) ; peduncle 
rather Blender and short (18 mm.); bracts Inconspicuous, concavej ftowera 
sesaUe, perfect; berries minute, obpyramldal, a-sided, glabrous; stigmas 3., 
minute. sessile. 
Type in the U. S. National Herbarium, no. 618920, collected on Loma de la 
Gloria, near Fat6, Province of ColOn, Panama, by H. Pittier (no. 8845). 
RANGE: Panama. 
CHIRIQUI: San FellI, Pill .... 5189, fill62. CocLt: 010, PUI .... 5075. CANAL 
ZoNE: Anron Hill, Slatullet! 25179; WUlI4...... 19. Borro Colorado Island, 
8ta'ildlell 31308. Fort Sherman. 81"",41611 31022. PAl'iIAllA: Juan Dlaz, Stand­
leu 30610. 
Ila. Piper pseudo .. varlabile pacbypus Trel., YOI'. nov. 
Nodose; leaves broadly elliptic, 10.5X20 em.; spikes 100 mm. long; peduncle 
stout. 
Type In tbe U. S. National Herbarium, no. 1,167,279, collected between Fri­
Joles and Mont. Llrlo, Ganal Zone, Pauama, by E. P. K1llip (no. 12145). 
RAI'I'olC: Central Panama. 
lIb. Piper pseudo~variAbne colltum Trel., var, nov. 
Leaves becoming elongate, 7 to 8 by 20 to 2:s cm.; passing into the type. 
Type In tbe U. 8. National Herbarium, no. 1,153,478, collected along tbe Rio 
Tapis, Province of Panama, Panama, by Paul C. Standley (no. 26130). 
RANGE: Central Panama. 
CANAL ZoNE: Callo Quebrado, P/ttler 6662. Frijoles, Standleu 27~fifi. Gatfin, 
Sta .. dle" 27191. La. Gaseadas Plantation, Standlell 211700, 25733, 29518, 20058. 
Obispo, Slandle" 31702. Rio Paralso, Slandley 29892. PANA ... ': Juan Dlas, 
Slandle" 30577. 
12. Piper crassi8picatum Opiz in Presl, ReI. Hnenk. 1: 152. 1880. 
Artant"" craBrilplcala Mlquel, Syst. Piper. 521. 1844. 
TyPE LOCAI.ITY: .. Panama." 
RANGB: Panama. 
13. Piper la:dspicum Trel., sp. nov. 
A.. small glabrous solt-wooded shrub ecarcely 50 cm. tall; stems slender, 
green; lea'fes lanceolate or lance-oblong, subacumtnate, equllaterally acute at 
bau, moderately small (4-5X12-16 em.), pinnately nerved tbrougbout, the 
ne"e8 about 8X2 with some shorter intermediates, drylng green and papery; 
petiole rather short (10 mm.), not winged; spikes opposite the leaves, In trult 
rather thick and short (7X50 mm.) ; peduncJe slender, scarcely 10 mm. long; 
bracts concave: ftowers sesstIe, pertect; berries not crowded, elUpsold, mucro­
nate, drying angular; stigmas 3, trom the mucro. 
Type in the U. S. National Herbarium, no. 1,215,963, collected on Barro Colo· 
rado Island. Gatt1n Lake, Canal Zone, Panama. by Paul C. Standley (no. 
31875). 
RANGE: East-central Panama. 
13a. Piper lu::lspicum latifollum (C. DC.) 1'1'eL 
Piper acum#·nat{uimum latifoH-um C. DC. In herb. 
Leaves bronder (6-SXl5-17 em.) and more heavily nerved. 
'£ype in U. S. National Herbarium. no. 679401, collected at Puerto Obaldfa, 
San BIas Coast. Province ot ColOn, Panama, by H. Pittter (no. 4288). 
RAffO':: Northeastern Panamn. 
14. Piper darien .... C. DC. In DC. Prodr. 16': 374. 1869. 
P'per 14naeolatum Auct. a8 to Panama. 
'IIPZ LOCALITY: "Veraguas" (Beemann). 
RANGE: South-central Panama. 
-
-I4&. PIper darl ... en .. trleu.pe (Mlquel) Trel. 
Arlattllle 1,Icu.piI Mlquel; Seem. Bot. Voy. Herald 200. pL ~2. 1800. 
Piper tn"""pe C. DC. In DC. Prodr. 16': 263. 1869. 
25 
Piper lonceolatum tricu'J)e C. DC. Candollc3. 1: 255. 1923. Name only, in 
index. 
TYPE LOCALlTi: .. Panama" (Seeman.n). 
RANGE: Panama to Ecuaclor. 
16. Piper trlcbopus TreJ., !!p. nov. 
A shrub; flowering internodes slender, rather short (2--3 em.). sparsely 
pubes<.'eut; leaves lanceolate, almost equally acute at both ends or oue sIde-
8ubobtuse at base, moderately small (3-3.5Xl2-13 em.), pinnately nerved nearly 
throughout, the looping Denes about 8X2, appressed·pubescent on the nerves 
beneath and somewhat granular; petiole rather short (5-10 mm.), very slender, 
not winged. pubescent; spikes opposite the leaves (2 X60 mm.) j peduncle ex­
tremely slender, 10 to 15 mm. long, sbort-hatry; bracts inconspicuous, concave, 
ciliate: flowers perfect, sessile; stigmas 3, minute, lJe8Sile; berries subcublcal, 
glabrous. 
Type in the Herbarium of Columbia University, New York Botanical Garden, 
collected In H Panama" by Sutton Hayes (no. 797) . 
RANOE: Central (1) Panama. 
16. Piper acutissimum Trel.. sp. nov. 
Piper lucaeanum Auct. as to Panama. 
A shrub, glabrous, nodose; flowering internodes short (2 cm.) and slender; 
leaves narrowly lanceolate or lance-Qblong, acute at both ends or subacuminate. 
rather small (2.5-3 X 10-14 em.), pinnately nerved nearly throughout, the larger 
nenes about 6 X2, but with ascending nenules upward; petiole rather short 
(10 mm.), not winged; spikes opposite the leuves, rather small (3 X50 mm.) , 
submucronate; peduncle slender and short (scarcely 10 mm.) : bracts concavely 
truncate, ciUate; flowers sessUe. perfect; berries depressed-globose, more or 
less angular from pressure, velvety; stigmas 3, sessile. 
Type In the U. S. National Herbarium, no. 38081, collected at Chagres, 
Province of CoJOn, Panama, by A. Fendler (no. 266). 
RANGE : Eastern centrnl Panama. 
CANAL ZoNE: Barro Colorado Island, StCOOM 1. Frijoles, 8ttJftdleJl 275Of., 
27500. 
17. PIper cordulatum C. DC. Joum. Bot. Brit. '" For. 4: 217. 11186. 
Typs LOCALITY: Chagie~t Province of Col6n (PtI,i4rer '267, the type"). 
RANGe: Eastern central Panama. 
CAN AI. ZoNE: Agua Clara, SteveM 580. Barro Colorado Island, S'afldle1l 
313M. Frijoles, 8'antUev 27430, 270'18. GatOn, H4fIe. ' "1391 If. Gorgona to 
GatOn, PiU£ef' 2267. Las Cruces, Pit tier 26241. Rfo Parafso, Statt4ley 2990tS. 
17a. Piper cordulatum granulatum Trel., var. nov. 
DUrers from the type In its very acute-based, granular-roughened leaves. 
'lype In the U. S. National Herbarium, no. 1,153.882, collected near FriJoles, 
Canal Zone, Panama. by Paul C. Standley (no. 27M7 ) . 
RANG~: East-central Panama. 
18. Piper lalclfollum Tre1., sp. nov. 
Piper tpenic1tlatum Auet. 88 to Panama. 
A shrub, glabrous, nodose; fl owering internodes Blender and short (2-3 em.). 
becomIng coarsely warty; l~aves falcately lanceolate, acuminate, unequal1~' 
aente at balJe with one sIde appreciably longer and decurrent, moderately sman 
(3-3.5X ll-14 em.), pinnately nerved throughout, the nerves nOOut 10X2 with 
22089-27 3 
26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
short tnterme<Uates; petiole rather long (10+5 or even 2C'i mm.), wmged 
throughout; spikes moderate (3X60 mm.) ; peduncle filiform, rather short (10 
mm.) ; !tracts truncate1,. sublunate, rusty-clliate j flowers sesslle. perfect. 
Type in the U. S. National Herbarium, no. 715322, collected at San Felix, 
Chiriqui, Panama, by H. Pittler (no. 5137). 
R.\NGE: Southwestern Panama. 
19. Piper laevibracteum Trel., sp. nov. 
A shrub, glabrous, nodose i flowering internodes slender and moderately short 
(2.5-4 em.), at length warty; leaves lance- or elliptic-oblong, subacuminate, very 
unequal at base with the longer side acute except on the largest, moderately 
large (4.5-7Xl5-22 em.), pinnately nerved throughout, the nerves about 12X2; 
petiole short (ts+l0 mm.), winged throughout; spikes opposite the leaves, 
moderate (2X60 mm.) ; peduncle slender Bod short (10 mm.) ; bracts truncate~ 
lunulate, microscopically cillolate; flowers sessile, perfect; stigmas 2 to 4, 
sessile. 
Type In the U. S. National Herbarium, no. 715545, collected on Cerro Vaca,. 
Chiriqui, Panama, by H. Plttler (no. 5324). 
BANGE: Southwestern Panama. 
CHIRIQuf: San Felix, Pittier 5131. EI Boquete, Malfon 5381. EI Boquete to 
-Caldera, Pitfier 8327. 
20. Piper Bubnudispicum Trel.. sp. nov. 
An essentially glabrous sllrub, resembling the preceding; leaves elliptlc­
oblong, acuminate, unequally obtuse at base, rather large (6-1X16 em.), drying 
membranaceous; spikes filiform, 2XSO mm. j peduncle short (5 mm.): bracts 
very Inconspicuous. 
Type In the U. S. National Herbarium, no. 679342, collected on Lorna de In 
GlorIa, Fat6, Province ot 0016n, Panama, by H. Plttier (no. 4241). 
RANGE: East-rentral Panama. 
CANAL ZoNE: Barro Colorado Island, Maa:on, Harvev & Valentine 6813. Fort 
Sherman, Slandiev 3111~. Frijoles, K lUiI> 3420; Slandlev 27519. 
21. Piper tuberculatum Jncq. CoIl. Bot. 5: 7. 1786. 
TyPE LOCALITY: "Tropical America" (Venezuela?). 
RANGE: Northern South America to Central AmeriCll. 
~.ANAL ZoNE: Balboa, Standley 25409, 32117. Empire. BteveM 1143. 
PANAMA: Corozal Road, Standletl 26781. Juan Dlaz, StandZev 30470. JuaD 
Franco Race Track, Panama, Standlev 27676. Rfo Tapia, Standley 28280. 
Tumba Muerta Road, Standley 29712. Taboga Island, Pittler 3560; 8tandlell 
27093. 
22. Piper calocoma (Mlquel) C. DO. In DC. Prodr. 16': 264. 1869. 
Artanthe caloooma Miquelin Seem. Bot. Voy. Herald 199. 1856. 
TYPE LOOAI.iTi : DarMn (Seemann 1067, the type). 
RANGE: Colombia to south-central Panama. 
COOLIt: Bismarck, above Penonom~, Williams 332. DARltN: Seemann 1067. 
23. Piper po}yneurnm C. DC. (Candollea 1: 270. 1923, name only, in index) ; 
Schroeder, Candollea 3: 138. 1926. 
A small tree 2 to 4 meters tall, coarsely glandular-granular throughout; 
flowering internodes moderately stout and long, sparingly tomentulose or 
glabrate j leaves elliptic-ovate, short-acuminate, subequally cordulate or 
rounded or exceptionally acute at baBe, large (9-11 X 20--23 cm.), pinnately 
nerved nearly throughout, the nerves about 12X2, appressed-pubescent on the 
nerves beneath; petiole short (5 mm.) and not winged or on the round-based 
leaves elongate (20--40 mm.) and winged to the end. glabrate; spikes opposite 
TBEI.EASE PIPEIlACEAE 01' PANAMA 27 
the leaves, arched. ratber large (5X1OO mm.) i peduncle thick and sbort 
(seo.reely 10 mm.) , glabrate; bracts large, deltoid-peltate, brown-tomeotuloae; 
ftowen sessile, perfect; stlgmas 3, large, sllgbtl7 stalked. 
TyPB LOCALITY: Cerro de Garagam., Samba Basin, Darl~ (Pi"ler GOa, the 
type). 
R .... NOE: Panama. 
CANAL ZoNE: FrUoles, Standley 27495. nro Indio de GatOn, PUtter 2790, 
2791. 
24. Piper gatunense Trel., &p. nov. 
A shrub, 2 to S meters tall; flowering internodes finally stout and long, greeD, 
from vlllou8 glabrescent and glandular-granular; leaves lance elUptle, short. 
acuminate and slightly inequi1ateraUy acute at base, or ovate-oblong BDd equl­
laterally 8ubtruneate at base, pinnately nerved nearly throughout, the nerves 
12 to 16X2, sparsely villous on both sides; petiole short (~10 mm. ) and not 
winged, or on the broader leaves 3 cm. long, vlllons; spikes opposite the leaves, 
arched, large (8 X 100 mm.); peduncle stout and moderately elongate (20 
mm.), trom villous globrescent; bracts triangular-.iobpeltate. the yellow mar­
gin c1l1ote; :flowers sessUe, perfect; stigmas 3, sessile; berrIes oblong. 
Type in the Torrey Herbarium, New York Boumleal Garden, collected at 
GatOn, Canal Zone. Panama, by Sutton Hayes (no. 5(2). 
RA.NGE: East-central Panama. 
CANAL ZoNE: FriJOles, 8/a1Odle!i 27465, 27491. 
2!S. Piper ladrillenae Trel., lip. nov. 
A moderate-sIzed shrub; flowering internodes rather slender &lid elongate, 
not granular, tronsiently erisp.pubernlent; leaves broadly ovate, shortly sub­
acuminate, somewhat InequUaterally rounded at base or the shorter side sub­
acute, large (ll X20-22 em.), pinnately nerved nearly througbout, the nerves 
12 to 15X2, sparsely whtte-erisp-hairy on the nerves beneath and obscurely 
and finely glandular·granular; petiole moderately short (10+2-1G+G mm.), 
winged, very sparsely whIte-hairy or glnbrate: spikes opposite the leaves, 
rather large (before flowering 4X55 mm.); peduncle short (10 mm.), glob­
rate; bracts round or lunate-subpeltate, the narrow pale margin ciUoJate; dow-
ers sesslle, perfect. . 
Type in the U. S. Notional Herbarium, no. 677563, collected along the Bto 
LadrUlo, Chiriquf. Panama, by H. PUtter (no. 3164) . The collection bas been 
referred to the glabrous peltote·leaved P. maatonit. 
RANG!: : Southwestern Panama. 
28. Piper stevensl TreJ.. sp. DOV. 
A shrub: flowering internodes rather stoot and. loog, gleeo, densely crisp­
hairy ; leaveH elllptic, sbort-acuminate, rounded at base 00 one aide with the 
other shorter and acute, large (l6-U X21-23 cm.), pinnately De"ed from be­
low the upper fourth, the nerves about 7X2, rather loosely crisp-pubescent on 
both faces with the nerves pale-hirsute beneath, scarcely granular; petiole 
moderately long (20+10 mm.), winged below the middle, crisp-bairy; spikes 
opposite the leaves, rather large (5XSO mm.) ; peduncle 2 cm. long, moderately 
slender, 8ubretrorsely crlsp.pubescent; bracts elliptical-subpeltate, pale-mar­
gined and clliate; flowers sesslle, perfect; stigmas 3, sessUe; berries 8ubglobose­
oblong. 
Type in the herbarium ot the University of Illinois, collected at FrljoleB. 
Canal Zone, Panoma, by F . L. Stevens (no. 1247). 
RANG!:: East-eentral Panama. 
28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
27. Piper latibracteum C. DC. Smiths. Misc. ColI. 71': 5. 1920. 
TyPE LOCALITY: Boca de Pauarand6, Sambl1 River, D8ri~n (PUtier 5588). 
RANGE: Southeastern Panama. 
28. Piper gonoearpum Trel., sp. nov. 
A shrub, 2 meters tall: flowering internodes moderately slender and elongate, 
din!'''Y-tomentulose; leaves lance-eUiptic or oblong, somewhat acuminate, rounded 
at base or subcordulate, with one side sllghtly shorter, moderate (6.5X17 em.), 
pinnately nerved trom below the middle, the nerves 5 or 6X 2, dull green. 
glabrous and slightly cellular-pitted above, hirsute on the nerves and paler 
beneath; petiole rather sbort (10+2 mm.), deeply concave, winged, tomentu­
lose; sl)ikes opposite the leaves, thick and rather short (8X45 mm.), caudate 
and sterile at tip; peduncle short (10 mm.), sparsely hairy i bracts broad, con­
cave; flowers sessile, perfect; berries depressed·quadrangular, 2 to 3 mm. In 
diameter, 4-ridged. obscurely puberulent; stigmas 3, broad, sessile. 
Type in the U. S. National Herbarium, no. 1,153,832, collected near Frijoles. 
Canal Zone, Panama, by Paul C. Standley (no. 27437). 
RANGE: East-central Panama. 
29. Piper culebranum C. DC. Candollea 1: 121, 243. 1923. 
TYPE LOCALITY: Rfo Culebra, Collin (PUtier 4154, the type). 
RANGE: East-central Panama. 
CANAL ZONE: Fort Randolph, Standley 28622, 28667. France Field to Catl· 
val, Standley 30315. PANAMA: Juan Diaz, Standley 30611. 
80. Piper publstlpulum C. DC. Smiths. Misc. Coli. 71': 5. 1920. 
TYPE LOCALITY: Agua Clara, GatOn, Cnnal Zone (Pittier 2661, the type). 
RANGE: East-central Panama. 
• 
30a. Piper publstipulum estylosum TreJ., var. nov. 
Differing in its sessile stigmas. 
Type in the U. S. National Herbarium, no. 1,215,959, collected on Barro 
Colorado Island, Canal Zone, Panama, by Paul C. Standley (no. 31364). 
RANGE: With the typt!. 
31. Piper pseudo-garagaranum Trel., sp. nov. 
A shrub, scarcely 2 meters tall; flowering internodes zigzng, rather slender 
and short, evanescently villous; leaves elliptic-Ianceolate, acuminate, inequJ­
laterally rounded at base, moderately large (5.5-7X 14-16 cm.), pinnately nerved 
from below about the middle, the alternate nerves about 4X2, with a few soft 
appressed long hairs on the dark green upper surface, the nerves villous be­
neath; petiole short (3+3 mm.), scarcely winged, villous; spikes opposite the 
leaves, as yet young and small (lX20 mm.), with short (5 mm.), sparsely 
pubescent peduncle; bracts rounded'subpeltute, the narrow pale margin cilia­
late; flowers sessile. 
Type in the U. S. National Herbarium, no. 1,215,804, collected on Barro 
Colorado Island, Canal Zone, Panama, by Paul C. Standley (no. 314(9). 
RANGE: East-central Panama. 
32. Piper garagaranum C. DC. Smiths. Misc. CoIl. 711: 15. 1920. 
TIPE LOCALITY: Garagarfi., Sambu Busin, Darien (Pittier 5618. the type). 
RANGE: Southeastern Panama. 
33. Piper magnantherum C. DC. Smiths. Misc. CoIl. 71': 16. 1920. 
TYPB LOCALITY: Cerro de Gal'agan1., Dari~n (PUtier 5666, the type). 
RANGE: Southeastern Panama. 
34. Piper wagneri C. DC. in DC. Prodr. 161 : 302. 1869. 
'l'ypE LOC"IJTY: .. Chiriquf" (Wagner). 
RANGE: Southwestern Panama. 
L 
TBELEASE PIPEBACEAE O:r PAN AMA 29 
34a. Piper wagner! var. minutlapicum (C. DC.) Trel. 
Piper minutl.plcum C. DC. Smith •. Misc. Coil. 71': 1. 1920. 
TYPE LOCALITY: Snbnna Grande, El Sulto, aboye El Boquete, Chiriqui (Pit· 
lior 3116, Ibe type). 
RA...~OE: Southwestern Panama. 
31S. Piper amphoricarpum Trol., sp. nov. 
A shrub, glabrous; ftowering internodes rather slender and short (2-3 em.) ; 
leaves lanceolnte. 8ubacuminate, equilaterally long·acute at base, moderately 
smen (3-4X12-14 em.), pinnately nened nearly to the end., the nerves about 
8X2; petiole short (5 mm.), wlBged at base; 8Plkes opposite tbe I ...... small 
(3-6X12-1fi mm.) , very blunt; peduncle filiform, short (searce17 10 mm.); 
bracts lunulate·subpeltate, goldcn..clUate; 1lowers sessile, pedect; ovary glab­
rous, granular, 1lask·shaped, in fruit attenuate Into a slender style bearing tbe 
3 sUgmas. 
Type In the U. S. National Herbarium, no. 1,010,185, collected near EI 
Boquete, Chiriqui, Panama, by E . P . KIlllp (no. 3611). 
R&NGE: Southwestern Panama. 
sa. Piper tenuimucronatum C. DC. Smiths. Misc. Coil. 71': 12. 1920. 
TypE LOCAl.n't: Los Siguas Camp, Cerro de 10. Horqueta, Chlriqu( (Maz01t 
5421, Ibe type). 
RANGE: Southwestern Panama. 
37. PJper tecumenae Trel. . sp. nov. 
A shrub, scarcely 2 meters tall, essentially glabrous j Internodes slender and 
elongate, evanescently velvety at the nodes; leaves lanCfH)vate to elUptlc-ovate, 
sbarply but gradually long-acuminate, rounded at base with one side sllgbtly 
longer, moderately small (4 X l1-6XI4 em.), pinnately nerved from below 
the middle, the nerves about 4 X 2, looping, very obscurely aDpressed-pubescent 
on the nerves beneath or quite glabrescent; peUole Vet'7 short (scarcely Ci+2 
mm.), winged at base; sUpules and stipular Hne somewhat pubescent; spikes 
opposite the leaves, when young rather small (I.CiX35 mm.) ; peduncle dUform, 
1 em. long; bracts rounded-subpeltate, with pale clliolate margin; flowers 
sessile, perfect. 
Type in the U. 8. National Herbarium, no. 1,154.361, collected along the Rio 
Tecumen, Province of Panamat Panama, by Paul C. Standley (no. 293fi8). 
RANGE: Eastern Panama. 
38. Piper san-Iorenzanllm Tre!., nom. nov. 
A.rtanthe tritlerv~a Mlquel; Seem. Bot. Voy. Herald 200. pl. ,,1. 1856. 
Piper trl"",'"" C. DC. in DC. Prodr. 16': 292. 1861), Nol P. IrI..."..," Vahl, 
1804. 
TypE LOCALITY: San Lorenzo, Veraguas (BeemoMl 1211t the type). 
RANGE: South-eentral Panama. 
39. Piper subtrlnerve Trel., sp. nov. 
A shrub, 2 to 3 meters tall, essentially glabrous; flowering internodes very 
slender, moderately short, finely wblte-dotted; leaves lanceolate. very gradD' 
• 
aU, attenuate or acnmlnate, characteristically somewhat unequally acute at 
base, moderately small (3 4X9-14 em.), pinnately nerved from below the 
middle, the nerves "bout 4X2 with the second pair greatly prolonged, glossy 
green, obscurely granular beneath; petiole very short (2 mm.), not winged; 
spikes opposite the leaves, moderate (4X5:i mm.); peduncle short (10 mm.), 
glabrous or sparsely pHose upward; bracts conspicuous, trtangular..subpeltate. 
elUate: ftowers sessile. perfect; berries BubprlsmatJc, glabrous; stigmas 3. 
• 
30 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Type In the U. S. National Herbarium. no. 678676. collected on Taboga 
Island, Panama. by H. PUtter (no. 3618). 
RANGE: East-central Panama. 
PANAlIA.: Corozal Road, Standley 26854. Juan Diaz. Standley 30600. Las 
Sabaoas, Bro. Oole,"ne 110. Las Saban8s to MaHas HernAndez, Bt4fldleV 
31836. Punta PaUllIa, Piper 5410. Rio Tecumen, Standtey _. Tumba 
Muerta Roo.d, Standley 29710. Taboga Island, Standley 27869. 
39a. Piper subtrinerve harveyannm Tre1., var. nov. 
Durers chiefly in having the spikes more dlsttnctly nnDulate. wJth the large 
triangular pale bracts alternating with the dark ovaries. 
Type In the U. S. National Herbarium, no. 1,180,427, collected along the 
Tapia River, ProvInce of Panama. Panama, by W. R. Maxon and A. D. Harvey 
(no. 6684). 
RANGE: East-eentral Panama. 
PANAMA: Tapln River, Mazon ~ Harvev 6734. 
40. Piper hirtelllpetiolum C. DC. Smiths. MIsc. ColI. 71': 3. 1920. 
TYPE LOCALITY: David, Chiriqut (PUti.er 2832 [not 2932]. the type) . 
RAN01t: Southwestern Panama. 
41. Piper tapianum Tre1., sp. nov. 
A shrub, 3 meters tall, nodose; ftowering internodes slender. moderately 
short. at first sparingly v11lous; leaves lanceoiate or lance-elllpUc, caudate, 
slightly lnequilaterally acute at base, moderate (5-6.5Xl5-16 em.), pinnately 
nerved trom below about the middle, the nerves about 4X2, glossy dark green 
and glabrous above, paler and with the nerves somewhat upeurved-pubescent 
beneath; petIole short (about 5 mm.), winged. belOW, hirsute; spikes opposite 
the leaves. rather slendf>r amI short (10 mm.). sparsely hlrsutej bracts COD· 
splcuous, roundish'8ubpeltate, v~y gray-ciUate; o.owers sessile, perfect; ovary 
depressed, gJabrous ; ~tigmas 3, filiform, sessile. 
Type In the U. S. National Herbarium, DO. 1,180,449, collected aloDg the Tapia 
River, Province of Panama, by W. R. Maxon and A. D. Harvey (no. 67(9). 
RANGE: East-central Panama. 
PAN AliA: Rto Tapia, Standley 26122. 
42. Piper turbeosEi Trel., sp. nov. 
A shrub; flowering internodes slender and rather short, loosely hairy or 
glabrate; leaves undulately lance·oblong, acuminate., obUquely rouuded at base 
or with one side shorter nnd subacute, moderately large (5-5.5Xl6-20 em.). 
plnnntely nerved from below the mhldl(', the nen'es 4 or 5 X 2, sparsely vUlous 
on the nerves beneath; petiole very short (5, or 4+4 mm.), Dot winged. somewhat 
hairy; spikes opposite the leaves. small (3 X20 mm.). mucronate ; peduncle 
slender, short ·(5 mm.), loosely pubescent; bracts inconspicuous, sublunate, 
c1l101ate; flowers sessile, perfect; ovary 6ask·sbaped, attenuate Into a style; 
stigmas 3, large. 
Type In the Torrey Herbarium, New York Botanical Garden, collected at 
Turbo, Colombia, by Schott (no. 3). 
RANGE: Western Colombia; and adjacent CoMo, Panama (1) 
43. l'Iper davldlanum C. DC. Smiths. Misc. CoIl. 71': 9. 1920. 
TYPB LOCALITY: Dnvid, ChirIqui (PitUer 2837, the type). 
RANGE: Southwestern Pannma. 
44. Piper call1bra.toum C. DC. Smiths. 1II18C. Call. 71': 13. 1920. 
Piver chamu.sonia rubellibractevm C. DC. Smiths. Misc. CoIl. 71': 13. 1920. 
1Pi·per aequale Auct. as to Panama. 
TypB LOCALITY: El Boquete, Chiriqui (Pittier 2940, the type) . 
CUUIQut: EI Boquete, Pi-tiler 2899, type ot P. cAam",onis rube'Ubract~m; 
KUlip 3549, In part. 
TBEI.EASE PlPERACEAE OF PAN AMA 
RANGE: Southwestern Panama. 
4~. Piper nltidlfolinm C. DC. Smiths. Misc. Coil 71': 14. 1920. 
TypE LOCALITY: El Boquete, Chiriqui ( Mazon. 4943, the type). 
RANGE: Southwestern Panama. 
46. Piper erectamentum C. DC. Smiths. Misc. Coil. 71': 10. 1920. 
TtPli LOCALITY: Cerro de 10. Horqueta, Chiriqui (PUti«' 3191, the type). 
RANGI!:: Southwestern Panama. 
·47. Piper papyracenm Trel., sp. nov. 
Piper pseudo·propinquum Auct. as to Panama. 
31 
A small, essentially glabrous tree; flowering internodes moderately slender 
and short, drying green, gleen-granular; leaves ovate-elliptic, acuminate. sub­
equllaterally acute at base, moderately large (6Xl4-9X20 em.), pinnately 
nerved trom below the middle, the nerves 5X2, drying green and papery; 
petiole rather long (2-3 em.), winged to above the middle; spikes opposite the 
leaves, moderate (5X60-SO mm.); peduncle short and thick, more or less 
velvety; bracts triangular~subpeltate, pale, ciliolate; flowers sessile, perfect; 
berries ovoid, with a short stout style; stigmas 3, large. 
'l'ype in the U. S. National Herbarium, no. 677410, collected at EI Boquete, 
Chiriqui, Panama, by H. Plttler (no. 3(23). 
RANGE: Southwestern Panama. 
48. Piper amphio::r:ys Trel.. ap. nov. 
A shrub; flowering internodes slender and short, at most very obscurely and 
transiently puberulent, scarcely granular; leaves elliptic, bluntly acuminate, 
equilaterally acute at base, rather small (4X10-7.5X15 em.), 8ubmultiple~ 
nerved from below the upper third, the nerves 3 or 4X2 with the lowest pair 
essentially basal, slightly puberulent on the nerves beneath; petiole very shOrt 
(5 mm.), channeled, not winged, glabrous; spikes opposite the leaves, slender 
and moderately elongate (2X50 mm.); peduncle slender, short (5 mm.), 
glabrous; bracts inconspicuous, round-peltate, cillolate; flowers sessile, pertect; 
berries minute, somewhat triquetrously obpyramidal, glabrous; stigmas 3, 
minute, sessile. 
Type in the U. S. National Herbarium, no. 670340, collected between Lorna 
de Ie. Gloria and Fat6, Province of Co16n, Panama, by H. Pittier (no. 4239). 
RANGE: East-central Panama. . 
48a. Piper amphio::r:ys laurifollum TreJ., var. nov. 
Dift'ers in its Darrow leaves (4X9---tiX13 em.), with the second pair ot midrib 
branches opposite and about 1 cm. from the base. 
Type in the U. S. National Herbarium, no. 71tS872, oolleeted at GaragarA, 
Sambl1 Basin, Dari~n, Panama, by H. PUtier (no. 5598). 
RANGE: Southeastern Panama. 
49. Piper subcaudatum Trel., sp. nov. 
A shrub, nodose; flowering internodes rather slender and short (2-3 em.), 
transiently crisp-bairy below the petioles; leaves obUquely elliptic or elliptic­
obovate, subcaudately long-acuminate, subobtnse at the narrowed base, rather 
large (6--7.5X14-16 cm.), pinnately nerved from below the middle, the nerves 
5 or OX2, more or less uppressed-hairy on the nerves beneath; petiole very 
short (5+3 mm.), somewhat crisp-hairy on the back, not winged; spikes 
opposite the leaves, as yet very young and small (lXIO mm.); peduncle 
sparsely velvety; bracts rounded-subpeltate, ctliolate; flowers sessIle. 
Type in the U. S. National Herbarium, no. 715886, collected at Garngarti., 
Sambfi. Basin, Darien, Panama, by H. PUtier (no. 5612). 
RANGI!:: Southeastern Panama . 
• 
, 
• 
• 
32 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
50. Piper subnudlbracteum C. DC. Smiths. Misc. Coli. 71': 14. 1920. 
TYPE LOCUJTY: Cerro Vaca to Hato del Loro, Chiriqui (PUtier 5390, the 
type). 
RANGE: Southwestern Panama. 
51. Piper sambnanum C. DC. Smiths. Misc. Coli. 71': 13. 1920. 
TYPE LOCAUTY: Garagari, Samhtl Basin, Darien (PUtier 5663, the type). 
RANGE: Southeastern Panama. 
DARIEN: Garagnrti, Pi-ttler 5726 . 
52. Piper storkii Trel., sp. nov. 
A shrub. essentially glabrous j fiowering internodes slender and short, green, 
granular; leaves lance elliptic, acuminate, very inequllateral, acute at base 
or the longer side somewhat rounded, moderate (4Xl0-6X15 em.), pinnately 
nerved from below about the middle, the nerves 4 or 6 X 2, drying green, rarely 
with a soft hair above and the nerves very obscurely appressed-pubescent be­
neath; petiole very short (scarcely 5 mm.), granular, not winged above the 
base; spikes opposite the leaves, dull red, 2X 60 mm.; peduncle very short 
(li mm.) and relatively thick; bracts roundlsh-subpeItate, glabrous; flowers 
sessile, perfect. 
Type in the U. S. National Herbarium, no. 1,166,892, collected at Rubber 
Tree Station, Bocas del Toro, Panama, by H. E. Stork (no. 124). 
RANGE: Northwestern Panama. 
tiS. Piper palUdibractenm C. DC. Smiths. Misc. Coli. 71': 8. 1920. 
TYPE LOCALITY: Puerto Obaldfa, San Bias Coast, Province ot Col6n (PUtter 
4396, the type). 
RANGE: Northeastern Panama. 
li4. Piper williamsii TreI., ap. nov. 
A shrub, 2 to 3 meters taU, nodose; flowering internodes slender and short 
(2-3 cm.), grayish-tomentulose; leaves inequilaterally lanceolate or lance­
elUpUc, acuminate, obliquely rounded at base with one side slightly shorter 
and more ncute, moderately lnrge (5--7Xl3-17 cm.), pinnately nerved from 
below the middle or upper third, the nerves 5 or 6X2, cancellate-veiny, lepi· 
dote but essentially glabrous above, appressed·tomentulose on the nerves 
beneath; petiole rather short (10+3 mm.), tomentulose, not winged; spikes 
opposite the leaves, slender, 2XOO mm., refracted·mucronate; peduncle slender 
and short (10 mm.), appressed-pubescent or tomentulose; bractslunately sub­
peltate, ciliolate; flowers sessile, perfect. 
Type in the U. S. National Herbarium, no. 678344, collected at MarraganU. 
Panama, by R. S. Williams (no. 986). 
55. Piper lucigaudens C. DC. Smiths. Misc. ColI. 71': 10. 1920. 
'.PVPE LOCALITY: Rfo Indio de Gattin, Canal Zone (PUtier 2791, the type). 
RANGE: East-central Panama. 
56. Piper viridicaule Trel., gp. nov. 
A shrub, 2 meters tall; llowering internodes green, slender and rather 
elongate, vUlous; leaves lanceolate, gradually sharp·acuminate, inequilaterally 
cordulate with the longer side covering the petiole, moderately small 
(3.5-4XIO-I2 em.), pinnately nerved from below about the middle, the nerves 
about 5X2, green, dull, sparsely villous and very white-scabrous above, paler 
and appressed-white-hirsute beneath especially on the nerves; petiole moder­
ately short (8+2-20+2 mm.), white-villous, winged below; inflorescence 
unknown. 
TUTJ!ASFl PlPERACEAE 01' PANAMA 33 
'1y;e in the U. S. National Herbarium, no. 1,215.801, collected on Barro 
Colorado I8Iand, GatQn Lake, Canal Zone, Panama, by Paul C. Standley (no. 
314(2). 
RANGE: East-central Panama. 
117. Piper pereubulatnm C. DC. Smiths. Misc. Call. 71': 4. 1920. 
'liP! LOCALITY: Loma de 18 Oloria. Fat6, CoMo (P1ttier 4240, the type). 
RANGE: East-central Panama. 
liS. Piper colonense C. DC. Smiths. Misc. Coli. 71': 11. 1920. 
TYPE LOCALITY: Dos Boeas, B(o Fat6, ColOn (PUtler 4221, the type). 
RANGE: East-central Panama. 
119. Piper panamense C. DC. Journ, Bot. Brit. & F or. 4: 216. 1866. 
Piper altrlfollum paMmen.e C. DC. Candollea 1: 114. 1923. 
'1 iPE LOCALITY: Chagl'es, Province of ColOn (Fendler 270, the type). 
RANGE: Northern Panama. 
BocA8 1)l3. TORO: Laguna de Chiriqui, Hart 124. 
60. Piper septupllnerv1um (Mlquel) C. DC. 10 DC. Prodr. 16': 318. 1889. 
Arlantll_ .eptMpli1lU'1Jla Miquel in Seem. Bot. VOY. Herald 199. pl. ~O. 1866. 
TIPI: LOCALITY: "Panama " (Seemann 1066, the t.rve), 
RANGE: Northern (1) Panama. 
CoWN: Porto Bello, PUtler 2477. 
61. Piper kllllpi Trel., sp. nov. 
A shrub; flowering internodes moderately slender and short, dlngy-8Ubto­
mentose; leaves lanceolate, gradually acuminate, obUquely subacute at base, 
moderately smaH (4-5Xll-15 em.), pInnately nerved from below the mIddle. 
the Impressed nerves 5 or 6 X2, dull. glabrous and scarcely scabrous above. 
appressed-hairy on the nenes beneath; petiole very short (5 mm.), crisp­
hairy; spikes opposite the leaves. elongate (3 X lOO mm.), booked-mucronate; 
peduncle short (5-10 mm.), glabrate; bracts triaugular-subpeltate, elllate; 
1l0wers sessile, perfect; stigmas 3, minute, sessile. 
Type In the U. S. Nationa} Herbarium, no. 1,012,175, collected near EI Bo· 
quete, Chiriqui. Panama. by E . P. Kllllp (no. 3549). 
62. Piper killlpi calderanum Trel., var. nov. 
Ditfers In somewbat larger (5 X l1S cm.) nnd rugose, cordulate leaves, and 
stralght·mucronate sp1kes. 
Type In the U. S. National Herbarium, no. 1,266,034. collected along Bfo 
Caldera. above EI Boquete, Chiriqui, Panama, by E. P. Klllip (no. 3644). 
RANOE: Southwestern Panama. 
63. Piper infraluteum Trel.. sp. nov. 
A shrUb, glabrous j flowering internodes slender and short; leaves laDce­
oblong or ovate-oblong, acumlnate, Bubequally acute at base, small (3-4 X IO 
em.), pInnately nerved from below the middle, the nerves nbout 4 X 2. drying 
yellow beneath; petiole short (7+2 mm.), not winged; spikes opposite the 
leaves, slender, 2 X 50 mm. before dowering; peduncle slender, short (scant 
10 mm.) ; bracts large, round- or trtangular-subpeltate, cll1~te; flowers sessUe, 
perteet. 
Type in the herbarium ot the New York BotanIcal Garden, collected to 
4. Panama" by Sutton Hayes (no. 798). 
RANOE: Central (?) Panama. 
M. Piper trachydermum Trel.. sp. nov. 
A shrub; flowering internodes sleDller and rather elongate, acabrld; leal'ee 
eWpUe'8ubol'ate or subobovRte, sharp-acuminate, tnequtlaterally obtnse at base, 
moderately large (7-8 X16 em.) , pInnately nerved trom below the upper third, 
34 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
the nerves about 5X2, microscopically white-scabrous, dingy appressed-pubes­
cent on the nerves beneath; petiole short (7+3 mm.) not winged, hlspld; 
spikes opposite the leaves, slender, 2X50-80 mm. before flowering; peduncle 
short (5 mm.), scabrld; bracts rounded-subpeltate, rnsty-cUioiate; flowers 
sessile. 
Type in the herbarium of the New York Botanical Garden, collected in 
.. Panama" by Sutton Hayes (no. 791). 
RANGE: East..central Panama. 
PANAllA: R10 Tapia, Stan.dley 28208. CocLt: Bismarck, WClliamB 343. 
65. Piper cativalense Trel., ap. nov. 
A shrub, 2 to 3 meters tall; flowering internodes slender and elongate, 
villous j leaves eUiptic-ovate or sUbovate. rather abruptly acuminate, inequi­
laterally cordulate, moderate (6.5X12-14 em.), pinnately nerved from below 
the middle or upper third, the nerves about 5 X 2, green, rather glossy and 
sparsely long-hairy above, more villous beneath, especially on the nerves; 
petiole rather short (scarcely 10+2 mm.), loosely villous, winged below; 
inflorescence unknown. 
Type in the U. S. National Herbarium, no. 1,154.579. collected between 
France Field, Canal Zone and Catival, Province of Co16n, Panama, by Paul C. 
Standley (no. 30206). 
RANGE: East--central Panama. 
66. Piper pseudo-cativalense Trel., sp. nov. 
A. shrub, 2 meters tall; flowering internodes slender and elongate, green~ 
sparingly crisp-pubescent; leaves ell1ptic-ovate to oblong, rather bluntly sub-­
acuminate, inequllaterally rounded at base or the broader deeply cordulate, 
moderate (5-6XI2-13 cm.), pinnately nerved from below the middle, the 
nerves 4 or 5X2, dull, crisp-hairy, and very scabrous above, grayer and ob­
scurely granular beneath with the nerves upcurved-pubescent; petiole short 
(scarcely 10+2 mm.), hispid or matted-hairy, winged below; spikes opposite­
the leaves, as yet minute and dingy-canescent. 
Type in the U. S. National Herbarium, no. 1,215,573, collected between 
France Field, Canal Zone, and Catival, Province ot Co16n, Panama, by Paul C. 
Standley (no. 30399). 
RANGE: East-central Panama. 
CANAL ZoNE: Barro Colorado Island, Standley 31288, 31387. Mount Hope­
Cemetery, Standley, 28776. 
67. Piper sperdinnm C. DC. Smiths. Misc. ColI. 711: 1. 1920. 
TYPE LOCAUTY: Puerto Obald(a, San BIas Coast, Co16n (Pittier 4348, the­
type) . 
RANGE: Northeastern Panama. 
COWN: Puerto Obald(a, Pittfer 4301. 
68. Piper diazannm Trel., sp. nov. 
A shrub, scarcely 2 meters tall; flowering internodes slender, moderately 
short, green, sparingly appressed-pubescent; leaves lanceolate to subovate or­
subobovate, gradually acuminate. rounded or subtruncate at base or shal, 
lowly subcordate, with one side sometimes shorter, moderately large (3.~ or-
7X13-16 cm.), pinnately nerved from below the middle, the nerves about 5X2. 
slightly pubescent on the nerves, yellowish green, the paler lower surface 
granular-punctate; petiole short (5 or 5+2-10 mm.), loosely crisp-hirsute or 
subvillous, winged only at base; spikes opposite the leaves, moderate (3X60-00 
mm.); peduncle short (5 mm.), with short appressed white hairs or glabrate; 
bracts rounded-snbpeltate, ciliate; fiowers sessile, perfect; berries tl'iquetrou~ly­
oblong; stigmas 3, minute, sessile. 
TREIJMBE PIPI!BACI!AE OF PANAMA 35 
Type In Ihe U. S. National Herbarium, no. 863136, collected eaat of Juan 
DraB. Province ot Panama. Panema, by E. P. KtIllp (no. 3114). 
RANOE: East-eentral Panama. 
CoWN: Porto Bello, Pitt{er 2428. CAN AL Zo N'E: Cerro Gordo, CuIebra, 
Btan41e1l 26006. Fort Sherman, St(UldleJl 30953, 30974, S0996. Fort San 
Lorenzo, Mazon <f Valentine 7012. Las Cascadas Plantation, Bfafl4lev ~798, 
29625. Matach1n to Las C8scadas, C()Wen 338. Monte Llno. MazOft, 6843. 
Mount Hope Cemetery, Bfarnllev 2SS02. Obispo, SfafldleJl 81671. TabernUla, 
Pitlier 3828. 
69. Piper subdilatatum Trel, sp. DOV. 
A.. shrub, scarcely 2 meters tall; flowering internodes rather slender and short, 
drying dark, at first loosely velvety or crisp-pubescent; leaves oblaoceolate­
oblong, acuminate, unequally truncate or rounded below or abruptly subacute 
at the very base, moderately large (8Xl8-22 em.), pinnately nerved from 
below the middle. the nerves I') or 6X2, scabrld and traDsiently or obscureJy 
appressed-pubescent above and velvety on the nerves, and somewhat appressed­
hairy or loosely velvety beneath; spikes opposite the leaves. as yet youPg and 
scarcely 1 X 10 mm .• eanescent; flowers sessUe. 
Type In the U. S. National Herbarium, no. 863177, collected on Taboga 
Island. Gull ot Panama, Panama. by E. P. Klllip (no. 3194) . 
RAl'fOZ: Soutb-central Panama. 
PANAlU.: Taboga Island, Standley 27019, 27920. 
70. Piper ohaldlanum C. DC. SmIths. Misc. Coil. 71': 7. 1920. 
TYPB LOCALITY: Puerto Obaldfa, San Bias Coast. ColOn (P'iUier 4366, the 
type). 
RANOE: Northeastern Panama. 
71. Piper peracnmlnatllm C. DC. Smitbs. Misc. Call. 71': 9. 1920. 
TYPE LOCALITY: Dos Boeas, Rfo Fat6, Co160 (Pittier 4210, the type). 
RANGz: East-central Panama. 
COWN: Cbagres, Fendler 269. Rfo Indio de Fat6, PUtter 4264. .. PaDama,'~ 
Hallel 771. 
72. Piper sancti .. telict. TreJ .• sp. nov. 
A shrub; ftowerlng internodes rather slender, elongate, nodose, retrorsely and 
somewhat hamately gray-hlsp1d: leaves elUptlc-oblanceolate or subobovate, grad­
ually acuminate, InequlIaterally rounded at base, rather large (~10X lrs-20 
em.), pinnately nerVed from below the middle, the nerves 6 or 7X 2. m1nutely 
scabrous on both faces and the nerves hirsute beneath i petJole short (5+4 mm.), 
not winged, subretrorsely hlspld; spites opposite the leaves, elongate (3X l20 
mm.) ; peduncle short (5 mm.), hlspid; bracts transversely snbpeltate. dUo­
late; dowers sessile, perted; stigma8 3, mlnnte, sessile; berries small, oblong. 
Type In the U. S. National Herbarium, no. 71ri300, collected at San Fellx~ 
ChiriquI, Panama, by H. Plttler (no. ~124). 
RANGE: Southwestern Panama. 
73. Piper barbinerve Trel.. Bp. nov. 
A shrub; ftowerlng Internodes moderately slender and elongate. tor a time 
dingy crisp-pubescent; leaves ellipUc-lanceolate or oblanceolate. bluntiy acum­
inate, unequally cordulate with the longer side covering the petiole, moder­
ately large (6-7.6 X I4-16 em.), pinnately nerved trom below the middle. the 
nerves at first hairy; petiole short (scarcely over 5+5 mm.). winged to the 
middle, hairy; spikes opposite the leaves. moderate (2 X60 mm. before fiower­
lng) j peduncle slender, short (10 mm.), sp8I'Hely velvety; bracts triangular­
IUbpeltate, pale-cl11ate; ftowers sessile, perfect. 
36 CONTRIBUTIONS FROM 'l'HE NATIONAL HERBAllIUM 
Type In the U. S. National Herbarium, no. 679247, collected above Santa 
Isabel, Culebra River, Province or Co16n, Panama, by H. Plttler (DO. 4148). 
RANGE: Northeastern Panama. 
74. Piper non·retronum Trel., sp. nov. 
A shrub, nodose; flowering internodes rather slender and sbort. huuate with 
upcurved hairs; leaves inequilaterally ellipUc-oblanceolate. sharply acuminate, 
the narrowed base obtuse or wtth the shorter side Bc-ute, moderately large 
(8-9 X 18-20 em.), pinnately nerved from below the upper. third, the nerves 
6 or 7 X 2, white-scabrous above, somewhat bairy beneath 011 the nerl'es; 
petiole moderate (10+7 mm.), bairy, Dot winged; spikes opposite the leaves. 
elongate (3X136 mm.), mucronate; peduncle short (scarcely 10 mm.), rough 
with upcurved pubescence; bracts round- or lunulate-subpeltate, pale-clllolate; 
flowers sessile, perfect. 
Type In the U. S. National Herbarium, no. 677170, collected near Porto Bello, 
Province or 0016n, Panama, by H. Plttler (no. 2429). 
RANOE: Northeastern Panama. 
CANAL ZoNC: Fort Sherman, Standley 31121. 
75, Piper scabrillmbum C. DC. Candollea 1: 121, 276. 1923. 
TYPE LOCALITY: Boca de P8u8randll, Sambti River, Darl~n (PUller 5584, 
the type). 
RANGE: Southeastern Panama. 
76. Piper vllllramulum C. DC. Smiths. MIsc. ColI. 71': 11. 1920. 
TYPE LOCALITY: l,oma de 10. Gloria, Fat6, CoMn (Pittier 4083, the type). 
RANGE: Enst-centraJ Panama. 
CANAL ZoNE: AnCOn. Bro. Celestine 32; Ptper 6028. Balboa, Slan4l6fl 26073, 
29267. Della Vista, Macbride 2732. Cerro Gordo, Culebra, Starullev 26012. 
El Par8fso, Pittier 2534. Empire to Mandlnga, Piper 5514. l!'rlJoles, Piper 
5867. Gamboa, Standley 28022. Las Cascadas Plantation, Standley 25002,20701. 
Mil'afiol'es to Corozal, Pittier 2193. Rfo Pedro Miguel, East Parafeo, Standleu 
29966. Summit, Standley 26937. COL6N: Fort Lorenzo. Rfo Chagres, Piper 
()971. PANAMA: Pacora River, Killip 3123. Corozo.l Road, Panama, 8tandle, 
26843. Sab8118 de Juan Corso, Chepo, PEttier 4537. 
76a. Piper vtlliramulum gamboanum (C. DC.) Trel. 
P ipo, Mspidtuu Da11lboanum C. DC. Smiths. Misc. Coil. 71': 12. 1920, 
TYPE LOCALITY : Gamboa, Canal Zone (Plttier 3410, the type) . 
RANGE: North- and east-cPutral Panama. 
BOCAS DEL ToB.O: Chunguinola Valley. Dunlap 234. CANAL ZoNE: CoroZ8~ 
Stevens 116, 117, 124. Cerro Goroo, Culebra, Standley 25998. Las C8scadas 
Plantation. BtandleJl 29639. Lns Cruces Trail, Standleu 29048, 29199. Obispo, 
Standley 31794. PANAMA: JU8n Dlaz, Standley 30623. Los Saoonas to Mattas 
Hcrnt\ndt'z, Standley 31818. R(o Tapia, Standley 28134. Rfo Tecumen, Stan .. 
lcll265D8. Cumino de 1ft GraDja, Bro. Heriberto 165, 
77. Piper elongatum Vahl, Enum, PI. 1: 812. 1804. 
TYPII LOCAl·I'lI: DrazU (1). 
RANGE: Around the northeastern pert of South America and, In aimllar 
If separable forms, through Central America Into Mexico. 
CANAL ZONE: GatOn, Standley 27256. Fort Sherman, MIJ~ft d VaJentfnO' 
6992; StatUllev 30941. CoLON: Fat6 to Playa de Damas, PUffer 3930. Fort 
Lorenzo, p .ipe." 598t5. PANAMA.: Tapia River, Alazopt, tl Har"1Je1J 6671. Tumba 
:Huerta Road, StamUelf 29803. 
... 
, 
• 
TRli:LIIIA8E PIPEIlACEAE 01' PAN 'M~ 
T7L Piper alongatum la.vitoltum (C. DC.) Trel. 
Piper adM""" ... 160";(011 .... C, DC. Smltbs. M .... CoIl. 71': 8. 1920. 
TYpm LOCAUTt: Porto Bello. Col60 (Pi/tier 2438. the type). 
RANOE: Nortb-ceotral Panama. 
?'rb. Piper eloaratum braehyarthrum Trel., var. nov. 
37 
Dltrers trom the type with sparsely 8ott-pubescent stem tn bavlng the latter 
very hinote-villous. 
Type In the U. 8. National Herbarium, no. 675501, collected near El Boquete, 
Chirlquf, Panama, by Wllllam R. :Maxon (no. 5139). 
RANGE: Panama. 
C .... AL ZoNE: Cblva·Chlva Trail. P,per 5749. 
78. Piper chagresJAnum Tret., sp. DOV. 
Piper Ckfuncvm Auet. 8S to Panama. 
A shrub; flowering internodes moderately slender and rather sbort, sparsely 
tawny crisp- or velvety-pubescent; leaves lance-oblong or elliptic-oblong, bluntly 
subacuminate, rounded or 8ubtruncate at base wIth one side sometimes a Utt1e 
shorter, rather large (5.6-6.5Xl~19 em.), pinnately De"ed from below the 
middle, the nervet:t 6 or 7X2, somewhat scabrous and hairy on the nerves above, 
ru.ther sparingly appressed-pubescent beneath; petiole moderately elongate 
(1~15+5 mm.), winged below the mJddle, dingy velvety-hirsute; spikes op­
posite the leaves, straight, moderate (3--4XM-SO mm.); peduncle short 
(10 mm. ), loosely velvety; bracts inconspicuous, lunulate-truncate, elllate; 
flowers sesene, perfect, stigmas 2 or 3, lInear, sessile; berries minute, trigon­
ousIy obpyramidal, glabrous. 
Type in the U. S. National Herbarium, no. 42617, collected at Chagres, 
Province ot Co16n, Panama, by A. Fendler (no. 268). 
RANOE: East-central Panama. 
COLON: Rro Indio de Fat6, Pittier 4271. 
79. Piper changuinoJanum Trel., sp. nov. 
A shrub, scabrous througbout, nodose; Internodes shOt't and rather slender, 
drying sulcate, densely but tlnely gray-hispid; leaves IneqoUaterally or Bub· 
falcately oblong, acuminate, rounded at base and shorter on one side, rather 
small (4X15 em.), pinnately nerved from below about the mtddle, the Derves 
about 5X2, the lowest somewhat approximate, sUghtly rugose, duU, paler and 
granular-punctulate beneath and appressed-puhescent on the Denel; petiole 
short (scarcely 5+5 mm.), winged at base, at ftrst appressed-halr7; spikes 
opposite the leaves, 2 becoming 3X80--100 mm.; peduncle rather short (10 mm.), 
appressed-blsptd; bracts small, round-subpeltate. clUate: dowers sea-llle, per­
fect; berries short-oblong, mostly elongate with the rachis, glabrous; stigmas 
8, minute, sessile. 
Type In the U. S. Natlonal Herbarium, no. 1,205,153, collected In the Chan­
,",'\1loola Valley, Bocas de-I Toro, Panama, by V. C. Dunlap (no. 283). 
RANGE: Northwestern Panama. 
80. Piper tatoa.num C. DC. Smiths. Misc. eoll. 71': 7. 1920. 
TypE. LOCALITY: Fat6 to Playa de Damns, Co16n (PUtier 8925, tbe type). 
RANGE: East-central Panama. 
81. Piper linearitolium C. DC. Linnaea 37: 3M. 1872. 
TUB LOC.t.I,ITY: Hacienda Santa Rosa, Costa Rica. 
RANGE: Costa Rica to central Panama. 
CHlRIQut: EI Boquete, Mazon. 5140. CoWN: Rfo Fat6, PHifer 3881. 
• 
• 
38 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
82. Piper tabogannm C. DC. Smiths. Misc. ColI. 71': 4. 1920. 
TYPE LOCAI,ITY: Taoop Island. Gulf of Panama (Pitff.er 3520, the type). 
RA.NOE; South--central Panama. 
88. Piper vitabundum 'I'cel., sp. noy. 
ClImbing; flowering internodes fatber slender and long, sparingly and tran­
siently appressed-hairy; leaves BUbe1l1ptic, l:mbacute at both ends or the base 
father rounded. moderate (4.5XIO--12 em.), pinnately Derved from below the 
middle, the nerves about 4X2, closely lepldote on both faces, the nerves ap. 
pressed-pubescent beneath; petiole short (l>+3 mm.+, sparsely hairy, scarcely 
winged; spIkes opposite the leaves, normally (1) arched. 3XSO mm.; peduncle 
short (scarcely 10 mm.), glabrate i bracts roundish-subpeltlite, the very narrow 
pale margin ciliolate; flowers perfect, sessile; berries 8ubcubical; stigmas 3, 
minute. sessile. 
Type tn the U. S. National Herbarium, no. 1,205,222. collected io the Cban­
pinola Valley. Bocas del Toro, Panama, by V. C. Dunlap (no. 338). 
84. Piper reptabundnm C. DC. Bot Gaz. 70: 169. 1920. 
TYPE: LOcALITY: Shirores, Talamanca, Costa Rtca. 
RANOE: Costa Rica and adjacent Panama. 
BOCAS nn TORO: Sibubl Falls. Rotolee 372. 
85. Piper breve C. DC., sp. DOV. 
A low shrub; flowering internodes slender and short, cancscently suOOp­
pressed·hairy when young; leaves lanceolnte or lance-oblong, gradually pointed. 
rounded. at base witb one side shorter, moderately small (3-4JSX9-13 cm.), 
pinnately nerved from below the middle, the nerves about 5X2, softly ap~ 
pressed-pubescent on both faces or becoming slightly scabrid, not rugose; 
petiole short (3+3 mm.). hairy, winged to the middle, spikes opposite the 
leaves, relati"ely rather tblck and sbort (4 X45-00 mm.); peduncle rather 
short (10 mm.), loosely pubescent, bracts lunate- or trlangular-subpeltate, 
ciliate; flowers sessile, perfect; stigmaa 3, sessile; berries subglobose, glabrous. 
TyPE tn the U. S. National Herbarium, no. 678858, collected between Gamboa 
and Cruces, Canal Zone, Panama, by H. PUtler (no. 3787). 
RANOE; East-central Panama. 
CANAL ZoNE: Cerro Gordo, Culebra, Standley 2597f1, 20018. Empire to 
Mandinga, Piper 5523. France Field to Catival, ColOn, Stevens 981. COLON: 
FatO to Playa de Damas, Pitlier 3923. FOl't Lorenzo, Piper 6944. 
85a. Piper breve palUdineurum TreT., var. nov. 
A rather larger, more scabrldulous-leaved torm, with tbe prominent nerves 
paler beneath, the spikes scarcely longer than In tbe type. 
Type in the U. S. National Herbarium, no. 1,154,564, collected between 
France Field, Canal Zone, and Catival, ColOn, Panama, by PaulO. Standley 
(no. 30164). 
RANOE: East-central Panama. 
CANAL ZoNE: AneOn Hill, Killfp 12078; Standley 25203, 26382. Brazos 
Brook. 81even6 752. Cbtva·Chiva Trail. Mazon 4 HarooV 659. Culebra, 
P/ttler 844~; 81""", .. 993. Gamboa, 81a"1<>1/ 28374, 28481. Joan MlDa, PI".,. 
5692. Obispo, Standley 81764. PANA1,(A.: Cbepo, Pit tier 4539. Juan Franco 
Race Trnck, Panama, Standley 27713, 27734, 27777. Tumba Huerta Road, 
Panama, Standley 29724. MaUas Herntlodez, Standley 28007. Las Snbanas to 
MaUns HernAndez, Statt4lev 31832. Las Sabanas, Bro. CeZe6tifte 110. Punta 
PaltUla. Standley 2630S; St ..... 349. 
TBEI.EABE PIPERACEAJI 01' PAN AMA 39 
85b. Piper breve pseudo-dumeticola Trel., nom. nov. 
Piper dtlmetlcola "" .... " .. " •• C. DC. Smiths. Misc. Coli. 71': 7. 1920. 
A form wttb lance-oblong, rugose. and scabrld leaves. the spikes scarcely 40 
mm. long. 
TtPm LOOAI.ITY: QUi, Cocl~ (Pittier:i034, the type), 
RANOI:: East-central Panama. 
85c. Piper breve beUae-vlatae Trel., var. nov. 
A torm with lanceolate-oblsoceolate leaves 7X16 em. 
Type In the U. S. National Herbarium, no. 1,167,181, collected at Bella 
Vista, Panama, by E. P. KIIUp (00. 1204li). 
RANGE: East-eentral Panama. 
PAN.UtA: Bella Vista, Kililp 12004; Piper 5331. 
Md. Piper breve divaricatum Trel., var. nov. 
A spreading form with lance elliptic or subovate leaves, and spikes becom. 
Ing 00 em. long. 
Type in the U. S. National Herbarium, no, 1,180,701, collected at Fort 
Shelman. Canal Zone, Panama, by William R. Maxon and A. T. ValentiDe (no. 
6986) • 
RANGE: East-central Panama. 
CANAL ZoNB: Fort Shermao, 8taftdlell 30918, 30982, 31149. 
Me. Piper breve paeudo-tatoanum Trel., var. nov. 
A distinct form with narrow leaves 4 X lti em .• and elongate spikes 3 to 4 by 
90 mm. 
T.fpe in the U. S. National Herbarium, no. 678777. collected at Gamboa, 
Conal Zone, Paoama, by H. Plttler (00.3700). 
RANGE: East-central Panama. 
CANAL ZoNE: Balboa. Standley 25420, 25565, 25603. Corozal. Standlefl 27341. 
Fort Raodolpb, Ma.,,,,, 6 Harvey 61)04, 6543; 8tanlll"1l 2861)9. Margarita 
Swamp, Mazon. If Harvell 7055. PANAIlA: Juan Dtnz, Standlev 80631. Rfo 
Tecum:en, Standley 26551, 29451. Tapia River, Maa-on 6 Haru6V 6620, 6626; 
8taMI"" 28103; 8t.".,.. 1033. 
86. Piper pervelutinum Trel., sp. DOV. 
A shrub, nodose; ftowerlng internodes short, dense1y red·velvety; leaves 00· 
long, 8ubacuminate, rounded at base with one side shorter, rather 8Dlall 
(5Xl2-15 em.), pinnately nerved from above the middle, the nerves 6X2, 
gradually approximate downward, rugose, gt~sy and very scabrous above, dull 
and red·hairy beneath; petiole abort (8+2 mm.), not W"inpd, red.bairy; 
sp1Jr:es opposite the leaves. wh11e very young 2X 15 mm.; pedoncle short, vel· 
vety j braets round·subpeltate. elliate; flowers 8e88l1e. 
Type in the herbarium ot the New York Botanical Garden, collected in 
., Panama" by Sutton Hayes (no. 796). 
RANOE: Panama. 
87. Piper pilosiusculnm Optz in Presl, ReI. Baenk. 1M. pl. 29. 1830. 
'lfP!: LOCALITY: .. Panama" (Haenke, the type). 
RANGE: Panama. 
• 
88. Piper hqesil Trel., sp. nov. 
A. shrub (?), at flr8t densely rusty-tomentose; leaves obliquely ovate, BUb-
• 
acuminate. subcordate, peltate, fully 20 by 30 ClD., 8ubmu1tlpl~nerved from be--
low the middle, the nerves 6 to 8X2, glabrate between the velvety nerves above. 
Tetvety between tbe balry nerves beneath, very bullstely rugose; pet101e elon· 
cate (fDl17 7 em.>, rusty·tomentose, winged; spikes large (4 X200 mm.); 
40 CONTRIBUTIONS FBOM THE NATIONAL HERBARIUM 
peduncle moderate (3.6 em.), puberulent and somewhat rusty-hairy; bracts trl ­
angular~8ubpeltate. bairy; flowers sessUe, perfect; ovary round-ovoid; stigmas 
8, .... 11 • . 
Type In tbe Torrey Herbarlum'ot the New York Botanical Garden, collected 
In" Panama" by Sutton Hayes (no. 760). 
RANGE: Panama. 
89. Piper veraguense C. DC. in DC. Prodr. 161: 294. 1869. 
TYPE LOCALITY: .. Costa Rica and VeraguQ" (Wor3cew\cz, the type In the 
Boissier Heroorlum). 
RANGE: Northwestern (7) Panama. 
110. Piper maxonll C. DC. Smiths. Misc. Coil. 71': 16. 1920. 
TYPE LOCALITY: EI Boquete, Chiriqui (M a;1)on 5MO, tbe type). 
RANOE: Southwestern Panama. 
CBlllIQU(: Rfo Ladrtllo, (l,bove El Boquete, Mazon 3163. 3164. 
91. Piper aurltum H. B. K. ~ov. Gen. &. Sp. 1: 54. 1816. 
TyPE LOCALITY: About Jalapa, Veracruz, Mexico. 
RANGE: Through Central America to Colombia in a number of scarcely sepa­
rable forms, the southernmost chiefty of the large-leaved variety s6emannianvm 
(!.Ilquel) Trei. (Artanth .... ,,,,,,nol/alta Miquel). 
PaoV1NCB (1): Beemann. 37, 1069. BocAS DEL ToBO: CbaDguinola Valley, 
Dtu,Jap 403. Laguna de Cblrlqui, Hart 125, 126. CHI&lQvl: San Felt!:, Pit­
tier ts159. CANAL ZoNE : Fort ShermaD, Jla~n d Valentine 6983; 8tandlev 
3093 j Stevens 263. GatOD, Standley 27244; 8teveWJ 832. Gorgona to GatOn, 
P4.Uler 22~. Monkey HUI, Gowen 22. Nuevo limOn, Jia;rQ1l. 6898. CoWN: 
Fat6 to Playa de Domas, Plttie!' 3933. Porto Beilo, Piliie!' 20141. Puerto 
Ob.ldla, Piliie!' 4380. RIo F.t6, Pilli"r 3909. Santa Isabel, Piliie!' 4177. PAN­
.u ... : Chepo, Pittie!' 4700. DAlUtN: Boca <Ii! Cupe, William. 78!5. 
92. Piper impedale (Mlquel) C. DC. in DC. Prodr. 16 ': 339. 1869. 
,Ar'a"tlle Imperia/II Miquei in Seem. B\)t. Voy. Heraid 198. 1800. 
Piper portobeUeme C. DC. Smiths. Misc. Coil. 71': 6. 1920. 
TyPIC LOCALITY: Cttco.gual Island, Dari~n (Seemnnn 1068, tbe typc). 
RANGE: East-central PaDama. 
CANAL ZoNE: Barro Colorado Island, 81andlell, 81273. Frijolest StandleJl 
27441. Frijoles to Monte Llrlo, KUli" 121~. CoLON: Porto Bellot Mazon 5795. 
Piper 8Olfcfnttn1o Opiz (in Presl, Ret HRenk. 100. 1830), ascribed when pub· 
IIsbed to Mexico Rod PftD8ma, Is clearly labeled at Prague as trom the Marl. 
anne blands, and 80 is to be dismissed definitely from nil consideration of the 
AmE:ricau fiora. 
4. PEPEROMIA Ruiz & Pay. 
KEY TO SPBC1ES 
1. Leaves alternate (the uppermost sometimes subopposlte because ot shortened Internodes) _________ ___ ____ ____ ____ ___ ______ _________________________ 2. 
Leaves opposi te or whorlcd ____ ___ ___ ________________ __ ____ ___ .. _______ AI. 
2. Leaves, or some of them, obviously pcltate; spikes never panlculate ___ ___ =1. 
T.eaves Dot peltate (the margin exceptlonully barely crossing the petiole) __ 7. 
3. Stem short and few-lea\·ed __________ __________________________________ 4. 
Stem elongate and leaty _____ _______ ____ _ ._____ .. __ . ___ .. __ .. ___________ __ 0. 
TREt,PAAI') PlPERACEAE 01' PAN uu 41 
., T.ea ... over 10 em, Iong ______ __ __________________ _______ 3fI. p, vutepta. 
U8.Ve8 .. to 6 em. loog _______ __ ____________________ _______ ______________ 5. 
5, ' ...... .ouoo"'.ate _______ __ h ______________ _ _ _ h h _hh ____ -8' p, k1IUpi. 
Lea ... elllptlc-o.ate ______ h __ h _____ h ___ h _ __ hhh __ A, p, !>3menod •• , 
6. Stem 8toOt; splke8 'ar"' ___ h h ___ ______ h ___ _ ___ h_h ___ 31, p, cordulata. 
Stem slender; spikes smaIL ________________________ 80. P. corduJ-tltormis. 
7. Spikes panlcled on a eeape; leaves larse ___ _ ...:.:. ____ ____________________ ___ B. 
Spikes subpaDlcled on the 'eafy stem ______________ ___ 38, p, oblonglbacco, 
Spikes 80lltary or palred __ ____ __ _________ --- ___ _______________________ 9, 
8. Leaves laDceolate. acuten ____ n ______ n ____________ n __ 44. P. mamefan&. 
Leaves 8ubobovate, acute _______________________________ 37. P. wil11am.l1. 
9. Leaves palmately Dened (the inner nerves exceptionally eon1iueDt toward the baae) _______________________________________________________ ___ 10. 
Teaves pinnately nened _____________________ ______ ___________________ 30. 
10. Leaves appreciably obtuse at base or cordate ______ :.. ____________________ ll. 
Leaves appN!dably acute at base ___________ ______ ____________________ 21. 
11. Lea"es at most elUate or obscurely puberulenL _____________________ __ 12. 
T eaves eTidenUy pubescent j creepingj leaves smalL _______________ ___ _ 20. 
12. Btem sbort or stouL ___ _______ ___ __ __________________ _____________ __ l3, 
Stem elongate and rather slender ________________ ____________________ 16. 
18. Leaves tew, at end ot the stem ___________ ____________________________ 14. 
l.ea .... distributed along tbe stem _____ _____ __ ____ ____________ ______ __ 15, 
14, Leaves ronnd"' .. te _______ _________________ ___ ____ __________ S, p, IdllIpL 
leaves eUlptlHvate ____ _______________________________ 4. P. h,menod.e •. 
1~, Petiole thick and 8bort _______ ________ ____ ________ ______ Sl, p, eordulata. 
Petiole slender and elongate _______________________ 2. P. brevtpeduncula. 
18. Leaves 4: to 6 em. 1011&; plant prostrate or pendenL ___________________ 17. , 
Leaves acarcely halt 88 long ___________________ ______________ ' _________ 19. 
17. I,eaves round, cordate ________________________ ____ 30. JI. eordulatuormie. 
Leaves obion,. Dot cordate ____________________________________ ________ 18. 
18. Spikes oPpo81te tbe '08 .. 8 ______________________ ____ 115, p, portobellensls, 
Spikes axUlary ____________________________________________ ,.26. P. elata. 
19. Plant Creeping; leaves obovate _______________ ____ ____ 15. P. panameDsis. 
Plant erect; leaves round-ovate _____________ , ___ ____ __ , ____ I. P. pelluclda. 
20. Leaves ovate __________ __ _________________ ____ ______ 29. P. urocarpoldes. 
Leaves round·subcordate __ __ ____________________________ 28. P .• CIIDdens. 
Leaves ronnd·subobovate __________________ _______ ___ 13. P. I'OtuDditoUa. 
21. T ... a"ea at most elUate or obscurely puberulenL _____ _______________ ____ .22. 
Leaves evidently pubeBCeoL _________________________ -M. P. ob8curtfoUa. 
22. T.eaVe8 gt"imular-puuctulate_ n ____ n __ __ _ __ n _ __ _ __ ________________ _ _ -23. 
Lea'e8 not granular ____ _________ ___ __ ___ _________ _________ ___ .. _______ 28. 
23. :Leaves 4 to 6 em. long __ ____ ____________ ____ ______ ________ _________ ___ 24. 
I.eaves 2.5 to 4: em. long ____________________ n n n _n _________________ 21. 
Leaves scarcely 2 em. long, blnck-grnnular ____ ________ 21. P. fUlco-ciliata. 
24. :Leaves laneeol·te _______ ____ __ ...; ___________ .;. __________________________ 25. 
:Leaves lance 81 UpUc ______ __ __________________ ______ _________________ _ 26. 
2ri. Leaves merely acute at base ________ _______________ 17. P. parietar1aetolia. 
Leayes long-CUne.ate _________ ________________________ __ 22. P . pel'CW1eata. 
26. Stem slemler ___ __ ______ __ __ _________________________ 18. P. conjunl'en.s. 
Stem rather stout ________ __ __ ________________ _________ 2T. P. paacuieola. 
27. Leaves pale-granular __ n ___ n __________ n ____ _ _ __ _ ___ 19. P . leucoaUcta. 
I eaves black-granular ________ _____ n_ n _____ _ :.. ____ ____ 20. P. leuc8ndra. 
42 CONTRmUTlONS FROM THE NATIONAL HERBARIUM 
28. leaves 4 to 6 em. 100g ______________________________________________ ...29. 
Leaves 1 to 4 em. long ____________________________ 14. P. hnlvpbiloldea. 
29. Spikes opposite the leaves __________________________ 25. P. portobellenal •• 
Spikes axUlary _____________________________________________ 26. P. elata. 
30. Leaves appreciably obtuse at base or cordate ___________________________ 31 . 
. Leaves appreciably acute at base _____________________________________ 36. 
31. Plants glabrous; leaves round-ovate ____________ ______ __ ___ l. P. pellucida.. 
Plants locally pubescent, nsually; Ieavea elougate ______________________ 32. 
82. Leaves 8 to 14 cm. long, velvety _____________________ 40. P. clUoUbractea. 
Ip8ves 3 to 7 cm.. 100g _______________________________________________ 33. 
33. Plants densely velvety _______ 7 __ _ ___ __________________ 16. P. atevenai. 
Plauts distinctly sllky-pubt>scenL _________ ____ ______ . __ 24. P. oblcurlfolla. 
Pubescence transient _________________________________________________ 34. 
34. Leaves drying thick i peduncle elongate ________________ 39. P. piperol''Ilm. 
Leaves drying thin __________________________________________________ .35. 
85. Peduncle elongate __________________________________ 41. P. cauduUlimba.. 
Peduncle very 8hort_h ____________ ~ ___________________ 43. P ... a tl1Den sla. 
36. Leaves at most elUate or obscurely puberulenL ________________________ 37. 
Leaves evidentlY pubescent _______________ _____ _______________________ 40. 
37. Leaves 12 to 20 em. long or more ______________________ 35. P. llavlspica. 
Leaves 6 to 10 or 12 cm. 10ng _________________________________________ 38. 
Leaves 4 to 6 cm. 10ng _______________________________________________ 39. 
Leaves scarcely 4 by 4 cm _________________________ 33. P. machaerodonta. 
88. Leaves sharply acumlnate _________________ ____ ___ .36. P. a.eumblattfoUa. 
Leaves blonL_nn __ _ n ________ n_nn_n ___ 34. P. dodecatheontoph:r1la. 
39. laves drying thick, subsessiJe ______________ ___________ 39. P. piperoMJm. 
Leaves drying thin, slender-peUoled ______ ____________ 41. P. caudulillmba. 
40. Petiole elongate ___________________________ ________ -41. P. caudulllimba. 
Petiole very short __ __________________ ___ ___________ 42. P. cyllndribacca. 
41. Leaves lanceolate or lance-elliptic _______ __ __________________________ 42. 
Leaves round-ooovate, whorled, smaIL __ ___ ____ ______________________ Ari 
Leaves round, opposite, smaIL ___ __________ ___________ 5. P. cyclophylla. 
Leaves oblong, whorled, mlnute _____ nnn _ ________________ 12. P. apoda. 
42. Leaves 6 to 10 em. ]ong ___________ __ ________________ lO. P. seem8Dn t ana. 
Leaves 4 to 6 em. long ________________________ ____ _____ 9. P. vIrldlsplca. 
LP.aves scarcely 4 em. long _________________________________________ A3. 
43. Stem stout __________________________ ___________________ 8. P. sarcoc:arpa. 
Stern slender ________________________________________________________ 44. 
44. Leaves an opposlte ______________________________________ l1. P. Irlabrlor. 
Leaves alternate on stem _____________________________ 23. P. allal'otacta. 
45. Plant glabrous _____________________________________ 6. P. aubquadrifolia. 
Plant pube8cenL ____ ____ _______________________________ 7. P. quaternata. 
1. Peperomla pellu.lda (L.) H. B. K. Nov. Gen. '" Sp. 1: 64. 181~. 
Piper pellucUlum L. Sp. PI. 30. 1753. 
PANAMA: Without loeaUty, Seemonn 113; Have, 195. Las Sabaoal. 8taf14.. 
le~ 25950. Tabaga Island, Bt"ndl"ll 27063, 27881. Chepo, pjU;er 4450. 
COWl'!: Chagre.. ~'en<ller 302. Porto Bello, Plttler 2469. Puerto Obaldla, 
p"t(er 4370. BocA-S Dn TOBO: Changulnola River, Stork 1. CANAL ZoNE: 
Summit, Standley 26906; Steven-If 32ri. Balboa, 8tandletl 2r5827, 27169; Mr,. 
Fader in 1924. AncOn Hill, Pittier 3962; BteveM 794. 
A common weed tbrough tbe West Indies (the type apparently from Mar­
tinique), Mexico, Central America, and northern South Ameriea, and Introduced 
Into tropical AfrIca. Tbe cooUneotal plant u8ually hos the leaves whlteoed 
TRI!J.]!ASI PIPEBACEAB 01' PANAMA 43 
beneath and often smener, var. PfgmaeG Kunth (STD. Pl. Aequln. 1: 117. 1m, 
the ts"" from Lou) and lIB f. minor Van Heorc\' 6; Mnell. (Obe. Bot. 1111. 
1870, the type trom Panama). 
2. Peperomla brevlpedonc:ula (C. DC.) Trel. 
Pep .. <>mI. /leI/del "I 1Ir •• Ipe ....... '" C. DC. Ann. Cons. Jard. Bot. Geuhe Ill: 
274. 1920. 
A. rather small, simple, erect. glabrous herb, perennial from a rhizome; 
800m 1IC8rre1) 10 em. high. with ehort internodes; leaves few (acarcely 10), 
alternate, round or roond-ovate or elliptic-ovate. rounded or barely anbaeute 
at apex. cordulate, moderate (4 to 7 em. long and wide), drying dark ,f'1SaD, 
dull and thin. 5 or 7 nerved; ""tiole 2 to--5-"m. long; spIkM Bevera!, clustered 
at the- end, comparatively stout and short (scarcely 2-3X80-1GO mm.), rather 
closely fiowered above; peduncle almost suppressed; bracts round·peltate; 
berries blackening, globo .. ; atigma apical ... aslJe. 
TTPS LOCALITY: Santa Marta, Colombia. 
DnrtN: Boca de Pauarand6, Samba River. PUtier MOO. 
RANGE: Eustern Panama and adjacent Colombia. 
3. Peperomia kUlipi Trel. Bot. Gas. 73: 143. 19~. 
CoWN: Alhajoela, Kl/Up 3218. Albajnela to El VlgI •• PUtier 4712. C .. "J.L 
ZoNE: ruo Parafso, East Para (so, Sta.ndlev 29890. 
4. Peperomia hymenodel Trel., gp. nov. 
A. small, nearly glabrou8 herb; stem short and subrbizomatous, with ver1 
abort 1Dternodes; leaves alternate, almost radtcal, elUptlc-ovate, somewbat 
acute, eordulate to subacute at base, moderate or rather small (2X4-6 or 8.5X6 
em.), eUlate and more or leSI minutely pubescent near tbe base, drying very thIn 
find membranaceous. 7-nerved; petiole very slender, 2 to 6 em. long; spikes 
seapose, slender and elongate (1 X 80 mm.), rather closely flowered, the :flowers 
8eparated by anastomosing papillate ridges j peduncle 3 to 5 em. long; braeta 
round-peltate; berries globose; 8tigma subapical. 
Type in the U. S. National Berbarlum, no. 715477, collected Dear San Felix, 
Chlriquf, Panama, by H. Plttler (no. 5266) . 
. . 
RA.NOE : Southwestern Panama. 
5. PeperomJa cyclophylla )liquelln Mart. FI. Bras. 41 ; 219. 1852. 
CA.NAL ZoNE: Gorgona, 88etiicmn. 607, the tl;e; WagMt". Gatunclllo, ~fJee ' 
5652. East Parntso, 814ndlell 29987. CBIBIQut; Puerto Remedios, PUtter 3381. 
Chorrera, [fillip 3408. PANAMA: Rfo Tecumen, 8101ldlell 26M2. Mat(as Ber­
nllnd .. to Juan Dlaz. Standl<v 31938. Rio Tapia. S,andlev 26195, 80670. 
6. Peperomia subquadrifolia Trel. lip. nov. 
Peperomia quadrifolit.J Auet. as to Panama. 
A small. essentially glabrous, arborieo]ous mountain herb j stem slender, 
rooting trom many nodes, repeatedly torking; leaves 3 to 7, commonly 4, at a 
node. obovate or obJong-obovate, revolutely emarglnate and sUghUy hairy In the 
notch, subacute at base. minute or small (5X 8-1 X14 mm .. ), drying opaque and 
• 
otten pe.le, scarcely nerved.; petiole very sbort (1-2 mm.); spIkes terDiinal, 
small (2G mm. long), rather closely flowered; peduncle slender, 10 to 16 mm. 
Ion,: bracts round-peltate, Blender, 10 to 15 mm. long; berries ovoId, with 
slender abrupt style and viscid peeudocupule j stigma apical. 
T,pe in the U. S. National Herbarium, no. 611591, collected at Los Slguas 
Camp, Cerro de la Horqueta, Cblrlquf, Panama, by H. Plttler (no. 3197). 
CBwQu1: Without locality, WClgtt.er. Cerro de la Horqueta, llOaJO'l ts2tt6, 
l5M9. EI Boquete, KIIUp 85S4; Jr....- 5013. 
• 
44 CONTRIBUTIONS FBOM THE NATIONAL HEBBARIUM 
7. l'ep"romia quat.mata }{Iquelln Seem. Bot. Voy. Herald 197. 18M. 
OUIlIIQut: Wag ... ,· 302, in part. Volcano ot ChiriquI, S .... ,."" 896. 
RANoa: Panama and adjacent South America. 
8. Poperomia sa.reocarpa Trel.. sp. nov. 
A moderately large, glabrous, 8SSUrgeot berb, drying greenish. on rocks: stem 
moderately atout (3 to 6 or 8 mm.). rooting from the lower or even the upper 
nodes; leaves opposite, rhombic-obovate or rbombtl!-Oblanceolat~ conspicuously 
blunt-.aeumJnate, 8ubcunenle, moderately small (U~X2.5-2XH or 6 em.), dry­
log coriaceoos, tndistinetly 3 or 5-nerved i petiole scarcely over 2 mm. or on the 
lower leaves 10 to 20 mm, long j spikes terminal, slender Bod moderately elongate 
(2X 70 mm,). rather loosely ft.owered; peduncle as thick as 'the rachis, 2 em. 
long; braeta elltpUc-peltate; berries globose. red, and suceulent; st:gma apical. 
Type 10 the herbarium of the New York BotanIcal Garden, collected near 
Penonom~, Province of Cocl~, Panama, by R. S. Williams (no. 882). 
R.a.l'OE: South-central Panama. 
9. Peperomia vlridispica Trel., ap. nov. 
A rather large, glabrous, pendent, succulent, epiphytic herb; stem rather 
thick (3-6 mm.) ; leaves opposlte or exceptionally 3 at a node, 8ubrhomblcally 
leceolate or oblanceolate, blunt·acuminate, 6ubcuneate, rather lar«e (1.t.iX3.5, 
2X4.6-3 X 6.6 em.), drying rather thick and opaque, Ind 'sUnetly about fi-nerved i 
petiole 5 to 10 mm. long. winged; spikes termilUll and occasIonally axHlary, 
Dliform (scarcely 2 X'lo-lOO mm.), loosely flowered. green; peduncle 1 to 2.5 
em. 10Dg; bracts BubelUptic·peltate; ovary ovoId, attenuate upward; stigma 
apical . 
Type In the U. S. National Herbarium, no. l,lM,147. collected at Balboa. 
Canal Zone. Panama, by Paul C. Standley (no. 28550). 
CANAL ZollB: Balboa, Standley 27117, 2&i68. PANAWA: Tumba Muerta, 
8ta"aleJ/ 29827. Las Sabanas to MaUas HernAndez, Stantlle,! 81893. 
RANOE: Central Panama. 
9a. Peperomla virldispica perejil Trel., var. nov. 
Erect, 60 em. taU; leaves more commonly 3, excepttonally 4, at a node, 
blunter-po:nted. 
Type tn tbe U. S. National Herbarium. no. 1,153,969, collected on Taboga 
Island, Panama, by Paul C. Standley (no. 21893) . 
PAl'U,I(A: Taboga Island, Macbride 2808. 
HANOIll: Southern Panama. 
10. Peperomla uflmannlana Miquelln Seem. Bot. Voy. Herald' 198. pl. j'f.1S54. 
CHIJUQUl : Volcano of Chiriquf, Seeman.,. 896. Cerro de la Horqneta. PUUer 
S26V. EI Boquete, Pllli .. · 2921 ; Ma"'" 0087 ; Killip 3542. 
11. Peperomla glabtlor (C. DC.) Trel., sp. nov. 
Peper Offila palmatta glabrior C. DC. Cundollea 1: 304. 1923. 
An erect herb of the aspect ot P. palma-na; lenves somewhat larger (1.~2X3-
5 cm.), glabrous; spikes larger (2X40 mm.). 
Type 1n the U. S. National Herbarium, no. 675770, collected on Cerro de la 
Horqueta, Ch 'riqut, Panama, by William R. Maxon (no. 5401). 
CHIBlQuf: Cerro de la Horqueta, Maxon 5541, 15553; Pittier 3217. EI Boquete, 
Killip 851V. 
RANGE: Southwestern Panama. 
12. Peperomia apoda Trel., sp. nov. 
A. small stoloniterous mountain herb; stem slender (1- 2 mm.). rooting from 
the lower nodes, minutely but densely puberulous; leaves 2 to 4 at a node, 
TRBT.EASE PIPERACEAE OB PAN .'u 
oblong. rounded at both ends, sometimes emarglnnlate, mlDute (2XIO mm.), 
somewhat pale-&lanular beneath, glabrous, I-nerved. i petiole .Imost IUppn lied; 
8p1kes termloal and from the upper axlls. ftllform (IX20-36 mm.), rather 
loosely flowered: pedoDCle nearly suppressed (1-2 mm.) ; bracts rouod-peltate; 
ovary ovoid; stlcma subapical. 
Type 10 the U. S. National Herbarium, no. 1,266.031, collected. In valley of 
R!o Caldera, Chlrlqu!, Panama, by E. P. KllIlp (no. 8lI14). 
B.A.}lf02: Soutbwestem Panama. 
13. Peperomia rotundlfolla (L.) H. B. K. Nov. Gen. '" Sp. 1: eG. 1816. 
Piper rotut04ltollum L. Sp. Pl. 30. 1753. 
00L0 .. : RIo FaW, Pitlior 3888. CANAL ZoNE: Frijoles, PIffl ... 871)8. 
A wld ... pread speeles, the prototype from Martinique, noel", trom the 
Antilles to and well lDto South America in puzzUng forms with leaves either 
orbleular or obovate. 
14. Peperomia h:verophllold •• C. DC. Candolle. 1: 329, 892. 1923. 
ClliBIQuf: Cerro de la Horqueta, Pittier 320~, .. 3203." Rio Calde:ra, K nlip 
3564. 
15. Peperomia pauamenol. C. DC. C.ndoll.. 1: 329, 401. 1923. 
CANAL Zon: R!o Indio de GatQn, Piltler 2789. 
16. Peperomf. stevenet Trel .. lIP. nov. 
A small, densely velvety, 8ssurgent herb in tree tops: stem IIbort, with abort 
internodes; leayes altuoate, clustered at end of the stem, elUpUc or elUptle­
oyate," subacute to acuminate, cordulate, rather small (2.6-SX~ em.). paler 
beneath, drying creen, submultiple-nerved from below the upp ar thlrd, the 
branches ot the midrib about 5 X 2; petiole 1.5 to 3 em. 101lJ; spikes terminal, 
moderately large (2X60-70 mm.). closely flowered; peduncle 10 to 15 mm. 
long; bracts round·peltate. drying thin and light yellow; berties &lobOse, 
obUquely subtruncate; stigma anterior. 
Type In the herbarium. of the University of llUnols, collected at Frijoles, 
Canal Zone, Panama, by F. L. Stevens (no. 1247). 
RANGK: Central Panama. 
17. Peperomia parietariaetolia Trel .• sp. nov. 
A rather large, forking, glabrous berb; stem slender (1 mm.), or stout 
(5 mm.) below; leaves alternate. lanceolate, acuminate. typleally acute at 
baae. moderate (1.~2.5X4-6 em.), conspicuously 5-nerved. broWD"lraDular 
"beneath; petiole about 6 mm. long; spikes a:dU.ry. u nt Immature and 'fery 
small. with short peduncle; bracts roond-pe1tate. 
Type In the U. S. National Herbarium, DO. 1,266,2118 082, collected 10 Talley ot 
the Rfo caldera, between El Boqoete and "tbe cordUlera, ChJrlqut, Panama, by 
E. P. Klllip (no. 8lI20). 
R.A.NOE: Southwestern Panama. 
18. Peperomia eonjuDgeDs Trel.. sp. nov. 
A moderate·sized, rppent-assurgent. merely emate. black 01' brown-punctulate 
herb; stem slender (1-2 mm.). ciliate-lined below the petiole margins, rooting 
from the nodes: leaves nlternate, lanceolate to subrhomblc-ell1pttc. acute or 
somewhat blunt-ncumlnate, rather acute at base, moderate (US XS. 2X4-e. or 
2.5X6 cm.), ~ or 7-nerved. rather firm; petiole ~ to 10 or Hi mm. long. coarsely 
c1l1ate; spikes terminal or also from the uppermost axil, rather small (as yet 
scarcely 2X5Q mm.), becoming loosely flowered; peduncle IS mm. long; brncts 
round-peltate; ovary ovoid; stigma oblique. 
46 CONTRmUTIONS FROM THE NATIONAL HERBARIUM 
Type In the U. S. National Herbarium, no. 1,215,9l!O, collected on Barro Colo­
rado Island in Gat'On Lake, Canal Zone, Panama, by Paul C. Standley (no. 
31342). 
CANAL ZoNE: Las Csscadas Plantation, standJell 25780, 29652. Frijoles, 
Siandleu 27411, In part. 
RANGI:: Central Panama. 
19. Peperomta leucosticta TreJ., sp. nov. 
A rather small, Resurgent, glabrous herb; stem slender (2 mm.), rooting 
from the lower nodes; leaves alternate, broadly rhombIc, rhomblc·lanceolate or 
oblanceolate, or obovate, blunt-acuminate, acute-bMed, rather small (US-2X2.6-
3.5 em.), somewhat paler-granular beneath, ti-oerved ; petiole 10 mm, long, decur­
rent; spikes terminal and axillary, rather slender and elongate (scarcely 
2 X 60 mm,), remotely flowered, the flowers In fusiform pits separated. by 
broad fiat anastomosing ridges; peduncle flUform, 10 to 15 mm, long; bracts 
small, round-peltate; ovary ovoid-pointed; stigma anterior. 
TyPE in the herbarium. of the New York Botanical Garden, collected in 
Panama by Sutton Hayes (no. 8ll). 
20. Peperomia leucandra Trel., sp. nov. 
Peperomia glabella Auct. in part as to Panama. 
A rather small, repent or 8ssurgent, black-punctulate herb; stem slender 
(about 1 mm.), occasionally slightly ciliate below the petiole margins, rooting 
from the nodes; leaves alternate, elliptic or lance elUptlc, more or less acumi­
nate, acute-based, ratber small (1.5-2 X 2.5--4.5 em.), somewhat ciliate above, 
IS-nerved; petiole 1llitorm, 5 mm. long, sometimes ciliate; spikes terminal and 
axillary, rather slender and elongate (1.5 X 00 mm.), moderately closely :flow­
ered; peduncle 5 mm. long; bracts roundlsh-peltate; anthers otten drying white­
(as In some other species) in contrast with the dark bracts; ovary ovoid~ 
acuminate; stigma oblique. 
Type in the U. S. National Herbarium, no. 715329, collected at San Felix. 
ChiriquI, Panama. by H. Pittier (no. 5148). 
RANGE: Southwestern Panama. 
21. Peperomia fusco-ciliata Trel .• sp. nov. 
A delicate black-punctulate repent herb; stem slender (1-2 mm.), rusty­
cil1ate below the petiole margins, rooting trom the nodes; leaves alternate. 
ovate, acute, rounded or subacute at base, minute as represented (scarcely 
7Xll mm.), 3-nerved, ciliate upward; petiole 5 mm. long, rusty-cUlate; spikes 
terminal and axillary, loosely :flowered. small (when partly in :flower lX7 
mm.); peduncle scarcely 5 mm. long; bracts round-peltate, black; ovaries 
ovoid, sunken in the rachis; stigma anterior. 
Type In the herbarium of the New York Botanical Garden, collected in 
Panama, 1859-60. by Sutton Hayes (no. t519). 
RANGE: Panama. 
22. Peperomia percuneata Trel.. sp. nov. 
A rather small. prostrate or stoloniferous. black-punctulate herb; stem 
slender (1-2 mm.), sometimes slightly cUiate below the petiole margins, root­
ing from the nodes; leaves alternate, rhombic-lanceolate or oblanceolate. 
acute or somewhat blunt-acuminate, cuneate, moderate or rather small (IX&-
1.5 X t5 em.), 3 or 5-nerved; petiole filiform, about 5 mm. long, occasIonally 
eiliate j spikes terminal, filiform (1XOO mm.), at length rather loosely flowered; 
peduncle 5 mm. long; bracts round-peltate; ovary ovoid; stigma oblique. 
Type In the U. S. National Herbarium, no. 715426, collected along the Rfo 
Dupf, Chlriquf, Panama, by H. Pittler (no. 5224). 
RANGE: Southwestern Panama. 
TRBI.BA8B PIPBRACEAB 01' PANAMA 47 
lIS. Pep.romla aUeg.tacta C. DC. Clwdolles 1: 817. 378. 1928. 
CR ...... qul: EI Pot .... ro. Chlrlqu1 Volcano, pjllier 3108. celro 4e 1& Hor. 
queta, Killip 8681. 
1M. Peperomla obocurlfolla C. DC. Condolles 1: M7. tOO. 1928. 
C.UUL ZoI'fS: Agua Clara, Trinidad RiTer, Pittier 3982. Barro Colorado 
Island. Statull,'I 81333. La. C .. e.da. Plantation. Statuller 26726. 
RANGE: Panama. 
:iJ4L Poperomla obocurltolla tatoana (C. DC.) Trel. 
Peperomla tatooll4 C. DC. Candollea 1: M7. 387. 1928. 
A small, softly white-balry, epiphytic herb; stem rather slender (2 mm.). 
rootlnC from the nodes; leaves alternate. but the uppermost congested, lance­
elliptic. acute at both end •• rather small (IX8 to scarcely 1.5X3.5 em.). 5-
nerved with the inDer laterals trom above the base, c1liolate upward, yellow· 
iDg and sparsely hairy on the nerves beneath j petiole very short (3 mm.); 
aplkea axillary and temdnal. dutorm and elonpte (lX1OO rom.); peduncle 
IIlltorm. e1oneote (2-3 cm.). hairy; bract. round_vote. peltote; berries 
globose, obUquely mucronulate; stigma subaplcat. 
CoLON: ruo FatO. Pilller 3002. CANAL ZoN.,: Acuo Clara. Pilller 3983. 
RANGI:: Central Panama. 
Mb. hperomla obseurifoUa minor Trel., var. DOV. 
A reduced form ot the Canal Zone. 
Type fn the herbarium. of the New York Botanical Garden, collected between 
lrIatachfn and Las Caseadas, Canal Zone, Panama. by J. F. Cowell (no. 336). 
CANAL ZoNII: Trlnfdad River, Plttier 3983. 
2:S. pep'romia portobellensts Beurl. Svensk. Vet. Akad. Bandt. 18ft': 147. l.856. 
CoLOI<: Porto Bello. Billberg. 
28. Peperomia elata C. DC. Caodollea 1: 35:S, 886. 1928. 
CHIBIQID: El Boquete, Jl ~<m M82. 
27. Peperomlo pueu1cola C. DC. Candollea 1: M2. 402. 1923. 
CRIBIqUi: El Boquete, PUller 291~a; KUllp 3547. 
28. Peperom1a Bendena Rulz. Pav6n. Fl. Peruv. 1: 32. pl. 51, b. 1198. 
CANAL ZolU:: Rfo Indio de GatOn, Jlazon 4867. 
Wide-spread from Peru, the type locality, to Nicaragua on the eontinent, and 
throuah the Weat Indies. A probably divisible spedes, ot many B1nonymB, 
29. Peperomla uroearpoldes C. DC. C.ndoll .. 1: 362. 414. 1928. 
DAutN: Cerro de Garagaril. SambO BaslD, PUller M70. 
30. Peperoml. cordulatltormlo ·Irel. BoL Gu. 78: 148. t. J9. 1922. 
CANAL ZoIU:: Mamel HIlI, ~tt~r 3806, 846T. 
81. Peperomia cordulata C. DC. Journ. Bot. Brit. 4:: For. 4: 187. 1866. 
Coww: Chagres, Fefldler 2M. 
32. Peperomla variegata Rul. & PavOll. Fl. Peru •. 1: 33. pl. 62. 1. a. 171l8. 
CHIMIqUi: EI Boquete. Pilller 298f. 
A Peruvian species; rarely foUDd In Central A Ii erica. 
33. PeperomJa maohaerodonta Trel., ap. nov. 
A.. rather small, Buberect, arborlcolous, glabrate herb: stem. moderate (8 
mm.) j leaves alternate, round~lUpttc. obtuse or abruptly Bhort-acuminate, 
rounded or acute at bane. rather Bmall (2.GX3.5-3X4 em.). drylng eor1aceous 
but greenlab. multiple-nei fed, the baBsl oen" and branchea of the midrib 
2X2 each. appressed-puberulent beneath, especially on the Denea; petiole 
48 CONTRmUTlONS FROM THE NATIONAL HERBARIUM 
about 5 mm, long. winged; spikes terminal, rather thick and elongate (2X70 
mm,) ; peduncle 2.5 em. loug; bracts round-peltate, about 0 In a whorl; berries 
elUpsold, with an equllonr. slightly curved. strong beak. at first In ellipsoid pits 
separated by narrow anastomosing ridges; stigma nnterIor, at base of the 
beak, 
Type In the U. S. National Herbarium. no. 1.266.003. collected In the valley 
ot Rio Piarn8stn, east at E1 Boquete, Chiriqui, Panama by E. P. Klllip 
(no. 3528). 
RANGE: Southwestern Panama. 
84. Peperomla dodecatheontoph:rlla Trel., sp. nov. 
A rather small, glabrous, epiphytIe berb; stem moderate (3 mm.), rooting 
trom the lower nodes; leaves alternate, elUptic-suboboyute, 8ubacute. 
acute at base. rather large (3.5X7.5 to usually 5.tHlXll-12 cm.). lIeshy. 
drying dull. cbnrtaceou8. and wrinkled beneath, obscurely pinnately veined to 
tbe middle. the branches of tbe midrib about 4X2; petiole 2.4 to 8.5 em. long. 
wInged above the middle; spikes axillary or terminal, 1 or 2 on a rather per~ 
slstently 1 or 2 bracted stalk, rather slender and elongate (2-3 X 70--100 mm.) ; 
peduncle rather long (4-5 em.); bracts round~peltatei berries oblong~tU8Ltorm, 
the conical beak flUform-atteuuate. 
Type In tbe U. S. National Herbarium. no. 677275. collected along RIo 
Indio de Gat110, Canal Zone, Panama, by H. PUtler (no. 28(0). 
COL6N: RIo Indio de ~'at6. Pittlet' 4263. 
RANOE: EaBt-central PRnama. 
35. Peperomia 1la vispiea Trel., sp. nov. 
A rather large, glabrous, arboricolous herb; stem stout (5-10 mm.), rooting 
trom the nodes, with very short, erect, leafy branches; leaves alternate, ell1p­
tlc-8ubobovate, 8UbaCUte, acuminately attenuate at baBe, large (6Xl2-10X20 
cm. or more), deshy, drying rather glossy, coriaeeons, and wl"lnkled beneath. 
obscurely pinnate-veined to beyond the middle, the delicate branches of the 
midrib about 6X2; petiole 5 to 12 em. long, winged upward; spikes Axillary 
(1) or terminal, slender and long (scarcely 3X100 mm. or more), yellow; 
peduncle rather short (2-.3 cm.), red; bracts round-peUate, small, about a 
dozen to the pseudo-whorl; ovary oblong, slender~beaked; stigma anterior at 
ba .. of the beak. 
Type in the U. S. National Herbarium no. 1,153,501, collected along the 
Rio Tapia, Province ot Panama, Panama, by Paul C. Standley (no. 26196). 
CAN AI. ZoNE: Las Cuscadas Plantation, 8tantllev 25729. 
RANGE: Central Panama. 
36. Peperomia acuminatifolia Tre1., sp. nov. 
An assurgent (1), essentially· glabrous herb; stem succulent, rather stout 
(3-4 mm.); leaves alternate, oblanceolat4Hlbovate, sharply acuminate, sub­
cuneate from about the middle, rather large (about 4X9 em.), drying rather 
coriaceous, very indistinctly pinnate-veined below about the middle, the dell­
cftte branches of the midrib about 5X2. sparingly ciUolate; petJole 1 to 2 cm. 
long, winged; spikes termlnsl, moderately tbJck BDd elongate (3X75 mm.). 
subacute: peduncle 2 cm. iong: bracts round-paitate. 
Type In the herbarium of Columbia eolle&,e, New York Botanical Garden. 
collected In Panama by Sutton Hayes (no. 817). 
RANGE: Panama. 
87. Peperomla williamlU Trel., sp. nov. 
A. glabrous, 8ubslmple, rather large, succulent herb growing on trees i stems 
stout (nearly 1 em.); leaves alternate, obovate-elltptlc, rounded or slightly 
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OF PANOlA • 49 
emorctnare at apex, ""JDe"bat abruptly contracted to the broadly wlnpd 
petiole, large (l~l1X15 CID. or more), impressed-ponctulate. very obscurely 
plnnat ... elned, slightly ",.olure; petiole long (11-12 em.); opll< .. mostly 
palred. at the upper nodes ot a stout zigzag l-Ieaved prolongation of the stem, 
mooerately thlclt and long (:HI X 100 mm.), obtuse; peduncle IOllJ (3-4.6 em.) ; 
bracts round-peltate; berries honey-colored, conlc-ovoid, tapering Into • nearly 
equllong, stout. conic beak, the sUrma anterior on the lower part ot the beak. 
'iype in the herbarium of the New York Botanical Garden, collected at Bia-. 
maret, Province ot Coel~, Panama, by R. S. W11l1ams (no. 483) . 
RANOE: Sonthern Panama. 
38. Feperomla oblongibacca C. DC. Candollea 1: 288, 400. 1928. 
COWN: Rio FaW, PiU/er 3891. 
89. Peperomta piperorum Trel., &p. nov. 
A. moderately large, succulent, glabrous, pendent. arboricolou8 berb; stem 
moderate (3 mm.), more or Jess scurty-e:doliatln~; leaves alternate, sub· 
BeaUe, lanceolate. sharply long·acumlnate, subacute at base. moderate 
(1.5X4 cm. ) or becomiDg rather large (fully 2X6 em.), drying opaque aDd 
thick and somewhat yellow18h and _ granular beneath, pinnately veiDed. the 
branches ot the midrib about 3X2; petiole thick, scarcely 2 mm. long; spikes 
terminal on the short narrow·bracted apex, moderately slender nod short 
(2X40-60 mm.), closely flowered; peduncle slender. 1 cm. long; bracts large, 
round·peltate; berry reddish, oblong, short nnd thick, acumlnately oblique­
truncate; stigma central on the scutulum. 
Type in the U. S. National Herbarium, no. 1,100,334, colleded aloog Rio Agua 
Salud, near Frijoles, Canal Zone, Panama, by C. V. Piper (no. OOM). 
RANOE: Central Panama. 
40. Foporomia elllolibracrea C. DC. Candollea 1: 360, 383. 1923. 
PANA1U.: Charara River Dear Chepo, PUtter 4712. CANAL ZoNE: Las Cns­
coda. Plantation, 8Ia"dle~ 25781. 
41. Peperomia eaudu1111mba C. DC. Ann. Coos. Jard. Bot. Gen~ve 21: 200. 
1920. 
C . , ... 1. ZoN.: Rio Indio de Gatdn, Plllier 2738. 
41a. Peperomia eaudulillmba marragantina C. DC. Ann. Cons. Jard. Bot. 
Oen~.e IU: 269. 1920. 
PANAlIA : MarraganU, WtU,ama 69 .. 
41b. Peperomta caudnnhuba cryptopoda TreL, var. nov. 
PeperomhJ glabrlromea- ~uct. 88 to PaDama. 
A moderate·slzed repent herb, rooting from the nodes: stem moderate 
(3-4 mm.), glabrous, or pubescent below the petiole margins; leaves alternate, 
lanceolate, gradually acuminate, somewhat rounded at base, rather large 
(2.5-4.G X8-U em.), pinnately veined from below about the middle, the 
delicate branches of the midrib 3 or 4X2, glabrous; petiole very short (scarcely 
IS mm.), sometimes clUate; spikes solitary or paired on sparsely hairy, bracted 
stalks less than :; mm. long, filiform and elongate (1 X 110 mm.); peduncle 
very short (5 mm.) ; bracts round·peltate; berries obconlcal·obloo,. truncately 
scutulate wJth short hOrizontal beak; stigma anterior. 
Type in the U. S. Notional Herbarium, no. 7Ui837, collected at Boca de 
Pauarand6, Samb11 River, DarMn, PaD8ma, by B. Pittter (no. lS071). 
CANAL Zoln:: RIo Indio de Gatlln, Muon 4864. 
R.6.NGE: Panama . 
, 
50 CONTRIBUTIONS PlIOM TRJ! NATIONAL HERBARIUM 
410. Peperomia e&udullllmba IODlI"pendula O. 00. Ann. 0008. Jard. Bot. 
Genhe Ill: 269. 1920. 
CUAL Zon: Fort Sherman, 81_1ev 81118. Barro Colorado Island, 810.dl"" 
31321. 
411. Peperomia eyllndrlbaeea C. 00. Cendollea 1: 370, 881). 1928. 
CnwQuf: San Felix, Piltier 5273. 
43. Peperomia tratunonm 0. DC. Cendollea 1: 863, 889. lD2S. 
PANA.1lA: 8eem4"" 608. CANAL Zorn!:: Gorgona to Gat11n, PUt ... 2281). Rfo 
Indio de Gatdn, M/WI" 4318. Barro Oolorado loland, 81andleu 318li6. CocLt: 
Pe.nonoml!. wtUl4ftW1 866. 
44. Peperomia mamaan. C. DC. Cnndolleo. 1: 290, 396. 1923. 
c...l'fAL Zol{J:: ?damet Hill, Pittier 3806 . 
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