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Botany 


FLORA  COSTARICENSIS 


William 


Family  #193  Scrophulariaceae 
Family  #193a  Schlegeliaceae 
Family  #194  Bignoniaceae 
Family  #195  Pedaliaceae 

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Family  #196  Martyniaceae 
Family  #197  Orobanchaceae 


PUBLI 


FIELDIANA 


Botany 

NEW  SERIES,  NO.  41 


FLORA  COSTARICENSIS 


William  Burger,  Editor 

Curator  Emeritus,  Department  of  Botany 
Field  Museum  of  Natural  History 
1400  South  Lake  Shore  Drive 
Chicago,  Illinois  60605-2496,  U.S.A. 


Family  #193  Scrophulariaceae     Family  #195  Pedaliaceae 

Kerry  Barringer  and  William  Burger  William  Burger 


Family  #193a  Schlegeliaceae 

William  Burger  and  Kerry  Barringer 

Family  #194  Bignoniaceae 

William  Burger  and  Alwyn  Gentryt 


Accepted  October  29,  1998 
Published  April  28,  2000 
Publication  1508 


Family  #196  Martyniaceae 

William  Burger 

Family  #197  Orobanchaceae 

Luis  D.  Gomez  and  William  Burger 


PUBLISHED  BY  FIELD  MUSEUM  OF  NATURAL  HISTORY 


Introduction 


This  is  the  eleventh  issue  in  the  Flora  Costar- 
icensis  series.  The  first  dealt  with  the  Piperaceae, 
family  number  41  (Fieldiana,  Bot.  35,  1971).  The 
second  included  families  numbered  42  through 
53,  Chloranthaceae  through  Urticaceae  (Fieldiana, 
Bot.  40,  1977).  The  third  issue  covered  the  Gra- 
mineae  (Poaceae)  and  was  authored  by  Richard 
Pohl  (Fieldiana,  Bot.  n.s.  No.  4,  1980).  The  fourth 
issue  included  families  numbered  54  through  70, 
Podostemaceae  through  Caryophyllaceae  (Fieldi- 
ana, Bot.  n.s.  13,  1983).  The  fifth  issue  covered 
families  200  and  201,  the  Acanthaceae,  authored 
by  L.  H.  Durkee,  and  the  Plantaginaceae  (Fieldi- 
ana, Bot.  n.s.  No.  18,  1986).  The  sixth  issue  in- 
cluded families  80  and  81,  the  Lauraceae  and  the 
Hernandiaceae  (Fieldiana,  Bot.  n.s.  No.  23,  1990). 
The  seventh  issue  included  families  numbered  97 
through  103,  Krameriaceae  through  Zygophylla- 
ceae  (Fieldiana,  Bot.  n.s.  No.  28,  1991).  The 
eighth  issue  included  family  202,  the  Rubiaceae 
(Fieldiana,  Bot.  n.s.  No.  33,  1993).  The  ninth  is- 
sue included  family  1 13,  the  Euphorbiaceae  (Fiel- 
diana, Bot.  n.s.  No.  36,  1995).  The  tenth  issue 
covered  Tribe  Maxillarieae  of  family  39,  the  Or- 
chidaceae  (Fieldiana,  Bot.  n.s.  No.  40,  1999). 

Illustrations  of  leafy  or  flowering  stems  are  all 
drawn  to  the  same  scale  with  the  exception  of 
Figures  1,  2,  and  28.  Enlarged  flowers,  fruits,  or 
seeds  are  drawn  to  the  same  scale  on  an  individual 
plate  unless  otherwise  noted.  Bignoniaceous  fruits 
are  drawn  to  the  same  scale  as  the  leafy  stems. 
The  closed  scales  represent  centimeters  and  the 


open  scales  represent  millimeters.  The  figures  are 
somewhat  diagramatic  and  represent  a  common  or 
characteristic  morphology  for  each  species.  Figure 
28  was  done  by  Luis  D.  G6mez;  the  others  are  by 
William  Burger. 

The  circumscription  of  the  Scrophulariaceae  is 
difficult  because  the  family  is  part  of  a  closely 
related  group  of  families  that  includes  Oroban- 
chaceae,  Bignoniaceae,  Myoporaceae,  Gesneri- 
aceae,  Pedaliaceae,  Martyniaceae,  and  Acantha- 
ceae. A  few  genera  have  been  difficult  to  place. 
Gibsoniothamnus  was  described  by  Williams 
(1970)  as  a  genus  of  Scrophulariaceae,  and  he 
noted  its  similarity  to  Schlegelia.  These  genera 
were  placed  in  the  Bignoniaceae  by  Gentry  but 
are  regarded  as  a  separate  family  here.  Reveal  has 
recently  (1995)  opted  for  a  finer  division  of  the 
Scrophulariales  and  proposed  segregating  families 
from  within  Scrophulariaceae.  We  adhere  to  a 
more  traditional  circumscription  of  the  family  be- 
cause of  its  long  history  in  providing  a  useful  in- 
formation retrieval  system. 

The  treatment  of  the  Bignoniaceae  presented 
here  is  publication  No.  2  in  the  Gentry  Invitation 
Series.  This  series  acknowledges  Dr.  Gentry's 
many  contributions  to  our  knowledge  and  under- 
standing of  the  Bignoniaceae.  The  late  Alwyn 
Gentry's  publications  and  his  many  authoritative 
determinations  have  served  as  the  foundation  for 
the  treatment  of  the  Bignoniaceae  presented  here. 
In  addition,  there  are  virtually  no  departures  from 
his  taxonomic  concepts  in  this  treatment,  and  he 
is  listed  as  co-author  for  that  reason. 


Acknowledgments 


We  thank  the  staff  of  the  Muse"o  Nacional  de 
Costa  Rica  for  their  assistance  in  collecting  pro- 
grams over  many  years.  The  National  Science 
Foundation  and  the  National  Geographic  Society 
helped  support  many  of  these  collecting  activities. 
NSF  grant  DEB-8103184  helped  support  the  work 
of  Kerry  Barringer  while  studying  Costa  Rica's 
Scrophulariaceae.  We  thank  Warren  Douglas  Ste- 
vens for  making  available  copies  of  treatments  of 
Bignoniaceae  and  Scrophulariaceae  prepared  for 


the  Flora  of  Nicaragua.  We  thank  Michael  Grayum 
for  providing  a  copy  of  the  treatment  of  the  Big- 
noniaceae for  the  Manual  Flora  of  Costa  Rica. 
Noel  Holmgren,  William  D'Arcy,  and  two  anony- 
mous reviewers  provided  many  helpful  comments 
and  corrections.  The  collections  of  the  Field  Mu- 
seum, the  Missouri  Botanical  Garden,  Duke  Uni- 
versity, and  the  U.S.  National  Herbarium  were  con- 
sulted in  preparing  this  treatment,  and  we  thank 
those  institutions  for  the  use  of  their  materials. 


FLORA  COSTARICENSIS 
Family  #193  Scrophulariaceae 
Family  #193a  Schlegeliaceae 
Family  #194  Bignoniaceae 
Family  #195  Pedaliaceae 
Family  #196  Martyniaceae 
Family  #197  Orobanchaceae 


SCROPHULARIACEAE 

By  Kerry  Barringer  and  William  Burger 

REFERENCES — W.  D'Arcy,  Scrophulariaceae,  in 
R.  Woodson  et  al.,  Flora  of  Panama.  Ann.  Mis- 
souri Bot.  Card.  66:  173-272.  1979.  F.  Pennell, 
The  Scrophulariaceae  of  eastern  temperate  North 
America.  Acad.  Nat.  Sci.  Philadelphia  Monogr.  1 : 
1-650.  1935.  P.  Standley  &  L.  O.  Williams,  Scro- 
phulariaceae, in  Flora  of  Guatemala.  Fieldiana, 
Bot.  24,  9(4):  319-416.  1973.  D.  Sutton,  A  Re- 
vision of  the  Tribe  Antirrhineae,  British  Museum 
(Natural  History)  &  Oxford  Univ.  Press,  London, 
576  pp.  1989.  D.  Sutton  &  R.  Hampshire,  Scro- 
phulariaceae, in  Flora  of  Nicaragua  (unpublished 
manuscript).  1995.  J.  Thieret,  The  tribes  and  gen- 
era of  Central  American  Scrophulariaceae.  Ceiba 
4:  164-184.  1954.  J.  Thieret,  Supraspecific  clas- 
sification of  Scrophulariaceae:  A  review.  Sida  3: 
87-106.  1967. 

Herbs  or  subshrubs  (rarely  woody  shrubs  or 
trees),  erect  to  decumbent  or  prostrate  (rarely 
climbing),  terrestrial  to  semiaquatic  or  aquatic, 
autotrophic  or  hemiparasitic,  bisexual,  stems 
without  internal  phloem,  glabrous  or  with  simple 
or  branched,  glandular  or  eglandular  uni-  or  mul- 
ticellular  hairs;  stipules  absent.  Leaves  alternate, 
opposite,  or  verticillate,  simple  (rarely  deeply 
pinnately  lobed),  distal  leaves  often  reduced  and 
intergrading  with  the  floral  bracts,  leaf  blades 
serrate  to  entire  or  deeply  pinnately  lobed,  ve- 
nation pinnate  or  less  often  palmate.  Inflores- 
cences racemes,  spikes,  thyrses,  or  panicles  of 
cymes  or  of  solitary  (several)  flowers  in  leaf  ax- 
ils, bracteoles  present  or  absent  on  the  pedicels, 
pedicels  usually  well  developed.  Flowers  bisex- 
ual, small  to  large,  often  showy,  calyx  4-  or  5- 
lobed,  or  deeply  divided  to  the  base  and  sepals 
4  or  5  (2),  imbricate  or  valvate  in  bud,  persisting 


and  often  enlarging  slightly  in  fruit;  corolla  unit- 
ed and  tubular  to  campanulate,  bilaterally  sym- 
metric and  usually  2-lipped  (rarely  radially  sym- 
metric), lobes  4-5  (8),  upper  (adaxial)  lip  2- 
lobed  to  emarginate,  sometimes  galeate,  lower 
(abaxial)  lip  usually  3-lobed,  tube  saccate  or 
spurred  in  some;  stamens  2-4  (5),  alternating 
with  the  lobes,  usually  of  2  unequal  pairs,  fila- 
ments borne  on  the  tube,  free,  anthers  with  2(1) 
equal  or  unequal  thecae,  distinct  or  confluent, 
rarely  awned,  a  staminode  present  or  absent 
(rarely  2  or  3),  a  disc  present  or  absent  around 
the  ovary;  pistil  solitary,  ovary  superior  (or  half 
inferior),  2-locular  (rarely  1-locular  near  the 
apex  or  3-locular),  ovules  usually  many  on  2  ax- 
ile  placentas,  unitegmic,  style  1,  terminal,  stig- 
mas simple  or  2-lobed.  Fruits  usually  dry  cap- 
sules (rarely  baccate),  dehiscence  loculicidal  or 
septicidal  or  both  (rarely  indehiscent),  placenta 
often  persisting;  seeds  usually  many  and  small, 
usually  with  an  ornamented  testa  or  exotesta 
(sometimes  smooth  or  winged),  endosperm  usu- 
ally present,  embryo  small. 

The  family  Scrophulariaceae  includes  285  gen- 
era with  between  4,000  and  5,000  species.  The 
family  is  primarily  north  temperate,  but  with  trop- 
ical montane  species  and  cosmopolitan  weeds. 
One  of  the  largest  genera.  Calceolaria  (300-400 
spp.),  is  almost  exclusively  South  American  and 
montane.  In  this  treatment  we  include  36  genera 
with  72  species,  1 8  of  these  being  introduced  or 
cultivated  (see  listing  at  end  of  family,  p.  69). 
Costa  Rica  lies  between  two  major  areas  of  di- 
versity for  the  family:  one  in  Andean  South 
America  and  one  in  Mexico  and  Guatemala. 
Among  the  southern  elements  are  Alonsoa,  Cal- 
ceolaria, Scoparia,  and  Siemodia,  while  northern 
elements  are  Castilleja,  Hemichaena,  Lamouroux- 
ia,  and  Russelia. 

Most  Scrophulariaceae  are  recognized  by  their 
sympetalous  tubular  (usually  two-lipped)  corollas, 


FIELDIANA:  BOTANY,  N.S.,  NO.  41,  APRIL  28,  2000,  PP.  1-174 


two-locular  ovaries  with  many  ovules  on  axile 
placentas,  capsular  fruits  with  many  small  seeds, 
and  stems  lacking  internal  phloem.  Most  species 
are  herbs  or  small  subshrubs,  and  the  corollas  are 
often  showy.  There  is  great  diversity  in  the  form 
of  corolla  and  pollination  vectors  (see  C.  M. 
Kampny,  Pollination  and  flower  diversity  in  Scro- 
phulariaceae,  Bot.  Rev.  61:  35-366,  1995).  Iridoid 
compounds  are  common  in  the  family.  The  family 
includes  many  species  cultivated  as  ornamentals 
and  an  important  drug  plant  (Digitalis)  but  no  im- 
portant food,  spice,  or  fiber  plants.  Interestingly, 


a  few  of  the  most  colorful  genera  (e.g.,  Castilleja 
and  Lamourouxia)  have  not  been  utilized  in  or- 
namental horticulture  because  of  difficulty  in 
propagating  their  hemiparasitic  species.  Species 
likely  to  be  mistakenly  identified  as  Scrophulari- 
aceae  are  found  in  Lamiaceae,  Acanthaceae,  and 
Gesneriaceae,  as  well  as  the  other  families  includ- 
ed in  this  volume.  Current  molecular  studies  in- 
dicate that  the  Scrophulariaceae,  as  traditionally 
defined,  are  polyphyletic  (Wagstaff  &  Olmstead, 
1997).  We  retain  the  traditional  circumscriptions 
of  families  in  this  series  to  facilitate  information 
retrieval. 


Key  to  Genera  and  Unusual  Species  of  Scrophulariaceae 

la.  Plants  fully  aquatic,  only  the  inflorescences  held  above  the  water  level;  leaves  submerged  or 

floating,  whorled,  with  pinnatisect  filiform  divisions  [rarely  collected] Benjaminia  reflexa 

Ib.  Plants  aquatic,  partly  aquatic,  or  terrestrial,  if  aquatic  the  leaves  floating  or  emergent  and  not  with 

submerged  filiform  pinnatisect  divisions 2 

2a.  Plants  slender-stemmed  twining  or  climbing,  usually  found  in  gardens  and  near  habitations;  leaves 

mostly  alternate  3 

2b.  Plants  erect  or  creeping  on  the  ground  or  aquatic,  not  twining  or  climbing,  found  in  a  great  many 

habitats;  leaves  alternate  or  opposite    5 

3a.  Leaf  blades  reniform  to  orbicular,  cordate  at  base,  10-35  mm  long,  alternate  or  rarely  op- 
posite; corollas  7-9  mm  long,  with  a  basal  spur Cymbalaria  muralis 

3a.  Leaf  blades  often  triangular-subcordate,  to  60  mm  long,  always  alternate;  corollas  30-80  mm 

long,  without  a  spur    4 

4a.  Corollas  60-70  mm  long;  calyx  15-24  mm  long;  seeds  with  lateral  wings  around  much  of 

the  seed Lophospermum  erubescens 

4b.  Corollas  30-40  mm  long;  calyx  9-15  mm  long;  seeds  without  thin  wings  ....  Maurandya 
5a.  (from  2b)  Leaves  deeply  dissected  or  compound,  the  lobes  or  leaflets  >  30%  the  width  of  the 

blade    6 

5b.  Leaves  entire  to  crenate,  dentate  or  lobed,  the  lobes  or  teeth  <  30%  the  width  of  the  blade    .  .  8 
6a.  Fruits  linear;  plants  small  (40  cm),  erect,  in  open  weedy  lowland  (0-600  m)  sites;  leaves 

opposite,  to  20  mm  long Schistophragma  mexicana 

6b.  Fruits  broader;  plants  usually  taller,  in  montane  (1 100-3800  m)  habitats;  leaves  opposite  or 

alternate,  10-100  mm  long 7 

7a.  Leaves  opposite;  corolla  bright  yellow  with  the  lower  lip  sac-like;  calyx  4-lobed,  green   .  .  . 

Calceolaria 

7b.  Leaves  alternate;  corolla  and  bracts  red,  orange,  yellow,  or  green,  the  lower  lip  usually 
narrowed  and  with  acute  lobes;  calyx  4-parted  but  usually  united  to  form  2  large  lateral  lobes, 

often  colored Castilleja 

8a.  (from  5b)  Lower  leaves  alternate,  plants  usually  with  leaves  alternate  all  along  the  stem 9 

8b.  Lower  leaves  opposite  or  whorled,  distal  leaves  opposite  or  alternate  along  the  stem  or  the  leaves 

rosulate  from  the  base 13 

9a.  Corolla  expanded  at  the  base  and  spurred  [yellow  or  blue,  5-30  mm  long;  lower  leaves  linear 

to  lanceolate;  cultivated  for  ornament] Linaria 

9b.  Corolla  tubular,  not  expanded  at  the  base  to  become  saccate  or  spurred;  wild,  naturalized,  or 

cultivated  species    10 

lOa.  Plants  repent  with  creeping  stems  rooting  at  most  nodes;  leaf  blades  rounded-orbicular  [to 
25  mm  wide];  corolla  4-8-lobed,  rotate,  3-5  mm  wide    Sibthorpia  repens 


FIELDIANA:  BOTANY 


lOb.  Plants  erect,  not  rooting  at  distal  nodes;  leaf  blades  not  rounded-suborbicular;  corolla  not 

rotate 11 

11  a.  Corolla  with  the  upper  lip  galeate,  elongate,  narrowly  1-lobed  and  entire,  lower  lip  shorter 
than  the  upper,  small  and  often  recurved;  distal  bracts  often  becoming  colorful  [not  found 

below  700  m  elevation]    Castilleja 

lib.  Corolla  not  as  above,  lobes  rounded  with  the  lower  lobes  usually  longer  than  the  upper; 

bracts  not  becoming  colorful    12 

12a.  Corolla  7-10  mm  long,  white,  campanulate;  stamens  5;  native  plants  at  0-500  m  elevation.  .  .  . 

Capraria  biflora 

12b.  Corolla  20-50  mm  long,  usually  pink  marked  with  purple  dots  within,  tubular;  stamens  4; 
in  gardens  or  naturalized  at  1800-3300  m  elevation  (if  at  lower  elevations  and  with  viscous 

hairs  or  slime  glands,  see  Martyniaceae  and  Pedaliaceae)    Digitalis  purpurea 

13a.  (from  8b)  Plants  lacking  erect  stems,  <  7  cm  tall;  found  only  in  wet  sites  above  3300  m  elevation 
in  Central  America;  leaves  linear  to  oblanceolate,  in  dense  fascicles  or  rosulate  at  the  nodes;  corolla 

2-3  mm  long,  rotate,  with  3-5  lobes    Limosella  acaulis 

13b.  Plants  with  erect  stems  or  without  the  above  combination  of  characteristics    14 

14a.  Leaf  blades  linear  to  linear-lanceolate,  often  scabrous,  larger  blades  not  exceeding  7  mm  in  width 

[plants  with  stiff  erect  stems  but  rarely  exceeding  0.7  m  height;  corollas  8-15  mm  long]    ...    15 

14b.  Leaf  blades  narrowly  lanceolate  to  broadly  ovate  or  rounded,  scabrous  or  smooth;  larger  blades 

usually  >  7  mm  in  width    18 

15a.  Corolla  20-45  mm  long  or  with  spreading  lobes  to  2  cm  wide,  magenta,  red,  blue-violet,  or 
purple  to  yellowish,  glabrous  or  densely  puberulent  externally;  flowers  pedicellate  in  leaf 
axils  or  in  distal  racemes;  plants  not  wiry  or  scabrous  herbs  or  subshrubs,  0.4-1.5  m  tall  .  . 

...    16 

15b.  Corolla  7-15  mm  long,  lavender  to  pale  purple,  glabrous  externally;  flowers  sessile  or  ped- 
icellate; plants  wiry  and  scabrous,  herbs  to  0.6  m  tall  17 

16a.  Corolla  densely  minutely  puberulent  externally,  narrowly  tubular  with  small  distal  lobes 

Lamourouxia  spp. 

16b.  Corolla  glabrous  externally,  with  short  tube  and  broadly  spreading  large  lobes   

Angelonia  angustifolia 

17a.  Flowers  borne  on  pedicels  10-35  mm  long;  fruits  4-7  mm  diam.;  erect  stems  usually  with 

several  prominent  lateral  branches Anisantherina  hispidula 

17b.  Flowers  sessile  or  subsessile  on  pedicels  <  3  mm  long;  fruits  2-3  mm  diam.;  erect  stems 

usually  with  few  or  no  prominent  lateral  branches   Buchnera  weberbaueri 

18a.  (from  14b)  Corollas  with  the  lower  lip  forming  a  distal  sac  and  slipper-like,  usually  bright  yellow 

[10-25  mm  long;  plants  both  wild  and  cultivated  for  ornament]    Calceolaria 

18b.  Corollas  not  slipper-like,  the  lower  lip  not  forming  a  distal  sac,  variously  colored,  usually  with  a 

short  or  long  tube  and  spreading  distal  lobes 19 

19a.  Corollas  becoming  >  15  mm  long  at  anthesis 20 

19b.  Corollas  usually  <  15  mm  long  at  anthesis   29 

20a.  Corollas  70-120  mm  long,  white  and  salverform  (with  a  narrow  tube  and  broad  subequal 

rotate  lobes) Escobedia  grandiflora 

20b.  Corollas  14-45  mm  long,  variously  colored,  usually  tubular  and  somewhat  2-lipped  with  the 

lobes  unequal    21 

2 la.  Corolla  yellow  or  yellow  and  white  [flowers  in  axillary  groupings;  leaves  sessile  and  auric- 

ulate  at  the  base,  blades  narrowly  lanceolate;  native  wild  plants] 22 

21b.  Corolla  pink  to  red  or  purple,  marked  with  dark  spots  if  white  or  yellowish  (note  that  Mar- 
tynia  and  Sesamum  may  key  out  in  this  dichotomy;  see  Martyniaceae  and  Pedaliaceae,  Fig. 

27) 24 

22a.  Plants  weak-stemmed  herbs  to  0.4  m  tall;  leaves  petiolate  with  blades  rounded  or  trun- 
cate at  the  base;  flowers  solitary  in  leaf  axils  [corolla  14-20  mm  long] 

Mimulus  glabratus 

22b.  Plants  erect  subshrubs,  often  to  1  (2)  m  tall;  leaves  sessile  with  blades  lanceolate  and 
gradually  narrowed  to  the  base;  flowers  usually  several  in  leaf  axils    23 

BURGER:  FLORA  COSTARICENSIS  3 


23a.  Corolla  25-45  mm  long,  calyx  11-16  mm;  fruits  dry,  brownish,  oblong,  13-17  mm 

long    Hemichaena  fruitcosa 

23b.  Corolla  14-23  mm  long,  calyx  5-9  mm  long;  fruits  fleshy,  white,  globose,  7-10  mm 

long    Leucocarpus  perfoliatus 

24a.  Corolla  densely  minutely  puberulent  or  glandular  puberulent  on  the  exterior   25 

24b.  Corolla  glabrous  on  the  exterior 26 

25a.  Corollas  densely  minutely  puberulent  externally,  the  throat  not  closed  by  a  palate;  native 

wild  species    Lamourouxia 

25b.  Corollas  sparsely  puberulent,  throat  usually  closed  by  a  palate;  cultivated  ornamentals 

Antirrhinum 

26a.  Corolla  tube  ca.  2-4  mm  diam.,  with  small  distal  lobes,  deep  red;  plants  shrub-like  orna- 
mentals, often  without  leaves  and  with  many  slender  green  stems       Russelia  equisetiformis 
26b.  Corolla  tube  7-10  mm  diam.,  with  prominent  distal  lobes,  purple  to  rose- white  or  marked 

with  dark  coloring;  plants  not  shrub-like,  rarely  without  leaves 27 

27a.  Plants  of  wet  forests;  stems  usually  unbranched  and  <  30  cm  tall;  leaf  blades  oblanceolate, 

to  3 1  cm  long Tetranema 

27b.  Plants  of  gardens  and  ornamental  plantings;  stems  usually  with  a  few  branches,  0.4-1  m  tall; 

leaf  blades  not  oblanceolate,  to  10  cm  long    28 

28a.  Plants  becoming  ca.  1  m  tall;  calyx  united  for  only  a  short  distance,  lobes  >  half  the  length 

of  the  calyx;  staminode  prominent;  grown  as  ornamentals  in  Costa  Rica    

Penstemon  gentianoides 

28b.  Plants  to  0.4  m  tall;  calyx  united  for  half  its  length  or  more,  lobes  <  half  the  length  of  the 

calyx;  wild  and  ornamental  species  in  Costa  Rica    Torenia 

29a.  (from  20b)  Calyx  united  for  >  half  its  length  at  anthesis,  a  calyx  tube  clearly  present  (note  that 
flowers  may  appear  to  have  separate  sepals  after  anthesis  as  the  fruit  develops  and  the  calyx  tube 

splits)  30 

29b.  Calyx  with  the  sepals  united  only  at  the  base,  a  calyx  tube  not  evident 35 

30a.  Stems  and  leaves  scabrid  or  hispid,  stems  often  becoming  1  m  tall;  flowers  sessile  or  sub- 
sessile  in  distal  spike-like  inflorescences   31. 

30b.  Stems  and  leaves  neither  scabrid  nor  hispid,  stems  rarely  >  40  cm  tall;  flowers  not  sessile 

in  spike-like  terminal  inflorescences  (except  in  Bacopa  sessiliflora)    32 

3 la.  Corolla  yellow,  campanulate;  flowers  subtended  by  large  bracts  differing  little  from  the 

leaves;  distal  leaves  narrowly  triangular   Alectra  aspera 

31b.  Corolla  white  or  marked  with  purple  or  blue,  salverform;  flowers  subtended  by  bracts 

much  smaller  than  the  leaves;  distal  leaves  narrowly  lanceolate   .  .  .    Buchnera  pusilla 

32a.  Lower  leaves  forming  a  rosette,  blades  narrowly  obovate  with  long-attenuate  base;  flowers 

in  lax  terminal  racemes;  rarely  encountered  introduced  weeds Mazus  pumila 

32b.  Lower  leaves  not  forming  dense  rosettes,  blades  narrowly  obovate  to  suborbicular;  flowers 

mostly  axillary  to  distal  leaves 33 

33a.  Calyx  4-lobed;  stems  terete;  leaf  blades  varying  from  obovate  to  suborbicular  (rarely  ovate 

with  truncated  base)   Bacopa  egensis 

33b.  Calyx  5-lobed;  stems  4-angled  or  with  2-4  longitudinal  ridges;  leaf  blades  mostly  ovate- 
triangular  or  ovate-ellipsoid  with  truncated  base 34 

34a.  Calyx  tube  <  4  mm  long,  usually  campanulate;  fruits  3-4  mm  long,  rounded  ovoid 

Lindernia 

34b.  Calyx  tube  usually  >  7  mm  long,  tubular;  fruits  8-10  mm  long,  narrowly  ellipsoid-oblong 

Torenia  thouarsii 

35a.  (From  29b)  Fruits  usually  lenticular  in  cross-section,  truncated  or  rounded  and  obcordate  at  the 
apex;  stamens  2;  corolla  tube  very  short,  with  corolla  lobes  usually  in  a  single  plane  (rotate); 

introduced  weeds,  1000-3300  m  elevation Veronica 

35b.  Fruits  usually  round  in  cross-section,  never  truncated  or  obcordate  at  the  apex,  and  without  the 

other  combination  of  characteristics   36 

36a.  Sepals  very  unequal,  the  outer  usually  much  broader  than  the  inner  and  imbricate  in  bud  (note 
that  small  linear  bracteoles  may  be  present  at  the  base  of  the  calyx)  37 

4  FIELDIANA:  BOTANY 


36b.  Sepals  equal  or  subequal,  valvate  or  less  often  imbricate  in  bud    38 

37a.  Flowers  yellow;  bracteoles  present  at  the  base  of  the  pedicel;  leaf  blades  broad  and  abruptly 

narrowed  to  a  short  petiole Mecardonia  procumbens 

37b.  Flowers  purple  or  blue  to  white  (sometimes  yellowish  in  age);  bracteoles  present  in  the  middle 
or  apex  of  the  pedicel  or  absent;  leaf  blades  usually  gradually  narrowed  to  the  base  or  sessile 

if  rounded  at  the  base Bacopa 

38a.  Corollas  bright  red  or  orange;  plants  erect  herbs  or  subshrubs  to  2  m  tall,  stems  often  stiff  and 

longitudinally  ridged   39 

38b.  Corollas  white  to  blue,  purple,  or  yellowish;  plants  mostly  weak-stemmed  herbs    41 

39a.  Leaf  blades  15-31  cm  long;  corollas  26-55  mm  long;  peduncles  9-24  cm  long,  axillary, 

slender  and  flexuous;  rarely  collected  endemic  species Tetranema 

39b.  Leaf  blades  1-11  cm  long;  corollas  5-16  mm  long;  peduncles  less  than  2  cm  long  or  the 

flowers  on  terminal  erect  racemes;  common  and  widespread  species 40 

40a.  Flowers  in  axillary  fascicles  of  2-20,  corolla  deep  red,  tube  9-12  mm  long  with  small  acute 

lobes;  seeds  surrounded  by  hairs  within  the  fruit    Russelia  sarmentosa 

40b.  Flowers  solitary  in  bract  or  leaf  axils  on  a  terminal  raceme,  corolla  red  to  orange,  tube  <  2 
mm  long  with  large  rounded  spreading-rotate  lobes;  seeds  not  associated  with  hair-like  struc- 
tures   Alonsoa  meridionalis 

4 la.  Corolla  <  4  mm  long,  rotate  or  campanulate 42 

41b.  Corolla  4-14  mm  long,  usually  tubular  and  bilabiate    43 

42a.  Plants  terrestrial,  erect,  to  1  m  tall;  leaf  blades  to  35  mm  long,  oblanceolate  and  dentate; 

fertile  stamens  4   Scoparia 

42b.  Plants  floating  or  prostrate,  small;  leaves  to  8  mm  long,  rounded  and  entire;  fertile  stamens 

2    Micranthemum  umbrosum 

43a.  Fertile  stamens  2;  leaves  sessile;  stems  glabrous   Lindernia  dubia 

43b.  Fertile  stamens  4;  leaves  petiolate  (sessile  and  amplexicaul  in  Stetnodia  durantifolia);  stems  gla- 
brous or  puberulent   44 

44a.  Anthers  glabrous;  stems  and  leaves  puberulent;  sepals  acute  at  the  apex   Stemodia 

44b.  Anthers  with  hairs;  stems  and  leaves  glabrous;  sepals  bluntly  acute  and  often  thickened  at  the  apex 
Darcya 


Agalinis  Rafinesque 
Nomen  conservandum 

Herbs  or  shrubs,  annual  or  perennial,  usually 
erect  and  branched,  glabrous  or  pubescent,  hem- 
iparasitic  on  roots,  turning  dark  when  dried. 
Leaves  opposite  or  alternate,  becoming  smaller  or 
bract-like  on  distal  stems,  subsessile  or  sessile, 
blades  usually  narrow,  entire  or  lobed,  glabrous 
or  puberulent.  Inflorescences  of  solitary  flowers 
in  axils  of  reduced  distal  leaves  (often  resembling 
open  racemes,  spikes,  or  panicles),  peduncles 
slender,  subtended  by  small  bracts,  pedicels  slen- 
der, bibracteolate  or  ebracteolate.  Flowers  showy, 
calyx  tubular  to  campanulate  or  hemispheric,  with 
5  prominent  lobes  or  teeth,  slightly  imbricate  or 
open  in  bud;  corolla  campanulate  and  often  some- 
what bilabiate,  glabrous  or  puberulent,  pink  to 
purple  (yellow  or  white),  tube  straight  or  curved, 
expanded  distally  into  a  broad  throat,  lobes  5, 
rounded,  subequal  or  the  posterior  (upper)  small- 
er; stamens  4,  of  2  unequal  pairs,  borne  from  the 


middle  of  the  corolla  tube,  shorter  than  the  corol- 
la, filaments  usually  pilose,  anthers  glabrous  or 
puberulent,  2-thecous  with  parallel  thecae,  un- 
equal in  some  species;  ovary  glabrous,  2-locular, 
style  straight  and  slender,  deciduous,  stigma  soli- 
tary and  linear.  Fruits  rounded  capsules,  woody 
to  chartaceous  or  leathery,  dehiscence  loculicidal 
(and  sometimes  septicidal);  seeds  many,  oblong  to 
angular,  testa  reticulate. 

A  distinctive  New  World  genus  with  ca.  40, 
mostly  North  American,  species.  Species  of  this 
genus  were  formerly  placed  in  Gerardia,  but  the 
Linnaean  type  proved  to  be  a  species  of  Acantha- 
ceae.  This  genus  has  not  been  collected  in  Costa 
Rica,  but  ranges  southward  as  far  as  northern  Nic- 
aragua. The  following  key  includes  Anisantherina 
hispidula,  which  occurs  in  Costa  Rica  (q.v.),  and 
two  species  of  Agalinis  known  from  Nicaragua: 
Agalinis  albida  Britton  &  Pennell  (?  =  Agalinis 
harperi  Pennell)  and  Agalinis  peduncularis 
(Benth.)  Pennell.  All  three  species  have  linear 
leaves. 


BURGER:  FLORA  COSTARICENSIS 


Bacopa 
monnieri 


Micranthemum  umbrosum 


.imosella  aquatica 
jar.  americana 


FIG.  1.     Scrophulariaccac:  aquatic  and  semiaquatic  species. 


FIELDIANA:  BOTANY 


Stemodia 
angulata 


Stemodi  a 
verticillata  r 


veronica 
serpyl li  folia 


Mecardonia 
procumbens  " 


Lindernia  diffusa' 

FIG.  2.     Scrophulariaccac:  small  herbs  with  opposite  leaves  and  broad  blades. 


V.  polita 


BURGER:  FLORA  COSTARICENSIS 


//  Darcya 
costaricensis 


Stemodi a 
peduncularis 


Calceolaria 
irazuensis 


FIG.  3.     Scrophulariaceae:  herbs  with  opposite  leaves  and  lanceolate  to  ovate  or  dissected  blades. 


FIELDIANA:  BOTANY 


Lindernia 
dubi  a 


Schistophragma  mexicana 


Anisanthenna 
hispidula 


Stemodi  a 
duranti folia 


Scoparia  dulcis 


FIG.  4.     Scrophulariaceae:  herbs  with  opposite  leaves  and  narrow  leaf  blades  or  narrow  pinnatifid  lobes. 


BURGER:  FLORA  COSTARICENSIS 


Alectra  aspera 


Russell  a 
sarmentosa 


Lamourouxia 
guttierrezii 


L.  lanceolata 

FIG.  5.     Scrophulariaccac:  herbs  with  stiff  opposite  leaves  and  erect  or  clambering  stems. 


10 


FIELDIANA:  BOTANY 


Calceolaria 
perfol i  ata 


Leucocarpus 
perfol iatus 


Escobedia 
grandi flora 


Stemodia 

duranti  folia 
v 


FIG.  6.     Scrophulariaccac:  erect  herbs  with  larger  sessile  or  pcrfoliatc  leaves. 


BURGER:  FLORA  COSTARICENSIS 


11 


Castilleja 
tayloriorum 


Castilleja 
irazuensis 


FIG.  7.     Scrophulariaccac:  erect  herbs  with  consistently  alternate  leaves  and  unusual  curved  flowers  (Castilleja)  or 
simple,  almost  regular  flowers  (Capraria). 


12 


FIELDIANA:  BOTANY 


Maurandya  barclaiana 


Torenia 
fournieri 


Lophospermum 
erubescens 


Antirrhinum 
majus 


Penstemon /gent i  anoi  des 
Angel onia'angusti folium      "       ^ 


FIG.  8.     Scrophulariaceae:  wild  and  cultivated  species  with  large  colorful  corollas  and  alternate  or  opposite  leaves. 


BURGER:  FLORA  COSTARICENSIS 


13 


Key  to  Species  of  Agalinis  and  Anisantherina 

la.   Flowers  usually  I/node,  pedicels  shorter  than  the  calyx;  distal  leaves  mostly  alternate;  stems  and 

leaves  smooth  to  the  touch  Agalinis  albida 

Ib.  Flowers  usually  2/node,  pedicels  much  longer  than  the  calyx;  distal  leaves  mostly  opposite  or 

suhopposite;  stems  and  leaves  slightly  scabrous 2 

2a.  Corolla  20-30  mm  long,  edge  of  corolla  lobes  with  multicellular  hairs;  anther  thecae  equal  in  size 

and  shape;  calyx  lobes  narrow;  pedicels  without  nodes  or  bracts  Agalinis  peduncularis 

2b.  Corolla  10-16  mm  long,  edge  of  corolla  lobes  glabrous;  anther  thecae  slightly  unequal  in  size  and 

shape;  calyx  lobes  triangular;  pedicels  usually  with  a  node  or  bracts  (reduced  leaves)  near  the  middle 
Anisantherina  hispidula 


Alectra  Thunberg 
nomen  conservandum 

REFERENCE — H.  Melchior,  Die  Gattung  Alectra 
Thunb.  Notizbl.  Hot.  Gart.  Berlin  15:  423-447. 
1941. 

Erect  annual  herbs,  hemiparasitic,  turning  dark 
when  dried,  unbranched  to  few-branched,  with 
hispid  or  scabrous  hairs  often  enlarged  at  the  base. 
Leaves  opposite  or  subopposite  (or  alternate 
above),  reduced  in  size  distally  (rarely  absent), 
simple  and  sessile  or  subsessile,  margins  serrate 
to  dentate  (entire),  often  with  3  prominent  veins 
from  the  base.  Inflorescences  racemose  or  spi- 
cate,  elongate,  of  solitary  flowers  in  axils  of  distal 
leaves,  pedicels  (peduncles)  short,  with  2  (1)  dis- 
tal bracts  or  bracts  absent.  Flowers  with  campan- 
ulate  calyx,  10- veined,  persisting  and  enclosing 
the  fruit,  with  5  acute  to  obtuse  lobes,  valvate  in 
bud,  equaling  the  tube  in  length;  corolla  usually 
yellow  or  orange,  personate  to  subglobose  or  cam- 
panulate,  as  long  or  slightly  longer  than  the  calyx, 
5-lobed  and  slightly  bilabiate,  lobes  rounded, 
shorter  or  equaling  the  tube;  stamens  of  2  shorter 
and  2  longer  pairs,  attached  near  the  base  of  the 
corolla,  included,  filaments  glabrous  or  puberu- 
lent,  anthers  2-thecous,  often  barbate  abaxially, 
disc  annular  and  fleshy;  ovary  ovate  or  com- 
pressed, 2-locular  with  thick  fleshy  placenta  and 
many  seeds,  style  linear,  elongate,  and  inflexed, 
stigma  entire  or  bifid,  thickened.  Fruits  loculici- 
dal  capsules  enclosed  within  the  dry  calyx,  round- 
ed or  compressed;  seeds  very  numerous,  small, 
with  transparent  exotesta  often  truncated  and  open 
at  the  2  ends. 

A  genus  of  41  species,  according  to  Melchior 
(1941,  above).  Nearly  all  the  species  are  from  Af- 
rica and  India.  The  one  American  species  is  native 
to  South  America  and  the  West  Indies;  it  differs 
from  the  Old  World  species  in  having  divergent 


rather  than  parallel  anther  thecae.  Alectra  is  close- 
ly related  to  Melasma  in  the  tribe  Buchnereae  and 
was  once  united  with  that  genus.  Few  characters 
reliably  distinguish  the  two.  In  Melasma  the  co- 
rolla is  twice  as  long  as  the  calyx  and  is  more 
campanulate  than  that  of  Alectra. 

Alectra  aspera  (Cham.  &  Schldl.)  L.  O.  Wil- 
liams, Fieldiana,  Bot.  34:  118.  1972.  Pedicu- 
laris  melampyroides  L.  C.  Rich.,  Actes  Soc. 
Hist.  Nat.  Paris  1:  111.  1792.  Glossostylis  as- 
pera Cham.  &  Schldl.,  Linnaea  3:  22.  1828. 
Scrophularia  fluminensis  Veil.,  Fl.  flumin.,  263. 
1829.  A.  brasiliensis  Benth.  in  DC.,  Prodr.  10: 
339.  1846.  A.  melampyroides  (L.  C.  Rich.) 
Kuntze,  Rev.  Gen.  PI.  2:  458.  1891,  non  A.  me- 
lampyroides Benth.  1846.  Melasma  melampy- 
roides (L.  C.  Rich.)  Pennell  in  Britton  &  Wil- 
son, Bot.  Porto  Rico  6:  188.  1925.  A.  fluminen- 
sis (Veil.)  Steam,  J.  Arnold  Arbor.  52:  636. 
1971.  Figure  5. 

Herbs  0.3-1.5  m  tall,  stems  erect  and  simple 
to  many-branched,  main  stems  1.7-6  mm  diam., 
scabrid  with  straight  stiff  whitish  hairs  0.3-1.5 
mm  long,  short  (0.1-0.2  mm)  thin  hairs  also  pre- 
sent. Leaves  becoming  gradually  smaller  distally, 
opposite,  subopposite  (rarely  alternate  distally), 
subsessile  with  petioles  0.5-4  mm  long;  leaf 
blades  l-5(-7)  cm  long,  4-16(-25)  mm  wide, 
narrowly  ovate-triangular  to  ovate-lanceolate  or 
lanceolate,  gradually  narrowed  to  the  acute  or 
acuminate  apex,  margin  coarsely  serrate  with 
teeth  0.3-2.5  mm  high,  base  truncated  or  obtuse, 
surfaces  scabrous  with  stiff  whitish  hairs  0.1-0.9 
mm  long,  venation  subpalmate  with  midvein  and 
2  prominent  ascending  basal  2°  veins  (triplivei- 
ned).  Inflorescences  of  spike-like  or  raceme-like 
distal  stems  or  with  solitary  flowers  in  the  axils 
of  smaller  leaves  (1-2  flowers/node),  pedicles  1- 
2(-7)  mm  long,  ca.  0.5  mm  diam.,  bracts  linear 


14 


FIELDIANA:  BOTANY 


or  absent,  at  the  base  of  the  calyx,  ca.  3  mm  long. 
Flowers  with  campanulate  calyx  6-9  mm  long  (to 
12  mm  in  fruit),  5-8  mm  wide  at  the  mouth,  calyx 
lobes -2-6  mm  long,  triangular  and  acute,  with 
scabrous  hairs  along  the  margins  and  veins;  co- 
rolla 10-13  mm  long,  included  within  the  calyx 
or  slightly  exserted,  campanulate,  yellow,  gla- 
brous, tube  5-8  mm  long,  lobes  subequal;  stamens 
with  pubescent  filaments,  anthers  1.5  mm  long, 
thecae  divergent,  without  awns;  ovary  ca.  3  mm 
long,  ovoid,  style  6-8  mm  long,  curved,  stigma 
lanceolate.  Fruits  5-8  mm  long,  4-6  mm  diam., 
oblate-globose,  included  within  the  persisting  ca- 
lyx; seeds  1-1.2  mm  long,  oblong-triangular  with 
truncated  ends,  translucent  yellowish,  exotesta  re- 
ticulate. 

Weeds  of  open  sunny  sites  in  lowland  Central 
America.  It  is  a  recent  introduction  and  is  spread- 
ing. It  has  been  collected  near  Upala  and  Villa 
Neilly,  at  Tarrazu,  and  on  the  lower  slopes  of  the 
Cerros  de  Puriscal.  The  species  ranges  from  Gua- 
temala and  the  West  Indies  to  Brazil,  Parguay,  and 
Bolivia. 

Alectra  aspera  is  recognized  by  its  short  erect 
stems,  stiff  pustulate-hispid  leaves,  tissues  drying 
dark,  solitary  flowers  in  distal  leaf  axils,  corolla 
tube  only  as  long  as  the  calyx,  persisting  campan- 
ulate-globose  calyx,  and  minute  seeds  with  trans- 
lucent testa.  The  corolla  of  this  species  is  open  for 
a  very  short  time.  It  soon  withers,  closing  over 
the  anthers  and  the  stigma.  The  anthers  are  borne 
close  to  the  stigma,  suggesting  that  the  species  is 
largely  self-pollinating. 


Alonsoa  Ruiz  Lopez  &  Pavdn 

REFERENCES — J.  L6pez  Guilldn,  El  Genero 
Alonsoa  en  el  Peru:  1.  Revision  de  las  especies 
endemicas.  Raymondiana  3:  155-246.  1970 
(1971).  Brian  Wrigley,  A  taxonomic  revision  of 
Alonsoa.  Ph.D.  diss.,  Univ.  Connecticut,  1968. 

Herbs  or  subshrubs,  annual  or  perennial,  often 
with  distal  branching,  usually  woody  at  the  base, 
stems  quadrangular  in  cross-section,  glabrous  or 
sparsely  minutely  puberulent.  Leaves  opposite  or 
ternate  (leaf-like  floral  bracts  alternate),  sessile  or 
petiolate,  leaf  blades  ovate  to  linear,  serrate  or 
rarely  entire,  venation  pinnate.  Inflorescences  ra- 
cemes, terminal  or  axillary  to  distal  leaves  (rarely 
flowers  axillary  to  distal  leaf  pairs),  floral  bracts 
alternate  along  the  rachis,  proximal  bracts  leaf- 
like,  pedicels  solitary,  well  developed,  and  be- 
coming twisted,  bracteoles  absent.  Flowers  small. 


resupinate  because  of  pedicel  twisting,  calyx 
deeply  5-parted.  lobes  narrow  and  slightly  un- 
equal, glabrous  or  sparsely  and  minutely  puberu- 
lent, valvate  in  bud;  corolla  rotate  with  very  short 
tube,  bilaterally  symmetric  with  the  2  lower  lobes 
small  and  divided  nearly  to  the  base,  lateral  lobes 
short  and  broad,  upper  lobe  much  larger  and  usu- 
ally held  erect  and  convex,  reddish  to  orange  or 
purple;  stamens  4,  subequal,  borne  on  the  base  of 
the  corolla,  filaments  thick  or  slender,  anthers 
closely  positioned  around  the  style,  thecae  parallel 
or  divergent;  ovary  2-locular,  style  curved  up- 
ward, stigma  capitate.  Fruits  capsules,  ovate  to 
oblong,  septicidal,  2-valved,  valves  chartaceous, 
entire  or  bifid  at  the  apex;  seeds  many,  punctate- 
rugose  or  longitudinally  ridged. 

Alonsoa  is  a  Neotropical  genus  of  six  to  fifteen 
species.  The  flowers  are  upside-down  (resupinate) 
because  of  the  twisted  pedicel,  with  the  result  that 
the  morphologically  lower  lobe  is  the  upper  lobe. 
The  reddish  to  yellow  resupinate  flowers  with  en- 
larged median  lobe  are  distinctive.  The  genus  is 
placed  in  the  tribe  Hemimeridae,  with  Angelonia, 
but  is  probably  more  closely  related  to  Scrophu- 
laria  and  Verbascum,  with  which  it  shares  char- 
acters of  seed  and  anther  morphology.  Alonsoa 
warscewiczii  Regel  is  often  used  as  a  potted  or- 
namental plant;  only  one  species  is  found  in  the 
higher  mountains  of  Central  America.  The  genus 
is  currently  being  studied  by  Fanny  Astholm 
(GB). 


Alonsoa  meridionalis  (L.f.)  Kuntze,  Rev.  Gen. 
PI.  2:  457.  1891. 

Scrophularia  meridionalis  L.f.,  Suppl.  280. 
1781.  Figure  3. 

REFERENCE — F  Astholm  &  Y.  Nyman,  Morpho- 
metric  variation  in  the  Alonsoa  meridionalis  com- 
plex (Scrophulariaceae).  Plant  Syst.  Evol.  193: 
53-68.  1994. 

Herbs  or  subshrubs  0.2-1.5  m  tall,  often 
woody  at  the  base,  usually  with  lateral  branches, 
leafy  stems  0.7-8  mm  diam.,  glabrous  or  sparsely 
minutely  puberulent  (rarely  densely  puberulent  at 
the  node),  with  4  prominent  longitudinal  ridges  or 
wings  (from  decurrent  petiole  margins).  Leaves 
opposite  (sometimes  alternate  below  the  flowering 
nodes),  petioles  3-15(-22)  mm  long,  0.4-1.7  mm 
diam.,  glabrous  or  with  few  hairs  less  than  0.3 
mm  long,  with  lateral  margins  continuous  with  the 
blade  margins;  leaf  blades  1.5-8(-ll)  cm  long, 


BURGER:  FLORA  COSTARICENSIS 


15 


1-4  cm  wide,  ovate- lanceolate  to  oblong-lanceo- 
late or  narrowly  ovate,  apex  acute,  margin  with 
4-7  acute  teeth/cm,  base  obtuse  to  slightly  cune- 
ate  and  decurrent,  drying  greenish  or  dark,  sparse- 
ly puberulent  on  both  surfaces,  2°  veins  5-10/side, 
strongly  ascending.  Inflorescences  terminal  ra- 
cemes 3-30  cm  long,  flowers  opposite  or  distally 
alternate,  subtended  by  progressively  smaller, 
leaf-like  to  subulate  bracts  4-14  mm  long,  pedi- 
cels 6-18(-25)  mm  long,  glabrous  or  with  few 
minute  (0.2  mm)  gland-tipped  hairs,  upcurved  in 
fruit.  Flowers  resupinate,  calyx  3-6  mm  long,  di- 
vided to  near  the  base,  lobes  1-1 .7  mm  wide,  sub- 
equal  or  unequal,  glabrous  or  with  few  minute 
hairs  near  the  base;  corolla  5-1 1  mm  long,  rotate, 
yellow  to  orange,  tube  ca.  2  mm  long,  with  prom- 
inent lobes,  median  lobe  to  15  mm  long,  larger 
than  the  lateral  lobes;  stamens  3-4  mm  long,  fil- 
aments 0.4-0.6  mm  diam.,  anthers  ca.  2  mm  long; 
ovary  1.5-4  mm  long,  narrowly  ovoid,  style  1.5- 
2  mm  long,  stigma  0.8  mm  wide.  Fruits  7-15  mm 
long,  3.7-6  mm  wide,  narrowly  ovoid  with  acute 
to  acuminate  apex,  smooth,  glabrous,  pale  brown, 
sulcate  along  the  plane  of  dehiscence,  septicidal; 
seeds  many,  1.1-1.4  mm  long,  1-1.2  mm  diam., 
oblong,  dark,  with  ca.  6  deep  longitudinal  sulci. 

Plants  of  open  sites  in  high  montane  forest  for- 
mations, 1600-3200  m  elevation.  Flowering  in 
October-December.  In  Costa  Rica  they  are  found 
in  the  Cordillera  Central  and  the  Cordillera  de  Tal- 
amanca.  The  species  ranges  from  southern  Mex- 
ico to  Bolivia. 

Alonsoa  meridionalis  is  recognized  by  its  ter- 
minal racemes  with  solitary,  usually  alternate 
flowers  subtended  by  bracts  gradually  diminishing 
in  size,  unusual  little  flowers  with  red-orange  co- 
rolla with  enlarged  median  uppermost  lobe,  short 
thick  filaments,  large  anthers,  and  capsules  grad- 
ually narrowed  to  the  apex.  Because  the  pedicels 
are  twisted  1 80°,  the  flowers  are  upside-down  (re- 
supinate). The  leaves  often  have  new  shoots  with 
small  leaves  in  their  axils. 


Angelonia  Humboldt  &  Bonpland 

REFERENCE — K.  Barringer,  A  Revision  of  An- 
gelonia (Scrophulariaceae).  Ph.D.  diss.  Univ. 
Connecticut,  1981. 

Erect  herbs  or  subshrubs,  annual  or  perennial, 
stems  terete  to  4-anguIate,  simple  or  branching 
from  the  base,  puberulent  with  multicellular  hairs 
or  glabrous.  Leaves  opposite  or  rarely  alternate 
distally,  petioles  short  or  absent;  blades  ovate  to 


linear,  serrate  or  rarely  entire,  apex  acute,  vena- 
tion pinnate.  Inflorescences  racemes,  terminal  or 
axillary,  bracts  leaf-like  to  rounded,  flowers  1-3/ 
axil,  pedicels  bibracteolate  or  ebracteolate.  Flow- 
ers showy,  calyx  of  5  free  or  partly  united  sepals, 
sepals  equal,  lanceolate,  entire,  acute  to  acumi- 
nate; corolla  strongly  bilaterally  symmetric,  cu- 
pular-campanulate  with  short  tube  and  2  broadly 
flaring  lips,  bisaccate  at  the  base  of  the  median 
lobe,  the  sacs  with  a  dense  mat  of  glandular  hairs 
within,  lobes  5,  upper  lobes  2,  the  lower  median 
(abaxial)  lip  3-lobed,  variously  ornamented  with 
a  ridge  or  a  crateriform  palate  and  a  bifid  tooth; 
stamens  4,  of  2  unequal  pairs,  held  against  the 
upper  part  of  the  corolla  tube,  filaments  short,  the- 
cae  divaricate,  without  spurs;  ovary  ovoid,  2-loc- 
ular,  ovules  many,  style  longer  than  the  ovary, 
stigma  entire  and  minute.  Fruits  dry  capsules, 
ovoid  to  broadly  ellipsoid,  chartaceous  or  leath- 
ery, septicidal,  often  secondarily  loculicidal;  seeds 
many,  obconical,  with  a  loose  reticulate  exotesta, 
endosperm  absent. 

A  genus  of  26  species  whose  major  concentra- 
tion is  in  the  dry  caatinga  and  cerrado  formations 
of  Brazil.  The  Central  American  and  Caribbean 
species  form  a  distinctive  group  within  the  genus. 
The  genus  is  easily  distinguished  by  the  bisaccate 
corolla  and  the  ornamented  median  corolla  lobe. 
One  species  is  found  in  Costa  Rica;  a  second,  An- 
gelonia ciliaris  B.  L.  Robinson  (with  puberulent 
leaves  auriculate  at  the  base),  ranges  from  south- 
ern Mexico  and  the  Antilles  to  Nicaragua. 

Angelonia  angustifolia  Bentham  in  DC.,  Prodr. 
10:  254.  1846.  Figure  8. 

Herbs  20-120  cm  tall,  erect,  few-branched  or 
unbranched,  leafy  stems  1.5-5  mm  diam.,  terete 
or  slightly  4-angled,  sparsely  puberulent  with  thin 
multicellular  hairs  0.5-2  mm  long  at  the  nodes  or 
along  longitudinal  lines.  Leaves  becoming  small- 
er distally  (intergrading  with  the  floral  bracts),  op- 
posite or  subopposite,  sessile  or  subsessile,  often 
clasping  the  stem;  leaf  blades  2-11  cm  long,  4- 
20  mm  wide,  linear  to  narrowly  elliptic-oblong  or 
lanceolate,  apex  acute,  margin  obscurely  serrate 
with  short  (0.2-0.6  mm)  teeth  1-4/cm,  base  cu- 
neate  to  acute,  drying  chartaceous  and  brown  or 
dark  grayish  green,  surfaces  subglabrous  with  few 
thin  hairs,  2°  veins  3-6/side  and  strongly  ascend- 
ing. Inflorescences  5-40  cm  long,  flowers  soli- 
tary in  distal  leaf/bract  axils  (usually  2/node), 
pedicels  4-18  mm  long,  0.2-0.4  mm  diam.,  gla- 
brous, ascending,  subtended  by  2  linear  bracteoles 
0.5-2  mm  long.  Flowers  glabrous,  calyx  2-4  mm 


16 


FIELDIANA:  BOTANY 


long,  1-2  mm  wide  at  the  base,  narrowly  ovoid 
to  campanulate,  calyx  lobes  2-3.5  mm  long,  lan- 
ceolate to  triangular;  corolla  8-25  mm  wide  at  the 
mouth,  purple  to  lavender  or  bluish,  minutely 
punctate,  tube  white  to  yellowish  or  green,  often 
marked  with  white  on  the  palate  (basal  entry  of 
the  throat),  corolla  lobes  to  9  mm  long,  rounded; 
stamens  included,  filaments  glandular-pubescent, 
anthers  2  mm  wide,  thecae  divergent,  equal,  gla- 
brous; ovary  glabrous.  Fruits  4-6  mm  long,  4-7 
mm  diam.,  globose  to  ovoid-rounded  with  trun- 
cated base,  glabrous;  seeds  1.3-1.5  mm  long,  con- 
ic to  oblong,  exotesta  strongly  reticulated  with 
prominent  thin  walls. 

Native  to  southern  Mexico,  this  species  has 
spread  into  much  of  Central  America,  where  it  is 
a  favorite  garden  ornamental.  In  Costa  Rica  it  has 
been  collected  in  both  the  deciduous  and  ever- 
green lowlands  and  from  gardens  in  the  Meseta 
Central;  it  flowers  throughout  the  year.  The  spe- 
cies is  now  cultivated  throughout  the  world. 

Angelonia  angustifolia  is  recognized  by  its 
small  stature,  narrow  subglabrous  opposite  leaves, 
showy  corolla  with  prominent  spreading  rounded 
lobes,  and  unusual  seed  surface.  This  species  is 
probably  pollinated  by  Centris  bees,  which  collect 
a  thick  oil  from  the  corolla  sacs  with  specialized 
combs  on  their  front  legs.  Common  names  used 
for  this  species  are  boca  de  la  vieja  (Guatemala), 
porto  hello  (Nicaragua),  and  angeldn  (Colombia). 


Anisantherina  Pennell 

Herbs,  annual,  erect  with  ascending  branches, 
hispidulous  with  multicellular  hairs  with  dark 
cross-walls,  hemiparasitic  on  roots,  turning  dark 
when  dried.  Leaves  opposite  or  subopposite  dis- 
tally,  becoming  smaller  or  bract-like  on  distal 
stems,  sessile,  blades  linear  and  entire,  scabrous. 
Inflorescences  of  solitary  flowers  in  the  axils  of 
reduced  distal  leaves  (raceme-like  with  well-sep- 
arated flowers),  pedicels  often  longer  than  the  ca- 
lyx, bibracteolate.  Flowers  showy,  calyx  campan- 
ulate with  5  prominent  equal  lobes;  corolla  tubu- 
lar-campanulate,  slightly  bilabiate,  glabrous,  pink 
to  purple,  tube  straight  or  curved,  expanded  dis- 
tally  into  a  broad  throat,  2-lipped,  upper  lip  2- 
lobed,  lower  lip  3-lobed,  lobes  rounded,  subequal, 
spreading;  stamens  4,  of  2  unequal  pairs,  inserted 
near  the  mouth  of  the  corolla  tube,  shorter  than 
the  corolla,  anterior  filaments  longer  than  the  pos- 
terior, pilose  above,  anthers  2-thecous  with  un- 
equal divergent  thecae,  glabrous;  ovary  ovoid, 
glabrous,  2-locular,  style  longer  than  the  ovary. 


straight  and  slender,  stigmas  linear  and  lateral  on 
the  liguliform  style  apex.  Fruits  globose  capsules, 
chartaceous,  loculicidal,  style  base  persistent,  pla- 
centa persistent;  seeds  many,  linear,  exotesta  re- 
ticulate. 

Anisantherina  is  a  monotypic  Neotropical  genus 
related  to  African  genera  in  the  tribe  Buchnereae. 
Pennell  distinguished  Anisantherina  from  Agalinis 
by  its  bibracteolate  pedicels,  unequal  anther  thecae, 
and  narrow  oblong-linear  seeds.  Canne  (1980) 
showed  that  the  structure  and  ornamentation  of  An- 
isantherina seeds  are  also  distinctive.  This  species 
was  once  placed  in  Gerardia,  but  that  name  is  no 
longer  valid;  see  the  discussion  under  Agalinis. 

Anisantherina  hispidula  (Man.)  Pennell,  Mem. 
Torrey  Bot.  Club  16:  106.  1920.  Gerardia  his- 
pidula Mart.,  Nov.  Gen.  Sp.  PI.  3:  13.  1829. 
Agalinis  hispidula  (Mart.)  D'Arcy,  Ann.  Mis- 
souri Bot.  Card.  65:  4.  1978  (1979).  Figure  4. 

Annual  herbs,  30-50  cm  tall,  unbranched  or 
with  few  branches  arising  in  the  lower  half,  leafy 
internodes  0.5-1.5  mm  diam.,  with  few  short 
(0.2-0.4  mm)  stiff  hispidulous  hairs;  usually  dry- 
ing dark.  Leaves  opposite  or  subopposite  (rarely 
alternate  distally),  sessile  or  subsessile,  sometimes 
clasping  the  stem;  leaf  blades  8-80  mm  long, 
0.5-4  mm  wide,  linear,  entire,  scabrous  with  short 
(ca.  0.2  mm)  whitish, hairs  above  and  along  the 
margin,  2°  veins  obscure.  Inflorescences  of  soli- 
tary flowers  in  axils  of  reduced  distal  leaves,  ra- 
ceme-like with  8-14  flowers,  pedicels  10-35  mm 
long,  0.3-0.4  mm  diam.,  glabrous,  usually  with  a 
node  or  a  pair  of  small  (0-2  mm)  bracts  near  the 
middle.  Flowers  glabrous,  drying  dark,  calyx  tube 
4-6  mm  long,  3-5  mm  diam.,  campanulate-tu- 
bular,  abruptly  rounded  and  truncated  at  the  base, 
calyx  lobes  1.2-2.5  mm  long,  triangular  and 
acute;  corolla  10-15  mm  long,  campanulate,  pink 
to  light  purple  with  darker  spots  within,  tube  8- 
10  mm  long,  lobes  2-3  mm  long,  glabrous,  round- 
ed; filaments  2-3  mm  long,  anther  thecae  unequal, 
divergent.  Fruits  5-9  mm  long,  4-7  mm  diam., 
abruptly  rounded  at  apex  and  base  (short-cupu- 
late),  glabrous,  drying  black,  slightly  exserted  be- 
yond the  thin  persisting  calyx;  seeds  0.6-0.8  mm 
long,  0.2-0.3  mm  diam.,  linear,  dark  brown. 

Rarely  collected  plants  of  open  sunny  seasonal 
pools  and  moist  savannas  in  deciduous  or  ever- 
green forest  areas,  0-600  m  elevation.  We  have 
seen  only  two  collections  from  Costa  Rica,  both 
from  near  La  Cruz  in  northern  Guanacaste  (L.  D. 
Gomez  18965  &  J.  Gomez-Laurito  9097);  flow- 


BURGER:  FLORA  COSTARICENSIS 


17 


ering  and  fruiting  in  November.  The  species  rang- 
es from  southern  Mexico  and  Cuba  to  Brazil. 

Anisaniherina  hispidula  is  recognized  by  its 
short  wiry  habit,  parts  drying  dark,  linear  sca- 
brous leaves,  glabrous  distant  upright  pink  flow- 
ers, and  rounded  fruits.  It  is  confined  to  wet  hab- 
itats in  open  sunny  savannas.  See  the  key  and  dis- 
cussion under  Agalinis. 


Antirrhinum  Linnaeus 

REFERENCE — D.  Sutton,  A  Revision  of  the  Tribe 
Antirrhineae.  British  Museum  (Natural  History) 
&  Oxford  Univ.  Press,  1988. 

Herbs,  annual  or  perennial,  erect  or  procum- 
bent, usually  few-branched,  stems  terete,  often 
glandular  pubescent.  Leaves  opposite  to  subop- 
posite,  sometimes  alternate  on  distal  stems,  sessile 
or  short-petiolate,  leaf  blades  lanceolate  to  ovate, 
usually  narrow,  entire  to  denticulate.  Inflores- 
cences usually  showy  terminal  racemes  or  with 
solitary  flowers  in  axils  of  distal  bracts  or  reduced 
leaves,  pedicels  ebracteolate.  Flowers  with  calyx 
united  at  the  base,  sepals  5,  imbricate  in  bud;  co- 
rolla bilaterally  symmetric,  2-lipped  and  very  ir- 
regular, corolla  tube  gibbous  or  saccate  at  the  base 
(not  spurred),  broader  than  high,  upper  lip  erect 
and  2-lobed,  lower  lip  spreading  and  3-lobed, 
base  of  the  lower  lobe  forming  a  palate  and  press- 
ing against  the  front  of  the  throat  (closing  off  easy 
entrance  to  the  tube);  stamens  4,  of  2  unequal 
pairs,  included  in  the  corolla  tube,  filaments  slen- 
der and  slightly  dilated  at  the  apex,  anther  cells 
divergent,  staminode  absent;  ovary  ovoid,  2-loc- 
ular,  many-ovulate,  style  filiform,  longer  than  the 
ovary,  stigma  bilobed.  Fruits  ovoid  or  globose 
capsules,  opening  below  the  apex  by  pores  or  slits 
(septicidal);  seeds  many,  oblong,  truncated,  exo- 
testa  smooth  or  rugose  (without  wings). 

A  genus  of  about  42  species  from  the  western 
United  States,  northwestern  Mexico,  Europe,  and 
the  Mediterranean  region  of  Eurasia.  The  genus  is 
a  member  of  tribe  Antirrhineae,  with  Cymbalaha 
and  Linaria.  A  number  of  species  and  many  va- 
rieties have  been  developed  as  garden  ornamen- 
tals with  bright  red,  purple,  yellow,  or  white 
forms.  The  mouth  of  the  corolla  is  closed  by  the 
curved  palate  of  the  lower  lip,  which  is  held 
against  the  base  of  the  upper  lip.  Thus,  bees  must 
force  their  way  into  the  interior  of  the  corolla  to 
gain  access  to  nectar.  The  following  species  is 
commonly  cultivated  in  cool  highland  gardens 
throughout  Central  America. 


Antirrhinum  ma  jus  L.,  Sp.  PL  617.  1753.  Figure  8. 

Herbs,  erect,  0.4-1  m  tall,  leafy  stems  2-8  mm 
diam.,  glabrous  and  terete.  Leaves  opposite  or  ter- 
nate  below,  alternate  or  rarely  ternate  distally,  pet- 
ioles 1-14  mm  long  but  poorly  differentiated  from 
the  blade;  leaf  blades  2-9  cm  long,  3-16  mm 
wide,  lanceolate  to  ovate-lanceolate  or  linear, 
apex  acute,  margin  entire,  base  gradually  nar- 
rowed, drying  stiffly  chartaceous  and  greenish. 
Inflorescences  to  30  cm  long,  racemes  of  alter- 
nate flowers,  rachis  usually  densely  puberulent 
with  gland-tipped  hairs  ca.  0.3  mm  long,  bracts 
ovate,  2-10  mm  long,  acute,  pedicels  2-10  mm 
long.  Flowers  showy,  calyx  glandular  puberulent, 
sepals  5,  6-8  mm  long,  ovate  to  rounded;  corolla 
2.5-4  cm  long,  pink  to  red  or  purple,  externally 
glandular  hairy,  the  palate  closing  the  throat,  yel- 
low within.  Fruits  1-1.5  cm  long,  oblong  cap- 
sules with  unequal  valves,  opening  near  the  apex, 
glandular  pubescent  to  glabrous;  seeds  light 
brown,  reticulate-tuberculate. 

Antirrhinum  majus  is  a  native  of  the  Pyrenees 
of  northern  Spain  and  southern  France.  It  is  wide- 
ly cultivated  in  temperate  and  subtropical  regions. 
It  can  be  recognized  by  its  showy  racemes  and 
slightly  saccate  corolla  with  an  enlarged  palate 
that  "closes"  the  throat.  In  some  horticultural 
forms  the  throat  is  open  or  the  flowers  are  highly 
modified.  Common  names  are  "snapdragon," 
boca  de  leon,  and  boca  de  dragon. 


Bacopa  Aublet 

REFERENCE — F.  W.  Pennell,  Reconsideration  of 
the  Bacopa-Herpestis  problem  of  the  Scrophular- 
iaceae.  Proc.  Acad.  Nat.  Sci.  Philadelphia  98:  83- 
98.  1946. 

Herbs,  erect  to  decumbent  or  procumbent,  usu- 
ally growing  in  moist  soil  or  in  standing  water, 
stems  simple  or  profusely  branched,  glabrous  or 
pubescent,  often  glandular  punctate.  Leaves  op- 
posite, sessile  or  petiolate,  leaf  blades  often  slight- 
ly succulent,  entire  to  dentate  (or  dissected),  ve- 
nation pinnate  or  palmate,  the  minor  venation  usu- 
ally obscure.  Inflorescences  of  1-6  flowers  in  leaf 
axils,  less  often  the  distal  flowering  stems  race- 
mose or  spicate  (cymose,  paniculate),  pedicels 
short  or  absent,  bracteoles  present  or  absent  at  the 
base  of  the  calyx.  Flowers  usually  with  5-parted 
calyx,  sepals  subequal  to  strongly  unequal  with 
the  3  adaxial  sepals  usually  much  wider  than  the 


18 


FIELDIANA:  BOTANY 


abaxial,  imbricate  in  bud  (if  4-parted,  the  calyx 
tube  equaling  the  lobes);  corolla  tubular,  strongly 
or  weakly  bilabiate,  blue-violet  to  white,  tube  cy- 
lindriq,  the  lips  spreading,  with  3,  4,  or  5  lobes, 
the  upper  lip  exterior  in  bud  and  2-lobed  or  emar- 
ginate  (1-lobed),  the  lower  lip  2-  or  3-lobed;  sta- 
mens (5)  4  (2,  3),  usually  in  2  unequal  pairs,  in- 
serted on  the  upper  half  of  the  corolla  tube,  in- 
cluded, anthers  approximate  or  distant,  thecae 
contiguous,  parallel  or  divergent,  staminode  ab- 
sent; ovary  2-locular,  style  usually  straight,  stigma 
terminal,  bilobed  or  entire.  Fruits  dry  capsules, 
globose  to  ovoid,  bisulcate,  loculicidal,  often  sec- 
ondarily septicidal  into  4  valves;  seeds  many, 
small,  oblong,  longitudinally  reticulate. 

Bacopa  is  a  genus  of  about  50  to  60  species, 
widespread  in  warm  temperate  and  tropical  areas 


throughout  the  world.  It  is  most  diverse  in  South 
America.  The  species  are  varied  but  they  can  usu- 
ally be  recognized  by  their  small  opposite  leaves, 
strongly  unequal  sepals,  and  preference  for  wet 
habitats.  They  are  often  found  in  open  sunny  sites 
in  shallow  standing  water  of  seasonal  pools,  along 
the  edges  of  watercourses,  and  in  moist  savannas. 
Most  of  the  species  described  below  are  rarely 
collected  in  Costa  Rica,  perhaps  because  of  their 
seasonally  inundated  habitats.  Because  the  sepals 
differ  so  much  in  size,  they  may  appear  to  be 
bracts  enclosing  the  flower.  The  inner  sepals  are 
usually  much  narrower  than  the  outer.  The  brae - 
teoles  are  borne  at  the  apex  of  the  pedicel  when 
present.  The  synonymy  of  this  genus  is  very  large; 
D'Arcy  (1979,  p.  183)  provides  a  long  list  of  ge- 
neric synonyms. 


Key  to  the  Species  of  Bacopa 

la.  Flowers  sessile  or  subsessile,  pedicels  up  to  2  mm  long;  plants  erect;  rarely  collected  in  Costa  Rica 

2 

Ib.  Flowers  borne  on  conspicuous  pedicels  >  2  mm  long;  plants  erect,  prostrate  or  floating;  rare  or 

common 4 

2a.  Stems  densely  puberulent  with  hairs  to  1.3  mm  long;  [flowers  often  in  dense  verticels  of  3-12 

flowers/node  in  the  axils  of  leaves;  outer  calyx  lobes  3-4  mm  long]    B.  axillaris 

2b.  Stems  glabrous  or  minutely  puberulent,  the  hairs  <  0.5  mm  long 3 

3a.   Stems  puberulent;  fruits  with  smooth  surfaces;  outer  sepals  1-2  mm  long;  flowers  in  distal 

fascicles  separated  by  conspicuous  slender  internodes;  leaves  subentire    ...    B.  monnierioides 

3b.  Stems  glabrous;  fruits  with  pitted-reticulated  surfaces;  outer  sepals  2-3.5  mm  long;  flowers 

often  in  distal  crowded  fascicles  and  spike-like  with  obscure  internodes;  leaves  serrate 

B.  sessiliflora 

4a.  Plants  mostly  erect  with  few  to  many  distal  branches,  rooting  only  at  the  base,  stems  glabrous, 
leaves  serrate,  sessile  and  auriculate  at  the  base  (petiolate  in  B.  lacertosa),  opposing  leaves  not 

united  and  usually  without  an  interpetiolar  line   5 

4b.  Plants  usually  prostrate,  floating  or  creeping,  with  few  or  no  distal  branches,  rooting  at  base  and 
lower  nodes,  stems  glabrous  or  puberulent;  leaves  serrate  or  entire,  sessile  or  petiolate,  opposing 
leaves  slightly  united  across  the  stem  to  form  an  interpetiolar  line  or  ridge  or  the  node  pubescent 

7 

5a.  Outer  sepals  to  5  mm  long,  sepals  usually  narrowed  at  the  base,  rarely  covering  the  fruits,  the 
venation  not  conspicuously  raised;  distal  flowers  subtended  by  greatly  reduced  leaves  (ca.  8 

mm  long)  and  easily  seen  [distal  internodes  usually  longer  than  the  adjacent  leaves] 

B.  laxiflora 

5b.  Outer  sepals  to  8  mm  long,  rounded  at  the  base,  usually  covering  the  fruits,  venation  often 
conspicuous  on  the  outer  surface;  distal  flowers  subtended  by  normal-size  or  reduced  leaves, 

usually  easily  visible    6 

6a.  Outer  sepals  not  developing  a  lustrous  surface,  venation  only  slightly  elevated;  plants  to  40 

(-50)  cm  tall;  distal  stems  glabrous  or  glandular  puberulent    B.  bacopoides 

6b.  Outer  sepals  with  smooth  lustrous  surface  and  conspicuous  elevated  venation;  plants  to  70  cm 

tall;  distal  stems  glabrous  [Belize  to  Nicaragua]   B.  lacertosa 

7a.  Internodes  densely  villous  with  hairs  to  1 .5  mm  long;  leaves  often  suborbicular  [outer  sepals  4-6 

mm  long,  ciliolate;  commonly  collected  in  Costa  Rica]    B.  salzmanii 

7b.  Internodes  glabrous  or  with  thin  hairs  <  0.5  mm  long;  leaves  broadly  to  narrowly  obovate  (rarely 
suborbicular)  8 


BURGER:  FLORA  COSTARICENSIS 


19 


8a.   Braeteoles  2,  slender,  below  the  calyx;  outer  sepals  5-7  mm  long;  fruits  4-7  mm  long;  flowers 

usual  1\  I/node;  pedicels  to  30  mm  long;  internodes  glabrous B.  monnieri 

8b.  Braeteoles  none;  outer  sepals  2.5-4  mm  long;  fruits  2.3-4  mm  long;  flowers  1-4/node;  pedicels  to 

1 8  mm  long;  internodes  glabrous  or  puberulent 9 

9a.  Sepals  (4)  5,  unequal  and  united  only  at  the  base;  leaves  usually  cuneate  or  rounded  at  the  base, 

sessile;  widespread  B.  repens 

9b.  Sepals  4,  subequal  and  united  in  the  lower  half;  leaves  cuneate  with  a  narrowed  petiole-like  base; 

rarely  collected    B.  egensis 


Bacopa  axillaris  (Benth.)  Standl.,  J.  Wash.  Acad. 
Sci.  15:  460.  1925.  Herpestis  axillaris  Benth.  in 
DC,  Prodr.  10:  396.  1846.  Monniera  axillaris 
(Benth.)  O.  Kuntze,  Rev.  Gen.  PI.  463.  1891. 
Caconapea  axillaris  (Benth.)  Pennell,  Proc. 
Acad.  Nat.  Sci.  Philadelphia  72:  152.  1920.  Fig- 
ure 1. 

Erect  herbs  10-30  cm  tall,  aquatic  or  paludal, 
rooting  mostly  at  the  base,  leafy  stems  1.3-4  mm 
diam.,  spongy,  villous  with  thin  multicellular  hairs 
to  1 .3  mm  long.  Leaves  opposite,  sessile  with  op- 
posing leaves  slightly  united  at  the  base  and  form- 
ing a  line  or  ridge  across  the  stem  (clasping  the 
stem);  leaf  blades  1.2-4.8  cm  long,  2-12  mm 
wide,  oblanceolate  to  narrowly  obovate  or  nar- 
rowly elliptic-oblong,  apex  bluntly  obtuse  to 
acute,  distal  %  of  the  margin  serrate,  teeth  0.2-0.5 
mm  high,  1-2.3  mm  wide,  base  gradually  nar- 
rowed and  cuneate,  drying  yellowish  brown  or  ol- 
ive green,  glabrous,  conspicuously  pellucid  punc- 
tate beneath,  venation  pinnate.  Inflorescences 
verticellate,  of  dense  axillary  fascicles  with  3-12 
flowers/node,  pedicels  0.2-1.5  mm  long,  expand- 
ed at  the  apex  and  with  2  small  (0.4-0.7  mm) 
slender  bracteoles.  Flowers  glabrous  externally, 
outer  sepals  3-4  mm  long,  1 .5-3  mm  wide,  ovate 
with  rounded  to  obtuse  apex,  palmately  veined, 
punctate,  glabrous  or  ciliate;  corolla  3-4  mm 
long,  white,  slightly  exserted,  the  upper  lip  1- 
lobed;  stamens  4,  inserted  in  the  upper  half  of  the 
tube;  ovary  ca.  1  mm  long,  style  1.5-2  mm  long. 
Fruits  2-3  mm  long,  ca.  1.4  mm  diam.,  very  nar- 
rowly ovoid  or  conical,  4-valved;  seeds  0.5-0.6 
mm  long,  0.2-0.3  mm  wide,  oblong  or  somewhat 
curved,  brown,  reticulate  with  parallel  longitudi- 
nal ridges. 

Plants  of  swamps  and  the  muddy  edges  of 
standing  water  in  seasonally  deciduous  and  ever- 
green forest  areas,  0-500  m  elevation  (to  1 500  m 
in  Guatemala).  Rarely  found  north  of  Panama,  the 
species  has  been  collected  near  Bagaces  and  along 
the  Rio  Grande  de  Tarcoles  in  Costa  Rica;  it  flow- 
ers in  September.  The  species  ranges  from  Gua- 
temala to  Colombia. 


Bacopa  axillaris  is  recognized  by  the  pubescent 
stems,  sessile  oblanceolate  serrate  leaves,  verticil- 
late  flower  clusters  with  small  subsessile  flowers, 
and  wet  habitat.  The  outer  sepals  are  often  flat, 
translucent,  and  conspicuously  punctate. 

Bacopa  bacopoides  (Benth.)  Pulle,  Enum.  PI.  Su- 
rinam. 415.  1906.  Herpestis  bacopoides  Benth. 
in  DC.,  Prodr.  10:  399.  1846.  B.  bracteolata 
Pennell  ex  Standl.,  Contrib.  U.S.  Natl.  Herb.  27: 
336.  1927. 

Erect  herbs  to  50  cm  tall,  aquatic  or  paludal, 
leafy  stems  1.4-4.5  mm  diam.,  glabrous,  terete  or 
slightly  4-angled,  with  minute  sessile  glands,  node 
lacking  interpetiolar  lines.  Leaves  sessile,  clasp- 
ing the  stem,  opposing  leaves  not  united  at  the 
base;  leaf  blades  1.5-3.8  cm  long,  2.5-7  mm 
wide,  linear-oblanceolate  to  linear-oblong  or 
ovate-elliptic,  gradually  narrowed  in  the  lower 
half  but  slightly  expanded  and  subauriculate  at  the 
base,  apex  acute,  margin  serrate  with  short  (0.2- 
0.3  mm)  broad  (1-3  mm)  teeth,  drying  dark 
brown,  glabrous,  minutely  punctate,  venation  pin- 
nate, obscure.  Inflorescences  of  1-2  axillary 
flowers  (2-4  flowers/node),  pedicels  2-7  mm 
long,  0.2-0.3  mm  diam.,  minutely  papillate-pu- 
berulent  with  whitish  hairs,  slightly  expanded  at 
the  apex,  paired  bracteoles  1-2  mm  long  or  ab- 
sent, linear.  Flowers  glabrous  externally  or  the 
outer  sepals  sometimes  puberulent,  outer  3  sepals 
4-8  mm  long,  3-5  mm  wide,  broadly  ovate, 
rounded  but  obtuse  at  the  base,  with  raised  ve- 
nation; corolla  4-7  mm  long,  white,  stamens  4, 
inserted  near  the  middle  of  the  corolla  tube,  an- 
thers ca.  0.6  mm  long;  ovary  glabrous,  style  ca. 
2  mm  long.  Fruits  ca.  3  mm  long,  globose,  en- 
closed within  the  enlarged  (to  15  mm)  wing-like 
sepals. 

Partly  aquatic  plants  of  marshes  and  wet  areas 
at  low  elevations.  The  species  has  not  been  col- 
lected in  Costa  Rica  but  is  common  in  central 
Panama  and  flowers  in  December-January  in  Nic- 
aragua. This  species  ranges  from  Guatemala  to 
Brazil. 


20 


FIELDIANA:  BOTANY 


Bacopa  bacopoides  is  recognized  by  its  erect 
habit,  sessile  narrow  serrate  leaves,  one  or  two 
pedicellate  flowers  in  leaf  axils,  large  rounded 
outer  sepals  enclosing  the  fruits,  and  wet  habitat. 

Bacopa  egensis  (Poepp.)  Pennell,  Proc.  Acad. 
Nat.  Sci.  Philadelphia  98:  96.  1946.  Hydran- 
thelium  egense  Poepp.  in  Poepp.  &  Endl.,  Nov. 
Gen.  Sp.  PI.  3:  75,  tab.  287.  1845. 

Aquatic  or  terrestrial  herbs,  floating  or  pros- 
trate on  wet  soil,  with  short  erect  flowering  stems, 
rooting  at  proximal  nodes,  leafy  stems  terete,  pu- 
berulent  on  the  upper  surfaces.  Leaves  opposite, 
larger  in  aquatic  plants,  petioles  not  clearly  distin- 
guished from  the  cuneate  base;  leaf  blades  7-23 
mm  long,  3-14  mm  wide,  obovate  to  spatulate, 
rhombic  or  suborbicular,  apex  obtuse  to  rounded, 
margin  serrate  distal  ly,  base  cuneate  from  the 
middle  of  the  blade,  puberulent  beneath,  venation 
palmate  with  5-7  1°  veins.  Inflorescences  of  sol- 
itary axillary  flowers  (usually  2/node),  pedicels  3- 
6  mm  long,  slender,  glabrous  or  puberulent,  brac- 
teoles  absent.  Flowers  with  4-parted  calyx  2-4 
mm  long,  lobes  united  below  the  middle,  sube- 
qual,  obtuse,  subglabrous  or  the  outer  surface  mi- 
nutely puberulent;  corolla  3-5  mm  long,  funnel- 
form,  white,  with  3  unequal  lobes  rounded  distal- 
ly;  fertile  stamens  3;  ovary  with  many  ovules, 
stigma  bilobed.  Fruits  3-4  mm  long,  ovoid,  bi- 
valved  with  membranaceous  walls;  seeds  many, 
cylindric-curved,  rugulose. 

Rarely  collected  plants  of  standing  water  and 
wet  depressions  in  southern  Nicaragua  and  north- 
eastern Costa  Rica,  0-200  m  elevation.  Flowering 
and  fruiting  in  September.  The  species  is  also 
found  in  Colombia  and  Brazil. 

Bacopa  egensis  is  recognized  by  its  unusual 
palmately  veined  leaves  with  cuneate  or  attenuate 
base,  flowers  with  four  equal  sepals,  three-lobed 
white  corollas,  and  curved  cylindric  seeds.  The 
terrestrial  plants  of  this  species  have  smaller 
leaves  with  more  puberulence  than  their  aquatic 
conspecifics.  We  have  not  seen  Costa  Rican  ma- 
terial and  follow  the  report  in  Flora  of  Nicaragua. 

Bacopa  lacertosa  Standley  is  found  along  the 
Caribbean  coast,  from  Belize  to  Nicaragua.  It  has 
erect  stems  to  70  cm  tall,  lanceolate  leaves  to  70 
X  15  mm,  axillary  flowers  on  short  pedicels,  and 
broadly  ovate  outer  sepals  (to  8  mm  long)  with  a 
parchment-like  texture  and  lustrous  surface  on 
which  the  venation  is  conspicuously  elevated. 
Sutton  and  Hampshire  (Flora  of  Nicaragua,  1995) 
suggested  that  this  species  may  be  conspecific 


with  the  African  B.  decumbens  (Fernald)  F.  N. 
Williams. 

Bacopa  laxiflora  (Benth.)  Wettst.  ex  Edwall.  Bol. 
Commiss.  Geogr.  Estado  Sao  Paulo  13:  180. 
1897.  Herpestis  laxiflora  Benth.  in  DC,  Prodr. 
10:  396.  1846.  H.  auriculata  Robinson,  Proc. 
Am.  Acad.  Arts  26:  172.  1891.  B.  auriculata 
(Robinson)  Greenman,  Publ.  Field  Mus.  Nat. 
Hist.  Bot.  Sen  2:  262.  1907.  Caconapea  auri- 
culata (Robinson)  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  72:  150.  1920.  Mella  laxiflora 
(Benth.)  Pennell,  Notul.  Nat.  Acad.  Nat.  Sci. 
Philadelphia  46:  1.  1940. 

Erect  herbs  10-40  cm  high,  aquatic  or  paludal, 
rooting  only  at  the  base,  main  stem  simple  or  with 
few  lateral  branches,  leafy  stems  0.5-3  mm  diam.. 
glabrous,  4-angled  with  longitudinal  ridges. 
Leaves  sessile,  smaller  and  bract-like  at  distal 
flowering  nodes,  often  appearing  perfoliate  but 
the  opposing  leaves  not  united  across  the  stem; 
leaf  blades  6-30  mm  long,  2-10  mm  wide,  lan- 
ceolate to  lanceolate-oblong  or  ovate-lanceolate, 
apex  acute,  margin  serrate,  teeth  ca.  0.3  mm  high 
and  1-2  mm  wide,  base  rounded  to  auriculate. 
drying  grayish  green,  glabrous  and  punctate,  ve- 
nation pinnate.  Inflorescences  of  solitary  axillary 
flowers  (1 -2/node)  or  racemose  and  terminal  (re- 
sembling panicles  in  some  collections),  pedicels 
3-8  mm  long,  minutely  papillate  puberulent, 
ebracteolate  at  base  of  calyx  or  with  2  slender 
bracteoles  1-1.5  mm  long.  Flowers  glabrous,  out- 
er sepals  3-5  mm  long,  1-3.3  mm  wide,  ovate, 
usually  narrowed  at  the  base,  acute,  palmately 
veined,  inner  sepals  linear;  corolla  7-10  mm 
long,  lilac  to  purple,  bilabiate,  tube  ca.  5  X  1.7 
mm,  lobes  2-3  mm  long;  stamens  4,  inserted  on 
the  upper  half  of  the  tube,  anthers  0.5  mm  long; 
style  2.5  mm  long.  Fruits  2.5-4  mm  long,  subgl- 
obose,  surface  slightly  reticulate  with  minute  pits; 
seeds  ca.  0.4  X  0.2  mm,  oblong  with  1  or  2  trun- 
cated ends,  longitudinally  ridged/reticulate, 
brown. 

Plants  of  wet  sites  and  shallow  standing  water, 
in  lowland  deciduous  and  evergreen  formations, 
0-300  m.  The  species  has  been  collected  near  La 
Cruz,  Guanacaste  Province;  it  flowers  in  Decem- 
ber-January. The  species  ranges  from  Mexico  to 
Brazil  but  is  rarely  collected  in  Central  America. 

Bacopa  laxiflora  is  recognized  by  its  short  erect 
habit,  wet  habitat,  sessile  serrate  leaves  appearing 
to  clasp  the  stem,  solitary  axillary  flowers  form- 
ing distal  racemes,  and  globose  or  ovoid  fruits. 


BURGER:  FLORA  COSTARICENSIS 


21 


Stems  and  leaves  are  usually  glabrous,  but  the 
pedicels  arc  minutely  papillate  puberulent. 

Bacopa  monnieri  (L.)  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  98:  94.  1946.  Lysimachia 
monnieri  L.,  Sp.  PI.  Cent.  2:  tab.  9.  1756.  Fig- 
ure 1. 

Prostrate  to  procumbent  herbs  to  30  cm  tall, 
rooting  from  the  lower  nodes,  leafy  stems  0.7-2.5 
mm  diam.,  glabrous,  nodes  marked  by  interpetio- 
lar  lines.  Leaves  slightly  succulent,  subsessile  or 
with  poorly  defined  petioles  0.5-2  mm  long, 
clasping  the  stem  at  the  base;  leaf  blades  3-18 
mm  long,  1 .5-7  mm  wide,  obovate  to  oblong-ob- 
ovate,  oblanceolate,  or  spatulate,  apex  rounded, 
margin  entire,  base  gradually  narrowed  and  cu- 
neate,  drying  yellowish  green,  glabrous,  often 
punctate,  venation  pinnate  but  obscure  (some- 
times appearing  tripliveined).  Inflorescences  of 
solitary  axillary  flowers,  usually  with  only  1  flow- 
er/node, pedicels  7-30  mm  long,  0.3-0.6  mm 
diam.,  glabrous,  bracteoles  2,  opposite,  1.5-3  mm 
long,  0.4-0.7  mm  wide,  near  the  apex  of  the  ped- 
icel and  resembling  the  larger  sepals  in  texture. 
Flowers  glabrous,  the  2  outer  sepals  5-7  mm 
long,  2-3  mm  wide,  3  inner  sepals  narrower, 
acute;  corolla  6-10  mm  long,  white  to  lavender, 
blue  or  pale  purple,  lobes  rotate  and  essentially 
regular,  ca.  2  cm  wide,  tube  3-5  mm  long;  sta- 
mens 4,  filaments  arising  near  the  apex  of  the 
tube,  glabrous,  anthers  ca.  1 .5  mm  long;  ovary  ca. 
4  mm  long,  ovate-oblong,  style  ca.  4  mm  long, 
stigma  flattened,  ca.  0.8  mm  wide.  Fruits  4-7  mm 
long,  ovoid,  enclosed  within  the  persisting  sepals; 
seeds  ca.  0.5  mm  long,  reddish  brown,  longitu- 
dinally reticulate. 

Plants  of  open  sunny  sites  in  marshes,  sandy 
stream  edges,  and  standing  water,  often  creeping 
on  wet  mud  and  tolerant  of  some  salinity,  0-1000 
m  elevation.  The  species  has  rarely  been  collected 
in  Costa  Rica,  where  it  is  usually  found  near  the 
coasts.  It  probably  flowers  throughout  the  year. 
We  have  seen  a  single  sterile  collection  (Crow 
9465  INBIO)  from  820  m  in  Alajuela.  The  species 
ranges  from  the  southeastern  United  States  to  Ar- 
gentina. 

Bacopa  monnieri  is  recognized  by  its  small  en- 
tire obovate  or  spatulate  leaves,  glabrous  parts, 
narrow  bracteoles  beneath  the  sepals,  somewhat 
larger  flowers,  and  ability  to  survive  in  slightly 
salty  water.  The  species  has  been  called  verdolaga 
in  Nicaragua. 


Bacopa  monnierioides  (Cham.)  Robinson,  Proc. 
Am.  Acad.  Arts  44:  614.  1909.  Ranaria  mon- 
nierioides Cham.,  Linnaea  8:  31.  1833.  Herpes- 
tis  ranaria  Benth.  in  J.  D.  Hook.,  Companion 
Bot.  Mag.  2:  57.  1836,  based  on  R.  monnierioi- 
des Cham.  B.  ranaria  (Benth.)  Chod.  &  Hassl., 
Bull.  Herb.  Boissier  ser.  2,  4:  288.  1904.  Ca- 
conapea  parviflora  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  72:  152.  1920.  B.  parviflora 
(Pennell)  Pennell  ex  Standl.,  Contrib.  U.S.  Natl. 
Herb.  27:  336.  1928.  B.  parviflora  (Pennell) 
Standl.  ex  L.  O.  Williams,  Fieldiana  Bot.  34: 
118.  1972. 

Erect  herbs  to  40  cm  tall,  aquatic  or  in  wet  soil, 
with  much  distal  branching,  rooting  mostly  at  the 
base,  leafy  stems  0.5-2.5  mm  diam.,  minutely  pu- 
berulent with  slender  crooked  hairs  0.1-0.4  mm 
long  on  upper  surfaces.  Leaves  sessile,  opposing 
leaves  not  united  across  the  stem  but  sometimes 
producing  a  line  across  the  stem,  distal  leaves 
conspicuously  smaller  than  leaves  of  the  main 
stem;  leaf  blades  8-30  mm  long,  2-7  mm  wide, 
oblong-lanceolate  to  narrowly  oblong,  apex  ob- 
tuse, margin  subentire  or  obscurely  dentate  with 
teeth  1-2  mm  wide,  drying  yellowish  brown  or 
dark  grayish,  glabrous,  glandular  punctate  on  both 
surfaces,  venation  subpalmate  or  tripliveined.  In- 
florescences mostly  fasciculate,  of  (1)2-4  axillary 
flowers,  2-8  flowers/node,  pedicels  ca.  0.5  mm 
long,  bracteoles  minute  (0.3  mm)  at  apex  of  ped- 
icel. Flowers  glabrous,  outer  sepals  1-2  mm  long, 
0.5-1  mm  wide,  narrowly  ovate-oblong,  with  an 
unusual  glandular-pitted  surface;  corolla  1.5-2.5 
mm  long,  tubular,  white  or  bluish  white,  upper 
lobe  entire,  lower  lip  3-lobed;  stamens  4,  inserted 
near  the  apex  of  the  tube;  ovary  glabrous.  Fruits 
ca.  1 .5  mm  long,  smooth,  enclosed  within  the  stiff 
persisting  sepals  (to  2.5  mm  long);  seeds  0.6-0.7 
mm  long,  0.2-0.3  mm  thick,  oblong-ellipsoid, 
longitudinally  ridged,  reddish  brown. 

Plants  of  inundated  areas,  the  margins  of  water 
bodies,  and  wet  savannas,  0-1500  m  in  Central 
America.  In  Costa  Rica,  this  species  has  been  col- 
lected only  in  lowland  Guanacaste.  The  species 
ranges  from  Guatemala  to  Paraguay. 

Bacopa  monnierioides  is  recognized  by  its  erect 
branching  habit,  sessile  lanceolate-oblong  leaves 
with  subserrate  margins,  small  fasciculate  flowers, 
and  smooth  fruit.  The  glandular  pit  at  the  base  of 
the  outer  sepal  is  unusual  but  difficult  to  see. 
Compare  B.  axillaris  and  B.  sessiliflora. 


22 


FIELDIANA:  BOTANY 


Bacopa  repens  (Sw.)  Wettst.  in  Engl.  &  Prantl, 
Nat.  Pflanzenfam.  4(3b):  76.  1895.  Gratiola  re- 
pens  Sw.,  Prodr.  14.  1788.  Herpestis  repens 
(Sw.)  Schldl.  &  Cham.,  Linnaea  5:  107.  1830. 
Macuillamia  limosa  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  72:  158.  1920.  Bacopa  limosa 
(Pennell)  Standl.,  Contrib.  U.S.  Natl.  Herb.  27: 
336.  1928.  M.  repens  (Sw.)  Pennell,  Acad.  Nat. 
Sci.  Philadelphia  Monogr.  1:  60.  1935.  B.  cur- 
tipes  Standl.  &  L.  O.  Williams.  Ceiba  3:  129. 
1952.  Figure  1. 

Herbs,  aquatic  or  terrestrial,  floating  to  pros- 
trate or  repent,  to  40  cm  long,  rooting  from  lower 
nodes,  stems  0.4-3  mm  diam.,  glabrous  or  with 
short  (0.1-0.3  mm)  thin  hairs  distally.  Leaves  ses- 
sile, bases  of  opposing  leaves  partly  united  and 
obscuring  the  node;  leaf  blades  6-20(-30)  mm 
long,  3-15(-20)  mm  wide,  obovate  to  obovate- 
oblong  or  suborbicular,  apex  rounded  or  obtuse, 
margin  entire,  base  cuneate,  drying  yellowish 
green  to  dark  green  or  blackish,  glabrous  except 
near  the  base,  venation  palmate  with  3-11  major 
veins.  Inflorescences  of  axillary  flowers,  2-4 
flowers/node,  pedicels  6-18  mm  long,  0.2-0.3 
mm  diam.,  glabrous  or  minutely  puberulent,  brac- 
teoles  absent  or  minute.  Flowers  with  4  or  5  se- 
pals, outer  sepals  2.5-4  mm  long,  0.7-1.5  mm 
wide,  narrowly  oblong,  rounded  to  obtuse  at  the 
apex,  margin  often  ciliolate,  inner  sepals  narrow- 
er; corolla  3-4  mm  long,  white  or  pale  violet, 
campanulate  and  regular  or  nearly  so,  throat 
sometimes  marked  with  yellow,  2-lipped,  the  up- 
per lip  2-lobed;  stamens  4,  filaments  inserted  near 
the  apex  of  the  tube,  anthers  ca.  0.7  mm  long; 
ovary  ca.  1.3  mm  long,  style  1-3  mm  long,  slen- 
der, stigma  capitate,  slightly  2-lobed.  Fruits  2.3- 
4  mm  long,  ca.  2  mm  diam.,  glabrous,  enclosed 
within  the  persisting  (3-4  mm  long)  sepals;  seeds 
ca.  0.4  mm  long,  oblong,  dull  whitish  to  brown, 
surface  reticulate. 

Aquatic  plants  of  lake  edges  and  marshes,  in 
both  evergreen  and  deciduous  forest  areas,  0- 
1000  m  elevation.  Rarely  collected  in  Costa  Rica, 
this  species  is  found  in  the  Caribbean  lowlands 
and  on  the  Pacific  coast  in  seasonal  ponds  at  Ba- 
gaces  and  Palo  Verde  National  Park.  Flowering  in 
August-October.  The  species  ranges  from  south- 
ern Mexico  and  the  West  Indies  to  Argentina. 

Bacopa  repens  is  recognized  by  its  aquatic  hab- 
itat, glabrous  or  sparsely  puberulent  stems,  ob- 
ovate sessile  leaves  slightly  united  (across  the 
stem)  at  the  base,  and  small  flowers.  The  distal 
leaves  and  stems  are  often  floating  and  can  be- 
come a  floating  mat.  This  species  may  be  vege- 


tatively  very  similar  to  B.  monnieri,  but  that  spe- 
cies has  larger  flowers  and  fruits  and  lacks  brac- 
teoles.  Compare  B.  salzmannii,  which  occurs  in 
similar  habitats  but  has  villous  stems,  larger  flow- 
ers, and  separate  leaf  bases. 

Two  collections  from  water  1 .5  m  deep  at  Palo 
Verde  National  Park  (G.  Crow  5977  &  60603)  are 
provisionally  placed  here.  These  collections  have 
larger  (20-40  mm)  leaves,  longer  fruits  (3-5 
mm),  and  linear-oblong  seeds  to  0.7  mm  long. 
They  conform  to  the  description  of  B.  valerii 
Standl.  &  L.  O.  Williams  (Ceiba  1:  163,  1950) 
based  on  material  from  20  m  elevation  in  we'stern 
Honduras.  Although  quite  distinctive,  it  seems 
likely  that  all  these  plants  represent  no  more  than 
an  unusually  robust  form  of  B.  repens. 

Bacopa  salzmannii  (Benth.)  Wettst.  ex  Edwall, 
Bol.  Commiss.  Geogr.  Estado  Sao  Paulo  13: 
176,  181.  \S91.ScrophulariaprocumbensVe\\., 
Fl.  tl nun n.  Ic.  6,  tab.  85.  1827,  non  B.  procum- 
bens  (Miller)  Greenman.  Herpestis  salzmannii 
Benth.  in  J.  D.  Hook.,  Companion  Bot.  Mag.  2: 
58.  1836.  B.  salzmannii  (Benth.)  Chod.  & 
Hassl.  Bull.  Herb.  Boissier  Ser  2,4:  290.  1904. 
Monocardia  humilis  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  72:  157.  1920.  M.  violacea 
Pennell,  loc.  cit.  156.  1920.  B.  humilis  (Pennell) 
Standl.,  J.  Wash.  Acad.  Sci.  15:  460.  1925.  B. 
violacea  (Pennell)  Standl.,  loc.  cit.  460.  1925. 
Herpestis  ciliata  Pennell,  Notul.  Nat.  Acad.  Sci. 
Philadelphia  46:  2.  1940.  Figure  1. 

Herbs,  aquatic  or  terrestrial,  prostrate  or  de- 
cumbent, sometimes  floating  or  forming  mats, 
rooting  from  lower  nodes,  stems  succulent,  1-2 
mm  diam.,  densely  hirsute  with  thin  multicellular 
hairs  0.7-1.5  mm  long.  Leaves  sessile  and  decus- 
sate, opposing  leaves  not  united  at  the  base,  often 
subtended  by  a  tuft  of  hairs;  leaf  blades  6-20  mm 
long,  5-18  mm  wide,  ovate  to  broadly  ovate-el- 
liptic or  ovate-orbicular,  apex  rounded  or  bluntly 
obtuse,  sometimes  slightly  notched,  margin  entire, 
base  rounded  and  truncate  to  subcordate  or  auric- 
ulate,  drying  greenish  to  yellowish,  glabrous  or 
hirsute  near  the  base  beneath,  venation  palmate 
with  3-7  poorly  defined  primary  veins.  Inflores- 
cences of  solitary  axillary  flowers,  1-2/node,  ped- 
icels 8-23  mm  long,  0.2-0.4  mm  diam.,  hirsutu- 
lous  with  slender  hairs  0.2-0.5  mm  long,  ebrac- 
teolate  (but  the  outer  sepals  resembling  bracts). 
Flowers  with  strongly  unequal  sepals,  outer  3  se- 
pals 4-6  mm  long,  2.5-5  mm  wide,  ovate  with 
truncated  or  subcordate  bases  and  resembling  the 
leaves,  ciliolate  along  the  margin,  inner  sepals 


BURGER:  FLORA  COSTARICENSIS 


23 


narrowly  triangular;  corolla  6-9  mm  long,  pale 
blue  to  lavender  or  violet  (white),  slightly  exsert- 
ed  beyond  the  sepals,  4-lobed  with  the  upper  lobe 
emerginate;  stamens  4,  included,  filaments  1-2 
mm  long,  glabrous,  larger  anthers  1.2-1.5  mm 
long;  ovary  ca.  1  mm  long,  narrowly  ovoid,  gla- 
brous, style  2-5  mm  long,  stigma  0.3  mm  wide. 
Fruits  ca.  3  mm  long  and  1  mm  wide,  narrowly 
ovoid-oblong;  seeds  many,  small,  reddish  brown, 
reticulate. 

Plants  of  shallow  water  and  moist  open  sunny 
sites  at  the  edges  of  water  in  both  evergreen  and 
deciduous  forest  areas,  40-1700  m  elevation.  This 
is  a  common  species  of  marshes,  swamps,  and  wet 
depressions,  growing  both  as  a  partly  submerged 
aquatic,  prostrate  and  mat-forming  on  wet  mud, 
or  erect  among  other  wetland  plants,  flowering 
throughout  the  year.  This  species  ranges  from 
Mexico  and  the  West  Indies  to  southern  Brazil. 

Bacopa  salzmannii  is  the  most  commonly  col- 
lected species  of  Bacopa  in  Costa  Rica.  It  is  rec- 
ognized by  its  often  aquatic  or  repent  habit  in  wet 
sites,  the  villous  stems,  sessile  rounded  leaves, 
and  leaf-like  (bract-like)  outer  sepals.  The  round- 
ed bases  of  the  leaf  blades  may  obscure  the  stem 
but  they  are  not  joined  (interpetiolar,  cf.  B.  re- 
pens)  nor  are  they  decurrent  on  the  stem. 

Bacopa  sessiliflora  (Benth.)  Pulle,  Enum.  PI.  Su- 
rinam 415.  1906.  Herpestis  sessiliflora  Benth. 
i/i  J.  D.  Hook.,  Companion  Bot.  Mag.  2:  58. 
1836.  B.  sessiliflora  (Benth.)  Edwall,  Bol.  Com- 
miss.  Geogr.  Estado  Sao  Paulo  13:  176,  181. 
1897,  incomplete  combination  fide  D'Arcy 
1979.  Caconapea  conferta  Pennell,  Proc.  Acad. 
Nat.  Sci.  Philadelphia  72:  153.  1920.  C.  sessi- 
liflora (Benth.)  Pennell,  loc.  cit.  75:  11.  1923. 

Erect  herbs  to  40  cm  tall,  with  short  lateral 
branches,  rooting  only  at  the  base,  leafy  stems 
0.8-3  mm  diam.,  glabrous,  with  2  opposite  lon- 
gitudinal ridges.  Leaves  sessile,  separate  at  the 
base  and  not  forming  an  interpetiolar  line,  decur- 
rent on  the  stem,  distal  leaves  smaller  (5  mm)  and 
bract-like;  leaf  blades  10-35  mm  long,  2-5  mm 
wide,  oblanceolate  to  linear-oblanceolate  or  nar- 
rowly elliptic-oblanceolate,  apex  acute,  distal  half 
of  the  margin  with  prominent  teeth  ca.  0.5  mm 
high,  1-2  mm  long,  cuneate  to  the  base,  drying 
yellowish  brown,  glabrous,  punctate  on  both  sur- 
faces, venation  pinnate.  Inflorescences  fasciculate 
in  distal  spiciform  arrangements  or  1-4  flowers  in 
leaf  axils,  distal  spikes  1-3  cm  long,  ca.  1  cm 
diam.,  flowers  sessile  or  subsessile  on  pedicels 
less  than  1.5  mm  long,  bracteoles  1-2  mm  long 


at  apex  of  pedicel  or  absent.  Flowers  glabrous 
externally,  calyx  united  at  base,  outer  calyx  lobes 
3-3.5  mm  long,  1.3-2  mm  wide,  ovate,  thin  with 
3  prominent  parallel  veins;  corolla  3-4  mm  long, 
tubular-campanulate,  exserted,  bluish  or  white, 
lobes  ca.  0.5  mm  long;  stamens  4,  attached  near 
middle  of  tube;  ovary  ca.  1 .5  mm  long,  style  ca. 
1  mm  long.  Fruits  3-4  mm  long,  1.5-2  mm 
diam.,  narrowly  ovoid,  surface  with  minute  (0.1 
mm)  capitate  hairs;  seeds  ca.  0.3  X  0.2  mm,  rect- 
angular with  truncated  ends,  brown,  longitudinal- 
ly reticulate/ridged. 

Rarely  collected  plants  of  open  sunny  sites  in 
wet  marshes  and  depressions.  The  plants  are  tol- 
erant of  brackish  water  and  can  be  found  near  the 
seashore.  The  species  has  not  been  collected  in 
Costa  Rica  but  is  known  from  Bluefields,  Nica- 
ragua, and  is  found  in  central  Panama.  The  spe- 
cies ranges  from  Guatemala  and  Belize  to  Ecua- 
dor and  the  West  Indies. 

Bacopa  sessiliflora  is  distinguished  by  its  erect 
habit,  general  lack  of  pubescence,  sessile  oblan- 
ceolate leaves  serrulate  along  their  distal  margins, 
and  sessile  flowers  often  in  axillary  fascicles  that 
may  be  arranged  in  terminal  spike-like  inflores- 
cences. Compare  B.  monnierioides. 


Benjaminia  Martius 

REFERENCE — L.  B.  Smith  &  J.  M.  Pires,  An 
evaluation  of  Benjaminia  Martius  ex  Benjamin.  J. 
Wash.  Acad.  Sci.  46:  86.  1956. 

Herbs,  aquatic  and  submerged,  rooting  at  lower 
nodes,  glabrous  or  puberulent,  glandular  punctate 
on  vegetative  parts.  Leaves  verticillate,  united  at 
the  base,  petiolate,  the  blades  pinnatifid  with  slen- 
der linear  segments.  Inflorescences  of  solitary  ax- 
illary flowers,  bracts  absent,  pedicels  elongating 
slightly  in  fruit,  bracteoles  absent  at  the  base  of 
the  calyx.  Flowers  with  calyx  lobes  united  near 
the  base,  calyx  lobes  (sepals)  5  and  subequal,  nar- 
row, valvate  in  bud;  corolla  tubular  and  2-lipped, 
upper  lip  slightly  2-lipped,  lower  lip  3-lobed;  sta- 
mens 4,  of  2  unequal  pairs,  included,  anthers 
equal,  thecae  similar;  ovary  subtended  by  a  ring 
of  filaments,  2-locular,  ovules  many,  style  simple, 
stigma  flat  and  slightly  curved.  Fruits  thin-walled 
capsules,  loculicidal,  surface  smooth;  seeds  many, 
oblong,  surface  longitudinally  reticulate. 

A  monotypic  genus  of  unusual  aquatic  plants, 
this  taxon  is  closely  related  to  Bacopa  but  differs 
in  having  nearly  equal  sepals,  unusual  pinnatifid 
leaves,  and  ovary  subtended  by  staminodial  fila- 


24 


FIELDIANA:  BOTANY 


ments  (present  in  some  Bacopa  species).  This  ge- 
nus has  been  confused  with  the  Old  World  Lim- 
nophila,  but  Benjaminia  is  distinct  because  of  its 
estipitate  anther-thecae  and  its  two-lobed  stigma. 
Also,  Limnophila  species  usually  have  emergent 
leaves  that  are  broad  and  dentate,  whereas  emer- 
gent leaves  are  never  found  in  Benjaminia.  For  a 
short  discussion  of  the  complex  nomenclature  of 
the  genus,  see  D'Arcy  (1979,  p.  194). 

Benjaminia  reflexa  (Benth.)  D'Arcy,  Ann.  Mis- 
souri Bot.  Card.  66:  194.  1979.  Herpestis  re- 
flexa Benth.  in  DC.,  Prodr.  10:  399.  1846.  Be. 
utriculariaeformis  Mart.,  Fl.  Bras.  10:  256. 
1847.  Quinquelobulus  utriculariaeoides  Benj., 
Linnaea  20:  316.  1847.  Monnierea  reflexa 
(Benth.)  Kuntze,  Rev.  Gen.  PI.  2:  463.  1891. 
Bacopa  reflexa  (Benth.)  Edwall,  Bol.  Commiss. 
Geogr.  Estado  Sao  Paulo  13:  176.  1897.  Naia- 
dothrix  longipes  Pennell,  Mem.  Torrey  Bot. 
Club  16:  105.  1920.  Bacopa  naias  Standl.,  Field 
Mus.  Bot.  Ser.  11:  141.  1932.  Limnophila  cos- 
taricensis  Suesseng.,  Bot.  Jahrb.  Syst.  72:  284. 
1942.  L.  costaricensis  forma  aquatica  Sues- 
seng., loc.  cit.  L.  costaricensis  forma  semiter- 
restris  Suesseng.,  loc.  cit.  Figure  1. 

Submerged  herbs,  only  the  flowers  and  fruits 
extending  above  the  water  surface,  branched, 
rooting  at  lower  nodes,  internodes  2-60  mm  long, 
0.5-2  mm  diam.,  glabrous  or  minutely  and  sparse- 
ly puberulent,  vegetative  parts  glandular  punctate. 
Leaves  in  verticels  of  6  or  8/node  (rarely  2  or  4), 
glabrous  or  sparsely  puberulent,  petioles  2-6  mm 
long  (to  the  first  pinna),  0.2-0.4  mm  wide;  leaf 
blades  4-35  mm  long,  5-25  mm  wide,  with  slen- 
der filiform  pinnate  lobes,  lobes  4-15/side  and  in 
a  single  plane,  central  rachis  0.2-0.4  mm  wide, 
lobes  0.05-0.15  mm  wide,  often  with  small  (0.1- 
0.2  mm)  punctate  glands.  Inflorescences  of  soli- 
tary axillary  flowers,  1  flower/node,  pedicels  5- 
9(-18)  mm  long,  0.2-0.4  mm  diam.,  glabrous  or 
sparsely  and  minutely  puberulent.  Flowers  with 
sepals  2-4  mm  long,  0.4-0.7  mm  wide,  linear- 
lanceolate  to  linear-oblong,  apex  usually  slightly 
rounded;  corolla  4-6  mm  long,  bluish  white  or 
purple,  yellowish  within  the  throat;  anthers  ver- 
satile and  similar;  ovary  ca.  2  X  1  mm,  narrowly 
ovoid,  style  ca.  1  mm  long,  slender.  Fruits  2-3 
mm  long,  narrowly  ovoid,  smooth,  with  a  persist- 
ing style  ca.  1  mm  long;  seeds  ca.  0.6  mm  long, 
oblong-fusiform,  reticulate. 

Submerged  aquatic  plants  of  shallow  ponds  and 
lakes,  0-800  m  elevation.  Only  three  collections 
from  Costa  Rica  have  been  seen,  all  from  the 


General  Valley  (near.  Buenos  Aires  and  between 
San  Isidro  and  Rivas),  flowering  and  fruiting  in 
November  (Crow  6176  &  6239),  December  (Nic- 
aragua), and  February  (Kupper  597).  The  species 
(in  a  wide  sense)  ranges  from  Mexico  and  Cuba 
to  Brazil. 

Benjaminia  reflexa  is  an  unusual  aquatic  plant 
distinguished  by  being  almost  entirely  submerged, 
having  whorls  of  feather-like  leaves  with  slender 
pinnate  lobes,  solitary  little  flowers  with  five  near- 
ly equal  narrow  sepals,  and  sympetalous  two 
lipped  corollas.  The  flowers  and  fruits  are  borne 
above  the  water  surface  on  stiff  pedicels.  The 
leaves  can  also  be  interpreted  as  being  opposite, 
with  each  leaf  having  three  or  four  primary  axes 
and  each  of  these  having  deeply  divided  pinnatifid 
divisions.  These  plants  resemble  the  submerged 
portions  of  species  of  Cabomba  (Nymphaeaceae) 
as  well  as  aquatic  Utricularia  species  (Lentibu- 
lariaceae).  The  only  similar  Neotropical  species  of 
Scrophulariaceae  is  Bacopa  myriophylloides 
(Benth.)  Pennell,  which  has  palmately  dissected 
leaves  and  differing  flowers.  This  species  was 
treated  as  Bacopa  naias  in  the  Flora  of  Guate- 
mala. Suessenguth's  Limnophila  costaricensis 
lacks  a  species  description  but  has  Latin  descrip- 
tions of  two  forms,  all  based  on  a  single  collection 
number  (Kupper  597,  fragments  at  F). 


Buchnera  Linnaeus 

REFERENCE — D.  Philcox,  Revision  of  the  New 
World  species  of  Buchnera  L.  Kew  Bull.  18:  275- 
316.  1965. 

Herbs,  annual  (in  ours)  or  perennial,  hemipar- 
asitic,  erect,  simple  or  branched,  hairs  with  a 
broad  base  and  stiff  straight  tip,  often  scabrous, 
drying  dark.  Leaves  opposite  or  subopposite, 
sometimes  alternate  distally,  sessile  to  short-peti- 
olate,  leaf  blades  narrow,  dentate  or  entire,  mar- 
gins often  revolute,  usually  scabrous,  venation 
pinnate  or  palmate,  with  1  or  3  (5)  major  veins. 
Inflorescences  terminal  compact  or  lax  spikes, 
flowers  sessile  or  subsessile,  subtended  by  a  bract, 
with  2  slender  lateral  bracteoles  beneath  the  calyx. 
Flowers  with  a  tubular  calyx,  slightly  enlarging 
in  fruit,  with  5  or  10  prominent  longitudinal  veins, 
lobes  4  or  5,  shorter  than  the  tube,  acute;  corolla 
radially  symmetric  and  usually  salverform,  tube 
longer  than  the  calyx,  cylindric  and  straight  or 
slightly  curved,  blue  to  white  or  purple,  glabrous 
(in  ours)  or  puberulent  externally,  lobes  5  and 


BURGER:  FLORA  COSTARICENSIS 


25 


subequal,  rotate-spreading,  shorter  than  the  tube, 
apex  of  the  tube  often  with  moniliform  hairs;  sta- 
mens 4,  of  2  unequal  pairs,  inserted  in  the  prox- 
imal hall  of  the  tube,  subsessile  or  with  short  fil- 
aments, anthers  ovoid,  versatile,  1-thecous;  ovary 
ellipsoid  to  ovoid,  2-locular  with  many  ovules, 
style  slender  and  included,  stigma  clavate.  Fruits 
dry  capsules  partly  enclosed  by  the  persisting  ca- 
lyx, splitting  loculicidal  into  2  equal  parts;  seeds 
many,  oblong  or  ellipsoid,  curved  or  angled,  lon- 
gitudinally reticulate. 

A  genus  of  ca.  100  species  in  tropical  and  tem- 


perate climates  throughout  the  world.  Most  of  the 
species  are  found  in  the  Old  World  tropics;  there 
are  about  1 6  species  in  the  New  World.  The  genus 
is  distinguished  by  its  slender  few-branched  habit, 
stiff,  narrow,  usually  scabrous  leaves,  spicate  in- 
florescences, tubular  calyx,  and  one-thecous  an- 
thers, all  parts  drying  dark  or  black.  All  species 
are  believed  to  be  hemiparasites,  attaching  by 
haustoria  to  the  roots  of  the  host  plants.  The  genus 
is  placed  in  the  tribe  Buchnerae,  along  with  many 
Old  World  genera.  Its  closest  Neotropical  relatives 
are  Alectra  and  Escobedia. 


Key  to  the  Species  of  Buchnera 

la.  Stems  hispid-scabrid,  rough  to  the  touch;  larger  leaves  with  prominent  teeth;  calyx  pubescent,  be- 
coming slightly  thickened  in  fruit,  but  the  areas  between  the  longitudinal  veins  usually  remaining 
flat  and  the  minor  venation  not  visible B.  pusilla 

Ib.  Stems  usually  smooth  to  the  touch  at  distal  internodes;  larger  distal  leaves  entire  and  linear;  calyx 
glabrous,  becoming  conspicuously  thickened  and  with  raised  minor  venation  between  the  longitu- 
dinal veins  in  fruiting  stages B.  weberbaueri 


Buchnera  pusilla  Kunth  in  H.B.K.,  Nov.  Gen. 
Sp.  2:  ed.  quarto  340.  1818.  B.  tinctoria  Bertol., 
Fl.  Guatemala  26.  1840.  B.  major  Polak.,  Lin- 
naea  41:  588.  1877.  B.  mexicana  Hemsl.,  Biol. 
Centr.  Am.  Bot.  2:  457.  1881.  Figure  4. 

Stiff  erect  herbs  (6-)10-70(-150)  cm  tall,  an- 
nual, unbranched  or  with  2-7  distal  branches,  dry- 
ing dark,  stems  0.4-3  mm  diam.,  terete,  hispid 
with  stiff  whitish  hairs  0.3-1.3  mm  long.  Leaves 
opposite  or  subopposite,  sessile  or  with  poorly 
differentiated  petioles  to  8  mm  long;  leaf  blades 
15-50  mm  long,  1-8  mm  wide,  linear  to  linear- 
oblong  or  linear-oblanceolate  (oblong  near  the 
base),  apex  acute,  margin  with  1-5  prominent 
teeth  (0.2-1  mm)  or  entire,  gradually  narrowed  to 
the  cuneate  base,  subcoriaceous,  with  scattered 
short  scabrid  hairs  above,  larger  (0.2-0.5  mm) 
stiff  hairs  along  the  margin  and  veins  beneath, 
with  1  or  3  major  veins.  Inflorescences  3-15  cm 
long,  bracts  4-9  mm  long,  ovate  to  lanceolate, 
pedicels  0-1  mm  long,  with  2  shorter  linear  brac- 
teoles  beneath  the  calyx,  bracts  and  bracteoles 
scabrid  and  ciliate.  Flowers  with  calyx  4-5  mm 
long,  enlarging  (to  8  mm)  and  becoming  more 
scabrid  in  fruit,  with  10  longitudinal  veins,  hispid 
only  along  the  veins,  lobes  1-2.5  mm  long,  equal 
or  unequal,  acute;  corolla  8-14  mm  long,  salver- 
form,  white  to  purplish  white  or  lilac,  tube  0.4- 
0.8  mm  diam.,  lobes  1-5  mm  long,  obovate  with 
narrowed  base  and  broadly  rounded  apex;  stamens 


inserted  near  the  middle  of  the  tube.  Fruits  4-6 
mm  long,  ca.  2  mm  wide,  oblong,  smooth,  partly 
enclosed  in  the  scabrid  calyx;  seeds  0.4-0.5  mm 
long,  oblong  or  curved. 

Plants  of  open  sunny  sites  in  fields  and  savan- 
nas in  deciduous  and  evergreen  forest  areas  of  the 
Pacific  slope  and  central  highlands,  10-1700  m 
elevation.  Flowering  and  fruiting  from  October  to 
early  March.  The  species  ranges  from  Mexico  to 
Ecuador  and  Brazil. 

Buchnera  pusilla  is  distinguished  by  its  short, 
stiff,  erect  (usually  few-branched)  habit,  hispid  to 
scabrous  vesture,  spicate  inflorescences,  and  sal- 
verform  white  to  pinkish  corollas,  all  parts  turning 
dark  when  dried.  This  is  a  frequently  collected 
species  in  Central  America.  We  have  included 
Herrera  801  within  our  concept  of  this  species, 
which  has  been  identified  as  B.  longifolia  Kunth 
by  Wilcox  (1965,  see  above).  Buchnera  pusilla 
and  B.  longifolia  form  a  variable  species  complex, 
widely  distributed  in  the  lowland  Neotropics. 
They  are  distinguished  by  the  amount  and  type  of 
pubescence  on  the  calyx,  but  that  character  can 
vary  with  the  age  of  the  flower,  the  fruits  becom- 
ing more  scabrid  as  they  mature.  See  the  illustra- 
tion in  Flora  of  Guatemala  (Standley  &  Williams, 
1973,  p.  337). 

Buchnera  weberbaueri  Diels,  Bot.  Jahrb.  Syst. 
37:  430.  1906.  B.  leiantha  Standl.,  Publ.  Field 
Mus.  Nat.  Hist.,  Bot.  Sen  22:  105.  1940. 


26 


FIELDIANA:  BOTANY 


Stiff  erect  herbs  30-70  cm  tall,  annual,  usually 
without  lateral  branches,  leafy  stems  1-2.5  mm 
diam.,  glabrous  or  with  minute  (0.1-0.2  mm)  sca- 
brid  hairs  in  longitudinal  files  or  at  lower  inter- 
nodes,  usually  smooth  to  the  touch.  Leaves  op- 
posite or  subopposite,  sessile  and  slightly  clasping 
the  stem,  a  petiole  not  differentiated;  leaf  blades 
2-7  cm  long,  0.5-4  mm  wide,  linear  to  linear- 
lanceolate,  apex  acute,  margins  entire  (except 
rarely  on  basal  leaves),  base  gradually  narrowed, 
drying  black  or  brown,  glabrous,  with  1  central 
vein  or  tripliveined.  Inflorescences  4-14  cm  long, 
flowers  separate  or  closely  congested  into  a  dense 
spike,  abaxial  bracts  2-3  mm  long,  acute,  enlarg- 
ing to  5  X  3  mm  in  fruit,  ciliolate  along  the  mar- 
gin, pedicels  0-1  mm  long.  Flowers  with  calyx 
5-7  mm  long,  to  10  mm  in  fruit,  sparsely  puber- 
ulent  with  few  white  hairs,  sepal  lobes  ca.  1 .5  mm 
long,  acute,  minor  venation  between  the  main 
veins  not  apparent  until  fruiting;  corolla  7-13  mm 
long,  salverform,  lilac  to  pink-purple  or  white, 
tube  ca.  0.6  mm  diam.,  lobes  1.5-2  mm  long, 
equal.  Fruits  5-7  mm  long,  ca.  3  mm  diam.,  sur- 
face smooth  and  glabrous,  included  within  the 
stiff  thickened  calyx. 

Rarely  collected  plants  of  open  sunny  sites, 
200-1300  m  elevation.  In  Costa  Rica  this  species 
has  only  been  collected  in  Guanacaste  Province, 
flowering  in  January  and  September-October, 
with  old  fruit  in  February.  The  species  has  been 
collected  in  Belize,  in  Izabal  (Guatemala),  and  on 
the  Cerro  de  Espiritu  Santo  near  Naranjo  in  Costa 
Rica;  it  ranges  southward  to  Trinidad  and  Peru. 

Buchnera  weberbaueri  is  recognized  by  its 
short,  usually  unbranched  stems,  very  sparse  pu- 
bescence, linear  entire  leaves,  and  sepals  that  be- 
come indurated  and  with  many  elevated  veins  in 
fruiting  stages.  Two  rather  different  specimens  are 
provisionally  placed  here:  Herrera  801,  with 
young  flowers,  and  Williams  &  Williams  24514, 
with  old  fruits.  Standley  and  Williams  (1973)  con- 
fused B.  weberbaueri  with  the  larger-flowered  B. 
palustris  (Aubl.)  Spreng.,  which  is  found  in  low- 
land South  America  in  wet  areas.  In  addition  to 
its  larger  flowers,  B.  palustris  has  longer  pedicels 
and  narrowly  lanceolate  bracts  and  bracteoles. 


Calceolaria  Linnaeus 

RKFERENCES — U.  Molau,  Scrophulariaceae,  Part 
1.  Calceolarieae.  Fl.  Neotropica,  Monogr.  47:  1- 
326.  1988.  L.  R.  Landrum  &  R.  McVaugh,  Cal- 
ceolaria mexicana  and  C.  tripartita  in  Mexico. 
Contrib.  Univ.  Michigan  Herb.  11:  273-309.  1978. 


Shrubs,  vines  or  herbs,  annual  or  perennial, 
usually  confined  to  high-elevation  habitats,  stems 
terete,  hairs  simple.  Leaves  opposite  (ternate  in 
some  Andean  species),  sessile  or  petiolate,  leaf 
blades  simple  and  without  prominent  lobes  or  be- 
coming deeply  lobed  or  pinnatifid.  margins  usu- 
ally serrate,  pinnatcly  veined.  Inflorescences  ter- 
minal or  axillary,  often  a  compound  thyrse  of 
cymes  with  3  or  4  flowers  or  of  1-3  axillary  flow- 
ers, often  subtended  by  smaller  leaves,  bracteoles 
absent  at  the  base  of  the  calyx.  Flowers  small  to 
large,  calyx  deeply  4-parted.  sepal  lobes  equal  or 
unequal,  valvate  in  bud,  usually  persisting  in  fruit; 
corolla  strongly  bilabiate  with  the  lower  lip  usu- 
ally developed  into  a  globose  or  slipper-like  sac- 
cate form,  the  upper  lip  usually  much  smaller  and 
arched  or  hooded  and  enclosing  style  and  sta- 
mens, yellow  to  red  or  purple,  sometimes  spotted 
or  mottled,  glabrous  on  the  exterior;  stamens  2, 
attached  near  the  base  of  the  tube,  filaments  short, 
anther  with  2  contiguous  thecae  opening  by  a  lon- 
gitudinal slit,  staminodes  absent;  ovary  superior 
and  conical  (partly  inferior  in  the  temperate  sub- 
genera  Cheiloncos  and  Rosula),  somewhat  2- 
lobed.  2-locular,  ovules  many,  style  short  and  of- 
ten recurved,  stigma  simple  or  capitate.  Fruits 
dry  capsules,  septicidal  and  loculicidal,  the  4 
valves  opening  from  the  apex;  seeds  many,  small 
(0.3-1  mm),  usually  with  longitudinal  and  trans- 
verse ridges  or  minute  tubercles. 

A  pantropical  genus  of  ca.  300  species  with  a 
majority  of  the  species  found  above  1000  m  ele- 
vation in  the  Andes.  A  number  of  species  range 
into  the  lowlands  of  temperate  Chile  and  Argen- 
tina. The  genus  is  classified  with  Jovellana  and 
Porodittia  in  the  tribe  Calceolarieae.  This  tribe 
exhibits  links  between  southern  South  America 
and  New  Zealand,  duplicating  the  disjunction 
found  in  the  tribe  Veronicae.  Molau  (1988)  pro- 
vided a  fine  treatment  of  this  complex  and  fasci- 
nating group  in  the  Neotropics;  his  discussions  of 
biology,  distribution,  speciation,  and  phylogeny 
are  especially  noteworthy.  Calceolaria  is  imme- 
diately recognizable  because  of  its  bright  yellow 
bilabiate  corollas  with  small  upper  lip  and  round- 
ed saccate  or  inflated  slipper-like  lower  lip.  The 
four-parted  calyx,  only  two  stamens,  and  restric- 
tion to  moist  high-elevation  habitat  (in  Central 
America)  are  further  distinctions.  Some  South 
American  species  have  corollas  that  are  somewhat 
S-shaped  in  lateral  view.  A  number  of  hybrids  are 
popular  as  potted  ornamentals,  called  "slipper- 
worts"  or  "slipper  flowers." 


BURGER:  FLORA  COSTARICENSIS 


27 


Key  to  the  Species  of  Calceolaria 

la.   Petioles  broadly  winged  to  the  base  and  united  across  the  node,  the  leaves  perfoliate  [larger  blades 

triangular  to  sagittate,  never  pinnatifid:  anther  cells  contiguous,  dark  brown;  uncommon]    

C.  perfoliata 

Ib.  Petioles  not  broadly  winged  to  the  base,  the  leaves  not  perfoliate    2 

2a.  Larger  leaves  lanceolate,  without  pinnate  lobes  or  deep  sinuses;  anther-thecae  contiguous,  not  sep- 
arated by  an  expanded  connective;  stems  slightly  woody  and  plants  clambering  to  2  m  high  (note 

that  these  2  species  are  very  similar)   3 

2b.  Larger  leaves  usually  with  prominent  pinnate  lobes  separated  by  deep  sinuses;  anther-thecae  sepa- 
rated by  the  slender  expanded  connective;  stems  mostly  herbaceous  and  semisucculent,  plants  to  1 

m  high   4 

3a.  Fruits  puberulent;  2°  veins  of  leaves  not  loop-connected  in  the  distal  half  of  the  blade,  usually 
strongly  ascending,  upper  surface  of  the  blade  with  very  short  hairs  and  glands,  lower  surface 
often  with  brown  punctations;  stems  usually  puberulent;  seeds  with  prominent  longitudinal 

ridges  and  smaller  transverse  ribs;  common  high-montane  plants    C.  irazuensis 

3b.  Fruits  glabrous;  2°  veins  of  leaves  usually  loop-connected  in  the  distal  half  of  the  blade,  upper 
surface  lacking  very  short  hairs  or  glands,  lower  surface  not  punctate;  stems  usually  glabrous 
(rarely  with  a  few  hairs  at  nodes  and  petioles);  seeds  with  prominent  longitudinal  ridges  but  no 

transverse  ribs;  rarely  collected  in  Costa  Rica C.  microbe/aria 

4a.  Corolla  5-15  mm  long,  mouth  open;  fruits  3-7  mm  long,  oblate;  seeds  0.4-0.5  mm  long;  sinuses 

of  larger  leaves  rarely  reaching  the  midvein;  lower  anther-theca  fertile    C.  mexicana 

4b.  Corolla  10-20  mm  long,  mouth  closed;  fruits  6-9  mm  long,  ovoid;  seeds  0.6-0.8  mm  long;  sinuses 
of  larger  leaves  sometimes  reaching  the  midvein;  lower  anther-theca  sterile    C.  tripartita 


Calceolaria  irazuensis  J.  D.  Smith,  Bot.  Gaz.  20: 
292.  1895.  C.  costaricensis  Kranzl.,  Ann.  K.  K. 
Naturhist.  Hofmus.  22:  192.  1907.  Figure  3. 

Subshrubs,  erect  or  scandent,  0.2-1  (-2)  m  tall, 
leafy  stems  1.2-5  mm  diam.,  densely  puberulent 
with  slightly  viscous  hairs  0.05-0.3  mm  long,  old- 
er stems  glabrescent.  Leaves  forming  a  line  across 
the  stem  at  their  base,  petioles  2-8(-15)  mm  long, 
0.5-1.8  mm  diam..  puberulent  with  longer  hairs 
adaxially;  leaf  blades  1.8-9(-12)  cm  long,  6-35(- 
45)  mm  wide,  lanceolate  to  ovate-lanceolate  or 
elliptic-lanceolate,  gradually  narrowed  to  the 
acute  apex,  margin  serrate  with  teeth  0.3-1  mm 
high,  2-5  teeth/cm,  base  acute  to  obtuse  or  slight- 
ly rounded,  asymmetric,  drying  much  paler  be- 
neath than  above,  minutely  (0.1-0.2  mm)  puber- 
ulent above,  glabrous  between  the  veins  beneath, 
2°  veins  4-9/side  and  strongly  ascending.  Inflo- 
rescences mostly  of  2-4  long-pedunculate  cymes 
subtended  by  a  pair  of  smaller  leaves  (bracts)  or 
with  2-4  flowers  terminal  on  the  distal  leafy  node, 
minutely  viscid  puberulent,  peduncles  1.5-12  cm 
long,  1-2  mm  diam.,  usually  bearing  3  or  4  ped- 
icellate flowers,  pedicels  1-4  cm  long,  0.4-0.6 
mm  diam.  Flowers  with  calyx  6-9  mm  long,  ca- 
lyx lobes  4-9  mm  long,  broadly  ovate-triangular, 
minutely  puberulent  along  the  margin;  corolla 


12-24  mm  long,  minutely  glandular-papillose  on 
the  exterior,  upper  lip  6-8  mm  long,  lower  lip  1 5- 
25  mm  long,  12-18  mm  wide,  bright  yellow;  fil- 
aments ca.  1.5  mm  long,  anthers  with  divaricate 
thecae  to  3.5  mm  wide;  style  2-3  mm  long, 
curved.  Fruits  6-9  mm  long,  5-8  mm  wide,  ovate 
with  truncated  base,  minutely  papillate-puberu- 
lent;  seeds  0.4-0.7  mm  long,  0.2-0.3  mm  wide, 
oblong-ellipsoid,  dark  brown,  with  prominent  lon- 
gitudinal ridges  and  minute  transverse  ribs  (X50). 

Plants  of  evergreen  high-montane  forest  for- 
mations, (1800-)2400-3500  m  elevation.  Flow- 
ering and  fruiting  throughout  the  year  but  with 
most  collections  made  between  November  and 
May.  The  species  ranges  from  Volcan  Barva 
southward  to  the  Chiriqui  highlands  of  Panama. 

Calceolaria  irazuensis  is  recognized  by  it  most- 
ly lanceolate  leaves,  slightly  viscid  hairs,  inflores- 
cences often  with  long  peduncles  and/or  pedicels, 
and  slipper-shaped  yellow  flowers.  This  species  is 
frequently  encountered  in  open  sites  at  higher  el- 
evations. Compare  the  closely  similar  but  rarely 
collected  C.  microbe/aria.  Common  names  are 
baton  de  oro  and  gallitos. 

Calceolaria  mexicana  Benth.,  PI.  Hartw.  47. 
1840.  C.  trachelifolia  Martens  &  Galeotii, 
Acad.  R.  Sci.  Bruxelles  12:  16.  1845.  C.  urti- 


28 


FIELDIANA:  BOTANY 


cina  Kranzl.,  Feddes  Repert.  Spec.  Nov.  Regni 
Veg.  1:  82.  1905.  Figure  3. 

Annual  herbs,  erect  to  decumbent,  10-60(- 
100)  cm  tall,  leafy  stems  0.6-3  mm  diam.,  slightly 
succulent,  often  reddish  in  color,  sparsely  to 
densely  puberulent  with  simple  or  viscid  gland- 
tipped  hairs  0.3-0.6  mm  long.  Leaves  slightly 
clasping  the  stem  and  forming  an  interpetiolar 
ridge,  petioles  2-40(-160)  mm  long,  0.4-1.4  mm 
diam.,  pubescence  similar  to  that  of  the  stem;  leaf 
blades  1-12  cm  long,  0.4-8(-13)  cm  wide,  vary- 
ing from  narrowly  ovate-triangular  in  smaller 
blades  to  broadly  ovate-triangular  in  outline  with 
2-4  prominent  pinnatifid  lobes  separated  by  deep 
sinuses  in  larger  blades,  apex  acute,  margins 
strongly  dentate-serrate  with  intermixed  larger  (2- 
4  mm)  and  smaller  (0.3-2  mm)  teeth,  base  obtuse 
to  subcordate,  drying  yellowish  green  to  grayish 
green,  upper  surface  with  slender  hairs  0.2-0.4 
mm  long,  lower  surface  with  few  thin  hairs  to  1 
mm  long  on  the  major  veins,  2°  veins  6-12/side. 
Inflorescences  of  solitary  axillary  flowers  (21 
node)  or  resembling  terminal  cymes  when  sub- 
tended by  reduced  bract-like  distal  leaves,  pedi- 
cels 4-14  mm  long,  elongating  in  fruit,  0.15-0.25 
mm  diam.,  minutely  puberulent  with  gland-tipped 
hairs.  Flowers  with  calyx  4-6  mm  long,  calyx 
lobes  2-5  mm  long,  1-3  mm  wide  at  the  base, 
acute,  with  glandular  hairs  at  the  base;  corolla  5- 
15  mm  long,  3-10  mm  wide,  the  upper  lip  1-3 
mm  long,  lower  lip  5-14  mm  long,  3-6  mm  wide, 
slipper-shaped,  yellow;  anthers  2-3.5  mm  wide, 
the  2  small  thecae  fertile,  separated  by  the  longer 
narrow  connective;  ovary  2.5  mm  diam.,  papil- 
late-puberulent.  Fruits  3-7  mm  long,  3-6  mm 
wide,  oblate  or  globose,  thin-walled;  seeds  0.4- 
0.5  mm  long,  ca.  0.3  mm  thick,  oblong,  dark. 

Plants  of  wet  sites  near  streams,  moist  depres- 
sions, and  wet  cliffsides,  1500-3200  m  elevation. 
Flowering  throughout  the  year,  but  with  most 
Costa  Rican  collections  made  in  April-August. 
This  species  is  the  commonest  Calceolaria  in 
Mexico  and  Central  America;  it  ranges  to  Bolivia. 

Calceolaria  mexicana  is  recognized  by  its  pref- 
erence for  wet  montane  habitats,  diverse  foliage 
with  the  large  leaves  having  prominent  pinnate 
lobes,  small  slipper-shaped  yellow  corollas,  and 
stamens  with  two  small  but  functional  thecae. 
This  species  can  be  found  growing  together  with 
the  very  similar  C.  tripartita,  and  the  two  are  eas- 
ily confused.  Both  species  have  a  strongly  devel- 
oped connective  separating  the  thecae.  In  C.  mex- 
icana the  corollas  are  open  and  the  lower  lip  is 


slightly  three-lobed.  In  addition,  the  larger  leaves 
do  not  have  the  deep  sinuses  of  C.  tripartita 
leaves. 

Calceolaria  microbefaria  Kranzl.,  Ann.  K  K. 
Natur.  Hofmus.  Wien  22:  193.  1907.  C.  storkii 
Standl.,  Publ.  Field  Mus.  Nat.  Hist.  Bot.  Ser.  18: 
1103.  1938. 

Shrubs  or  subshrubs  0.3-3  m  tall,  erect  or 
clambering,  much  branched,  stems  glabrous  or 
with  a  few  hairs  at  the  nodes.  Leaves  with  peti- 
oles 3-1 1  mm  long,  glabrous  or  with  a  few 'hairs 
along  the  adaxial  margins;  leaf  blades  4-9(-13) 
cm  long.  l-2.3(-2.8)  cm  wide,  lanceolate  to  nar- 
rowly oblong-lanceolate,  apex  acute,  margin  ser- 
rate with  5-7  gland-tipped  mucronulatc  teeth/cm, 
base  acute,  upper  surface  glabrous  or  minutely  pa- 
pillate, lower  surface  often  drying  pale  grayish  or 
reddish,  2°  veins  5-1  I/side,  loop-connected  dis- 
tally.  Inflorescences  3-12  cm  long,  usually  with 
2  or  3  pairs  of  4-8  flowered  cymes,  peduncles  1- 
5  cm  long,  bracts  often  lacking  at  the  base  of  the 
cymes,  pedicels  0.5-3.4  cm  long,  glabrous  to  mi- 
nutely papillate-puberulent  (to  tomentose  in  South 
America).  Flowers  with  sepals  3.3-6  mm  long, 
2.8-4  mm  wide  at  anthesis,  minutely  puberulent 
along  the  margins;  corolla  10-20  mm  long,  bright 
yellow,  the  upper  lip  hooded  or  flattened  and  sub- 
circular,  the  lower  lip  projecting  or  pendant,  sac- 
cate; anthers  2-3.7  mm  wide.  Fruits  5-9  mm 
long,  5-9  mm  wide,  broadly  ovoid  with  truncated 
base,  glabrous;  seeds  0.7-0.9  mm  long,  ca.  0.3 
mm  thick,  with  prominent  longitudinal  ridges, 
surface  smooth. 

Plants  of  subparamo  formations  near  the  high- 
est point  along  the  Interamerican  Highway  in  the 
western  part  of  the  Talamanca  range.  3100-3500 
m  elevation.  Flowering  in  March-August;  fruiting 
in  August.  This  species  is  known  from  only  four 
collections  in  Costa  Rica  (Barringer  el  al.  2913, 
Grayum  &  Affolter  8190,  Skutch  5188,  and  Stork 
3048,  type  of  C.  storkii).  The  species  ranges  from 
Venezuela  to  central  Ecuador  in  the  Andes,  with 
a  small  disjunct  population  in  Costa  Rica. 

Calceolaria  microbefaria  is  recognized  by  its 
lanceolate  leaves,  generally  glabrous  vegetative 
parts  and  fruits,  bright  yellow  slipper-shaped  co- 
rollas, and  very  limited  range  in  our  area.  Molau 
(1988)  divides  this  species  into  three  geographi- 
cally and  morphologically  distinct  subspecies, 
with  the  Costa  Rican  collections  placed  in  subsp. 
microbefaria.  In  Costa  Rica,  this  species  is  easily 
mistaken  for  the  much  more  common  C.  irazuen- 


BURGER:  FLORA  COSTARICENSIS 


29 


sis;  the  characteristics  used  in  the  key  to  separate 
the  two  species  usually  allow  confident  identifi- 
cation. 

Calceolaria  perfoliata  L.f.,  Suppl.  86.  1781.  Fa- 
gelia  perfoliata  (L.f.)  Kuntze,  Rev.  Gen.  PI.  2: 
460.  1891.  C.  sciadephora  J.  D.  Smith,  Bot. 
Gaz.  25:  151.  1898.  Figure  6. 

Herbs  or  weak-stemmed  subshrubs  0.5-1.5  m 
high  (to  5  m  long),  often  scandent  on  other  plants, 
leafy  stems  1.5-6  mm  diam.,  often  with  4  prom- 
inent longitudinal  ridges,  slightly  succulent,  with 
slender  crooked  multicellular  hairs  0.3-1  mm 
long.  Leaves  sessile  and  perfoliate,  opposing 
leaves  united  at  the  base  with  lateral  tissue  to  6 
mm  wide,  leaves  subtending  the  inflorescences 
not  perfoliate,  petioles  broadly  winged  and  blade 
like,  0.5-6  cm  long,  3-12  mm  wide  and  expanded 
near  the  stem;  leaf  blades  2-11  cm  long  (beyond 
the  blade-like  petiole),  2-7  cm  wide,  narrowly  tri- 
angular or  sagittate,  apex  acute,  margins  serrate- 
dentate  with  larger  (2-4  mm)  and  smaller  (1-2 
mm)  teeth  intermixed,  base  truncated  to  the 
winged  petiole-like  part,  drying  yellowish  brown, 
often  silvery  below  (in  life),  upper  surface  with 
short  (0.2  mm)  scattered  hairs,  lower  surface  with 
thin  hairs  to  1  mm  long,  2°  veins  5-7/side.  Inflo- 
rescences with  8-12  (4-24)  flowers,  borne  on  ax- 
illary peduncles  6-15  cm  long,  1-1.5  mm  diam., 
terminated  by  a  pair  of  asymmetric  leaves/bracts 
2-5  cm  long,  pedicels  1 .5-5  cm  long,  densely  pu- 
berulent.  Flowers  with  calyx  8-12  mm  long, 
lobes  6-8  mm  long,  puberulent,  broadly  ovate, 
rounded  at  base,  apex  obtuse,  enlarging  slightly 
in  fruit;  corolla  to  22  mm  long,  bright  yellow, 
upper  lip  4-9  mm  long,  lower  lip  1 5-20  mm  long, 
8-18  mm  wide,  anthers  U-  or  C-shaped,  2-4  mm 
wide;  ovary  pubescent,  style  3-5  mm  long.  Fruits 
5-7  mm  long,  ca.  5  mm  diam.,  ovoid  with  trun- 
cated base,  pubescent;  seeds  0.5-0.7  mm  long, 
0.1-0.2  mm  wide. 

Infrequently  collected  plants  of  evergreen  mon- 
tane forest  formations,  2700-3300  m  elevation  (as 
low  as  1800  m  in  Chiriqui).  Flowering  and  fruit- 
ing appear  to  be  restricted  to  January-February  in 
Costa  Rica.  This  species  ranges  from  the  Cordil- 
lera de  Talamanca  and  the  Chiriqui  highlands  to 
southern  Ecuador. 

Calceolaria  perfoliata  is  distinguished  by  its 
perfoliate  leaves  with  broadly  winged  petioles  and 
distal  narrowly  triangular  or  sagittate  blades.  The 
anthers  are  attached  at  the  center  and  decurved 
laterally  to  produce  a  rounded  C-shaped  or  horse- 


shoe-shaped form.  The  restricted  flowering  period 
may  account  for  the  relatively  few  Costa  Rican 
collections.  A  related  species,  C.  trilobata  Hem- 
sley,  is  native  to  Mexico-Guatemala  and  Vene- 
zuela-Bolivia but  is  not  known  from  southern 
Central  America.  That  species  has  a  more  defi- 
nitely contracted  petiole  between  the  distal  blade 
and  the  expanded  perfoliate  base,  and  the  anthers 
are  not  horseshoe-shaped. 

Calceolaria  tripartita  R.  &  P.,  Fl.  Peruv.  Prodr. 
1:  14,  tab.  22.  1798.  C.  heterophylla  Wild., 
Enum.  PI.  1:  29.  1809  (non  C.  heterophylla  R. 
&  P.).  Figure  3. 

Herbs,  erect  or  decumbent,  9-90  cm  tall,  stems 
1-4  mm  diam.,  slightly  succulent,  puberulent  with 
multicellular,  often  gland-tipped  hairs  0.2-0.7  mm 
long.  Leaves  simple  and  serrate  to  deeply  pinnat- 
ifid,  slightly  clasping  the  stem  and  forming  a  ridge 
across  the  node,  petioles  1-5  cm  long,  0.5-1.5 
mm  wide,  glandular  puberulent;  leaf  blades  1-16 
cm  long,  0.3-1 1  cm  wide,  ovate-lanceolate  in 
smaller  leaves  to  broadly  ovate-triangular,  larger 
blades  with  deep  proximal  sinuses  separating  1-3 
pairs  of  subopposite  lobes,  apex  acute,  margin 
with  larger  (3-10  mm)  and  smaller  (0.3-3  mm) 
teeth,  base  truncated  in  larger  leaves,  drying 
greenish  or  brown,  upper  surface  with  transparent 
multicellular  hairs  to  1  mm  long.  Inflorescences 
of  terminal  or  axillary  flowers,  2-4/node  (if  small 
distal  leaves  are  interpreted  as  bracts,  the  sub- 
tending internodes  can  be  interpreted  as  peduncles 
of  thyrse-like  inflorescences),  pedicels  4-45  mm 
long,  0.2-0.4  mm  diam.,  puberulent  with  simple 
or  glandular  hairs  0.2-0.4  mm  long.  Flowers  with 
calyx  5-7  mm  long,  lobes  ca.  4  mm  long,  with 
small  glandular  hairs;  corolla  10-20  mm  long, 
bright  yellow,  upper  lip  2-5  mm  long,  lower  lip 
10-25  mm  long,  6-15  mm  wide,  slipper-shaped, 
unlobed;  anthers  2-5  mm  long,  one  theca  sterile, 
reduced  to  a  nob,  connective  slender.  Fruits  6-9 
mm  long,  ovoid,  rounded  at  the  apex;  seeds  0.6- 
0.8  mm  long,  ca.  0.4  mm  diam.,  reddish  brown, 
with  longitudinal  ridges. 

Plants  of  wet  or  moist  open  sites  in  evergreen 
montane  forest  formations,  1000-3200  m  eleva- 
tion. Flowering  throughout  the  year,  but  with  most 
collections  made  between  October  and  December. 
This  species  ranges  from  Mexico  to  Peru. 

Calceolaria  tripartita  is  recognized  by  it  high- 
land habitats,  glandular  viscid  pubescence,  larger 
leaves  deeply  pinnatisect  (almost  pinnately  com- 
pound), yellow  slipper-shaped  corollas,  and  un- 


30 


FIELDIANA:  BOTANY 


usual  stamens.  This  species  is  similar  to  C.  mex- 
icana  (q.v.),  and  the  two  have  often  been  con- 
fused, as  in  the  Flora  of  Guatemala.  In  addition 
to  the  differences  used  in  the  key,  the  pedicels  of 
C.  tripartita  usually  have  more  conspicuous 
gland-tipped  hairs. 


Capraria  Linnaeus 

REFERENCES — T.  A.  Sprague,  A  revision  of  the 
genus  Capraria.  Kew  Bull.  1921:  205-212.  1921. 
C.  Niezgoda  &  S.  Tomb,  Systematic  palynology 
of  the  tribe  Leucophylleae  (Scrophulariaceae)  and 
selected  Myoporaceae.  Pollen  et  Spores  17:  497- 
516.  1975. 

Herbs  or  subshrubs,  annual  or  perennial, 
branches  usually  slender  and  terete,  pubescent  or 
glabrous.  Leaves  alternate,  sessile  or  petiolate, 
blades  usually  serrate,  glandular  punctate  (viewed 
by  transmitted  light).  Inflorescences  of  1-3  axil- 
lary flowers  borne  on  slender  pedicels  at  distal 
nodes  (racemose  when  distal  leaves  are  reduced 
and  bract-like),  bracteoles  absent  beneath  the  ca- 
lyx. Flowers  radially  symmetric,  calyx  united 
only  near  the  base,  sepals  5,  equal  or  subequal, 
narrow,  valvate  in  bud;  corolla  funnelform  or 
campanulate,  white,  purple,  or  greenish  yellow, 
glabrous  externally,  tube  as  long  as  the  lobes  or 
slightly  shorter,  lobes  5,  throat  sometimes  bearded 
within;  stamens  4  or  5,  equal  or  subequal,  includ- 
ed, borne  proximally  or  distally  within  the  tube, 
filaments  glabrous,  anthers  sagittate,  basifixed, 
versatile,  introrse,  thecae  divergent  but  basal  parts 
confluent;  ovary  2-locular,  ovules  many,  style 
slender,  apically  flattened  with  stigmas  on  the  lat- 
eral faces.  Fruits  capsules,  ellipsoid  to  ovoid, 
glandular-punctate,  dehiscing  loculicidally  and 
secondarily  septicidal,  4-vaIved,  placenta  remain- 
ing as  a  conspicuous  column  with  pitted-reticulate 
surface;  seeds  many,  small,  yellow  to  brown. 

Capraria  is  a  genus  of  five  species  ranging 
from  the  southern  United  States  to  Peru;  one  spe- 
cies is  adventive  in  West  Africa.  The  narrow  al- 
ternate leaves,  flowers  that  are  radially  symmetric, 
and  fruit  with  persisting  reticulated  column  (pla- 
centa) make  these  plants  distinctive.  The  campan- 
ulate five-lobed  corolla  and  five  stamens  are  un- 
usual in  the  Scrophulariaceae,  so  that  the  genus 
has  never  been  satisfactorily  classified  within  the 
family.  Niezgoda  and  Tomb  (reference  above) 
showed  that  the  pollen  of  Capraria  is  diorate  and 
more  closely  related  to  types  found  in  the  My- 


oporaceae than  to  the  pollen  of  other  Scrophular- 
iaceae. 

Capraria  biflora  L.,  Sp.  PI.  628.  1753.  C.  biflora 
var.  pilosa  Griseb.,  Fl.  Brit.  W.I.  427.  1861.  C. 
biflora  form  hirta  Loes.,  Bull.  Herb.  Boissier  2, 
3:  284.  1903.  Figure  7. 

Herbs  or  subshrubs.  perennial,  erect  or  clam- 
bering, 0.5-2  m  tall,  distal  stems  usually  few- 
branched,  leafy  stems  0.7-5  mm  diam.,  densely 
puberulent  to  glabrescent  with  thin  erect  whitish 
hairs  0.2-0.7  mm  long.  Leaves  alternate,  gradu- 
ally becoming  smaller  on  distal  stems,  petioles  to 
8  mm  long  but  usually  not  clearly  differentiated; 
leaf  blades  1.5-11  cm  long,  6-30  mm  wide,  el- 
liptic to  elliptic-oblanceolate  or  oblanceolate, 
apex  acute  and  sharp-tipped,  distal  half  of  the 
margin  with  sharp-pointed  teeth  0.5-3  mm  high. 
1-6  teeth/cm,  gradually  narrowed  to  the  cuneate 
base,  drying  grayish  green,  both  surfaces  with 
thin  white  hairs  0.1-0.4  mm  long,  2°  veins  3  or 
4/side,  strongly  ascending.  Inflorescences  of  2  or 
3  flowers  in  distal  leaf  axils,  pedicels  4-12  mm 
long,  ca.  0.2  mm  diam..  minutely  puberulent,  not 
articulated  at  the  base  of  the  calyx.  Flowers  with 
calyx  4-6  mm  long,  sepals  1-1.5  mm  wide  at  the 
base,  narrowly  triangular  to  linear,  to  7  mm  long 
in  fruit,  with  short  (0.3  mm)  thin  whitish  hairs; 
corolla  7-10  mm  long,  4-7  mm  wide  at  the 
mouth,  tubular-campanulate,  white,  lobes  3-5  mm 
long,  2.5-3.5  mm  wide  at  base,  triangular;  fila- 
ments attached  at  the  base  of  the  tube.  3-4  mm 
long,  anthers  ca.  3  mm  long.  Fruits  4-6  mm  long. 
2.5-4  mm  wide,  ovoid-ellipsoid  and  bisulcate, 
surface  smooth,  yellowish  brown  and  punctate; 
seeds  0.3-0.4  mm  long,  oblong  or  angular. 

Plants  of  open  sunny  sites  in  lowland  situations 
in  both  the  Caribbean  (evergreen)  and  northern 
Pacific  (deciduous)  areas  of  Costa  Rica,  0-500  m 
elevation  (to  1000  m  in  Nicaragua).  Flowering 
and  fruiting  throughout  the  year  but  collected  pri- 
marily in  July-January.  The  species  ranges  from 
Florida,  Mexico,  and  the  West  Indies  to  Argenti- 
na; it  is  found  on  Cocos  Island  and  has  become 
naturalized  in  West  Africa. 

Capraria  biflora  is  recognized  by  its  long  slen- 
der stems,  alternate  leaves  that  are  smaller  distal- 
ly, oblanceolate  leaf  blades  serrate  in  the  distal 
half,  flowering  nodes  with  two  or  three  flowers  on 
slender  pedicels,  and  flowers  with  narrow  sepals 
and  radially  symmetric  campanulate  white  corol- 
la. The  glandular  punctations  of  the  leaves  can 
best  be  seen  by  transmitted  light.  This  species  is 


BURGER:  FLORA  COSTARICENSIS 


31 


uncommon  in  Central  America.  In  Panama  it  is 
used  to  make  a  tea  that  may  be  a  dangerous  de- 
pressant in  large  quantities  (D'Arcy,  1979). 


Castilleja  Mutis  ex  Linnaeus  filius 

RHFKRHNCE — N.  H.  Holmgren,  Castilleja  (Scro- 
phulariaceae)  in  Costa  Rica  and  Panama.  Brittonia 
30:  182-194.  1978. 

Herbs  or  subshrubs,  annual  or  perennial,  erect, 
branching  mostly  from  the  base,  hemiparasitic  on 
roots  of  hosts,  drying  dark.  Leaves  alternate,  cau- 
line,  sessile  or  the  petioles  poorly  differentiated, 
leaf  blades  entire  to  deeply  dissected  or  pinnately 
lobed.  Inflorescences  terminal  spikes  or  racemes, 
spikes  usually  with  flowers  hidden  by  colorful 
bracts,  the  racemes  often  with  conspicuous  flow- 
ers arranged  along  one  side  (secund),  subtending 
leaves  intergrading  with  bracts,  distal  bracts  often 
more  conspicuous  than  the  flowers,  bracteoles  ab- 
sent at  the  base  of  the  calyx.  Flowers  bilaterally 
symmetric,  calyx  tubular  with  4  equal  or  unequal 
lobes  or  united  into  2  entire  lateral  lobes,  slightly 
enlarging  in  fruit;  corolla  bilaterally  symmetric 
and  strongly  2-lipped,  greenish  yellow  to  brightly 
colored,  tube  usually  elongate  and  narrow,  upper 
(adaxial)  lip  entire  with  united  lobes  hooded  and 
forming  a  beak-like  galea,  enclosing  the  anthers, 
lower  lip  slightly  3-saccate,  with  3  rudimentary 


teeth  or  petaloid  lobes;  stamens  4,  of  2  unequal 
pairs,  attached  near  or  above  the  middle  of  the 
corolla  tube,  anther  sacs  (thecae)  unequally  placed 
on  the  connective,  the  outer  longer  and  attached 
near  the  center,  the  inner  theca  smaller  and  borne 
on  the  apex  of  the  connective;  ovary  2-locular, 
ovules  many,  style  slender,  stigma  capitate  or 
slightly  2-lobed.  Fruits  dry  capsules,  usually 
asymmetric,  ovate  to  globose,  opening  loculici- 
dally;  seeds  many,  with  a  loose  reticulate  exotesta. 
A  genus  of  ca.  200  species,  ranging  from  North 
America  into  higher  elevation  Central  America 
and  the  Andes,  and  with  a  few  northern  Asian  and 
European  species.  These  plants  are  easily  recog- 
nized because  of  their  colorful  inflorescences  in 
which  bracts,  sepals,  and  petals  may  all  be  vari- 
ously colored  and  showy.  The  calyx  tube  is  some- 
what gibbous  at  the  base  (abaxially)  and  often 
with  only  two  large  lateral  lobes;  the  narrow, 
curved  corolla  has  a  beaked  upper  lip  that  is  usu- 
ally much  longer  than  the  lower  lip.  In  many  spe- 
cies all  parts  become  blackish  on  drying.  Many 
species  of  Castilleja  have  the  reputation  of  being 
very  variable  and  difficult  to  distinguish,  making 
the  delimitation  of  some  species  quite  arbitrary. 
This  problem  manifests  itself  in  Costa  Rica, 
where  C.  irasuensis,  C.  quirosii,  and  C.  talaman- 
censis  form  a  closely  related  complex  that  de- 
serves further  study.  Our  treatment  is  based  on  the 
publication  (1978,  cited  above)  and  determina- 
tions of  Noel  Holmgren. 


Key  to  the  Species  of  Castilleja 

la.  Calyx  9-12  mm  long,  corolla  9-12  mm  long;  flowers  sessile  and  difficult  to  see  within  the  densely 
congested  distal  portion  of  the  spicate  inflorescence;  specimens  drying  pale  grayish  green  to  dark 
gray;  700-2500  m  elevation  in  Costa  Rica  C.  arvensis 

Ib.  Calyx  13-24  mm  long,  corolla  18-34  mm  long;  flowers  short-  to  long-pedicellate  and  usually  easily 
seen  among  the  bracts;  specimens  drying  grayish  to  blackish;  1000-3800  m  in  Costa  Rica  ....  2 

2a.  Leaves  to  2  cm  long  and  nearly  as  wide,  pinnatifid  with  narrow  (1-2  mm)  central  rachis  and  slender 
filiform  (1  mm)  pinnate  lobes;  clefts  of  the  calyx  tube  more  nearly  the  same  (adaxial  4-6.5  mm 
deep,  abaxial  8-12  mm  deep);  rarely  collected,  1 100-1600  m  C.  tayloriorum 

2b.  Leaves  much  longer  than  wide,  usually  lanceolate  to  linear,  the  pinatifid  lobes  short  or  >  1  mm 
wide  when  present;  clefts  of  the  calyx  tube  differing  greatly  in  length  (adaxial  1-4  mm  deep,  abaxial 
9-17  mm  deep);  rare  to  common,  1400-3500  m 3 

3a.  Leaves  lanceolate,  mostly  3-6  cm  long;  upper  galeate  lip  equal  to  the  corolla  tube  or  up  to  1.5 
times  as  long;  rarely  encountered  at  1400-1700  m  elevation  C.  lentii 

3b.  Leaves  mostly  linear  or  pinnatifid,  1-4  cm  long;  upper  galeate  lip  2-3  times  as  long  as  the  corolla 
tube;  not  collected  from  below  1600  m  elevation  [the  following  three  species  are  part  of  a  closely 
related  complex]  4 

4a.  Plants  usually  densely  puberulent  with  stiff  slender  whitish  hairs  in  most  parts,  the  stems  terete  or 
with  poorly  defined  longitudinal  ridges,  parts  drying  dark  grayish;  corolla  tube  7-10  mm  long; 
uncommon  in  Costa  Rica,  1600-3500  m  C.  quirosii 


32 


FIELDIANA:  BOTANY 


4b.  Plants  glabrous  to  sparsely  pubemlent  with  slender  whitish  hairs,  the  stems  usually  with  well-defined 
longitudinal  ridges  continuous  with  the  leaf  bases,  parts  usually  drying  black;  corolla  tubes  7-16 
mm  long;  commonly  collected  in  Costa  Rica,  2600-3800  m  5 

5a.  Le.aves  mostly  entire,  with  short  lobes  near  the  apex;  calyx  red  to  brownish  purple  with  yellow 
distal  tips;  2600-3800  m  in  the  Cordillera  de  Talamanca  C.  talamancensis 

5b.  Leaves  usually  with  prominent  pinnate  lobes  in  the  distal  half;  calyx  reddish  throughout;  3000- 
3700  m  on  Volcan  Irazu,  Volcan  Turrialba,  and  rarely  collected  in  the  Cordillera  de  Talamanca  .  . 

C.  irasuensis 


Castilleja  arvensis  Schldl.  &  Cham.,  Linnaea  5: 
103.  1830.  C.  communis  Benth.  in  DC.,  Prodr. 
10:  529.  1846.  C.  agrestis  Pennell,  Fieldiana, 
Bot.  28:  519.  1953.  Figure  7. 

Erect  herbs,  annual,  1 5-80  cm  tall,  unbranched 
or  with  few  lateral  branches,  leafy  stems  0.7-4 
mm  diam.,  sparsely  to  densely  hirsutulous  with 
thin  whitish  hairs  0.2-1.5  mm  long,  gland-tipped 
hairs  also  present.  Leaves  sessile  and  lacking  a 
clearly  defined  petiole,  gradually  smaller  distally 
and  intergrading  with  the  floral  bracts;  leaf  blades 
15-60(-90)  mm  long,  4-14(-22)  mm  wide,  ob- 
lanceolate  to  narrowly  elliptic-obovate  or  elliptic, 
apex  bluntly  acute,  margin  entire,  gradually  nar- 
rowed to  the  cuneate  base,  drying  greenish  gray 
or  dark  gray,  with  thin  hairs  0.2-0.5  m  long  on 
both  surfaces,  tripliveined.  Inflorescences  4-15 
cm  long,  2-3  cm  diam.,  flowers  subsessile  and 
densely  congested  (becoming  more  distant  in 
fruit),  pubescent  throughout  with  villous  and  glan- 
dular hairs,  bracts  12-20  mm  long,  leaf-like  to 
ovate-oblong,  often  green  tipped  with  red  or  yel- 
low, enlarging  slightly  in  fruit,  pedicels  0-1  mm 
long.  Flowers  hidden  within  the  bracts,  calyx  9- 
12  mm  long,  ovate-oblong  and  slightly  curved, 
viscid-villous,  adaxial  and  abaxial  clefts  subequal, 
3-5  mm  deep;  corolla  9-12  mm  long,  greenish 
yellow,  usually  hidden  within  the  calyx,  tube  6- 
8.5  mm  long,  galeate  upper  lip  2.5-5  mm  long, 
lower  lip  reduced,  with  3  lanceolate  lobes;  ovary 
glabrous.  Fruits  6-8  mm  long,  3.5-5  mm  wide, 
ovoid-oblong,  dark  brown,  producing  many  seeds; 
seeds  0.6-1.2  mm  long,  0.2-0.3  mm  wide,  wedge- 
shaped  to  narrowly  rectangular  with  translucent 
truncated  ends. 

Weedy  plants  of  moist  open  sunny  sites  in  ev- 
ergreen or  partly  deciduous  forest  areas,  700- 
2500  m  elevation  (to  3000  m  in  Guatemala). 
Flowering  and  fruiting  throughout  the  year,  but 
collected  most  often  in  November-March.  The 
species  ranges  from  central  and  northeastern  Mex- 
ico to  Paraguay;  it  is  adventive  in  Hispaniola  and 
Hawaii. 


Castilleja  arvensis  is  distinguished  by  its  an- 
nual growth,  alternate  grayish  leaves  (when 
dried),  densely  congested  spikes  in  which  the  co- 
rollas are  difficult  to  see,  green  bracts  often  red- 
dish at  the  tips,  and  the  subsessile  fruit  producing 
numerous  wedge-shaped  seeds.  The  entire,  nar- 
rowly elliptic  leaves  give  this  species  a  quite  dif- 
ferent appearance  from  Costa  Rica's  other  Castil- 
leja species.  Small  (3  mm)  fleshy  tubercle-like 
leaves  may  be  present  at  the  base  of  the  stems 
near  ground  level.  This  is  a  well-defined  species 
and  the  widest  ranging  of  the  genus;  it  belongs  to 
section  Epichroma. 


Castilleja  irasuensis  Oersted,  Vidensk.  Meddel. 
Dansk  Naturhist.  Foren.  Kjobenhavn  1853:  27. 
1854.  Figure  7. 

Herbs  or  subshrubs,  10-50  cm  tall,  branching 
mostly  from  the  woody  base,  lateral  branches  usu- 
ally short,  leafy  stems  1-2  mm  diam.,  mostly  gla- 
brous, with  longitudinal  ridges.  Leaves  sessile,  ar- 
ticulated at  the  base,  with  fewer  lobes  distally  and 
becoming  bract-like  in  the  inflorescence;  leaf 
blades  6-30  mm  long,  the  central  portion  1-2.5 
mm  wide,  linear  and  entire  to  pinnatifid  with  1-3 
pairs  of  lobes  0.5-8  mm  long  and  ca.  0.8  mm 
wide  (the  longest  pair  often  also  pinnately  lobed), 
apex  bluntly  acute,  margin  usually  revolute,  base 
with  parallel  margins  to  the  stem,  drying  black, 
minutely  papillate  puberulent  above,  with  larger 
(0.1-0.2  mm)  hairs  beneath,  venation  obscure. 
Inflorescences  3-15  cm  long,  racemose  and  1 -sid- 
ed, rachis  hispid-villous,  proximal  bracts  little  dif- 
ferentiated from  the  leaves,  distal  bracts  oblanceo- 
late,  ca.  2  cm  long,  bright  red,  pedicels  2-6(-ll) 
mm  long,  ca.  0.3  mm  diam.,  pubescent.  Flowers 
with  calyx  14-20  mm  long,  4-6  mm  wide,  ovoid- 
tubular,  adaxial  cleft  1-4  mm  deep,  abaxial  cleft 
9-15  mm  deep,  lobes  rounded  and  entire,  becom- 
ing red,  puberulent;  corolla  22-30  mm  long,  tube 
8-12  mm  long,  galeate  upper  lip  12-19  mm  long, 
puberulent  and  green  above,  red  along  the  mar- 


BURGER:  FLORA  COSTARICENSIS 


33 


gins;  anthers  ca.  2  mm  long.  Fruits  7-12  mm 
long. 

Plants  of  open  habitats  in  high-montane  forest 
and  paramo  formations,  3000-3700  m  elevation. 
Flowering  and  fruiting  throughout  the  year.  The 
species  is  endemic  to  the  eastern  volcanoes  (Irazu, 
Turrialba)  and  the  western  part  of  the  Cordillera 
de  Talamanca. 

Castilleja  irasuensis  is  recognized  by  its  re- 
striction to  high-elevation  habitats,  alternate  pin- 
natifid  leaves,  and  colorful  racemes.  Plants  often 
differ  because  they  grow  on  volcanic  ash,  open 
disturbed  sites,  or  moist  depressions,  and  the  spe- 
cies exhibits  considerable  variation.  More  impor- 
tant, the  characters  used  to  separate  this  species 
from  C.  talmancensis  vary  greatly  within  and  be- 
tween populations.  Also,  the  two  species  may  hy- 
bridize in  the  Talamanca  Mountains,  and  it  ap- 
pears that  the  separation  of  the  two  species  may 
be  an  arbitrary  distinction.  This  problem  is  worthy 
of  careful  study  in  the  field.  The  name  gallito  has 
been  recorded  for  this  species. 


Castilleja  lentii  N.  Holmgren,  Brittonia  30:  191. 
1978.  Figure  7. 

Erect  herbs,  20-60  cm  tall,  simple  or 
branched,  leafy  stems  0.7-3  mm  diam.,  with  thin 
whitish  hairs  to  0.8  mm  long  or  glabrescent,  terete 
and  with  longitudinal  ridges.  Leaves  sessile,  often 
slightly  thickened  (articulated)  at  the  base;  leaf 
blades  (1.4-)3-6  cm  long,  2-7  mm  wide,  linear- 
oblanceolate  to  narrowly  elliptic,  apex  acute  or 
slightly  rounded,  distal  half  of  the  margin  with  1- 
4  short  (0.7-3  mm)  pairs  of  lateral  lobes,  gradu- 
ally narrowed  to  the  cuneate  base,  drying  black, 
mostly  glabrous  above,  with  thin  whitish  hairs  be- 
neath, tripliveined  from  near  the  base.  Inflores- 
cences 4-9  cm  long,  terminal  1 -sided  racemes, 
proximal  bracts  leaf-like  and  green,  distal  bracts 
oblanceolate  to  obovate  and  distally  reddish,  ped- 
icels 3-12  mm  long,  glabrous  or  pubescent.  Flow- 
ers with  calyx  12-24  mm  long,  4-6  mm  wide, 
glabrous,  adaxial  cleft  2-4  mm  deep,  abaxial  cleft 
12-17  mm  deep,  lobes  usually  entire,  pale  green 
below,  yellow  or  reddish  distally;  corolla  27-33 
mm  long,  tube  11-15  mm  long,  curved,  galeate 
upper  lip  12-18  mm  long,  1-2  mm  wide,  green 
above  and  reddish  along  the  sides,  lower  lip  with 
short  (1  mm)  teeth.  Fruits  9-12  mm  long,  4-5 
mm  diam.,  narrowly  obovoid;  seeds  not  seen. 

Plants  of  open  sunny  wet  sites  or  moist  cliff- 
sides  in  wet  evergreen  cloud  forest  formations. 


1400-1700  m  elevation.  Flowering  material  has 
been  collected  in  April,  August,  and  November. 
The  species  is  only  known  from  near  Cachi  (Car- 
tago  Prov.)  and  the  Rio  Cascajal  (San  Jose  Prov.). 
The  species  is  endemic  to  the  Caribbean  slope  of 
central  Costa  Rica. 

Castilleja  lentii  is  recognized  by  its  lower  ele- 
vation habitat  (among  our  species  of  Castilleja), 
the  alternate  oblanceolate  leaves  with  short  pin- 
nate distal  lobes,  glabrous  sepals,  and  long  slender 
upper  corolla  lip  reddish  along  the  lateral  margins. 

Castilleja  quirosii  Standl.,  Publ.  Field  Mus.  Nat. 
Hist.,  Hot.  Sen  18:  1104.  1938.  C.  aurantiaca 
Pennell,  Ann.  Missouri  Hot.  Card.  27:  338. 
1940.  C.  chiriquiensis  Pennell,  loc.  cit.  338. 
1940.  C.  seibertii  Pennell,  loc.  cit.  339.  1940. 
C.  bicolor  Pennell,  loc.  cit.  340.  1940.  Figure  7. 

Herbs  or  subshrubs,  30-90  cm  tall,  main  stems 
with  few  to  many  branches,  leafy  stems  0.7-3  mm 
diam.,  terete  or  with  poorly  defined  longitudinal 
ridges,  densely  hirsutulous  with  straight  or  curved 
hairs  0.2-0.5  mm  long.  Leaves  sessile,  a  petiole 
not  clearly  differentiated,  slightly  thickened  at  the 
base  (abaxially);  leaf  blades  6-30(-40)  mm  long, 
0.7-3  mm  wide  (not  including  the  lobes),  linear 
to  linear-oblong,  rounded  at  the  apex,  margin  en- 
tire or  with  1  or  2  pairs  of  short  ( 1-2  mm)  narrow 
(0.3-0.7  mm)  lobes  distally,  base  parallel-cuneate, 
drying  dark  grayish  green,  densely  to  sparsely  pu- 
bescent with  stiff  hairs  ca.  0.2  mm  long,  often 
scabrous,  venation  obscure.  Inflorescences  4-12 
cm  long,  1 -sided  racemes,  rachis  and  bracts  mi- 
nutely (0.3  mm)  puberulent,  bracts  ca.  25  mm 
long,  1-2  mm  wide,  proximal  bracts  leaf-like, 
marked  with  red  or  orange,  pedicels  2-12  mm 
long,  puberulent  (sometimes  with  gland-tipped 
hairs).  Flowers  with  calyx  17-20  mm  long,  3—5 
mm  wide,  minutely  puberulent,  red  to  orange  or 
yellowish  at  the  tip,  adaxial  cleft  1-3  mm  deep, 
abaxial  cleft  11-16  mm  deep;  corolla  (18-)22- 
28(-32)  mm  long,  yellow  to  yellowish  green  with 
red  margin,  tube  7-10  mm  long,  curved,  galeate 
upper  lip  15-23  mm  long,  ca.  1.5  mm  wide, 
densely  puberulent  on  the  dorsal  (adaxial)  margin; 
anthers  ca.  1.7  mm  long.  Fruits  8-13  mm  long, 
3-6  mm  diam.,  ovoid-oblong,  terminated  by  the 
persisting  style  base. 

Plants  of  open  sunny  sites  or  partly  shaded  sites 
in  evergreen  montane  and  open  paramo  forma- 
tions, 1600-3500  m  elevation.  The  species  ap- 
pears to  do  best  in  volcanic  soils  and  burnt  areas. 
Flowering  and  fruiting  throughout  the  year.  The 


34 


FIELDIANA:  BOTANY 


species  ranges  from  Volcan  Irazu  eastward  to  the 
Chiriqui  highlands  of  Panama. 

Castilleja  quirosii  is  recognized  by  its  generally 
dense  .short  pubescence,  small  alternate  linear 
leaves  with  few  short  distal  lobes,  colorful  flowers 
in  one-sided  racemes,  and  unusual  corollas. 
Among  our  species  of  Castilleja,  this  species  has 
the  widest  altitudinal  range  within  Costa  Rica,  al- 
though it  is  not  frequently  collected.  This  species 
and  its  close  congeners  should  be  studied  in  the 
field;  it  seems  possible  that  C.  quirosii  might  be 
included  within  a  more  broadly  defined  C.  ira- 
suensis. 

Castilleja  talamancensis  N.  Holmgren,  Brittonia 
30:  187.  1978. 

Herbs  and  shrubs,  erect  or  clambering,  0.1- 
1 .5(-2)  m  tall,  simple  or  with  many  distal  branch- 
es, woody  at  the  base,  stems  0.8-3.5  mm  diam., 
glabrous  or  minutely  (0.1  mm)  puberulent,  with 
prominent  longitudinal  ridges.  Leaves  sessile  or 
subsessile,  slightly  articulated  at  the  base,  bract- 
like  at  the  base  of  the  inflorescence;  leaf  blades 
5-18(-25)  mm  long,  0.5-2  mm  wide,  linear  to 
linear-oblong,  often  with  short  distal  lateral  lobes 
0.5-1.5  mm  long,  apex  rounded,  margins  entire  or 
with  small  lobes  near  the  apex,  base  decurrent  on 
the  stem,  drying  black,  glabrous  or  minutely  (0.05 
mm)  puberulent  beneath,  venation  obscure.  Inflo- 
rescences 2-8  cm  long,  1 -sided  racemes,  rachis 
minutely  hispidulous,  proximal  bracts  leaf-like, 
distal  bracts  obovate  with  rounded  apex,  marked 
with  red,  purple,  or  green,  pedicels  2-1 1  mm 
long,  0.2-0.4  mm  diam.,  with  erect  thin  hairs  0.1- 
0.2  mm  long.  Flowers  with  calyx  14-20  mm 
long,  3-4  mm  diam.,  adaxial  cleft  1-3.5  mm 
deep,  abaxial  cleft  9-17  mm  deep,  rounded  at  the 
apices,  minutely  puberulent  along  the  veins,  red 
to  brownish  purple  with  yellow  distal  margin;  co- 
rolla 20-32  mm  long,  tube  7-12  mm  long,  hood- 
ed upper  lip  13-22  mm  long,  green  above,  mar- 
gins red  to  orange,  lower  lip  reduced  with  3  nar- 
row teeth  ca.  1  mm  long.  Fruits  6-9  mm  long, 
4-5  mm  diam.,  dark  brown  and  smooth;  seeds  ca. 
2  mm  long,  C-shaped. 

Plants  of  open  or  partly  shaded  sites  in  high- 
montane  forest  and  paramo  formations,  2600- 
3800  m  elevation.  Flowering  and  fruiting  through- 
out the  year.  The  species  is  endemic  to  the  Cor- 
dillera de  Talamanca  of  Costa  Rica. 

Castilleja  talamancensis  is  recognized  by  its  re- 
striction to  higher  elevations,  narrow  linear-ob- 
long leaves  that  may  have  three  small  lobes  at  the 


apex,  and  the  colorful  inflorescences.  The  distal 
bracts  subtending  undeveloped  flowers  are  often 
bright  red.  Plants  in  protected  sites  become  large 
and  shrubby,  while  those  on  exposed  ridges  or 
open  sites  are  small  and  have  few  branches.  This 
species  is  particularly  successful  along  the  dis- 
turbed margins  of  the  Carretera  Interamericana. 
However,  there  is  so  much  variation  in  this  spe- 
cies, and  within  C.  irasuensis  and  C.  quirosii  as 
well,  that  there  is  the  strong  possibility  that  all 
three  would  be  better  considered  a  single  variable 
species,  with  C.  irasuensis  having  priority.  Care- 
ful field  work  with  the  living  populations  should 
help  resolve  this  problem.  The  name  ./for  de  Indio 
has  been  used  for  this  species. 

Castilleja  tayloriorum  N.  Holmgren,  Brittonia 
30:  193.  1978. 

Slender  herbs,  annual  or  short-lived  perennials, 
erect,  25-150  cm  high,  stems  0.4-1.5  mm  diam., 
glabrous  or  with  few  minute  (0.1  mm)  appressed 
whitish  hairs.  Leaves  sessile  or  appearing  to  have 
winged  petioles  (to  the  first  lobes),  deeply  pin- 
natisect  from  a  narrow  (0.7-2  mm)  rachis;  leaf 
blades  5-20  mm  long,  3-14  mm  wide,  with  a 
slender  (petiole-like)  proximal  half  and  the  distal 
rachis  bearing  2  or  3  pairs  of  narrow  (1-2  mm) 
lateral  lobes,  the  proximal  lobes  often  with  small- 
er distal  lobes,  apices  rounded,  margin  entire  and 
revolute,  drying  dark  grayish,  glabrous,  venation 
obscure.  Inflorescences  3-7  cm  long,  racemes 
(often  1 -sided),  lower  bracts  leaf-like  and  green, 
distal  bracts  shorter  and  reddish,  often  deciduous, 
pedicels  3-8  mm  long,  ca.  0.4  mm  diam.,  mi- 
nutely puberulent.  Flowers  with  calyx  13-17  mm 
long,  glabrous,  adaxial  cleft  4-6.5  mm  deep,  ab- 
axial cleft  8-12  mm  deep,  major  lobes  usually 
rounded  and  yellowish;  corolla  22-25  mm  long, 
curved,  tube  8-12  mm  long,  ca.  1  mm  diam., 
curved,  galeate  upper  lip  13-15  mm  long,  finely 
puberulent  and  green  dorsally.  margins  reddish, 
lower  lip  short.  Fruits  7-10  mm  long,  ca.  4  mm 
wide,  obovoid  or  oblong  with  abruptly  truncated 
apex. 

Plants  of  open  sites  or  cliffsides  in  wet  ever- 
green cloud  forest  formations,  1 100-1600  m  ele- 
vation. Flowers  were  collected  in  August  and  Oc- 
tober, fruiting  in  October.  The  species  is  known 
from  only  two  collections:  G6mez-Laurito  10180, 
from  near  the  highway  tunnel  at  Zurqui,  and  J. 
Taylor  &  C.  Taylor  11901,  from  below  Bajo  La 
Hondura.  Presently  the  species  is  only  known 
from  the  upper  Rio  Zurqui  drainage  in  Braulio 


BURGER:  FLORA  COSTARICENSIS 


35 


Carrillo  National  Park  on  the  Caribbean  slope  of 
central  Costa  Rica. 

Castilleja  tayloriorum  is  recognized  by  its  low- 
er elevation  habitat,  small  pinnatifid  leaves  with 
slender  lateral  lobes,  glabrous  calyx,  slender 
curved  corolla  tube,  and  long  narrow  upper  lip. 
Castilleja  tayloriorum  is  probably  most  closely 
related  to  species  of  section  Epichroma  from 
south-central  Mexico. 


Cymbalaria  Hill 

REFERENCES — G.  Cufodontis,  Die  Gattung 
Cymbalaria  Hill.  Nachtrage  und  Zusammenfas- 
sung.  Bot.  Not.  1947:  135-156.  1947.  D.  Sutton, 
A  Revision  of  the  Tribe  Antirrhineae.  British  Mu- 
seum (Natural  History)  &  Oxford  Univ.  Press. 
1988. 

Weak-stemmed  herbs,  creeping  or  twining,  pe- 
rennials, simple  or  much-branched.  Leaves  op- 
posite or  alternate,  simple,  long-petiolate,  leaf 
blades  reniform  to  suborbicular,  palmately  lobed 
or  toothed,  venation  palmate.  Inflorescences  ra- 
cemes, cymes,  or  solitary  flowers  in  leaf  axils, 
pedicels  well  developed,  elongating  in  fruit,  brac- 
teoles  absent  at  the  base  of  the  calyx.  Flowers 
small,  calyx  united  only  at  the  base,  sepals  5,  im- 
bricate (valvate)  in  bud,  enlarging  slightly  in  fruit; 
corolla  strongly  bilabiate,  usually  blue  or  purple, 
tube  with  a  short,  backward-pointing  abaxial  spur 
at  the  base,  upper  lip  with  2  lobes,  lower  lip  with 
3  lobes  and  elevated  transverse  ridge  (upper  pal- 
ate) restricting  access  to  the  throat;  stamens  4,  of 
2  unequal  pairs,  included,  thecae  separate  and  par- 
allel; ovary  2-locular,  ovules  many,  stigma  2- 
lobed.  Fruits  dry  capsules,  opening  by  2  pore-like 
slits,  the  2  valves  often  splitting  into  3;  seeds  el- 
lipsoid or  crested,  rugulose. 

A  genus  of  nine  species  native  to  western  Eu- 
ropean and  the  Mediterranean.  It  is  classified  in 
the  tribe  Antirrhineae,  and  its  species  were  once 
placed  in  Linaria,  but  they  differ  in  having  soli- 
tary axillary  flowers  and  leaves  with  palmate  ve- 
nation. Several  species  are  used  in  ornamental 
horticulture  as  groundcover  and  in  hanging  bas- 
kets. One  adventive  species  is  sometimes  found 
on  moist  walls  in  Mexico  and  Central  America. 

Cymbalaria  muralis  P.  Gaertn.,  B.  Meyer  &  J. 
Sherb.,  Oekon.  Fl.  Wetterau  2:  397.  1800.  An- 
tirrhinum cymbalaria  L.,  Sp.  PI.,  612.  1753. 


Linaria  cymbalaria  (L.)  Miller,  Gard.  Diet.,  ed. 
8,  #17.  1768. 

Herbs  with  slender  creeping  or  pendent  stems, 
often  rooting  from  the  nodes,  internodes  0.5-10 
cm  long,  0.5-1.7  mm  diam.,  glabrous.  Leaves 
mostly  alternate,  glabrous,  petioles  10-45  mm 
long,  0.3-0.5  mm  diam.;  leaf  blades  10-35  mm 
long,  9-40  mm  wide,  orbicular-oblate  to  reniform 
in  outline,  apex  obtuse  to  rounded  with  apiculate 
tip,  margins  entire  with  usually  5  (3,  7)  rounded 
lobes  separated  by  short  narrow  sinuses,  base  sub- 
cordate  to  deeply  cordate,  drying  dark  greenish  or 
grayish,  with  usually  3  major  veins.  Inflorescenc- 
es of  solitary  axillary  flowers,  pedicels  1 2-40  mm 
long,  0.4-0.7  mm  diam.,  glabrous,  elongating  to 
6  cm  and  negatively  phototropic  after  anthesis. 
Flowers  glabrous  externally,  sepals  0.8-1.5  mm 
long,  0.5-0.8  mm  wide,  lanceolate  to  narrowly 
oblong,  elongating  to  2.5  mm  long  in  fruit;  co- 
rolla 7-9  mm  long,  lilac  or  pale  violet  to  blue 
with  yellow  interior,  tube  3.5-4  mm  long  (includ- 
ing the  spur  to  2  mm  long),  lower  lip  ca.  3  mm 
long.  Fruits  2.7-4  mm  long,  3-4  mm  wide,  glo- 
bose or  rounded-oblong,  thin-walled. 

Cymbalaria  muralis,  as  its  Latin  epithet  im- 
plies, is  usually  found  on  damp  shaded  walls.  A 
native  of  Europe,  this  species  has  only  been  col- 
lected in  San  Jose  and  Tres  Rios  (1100-1300  m) 
in  Costa  Rica.  The  slender  stems,  small  thin-tex- 
tured five-lobed  cordate  leaves,  and  solitary  flow- 
ers with  strongly  bilabiate  bluish  corolla  and 
backward-oriented  spur  make  these  plants  quite 
distinctive.  In  addition,  the  pedicels  are  negatively 
phototropic  after  anthesis,  moving  the  capsules 
into  dark  corners  as  they  elongate.  They  are  some- 
times planted  as  cover  plants  in  gardens  and  are 
called  Kenilworth  ivy,  Coliseum  ivy,  and  penny- 
wort. 


Darcya  Turner  &  Cowan 

REFERENCE — B.  Turner  &  C.  Cowan,  Darcya 
(Scrophulariaceae),  a  new  genus  from  Central  and 
South  America.  Phytologia  74:  267-270.  1993. 

Herbs  or  weak-stemmed  subshrubs  to  1  m  tall, 
annual  or  perennial,  glabrous  or  puberulent,  dry- 
ing greenish  to  dark  brown.  Leaves  opposite,  sim- 
ple, petiolate,  blades  with  serrulate  margins,  ve- 
nation pinnate  or  subpalmate  with  3-5  major 
veins  from  near  the  base.  Inflorescences  terminal 
racemes  with  2  flowers/node  separated  by  well- 


36 


FIELDIANA:  BOTANY 


developed  internodes  (flowers  solitary  in  axils  of 
distal  leaves  or  bracts),  pedicels  slender,  bracte- 
oles  absent  at  the  base  of  the  calyx.  Flowers  with 
calyx  deeply  5-parted,  calyx  lobes  equal  or  sub- 
equal,  valvate  in  bud,  enlarging  slightly  in  fruit; 
corolla  tubular  and  strongly  2-lipped  (bilaterally 
symmetric),  white  to  blue  or  purple,  tube  shorter 
than  the  lips,  lower  (abaxial)  lip  3-lobed  and  larg- 
er, upper  (adaxial)  lip  2-lobed;  stamens  4,  sube- 
qual  or  unequal,  filaments  borne  at  2  levels  on  the 
corolla  tube,  anthers  with  straight  white  hairs  on 
dorsal  side;  ovary  ovoid,  style  short.  Fruits  thin- 
walled  capsules,  dehiscence  loculicidal  and  sep- 


ticidal  producing  4  valves  (or  irregular);  seeds 
small,  oblong  to  trapezoidal,  testa  reticulate. 

Darcya  is  a  genus  of  three  narrowly  endemic 
species,  restricted  (respectively)  to  Costa  Rica, 
Panama,  and  Colombia.  These  plants  resemble 
species  of  Stemodia  but  differ  in  having  pubescent 
anthers,  very  short  styles,  more  clearly  racemose 
inflorescences,  three  or  five  prominent  veins  from 
the  base  of  the  blade,  and  oblong  trapezoidal 
seeds.  The  decurrent  leaf  bases  merge  with  the 
longitudinal  ridges  (angles)  of  the  stem  in  a  dis- 
tinctive manner.  This  genus  also  resembles  the 
Asiatic  genus  Limnophila.  which  has  become  an 
invasive  weed  in  some  areas. 


Key  to  the  Species  of  Darcya 

la.  Calyx  and  pedicels  glabrous;  leaf  blades  15-45  mm  long,  ovate-triangular  to  ovate-elliptic;  stamens 
4,  subequal;  plants  of  the  wet  Caribbean  slope  in  central  Costa  Rica D.  costaricensis 

Ib.  Calyx  and  pedicels  puberulent;  leaf  blades  17-60  mm  long,  ovate-lanceolate;  stamens  4  in  2  unequal 
pairs;  collected  in  the  area  of  Boquete,  Chiriquu  Panama D.  reliquiarum 


Darcya  costaricensis  (B.  L.  Turner)  B.  L.  Turner, 
Phytologia  74:  268.  1993.  Stemodia  costaricen- 
sis B.  L.  Turner,  Phytologia  73:  253.  1992.  Fig- 
ure 3. 

Erect  or  sprawling  herbs  to  1  m  tall  with  many 
internodes  3-6  cm  long,  leafy  stems  0.7-2.5  mm 
diam.,  glabrous,  terete  but  with  4  longitudinal 
ridges,  nodes  without  an  interpetiolar  ridge. 
Leaves  opposite,  petioles  2-10  mm  long,  0.4-1.2 
mm  wide,  lateral  margins  decurrent  on  the  stem 
and  continuous  with  the  stem  ridges;  leaf  blades 
15-45  mm  long,  10-25  mm  wide,  ovate-triangu- 
lar to  ovate-elliptic,  apex  acute,  margin  with  4-6 
teeth/cm,  often  revolute  (dried),  base  obtuse  to 
truncate,  minutely  punctate,  glabrous  or  with  few 
minute  (0.1  mm)  hairs  beneath,  subpalmate  with 
3  prominent  basal  veins,  2°  veins  2  or  3/side.  In- 
florescences 5-10  cm  long,  glabrous  racemes, 
basal  flower  pair  subtended  by  reduced  leaves 
(12-16  mm  long),  distal  flowers  subtended  by  ses- 
sile lanceolate  or  linear  bracts  2-6  mm  long,  ped- 
icels 6-16  mm  long,  glabrous.  Flowers  glabrous 
externally,  calyx  lobes  2.5-3  mm  long,  to  4  mm 
in  fruit,  0.3-0.5  mm  wide,  narrowly  oblong,  apex 
truncated  and  thickened,  with  3  prominent  parallel 
veins;  corolla  4-7  mm  long,  blue  or  purple,  tube 
3-4  mm  long,  puberulent  internally  near  the 
mouth,  central  lobe  of  lower  lip  3-6  mm  long; 
stamens  4,  subequal,  anthers  similar;  stigma  in- 


cluded. Fruits  3-4  mm  long,  1.5-2  mm  wide, 
narrowly  ovate  to  oblong,  persisting  style  0.4-0.8 
mm  long;  seeds  0.5-0.7  mm  long,  yellowish 
brown,  reticulate  with  longitudinal  ridges  separat- 
ed by  rounded  pits. 

Plants  of  open  sites  and  along  stream  edges  in 
very  wet  lower  montane  forest  formations  of  the 
Caribbean  slope,  1400-1600  m  elevation.  Flow- 
ering and  fruiting  throughout  the  year.  This  spe- 
cies is  known  only  from  along  the  upper  Rio 
Grande  de  Orosf,  Tapantf  Refuge,  Cartago  Prov- 
ince, central  Costa  Rica. 

Darcya  costaricensis  is  recognized  by  its  weak- 
stemmed  herbaceous  habit,  opposite  leaves  with 
broad  serrulate  blades,  racemose  inflorescences, 
narrow  calyx  lobes,  two-lipped  blue  or  purple  co- 
rollas with  short  tube,  and  restricted  geographical 
range.  This  material  had  earlier  been  placed  in 
Stemodia  reliquiarum  (see  the  following  species). 

Darcya  reliquiarum  (D'Arcy)  B.  L.  Turner  &  C. 
C.  Cowan,  Phytologia  74:  269.  1993.  Stemodia 
reliquiarum  D'Arcy,  Ann.  Missouri  Bot.  Card. 
66:  258.  1979. 

Herbs  with  sprawling  stems  to  40  cm  tall,  leafy 
stems  0.7-3  mm  diam.,  with  2  prominent  ridges, 
glabrous  except  near  the  nodes  with  erect  5-celled 
hairs,  glabrescent,  drying  brown.  Leaves  oppo- 
site, petioles  0-4(-8)  mm  long,  with  narrow  lat- 


BURGER:  FLORA  COSTARICENSIS 


37 


eral  margins  dccurrent  on  the  stem  and  poorly  dif- 
ferentiated from  the  blade;  leaf  blades  1 .7-6  cm 
long,  10-26  mm  wide,  ovate-lanceolate  to  nar- 
rowly ovate,  apex  acute,  margins  serrulate  distally 
with  10-15  teeth/side,  base  obtuse  or  truncate, 
punctate  on  both  surfaces,  glabrous,  subpalmate 
with  3  major  veins  from  the  base,  midvein  with  2 
or  3  lateral  veins/side.  Inflorescences  terminal 
open  racemes,  rachis  0.5-0.8  mm  diam.,  bracts 
leaf-like  to  scale-like,  mostly  linear  (ca.  3  X  1 
mm),  ciliolate  along  the  edge  with  multicellular 
gland-tipped  hairs  0.1-0.3  mm  long,  pedicels  4- 
12  mm  long,  puberulent.  Flowers  small,  calyx 
lobes  2-4  mm  long,  ca.  0.5  mm  wide,  minutely 
puberulent  externally,  with  3  longitudinal  veins, 
drying  brown;  corolla  4-6  mm  long,  salverform, 
blue  to  lavender  or  purple,  tube  3-4  mm  long, 
slightly  exceeding  the  calyx,  subglabrous  exter- 
nally, lobes  2-3  mm  long,  subequal,  minutely  pu- 
berulent at  the  base  within;  stamens  4,  the  lower 
2  filaments  ca.  0.5  mm  long  and  with  reduced 
thecae.  the  upper  filaments  0.7-1  mm  long,  in- 
serted near  the  middle  of  the  tube,  one  theca  ses- 
sile on  the  connective  with  the  other  on  a  stipe- 
like  arm  of  the  connective;  ovary  conical,  style 
0.5  mm  long,  stigma  club-like.  Fruits  4-5  mm 
long,  2.4-3  mm  wide,  dehiscing  apically  into  4 
valves,  placenta  broad,  unwinged;  seeds  many, 
0.4-0.6  mm  long,  oblong  to  trapezoidal,  longitu- 
dinally reticulate,  dark  brown. 

Plants  of  evergreen  montane  cloud  forests 
around  Boquete,  1500-2500  m  elevation.  Proba- 
bly flowering  throughout  the  year.  Endemic  to  the 
Chiriqui  highlands  of  western  Panama. 

Darcya  reliquiarwn  is  recognized  by  its  her- 
baceous sprawling  habit,  opposite  serrulate  leaves 
with  subpalmate  venation,  racemose  puberulent 
inflorescences,  narrow  calyx  lobes,  strongly  2- 
lipped  blue  or  purple  corolla,  pubescent  anthers, 
and  restricted  geographical  range. 


Digitalis  Linnaeus 

REFERENCE — K.  Werner,  Zur  Nomenclature  und 
Taxonomie  von  Digitalis  L.  Bot.  Jahrb.  Syst.  79: 
218-254.  1960. 

Erect  herbs  or  rarely  shrubs,  biennial  or  peren- 
nial, stems  simple  or  branched  from  the  base. 
Leaves  alternate,  sessile  or  petiolate,  entire  to 
dentate,  pinnately  veined,  drying  greenish  or 
grayish.  Inflorescences  terminal  racemes,  the 
flowers  often  aligned  along  one  side,  bracts  sub- 


tending the  pedicels,  lacking  bracteoles  at  the  base 
of  the  calyx.  Flowers  large  and  colorful,  calyx 
with  5  sepals  united  only  near  the  base,  imbricate 
in  bud,  enlarging  slightly  in  fruit;  corolla  tubular 
to  campanulate,  longer  than  the  calyx,  purple  to 
yellow  or  white,  bilaterally  symmetric  with  the 
lower  lip  slightly  longer  than  the  upper,  upper  lip 
entire  or  2-cleft,  lower  lip  3-lobed,  lateral  lobes 
exterior  in  bud;  stamens  4  in  2  similar  pairs,  in- 
cluded within  the  tube,  anthers  with  2  divergent 
thecae;  ovary  2-locular,  ovules  many,  style  sim- 
ple, stigma  2-lobed.  Fruits  dry  capsules,  dehisc- 
ing septicidally;  seeds  many,  minute,  rugulose. 

Digitalis  is  a  genus  of  about  20  species  ranging 
from  western  Europe  into  central  Asia.  They  are 
classified  in  the  subtribe  Digitalieae  of  the  the 
tribe  Veroniceae  and  are  distinguished  by  their 
erect  habit,  one-sided  racemes,  and  large  tubular 
corollas.  The  common  foxglove  is  found  in  gar- 
dens and  as  an  escape  in  the  cooler  highlands  of 
Central  America. 

Digitalis  purpurea  L.,  Sp.  PI.  621.  1753.  Figure  8. 

Stout  herbs,  0.5-1.8  m  tall,  biennial,  leafy 
stems  2-8  mm  diam.,  minutely  puberulent  with 
thin  hairs  0.1-0.3  mm  long.  Leaves  becoming 
progressively  shorter  from  base  to  inflorescence, 
petioles  1-11  cm  long  but  not  well  differentiated 
from  the  blade,  2-12  mm  wide  with  winged  mar- 
gins; leaf  blades  2.5-20  cm  long,  1-8  cm  wide, 
narrowly  elliptic  to  ovate-elliptic  or  elliptic-ob- 
ovate,  apex  acute  to  obtuse,  margin  serrate  with 
5-10  teeth/cm,  gradually  narrowed  to  the  cuneate 
base,  minutely  puberulent  on  both  surfaces,  2° 
veins  3-5/side,  strongly  ascending.  Inflorescenc- 
es 15-40  cm  long,  flowers  mostly  along  1  side, 
bracts  8-18  mm  long,  lanceolate,  sessile,  pedicels 
5-10  mm  long,  0.4-0.7  mm  diam.,  puberulent. 
Flowers  with  calyx  8-16  mm  long,  sepals  sepa- 
rate to  the  base,  4-7  mm  wide,  broadly  ovate  to 
ovate-oblong,  ciliolate  along  the  edge;  corolla 
20-50  mm  long,  tube  1.3-2  cm  wide,  tubular- 
campanulate,  pink  to  purple  (white),  lower  lip  ex- 
tending 3-8  mm  beyond  the  upper  lip,  usually 
with  dark  spots  within  the  throat;  thecae  divergent 
and  equal,  ca.  3  mm  long.  Fruits  9-1 1  mm  long, 
9-10  mm  diam.,  ovoid  with  broadly  truncated 
base,  minutely  puberulent;  seeds  0.4-0.7  mm 
long,  narrowly  rectangular. 

Digitalis  purpurea,  native  to  Europe,  occurs  as 
a  garden  plant  and  in  small  naturalized  popula- 
tions in  high-montane  areas,  1800-3300  m  ele- 
vation. This  species  is  distinctive,  with  its  alter- 


38 


FIELDIANA:  BOTANY 


nate  leaves  gradually  narrowed  into  the  long  pet- 
iole, conspicuous  (often  one-sided)  racemes  of 
large  colorful  flowers,  and  the  open  corolla  tubes 
marked  with  dark  spots  within.  Many  varieties  are 
used  in  ornamental  horticulture.  The  plants  are 
poisonous  to  livestock  and  contain  a  number  of 
potent  compounds.  This  species  is  the  principal 
source  of  digitalin,  an  important  drug  in  treating 
some  kinds  of  heart  ailments.  Common  names  are 
digital,  manga  de  la  Senora,  and  foxglove. 


Escobedia  Ruiz  &  Pav6n 

REFERENCES — F.  Pennell,  Escobedia,  a  Neotrop- 
ical genus  of  Scrophulariaceae.  Proc.  Acad.  Nat. 
Sci.  Philadelphia  83:  411-426.  1931.  J.  Thieret, 
The  Scrophulariaceae — Buchnereae  of  Central 
America.  Ceiba  8:  92-101.  1961. 

Perennial  herbs,  stems  simple  or  branched,  gla- 
brous or  puberulent,  striate  or  angled,  roots  yel- 
low to  orange.  Leaves  opposite,  sessile  or  sub- 
sessile,  base  cuneate  to  clasping,  coriaceous,  en- 
tire to  serrulate,  venation  pinnate  to  subpalmate, 
often  scabrous.  Inflorescences  racemose  or  with 
solitary  flowers  in  axils  of  distal  leaves,  pedicels 
(peduncles)  with  2  bracteoles  distally  or  lacking 
bracteoles.  Flowers  large,  calyx  tubular  to  fun- 
nelform,  with  5-10  longitudinal  veins,  lobes  3-6, 
shorter  than  the  tube,  triangulate  to  rounded;  co- 
rolla salverform  with  a  long  slender  tube,  usually 
white,  externally  glabrous  or  puberulent,  lobes  5, 
small  to  large,  equal  and  rounded;  stamens  4,  sub- 
equal,  inserted  in  the  middle  of  the  tube,  filaments 
ciliate  or  glabrous,  anthers  glabrous  and  aristate; 
ovary  2-locular,  ovules  many,  style  elongate,  stig- 
mas linear  on  the  side  of  the  style  apex.  Fruits 
dry  capsules,  ellipsoid,  hard,  included  within  the 
persisting  calyx;  seeds  many,  narrowly  conical  to 
linear-oblong,  surface  reticulate. 

Escobedia  is  a  genus  of  six  to  eight  species 
ranging  from  Mexico  to  Bolivia  and  Brazil.  The 
genus  is  classified  in  the  tribe  Buchnereae,  sub- 
tribe  Melasmineae.  These  plants  are  distinctive 
among  Neotropical  Scrophulariaceae  because  of 
their  large  salverform  white  corollas  with  long 
narrow  tubes.  The  roots  have  been  used  by  indig- 
enous peoples  as  a  source  of  a  yellow  dye  for  food 
coloring.  Three  species  are  found  in  northern  Cen- 
tral America  (Standley  &  Williams,  1973),  but 
only  one  of  these  has  been  found  in  Costa  Rica. 
Escobedia  laevis  Schldl.  &  Cham,  (with  long  lin- 
ear leaves  and  calyx  4-7  cm  long  with  narrow 


calyx  lobes)  ranges  from  southern  Mexico  to  cen- 
tral Nicaragua. 

Escobedia  grandiflora  (L.f.)  O.  Kuntze,  Rev. 
Gen.  PI.  3:  231.  1893.  Buchnera  grandiflora 
L.f.,  Suppl.  PI.  287.  1781.  £.  scabrifolia  Ruiz 
&  Pav6n,  Syst.  Veg.  Peruv.  Chil.  159.  1798. 
Micalia  grandiflora  (L.f.)  Raf.,  Fl.  Tell.  2:  104. 
1837.  E.  curialis  Pennell.  Proc.  Acad.  Nat.  Sci. 
Philadelphia  83:  417.  1931.  E.  longiflora  Pen- 
nell, loc.  cit.  423.  1931.  E.  reticulata  Pennell. 
loc.  cit.  420.  1931.  Figure  6. 

Erect  perennial  herbs,  0.6-1. 5(-2)  m  tall,  leafy 
stems  1.5-7  mm  ilium.,  with  stiff  whitish  hairs 
0.1-0.3  mm  long,  smooth  or  scabrous,  roots 
bright  yellow-orange.  Leaves  opposite  or  subop- 
posite,  gradually  becoming  smaller  distally,  sub- 
sessile  with  petioles  1-4  mm  long,  ca.  2  mm 
wide;  leaf  blades  5-12  cm  long,  1.5-3  cm  wide, 
lanceolate  to  narrowly  triangular  or  ovate-elliptic, 
apex  acute,  margin  serrate  with  1-3  teeth/cm, 
base  rounded  and  truncate  to  auriculate.  drying 
pale  to  dark  gray  and  coriaceous,  scabrous  above 
and  below  with  short  (0.1-0.2  mm)  hairs,  tripli- 
veined  from  the  base  or  palmately  5-veined.  In- 
florescences of  solitary  flowers  (2/node)  in  axils 
of  smaller  distal  leaves  (or  the  inflorescences  ra- 
cemose if  the  smaller  leaves  are  interpreted  as 
bracts),  pedicels  11-35  mm  long,  0.7-2  mm 
diam.,  slightly  curved,  often  with  linear  bracteoles 
to  4  mm  long  in  the  upper  third.  Flowers  with 
calyx  34-44  mm  long,  6-10  mm  diam.,  scabrous, 
lobes  2-5  mm  long,  triangular;  corolla  7-12  cm 
long,  6-8  cm  wide  at  the  mouth,  salverform  with 
a  long  narrow  tube  to  10  cm  long,  3-8  mm  diam., 
lobes  1-3  cm  long  and  broadly  rounded,  white, 
minutely  puberulent;  stamens  inserted  on  the  mid- 
dle of  the  tube;  ovary  glabrous,  style  5-8  cm  long. 
Fruits  20-28  mm  long,  ca.  10  mm  wide,  smooth; 
seeds  3-4  mm  long,  0.3-0.5  mm  wide,  linear-rect- 
angular, translucent  except  for  the  center. 

Uncommon  plants  of  open  sites,  in  marshy  or 
wet  situations  of  lower  montane  evergreen  forest 
formations,  1000-1500  m  elevation.  Collected  in 
flower  in  July-February.  The  species  ranges  from 
Mexico  to  Brazil. 

Escobedia  grandiflora  is  recognized  by  its  op- 
posite scabrous  coriaceous  subsessile  leaves  with 
three  (five)  major  veins,  distal  nodes  with  two 
large  opposite  flowers,  and  large  white  corollas 
with  long  narrow  tubes  and  widely  flaring  limb. 
The  color,  form,  and  size  of  the  corolla  suggest 
nocturnal  pollination  by  hawkmoths.  Escobedia 


BURGER:  FLORA  COSTARICENSIS 


39 


reticiilata  (from  near  Canas  Gordas,  Pitiier  11118 
holotvpe  t  S)  was  distinguished  by  Pennell  and 
Thicrct  on  the  basis  of  a  pustulate  calyx  and  the 
size  and  position  of  the  bracteoles.  This  is  most 
likely  a  local  variation  and  not  deserving  of  spe- 
cies rank  (D'Arcy,  1979,  p.  220). 

Gerardia  species  are  now  placed  in  Agalinis 
(q.v.). 

Gibsoniothamnus  is  now  placed  in  the  Schle- 
geliaceae. 


Hemichaena  Bentham 

REFERENCE — J.  Thieret,  Synopsis  of  Hemichae- 
na, including  Berendtiella  (Scrophulariaceae). 
Fieldiana,  Bot.  34:  89-99.  1972. 

Herbs  or  shrubs,  simple  or  branched,  glabrous 
or  viscid-pubescent,  drying  brownish.  Leaves  op- 
posite or  fasciculate,  sessile  or  subsessile,  margins 
dentate,  surface  often  rugose,  venation  pinnate. 
Inflorescences  axillary,  of  solitary  flowers  or  1  or 
2  cymes/axil,  usually  pedunculate  with  bracts  sub- 
tending the  well-developed  pedicels,  bracteoles 
absent  at  the  base  of  the  calyx.  Flowers  showy, 
calyx  campanulate  to  tubular,  with  5  prominent 
longitudinal  costae,  5-lobed  or  5-toothed,  lobes 
unequal,  shorter  than  the  tube;  corolla  funnelform 
to  tubular-campanulate,  2-lipped,  bright  yellow  to 
orange  or  red,  tube  exceeding  the  calyx,  lobes 
equal  to  or  shorter  than  the  tube,  upper  lip  2-lobed 
or  emarginate,  lower  lip  3-lobed;  stamens  4,  sub- 
equal  or  of  2  unequal  pairs,  included  or  exserted, 
inserted  at  the  middle  or  on  the  lower  half  of  the 
tube,  anthers  2-thecous,  parallel  but  becoming  di- 
vergent, staminode  absent;  ovary  2-locular,  pla- 
centas bilamellate,  ovules  many,  style  slender, 
with  2  flattened  stigmatic  areas  or  lobes  at  the 
apex.  Fruits  dry  capsules,  ovoid  to  oblong,  lo- 
culicidally  dehiscent  and  secondarily  septicidal; 
seeds  very  many,  minute,  linear-oblong  to  fusi- 
form, testa  reticulate  with  thin  translucent  walls. 

A  genus  of  five  species,  ranging  from  northern 
Mexico  to  the  Talamanca  mountains  of  Costa 
Rica.  Hemichaena  is  classified  in  the  tribe  Do- 
dartieae,  with  Mimulus,  Mazus,  Leucocarpus,  and 
Berendtiella.  The  genus  was  monotypic,  but 
Thieret  (1972,  cited  above)  broadened  the  concept 
to  include  the  four  species  of  Berendtiella.  Ge- 
neric delimitations  within  the  tribe  are  problem- 
atic. If  characters  of  the  fruit  are  emphasized,  then 


Hemichaena  and  Berendtiella  appear  closely  re- 
lated, but  Hemichaena  can  be  distinguished  by  its 
ampliate  corolla  tube,  included  stamens  that  are 
attached  near  the  base  of  the  corolla,  and  cordate- 
amplexicaul  leaves.  The  generic  classification  of 
the  tribe  Dodartieae  will  remain  obscure  until  Mi- 
mulus is  revised  and  the  classification  of  the  entire 
group  is  compared.  Only  the  following  disjunct 
species  is  found  south  of  northern  Nicaragua. 

Hemichaena  fruticosa  Ik-nib..  PI.  Hartw.  78. 
1841.  Leucocarpus  fruticosus  (Benth.)  Benth. 
in  DC.,  Prodr.  10:  336.  1846. 

Herbs  or  subshrubs  to  2  m  tall,  distal  stems 
with  few  or  no  lateral  branches,  leafy  stems  2-1 1 
mm  diam.,  terete  or  slightly  quadrangular,  densely 
pubescent  with  gland-tipped  hairs  0.5-1.3  mm 
long.  Leaves  opposite,  sessile,  of  similar  size 
along  the  stems,  equal  or  subequal  at  the  node; 
leaf  blades  5-17  cm  long,  2-5  cm  wide,  lanceo- 
late to  narrowly  elliptic-lanceolate  or  oblanceo- 
late,  apex  acute  or  acuminate,  margin  serrate  with 
3—6  small  or  prominent  teeth/cm,  base  rounded 
and  auriculate  to  cordate-amplexicaul,  drying 
dark  above  and  paler  beneath,  minutely  glandular 
puberulent  with  hairs  0.2-0.6  mm  long,  2°  veins 
4-9/side.  Inflorescences  3-8  cm  long,  usually  of 
1  or  2  axillary  cymes  (1-4/node),  peduncles  10- 
28  mm  long,  bracts  6-22  mm  long,  1-5  mm  wide, 
lanceolate,  viscid  puberulent  with  hairs  ca.  0.5 
mm  long,  pedicels  5-18  mm  long.  Flowers  with 
calyx  11-16  mm  long,  4-6  mm  diam.,  densely 
puberulent  with  gland-tipped  hairs,  lobes  3-8  mm 
long,  narrowly  triangular  and  acute;  corolla  25- 
45  mm  long,  14-28  mm  wide  at  the  mouth,  tu- 
bular-campanulate, tube  6-11  mm  wide  at  the 
mouth,  minutely  puberulent  externally,  lobes  5-9 
mm  long;  stamens  subequal,  inserted  in  the  lower 
¥4  of  the  tube,  filaments  12-15  mm  long,  glabrous, 
anthers  3.3-3.8  mm  long;  style  15  mm  long,  stig- 
mas flattened.  Fruits  13-17  mm  long,  4-6  mm 
diam.,  oblong,  becoming  brown;  seeds  0.6-1  mm 
long,  0.3-0.4  mm  wide,  narrowed  at  each  end. 

Plants  of  open  moist  to  wet  sunny  sites  along 
streams,  on  steep  slopes,  and  in  disturbed  sites  in 
montane  evergreen  forest  formations,  1600-3100 
m  elevation.  Probably  flowering  and  fruiting 
throughout  the  year,  but  collected  most  often  in 
December-March.  The  species  is  found  in  south- 
ern Mexico  and  Guatemala  and  in  the  western 
portion  of  the  Cordillera  de  Talamanca. 

Hemichaena  fruitcosa  is  recognized  by  its  stout 
viscid  puberulent  stems,  sessile  opposite  serrate 


40 


FIELDIANA:  BOTANY 


leaves,  axillary  cymes,  bright  yellow  corollas  that 
are  only  slightly  bilabiate,  and  its  restricted  geo- 
graphical range  (in  Costa  Rica).  It  is  surprising 
that  this  species,  which  is  so  common  in  the  west- 
ern half  of  the  Cordillera  de  Talamanca,  has  not 
been  collected  elsewhere  in  Costa  Rica.  D'Arcy 
(1979)  placed  this  species  in  synonomy  under 
Leucocarpus  perfoliatus  (H.B.K.)  Benth.,  but  that 
species  has  smaller  flowers,  smaller  leaves,  and 
baccate  white  fruits. 


Lamourouxia  Kunth  in  H.B.K. 
Nomen  conservandum 

REFERENCE — W.  Ernst,  Floral  morphology  and 
systematics  of  Lamourouxia  (Scrophulariaceae: 
Rhinanthoideae).  Smithson.  Contrib.  Bot.  6:  1-63. 
1972. 

Perennial  herbs  or  subshrubs,  usually  woody  at 
the  base,  stems  erect  or  scandent,  distal  stems  of- 
ten few-branched  and  arched,  glabrous  or  pubes- 
cent, gland-tipped  hairs  often  present,  probably 
hemiparasitic  on  roots.  Leaves  opposite  (verticil- 
late),  sessile  or  short-petiolate,  often  smaller  and 
bract-like  distally,  blades  serrate  to  deeply  dis- 
sected, drying  brownish  or  black,  pubescent  with 
simple,  branched,  or  gland-tipped  hairs.  Inflores- 
cences of  solitary  flowers  in  distal  leaf  axils  (1  or 
2/node)  or  appearing  racemose  (paniculate  or  cor- 
ymb-like) when  distal  leaves  are  bract-like,  pedi- 
cels ebracteolate  at  the  apex.  Flowers  large  and 
showy,  calyx  tubular-campanulate,  4-lobed,  lobes 
subequal  or  of  2  unequal  pairs  (rarely  cleft  to 


base),  obtuse  to  deltoid  or  linear,  with  10  promi- 
nent longitudinal  veins;  corolla  tubular  and  bila- 
biate, red  to  orange  or  white,  usually  puberulent 
on  the  exterior,  upper  lip  often  erect  (rarely  hood- 
like),  entire  or  bilobed,  lower  lip  usually  shorter, 
narrow,  3-lobed,  biplicate  within  below  the  apex; 
stamens  4  in  2  subequal  pairs  or  with  2  fertile 
stamens  and  2  staminodes,  attached  near  base  of 
corolla  tube,  filaments  swollen  and  puberulent  at 
the  base,  included  in  the  upper  lip,  fertile  anthers 
basi  fixed  and  pilose;  ovary  2-locular,  ovules 
many,  stigma  terminal,  exserted  and  2-lipped. 
Fruits  dry  capsules,  ovoid  to  ellipsoid  with  per- 
sisting style-base,  opening  loculicidally  into  2  en- 
tire valves;  seeds  many,  ellipsoid  to  oblong,  mi- 
nutely bullate  to  reticulate. 

Lamourouxia  is  a  genus  of  ca.  24  species  found 
in  Neotropical  highlands  from  Mexico  to  Peru. 
The  center  of  species  richness  is  in  Mexico,  with 
only  three  species  found  in  southern  Central 
America  and  two  species  in  the  Andes.  Although 
a  very  distinctive  genus,  the  individual  species 
can  be  difficult  to  separate  from  each  other.  In 
Costa  Rica  the  plants  are  restricted  in  distribution 
and  not  often  collected.  The  large  showy  flowers 
with  bright  red  or  orange  narrowly  tubular  corol- 
las, spicate  or  racemose  inflorescences,  four-lobed 
calyx,  subsessile  leaves,  and  preference  for  open 
habitats  at  mid-elevations  help  distinguish  this  ge- 
nus. Our  species  belong  to  section  Hemispadon, 
according  to  Ernst's  fine  monograph  (1972,  cited 
above).  Lamourouxia  is  in  the  subfamily  Rhin- 
anthoideae. Every  member  of  this  subfamily  that 
has  been  tested  is  hemiparasitic,  perhaps  explain- 
ing why  these  lovely  plants  are  not  seen  in  gar- 
dens. 


Key  to  the  Species  of  Lamourouxia 

la.  Calyx  glabrous;  young  stems  glabrous  except  for  longitudinal  lines  of  minute  hairs;  leaves  linear- 
lanceolate;  1600-2800  m  elevation L.  lanceolata 

Ib.  Calyx  puberulent  externally;  young  stems  pubescent;  leaves  narrowly  elliptic  to  ovate-triangular; 
900-1800  m  elevation 

2a.  Stems  and  flowers  with  simple  (non-gland-tipped)  hairs,  leaves  7-24  mm  long;  flowering  mostly 
November-February L.  gutierrezii 

2b.  Stems  and  flowers  with  gland-tipped  hairs;  leaves  10-110  mm  long;  flowering  in  July-December 

.  L.  viscosa 


BURGER:  FLORA  COSTARICENSIS 


41 


Lamourouxia  gutierrezii  Oerst.  in  Benth.  & 
Ocrst.,  Vidcnsk,  Meddel.  Dansk  Naturhist.  For- 
en.  Kjobenhavn  1853:  29.  1853.  L  scabra  See- 
mann,  Bot.  Voy.  Herald  177:  pi.  33.  1854.  Fig- 
ure 5. 

Herbs  or  subshrubs,  0.5-1. 3(-2)  m  tall,  distal 
branches  arched  and  often  with  short  lateral 
branching,  leafy  stems  1-4  mm  diam.,  sparsely  to 
densely  puberulent  with  thin  straight  or  retrorse 
hairs  0.2-0.5  mm  long.  Leaves  opposite  (verticil- 
late),  sessile  or  subsessile  with  poorly  differenti- 
ated petioles  to  2  mm  long;  leaf  blades  7-24  mm 
long,  2-10  mm  wide,  elliptic  to  elliptic-oblong  or 
ovate-elliptic,  apex  obtuse  or  rounded,  margin 
with  4-6  rounded  teeth/cm,  base  cuneate,  drying 
dark  grayish  green,  slightly  scabrous  above  and 
below  with  short  stiff  hairs  ca.  0.2  mm  long,  2° 
veins  3-5/side.  Inflorescences  5-20  cm  long,  spi- 
cate  or  racemose,  bracts  8-12  mm  long,  5-6  mm 
wide,  leaf-like  and  lanceolate  to  ovate,  sometimes 
caducous  and  the  flowers  appearing  ebracteate, 
pedicels  2-5  mm  long,  ca.  0.7  mm  diam.,  puber- 
ulent. Flowers  with  calyx  6-9  mm  long,  enlarging 
to  12  mm  in  fruit,  2-4  mm  diam.,  cupulate,  5- 
lobed,  lobes  2-4  mm  long,  triangular  to  lanceo- 
late; corolla  30-44  mm  long,  4-6  mm  diam.,  tu- 
bular, bright  red  to  orange-red,  densely  puberulent 
externally,  upper  lip  ca.  14  mm  long;  functional 
stamens  2,  anthers  connivent  and  positioned 
against  the  upper  lobe,  3-4  mm  long  with  hairs 
1-2  mm  long;  ovary  glabrous,  style  2-4  cm  long. 
Fruits  8-13  mm  long,  4-8  mm  diam.,  ovoid, 
short-beaked,  dark  and  smooth;  seeds  0.8-1  mm 
long,  0.3-0.4  mm  thick. 

Infrequently  collected  plants  of  open  sites  in 
lower  montane  evergreen  forest  formations,  900- 
1900  m  elevation.  Probably  flowering  throughout 
the  year,  with  most  collections  made  in  Novem- 
ber-February and  June-July.  The  species  ranges 
from  the  western  part  of  the  Meseta  Central  in 
Costa  Rica  to  the  Chiriqui  highlands  in  Panama. 

Lamourouxia  gutierrezii  is  recognized  by  its 
smaller  narrow  scabrous  leaves,  simple  pubes- 
cence, and  puberulent  calyx.  The  structure  and 
position  of  the  staminodes  suggest  that  they  may 
function  as  a  barrier  to  keep  visiting  insects  from 
the  ovary.  This  is  the  only  species  of  the  genus 
endemic  to  southern  Central  America. 

Lamourouxia  lanceolata  Benth.  ///  DC.,  Prodr. 
10:  542.  1846.  L.  longiflora  var.  lanceolata 
(Benth.)  L.  O.  Williams,  Fieldiana,  Bot.  34: 
121.  1972.  Figure  5. 


Herbs  or  shubshrubs  to  1 .5(-3)  m  tall,  erect  or 
scandent,  stems  often  arching  and  with  multiple 
branching,  leafy  stems  1-3  mm  diam.,  with  mi- 
nute (0.2  mm)  crooked  hairs  in  2  opposing  lon- 
gitudinal lines  along  the  stems.  Leaves  sessile  or 
subsessile  with  poorly  defined  petioles  to  2  mm 
long,  not  conspicuously  smaller  distally;  leaf 
blades  1 1-48(-57)  mm  long,  2-8(-14)  mm  wide, 
narrowly  lanceolate  to  narrowly  linear-lanceolate 
or  narrowly  elliptic-oblong,  apex  acute,  margin 
with  4-6  teeth/cm,  base  narrowly  cuneate,  drying 
brownish  to  blackish,  glabrous  or  with  minute 
(0.1  mm)  papillate-puberulent  hairs  on  the  mid- 
vein  above,  2°  veins  4-7/side.  Inflorescences  3- 
15  cm  long  or  of  flowers  axillary  to  distal  leaves, 
flowers  subtended  by  leaves  very  similar  to  those 
lower  on  the  stems,  pedicels  3-9(-14)  mm  long, 
0.4-0.8  mm  diam.,  glabrous  or  with  a  line  of  hairs 
along  one  side.  Flowers  with  calyx  7-18  mm 
long,  3-5  mm  diam.,  glabrous  externally,  veins 
elevated,  lobes  6-15  mm  long  and  becoming  re- 
flexed;  corolla  25-47  mm  long,  4-7  mm  diam., 
tubular,  bright  red  to  red-orange,  densely  puber- 
ulent externally,  upper  lip  14-18  mm  long;  sta- 
mens 2,  with  larger  pubescent  anthers  often  co- 
herent, staminodes  2,  borne  above  the  fertile  sta- 
mens, thickened  at  the  apex.  Fruits  8-14  mm 
long,  5-8  mm  diam.,  ovoid-rounded,  beaks  1-2 
mm  long;  seeds  1-1.3  mm  long,  oblong  to  wedge 
shaped,  surface  reticulate. 

Infrequently  collected  plants  of  open  sites  in 
evergreen  montane  forest  formations,  1600- 
2800(-3200)  m  elevations.  Flowering  in  Decem- 
ber-March. In  Costa  Rica  this  species  is  known 
only  from  the  western  portion  of  the  Cordillera  de 
Talamanca.  The  species  ranges  from  central  Mex- 
ico to  Costa  Rica. 

Lamaourouxia  lanceolata  is  recognized  by  its 
narrow  opposite  leaves,  glabrous  calyx  with  re- 
flexed  lobes,  narrow  bright  red  corolla  tubes,  an- 
droecium  of  two  stamens  with  pubescent  anthers 
and  two  staminodes,  and  higher  elevation  habitats. 
This  species  has  been  confused  with  L.  longiflora 
Benth.  (Williams,  1972;  Standley  &  Williams, 
1973),  but  L.  longiflora  has  four  fertile  stamens 
and  the  calyx  is  puberulent  on  the  exterior. 

Lamourouxia  viscosa  Kunth  in  H.B.K.,  Nov. 
Gen.  Sp.  2  ed.,  folio  272.  1817;  ed.  quarto  338. 
1818.  L.  veijensis  Oerst.  in  Benth.  &  Oerst.,  Vi- 
densk.  Meddel.  Dansk.  Naturhist.  Foren.  Kjob- 
enhavn  1853:  28.  1853. 


42 


FIELDIANA:  BOTANY 


Erect  perennial  herbs  0.5-1. 3(-3)  m  tall,  leafy 
stems  1 .5-7  mm  diam.,  terete,  densely  puberulent 
with  gland-tipped  hairs  0.2-0.7  mm  long.  Leaves 
gradually  diminishing  in  size  distally,  sessile  or 
subsessile  with  petioles  ca.  1  mm  long,  appearing 
amplexicaul;  leaf  blades  l-7(-ll)  cm  long,  6- 
28(-53)  mm  wide,  ovate-triangular  to  narrowly 
triangular-oblong,  apex  acute,  margin  serrate  with 
2-6  teeth/cm,  base  truncate  to  cordate-auriculate 
and  amplexicaul,  drying  dark  grayish  to  dark 
brown  and  coriaceous,  with  short  (0.3  mm)  hairs 
above  and  more  densely  puberulent  beneath,  2° 
veins  3-7/side.  Inflorescences  10-20  cm  long,  ra- 
cemes or  spicate,  glandular  puberulent,  flowers 
crowded  or  separated  by  internodes  to  12  mm 
long,  bracts  5-16  mm  long,  3-1 1  mm  wide  at  the 
base,  ovate  to  lanceolate,  sessile,  pedicels  l-3(-6) 
mm  long.  Flowers  glandular-puberulent  external- 
ly, calyx  4-8  mm  long,  3-5  mm  diam.,  lobes  2- 
4  mm  long,  triangular  and  acute;  corolla  14-38(- 
60)  mm  long,  3.5-6(-10)  mm  diam.,  tubular,  dark 
red  to  red-orange  (pink  or  purple),  upper  lip  6-12 
mm  long,  lower  lobes  ca.  2  mm  long;  functional 
stamens  2;  thecae  glabrous.  Fruits  6-12  mm  long, 
4-7  mm  diam.,  ovoid,  dark;  seeds  0.8-1.2  mm 
long,  0.4-0.6  mm  thick,  wedge  shaped,  with  a 
deeply  reticulated  surface. 

Plants  of  open  sites  in  lower  montane  evergreen 
or  partly  deciduous  formations,  1000-1800  m  el- 
evation (500-2800  m  in  Mexico).  Flowering  pri- 
marily in  July-December  in  Costa  Rica  (flowering 
and  fruiting  throughout  the  year  in  Nicaragua).  In 
Costa  Rica  this  species  has  been  collected  only  at 
the  eastern  and  western  edges  of  the  Meseta  Cen- 
tral and  in  the  region  between  Paraiso  and  Orosi. 
This  species  ranges  from  northern  Mexico  to  Pan- 
ama. 

Lamourouxia  viscosa  is  recognized  by  its  mi- 
nute gland-tipped  hairs  on  stems  and  flowers,  stiff 
sessile  opposite  leaves  that  often  surround  the 
stem  at  their  cordate  bases,  and  lower  montane 
habitats.  This  species  is  most  closely  related  to 
the  Mexican  species  L  smithii  Pringle  and  L 
rhinanthifolia  Kunth  in  H.B.K.  Costa  Rican  col- 
lections of  L.  viscosa  do  not  possess  leaves  and 
flowers  as  large  as  some  found  in  Mexico. 


Leucocarpus  D.  Don  in  Sweet 

Herbs,  stems  erect,  quadrangular  in  cross-sec- 
tion and  often  with  4  longitudinal  wings,  glabrous 
or  sparsely  puberulent.  Leaves  opposite,  sessile, 
narrow  and  serrulate,  cordate-amplexicaul  at  the 


base,  venation  pinnate,  drying  yellowish  brown. 
Inflorescences  cymose.  solitary  in  distal  leaf  ax- 
ils, peduncles  with  bracteoles  subtending  the  ped- 
icels or  secondary  inflorescence  branches,  brac- 
teoles absent  at  the  base  of  the  calyx.  Flowers 
with  cupulate  thin-walled  calyx,  5-veined.  per- 
sisting in  fruit,  calyx  lobes  narrow  and  tooth-like; 
corolla  tubular  or  tubular-campanulate.  somewhat 
bilaterally  symmetrical  with  subequal  upper  and 
lower  lips,  glabrous  externally,  upper  lip  exterior 
in  bud,  2-lobed,  lower  lip  3-lobed,  puberulent  at 
the  mouth  within;  stamens  4  in  2  unequal  pairs 
(or  with  only  2  fertile  stamens),  anthers  with  2 
slightly  diverging  thecae,  staminodes  rarely  pres- 
ent; ovary  conical,  2-locular.  ovules  many,  style 
thickened  at  the  apex,  stigma  2-lobed.  Fruits 
fleshy  indehiscent  berries,  white,  pericarp  thin, 
style  base  forming  a  small  beak;  seeds  immersed 
in  pulp,  minutely  reticulate. 

Leucocarpus  is  a  monotypic  genus,  ranging 
from  Mexico  to  Bolivia.  It  is  included  in  the  tribe 
Dodartieae,  with  Hemichaena,  Mimulus.  and  Ma- 
zus.  At  one  time,  Bentham  included  Hemichaena 
in  Leucocarpus,  but  the  genera  are  now  consid- 
ered distinct. 

Leucocarpus  perfoliatus  (Kunth  in  H.B.K.) 
Benth.  in  DC.,  Prodr.  10:  335.  1846.  Mimulus 
perfoliatus  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  2: 
371.  1817.  Conobea  alata  Graham,  Edinburgh 
New.  Philos.  J.  10:  168.  1830.  L  alatus  (Gra- 
ham) D.  Don  in  Sweet,  Brit.  Fl.  Card.  2:  tab. 
124.  1833.  Figure  6. 

Erect  herbs  or  shrubs  0.5-2.5  m  tall,  flowering 
stems  usually  unbranched.  leafy  stems  3-8  mm 
wide,  with  4  prominent  longitudinal  wings  (ridg- 
es), wings  0.3-2  mm  wide,  usually  glabrous. 
Leaves  of  the  same  node  slightly  united  at  the 
base  to  form  a  short  interpetiolar  ridge,  sessile  and 
amplexicaul;  leaf  blades  8-23(-28)  cm  long,  1- 
4.5(-6)  cm  wide,  narrowly  lanceolate  to  very  nar- 
rowly elliptic-oblong,  tapering  gradually  to  the 
acute  or  acuminate  apex,  margin  serrulate  with  3- 
5  teeth/cm,  cuneate  in  the  lower  V4  but  slightly 
expanded  at  the  rounded-auriculate  base,  drying 
thin-chartaceous,  usually  glabrous.  2°  veins  10- 
14/side.  Inflorescences  2-6  cm  long,  axillary 
cymes  or  with  2°  branching,  with  2-12  flowers, 
peduncles  1-3  cm  long,  0.7-1.3  mm  diam.,  gla- 
brous or  minutely  puberulent  with  thin  hairs  0.1- 
0.2  mm  long,  bracts  3-8  mm  long,  lanceolate, 
pedicels  4-14  mm  long,  minutely  puberulent. 
Flowers  with  calyx  5-9  mm  long,  2.5-3.5  mm 


BURGER:  FLORA  COSTARICENSIS 


43 


diam..  campunulate,  with  narrow  acute  lobes  1-3 
mm  long,  sparsely  minutely  puberulent;  corolla 
(12-)14-17(-23)  mm  long,  tubular  with  slightly 
expanded  lobes,  yellow  to  white,  marked  with 
yellow  and  barbate  within,  tube  2-5  mm  diam., 
lobes  slightly  unequal;  filaments  5-10  mm  long; 
style  5-7  mm  long.  Fruits  globose,  7-10  mm 
diam..  white,  fleshy;  seeds  0.4-0.5  mm  long,  0.2- 
0.3  mm  wide,  ellipsoid-oblong,  yellowish  brown, 
reticulation  minute  (visible  at  50X). 

Plants  of  open  moist  to  wet  sites  in  lower  mon- 
tane evergreen  forest  formations,  800-2200  m  el- 
evation. Probably  flowering  and  fruiting  through- 
out the  year,  but  with  most  Costa  Rican  collec- 
tions made  in  December-January  and  April-June. 
The  species  ranges  from  Veracruz,  Mexico,  to  the 
highlands  of  western  Panama  and  Bolivia. 

Leucocarpus  perfoliatus  is  recognized  by  the 
unbranched  flowering  stems  with  winged  margins, 
sessile  leaves  with  auriculate  bases  clasping  the 
stems,  axillary  inflorescences,  always  shorter  than 
the  leaves,  smaller  tubular  yellow  corollas,  and 
baccate  fruit.  Superficially,  these  plants  are  simi- 
lar to  Hemichaena  fruticosa  (q.v.),  but  that  spe- 
cies has  larger  flowers  and  dry  dehiscent  fruits. 


Limosella  Linnaeus 

REFERENCE — H.  Gliick,  Limosella  studien.  Bei- 
trage  zur  Systematik,  Morphologic,  und  Biologic 
der  Gattung  Limosella.  Bot.  Jahrb.  Syst.  66:  490- 
563.  1934.  A.  Lourtieg,  Etude  sur  Limosella. 
Comite  Nacional  Fran^ais  des  Researches  Antarc- 
tiques:  Biologic  1(10):  165-173.  1964. 

Very  small  herbs,  annual  or  perennial,  growing 
in  shallow  water  or  on  moist  soil,  caespitose  or 
with  short  stolon-like  stems  rooting  from  the 
nodes,  glabrous  or  puberulent.  Leaves  from  the 
apex  of  the  rootstock  or  fasciculate  at  the  nodes, 
densely  clustered,  sessile  or  with  poorly  differ- 
entiated petioles,  blades  linear  to  obovate,  entire, 
flat  or  cylindric,  drying  brown  or  yellowish,  gla- 
brous, venation  obscure  or  with  a  midvein.  Inflo- 
rescences of  1  to  several  flowers  from  the  leaf 
axils,  glabrous,  bracts  absent,  pedicels  usually 
shorter  than  the  leaves,  bracteoles  absent  at  the 
base  of  the  calyx.  Flowers  very  small,  glabrous 
externally,  calyx  campanulate,  usually  thin- 
walled,  equal  to  or  slightly  shorter  than  the  corolla 
tube,  lobes  5  (4)  with  the  posterior  lobe  exterior; 
corolla  funnelform  to  campanulate,  white  to  blu- 
ish or  pinkish,  tube  short,  lobes  3-5,  subequal  and 


imbricate  in  bud,  equal  to  or  shorter  than  the  co- 
rolla tube;  stamens  4  (2,  5),  included,  filaments 
simple,  borne  near  the  middle  of  the  tube,  fila- 
ments of  lower  stamens  crossing  over  the  fila- 
ments of  upper  stamens,  anther  thecae  confluent 
(1-thecous),  not  mucronate;  ovary  ovoid  to  ellip- 
soid, 2-locular  near  the  base,  style  short,  stigma 
capitate.  Fruits  capsules  or  indehiscent,  usually 
bivalvate;  seeds  small,  numerous,  ovoid,  striate 
and  reticulate. 

A  genus  of  ca.  1 1  species  distributed  widely 
over  temperate  and  tropical  montane  regions  but 
most  diverse  in  the  Southern  Hemisphere.  The 
very  small  size  of  the  plants,  their  small  flowers, 
and  their  aquatic  or  moist  habitats  help  distinguish 
the  genus.  The  genus  is  classified  in  the  tribe  Gra- 
tioleae  and  is  often  placed  in  the  subtribe  Limo- 
sellineae. 

Limosella  acaulis  Sesse  &  Mocino,  Fl.  Mex.  ed. 
2:  143.  1894.  L.  americana  Gliick,  Notizbl.  Bot. 
Gart.  Berlin  Dahlem  12:  75.  1934.  L.  americana 
f.  submersa  Gliick,  loc.  cit.  75.  1934.  L.  amer- 
icana f.  natans  Gliick,  loc.  cit.  76.  1934.  L. 
americana  f.  terrestris  Gliick,  loc.  76.  cit.  1934. 
Figure  1. 

Small  herbs  to  5  cm  tall,  stems  to  6  cm  long 
or  little  developed  (and  plants  acaulescent),  root- 
ing at  the  nodes,  glabrous.  Leaves  rosulate  or  fas- 
ciculate, sessile  or  with  poorly  differentiated  pet- 
ioles to  4  cm  long,  clasping  the  stem  at  their  base, 
glabrous  throughout;  leaf  blades  4-15  mm  long 
(-11  cm  when  linear),  0.7-4  mm  wide,  linear  to 
narrowly  oblong  to  spatulate  or  oblanceolate, 
apex  usually  rounded,  margins  entire,  gradually 
narrowed  at  the  base,  drying  brown  or  yellowish, 
midvein  usually  visible  only  near  the  base.  Inflo- 
rescences of  1-3  flowers  in  leaf  axils  (but  difficult 
to  see  among  the  crowded  leaf  bases),  pedicels 
10-30  mm  long,  ca.  0.4  mm  wide,  glabrous. 
Flowers  glabrous  externally,  calyx  1.5-2  mm 
long,  lobes  5,  0.3-0.7  mm  long,  narrowly  trian- 
gular to  oblong,  dark  coloring  sometimes  present 
between  the  lobes  distally;  corolla  ca.  4  mm  long, 
campanulate  or  subrotate,  slightly  zygomorphic 
(bilaterally  symmetric),  white,  tube  ca.  2.5  mm 
long,  ca.  1.3  mm  diam.,  lobes  3-5,  1-1.5  mm 
long,  rotate;  stamens  4  in  2  opposing  pairs.  Fruits 
1.5—4  mm  long,  1-3  mm  diam.,  ovoid-ellipsoid, 
glabrous;  seeds  0.5-1  mm  long,  0.2-0.5  mm  wide, 
striate. 

Rarely  collected  plants  in  shallow  water  or  wet 
soils,  found  only  above  3300  m  elevation  in  Cen- 


44 


FIELDIANA:  BOTANY 


tral  America.  Probably  flowering  and  fruiting 
throughout  the  year.  Known  in  Costa  Rica  from  a 
single  collection  made  in  the  Valle  de  los  Conejos 
on  Chirjipo  Grande  (3400  m)  in  February  (Weston 
12360).  The  species  ranges  disjunctly  from  Mex- 
ico to  Venezuela  and  Bolivia. 

Limosella  acaulis  is  distinguished  by  its  moist 
and  very  high  altitude  habitat,  the  small  size  of 
the  plants,  mostly  fasciculate  linear  sedge-like 
leaves,  and  very  small  flowers  with  white  corollas 
that  have  three  to  five  lobes.  When  seen  from 
above,  the  flowers  may  be  reminiscent  of  Hous- 
tonia  (Rubiaceae),  with  four  corolla  lobes  held  in 
a  single  horizontal  plane.  Species  are  variable  and 
individual  populations  are  often  isolated,  so  local 
variation  is  common.  Also,  the  plant's  habit 
changes  with  water  depth  and  length  of  immer- 
sion. When  submersed,  leaves  develop  to  be  lon- 
ger and  more  linear  than  spatulate;  flowers  remain 
closed  and  are  cleistogamous.  The  Costa  Rican 
collection  describes  the  plants  as  forming  small 
mats  in  a  creek  bed.  Lourtieg  (1964,  cited  above) 
placed  most  taxa  as  synonyms  of  L  australis  R. 
Br.,  but  we  follow  the  decisions  of  Louis  Williams 
regarding  the  circumscription  of  this  species  (Fiel- 
diana,  Bot.  34:  121,  1972). 


Linaria  Miller 

REFERENCE — D.  Sutton,  A.  Revision  of  the 
Tribe  Antirrhineae.  British  Museum  (Natural  His- 
tory) &  Oxford  Univ.  Press,  1988. 

Herbs  or  subshrubs,  annual,  biennial  or  peren- 


nial, erect,  branching,  mostly  from  the  base,  gla- 
brous or  less  often  glandular-puberulent,  drying 
grayish  to  dark  brown.  Leaves  alternate,  some- 
times opposite  or  verticellate  near  the  base,  leaf 
blades  usually  sessile,  entire,  linear  to  reniform, 
pinnately  veined.  Inflorescences  colorful  racemes 
or  spikes  (rarely  flowers  solitary  in  leaf  axils),  al- 
ternate along  the  rachis,  bracts  usually  small, 
bracteoles  absent  at  base  of  calyx.  Flowers  usu- 
ally glabrous,  calyx  campanulate,  with  5  sepals  or 
5  calyx  lobes,  equal  or  subcqual,  imbricate  in  bud, 
persisting  in  fruit;  corolla  tubular  and  strongly  bi- 
labiate, tube  with  a  backward-pointing  abaxial 
spur,  upper  (adaxial)  lip  bilobed  and  exterior  in 
bud,  lower  (abaxial)  lip  3-lobed  and  with  a  prom- 
inent raised  palate  near  the  entrance  to  the  tube; 
stamens  4  in  2  pairs,  included,  filaments  attached 
near  the  base  of  the  tube,  anthers  with  2  parallel 
or  divergent  thecae,  a  staminode  sometimes  pre- 
sent; ovary  2-locular,  ovules  many,  style  slender, 
stigma  small,  2-lobed  or  capitate.  Fruits  thin- 
walled  capsules,  ovoid  to  globose,  dehiscing  lo- 
culicidally  into  2  valves  or  forming  distal  pores; 
seeds  ovoid  to  discoid  or  C-shaped,  testa  thin, 
smooth  to  rugose. 

Linaria  is  a  genus  of  ca.  100  species,  native  to 
north  temperate  regions  and  the  Mediterranean 
area.  They  are  often  grown  in  gardens  for  their 
colorful  and  complex  flowers  or  their  foliage.  The 
corollas  with  backward-pointing  spur  and  strongly 
two-lipped  lobes  are  distinctive.  The  genus  is 
classified  in  the  tribe  Antirrhineae  and  is  closely 
related  to  Cymbalaria  (q.v.).  Although  not  known 
to  be  naturalized  in  Costa  Rica,  the  following  two 
species  are  likely  to  be  present. 


Key  to  the  Species  of  Linaria 

la.  Flowers  yellow,  corollas  20-30  mm  long;  planted  in  gardens  and  rarely  escaping    .  .  .    L.  vulgaris 

Ib.  Flowers  blue  to  purple,  corollas  5-15  (20)  mm  long;  weedy  plants  of  cooler  climates 

.  L  canadensis 


Linaria  canadensis  (L.)  Dumort.,  Bot.  Cult.  2: 
96.  1802.  Antirrhinum  canadensis  L.,  Sp.  PI. 
618.  1753.  L.  texana  Scheele,  Linnaea  21:  761. 
1848. 

Herbs,  annual  or  biennial,  flowering  stems 
erect  from  a  basal  rosette  of  leafy  stems,  15-50 
cm  tall,  branching  from  the  base,  leafy  stems  0.5- 
2.5  mm  diam.,  glabrous.  Leaves  opposite  or 
whorled  near  the  base,  alternate  on  erect  stems. 


sessile;  leaf  blades  of  erect  stems  5-20  mm  long, 
1-2  mm  wide,  linear,  apex  acute  to  bluntly  acute, 
margin  entire,  base  cuneate,  venation  obscure.  In- 
florescences racemes  (or  appearing  spicate),  usu- 
ally making  up  the  distal  half  of  erect  stems,  gla- 
brous, bracts  ca.  2  mm  long,  pedicels  1-4  mm 
long.  Flowers  glabrous  externally,  calyx  2-3.5 
mm  long,  sepals  5,  0.5-1  mm  wide;  corolla  5- 
15(-20)  mm  long,  blue  to  purple,  spur  2-9  mm 
long,  tube  strongly  2-lipped,  the  throat  open, 


BURGER:  FLORA  COSTARICENSIS 


45 


white:  stamens  2-3  mm  long;  ovary  1-2  mm  long, 
ovoid,  style  1-2  mm  long.  Fruits  2-3.5  mm  long, 
subglobose  to  rounded  oblong,  with  persisting 
style  0.5  mm  long;  seeds  0.3-0.5  mm  long,  an- 
gled, smooth. 

Linaria  canadensis,  native  from  southern  Can- 
ada to  northern  Mexico,  is  a  weed  of  open  sunny 
sandy  sites  in  cooler  temperate  or  montane  trop- 
ical climates  (ca.  2000  m  elevation  in  Mexico). 
Although  not  yet  recorded  for  Central  America,  it 
is  naturalized  in  South  America  and  is  likely  to 
become  established  in  our  area.  The  unbranched 
erect  stems,  linear  leaves,  slender  racemes,  and 
blue  corolla  with  spur  help  to  distinguish  this  spe- 
cies. 

Linaria  vulgaris  Miller,  Card.  Diet.  ed.  8.  1768. 
Antirrhinum  linaria  L.,  Sp.  PI.  616.  1753. 

Erect  perennial  herbs  0.2-0.8  m  tall,  spreading 
by  underground  stems  to  form  persisting  clumps, 
erect  stems  usually  unbranched,  1-3  mm  diam., 
usually  glabrous.  Leaves  alternate  and  numerous 
along  the  stem,  glabrous,  sessile;  leaf  blades  3-5 
cm  long,  1-6  mm  wide,  linear  to  linear-lanceolate, 
secondary  veins  obscure.  Inflorescences  spicate 
racemes  with  crowded  flowers,  bracts  ca.  1  cm 
long,  similar  to  the  leaves,  pedicels  1-4  mm  long, 
pedicels  and  flower  oriented  upward  at  an  acute 
angle  to  the  stem.  Flowers  with  glabrous  calyx, 
sepals  2-3.5  mm  long,  narrowly  ovate,  acute;  co- 
rolla 2-3  cm  long,  yellow  with  orange  near  the 
mouth,  tube  with  a  spur  equaling  the  tube  and 
lobes  in  length,  mouth  of  the  corolla  strongly  bi- 
labiate, upper  (adaxial)  lip  8-12  mm  long,  lower 
(abaxial)  lip  6-9  mm  long,  with  a  puberulent 
rounded  palate.  Fruits  5-10  mm  long,  glabrous; 
seeds  ca.  2  mm  long,  discoid  and  winged. 

Linaria  vulgaris,  a  native  of  Eurasia,  is  occa- 
sionally grown  in  gardens  for  its  unusual  yellow 
flowers,  but  it  is  more  likely  to  be  seen  as  a  weed 
in  cooler  climates  of  the  world.  It  is  recognized 
by  its  often  linear  alternate  leaves  and  the  unusual 
two-lipped  yellow  corolla  with  basal  spur.  This 
species  has  not  been  recorded  as  naturalized  in 
Central  America.  These  plants  sometimes  produce 
abnormal  flowers  that  are  radially  symmetric  (not 
two-lipped)  and  have  five  spurs  or  none.  Linnaeus 
mistakenly  erected  the  genus  Peloria  for  such 
plants,  and  the  word  peloria  is  now  used  to  de- 
scribe abnormal  radially  symmetric  forms  of  flow- 
ers that  are  normally  strongly  bilaterally  symmet- 
ric (zygomorphic). 


Lindernia  Allioni 

REFERENCES — D.  Miranda,  Flavonoid  and  mor- 
phological studies  of  Lindernia  Allioni  (Scrophu- 
lariaceae)  in  South  America.  Bot.  J.  Linn.  Soc.  75: 
47-67.  1977.  D.  Philcox,  Revision  of  the  Male- 
sian  species  of  Lindernia  All.  (Scrophulariaceae). 
Kew  Bull.  22:  1-72.  1968.  T.  Yamazaki,  Revision 
of  the  Indo-Chinese  species  of  Lindernia  All.  J. 
Fac.  Sci.  Univ.  Tokyo,  Bot.  13:  1-64.  1981. 

Small  annual  herbs,  terrestrial  or  semiaquatic, 
erect  to  prostrate  or  creeping  and  rooting  at  the 
nodes,  stems  simple  or  branched,  glabrous  or  pu- 
berulent, usually  4-angled,  drying  yellowish  or 
brown.  Leaves  opposite,  simple,  small,  sessile  or 
petiolate,  blades  entire  or  denticulate,  base  often 
truncated  to  cordate,  often  glandular-punctate,  ve- 
nation pinnate  or  3-5-veined  from  the  base.  Inflo- 
rescences of  solitary  flowers  in  the  axils  of  distal 
leaves,  less  often  terminal  or  axillary  racemes  or 
umbel-like  clusters,  bracts  and  bracteoles  absent, 
pedicels  slender,  usually  longer  than  the  calyx. 
Flowers  small,  calyx  tubular  with  5  short  spread- 
ing lobes  or  deeply  parted  to  the  base  with  5  se- 
pals, the  sepals  or  lobes  subequal,  glabrous  or  pu- 
berulent externally,  tube  with  5  longitudinal  ridg- 
es (sometimes  winged);  corolla  tubular  to  cam- 
panulate,  longer  than  the  calyx  (equaling  the  calyx 
and  not  opening  in  cleistogamous  flowers),  blue 
to  purple  or  white,  bilabiate,  upper  lip  usually 
shorter  and  erect,  entire  or  2-lobed,  lower  lip  3- 
lobed  and  spreading;  stamens  4  or  with  2  fertile 
and  2  staminodes  present,  filaments  often  inserted 
at  2  levels  in  the  distal  half  of  the  tube,  anterior 
filaments  often  with  a  distinct  spur  near  the  base, 
upper  filaments  often  bent  just  near  the  apex,  an- 
thers free  or  united  in  pairs,  2-thecous,  thecae  of- 
ten widely  divergent  and  X-shaped;  ovary  gla- 
brous, 2-locular,  ovules  many,  style  slender,  stig- 
mas 2-lobed.  Fruits  dry  capsules,  ovoid  to  ellip- 
soid, globose  or  cylindric,  dehiscing  septicidally 
and  2-valved,  the  placentae  winged;  seeds  many, 
oblong-elliptic,  smooth  to  reticulate  or  alveolate 
with  prominent  transverse  or  longitudinal  raised 
ridges. 

A  genus  of  50  to  70  species  widespread  in  trop- 
ical and  temperate  areas  throughout  the  world, 
with  the  majority  of  species  in  tropical  parts  of 
Africa,  Asia,  and  Australia.  The  genus  is  classi- 
fied in  the  tribe  Gratioleae  and  placed  in  subtribe 
Lindernieae  with  Torenia.  It  seems  likely  that  L. 
crustacea  is  indigenous  to  Costa  Rica  and  that  the 


46 


FIELDIANA:  BOTANY 


other  two  species  have  been  introduced.  These 
plants  are  generally  found  in  marshes,  at  the  edges 
of  standing  water,  or  in  moist  depressions,  in  open 
sunny  or  partly  shaded  sites.  The  preference  for 


moist  soils,  the  short  slender  stems,  small  opposite 
leaves,  small  bluish  flowers,  tubular  bilabiate  co- 
rollas, four  or  two  fertile  stamens,  and  appenda- 
ged  anther  filaments  help  distinguish  the  genus. 


Key  to  the  Species  of  Lindernia 

la.  Flowers  subsessile,  pedicels  <  3  mm  long,  flowers  solitary  in  leaf  axils;  corollas  white  or  yellowish 

to  pale  lilac;  leafy  stems  usually  with  puberulent  longitudinal  ridges;  fruits  6-10  mm  long  

L.  diffusa 

Ib.  Flowers  borne  on  pedicels  3-23  mm  long  (longer  pedicels  at  distal  nodes),  flowers  often  in  terminal 
cyme-like  groupings;  corollas  bluish  to  lilac,  rose,  or  white  marked  with  yellow;  leafy  stems  gla- 
brous or  puberulent;  fruits  3-4  mm  long  2 

2a.  Calyx  5-lobed,  united  ca.  50%  to  form  a  tube  (but  splitting  after  anthesis  and  appearing  as  separate 
sepals  in  fruit);  fruits  ovoid,  seeds  0.4-0.5  mm  long;  leaves  petiolate  L  Crustacea 

2b.  Calyx  with  5  sepals  united  only  at  the  base;  fruits  ellipsoid,  seeds  0.3-0.4  mm  long;  leaves  sessile 

.  /  diihni 


Lindernia  Crustacea  (L.)  F.  v.  Muell.,  Syst.  Cen- 
sus Aust.  PL  1:  97.  1882.  Capraria  Crustacea 
L.,  Mant.  PL  1:  87.  1767.  Torenia  Crustacea 
(L.)  Cham.  &  Schldl.,  Linnaea  2:  570.  1827. 
Vandelila  Crustacea  (L.)  Benth.,  Scroph.  In.  35: 
1835.  Pyxidaria  Crustacea  (L.)  Kuntze,  Rev. 
Gen.  PL  2:  464.  1891.  Figure  2. 

Prostrate  or  procumbent  herbs  to  15  cm  tall, 
rarely  rooting  at  distal  nodes,  leafy  stems  0.2-1.3 
mm  diam.,  longitudinally  ribbed,  glabrous  or  with 
thin  crooked  hairs  0.1-1  mm  long.  Leaves  op- 
posite, petioles  1-7  mm  long,  0.4-0.7  mm  wide, 
glabrous  or  sparsely  puberulent,  leaf  bases  not 
united  across  the  stem;  leaf  blades  6-16  mm  long, 
4-15  mm  wide,  ovate  to  ovate-rounded  or  ovate- 
triangular,  apex  obtuse  or  rounded,  margin  serrate 
with  4-6  teeth/cm,  base  truncate  to  rounded-sub- 
cordate,  drying  membranaceous  and  yellowish 
green,  minutely  punctate,  glabrous  or  very  sparse- 
ly puberulent  beneath  and  along  margin,  2°  veins 
3-5/side.  Inflorescences  solitary  axillary  flowers 
or  terminal  cymes,  pedicels  4-18(-25)  mm  long, 
0.1-0.2  mm  diam.,  glabrous  or  with  few  thin  hairs 
to  0.4  mm  long.  Flowers  with  calyx  2-4  mm 
long,  1-1.8  mm  diam.,  lobes  0.7-1.5  mm  long, 
triangular  and  acute,  glabrous  or  very  sparsely  pu- 
berulent; corolla  4-7  mm  long,  blue  to  violet  or 
blue-purple  (white),  tube  ca.  5  mm  long,  1-1.5 
mm  diam.;  fertile  stamens  4  in  2  pairs,  anthers 
connivent;  ovary  glabrous.  Fruits  3-4  mm  long, 
2-2.5  mm  wide,  rounded-ovoid,  central  column 
(placenta)  to  2  mm  wide  with  smooth  lateral 
wings  and  central  reticulated  area;  seeds  0.4-0.5 


mm  long,  0.3-0.4  mm  diam.,  oblong,  yellowish, 
obscurely  reticulate. 

Uncommon  plants  of  moist  open  sunny  or  part- 
ly shaded  sites  in  lowland  evergreen  forest  for- 
mations, 0-300(-1000)  m  elevation.  It  is  also 
found  along  stream  edges  in  seasonally  dry  areas. 
Flowering  and  fruiting  throughout  the  year.  This 
species  is  a  widespread  weed,  now  found  in  trop- 
ical and  warm-temperate  climates  throughout  the 
world. 

Lindernia  crustaced  is  recognized  by  its  small, 
mostly  prostrate  habit,  mostly  glabrous  parts, 
small  ovate-serrate  leaves,  small  blue  flowers  on 
prominent  slender  pedicels,  calyx  lobes  united  for 
half  their  length,  small  rounded  fruit,  and  unusual 
persisting  placenta  with  smooth  lateral  wings.  In 
addition,  the  lower  pair  of  anthers  is  inserted  on 
the  lower  corolla  lip.  The  persisting  and  expanded 
calyx  often  splits  along  the  thin  intercostal  areas 
to  produce  separate  sepals,  appearing  very  differ- 
ent from  the  calyx  in  anthesis. 

Lindernia  diffusa  (L.)  Wettst.  in  Engl.  &  Prantl, 
Naturlichen  Pflanzenfam.  4,  3b:  79.  1891.  Van- 
delia  diffusa  L.,  Mant.  PL  1:  89.  1767.  Pyxi- 
daria diffusa  (L.)  Kuntze,  Rev.  Gen.  PL  2:  464. 
1891.  Figure  2. 

Prostrate  or  decumbent  herbs  2-15  cm  tall, 
sometimes  forming  small  mats  to  25  cm  wide, 
often  rooting  at  the  nodes,  leafy  stems  0.4-1.3 
mm  diam.,  quadrangular  with  4  puberulent  lon- 
gitudinal ribs,  the  thin  whitish  hairs  0.2-0.3  mm 
long.  Leaves  opposite,  united  at  the  base  to  form 


BURGER:  FLORA  COSTARICENSIS 


47 


an  interpetiolur  line  or  ridge,  subsessile  or  with 
petioles  to  4  mm  long;  leaf  blades  6-25  mm  long, 
4-24  nun  \vide.  ovate  to  broadly  ovate  or  rhom- 
bic, apex  obtuse  to  rounded,  margin  serrate  with 
6-10  teeth/cm,  base  broadly  obtuse  to  rounded 
and  truncate,  minutely  punctate,  glabrous  above, 
with  minute  (0.1-0.2  mm)  hairs  along  the  veins 
beneath,  venation  subpalmate  with  2-3  2°  veins/ 
side.  Inflorescences  of  solitary  axillary  flowers, 
usually  2/node,  pedicels  1-3  mm  long,  ca.  0.3  mm 
diam.,  minutely  puberulent.  Flowers  with  calyx 
4-7  mm  long,  1-1.5  mm  diam.,  lobes  2-4  mm 
long,  linear-lanceolate,  glabrous  or  minutely  pu- 
berulent on  the  major  veins  and  margins  of  lobes; 
corolla  6-9  mm  long,  white,  yellowish,  or  pale 
lilac  with  yellow  interior;  fertile  stamens  4;  ovary 
glabrous,  style  ca.  3  mm  long.  Fruits  6-10  mm 
long  (including  the  1-2  mm  beak),  2.3-4  mm 
diam.,  ellipsoid  or  narrowly  ovoid-ellipsoid;  seeds 
0.5-0.6  mm  long  ca.  0.3  mm  diam.,  oblong,  with 
minute  spines  and  a  reticulate-pitted  surface. 

Plants  of  open  sunny  sites  in  evergreen  forest 
formations,  10-1200  m  elevation.  Probably  flow- 
ering and  fruiting  throughout  the  year.  Probably 
originating  in  the  Old  World,  this  species  is  now 
widespread  in  warmer  climates. 

Lindernia  diffusa  is  recognized  by  its  small 
size,  stems  with  usually  four  puberulent  ridges, 
small  subsessile  serrate  leaves  with  broad  blades, 
solitary  axillary  flowers  on  very  short  pedicels, 
and  white  corollas  marked  with  yellow. 

Lindernia  dubia  (L.)  Pennell,  Acad.  Nat.  Sci. 
Philadelphia  Monogr.  1:  141.  1935.  Gratiola 
dubia  L.,  Sp.  PI.  17.  1753.  Capraria  gratiolo- 
ides  L.,  Syst.  Veg.  ed.  10:  1117.  1759.  G.  in- 
aequalis  Walt.,  Fl.  Carol.  61.  1788.  G.  anagal- 
lidea  Michx.,  Fl.  Bor.  Am.  1:  6.  1803.  Ilysan- 
thes  gratioloides  (L.)  Benth.  in  DC,  Prodr.  10: 
419.  1846.  L.  gratioloides  (L.)  Lloyd  &  Fouc., 
Fl.  Quest  Fr.  ed.  4:  246.  1886.  L.  inaequalis 
(Walt.)  Pennell,  Torreya  19:  149.  1919.  /.  dubia 
(L.)  Barnhart,  Bull.  Torrey  Bot.  Club.  26:  376. 
1899.  L.  anagallidea  (Michx.)  Pennell,  Acad. 
Nat.  Sci.  Philadelphia  Monogr.  1:  152.  1935. 
Figure  4. 

Herbs,  stems  procumbent  or  erect,  5-30  cm 
tall,  simple  or  much-branched,  rooting  mostly  at 
the  base,  leafy  stems  0.2-1  mm  diam.,  usually 
with  4  longitudinal  ridges,  glabrous.  Leaves  op- 
posite, sessile,  free  or  united  at  the  base  and  form- 
ing an  interpetiolar  line;  leaf  blades  4-14(-25) 
mm  long,  3-8(-10)  mm  wide,  narrowly  ovate  to 


narrowly  ovate-triangular  or  lanceolate,  apex 
acute  to  obtuse,  margin  entire  or  with  2  or  3  teeth/ 
side,  base  acute  to  somewhat  rounded,  drying 
membranaceous,  glabrous,  venation  usually  pal- 
mate with  3  major  veins.  Inflorescences  of  soli- 
tary axillary  flowers  or  cymes  subtended  by  leaf- 
like  bracts,  pedicels  3-23  mm  long,  ca.  0.2  mm 
diam.,  glabrous,  distal  pedicels  much  longer  than 
those  at  lower  nodes.  Flowers  with  calyx  deeply 
5-parted,  sepals  1.8-2.5  mm  long,  0.3-0.4  mm 
wide,  linear-oblong,  becoming  3-5  mm  long  in 
fruit  but  remaining  narrow;  corolla  5-10  mm 
long,  blue  or  white  with  purplish  throat  or  lilac  to 
rose;  fertile  stamens  2,  staminodes  2  and  bifid  at 
the  apex;  ovary  glabrous,  style  3  mm  long.  Fruits 
3-6  mm  long,  ca.  2  mm  wide,  narrowly  ovoid- 
oblong  to  oblong-ellipsoid;  seeds  0.3-0.4  mm 
long,  0.2-0.3  mm  diam.,  oblong,  with  longitudinal 
rows  of  pits. 

Plants  of  open  wet  sites  and  marshes  in  ever- 
green or  seasonally  deciduous  areas,  5-1100  m 
elevation.  Probably  flowering  primarily  at  the  end 
of  the  rainy  season  (November-January).  Rarely 
encountered  in  Costa  Rica,  this  species  ranges 
from  southern  Canada  and  the  eastern  United 
States  to  southern  South  America  and  the  West 
Indies. 

Lindernia  dubia  is  recognized  by  its  small  stat- 
ure, moist  habitats,  lack  of  pubescence,  opposite 
sessile  leaves,  distal  flowers  on  longer  pedicels, 
white  and  blue  two-lipped  corollas,  and  separate 
sepals.  There  is  considerable  diversity  of  leaf 
form,  with  some  plants  having  narrowly  ovate 
leaves  (van  anagallidea  (Michx.)  Cooperrider) 
and  other  plants  with  more  elongate  narrow  leaves 
(var.  dubia).  Holmgren  (Flora  of  the  Great  Plains, 
1 986,  p.  769)  found  that  L.  anagallidea  cannot  be 
effectively  separated  from  L.  dubia.  Likewise,  in 
1984  D.  A.  Quails  annotated  many  North  Amer- 
ican collections  as  L.  dubia.  We  follow  their  de- 
cision to  consider  the  two  as  elements  of  the  same 
species.  Lindernia  microcalyx  Pennell  &  Stehle  is 
very  similar  but  has  corollas  9-14  mm  long;  it 
has  been  collected  along  the  Caribbean  shore  of 
Guatemala  and  Honduras  (determinations  by  D. 
A.  Quails,  1984). 


Lophospermum  D.  Don 

REFERENCES — W.  Elisens,  Monograph  of  the 
Maurandyinae  (Scrophulariaceae — Antirrhineae). 
Syst.  Bot.  Monogr.  5:  1-97.  1985.  D.  Sutton,  A 
Revision  of  the  Tribe  Antirrhineae.  British  Mu- 


48 


FIELDIANA:  BOTANY 


seum  (Natural  History)  &  Oxford  Univ.  Press. 
1988. 

Perennial  herbs  or  climbers,  stems  often  scan- 
dent  or  clambering,  branches  often  arising  from  a 
woody  base,  sparsely  puberulent  to  glandular-vil- 
lous,  drying  grayish  or  greenish.  Leaves  alternate, 
petioles  well  developed  and  often  bending  to  fa- 
cilitate climbing;  leaf  blades  deltoid  to  reniform, 
apex  acute  to  mucronate,  margins  dentate  to 
broadly  crenate,  venation  palmate.  Inflorescences 
of  solitary  flowers  in  axils  of  leaves,  pedicels  hor- 
izontal to  ascending,  bracts  and  bracteoles  absent. 
Flowers  large,  calyx  urceolate  or  campanulate 
and  expanded,  sepals  free  or  united  at  the  base, 
narrowly  to  broadly  ovate,  glandular  puberulent, 
often  enlarging  in  fruit;  corolla  tubular-campan- 
ulate,  somewhat  bilabiate  and  open-throated,  red- 
dish to  violet  or  purple  distally  (whitish  or  pale 
near  the  base),  5-lobed,  upper  2  lobes  recurved, 
lower  3  lobes  projecting  forward;  stamens  4,  sub- 
equal  or  in  2  pairs,  included,  connective  often  ex- 
panded, a  staminode  usually  present;  ovary  gla- 
brous or  glandular  puberulent,  2-locular,  ovules 
many,  stigma  recurved  or  straight,  forked  and  di- 
vergent (rarely  conical  and  lobed).  Fruits  dry 
capsules,  ovoid  to  globose,  symmetric  or  asym- 
metric, dehiscing  irregularly  or  poricidal,  bi- 
valved;  seeds  rounded  with  a  wing  around  the 
margin. 

A  genus  of  6  or  20  species,  depending  on 
whether  the  Mexican  genus  Rhodochiton  Zucc.  is 
included  (following  Elisens,  1985)  or  excluded 
(following  Sutton,  1988).  Species  range  from 
north-central  Mexico  to  Guatemala;  a  few  species 
are  planted  as  ornamental  climbers.  These  plants 
are  characterized  by  the  scandent  or  clambering 
habit,  alternate  leaves,  twining  petioles,  deltate  to 
cordiform  blades,  large  corollas,  and  round  seeds 
with  thin  peripheral  wing.  The  one  species  found 
in  Costa  Rica  was  formerly  placed  in  the  genus 
Maurandya;  both  are  placed  in  the  tribe  Antirrhi- 
neae. 

Lophospermum  erubescens  D.  Don  in  Sweet, 
Brit.  Fl.  Card,  sen  2,  1:  tab.  68  and  after  tab. 
75.  1830.  Maurandya  erubescens  (D.  Don) 
Gray,  Proc.  Am.  Acad.  Arts  7:  377.  1868.  Asar- 
ina  erubescens  (D.  Don)  Pennell,  Proc.  Acad. 
Nat.  Sci.  Philadelphia  99:  174.  1947.  Figure  8. 

Climbing  herbs,  leafy  stems  1-4  mm  diam., 
pubescent  with  thin  straight  or  crooked  multicel- 
lular  hairs  0.3-0.8  mm  long,  some  hairs  with 


gland  tips.  Leaves  alternate,  petioles  3-6  cm  long, 
0.6-1.2  mm  diam..  often  bent  or  curving  (some- 
times helping  to  support  the  stem),  pubescent;  leaf 
blades  3-7(-15)  cm  long,  2-6.5(-15)  cm  wide, 
triangular  to  triangular-rhombic  or  sagitate,  apex 
acute,  margin  prominently  crenate-serrate  with 
teeth  0.5-6  mm  high  and  1-8  mm  wide,  base  cor- 
date, with  thin  whitish  hairs  0.3-0.5  mm  long 
above  and  below,  major  veins  3  or  5.  Inflores- 
cences of  solitary  axillary  flowers,  pedicels  2-7(- 
11)  cm  long,  0.7-1.4  mm  diam.,  puberulent,  not 
bent  or  twining.  Flowers  puberulent  externally, 
calyx  15-24  mm  long,  sepals  subequal,  10-14 
mm  wide,  broadly  ovate;  corolla  6-7  cm  long, 
tubular-campanulate,  bright  pink  to  red  distally. 
tube  constricted  above  the  base,  whitish  within 
and  with  yellow  hairs  in  the  throat,  ca.  2  cm  wide 
at  the  mouth,  lobes  10-14  mm  long,  12-16  mm 
wide,  rounded  distally.  Fruits  15-20  mm  long, 
subtended  by  persisting  sepals,  surface  with  thin 
multicellular  hairs;  seeds  ca.  2.5  X  2.5  mm,  wings 
lateral  with  a  narrow  notch  at  apex  and  truncated 
base,  body  of  the  seed  ca.  1 .5  X  0.8  mm,  tuber- 
culate. 

Lophospermum  erubescens  is  characterized  by 
its  vining  habit,  twisted  petioles,  triangular  and 
coarsely  dentate  leaf  blades,  broad  sepals,  and 
large,  tubular,  slightly  asymmetric  pink  corollas. 
This  species  is  native  to  the  oak  forests  of  the 
Sierra  Madre  Oriental  of  Mexico  and  is  now  com- 
monly grown  in  gardens  as  a  climbing  ornamen- 
tal. It  has  been  collected  as  an  escape  at  Monte- 
verde,  where  it  was  flowering  in  January  and  Sep- 
tember at  ca.  1400  m,  and  north  of  San  Isidro  del 
General  at  1500  m  elevation,  where  it  was  flow- 
ering in  April.  Compare  Maurandya  barclaiana 
and  M.  scandens. 


Maurandya  Ortega 

RHFERHNCKS — W.  Elisens,  Monograph  of  the 
Maurandyinae  (Scrophulariaceae — Antirrhineae). 
Syst.  Bot.  Monogr.  5:  1-97.  1985.  D.  Sutton,  A 
Revision  of  the  Tribe  Antirrhineae.  British  Mu- 
seum (Natural  History)  &  Oxford  Univ.  Press. 
1988. 

Scandent  herbs,  annual  or  perennial,  with  thin 
flexible  stems  from  a  fibrous  base  or  taproot,  gla- 
brous. Leaves  alternate,  petioles  long  and  often 
twisting  to  support  the  twining  habit;  leaf  blades 
hastate  to  sagittate  or  rarely  cordiform,  margins 
usually  entire,  glabrous,  venation  palmate.  Inflo- 


BURGER:  FLORA  COSTARICENSIS 


49 


rescences  of  solitary  flowers  in  leaf  axils,  pedi- 
cels long,  terete  or  winged  near  the  base,  glabrous 
or  glandular-puberulent  distally,  bracts  and  brac- 
teoles  absent.  Flowers  with  5  sepals  united  only 
at  the  base,  equal,  lanceolate,  imbricate  near  the 
base  ( urceolate),  margins  entire,  glabrous  or  glan- 
dular-pubescent; corolla  tubular  and  prominently 
2-lipped,  with  open  or  closed  throat,  pink  to  red, 
violet,  or  blue,  often  whitish  near  the  base,  gla- 
brous to  glandular-puberulent  externally,  upper  lip 
with  2  recurved  lobes,  lower  lip  with  3  recurved 
or  projecting  lobes,  often  closing  the  mouth  with 
the  well-developed  palate;  stamens  4  in  2  pairs, 
included,  inserted  near  the  base  of  the  tube,  fila- 
ments villous  at  the  base;  ovary  2-locular,  locules 
subequal  or  unequal,  glabrous  or  with  glandular 
trichomes,  ovules  many,  stigma  conical  and  re- 
curved. Fruits  dry  2-valved  capsules,  ovoid  to 


globose,  dehiscence  irregular,  irregular-transverse 
or  poricidal;  seeds  rectangular  with  tuberculate- 
cristate  surface  (ovoid  with  a  peripheral  wing  and 
minute  tuberculate  surface  sculpturing  in  M.  wis- 
lizeni),  dark  brown. 

As  revised  by  Elisens  (1985,  cited  above), 
Maurandya  is  a  genus  of  four  species  ranging 
from  the  southwestern  United  States  to  central 
Mexico.  Sutton  (1988,  cited  above)  recognizes 
only  two  species  in  the  genus,  transferring  two 
species  found  in  the  southwestern  United  States 
to  splinter  genera.  The  genus  is  closely  related  to 
Lophospermum,  and  they  are  placed  in  the  tribe 
Antirrhineae.  Two  species  are  likely  to  be  found 
as  garden  ornamentals  in  Central  America,  and 
one  has  recently  been  collected  as  an  escape. 
These  plants  have  also  been  placed  in  the  genus 
Asarina. 


Key  to  the  Species  of  Maurandya 

la.  Seeds  with  lateral  wings;  corolla  to  6  cm  long    (see  Lophospermum  erubescens) 

Ib.  Seeds  without  wings;  corolla  to  4  cm  long    2 

2a.  Calyx  covered  with  short  gland-tipped  hairs;  corolla  usually  blue-violet    M.  barclaiana 

2b.  Calyx  glabrous  or  with  few  gland-tipped  hairs;  corolla  usually  pink M.  scandens 


Maurandya  barclaiana  Lindley,  Hot.  Reg.  13, 
tab.  1108.  1827.  Figure  8. 

Vines  with  slender  herbaceous  climbing  stems 
and  twisting  petioles,  leafy  stems  0.6-1  mm  diam., 
glabrous  or  minutely  papillate-puberulent  at  the 
nodes.  Leaves  alternate,  petioles  12-32  mm  long, 
0.3-0.5  mm  diam.,  glabrous,  often  coiling  around 
objects  for  support;  leaf  blades  16-24(-35)  mm 
long,  14-35  mm  wide,  triangular-hastate  or  sagit- 
tate, apex  acute,  usually  with  2  basal  lobes,  gla- 
brous, venation  subpalmate  with  3  (5)  major  veins 
from  the  base.  Inflorescences  of  solitary  flowers 
in  leaf  axils,  pedicels  15-55  mm,  ca.  0.5  mm 
diam.,  straight  or  curved,  glabrous  except  near  the 
calyx.  Flowers  with  calyx  8-16  mm  long,  2-3  mm 
wide  at  base,  lobes  triangular  with  long  narrow 
apex,  covered  with  gland-tipped  hairs  0.3-0.5  mm 
long;  corolla  2.5-4  cm  long,  ca.  15  mm  wide, 
blue- violet,  lobes  6-10  mm  long,  filaments  ca.  18 
and  14  mm  long.  Fruits  11-17  mm  long,  ovoid; 
seeds  with  angular  projections. 

Maurandya  barclaiana  is  a  vining  ornamental 
often  planted  in  tropical  gardens.  It  is  recognized 
by  its  colorful  tubular  two-lipped  corollas,  glan- 
dular-puberulent calyx,  often  sagittate  leaves,  and 
twisting  petioles. 


Maurandya  scandens  (Cav.)  Pers.,  Syn.  PI.  2: 
160.  1806.  Usteria  scandens  Cav.,  Icon.  2:  15, 
tab.  116.  1793.  Maurandya  semperflorens  Or- 
tega, Nov.  PI.  Descr.  Dec.  21.  1797.  Reichardia 
scandens  (Cav.)  Roth,  Catal.  bot.  2:  65.  1800. 
Asarina  scandens  (Cav.)  Pennell,  Proc.  Acad. 
Nat.  Sci.  Philadelphia  99:  175.  1947.  Figure  8. 

Herbaceous  climbers  with  slender  twining 
stems,  often  with  adventitious  roots,  leafy  stems 
0.5-2  mm  diam.,  glabrous.  Leaves  alternate,  pet- 
ioles 8-35(-42)  mm  long,  0.4-0.8  mm  diam.,  gla- 
brous, often  bent  or  twisted  along  their  length; 
leaf  blades  2-5(-6)  cm  long,  l-3.5(-4.5)  cm 
wide,  sagittate  to  hastate  or  triangular,  apex  acute, 
margin  entire,  base  cordate  and  usually  with 
prominent  lateral  lobes,  drying  membranaceous, 
glabrous  or  minutely  papillate-puberulent  near  the 
petiole  attachment,  major  veins  5  or  7,  the  3  cen- 
tral veins  reaching  the  middle  of  the  blade.  Inflo- 
rescences of  solitary  axillary  flowers,  pedicels  2- 
6(-8.5)  cm  long,  0.4-1  mm  diam.,  slightly  thick- 
ened near  the  base,  glabrous.  Flowers  with  calyx 
9-15  mm  long,  sepals  2-4  mm  wide  near  the 
base,  lanceolate,  glabrous  (rarely  sparsely  glan- 
dular puberulent);  corolla  to  4  cm  long,  tubular- 


50 


FIELDIANA:  BOTANY 


campanulate,  pink  to  pale  purple,  tube  2-3  cm 
long,  lobes  6-10  mm  long,  rounded;  filaments  16 
and  12  mm  long,  lower  filaments  shorter;  ovary 
glabrous  or  with  glandular  hairs  near  apex.  Fruits 
9-12  mm  long,  ovoid  to  oblong,  locules  subequal; 
seeds  with  rounded  projections. 

Maurandya  scandens,  a  native  of  central  Mex- 
ico, is  planted  as  an  ornamental  climber  in  Central 
America.  The  thin  twining  stems,  lack  of  pubes- 
cence, often  twisted  petioles,  sagittate-triangular 
leaves,  solitary  axillary  flowers,  and  tubular  two- 
lipped  pinkish  corollas  make  the  plants  distinc- 
tive. 


Mazus  Loureiro 

Small  annual  herbs,  diffusely  branched  from  a 
basal  rosette,  stems  with  longitudinal  ridges,  pu- 
bescent or  subglabrous.  Leaves  opposite  and 
crowded  near  the  base,  alternate  distally,  petioles 
poorly  differentiated;  leaf  blades  obovate  with 
crenulate  or  coarsely  dentate  margins,  venation 
pinnate.  Inflorescences  terminal  racemes,  flowers 
alternate  and  solitary  along  the  rachis  (often  on  1 
side),  bracts  minute  or  absent,  pedicels  well  de- 
veloped, lacking  bracteoles.  Flowers  with  cam- 
panulate or  tubular  calyx,  5-lobed,  tube  usually 
equaling  the  lobes  in  length,  lobes  equal  in  size 
and  shape,  glabrous  or  puberulent,  slightly  ex- 
panded and  enclosing  the  fruit;  corolla  2-lipped, 
light  blue  to  white  or  lavender,  tube  short,  upper 
lip  2-lobed,  lower  lip  3-lobed,  larger  than  the  up- 
per and  spreading;  stamens  4  in  2  pairs,  filaments 
inserted  at  the  base  of  the  tube,  anthers  divaricate, 
thecae  contiguous,  a  staminode  absent;  ovary  2- 
locular,  ovules  many,  style  longer  than  the  ovary, 
stigma  broadly  bilabiate.  Fruits  capsules,  globose 
or  slightly  compressed,  loculicidal,  producing  2 
entire  valves;  seeds,  ovoid  to  oblong  or  angled, 
rugulose  or  reticulate,  black. 

Mazus  is  an  Asian-Australian  genus  of  30  to  40 
species  that  is  placed  in  the  tribe  Dodartieae  with 
Mimulus  and  Leucocarpus.  The  following  species 
is  now  an  occasional  weed  in  temperate  and  trop- 
ical montane  climates. 

Mazus  pumilus  (Burm.f.)  Steenis,  Nova  Guinea 
N.S.  9:  31.  1958.  Lobelia  pumila  Burm.f.,  Fl. 
Ind.  186,  t.  60,  f.  3.  1768.  M.  japonicus 
(Thunb.)  Kuntze,  Rev.  Gen.  PI.  1:  462.  1891. 
Lindernia  japonica  Thunb.,  Fl.  Jap.  253.  1784. 
M.  rugosus  Lour.,  Fl.  Cochinch.  385.  1790. 


Annual  herbs  3-15  cm  tall,  ascending  or  de- 
cumbent, sometimes  forming  small  mats,  stems  to 
20  cm  long,  leafy  stems  0.8-1.5  mm  diam., 
sparsely  puberulent  with  hairs  to  0.5  mm  long. 
Leaves  rosulate  or  crowded  at  the  base,  opposite 
leaf  bases  united  across  the  stem,  distal  leaves 
smaller  and  alternate,  petioles  2-15  mm  long, 
with  lateral  margins  not  clearly  differentiated 
from  the  blades;  leaf  blades  6-60  mm  long,  4- 
20  mm  wide,  obovate  to  cuneate-oblong  or  oblan- 
ceolate,  apex  obtuse  or  rounded,  margins  of  larger 
leaves  with  prominent  lobes  0.5-3  mm  long  (sep- 
arated by  wide  sinuses),  base  cuneate  and  decur- 
rent,  sparsely  puberulent  with  hairs  to  0.4  mm 
long,  2°  veins  2  or  3/side.  Inflorescences  3-8  cm 
long,  racemes  with  3-13  flowers  separated  by  in- 
ternodes  to  12  mm  long,  bracts  1-2  mm  long,  lin- 
ear-filiform, pedicels  2-6(-l  1)  mm  long,  sparsely 
to  densely  papillate  puberulent.  Flowers  with  ca- 
lyx 4-7  mm  long,  with  5  prominent  veins,  tube 
1-2  mm  diam.,  lobes  2-3  mm  long,  narrowly  tri- 
angular or  oblong,  with  5  prominent  longitudinal 
veins,  glabrous  or  minutely  papillate  puberulent; 
corolla  7-10  mm  long,  pale  violet  and  white,  tu- 
bular, upper  lobes  erect,  lower  lip  ca.  2  mm  long 
with  2  raised  longitudinal  ridges  adaxially,  palate 
yellow  or  whitish;  stamens  included;  stigma  lobes 
closing  after  pollination.  Fruits  2-4  mm  long,  2- 
3  mm  diam.,  obovoid  and  bisulcate,  surface  gla- 
brous, smooth,  enclosed  within  the  persisting  ca- 
lyx cup;  seeds  oblong,  minutely  reticulate. 

Mazus  pumilus,  native  to  eastern  Asia,  is  rarely 
collected  in  the  Neotropics.  It  is  often  found  as  a 
weed  in  gardens  and  open  places  (1000-2000  m). 
The  species  is  recognized  by  its  small  size,  nar- 
rowly obovate  opposite  leaves  with  crenulate  mar- 
gins and  long-decurrent  blades,  few-flowered  ra- 
cemes with  long  pedicels,  partly  united  calyx,  and 
blue  or  pink  two-lipped  corolla.  These  plants  may 
resemble  species  of  Veronica,  but  that  genus  lacks 
the  strongly  two-lipped  flowers  and  four  stamens. 


Mecardonia  Ruiz  &  Pav6n 

RHFHRKNCE — F.  Pennell,  Reconsideration  of  the 
Bacopa-Herpestis  problem.  Proc.  Acad.  Nat.  Sci. 
Philadelphia  98:  83-98.  1946.  R.  Rossow,  Revi- 
si6n  del  g£nero  Mecardonia  (Scrophulariaceae). 
Candollea  42:  431-474.  1987. 

Erect  or  prostrate  herbs,  annual  or  perennial, 
stems  with  few  to  many  branches,  mostly  gla- 
brous, drying  brown  or  blackish.  Leaves  opposite, 


BURGER:  FLORA  COSTARICENSIS 


51 


sessile  or  short-petiolate,  blades  with  serrate  mar- 
gins, usually  glabrous,  glandular-punctate,  vena- 
tion pinnate  with  most  veins  diverging  in  the 
proximal  half  of  the  blade.  Inflorescences  of  sol- 
itary axillary  flowers  (2/node),  2  small  linear 
bracts  subtending  the  pedicel,  bracteoles  absent  at 
the  base  of  the  calyx,  pedicels  slender.  Flowers 
with  deeply  5-parted  calyx,  sepals  unequal  in 
width  with  3  broad  outer  ones;  corolla  tubular- 
campanulate,  slightly  2-lipped  and  sometimes 
with  a  prominent  palate  (personate),  yellow  or 
white,  upper  lip  slightly  2-lobed,  lower  lip  3- 
lobed;  stamens  4  in  2  pairs,  inserted  near  the  base 
of  the  tube,  thecae  stipitate  on  short  arms  of  the 
connective,  a  short  staminode  sometimes  present; 
ovary  2-locular,  ovules  many,  stigma  flattened,  2- 
lobed,  slightly  bent.  Fruits  thin-walled  capsules, 
dehiscence  loculicidal;  seeds  oblong,  surface  re- 
ticulate and  ridged. 

A  tropical  and  warm-temperate  American  ge- 
nus of  ca.  15  species,  the  majority  in  South  Amer- 
ican. The  species  have  sometimes  been  united 
with  those  of  Bacopa,  but  Pennell  (1946)  sepa- 
rated them  because  of  the  stipitate  anther-thecae 
and  the  yellow  pigmentation  of  the  corolla.  The 
anthers,  the  gland-dotted  foliage,  the  slightly  bi- 
lobed  deflexed  stigma,  and  the  loculicidal  capsule 
strengthen  the  distinction  and  indicate  a  close  re- 
lationship with  Stemodia.  In  Costa  Rica,  Mecar- 
donia  differs  from  Stemodia  in  having  yellow  co- 
rollas and  pedicels  with  small  bracts  at  the  base. 
However,  these  distinctions  do  not  hold  over  the 
entire  range  of  all  species,  and  it  may  be  neces- 
sary to  reevaluate  the  genera. 

Mecardonia  procumbens  (Mill.)  Small,  Fl.  S.E. 
U.S.  1065  &  1338.  1903.  Erinus  procumbens 
Mill.,  Card.  Diet.  ed.  8.  1768.  Lindernia  dianth- 
era  Sw.,  Prodr.  92.  1788.  Herpestis  caparioides 
Kunth  i/i  H.B.K.,  Nov.  Gen.  Sp.  2:  368.  1818. 
H.  peduncularis  Benth.,  Bot.  Mag.  2:  56.  1836. 
Bacopa  procumbens  (Mill.)  Greenman,  Publ. 
Field  Columb.  Mus.  Bot.  Ser.  2:  261.  1907.  Fig- 
ure 2. 

Herbs,  prostrate  to  procumbent,  stems  to  40  cm 
long,  distal  stems  few-branched,  sometimes  root- 
ing at  the  proximal  nodes,  leafy  stems  0.4-2  mm 
diam.,  glabrous,  with  4  longitudinal  ridges. 
Leaves  opposite,  subsessile  or  with  poorly  de- 
fined petioles  to  4  mm  long;  leaf  blades  6-24  mm 
long,  4-16  mm  wide,  ovate-elliptic  to  broadly  or 
narrowly  elliptic,  apex  obtuse,  margin  with  6-9 
teeth/cm,  base  obtuse  to  cuneate,  drying  charta- 


ceous  and  often  blackish,  glabrous,  2°  veins  2  or 
3/side.  Inflorescences  of  solitary  axillary  flowers, 
linear  bracts  ca.  2  mm  long  at  base  of  pedicel, 
pedicels  2-16(-24)  mm  long,  0.2-0.4  mm  diam., 
glabrous.  Flowers  glabrous  externally,  calyx  5-9 
mm  long,  outer  3  sepals  2.7-4  mm  wide,  ovate  to 
lanceolate;  corolla  8-10  mm  long,  tube  ca.  7  mm 
long,  yellow,  purplish  in  the  throat,  barbate  in  the 
mouth,  lobes  1-2  mm  long;  stamens  4,  filaments 
borne  on  the  lower  half  of  the  tube,  a  staminode 
sometimes  present.  Fruits  4-6  mm  long,  ovoid, 
dehiscent  from  the  apex,  placenta  spongy;  seeds 
0.4-0.5  mm  long,  ca.  0.2  mm  diam.,  oblong  or 
ovoid,  minutely  reticulate,  light  brown. 

Plants  of  open  sunny  or  partly  shaded  sites  on 
wet  soils  or  wet  sand  in  evergreen  or  deciduous 
areas,  0-1200(-2400)  m  elevation.  Flowering  and 
fruiting  throughout  the  year.  The  species  is  partic- 
ularly common  in  areas  supporting  evergreen  low- 
land rain  forest  formations.  The  species  ranges 
from  Mexico  to  Uruguay  and  Argentina. 

Mecardonia  procumbens  is  recognized  by  its 
short  herbaceous  stems,  general  lack  of  pubes- 
cence, subsessile  opposite  serrate  leaves,  solitary 
axillary  flowers  on  prominent  pedicels,  calyx  with 
wide  outer  sepals  and  narrow  inner  sepals,  and 
yellow  tubular  two-lipped  corolla.  These  plants 
resemble  species  of  Bacopa,  Lindernia,  and  Ste- 
modia, both  in  growth  habit  and  in  preference  for 
moist  open  sites.  This  species  can  be  separated 
from  those  in  the  other  genera  by  the  bright  yel- 
low corolla,  glabrous  gland-dotted  foliage  and 
bracteolate  pedicels.  This  species  is  very  closely 
related  to  M.  montevidensis  (Spreng.)  Kuntze  of 
southern  South  America. 


Micranthemum  Michaux 

REFERENCE — L.  O.  Williams,  Tropical  Ameri- 
can plants  XII.  Fieldiana,  Bot.  34:  101-132.  1972 
(Micranthemum  pp.  122-124). 

Small  slender  stemmed  herbs,  aquatic  or  grow- 
ing in  wet  places,  diffuse  and  much-branched, 
nodes  often  with  adventitous  roots.  Leaves  op- 
posite, sessile,  small,  blades  with  entire  margins, 
thin,  usually  glabrous,  venation  palmate.  Inflores- 
cences of  solitary  (rarely  2)  axillary  flowers,  usu- 
ally with  1  flower/node,  bracts  and  bracteoles  ab- 
sent, pedicels  short.  Flowers  minute,  calyx  with 
4  or  5  lobes  or  4  or  5  sepals,  lobes  equal,  tube 
short  or  absent,  glabrous  or  sparsely  puberulent; 
corolla  2-lipped,  tube  very  short,  adaxial  lip  short 


52 


FIELDIANA:  BOTANY 


and  2-lobed,  abaxial  lip  prominent  and  3-lobed; 
stamens  2,  filaments  inserted  at  the  mouth  of  the 
tube  abaxially,  anthers  2-thecous,  staminodes  ab- 
sent; ovary  1-  or  2-locular,  ovules  many,  style 
short,  stigma  2-lobed.  Fruits  dry  capsules,  glo- 
bose or  ovoid,  splitting  irregularly;  seeds  oblong, 
with  longitudinal  ridges. 

Micranthemum  is  a  genus  of  perhaps  10  to  12 
species,  ranging  from  the  southeastern  United 
States  to  southern  South  America.  It  is  most  di- 
verse in  Cuba.  The  genus  is  recognized  by  its 
small  slender-stemmed  aquatic  or  semiaquatic 
habit,  small  sessile  rounded  leaves,  minute  soli- 
tary flowers,  and  small  two-lipped  corolla  with 
two  distal  stamens.  These  delicate  little  plants  are 
sometimes  grown  in  freshwater  aquaria.  Although 
rarely  collected  in  Central  America,  it  is  repre- 
sented by  the  following  species  in  Costa  Rica. 

Micranthemum  umbrosum  (J.  F.  Gmelin)  Blake, 
Rhodora  17:  131.  1915.  Anonymous  umbrosa 
Walter,  Fl.  Carol.  63.  1788.  Globifera  umbro- 
sum (Walter)  J.  F.  Gmelin,  Syst.  Nat.  32.  1791. 
Figure  1. 

Herbs,  aquatic  or  in  wet  sites,  creeping  or  often 
forming  small  mats,  stems  to  ca.  1 2  cm  long,  leafy 
stems  0.2-0.8  mm  diam.,  glabrous,  with  2  longi- 
tudinal ridges.  Leaves  sessile,  base  of  opposing 
leaves  often  forming  a  line  or  ridge  across  the 
stem;  leaf  blades  3-8  mm  long,  2-7  mm  wide, 
ovate-orbicular  to  rounded-obovate,  apex  obtuse 
to  rounded,  margins  entire,  obtuse  or  rounded  at 
the  contracted  base,  drying  greenish  and  translu- 
cent, glabrous,  with  3  major  veins  and  2  lateral 
veins.  Inflorescences  of  solitary  axillary  flowers, 
usually  with  1  flower/node,  pedicels  0.5-1  mm 
long,  0.2-0.4  mm  diam.,  glabrous.  Flowers  with 
calyx  0.8-1.3  mm  long,  sepals  4,  equal  and  sep- 
arate nearly  to  the  base,  narrowly  oblong,  with 
few  minute  hairs  or  glabrous;  corolla  ca.  1 .5  mm 
long,  white,  tube  very  short;  stamens  2.  attached 
at  the  base  of  the  abaxial  sinuses.  Fruits  ca.  1 
mm  diam.,  globose,  thin-walled,  1-locular;  seeds 
0.3-0.4  mm  long,  ca.  0.2  mm  diam.,  oblong,  yel- 
lowish, with  minutely  ribbed  longitudinal  ridges 
(50X). 

Micranthemum  umbrosum,  ranging  from  the 
southeastern  United  States  to  South  America,  is 
rarely  collected  in  Central  America,  but  the  very 
small  size  of  the  plants  and  their  aquatic  or  wet 
habitats  may  have  resulted  in  the  species  being 
overlooked.  The  species  has  been  collected  in 
Honduras,  Nicaragua,  and  the  Tortuguero  region 
of  northeastern  Costa  Rica.  The  slender  glabrous 


stems,  small  opposite  .sessile  rounded  leaves,  mi- 
nute solitary  flowers,  white  two-lipped  corolla 
with  two  stamens,  and  thin-walled  capsules  with 
one  locule  help  distinguish  these  plants.  Micran- 
themum standleyi  Williams,  an  endemic  of  Gua- 
temala, has  smaller  flowers,  ovoid  fruits,  and  pi- 
lose calyx  lobes.  Micranthemum  pilosum  Ernst, 
from  Venezuela,  is  also  similar,  but  it  is  possible 
that  these  are  only  variants  of  M.  umbrosum  in  a 
broad  sense. 


Mimulus  Linnaeus 

Ri  H:RKNCK — A.  Grant,  A  monograph  of  the  ge- 
nus Mimulus.  Ann.  Missouri  Bot.  Gard.  11:  98- 
388.  1924. 

Annual  or  perennial  herbs  (rarely  shrubs),  of- 
ten preferring  moist  sites,  decumbent  or  erect, 
with  viscid  pubescence  or  glabrous.  Leaves  op- 
posite, simple,  sessile  or  petiolate,  blades  entire  or 
dentate,  venation  pinnate  or  palmate.  Inflores- 
cences of  solitary  flowers  in  leaf  axils  or  forming 
terminal  racemes,  bracts  absent  or  leaf-like,  ped- 
icels without  bracteoles  at  the  apex.  Flowers 
showy,  calyx  tubular  to  campanulate,  almost  as 
long  as  the  corolla,  with  5  prominent  veins,  tube 
longer  than  the  5  short  lobes  or  teeth,  lobes  usu- 
ally unequal;  corolla  tubular  or  narrowly  campan- 
ulate, 2-lipped,  lips  shorter  than  the  tube,  adaxial 
lip  2-lobed,  abaxial  lip  3-lobed.  lobes  subequal. 
usually  with  2  yellowish  protuberances  in  the 
throat;  stamens  4  in  2  unequal  pairs,  included  or 
exserted,  filaments  attached  near  the  base  of  the 
tube,  linear,  anthers  2-thecous  with  the  thecae  di- 
varicate and  confluent  at  the  apex;  ovary  2-locu- 
lar, ovules  many,  style  included,  stigma  flattened 
and  2-lobed.  Fruits  capsules,  oblong  to  linear, 
thin-  or  thick-walled,  dehiscence  loculicidal,  en- 
closed within  the  persisting  and  enlarged  calyx 
tube;  seeds  small,  smooth  or  reticulate. 

Mimulus  is  a  genus  of  ca.  150  species  ranging 
from  the  Americas  to  Africa  and  Asia  but  with 
most  of  the  species  in  western  North  America. 
Colorful  species  (monkey  flowers)  are  grown  in 
temperate  gardens.  No  specimens  of  this  genus 
have  been  collected  in  Costa  Rica,  but  the  follow- 
ing species  may  occur. 


Mimulus  glabratus  Kunth  in  H.B.K.,  Nov.  Gen. 
Sp.  2:  ed.  quarto  370.  1818.  Figure  1. 

Perennial  herbs,  procumbent  or  prostrate,  4-40 
cm  long,  rooting  at  many  lower  nodes,  leafy  stems 


BURGER:  FLORA  COSTARICENSIS 


53 


0.3-4  mm  diam.  (dried),  succulent  in  life,  gla- 
brous, nodes  with  an  interpetiolar  line  formed  by 
the  clasping  leaf  bases,  rooting  at  lower  nodes. 
Leaves  opposite,  subsessile  near  the  apex  of 
stems  or  with  petioles  2-18  mm  long,  0.4-2  mm 
wide,  with  lateral  margins  continuous  with  the 
blade  margins;  leaf  blades  8-45  mm  long,  4-35 
mm  wide,  ovate  to  ovate-oblong  or  ovate-orbic- 
ular, apex  obtuse  to  rounded,  margins  with  3-5 
prominent  teeth/cm,  base  obtuse  or  truncated,  dry- 
ing grayish  green,  glabrous,  venation  palmate 
with  3-5  major  veins.  Inflorescences  of  solitary 
flowers  axillary  to  foliage  leaves  (2/node),  pedi- 
cels 12-42  mm  long,  0.4-0.8  mm  diam.,  gla- 
brous. Flowers  with  calyx  5-10  mm  long  (14  mm 
in  fruit),  2.5-4  mm  diam.,  tubular,  upper  lobe 
larger  (ca.  4  mm)  than  lower  lobes  (ca.  2  mm), 
glabrous;  corolla  7-18  mm  long,  yellow  with  red 
spots  within  the  throat,  tube  2.5-4  mm  diam., 
lobes  1-2  mm  long.  Fruits  6-10  mm  long,  5-6 
mm  diam.,  included  within  the  calyx  tube;  seeds 
0.4-0.5  mm  long,  ca.  0.3  mm  diam.,  oblong, 
brown,  with  poorly  defined  longitudinal  ridges. 

Mimulus  glabratus  ranges  from  the  southern 
United  States  and  Mexico  to  Nicaragua  and  from 
Colombia  to  Chile.  In  tropical  environments  it 
grows  in  wet  sites  at  100-2200  m  elevation. 
These  plants  are  recognized  by  their  wet  habitat, 
creeping  stems  with  adventitous  roots,  lack  of  pu- 
bescence, opposite  petiolate  leaves,  solitary  axil- 
lary flowers  on  slender  pedicels,  tubular-plicate 
calyx  with  unequal  lobes,  and  yellow  two-lipped 
corolla  with  short  lobes.  Although  not  collected 
between  Nicaragua  and  Colombia,  this  species 
may  occur  in  our  area. 


Penstemon  Mitchell 

REFERENCE — R.  Straw,  The  penstemons  of 
Mexico.  II.  Penstemon  hartwegii,  Penstemon  gen- 
tianoides,  and  their  allies.  Bol.  Soc.  Bot.  Mexico 
27:  1-25.  1962. 

Perennial  herbs  or  subshrubs,  mostly  erect, 
stems  simple  or  branching  from  near  the  base, 
glabrous  or  pubescent.  Leaves  opposite,  the  lower 
leaves  often  petiolate,  distal  leaves  often  sessile 
and  clasping  the  stem  or  bract-like;  leaf  blades 
serrate  or  entire,  acute  to  rounded,  venation  pin- 
nate. Inflorescences  terminal  panicles  or  thyrses 
(often  racemiform),  the  partial  inflorescences  usu- 
ally cymose,  pedunculate  with  dichotomous 
branching,  subtended  by  leaf-like  or  small  bracts, 


glabrous  or  puberulent,  bracteoles  present  or  ab- 
sent. Flowers  small  or  large,  calyx  deeply  5-part- 
ed,  sepals  imbricate  in  bud,  equal  or  subequal; 
corolla  tubular  to  tubular-campanulate,  often  nar- 
rowed at  the  base  and  abruptly  expanded,  2- 
lipped,  mostly  blue  to  purple,  sometimes  red  or 
white,  upper  (adaxial)  lip  2-lobed  and  often  erect, 
lower  (abaxial)  lip  spreading  or  reflexed  and  3- 
lobed;  stamens  4  in  2  pairs,  shorter  than  the  co- 
rolla, bases  glandular  and  nectariferous,  filaments 
attached  near  the  base  of  the  tube,  anthers  gla- 
brous or  villous,  thecae  distinct  or  confluent,  a 
staminode  present  and  conspicuous;  ovary  ovoid, 
2-locular,  ovules  many,  style  slender,  stigma 
slightly  2-lobed  or  subcapitate.  Fruits  dry  cap- 
sules, ovoid  to  globose,  dehiscence  septicidal  and 
often  loculicidal,  valves  entire  or  bifid;  seeds 
many,  dark,  angular,  rugulose  to  smooth,  rarely 
winged  along  the  margins. 

Penstemon  is  one  of  the  largest  genera  of  Scro- 
phulariaceae,  with  ca.  300  species  ranging  from 
temperate  North  America  into  the  mountains  of 
Mexico  and  Guatemala.  The  greatest  number  of 
species  are  found  in  the  mountains  of  the  western 
United  States.  The  genus  is  identified  by  the  tu- 
bular to  campanulate  flowers  with  a  conspicuous 
staminode  oriented  on  the  lower  (abaxial)  side  of 
the  tube.  Several  species  are  colorful  ornamentals, 
grown  widely  in  temperate  or  tropical  montane 
gardens.  The  following  species  is  likely  to  be  seen 
in  Costa  Rican  gardens. 

Penstemon  gentianoides  (Kunth  in  H.B.K.)  Poir- 
et,  Diet.  Sci.  Nat.  38:  385.  1825.  Chelone  gen- 
tianoides Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  2: 
363.  tab.  172.  1818.  P.  skutchii  Straw,  Bol.  Soc. 
Bot.  Mexico  27:  13.  1962.  Figure  8. 

Erect  perennial  herbs  or  subshrubs  0.6-1.5  m 
tall,  branched  mostly  in  the  lower  half,  leafy 
stems  2-6  mm  diam.,  terete,  glabrous  or  with  thin 
straight  hairs  0.1-0.2  mm  long,  nodes  with  inter- 
petiolar ridges.  Leaves  opposite  (but  often  with 
axillary  short-shoots  and  giving  a  verticillate  ap- 
pearance), sessile  or  subsessile;  leaf  blades  4-12 
cm  long,  5-20  mm  wide,  lanceolate  to  linear-lan- 
ceolate, apex  acute,  margin  entire,  base  cuneate  to 
rounded,  drying  stiffly  chartaceous,  usually  gla- 
brous (except  at  the  base),  punctate  above  and  be- 
low, 2°  veins  often  obscure.  Inflorescences  10- 
35  cm  long,  racemiform  thyrses  with  axillary 
groups  of  (l-)3-9  flowers,  bracts  (reduced  leaves) 
1 2-30  mm  long,  usually  lanceolate,  peduncles  4- 
12  mm  long,  often  terminated  by  opposite  brac- 


54 


FffiLDIANA:  BOTANY 


teoles  3-8  mm  long,  pedicels  3-12  mm  long,  mi- 
nutely puberulent  or  subglabrous.  Flowers  with 
calyx  6-10  mm  long,  sepals  3-5  mm  wide,  broad- 
ly imbricate,  glabrous  or  puberulent;  corolla  2.5- 
4  cm  long,  tube  7-10  mm  diam.,  narrowed  near 
the  base,  deep  purple  to  rose,  red.  or  white,  gla- 
brous or  minutely  puberulent,  lobes  6-10  mm 
long,  throat  white;  filaments  glabrous,  anthers  in- 
cluded, staminode  ca.  22  mm  long;  ovary  5-7  mm 
long.  Fruits  8-12  mm  long,  5-6  mm  wide,  ovoid, 
style  base  persisting,  septicidal  and  loculicidal; 
seeds  black,  angular,  rugulose. 

Penstemon  gentianoides,  native  to  the  higher 
(2400-4000  m)  mountains  of  Mexico  and  Gua- 
temala, is  sometimes  cultivated  in  gardens  of  the 
Meseta  Central  and  at  higher  elevations.  The  spi- 
cate  or  racemiform  arrangement  of  verticillate 
groups  of  large  colorful  flowers,  with  tubular  co- 
rollas slightly  expanded  above  the  base,  and  the 
prominent  staminode  help  to  distinguish  these 
plants. 


Russelia  Jacquin 

REFERENCE — M.  Carlson,  Monograph  of  the  ge- 
nus Russelia.  Fieldiana,  Bot.  29:  231-292.  1957. 

Perennial  shrubs  or  herbaceous  subshrubs  from 
a  woody  base,  erect  or  with  pendent  or  scandent 
stems,  distal  branches  few  to  many,  stems  angular 
or  terete,  puberulent  or  glabrous.  Leaves  opposite 
or  whorled  (caducous  in  R.  equisetiformis),  sub- 


sessile  or  short-petiolate,  blades  usually  ovate  to 
lanceolate,  margin  entire  or  dentate  to  deeply  in- 
cised, glandular  (resinous)  peltate  scales  (dots)  of- 
ten present,  venation  pinnate.  Inflorescences  di- 
chotomous  cymes,  1-3  in  leaf  axils  or  subtended 
by  reduced  bract-like  leaves,  often  verticillate. 
subsessile  to  long  pedunculate,  linear  bracts  sub- 
tending the  slender  pedicels,  bracteoles  absent  at 
the  base  of  the  calyx.  Flowers  mostly  small  (1-2 
cm),  calyx  deeply  5-lobed  or  with  5  sepals,  ovate 
to  lanceolate,  imbricate  in  bud,  with  or  without 
peltate  glands;  corolla  tubular  to  funnelform, 
slightly  2-lipped.  red  to  pink  (white),  the  5  lobes 
slightly  unequal,  the  2  (upper)  adaxial  lobes  out- 
side in  bud,  lower  lip  3-lobed;  stamens  4  in  2 
unequal  pairs,  included,  filaments  borne  in  the 
lower  half  of  the  tube,  thecae  divaricate  and  con- 
fluent, a  short  staminode  usually  present;  ovary  2- 
locular,  ovules  many,  style  slender,  stigma  slightly 
thickened.  Fruits  dry  capsules,  ovoid  to  subglo- 
bose,  glabrous,  dehiscence  septicidal  (secondarily 
loculicidal);  seeds  small,  slightly  rugulose,  devel- 
oping among  translucent  fragmenting  hairs  (ela- 
ters)  produced  by  the  placenta. 

Russelia  is  a  genus  of  51  species  centered  in 
Mexico  and  Guatemala,  with  one  species  reaching 
Cuba  and  Colombia.  The  genus  has  been  placed 
in  its  own  tribe  because  of  the  unusual  hair-like 
fragments  in  the  capsule.  The  small  tubular  red 
flowers  (only  slightly  two-lipped)  in  axillary 
cymes  and  the  peltate  glands  (when  present)  also 
help  to  distinguish  members  of  this  genus.  Two 
species  are  found  in  Costa  Rica. 


Key  to  the  Species  of  Russelia 

la.  Common  wild  plants  of  seasonally  dry  habitats;  leaves  ovate,  subsessile,  1-9  cm  long;  stems  with 
opposite  branches;  inflorescences  >  2-flowered;  corollas  10-16  mm  long R.  sarmentosa 

Ib.  Plants  grown  in  gardens;  leaves  falling  off  early  and  stems  and  verticillate  branches  leafless;  inflo- 
rescences 2-flowered;  corollas  15-30  mm  long  R.  equisetiformis 


Russelia  equisetiformis  Schltdl.  &  Cham.,  Lin- 
naea  6:  377.  1831.  R.  juncea  Zucc.,  Flora  15, 
Beibl.  2:  99.  1832. 

Erect  subshrubs  to  1  m  tall,  much-branched, 
lower  nodes  with  whorls  of  4-8  branches,  distal 
nodes  with  2  branches  or  peduncles  at  each  node, 
longitudinal  ridges  2,  4,  or  6,  distal  stems  usually 
leafless,  0.7-6  mm  diam.,  glabrous  or  minutely 
puberulent  near  the  nodes  with  thin  hairs  0.1-0.3 
mm  long.  Leaves  opposite  or  verticillate,  usually 


caducous,  and  absent  at  distal  stems,  petioles  3- 
4  mm  long  and  often  remaining  appressed  to  the 
stems  when  blades  fall;  lower  larger  leaf  blades 
8-15  mm  long.  6-9  mm  wide,  ovate  to  elliptic. 
Inflorescences  2-16  cm  long,  often  resembling 
open  racemose  panicles,  usually  with  2-4  pedun- 
cles/node, each  peduncle  with  1  or  2  flowers,  bas- 
al peduncles  0.5-1.5  mm  diam.,  glabrous,  with  2 
or  4  prominent  longitudinal  ridges,  bracteoles  1- 
2  mm  long,  linear,  pedicels  6-15  mm  long,  0.2- 
0.3  mm  diam.  Flowers  glabrous  externally,  calyx 


BURGER:  FLORA  COSTARICENSIS 


55 


2-3  mm  long,  imbricate,  acute  or  acuminate,  gla- 
brous; corolla  15-30  mm  long,  2-4  mm  diam., 
tubular-funnelform,  bright  red,  glabrous  within; 
stamens  18  and  20  mm  long,  anthers  near  mouth 
of  the  tube,  staminode  present;  ovary  2  mm  long, 
ovoid,  style  15  mm  long,  stigma  minute.  Fruits 
3-6  mm  diam.,  globose;  seeds  light  brown, 
among  white  hairs  within  the  capsule. 

Russelia  equisetiformis  is  a  popular  garden 
plant  because  of  its  slender  green  leafless  stems 
and  bright  red  flowers.  It  is  distinctive  because  of 
the  ridged  stems  and  verticillate  arching  branches, 
which  resemble  those  of  Equisetum.  Common 
names  in  Central  America  are  coral  and  lluvia  de 
coral.  The  species  is  native  to  Mexico  and  is  an 
occasional  roadside  escape  in  the  Meseta  Central. 

Russelia  sarmentosa  Jacq.,  Enum.  PI.  Syst.  6:  25. 
1760.  R.  colombiana  Pennell,  Proc.  Acad.  Nat. 
Sci.  Philadelphia  72:  186.  1920.  R.  flavoviridis 
Blake,  Contrib.  U.S.  Natl.  Herb.  24:  22.  1922. 
R.  tabacensis  Lundell,  Contrib.  Univ.  Michigan 
Herb.  6:  59.  1941.  R.  oxyphylla  Lundell,  loc. 
cit.  7:  51.  1942.  Figure  5. 

Herbs  or  weak-stemmed  shrubs  0.5-2  m  tall, 
erect  or  spreading,  base  woody,  distal  flowering 
stems  usually  unbranched,  leafy  stems  1-5  mm 
diam.,  strongly  4-  or  6-angled  with  raised  longi- 
tudinal ridges,  puberulent  with  thin  whitish  hairs 
0.1-0.3  mm  long  or  glabrous  (except  at  the 
nodes).  Leaves  2  or  3  at  each  node,  subsessile 
with  petioles  0.5-4  mm  long,  often  forming  an 
interpetiolar  line  across  the  stem,  lateral  margins 
puberulent;  leaf  blades  l-7(-9)  cm  long,  l-5(-6) 
cm  wide,  ovate  to  broadly  ovate  or  ovate-trian- 
gular, apex  acute,  margin  with  2-5  teeth/cm,  teeth 
0.5-2  mm  long,  obtuse  or  truncated  at  the  base, 
drying  dark  or  grayish,  glabrous  or  puberulent 
above,  usually  more  pubescent  beneath  with  hairs 
0.2-0.4  mm  long,  peltate  glands  (ca.  0.2  mm 
diam.)  often  present  on  both  surfaces,  2°  veins  3- 
5/side  and  strongly  ascending.  Inflorescences  2- 
3  cm  long,  congested  verticels  of  cymes  in  distal 
leaf  axils,  12-20  flowers/cyme,  peduncles  2-18 
mm  long,  resembling  the  stems,  bracts  3-4  mm 
long,  linear,  pedicels  5-8  mm  long.  Flowers  with 
calyx  3-4  mm  long,  ca.  1.3  mm  diam.,  lobes  2- 
3  mm  long,  with  thin  lateral  margins  and  acumi- 
nate apex,  often  with  peltate  glands;  corolla  10- 
14(-16)  mm  long,  1.5-3  mm  diam.,  tubular,  deep 
red,  puberulent  within,  lobes  subequal,  1 .3-3  mm 
long;  stamens  ca.  8  and  6  mm  long,  inserted  near 
the  middle  of  the  tube,  anthers  white;  ovary  1-2 


mm  long,  style  4-8  mm  long.  Fruits  4-5  mm 
long  (not  including  the  0.5-4  mm  style  base),  4- 
5  mm  diam.,  rounded-ovoid  to  subglobose,  drying 
dark;  seeds  0.4-0.5  mm  long,  ca.  0.2  mm  wide, 
dark,  among  yellowish  hairs. 

Common  plants  of  open  weedy  sites  and  forest 
edges  in  deciduous  and  partly  deciduous  (rarely 
evergreen)  forest  formations,  0-1300  m  elevation. 
Flowering  and  fruiting  throughout  the  year  but 
collected  most  often  in  July-August  and  Decem- 
ber-January. The  species  ranges  from  Mexico  to 
Cuba  and  Colombia. 

Russelia  sarmentosa  is  recognized  by  the  often 
shrub-like  habit,  stiff  few-branched  distal  stems 
with  two  or  three  subsessile  leaves  at  each  node, 
short  cymes  often  forming  compact  verticels  of 
flowers  and  fruits,  and  bright  red  tubular  corollas. 
The  prominent  longitudinal  ridges  on  stems,  pel- 
tate glands  on  leaf  surfaces  and  calyx  (when  pre- 
sent), and  sepals  broadly  overlapping  at  the  base 
are  also  helpful  in  recognizing  this  common  spe- 
cies. We  agree  with  Margery  Carlson's  delimita- 
tion (1957,  cited  above)  of  this  wide-ranging  and 
variable  species,  but  we  do  not  believe  that  the 
recognition  of  forms  and  varieties  is  useful.  The 
common  name  is  coralillo. 


Schistophragma  Bentham  ex  Endlicher 

Small  annual  herbs,  erect  or  procumbent,  stems 
quadrangular  and  with  longitudinal  ridges,  gla- 
brous or  sparsely  minutely  puberulent,  drying  yel- 
lowish brown.  Leaves  opposite,  petiolate,  blades 
deeply  pinnatisect  with  few  opposite  or  alternate 
narrow  lobes  (smaller  leaves  sometimes  linear-ob- 
lanceolate),  margins  entire  or  with  a  few  distal 
teeth,  venation  pinnate.  Inflorescences  of  solitary 
flowers  axillary  to  distal  leaves,  bracts  and  brac- 
teoles  absent,  pedicels  slender.  Flowers  small,  ca- 
lyx divided  to  near  the  base,  sepals  or  calyx  lobes 
5,  narrow,  acute,  subequal;  corolla  tubular  and 
slightly  2-lipped,  upper  (adaxial)  lip  with  1  emar- 
ginate  lobe,  lower  (abaxial)  lip  3-lobed;  stamens 
4,  in  2  subequal  pairs,  included;  ovary  narrowly 
ellipsoid,  2-locular,  ovules  many,  style  slender,  di- 
lated at  apex.  Fruits  linear  capsules,  somewhat 
compressed  and  bisulcate,  septicidal;  seeds  ob- 
long, surface  with  compressed  spiral  ridges. 

The  delimitation  of  Schistophragma  and  its  dis- 
tinction from  Leucospora  Nuttall  and  Conobea 
Aubl.  is  the  subject  of  disagreement.  B.  L.  Turner 
(Turner  &  Cowan,  Phytologia  74:  61-103,  1993) 
recognizes  three  species  of  Schistophragma  and 


56 


FIELDIANA:  BOTANY 


includes  Leucospora  within  Stemodia.  D.  Keil 
(pers.  comm.)  includes  the  species  of  Schisto- 
phragma  in  his  concept  of  Leucospora,  which  he 
retains  a.s  distinct  from  Stemodia.  Schistophragma 
is  placed  in  the  tribe  Gratioleae,  subtribe  Stemo- 
diinae.  The  spirally  striate  seeds  are  very  unusual 
in  the  Scrophulariaceae,  but  seeds  of  this  type  are 
sometimes  found  in  the  Gesneriaceae.  The  small 
size  of  the  plants,  narrow  or  pinnatisect  leaves, 
small  tubular  flowers,  and  very  narrow  capsules 
make  these  plants  distinctive.  The  linear  fruits  re- 
semble those  of  some  Brassicaceae.  Among 
American  Scrophulariaceae,  only  Mabrya  Elisens, 
from  Mexico,  has  similar  linear  fruits. 

Schistophragma  mexicana  Benth.  ex  D.  Dietr, 
Syn.  PI.  3:  513.  1842.  S.  pusilla  Benth.  in  DC, 
Prodr.  10:  392.  1846.  Conobea  pusilla  (Benth.) 
Benth.  &  Hook,  ex  Jackson,  Index  Kew  1,  fasc. 
1:  596.  1893.  Stemodia  siliguosa  Sesse"  &  Mo- 
cino,  PI.  Nov.  Hisp.  ed.  1:  98.  1887-1890,  ed. 
2:  91.  1893.  Figure  4. 

Erect  to  decumbent  annual  herbs  6-25  cm  tall, 
with  few  or  many  branches,  leafy  stems  0.3-1  mm 
diam.,  with  4  longitudinal  ridges  (quadrangular  in 
cross-section),  glabrous  or  with  few  minute  hairs, 
nodes  lacking  interpetiolar  lines.  Leaves  opposite, 
deeply  pinnatifid,  petioles  2-7  mm  long  (to  first 
lobe),  undifferentiated  from  the  narrow  rachis,  0.2- 
0.5  mm  wide,  sulcate  adaxially;  leaf  blades  4-18 
mm  long,  to  12  mm  wide,  deeply  pinnatisect  with 
3-7  narrow  lobes  (including  the  distal  extension  of 
the  rachis),  lobes  0.3-2  mm  wide,  entire  or  the 
larger  with  1  or  2  lateral  teeth,  glabrous  or  mi- 
nutely puberulent  on  the  veins  beneath,  proximal 
lobes  rarely  with  1  or  2  2°  lobes  (bipinnatifid).  In- 
florescences of  solitary  axillary  flowers,  usually  2/ 
node,  pedicels  3-5  mm  long,  ca.  0.15  mm  diam., 
glabrous  or  subglabrous.  Flowers  with  calyx  2-4 
mm  long,  sepals  united  only  at  the  base,  ca.  0.3 
mm  wide,  glabrous;  corolla  5-6.5  mm  long,  violet 
to  purple  or  bluish  purple,  tube  ca.  4  mm  long,  1 .5 
mm  diam.;  ovary  glabrous,  style  2-3  mm  long. 
Fruits  8-12  mm  long,  0.9-1.2  mm  wide,  linear, 
smooth  and  bisulcate,  drying  blackish,  apex  acute, 
style  deciduous  or  sometimes  persisting;  seeds  0.7- 
0.9  mm  long,  0.3-0.4  mm  diam.,  cylindric  with 
truncated  ends,  brown,  the  closely  spaced  ridges 
forming  spirals  around  the  surface. 

Rarely  collected  plants  of  moist  areas  in  open 
sunny  sites,  0-1 100  m  elevation  in  Central  Amer- 


ica. Probably  flowering  primarily  in  the  wet  sea- 
son, June-December.  Only  two  Costa  Rican  col- 
lections have  been  seen:  A.  Jimenez  984,  from 
near  Playa  Coco,  Guanacaste,  and  Tonduz  13790, 
from  Nicoya,  Puntarenas.  The  species  ranges  from 
Mexico  to  northwestern  Costa  Rica  and  has  been 
collected  in  Colombia. 

Schistophragma  mexicana  is  very  distinctive 
because  of  its  small  stature,  small  opposite  pin- 
natifid leaves  with  narrow  lobes,  small  flowers 
with  tubular  purple  or  violet  corollas,  and  dark 
linear  fruits.  This  is  the  only  Costa  Rican  species 
of  Scrophulariaceae  with  linear  fruits  and  one  of 
the  few  with  pinnatifid  leaves  (compare  Benja- 
min ia  and  Castilleja). 


Scoparia  Linnaeus 

Herbs  or  subshrubs,  annual  or  perennial,  pro- 
fusely branched,  leafy  stems  slender,  4-  to  6-an- 
gled.  Leaves  opposite  or  verticillate,  2-4/node, 
sometimes  caducous  or  reduced  to  scales,  petioles 
little  differentiated  from  the  blade,  blades  dentate 
to  pinnatifid  or  entire,  base  cuneate,  often  glan- 
dular-punctate, venation  pinnate.  Inflorescences 
of  1  or  2  flowers  in  leaf  axils  (1-8/node),  bracts 
and  bracteoles  absent,  pedicels  filiform.  Flowers 
small,  calyx  4-  or  5-parted,  sepals  ovate  to  lan- 
ceolate, 1  often  larger  than  the  others;  corolla 
subrotate  with  very  short  tube  and  4  spreading  or 
reflexed  lobes  (upper  lip  emarginate),  white  to 
blue,  rose,  or  yellow,  with  a  ring  of  hairs  at  the 
mouth  of  the  tube;  stamens  4,  subequal  or  equal, 
inserted  at  the  base  of  the  tube,  anthers  sagittate 
with  parallel  or  subequal  thecae;  ovary  2-locular, 
ovules  many,  style  short,  stigma  subcapitate,  ex- 
serted.  Fruits  thin-walled  capsules,  globose  to 
ovoid,  dehiscence  septicidal;  seeds  small,  reticu- 
late with  pitted  surface. 

Scoparia  is  a  Neotropical  genus  of  ca.  1 7  species 
with  centers  of  diversity  in  Mexico  and  southern 
South  America.  The  plants  are  native  to  dry  scrub 
and  grasslands,  and  some  species  have  become  suc- 
cessful weeds.  Scoparia  is  distinctive  within  the 
family  because  of  its  usually  four-lobed  corolla  with 
very  short  tube  and  conspicuous  hairs.  Unfortunate- 
ly, the  corollas  are  fugaceous,  and  the  characters  of 
erect  habit,  solitary  or  paired  flowers,  four  calyx 
lobes,  and  crenatc  or  pinnatifid  leaves  must  be  used 
for  identification.  One  species  occurs  in  Costa  Rica, 
and  a  second  is  likely  to  be  found  here. 


BURGER:  FLORA  COSTARICENSIS 


57 


Key  to  the  Species  of  Scoparia 

la.  Corolla  white  or  pale  lilac;  sepals  4,  obtuse  or  rounded  at  the  apex;  leaf  blades  to  35  mm  long; 

plants  usually  becoming  >40  cm  tall;  Mexico  to  Costa  Rica   S.  dulcis 

Ib.  Corolla  yellow;  sepals  4  or  5,  acute  at  the  apex;  leaf  blades  to  20  mm  long;  plants  <30  cm  tall; 

Mexico  to  Nicaragua .5.  annua 


Scoparia  annua  Schldl.  &  Cham.,  Linnaea  6: 
375.  1831. 

Small  erect  annual  herbs  10-20(-30)  cm  tall, 
much-branched  from  the  base  or  lower  half,  leafy 
stems  0.4-1.5  mm  diam.,  glabrous,  with  4  prom- 
inent longitudinal  ridges.  Leaves  opposite,  peti- 
oles not  clearly  differentiated  from  the  blades, 
glabrous;  leaf  blades  5-20  mm  long  (including 
the  narrowed  base),  lower  blades  ovate  to  rhombic 
and  pinnatifid  with  rounded  lobes  to  4  mm  long, 
distal  blades  linear-oblanceolate  and  entire  or  with 
a  few  distal  teeth,  glandular  punctate  beneath.  In- 
florescences of  solitary  flowers  in  leaf  axils  (21 
node),  pedicels  6-18  mm  long,  filiform,  glabrous. 
Flowers  with  calyx  2  mm  long,  sepals  4-  or  5- 
parted,  lanceolate-oblong  with  acute  apices,  gla- 
brous; corolla  to  3  mm  long,  rotate,  yellow,  fu- 
gaceous,  lobes  2  mm  long,  rounded;  stamens 
equal,  filaments  glabrous;  ovary  ellipsoid,  style 
shorter  than  the  ovary,  stigma  minute.  Fruits  2.5- 
3  mm  long,  1 .5-2  mm  wide,  ovoid,  subtended  by 
sepals  ca.  3  mm  long;  seeds  reticulate. 

Scoparia  annua  ranges  from  Mexico  to  central 
Nicaragua  in  open  sunny  habitats  from  sea  level 
to  1000  m  elevation;  it  is  also  found  in  South 
America.  These  small  weedy  plants  are  likely  to 
be  overlooked,  and  it  is  possible  that  they  will  be 
found  in  northwestern  Costa  Rica. 


Scoparia  dulcis  L.,  Sp.  PI.  116.  1753.  Figure  4. 

Herbs  or  subshrubs  0.3-1  m  tall,  woody  at  the 
base,  often  with  many  lateral  branches,  leafy 
stems  0.5-4  mm  diam.,  4-  or  6-angled  with  4  or 
6  longitudinal  ridges,  glabrous  or  puberulent  at 
the  nodes,  nodes  with  interpetiolar  line.  Leaves  2 
or  3  (4)  at  each  node,  petioles  1-8  mm  long  (poor- 
ly differentiated),  0.3-0.9  mm  wide;  leaf  blades 
6-35  mm  long,  2-15  mm  wide,  narrowly  elliptic 
to  oblanceolate  (basal  leaves  ovate  to  ovate-rhom- 
bic), apex  obtuse  to  acute,  with  5-8  teeth/side  dis- 
tally,  teeth  0.3-3  mm  long,  base  obtuse  to  cune- 
ate,  drying  grayish  green,  glabrous,  glandular 
punctate  on  both  surfaces,  2°  veins  2  or  3/side. 


Inflorescences  of  1-3  flowers  axillary  to  distal 
leaves  (leaves  often  reduced  to  6  mm  long  and 
bract-like),  nodes  with  2-6  flowers,  pedicels  3-9 
mm  long,  0. 1-0.2  mm  diam.,  glabrous  or  minutely 
puberulent.  Flowers  with  4-parted  calyx,  2  mm 
long  (-3  mm  in  fruit),  lobes  ca.  1.3  mm  long, 
elliptic  to  oblong,  apex  obtuse  or  rounded;  corolla 
rotate  and  3-4  mm  wide  distally,  pale  lilac  or 
white,  white-barbate  within  the  throat;  stamens  ca. 
1.5  mm  long,  anthers  0.7  mm  long.  Fruits  3  mm 
long,  2-3  mm  diam.,  broadly  ovoid,  smooth,  yel- 
lowish gray;  seeds  0.3-0.4  mm  long,  0.2-0.3  mm 
diam.,  oblong  or  irregularly  angled,  pale  brown, 
pits  in  longitudinal  rows. 

Common  plants  of  open  sunny  sites  in  decid- 
uous, partly  deciduous,  and  wet  evergreen  forest 
formations,  0-1300  m  elevation.  Flowering  and 
fruiting  throughout  the  year  but  collected  most  of- 
ten in  the  wet  season,  June-December.  The  spe- 
cies ranges  from  the  southeastern  United  States  to 
southern  South  America  and  is  a  pantropical 
weed. 

Scoparia  dulcis  is  recognized  by  its  short, 
much-branched  herbaceous  habit,  nodes  with  usu- 
ally two  or  three  leaves,  dentate  leaf  blades,  small 
flower  with  white  rotate  corollas,  and  thin-walled 
fruits  with  pitted  seeds.  This  species  is  a  common 
weed  in  the  lowlands  of  Central  America.  The 
bushy  stems  have  been  used  to  make  small 
brooms  and  are  believed  to  repel  fleas  (Standley 
&  Williams,  1973).  The  plants  have  an  aromatic 
odor  and  are  called  sweet  broom;  culantrillo,  es- 
coba  amarga,  escobeta,  and  escobilla  amarga  are 
Spanish  names. 


Sibthorpia  Linnaeus 

REFERENCE — O.  Hedberg,  A  taxonomic  revision 
of  the  genus  Sibthorpia.  Bot.  Not.  108:  161-183. 
1955. 

Prostrate  or  creeping  herbs  rooting  at  many 
nodes,  perennial,  internodes  often  elongated, 
stems  terete,  with  multicellular  hairs,  plants  dry- 
ing greenish  or  brown.  Leaves  alternate  or  crowd- 


58 


FIELDIANA:  BOTANY 


ed,  petioles  often  long,  slender,  blades  orbicular 
to  reniform,  margin  crenate  with  broad  rounded 
or  truncated  lobes,  base  cordate,  villous  to  gla- 
brous, venation  palmate.  Inflorescences  of  1-3 
flowers  in  axils  of  leaves,  bracts  and  bracteoles 
absent,  pedicels  long  and  slender.  Flowers  very 
small,  almost  radially  symmetric,  calyx  campan- 
ulate,  united  in  the  basal  half  with  5  (4-8)  lobes, 
lobes  subequal,  acute,  persisting  in  fruit;  corolla 
subrotate  with  very  short  tube  and  5  (4-8)  spread- 
ing lobes,  white,  yellow,  pink,  or  reddish  purple, 
lobes  rounded;  stamens  2-8,  as  many  as  the  co- 
rolla lobes  or  1  or  2  fewer,  borne  near  the  apex 
of  the  tube,  filaments  slender,  subequal,  anthers 
slightly  sagittate,  thecae  parallel  or  slightly  diver- 
gent, confluent  near  the  apex,  staminodes  absent; 
ovary  pubescent,  ovules  few,  style  short,  stigma 
simple  or  capitate.  Fruits  thin-walled  capsules, 
dehiscence  loculicidal;  seeds  few,  oblong  to 
ovate,  convex  on  1  surface,  reticulate  or  smooth. 
Sibthorpia  is  a  genus  of  six  species  found  in 
Europe,  the  high  mountains  of  Africa,  and  higher 
elevations  in  the  Neotropics.  The  creeping  growth 
form,  small  alternate  cordate  reniform  or  orbicular 
leaves  with  crenate  margins,  very  small  axillary 
flowers,  almost  radially  symmetric  corolla,  short 
corolla  tube,  and  few-seeded  capsules  distinguish 
this  genus.  Seed  morphology  indicates  that  this 
genus  is  related  to  Veronica. 

Sibthorpia  repens  (Mutis  ex  L.)  Kuntze,  Rev. 
Gen.  PI.  3;  239.  1898.  Willichia  repens  Mutis 
ex  L.,  Mant.  PI.  2:  558.  1771.  5.  pichinchensis 
Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  2:  390,  tab. 
176.  1817.  5.  triandra  Suesseng.,  Repert.  Sp. 
Nov.  Regni  Veg.  39:  18.  1935. 

Creeping  herbs,  usually  with  long  internodes 
and  few  lateral  branches,  rooting  at  most  nodes, 
leafy  stems  0.5-1  mm  diam.,  with  slender  trans- 
lucent multicellular  hairs  0.2-0.8  mm  long,  older 
stems  usually  glabrescent.  Leaves  alternate,  peti- 
oles 2-22(-30)  mm  long,  0.3-0.8  mm  diam.,  with 
translucent  hairs  to  0.8  mm  long;  leaf  blades  6- 
22  mm  long,  6.5-25  mm  wide,  ovate-suborbicular 
to  rounded-reniform,  apex  a  rounded  lobe,  margin 
crenate  with  5-9  rounded  or  truncated  lobes/side, 
lobes  0.5-4.5  mm  wide,  base  cordate  with  sinus 
1-7  mm  deep,  drying  membranaceous,  translu- 
cent hairs  sparse  to  dense  on  both  surfaces,  major 
veins  3  or  5.  Inflorescences  of  1  (2)  flowers  in 
axils  of  leaves,  pedicels  3-12  cm  long  (to  4  cm 
long  in  Mexico),  filiform,  puberulent.  Flowers 
with  calyx  1.7-2  mm  long,  united  and  campanu- 


late,  lobes  4  or  5,  0.5-1  mm  long,  acute;  corolla 
2-3  mm  long,  3.5-5  mm  wide,  subrotate,  pale  li- 
lac to  dull  purple  to  brown-purple  or  dark  wine- 
red  (white  in  Ecuador),  tube  very  short,  lobes  0.7- 
1 .3  mm  wide;  stamens  2-4;  style  2  mm  long,  stig- 
ma capitate.  Fruits  ca.  2  mm  long,  subglobose, 
lower  part  enclosed  within  the  calyx  cup,  surface 
with  straight  ascending  hairs  0.1-0.2  mm  long, 
persisting  style  ca.  0.8  mm  long;  seeds  ca.  0.7  mm 
long,  oblong,  dark  with  a  whitish  reticulum. 

Rarely  collected  herbs  of  partly  shaded  to  deep- 
ly shaded  sites  in  evergreen  montane  forest  for- 
mations, 1600-3100  m  elevation.  Collected  with 
flowers  in  April-June  (flowering  primarily  in  De- 
cember-March in  northern  Central  America). 
These  plants  are  locally  common  in  open  high- 
elevation  forests  of  Guatemala,  but  we  have  seen 
only  five  collections  from  Costa  Rica.  This  spe- 
cies ranges  from  central  Mexico  to  Costa  Rica  and 
is  found  in  the  Andes. 

Sibthorpia  repens  is  recognized  by  the  creeping 
habit,  slender  stems  with  many  adventitous  roots, 
small  suborbicular  leaves  with  lobed  crenate  mar- 
gins, small  axillary  flowers,  subrotate  corollas 
with  two  epipetalous  stamens,  and  few-seeded 
capsules  enclosed  at  their  base  by  the  campanu- 
late  calyx.  This  species  is  quite  variable  in  a  num- 
ber of  significant  taxonomic  features.  Despite  this 
variability,  it  seems  probable  that,  as  Hedberg 
(1955,  cited  above)  and  Williams  (1972)  have 
suggested,  the  Neotropical  material  represents  a 
single  species.  These  plants  are  easily  mistaken 
for  species  of  Hydrocoryle  (Apiaceae)  or  Dichon- 
dra  (Convolvulaceae)  with  similar  growth  form. 


Stemodia  Linnaeus 
Nomen  conservandum 

RKPKRKNCKS — B.  Turner  &  C.  Cowan,  Taxo- 
nomic overview  of  Stemodia  (Scrophulariaceae) 
for  North  America  and  the  West  Indies.  Phytolo- 
gia  74:  61-103.  1993.  Taxonomic  overview  of 
Stemodia  (Scrophulariaceae)  for  South  America. 
Phytologia  75:  281-324.  1993. 

Erect  or  procumbent  herbs  or  subshrubs  to  3 
m  tall,  annual  or  perennial,  stems  terete  or  angu- 
lar, often  much-branched,  usually  puberulent,  of- 
ten with  gland-tipped  hairs,  drying  greenish  or 
brown.  Leaves  opposite,  subopposite,  or  verticil- 
late,  simple,  sessile  or  petiolate,  blades  serrulate 
to  pinnately  lobed  (rarely  subentire),  pinnately 
veined  or  subpalmate,  often  glandular-punctate. 
Inflorescences  of  1-3  flowers  in  axils  of  leaves 
or  bracts  (1-6/node),  sometimes  forming  distal  ra- 
cemes, thyrses,  or  panicles,  bracts  absent  when 


BURGER:  FLORA  COSTARICENSIS 


59 


flowers  are  subtended  by  leaves,  pedicels  slender, 
bracteoles  present  or  absent  at  the  base  of  the  ca- 
lyx. Flowers  with  deeply  5-parted  calyx,  sepals 
narrow  and  acute,  equal  or  more  often  slightly 
unequal  (adaxial  sepal  often  larger),  valvate  in 
bud,  persisting  and  enlarging  slightly  in  fruit;  co- 
rolla tubular  to  narrowly  campanulate,  2-lipped, 
white  to  blue,  lilac,  or  purple  (yellowish  when 
faded),  4-  or  5-lobed,  bilabiate,  upper  (adaxial)  lip 
2-lobed  or  entire,  lower  lip  3-lobed,  often  bearded 
at  the  base  of  the  lobes;  stamens  4,  equal  or  of  2 
unequal  pairs,  included,  borne  on  the  tube,  fila- 
ments glabrous  or  puberulent,  anthers  glabrous, 
with  an  enlarged  connective  separating  the  2  par- 
allel or  divergent  thecae,  a  staminode  sometimes 
present;  ovary  ovoid,  style  terete,  stigma  ligulate, 
often  minutely  2-lobed  or  capitate  and  reflexed. 
Fruits  dry  capsules,  often  bisulcate,  dehiscing  lo- 
culicidally  and  partly  septicidally  from  the  apex 
(2-  or  4-valved),  placenta  drying  to  form  a  peg 
that  is  free  from  the  apex  of  the  locule;  seeds 
many,  small,  oblong  to  pyriform  or  irregular,  often 
longitudinally  sulcate  or  ridged,  usually  with  a 
short  stipe  at  one  end. 


According  to  Turner  and  Cowan  (1993)  Ste- 
modia  is  a  genus  of  29  Neotropical  and  20  Old 
World  species,  including  several  widespread 
weeds.  They  are  placed  in  subfamily  Antirrhino- 
ideae,  tribe  Gratioleae,  subtribe  Stemodiinae.  Be- 
cause of  the  variability  of  important  taxonomic 
characteristics  within  the  genus,  it  is  likely  that 
Stemodia  may  be  broken  up  into  smaller  genera 
(see  Darcyd).  Mined  (1918)  split  the  genus  into  a 
number  of  genera  that  have  generally  not  been 
accepted.  His  characters  were  often  variable  and 
his  nomenclature  was  flawed,  which,  as  much  as 
his  splitting,  accounts  for  the  lack  of  interest  in 
his  classification.  We  follow  Turner  and  Cowan's 
treatment  and  their  annotations.  These  generally 
short,  weedy  plants  are  recognized  by  their  op- 
posite or  ternate  leaves,  one-  to  six-flowered 
nodes,  calyx  with  narrow  sepals  united  only  near 
the  base,  tubular  two-lipped  bluish  to  purple  or 
white  corollas,  and  style  often  curved  at  the  ter- 
minal stigma.  Other  important  characters  are  the 
anthers  with  thecae  slightly  separated  by  an  ex- 
panded connective,  and  the  seeds  often  with  lon- 
gitudinal ridges  or  sulci. 


Key  to  the  Species  of  Stemodia 

la.  Flowers  and  fruits  borne  on  slender  pedicels  6-19  mm  long;  corollas  white  to  pink  or  yellowish  (if 
flowers  have  a  prominent  calyx  tube,  see  Torenia)  2 

Ib.  Flowers  and  fruits  subsessile  or  on  pedicels  <  4  mm  long;  corollas  blue  to  purple 3 

2a.  Flowers  7-9  mm  long;  fruits  4-5  mm  long;  plants  to  20  cm  tall 5.  angulata 

2b.  Flowers  12-15  mm  long;  fruits  5-7  mm  long;  plants  to  80  cm  tall 5.  peduncularis 

3a.  Fruits  at  least  3  mm  long;  calyx  4-6  mm  long;  flowers  on  spike-like  axes;  leaves  subsessile,  larger 
leaves  to  6  cm  long  and  oblanceolate 5.  durantifolia 

3b.  Fruits  1.8-2.5  mm  long;  calyx  2-4  mm  long;  flowers  in  leaf  axils;  leaves  petiolate,  larger  leaves  to 
2  cm  long  and  ovate-triangular S.  verticillata 


Stemodia  angulata  Oerst.,  Vidensk.  Meddel. 
Dansk  Naturhist.  Foren.  Kjobenhavn  1853:  22. 
1854.  Stemodiacra  angulata  (Oerst.)  Kuntze, 
Rev.  Gen.  PI.  2:  466.  1891.  Stemodia  jorullensis 
subsp.  reptans  Minod,  Bull.  Soc.  Bot.  Geneve, 
ser  2,  10:  190.  1918.  Figure  2. 

Small  herbs  5-20  cm  tall,  stems  to  40  cm  long, 
often  much-branched,  leafy  stems  0.3-1.7  mm 
diam.,  pubescence  of  thin  multicellular  translucent 
hairs  0.2-1.2  mm  long,  slightly  viscid.  Leaves  op- 
posite, petioles  2-9  mm  long,  0.2-0.5  mm  wide, 
puberulent  with  thin  whitish  hairs;  leaf  blades  6- 
20  mm  long,  3-18  mm  wide,  ovate  to  ovate-tri- 
angular or  ovate-rhombic,  apex  obtuse  or  round- 


ed, margin  with  7-1 1  teeth/side,  base  broadly  ob- 
tuse or  truncate,  glandular  punctate  and  sparsely 
puberulent  above  and  below,  2°  veins  3-5/side, 
ascending.  Inflorescences  of  1  (2  or  3)  flowers  in 
leaf  axils,  pedicels  4-12  mm  long  (-19  mm  in 
fruit),  0.1-0.2  mm  diam.,  puberulent,  bracteoles 
absent.  Flowers  with  calyx  4-6  mm  long,  sepals 
3-5  mm  long,  ca.  0.8  mm  wide  at  the  base,  pu- 
berulent; corolla  7-9  mm  long,  white  with  purple 
lines  within  (yellowish  in  age),  tube  to  7  mm  long, 
1.4-2  mm  diam.,  lobes  to  2  mm  long;  stamen 
pairs  very  unequal,  2  borne  at  the  base  and  2  in 
the  upper  part  of  the  tube.  Fruits  4-5  mm  long, 
ca.  2  mm  diam.,  oblong,  pale  yellowish  brown, 
glabrous;  seeds  0.3-0.4  mm  long,  0.2-0.3  mm 


60 


FIELDIANA:  BOTANY 


diam.,  variously  shaped,  brown  to  black,  longi- 
tudinally sulcate  with  6-8  ridges. 

Plants  of  open  sunny  sites  in  both  seasonally 
dry  and  wet  evergreen  forest  areas,  10-1400  m 
elevation.  Probably  flowering  and  fruiting 
throughout  the  year,  but  collected  most  often  in 
August-March.  The  species  ranges  from  Mexico 
and  the  West  Indies  to  northwestern  South  Amer- 
ica. 

Stemodia  angulata  is  recognized  by  its  short 
stature,  opposite  ovate-triangular  leaves,  solitary 
axillary  flowers  on  slender  pedicels,  nearly  sepa- 
rate sepals,  and  small  white  to  rose  corollas. 
Standley  (1938)  included  this  species  under  S.  pe- 
duncularis,  but  that  species,  although  similar,  is 
larger  in  almost  all  its  dimensions.  In  the  Flora 
of  Panama,  specimens  of  this  species  were  placed 
under  5.  jorullensis  Kunth  in  H.B.K.,  a  species  of 
western  Mexico. 

Stemodia  durantifolia  (L.)  Swartz,  Obs.  Bot. 
240.  1791.  Capraria  durantifolia  L.,  Syst.  Nat. 
ed.  10:  1116.  1759.  Figures  4  and  6. 

Herbs  0.2-0.9(-1.5)  m  tall,  annual  or  becom- 
ing woody  at  the  base,  usually  with  many  branch- 
es, leafy  stems  0.5-4.5  mm  diam.,  pubescent  with 
crooked  translucent  multicellular  viscid  glandular 
hairs  0.1-0.5  mm  long,  stems  4- angled  with  4  lon- 
gitudinal ridges.  Leaves  2-3/node,  diminishing  in 
size  distally  and  intergrading  with  the  bracts,  ses- 
sile and  clasping  the  stem;  leaf  blades  1 .4-7  cm 
long,  4-18  mm  wide,  smaller  leaves  narrowly  el- 
liptic-oblong to  narrowly  ovate,  larger  leaves  ob- 
lanceolate,  apex  acute,  margin  with  3-12  teeth/ 
side,  base  auriculate  in  larger  leaves,  both  surfac- 
es with  minute  (0. 1  mm)  glandular  hairs  and  lon- 
ger thin  hairs,  2°  veins  2-5/side.  Inflorescences 
4-17  cm  long,  spicate  or  paniculate  to  50  cm  long 
with  opposite  spicate  branches  and  internodes  5- 
15  mm  long,  puberulent  like  the  stems,  bracts  (re- 
duced leaves)  5-16  mm  long,  2-5  mm  wide,  ped- 
icels 0-3  mm  long,  bracteoles  1  or  2,  subtending 
the  calyx,  linear,  2-5  mm  long.  Flowers  with  ca- 
lyx 4-6  mm  long,  ca.  1  mm  diam.,  lobes  2-4  mm 
long,  linear-lanceolate,  densely  glandular-puberu- 
lent;  corolla  5-7  mm  long,  blue  to  purple,  glan- 
dular-puberulent,  tube  ca.  1  mm  diam.,  yellow 
within  the  throat,  lower  lobes  ca.  1.5  mm  long; 
stamens  inserted  at  the  same  level  near  the  mouth 
of  the  tube,  connective  globose;  style  3-4  mm 
long.  Fruits  3-4  mm  long,  2  mm  diam.,  narrowly 
ovoid,  smooth,  yellowish  brown;  seeds  0.2-0.3 
mm  long,  0.15  mm  diam.,  narrowly  oblong  or  nar- 


rowed at  one  end,  dark  brown,  with  longitudinal 
rows  of  rounded  tubercles  (50X). 

Weedy  plants  of  open  sunny  moist  sites  or  part- 
ly shaded  woodland  in  seasonally  dry  vegetation 
of  the  Pacific  slope  (rarely  collected  on  the  Carib- 
bean slope),  0-900  m  elevation.  Flowering  and 
fruiting  throughout  the  year,  but  collected  most 
often  in  November-March.  The  species  ranges 
from  the  southwestern  United  States,  Mexico,  and 
the  West  Indies  to  Chile  and  Brazil. 

Stemodia  durantifolia  is  recognized  by  the 
erect,  much-branched,  weedy  habit,  viscid  pubes- 
cence, sessile  leaves  with  basal  blades  oblanceo- 
late,  long  distal  spikes  with  short  internodes,  small 
flowers,  and  blue  or  purple  corollas.  The  flowers 
are  usually  two  per  node  but  may  be  as  many  as 
six  per  node.  Distal  bracts  may  be  alternate  or 
tightly  congested  on  flowering  stems.  The  brac- 
teoles are  difficult  to  distinguish  from  the  sepals 
but  are  an  unusual  character.  These  plants  resem- 
ble species  of  Hyptis  (Lamiaceae). 

Stemodia  peduncularis  Benth.  in  DC.,  Prodr.  10: 
382.  1846.  Stemodiacra  peduncularis  (Benth.) 
Kuntze,  Rev.  Gen.  PI.  2:  466.  1891.  Figure  3. 

Erect  or  decumbent  herbs  10-80  cm  tall,  usu- 
ally with  few  or  no  lateral  branches  on  distal 
stems,  leafy  stems  0.5-2  mm  diam.,  pubescence 
of  thin  white  or  viscid  hairs  0.2-0.5  mm  long, 
glabrescent.  Leaves  opposite,  petioles  4-14  mm 
long.  0.3-0.9  mm  wide,  puberulent  with  thin 
whitish  hairs,  leaf  base  decurrent  on  the  petiole; 
leaf  blades  8-40(-50)  mm  long,  6-25(-35)  mm 
wide,  ovate  to  ovate-triangular  or  ovate-oblong, 
apex  acute,  margin  with  8-12  teeth/side,  base  ob- 
tuse to  truncate,  punctate  and  minutely  (0.1-0.2 
mm)  puberulent  on  both  sides,  2°  veins  3  or  4/ 
side,  strongly  ascending.  Inflorescences  of  1  or  2 
flowers  in  leaf  axils  (1-4/node),  pedicels  6-18 
mm  long  (-50  mm  in  fruit),  0.1-0.2  mm  diam., 
with  thin  whitish  hairs  0.1-0.2  mm  long.  Flowers 
with  sepals  4-7  mm  long.  0.7-1  mm  wide  at  the 
base,  linear  to  linear-triangular,  minutely  puberu- 
lent (sometimes  glandular  punctate);  corolla  1 1- 
15  mm  long,  white  or  red-veined  (yellowish),  tube 
8-10  mm  long,  2-3  mm  diam..  externally  gla- 
brous, lobes  2-3  mm  long;  stamens  borne  near  the 
base  of  the  tube;  style  4-7  m  long.  Fruits  5-7 
mm  long,  narrowly  ovoid;  seeds  0.5-1  mm  long, 
irregular  in  shape,  black. 

Infrequently  encountered  plants  of  shaded  sites 
in  moist  evergreen  montane  forest  formations 
along  the  Continental  Divide  and  Pacific  slope  in 


BURGER:  FLORA  COSTARICENSIS 


61 


the  central  volcanic  highlands  and  Cordillera  de 
Talamanca.  1300-2400  m  elevation.  Flowering 
and  fruiting  throughout  the  year,  but  collected 
mostly  in  December-March.  The  species  ranges 
from  western  Mexico  to  western  Panama. 

Stemodia  peduncularis  is  recognized  by  its 
short,  erect,  few-branched  habit,  opposite  serrate 
leaves,  mostly  solitary  flowers  on  long  slender 
pedicels,  separate  linear  sepals,  and  white  tubular 
corollas.  Standley  (1938)  incorrectly  included  5. 
angulata  under  this  species.  This  species  resem- 
bles our  species  of  Darcya  in  general  appearance. 

Stemodia  verticillata  (Miller)  Hassler,  Trab. 
Mus.  Farmacol.  21:  110.  1909.  Erinus  verticil- 
lotus  Miller,  Card.  Diet.  ed.  8.  1768.  S.  parvi- 
flora  W.  T.  Aiton,  Hortus  Kew.,  ed.  2,  4:  52. 
1812.  S.  arenaria  Kunth  in  H.B.K.,  Nov.  Gen. 
Sp.  2:  357.  1817.  Lindernia  verticillata  (Miller) 
Britton  in  Britton  &  Wilson,  Bot.  Porto  Rico  6: 
184.  1925.  Figure  2. 

Small  herbs  5-15(-25)  cm  tall,  prostrate  to  as- 
cending, rooting  only  near  the  base,  internodes 
mostly  4-14  mm  long,  leafy  stems  0.5-1  mm 
diam.,  densely  puberulent  with  straight  or  curved 
(often  glandular)  hairs  0.2-0.8  mm  long.  Leaves 
usually  2  or  3  (4)  per  node,  petioles  2-1 1  mm 
long,  0.2-0.5  mm  wide,  puberulent,  leaf  base  de- 
current  on  the  petiole;  leaf  blades  5-12(-18)  mm 
long,  3-10(-14)  mm  wide,  ovate  to  ovate-trian- 
gular or  ovate-oblong,  apex  obtuse,  margin  with 
5-9  teeth/side,  teeth  ca.  0.5  mm  long,  base  obtuse 
or  truncated,  both  surfaces  with  hairs  0.1-0.5  mm 
long,  2°  veins  3-5/side.  Inflorescences  of  usually 
2  subsessile  axillary  flowers  (2-6/node),  pedicels 
0.5-2  mm  long,  minutely  puberulent,  bracteoles 
absent.  Flowers  with  calyx  2-4  mm  long,  sepals 
equal  or  subequal,  0.5  mm  wide  near  the  base, 
with  thin  whitish  and  glandular  hairs;  corolla  3- 
5  mm  long,  blue-violet  or  purple  with  white 
throat,  tube  1  mm  diam.,  lobes  1-2  mm  long;  sta- 
mens borne  in  the  lower  half  of  the  tube,  anther 
thecae  0.3  mm  long;  style  1-1.5  mm  long  with 
recurved  stigma.  Fruits  1.8-2.5  mm  long,  1.4-2 
mm  wide,  ovoid-subglobose,  glabrous,  pale  yel- 
low-brown; seeds  ca.  0.5  mm  long,  0.2-0.3  mm 
diam.,  narrowly  ovoid,  yellowish,  with  dark  tip 
0.05  mm  long,  surface  with  longitudinal  ridges 
and  sulci. 

Uncommon  weeds  of  open  sunny  or  shaded 
sites  (often  on  sand  or  wet  gravel)  in  evergreen 
or  deciduous  vegetation,  0-1600  m  elevation. 
Probably  flowering  and  fruiting  throughout  the 


year,  but  collected  in  Costa  Rica  primarily  in 
July-August.  The  species  ranges  from  Mexico 
and  the  West  Indies  to  Argentina. 

Stemodia  verticillata  is  recognized  by  its  small, 
often  prostrate  habit,  mixture  of  both  glandular 
and  eglandular  hairs,  small  petiolate  leaves  sub- 
tending solitary  subsessile  flowers,  narrow  sepals, 
and  bluish  or  purple  corollas.  Plants  with  three  or 
four  leaves  per  node  are  distinctive.  The  glandular 
punctations  of  the  leaf  surfaces  are  often  obscure. 
These  plants  have  been  called  hierba  santa  (Costa 
Rica)  and  corrimiento  (El  Salvador). 


Tetranema  Bentham  ex  Lindley 

REFERENCES — M.  Grayum  &  B.  Hammel,  The 
genus  Tetranema  (Scrophulariaceae)  in  Costa 
Rica,  with  two  new  species.  Phytologia  79:  269- 
280.  1995  (1996).  T.  Mendez-Larios  &  J.  Villa- 
senor.  Revision  taxonomica  del  genero  Tetranema 
(Scrophulariaceae).  Acta  Bot.  Mexico  32:  53-68. 
1995. 

Herbs,  erect  or  decumbent,  perennial,  some- 
times woody  at  the  base  but  the  stems  usually 
short,  glabrous  or  pubescent.  Leaves  opposite  and 
often  crowded  near  the  base,  equal  or  unequal  at 
a  node,  subsessile  and  clasping  the  stem  or  peti- 
olate, blades  dentate  to  subentire,  venation  pin- 
nate. Inflorescences  axillary,  cymose  to  subum- 
bellate  or  congested,  long-pedunculate,  bracteate, 
pedicels  short  but  often  elongating  in  fruit.  Flow- 
ers with  the  calyx  deeply  5-parted,  campanulate, 
lobes  narrow  and  attenuate-acuminate,  with  prom- 
inent longitudinal  ribs;  corolla  tubular  to  campan- 
ulate, 2-lipped,  glabrous,  white  to  rose,  purple  or 
less  often  red,  tube  longer  than  the  lobes,  upper 
lip  emarginate  ( 1  -lobed),  lower  lip  with  3  spread- 
ing lobes,  imbricate  in  bud;  stamens  4  (rarely  3), 
filaments  of  2  longer  and  2  shorter  pairs,  anthers 
2-thecous,  a  staminode  very  small  (in  ours)  or  ab- 
sent; ovary  2-locular,  style  slender  or  thick,  stig- 
ma capitate  or  bilobed.  Fruits  capsules,  ovoid  to 
globose,  glabrous,  dehiscence  loculicidal;  seeds 
many,  angled  or  tetrahedral,  surface  reticulate  or 
foveolate. 

Tetranema  is  a  genus  of  six  species  from  Mex- 
ico and  Central  America.  It  is  distinguished  by  its 
opposite  leaves,  axillary  and  cymose  inflorescenc- 
es, separate  sepals,  four-lobed  and  two-lipped  co- 
rolla, and  loculicidal  capsules.  Our  recently  dis- 
covered species  are  unusual  in  having  red  (prob- 
ably hummingbird-pollinated)  flowers.  These  dis- 
tinctive plants  (Fig.  27)  may  resemble  Razisea 


62 


FffiLDIANA:  BOTANY 


spicata  Oerst.  and  Odontonema  tubaeforme  (Ber- 
tol.)  Kuntze  of  the  Acanthaceae.  The  Costa  Rican 
species  are  similar  to  T.  megaphyllum  (Brande- 
gee)  L.  O.  Williams,  but  that  species  has  smaller 
(2-4  mm)  corolla  lobes  and  larger  (6-10  mm)  flo- 
ral bracts  and  is  endemic  to  Mexico.  Species  of 
Tetranema  are  sometimes  cultivated  as  ornamen- 


tals, especially  T.  roseum  (M.  Martens  &  Galeoti) 
Standl.  &  Steyerm.  The  new  species  come  from 
the  Pacific  slope  of  central  Costa  Rica  and  both 
slopes  of  southern  Costa  Rica.  The  dichotomy  in 
Flora  of  Guatemala  separating  Tetranema  from 
Penstemon  and  Uroskinnera  is  inverted  (Standley 
&  Williams,  1973,  p.  321). 


Key  to  the  Species  of  Tetranema 

la.  Inflorescences  with  14-30  flowers,  peduncle  purple;  corollas  26-35  mm  long,  pubescent  internally 
with  a  band  of  flat  yellow  hairs  along  the  entire  ventral  (abaxial)  surface  and  onto  the  lower  lobe; 
leaf  apex  rounded  to  short-acuminate;  1200-1600  m  on  Cerro  Turrubares T.  floribundum 

Ib.  Inflorescences  with  2-12  flowers,  peduncle  green;  corollas  44-55  mm  long,  glabrous  throughout  or 
rarely  pubescent  on  the  lower  lobe  and  mouth;  leaf  apex  long-acuminate;  500-1200  m  elevation  on 
both  Caribbean  and  Pacific  slopes  T.  gamboanum 


Tetranema  floribundum  B.  Hammel  &  M.  Gra- 
yum,  Phytologia  79:  269-280.  1995  (1996). 
Figure  27. 

Erect  herbs  (0.3-)0.8-2  m  tall,  decumbent  near 
the  base  and  often  rooting  at  the  lower  nodes,  in- 
ternodes  to  5  cm  long,  arachnoid  or  wooly-pu- 
bescent  when  young,  leafy  stems  1 .5-5  mm  thick, 
glabrescent.  Leaves  opposite,  subsessile  with  pet- 
ioles 0-6  mm  long,  ca.  2  mm  thick;  leaf  blades 
14-24  cm  long,  7-13  cm  wide,  broadly  elliptic  to 
oblanceolate  or  spatulate,  apex  abruptly  acute  or 
short-acuminate,  margins  coarsely  serrate  or  un- 
dulate-toothed with  teeth  ca.  1  mm  high,  base 
gradually  cuneate,  mostly  glabrous  above,  mi- 
nutely puberulent  on  the  major  veins  beneath,  lus- 
trous beneath  when  dry,  2°  veins  8-1  I/side,  as- 
cending. Inflorescences  15-30  cm  long,  14-30- 
flowered,  peduncles  13-23  cm  long,  purple,  with 
4  longitudinal  ridges,  bracts  1-5  mm  long,  nar- 
rowly triangular,  margins  ciliate  (often  only  at  the 
base),  pedicels  10-20  mm  long,  glabrous.  Flow- 
ers with  calyx  ca.  3  mm  long  (-5  mm  in  fruit), 
divided  nearly  to  base,  striate  and  glabrous  (ex- 
cept along  margins),  with  a  short  apical  projec- 
tion; corolla  26-35  mm  long,  5-6  mm  diam.,  tu- 
bular and  slightly  expanded  distally  and  curved 
upward,  red-orange,  lobes  4,  ca.  1 3  mm  long,  2.5- 
5.5  mm  wide,  the  3  lower  rounded  apically  and 
reflexed-spreading,  puberulent  internally  abaxial- 
ly;  stamens  slightly  exserted,  anthers  0.8-0.9  mm 
long,  staminode  ca.  0.5  mm  long;  ovary  ca.  3.5 
mm  long.  Fruits  not  seen  at  maturity,  ca.  8  mm 
long,  ovoid,  borne  on  pedicels  ca.  20  mm  long. 

Plants  of  evergreen  lower  montane  forest  for- 


mations at  1200-1600  m  elevation  on  Cerro  Tur- 
rubares. Flowering  in  December-January  and 
March.  This  species  is  known  only  from  an  iso- 
lated peak  on  the  Pacific  slope  of  the  southeastern 
part  of  the  central  plateau  in  the  province  of  San 
Jose". 

Tetranema  floribundum  is  recognized  by  its 
larger  subsessile  elliptic  to  obovate  leaves  with 
many  ascending  secondary  veins,  inflorescences 
with  long  purple  peduncles,  five  separate  acute 
sepals,  curved-tubular  red  corollas,  and  isolated 
habitat.  Cerro  Turrubares  is  also  the  location  of  a 
recently  discovered  and  unusual  species  of  Psy- 
chotria  (Burger  &  Jime'nez,  1994). 

Tetranema  gamboanum  M.  Gray  urn  &  B.  Ham- 
mel, Phytologia  79:  269-280.  1995  (1996).  Fig- 
ure 27. 

Erect  herbs  1-2  m  tall,  internodes  to  1 1  cm  (or 
more)  long,  at  first  minutely  strigulose,  1 .7-5  mm 
thick,  glabrous  but  with  a  puberulent  interpetiolar 
line.  Leaves  opposite,  sessile  to  subsessile  or  with 
petioles  to  1  cm  long,  canaliculate  above,  margins 
ciliate  at  the  base;  leaf  blades  14-31  cm  long,  5- 
1 1  cm  wide,  elliptic  to  elliptic-obovate,  apex  long- 
acuminate,  margin  coarsely  serrate  with  teeth  1- 
2  mm  high,  base  cuneate  and  abruptly  narrowed 
at  the  base,  glabrous  above  or  with  few  hairs 
along  midvein  and  major  veins,  minutely  puber- 
ulent along  the  veins  beneath,  2°  veins  9-13/side. 
Inflorescences  axillary  cymes  with  2-13  flowers, 
peduncles  9-24  cm  long,  green,  quadrangular 
with  4-winged  ribs,  bracts  0.5-2  mm  long,  subu- 
late to  narrowly  triangular,  ciliate  along  the  mar- 
gins, pedicels  9-12  mm  long  (-20  mm  in  fruit), 


BURGER:  FLORA  COSTARICENSIS 


63 


glabrous.  Flowers  with  5-parted  calyx,  sepals  3- 
5  mm  long  (-6  mm  in  fruit),  inner  2  sepals  slight- 
ly smaller  than  the  outer,  narrowly  to  broadly 
ovate,  imbricate  in  bud,  minutely  ciliolate  along 
the  margin,  apex  acute,  venation  parallel;  corolla 
44-55  mm  long,  ca.  5  mm  diam.,  narrowly  tu- 
bular, gradually  expanded  distally  and  slightly 
curved,  bright  red,  glabrous  throughout  or  with 
hairs  at  throat  and  lower  lobes,  lobes  8-14  mm 
long,  3-4  mm  wide,  lanceolate  or  oblong,  the  3 
lower  lobes  obtuse  to  rounded  at  the  apex  and 
spreading-reflexed;  stamens  slightly  exserted,  fil- 
aments dilated  near  base,  glabrous,  anthers  0.8-1 
mm  long,  with  divergent  thecae  confluent  at  the 
base,  staminode  1.5-2  mm  long;  ovary  3-4  mm 
long,  narrowly  ovoid,  style  exserted,  stigma  cap- 
itate. Fruits  capsules,  6-9  mm  long  (not  including 
the  1-2  mm  style  base),  subglobose-apiculate, 
glabrous;  seeds  0.5-0.7  mm  long,  0.4-0.5  mm 
diam.,  rectangular  or  oblong-rounded,  surface 
with  prominent  pits  (foveolate),  dark  yellow  to 
brown  or  blackish. 

Newly  discovered  plants  of  evergreen  rain  for- 
est formations  on  the  Caribbean  slope  of  the  Cor- 
dillera de  Talamanca  and  on  the  southern  Pacific 
slope,  500-1200  m  elevation.  Flowering  and  fruit- 
ing in  December-May.  This  species  has  been  col- 
lected only  from  between  83°W  and  84°W  in  the 
southern  half  of  Costa  Rica. 

Tetranema  gamboanum  is  distinguished  by  its 
erect  stems  with  larger  ellipsoid  leaves  with  many 
lateral  veins,  axillary  cymes  on  long  peduncles, 
five-parted  calyx,  and  bright  red,  slightly  curved 
corolla  tubes. 


Torenia  Linnaeus 

Annual  or  perennial  herbs,  erect  or  procum- 
bent, branching  from  both  basal  and  distal  nodes, 


stems  4-angled  with  longitudinal  ridges,  puberu- 
lent  to  hirsute  or  glabrous.  Leaves  opposite,  pet- 
iolate,  blade  margins  entire  to  serrate  or  crenulate, 
venation  pinnate.  Inflorescences  of  1-3  flowers 
in  axils  of  distal  leaves  or  in  short  terminal  or 
axillary  few-flowered  racemes,  linear  bracts 
sometimes  present,  pedicels  usually  held  at  a  45° 
angle  to  the  stem.  Flowers  showy  or  small,  calyx 
usually  somewhat  shorter  than  the  corolla  tube, 
united  to  form  a  long  tube  with  3-5  short  lobes 
(or  2-lipped),  with  3-5  longitudinal  ribs  or  broad 
wings,  persisting  and  enlarging  in  fruit;  corolla 
campanulate  to  tubular  or  salverform,  bilabiate 
with  the  upper  lip  erect  and  2-lobed,  lower  lip 
with  3  lobes,  tube  usually  widened  distally,  beard- 
ed in  the  throat;  stamens  4,  in  2  unequal  pairs, 
filaments  inserted  near  the  top  of  the  tube  and 
usually  with  a  tooth-like  appendage  at  the  base, 
anterior  2  arching  upward  over  the  stigma  (with 
anthers  connivent  or  reduced),  thecae  oblong  to 
linear,  divaricate  by  the  enlarged  connective;  disc 
prominent  and  saucer-shaped  or  cupulate;  ovary 
oblong,  style  straight,  stigma  slightly  2-lobed. 
Fruits  capsules,  usually  enclosed  within  the  per- 
sisting perianth  tube,  oblong-ellipsoid,  dehiscing 
septicidally  to  the  base,  placenta  linear  with  sep- 
tum forming  wings;  seeds  globose  to  variously 
angled,  tuberculate  or  reticulate. 

Torenia  is  a  genus  of  40  to  80  species  of  trop- 
ical Africa  and  Asia.  A  few  species  are  cultivated 
ornamentals  that  have  become  naturalized  in  the 
Americas.  Torenia  is  closely  related  to  Lindernia, 
sharing  the  curved  appendaged  filaments  often 
found  in  that  genus.  Species  of  Torenia  in  the 
New  World  can  be  distinguished  from  Lindernia 
by  their  larger  flowers,  tubular  calyx,  elongate 
fruits,  and  globose  pitted  seeds. 


Key  to  the  Species  of  Torenia 

la.  Corolla  30-40  mm  long,  lower  lobes  dark  purple  or  blue-violet;  calyx  broadly  winged  and  5-10 
mm  wide;  leaves  1 6-50  mm  long,  often  truncate  at  the  base  T.  fournieri 

Ib.  Corolla  7-12  mm  long,  lower  lobes  pale  purple  or  pale  blue- violet;  calyx  not  winged,  2-3  mm 
diam.;  leaf  blades  8-22  mm  long,  obtuse  to  rounded  at  the  base  T.  thouarsii 


Torenia  fournieri  Linden  in  Fourn.,  Illustr.  Hor- 
tic.  23:  129,  tab.  249.  1876.  Figure  8. 

Weak-stemmed  erect  herbs  15-50  cm  tall,  of- 
ten reddish  near  the  base,  main  stems  with  2-5 
branching  nodes,  leafy  stems  1-3  mm  diam.,  4- 


angled  with  prominent  longitudinal  ridges,  nodes 
with  thin  whitish  hairs  0.2-0.5  mm  long  (inter- 
nodes  often  glabrous).  Leaves  opposite  through- 
out, petioles  3-17  mm  long,  0.7-1.4  mm  wide, 
with  thin  whitish  hairs  along  the  adaxial  margins; 
leaf  blades  16-50  mm  long,  6-25  mm  wide,  tri- 


64 


FIELDIANA:  BOTANY 


angular  to  ovate-triangular  or  ovate-elliptic,  apex 
acute,  margin  with  prominent  teeth,  base  truncate 
to  obtuse,  upper  surface  with  few  thin  straight 
hairs  0.2-0.5  mm  long,  2°  veins  3-6/side,  strongly 
ascending.  Inflorescences  of  solitary  flowers  in 
distal  leaf  axils  (2/node)  or  more  often  in  short 
terminal  racemes  with  4-8  flowers,  bracts  2-5(-12) 
mm  long,  linear,  pedicels  4-18  mm  long,  0.6-1 
mm  wide,  glabrous  or  very  sparsely  puberulent. 
Flowers  with  calyx  13-22  mm  long,  5-10  mm 
wide,  tube  more  than  half  the  calyx  length,  broad- 
ly winged  to  produce  an  ovate  form  with  acute  or 
acuminate  apices,  ciliolate;  corolla  3-4  cm  long, 
2-3  cm  wide  distally,  tube  7-9  mm  diam.  in  the 
center,  whitish  near  the  base,  upper  lip  often  pale 
blue  with  the  lower  lobes  very  dark  blue-violet,  a 
yellow  patch  present  within  at  the  base  of  the  cen- 
tral lobe.  Fruits  1-2  cm  long,  included  within  the 
slightly  enlarged  calyx. 

Torenia  fournieri,  a  native  of  southeastern 
Asia,  is  widely  planted  as  an  ornamental.  This 
species  may  become  naturalized  in  lowland  and 
mid-elevation  areas  of  Central  America,  but  it 
seems  doubtful  that  the  populations  persist  for 
long.  These  plants  are  easily  distinguished  by 
their  small  size,  opposite  (often  triangular)  leaves, 
few-flowered  racemes,  broadly  winged  calyx,  and 
relatively  large  corollas  with  dramatic  dark  pur- 
ple-violet coloring.  This  species  is  unusual  within 
the  genus  in  having  broadly  winged  sepals  and  in 
lacking  basal  appendages  on  the  lower  filaments. 
Torenia  asiatica  L.  is  similar  but  has  prostrate 
stems.  The  large  colorful  corollas  make  these 
plants  especially  attractive.  They  can  be  mistaken 
for  members  of  the  Gesneriaceae. 

Torenia  thouarsii  (Cham.  &  Schldl.)  Kuntze, 
Rev.  Gen.  PI.  2:  468.  1891.  Nonenia  thouarsii 
Cham.  &  Schldl.,  Linnaea  3:  18.  1828.  Linder- 
nia  thouarsii  (Cham.  &  Schldl.)  Edwin,  Phy- 
tologia  19:  361.  1970.  Figure  3. 

Procumbent  or  erect  herbs  10-30  cm  tall,  leafy 
stems  0.6-1.3  mm  diam.,  quadrangular,  internodes 
glabrous  or  with  thin  whitish  hairs  0.2-0.5  mm 
long,  nodes  usually  with  thin  hairs.  Leaves  op- 
posite throughout,  petioles  1-7  mm  long,  ca.  1 
mm  wide  with  thin  lateral  margins,  sparsely  pu- 
berulent; leaf  blades  8-22  mm  long,  5-12  mm 
wide,  ovate-elliptic  to  ovate-triangular,  apex 
acute,  margins  with  4  or  5  prominent  teeth/cm, 
base  obtuse  to  rounded,  surfaces  glabrous  or 
sparsely  puberulent,  2°  veins  3-5/side,  arcuate  as- 
cending. Inflorescences  of  1  or  2  flowers  axillary 


to  foliage  leaves  (l-r4/node),  bracts  ca.  2  mm 
long,  linear,  pedicels  2-15  mm  long,  ca.  0.5  mm 
diam.,  glabrous  or  sparsely  puberulent.  Flowers 
with  calyx  4-12  mm  long,  1.3-2  mm  diam.  (to 
18X3  mm  in  fruit),  narrowly  tubular  to  tubular- 
ellipsoid,  glabrous,  lobes  2-3  mm  long,  acute;  co- 
rolla 7-12  mm  long,  bluish,  purple,  or  white, 
lobes  crenate;  filaments  with  a  linear  appendage 
at  base,  staminode  with  a  rounded  tip.  Fruits  8- 
10  mm  long.  2-3  mm  wide,  narrowly  oblong-el- 
lipsoid, included  within  the  enlarged  calyx  tube 
or  the  tube  splitting. 

Torenia  thouarsii,  a  native  of  India,  is  natural- 
ized in  parts  of  Central  and  South  America.  In 
Costa  Rica  it  is  occasionally  found  in  wet  sites  at 
10-2000  m  elevation,  from  the  Caribbean  low- 
lands to  the  central  highlands.  These  plants  resem- 
ble species  of  Lindernia  and  Stemodia  but  differ 
in  the  tubular  calyx  and  narrowly  oblong  fruits. 


Veronica  Linnaeus 

Annual  or  perennial  herbs  (rarely  shrubs), 
prostrate  or  ascending  to  erect,  often  branching 
from  the  base,  stems  terete  or  with  longitudinal 
ridges,  glabrous  or  puberulent.  Leaves  opposite 
near  the  base  and  usually  alternate  distally,  some- 
times intergrading  with  smaller  floral  bracts,  ses- 
sile or  petiolate.  blades  subentire  to  dentate,  cre- 
nate or  divided,  pubescence  of  simple  multicel- 
lular  or  glandular  hairs,  venation  usually  palmate 
or  subpalmate.  Inflorescences  of  solitary  flowers 
in  leaf  axils  or  of  elongate  racemes  (or  spike-like) 
with  flowers  solitary  in  the  axils  of  bracts,  usually 
terminal,  pedicels  very  short  to  long  and  slender. 
Flowers  mostly  small,  calyx  deeply  4-lobed  (rare- 
ly 5-lobed),  the  lobes  (sepals)  subcquai  to  un- 
equal, slightly  overlapping  in  bud,  usually  per- 
sisting and  slightly  enlarged  in  fruit,  glabrous  or 
puberulent;  corolla  rotate  (rarely  campanulate), 
tube  very  short,  corolla  lobes  4  (5).  unequal,  with 
the  lower  lobe  the  smallest,  lateral  lobes  exterior 
in  bud,  blue  to  purple  or  white  (rarely  reddish); 
stamens  2,  borne  at  either  side  of  the  upper  lobe, 
exsertcd,  filaments  attached  at  the  base  of  the  tube 
and  free,  anthers  2-thecous.  thecae  confluent  at 
the  apex;  ovary  rounded,  2-locular,  ovules  few  to 
many,  style  simple,  persisting  in  fruit  but  not  en- 
larging, stigma  capitate.  Fruits  capsules,  flattened 
at  right  angles  to  the  plane  of  the  septum,  obovoid 
or  2-lobed  (obcordate)  with  a  depressed  or  emar- 
ginate  apex,  dehiscence  loculicidal  (also  some- 
times septicidal);  seeds  few  to  many,  ovate  to  or- 


BURGER:  FLORA  COSTARICENSIS 


65 


bicular,  surfaces  smooth  to  rugulose,  often  with 
the  inner  face  concave,  embryo  often  U-shaped. 
Veronica  is  a  genus  of  150  to  250  species  con- 
fined to  cool  climates.  The  great  majority  of  spe- 
cies are  found  in  Europe  and  Asia,  but  a  number 
have  become  widespread  weeds;  it  seems  likely 
that  none  are  native  to  Central  America.  The  ge- 
nus is  distinguished  (in  our  area)  by  its  small  her- 
baceous growth  form,  small  leaves  becoming  al- 
ternate distally,  small  flowers  with  prominent  ca- 
lyx lobes  (sepals),  bluish  to  lilac  or  white  corollas 
with  the  four  lobes  usually  held  in  a  single  plane 
(rotate),  two  stamens,  and  unusual  fruits.  The  cap- 


sules are  flattened  perpendicular  to  the  plane  of 
the  septum  and  often  have  an  emarginate  or  deep- 
ly notched  apex.  The  rounded  lateral  lobes  of  the 
fruits  (carpels)  can  be  lenticular  or  rounded.  The 
genus  is  placed  in  the  tribe  Veroniceae  and  is 
closely  related  to  Sibthorpia  and  the  Australasian 
Hebe.  Unfortunately,  some  species  of  Veronica 
are  very  similar,  and  a  number  of  Central  Amer- 
ican collections  have  been  misidentified  in  the 
past.  In  Costa  Rica  the  genus  has  not  been  found 
below  1200  m  elevation,  and  only  two  species  (V. 
polita  and  V.  serpyllifolia)  are  commonly  collect- 
ed. 


Key  to  the  Species  of  Veronica 

la.  Rowers  borne  on  conspicuous  pedicels,  the  pedicels  much  longer  than  the  calyx  lobes,  flowers 
arising  from  the  axils  of  leaves  or  from  short  axillary  racemes;  leaves  with  prominent  teeth  along 

the  margins;  styles  1.3-3  mm  long 2 

Ib.  Flowers  borne  on  short  pedicels,  the  pedicels  much  shorter  to  slightly  longer  than  the  calyx  lobes, 
flowers  arising  from  the  axils  of  leaves  or  from  the  axils  of  floral  bracts  differing  greatly  in  size 
and  form  from  the  basal  leaves;  leaves  with  small  or  inconspicuous  teeth  along  the  margin  (except 

in  V.  arvensis);  styles  0.5-2.5  mm  long 3 

2a.  Corolla  <  8  mm  wide;  fruits  about  as  broad  as  long,  ca.  4  mm  wide,  venation  obscure;  fruiting 
sepals  ca.  5  mm  long;  leaf  blades  often  triangular-ovate;  styles  1.3-1.6  mm  long;  occasional  at 

1400-3300  m  elevation    V.  polita 

2b.  Corolla  8-13  mm  wide;  fruits  distinctly  broader  than  long,  5-9  mm  wide,  venation  prominent; 
fruiting  sepals  ca.  6  mm  long;  leaf  blades  usually  ovate-oblong;  styles  1.8-3  mm  long;  rarely 

collected  in  Costa  Rica V.  persica 

3a.  Lower  leaf  blades  8-28  mm  long,  narrowly  oblanceolate,  leaves  gradually  intergrading  into  the 

smaller  distal  floral  bracts;  erect  stems  to  45  cm  tall;  corolla  white  with  purple  lines  [pedicels  ca. 

1  mm  long;  styles  ca.  0.5  mm  long;  rarely  collected  in  southern  Central  America]  .  .  V.  peregrina 

3b.  Lower  leaf  blades  up  to  12  mm  long,  usually  broadly  ovate;  leaves  usually  clearly  differentiated 

from  the  smaller  floral  bracts;  erect  stems  rarely  exceeding  20  cm  in  height;  corolla  usually  bluish 

4 

4a.  Pedicels  becoming  6  mm  long;  styles  1.8-2.5  mm  long;  margins  of  leaf  blades  subentire  to  slightly 

crenate;  frequent  in  Costa  Rica V.  serpyllifolia 

4b.  Pedicels  0.5-1 .2  mm  long;  styles  ca.  0.6  mm  long;  margins  of  leaf  blades  minutely  to  conspicuously 
dentate-crenate;  rarely  collected  in  Costa  Rica V.  arvensis 


Veronica  arvensis  L.,  Sp.  PI.  13.  1753. 

Annual  herbs,  5-20(-40)  cm  tall,  stems  usually 
branching  only  near  the  base,  leafy  stems  0.4-1.3 
mm  diam.,  puberulent  with  thin  curved  whitish 
hairs  0.2-0.7  mm  long.  Leaves  opposite  (but 
bracts  of  the  inflorescences  alternate),  sessile  or 
the  basal  leaves  with  petioles  1-4  mm  long;  leaf 
blades  4-12  mm  long,  3-10  mm  wide,  broadly 
ovate  to  ovate-oblong,  apex  obtuse  or  rounded, 
margin  crenate-dentate  with  2-6  teeth  0.3-0.8 
mm  high,  base  obtuse  or  truncate,  with  thin  hairs 


ca.  0.5  mm  long  on  both  surfaces,  venation  pal- 
mate with  3  or  5  major  veins.  Inflorescences  erect 
spike-like  racemes,  2-15  cm  long,  bracts  4-7  mm 
long,  1-2  mm  wide,  narrowly  oblong  to  narrowly 
ovate,  narrowed  at  the  base,  rachis  0.5-1.2  mm 
diam.,  puberulent,  pedicels  0.5-1.2  mm  long. 
Flowers  with  unequal  calyx  lobes  2.5-4  mm  long 
(becoming  5  mm  long  in  fruit),  ca.  1  mm  wide, 
narrowly  oblong  to  narrowly  elliptic-oblong,  pu- 
berulent with  short  thin  hairs;  corolla  blue,  upper 
lobe  ca.  2  mm  long;  anthers  ca.  0.4  mm  long; 
styles  ca.  0.6  mm  long.  Fruits  3.5-4  mm  long, 


66 


FffiLDIANA:  BOTANY 


ca.  3.5  mm  wide,  lenticular-obovoid,  distally  bi- 
lobed  with  an  apical  notch  0.5-1  mm  deep,  pu- 
berulent  along  the  margin. 

Veronica  arvensis  is  found  on  the  slopes  of 
Volcan  Turrialba  (Khan  et  al.  977),  but  it  is  rarely 
collected  in  Central  America.  This  species  is  rec- 
ognized by  its  short  stature,  leaves  with  subentire 
margins,  very  short  pedicels,  and  blue  corollas. 
The  closely  clustered  bracts  and  calyx  lobes  give 
the  inflorescences  of  this  species  a  spike-like  ap- 
pearance. Also,  there  are  few  intermediate  leaves 
along  the  stem  between  the  lower  foliage  leaves 
and  the  much  narrower  floral  bracts  in  this  spe- 
cies. 

Veronica  peregrina  L.,  Sp.  PI.  14.  1753.  V.  xal- 
apensis  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  2:  389. 
1817.  V.  peregrina  var.  xalapensis  (Kunth  in 
H.B.K.)  Pennell,  Torreya  19:  167.  1919. 

Erect  or  spreading  herbs  5-45  cm  tall,  branch- 
ing mostly  near  the  base,  leafy  stems  0.3-3  mm 
diam.,  glabrous  (in  var.  peregrina)  or  with  minute 
(0.2  mm)  gland-tipped  hairs  (in  var.  xalapensis). 
Leaves  opposite  near  the  base  and  alternate  dis- 
tally, gradually  becoming  smaller  and  bract-like 
distally,  narrowed  at  the  base  but  a  petiole  not 
clearly  differentiated;  leaf  blades  8-28  mm  long, 
1-8  mm  wide,  narrowly  oblong  to  narrowly  ellip- 
tic-oblong or  oblanceolate  (spatulate),  apex  blunt- 
ly acute  or  rounded,  margin  entire  or  with  few 
small  teeth,  base  gradually  narrowed  and  cuneate, 
surfaces  glabrous,  venation  subpalmate  with  a 
prominent  midvein.  Inflorescences  of  solitary 
flowers  in  leaf  axils  or  flowers  in  the  axils  of  re- 
duced bract-like  distal  leaves,  spike-like  with  ped- 
icels ca.  1  mm  long,  rachis  sparsely  puberulent. 
Flowers  with  subequal  or  strongly  unequal  calyx 
lobes  ca.  4  X  1  mm  (-6  mm  long  in  fruit),  nar- 
rowly oblong,  sparsely  puberulent;  corolla  2-3 
mm  wide,  white  marked  with  purple  lines;  sta- 
mens 2-3  mm  long;  style  ca.  0.5  mm  long.  Fruits 
2.8-4  mm  long,  4-5  mm  wide,  obovoid-triangular 
with  a  slight  indentation  at  the  apex,  compressed- 
lenticular,  glabrous  or  puberulent. 

Plants  of  open  moist  sites,  often  along  river- 
beds, 1000-3000  m  elevation.  Probably  flowering 
throughout  the  year  in  northern  Central  America, 
but  rarely  collected  in  southern  Central  America. 
This  species  is  native  to  southern  Europe  and  is 
now  widely  naturalized  in  temperate  climates. 

Veronica  peregrina  is  recognized  by  its  white 
flowers,  longer  erect  spicate  stems,  and  narrowly 
oblong  subentire  leaves.  Although  glabrous  in  Eu- 


rope, plants  in  the  Americas  range  from  glabrous 
to  glandular  puberulent  (var.  xalapensis).  The  typ- 
ical glabrous  variety  is  more  common  in  the  east- 
ern United  States,  whereas  var.  xalapensis  is  most 
common  in  the  western  United  States,  Mexico, 
and  Central  America,  but  they  intergrade,  and 
both  varieties  can  be  found  in  a  broad  band  across 
the  central  United  States.  The  longer  distal  stems 
arc  unusual  in  that  there  is  a  gradual  change  from 
flowers  subtended  by  narrow  leaves  (to  3  cm 
long)  to  flowers  subtended  by  linear  bracts  only  6 
mm  long. 

Veronica  persica  Poiret  in  Lam.,  Encycl.  8:  542. 
1808.  V.  rotundifolia  Sesse  &  Mocifto,  Fl.  Mex. 
5.  1892. 

Creeping  herbs,  stems  5-30(-50)  cm  long 
(-15  cm  tall),  leafy  stems  0.5-2  mm  diam.,  pu- 
berulent with  thin  whitish  hairs  0.1-0.5  mm  long, 
often  in  longitudinal  rows.  Leaves  opposite  near 
the  base  and  alternate  throughout  the  distal  stems, 
petioles  1-7  mm  long,  puberulent;  leaf  blades 
ovate  to  ovate-orbicular,  6-24  mm  long,  4-16  mm 
wide,  apex  obtuse  to  rounded,  margin  with  3-6 
prominent  (0.5-2  mm)  serrate  lobes/side,  base  ob- 
tuse or  truncate,  surfaces  with  whitish  hairs  to  0.7 
mm  long,  venation  palmate  with  3  or  5  major 
veins.  Inflorescences  of  solitary  flowers  in  leaf 
axils  (bracts  absent),  pedicels  10-35  mm  long, 
0.2-0.3  mm  diam.,  puberulent.  Flowers  with  4- 
parted  calyx,  sepals  4-5  mm  long,  subequal,  to  6 
mm  long  in  fruit,  1-1.5  mm  wide  at  the  base, 
lanceolate,  surfaces  glabrous  but  the  margin  cili- 
olate;  corolla  8-13  mm  wide,  upper  lobe  ca.  4 
mm  wide,  pale  blue  or  lilac  with  darker  blue  lines; 
filaments  2-3  mm  long,  anthers  0.9  mm  long; 
style  1.8-3  mm  long.  Fruits  3.5-5  mm  long,  5- 
9  mm  wide,  broadly  obcordate  with  apical  notch 
and  divergent  lateral  sides,  surface  glabrous  or 
with  thin  hairs,  the  veins  becoming  prominent; 
seeds  ovoid,  flattened,  rugulose. 

Veronica  persica,  probably  a  native  of  the  Cau- 
casus and  southwestern  Asia,  is  now  naturalized 
over  much  of  northern  North  America  and  is  rare- 
ly found  at  higher  elevations  in  Mexico  and  Cen- 
tral America.  Standley  reported  this  species  as 
growing  on  the  slopes  of  Volcdn  Irazii  in  Costa 
Rica.  This  species  is  recognized  by  its  creeping 
habit,  solitary  flowers  in  leaf  axils,  and  larger  ob- 
cordate capsules  that  are  distinctly  broader  than 
long.  The  broad  fruit  causes  the  persisting  sepals 
to  spread  wide  as  the  fruit  matures.  This  species 
has  often  been  confused  with  V.  polita,  but  that 


BURGER:  FLORA  COSTARICENSIS 


67 


species  has  smaller  flowers  and  different  fruits. 
Material  cited  as  V.  polita  in  Flora  of  Guatemala 
and  Flora  of  Panama  is  actually  V.  persica. 

Veronica  polita  Fries,  Novit.  Fl.  Suec.  63.  1819. 
Figure  2. 

Creeping  or  vining  herbs,  stems  5-70(-120) 
cm  long,  branching  at  both  distal  and  basal  nodes, 
leafy  stems  0.4-1.5  mm  diam.,  appressed  puber- 
ulent  with  thin  whitish  hairs  0.1-0.2  mm  long. 
Leaves  opposite,  petioles  4-18  mm  long,  0.3-0.6 
mm  wide  and  expanding  at  the  blade,  appressed 
puberulent;  leaf  blades  6-22  mm  long,  6-22  m 
wide,  triangular  to  ovate-triangular,  apex  acute  to 
obtuse,  margins  with  4-8  prominent  teeth  0.3-3 
mm  high,  base  truncate  to  subcordate,  surfaces 
with  scattered  stiff  sharp  hairs  0.1-0.7  mm  long, 
venation  palmate  with  3  major  veins.  Inflores- 
cences of  solitary  flowers  in  axils  of  leaves  or  of 
short  (2-4  cm)  axillary  racemes  with  narrowly  el- 
liptic bracts  2-4  mm  long,  pedicels  3-15  mm 
long,  0.2-0.3  mm  diam.,  puberulent.  Flowers 
with  calyx  lobes  (sepals)  4,  subequal  or  with  1 
distinctly  smaller,  lobes  3-4  mm  long  (-5  mm  in 
fruit),  1.5-2  mm  wide,  narrowly  obovate  or  ob- 
ovate,  surfaces  subglabrous  but  margins  with  stiff 
curved  hairs;  corolla  4-5  mm  long,  less  than  8 
mm  wide  at  anthesis,  blue  with  linear  darker 
markings,  filaments  ca.  1.3  mm  long,  anthers  0.4 
mm  long;  styles  1.3-1.6  mm  long.  Fruits  ca.  3 
mm  long,  3-4  mm  wide,  obovate-triangular  or 
slightly  obcordate,  veins  usually  indistinct;  seeds 
obvoid,  with  one  flattened  face,  rugulose  to 
ridged. 

Plants  of  open  moist  slopes,  damp  disturbed 
sites,  and  partly  shaded  forest  floor,  1400-3000  m 
elevation.  Flowering  and  fruiting  throughout  the 
year.  This  species  is  a  native  of  Europe  that  is  now 
widely  naturalized  in  North  America  and  montane 
areas  in  Central  America. 

Veronica  polita  is  distinguished  by  its  often  tri- 
angular leaves  with  prominent  teeth,  flowers  in 
axils  of  leaves  or  on  short  axillary  racemes,  well- 
developed  pedicels,  and  blue  corollas  not  extend- 
ing much  beyond  the  obovate  sepals.  In  addition, 
the  stems  are  often  long  and  much-branched,  and 
many  parts  have  short  sharp  hairs.  There  has  been 
a  controversy  over  the  correct  name  for  this  spe- 
cies. An  earlier  name,  V.  didyma  Tenore,  has  been 
used  for  this  species,  but  many  European  authors 
consider  that  name  a  nomen  ambiguum  and  reject 
it  in  favor  of  V.  polita.  This  species  is  closely 
related  to  V.  persica. 


Veronica  serpyllifolia  L.,  Sp.  PI.  12.  1753.  V. 
tenella  All.,  Fl.  Pedemont.  1:  75.  1785.  V.  hum- 
ifusa  Dickson,  Trans.  Linn.  Soc.  2:  228.  1794. 
V.  serpyllifolia  var.  humifusa  (Dickson)  Vahl, 
Enum.  PI.  1:  65.  1805.  V.  crenulata  Sesse  & 
Mocino,  Fl.  Mex.  5.  1892,  non  Ruiz  &  Pavon 
1798.  Figure  2. 

Small  herbs  5-20  cm  tall,  erect  stems  un- 
branched,  sometimes  with  repent  sterile  leafy 
branches  forming  small  mats,  leafy  stems  0.5-0.9 
mm  diam.,  puberulent  with  thin  ascending  whitish 
hairs  0.1-0.2  mm  long.  Leaves  opposite  in  the 
lower  half  of  the  stem,  alternate  and  smaller  be- 
low the  flowering  nodes,  petioles  0.5-2  mm  long 
or  the  leaves  subsessile;  leaf  blades  3.5-12  mm 
long,  1.5-9  mm  wide,  broadly  ovate  to  broadly 
oblong,  becoming  narrower  distally  (6X3  mm), 
apex  rounded  to  obtuse,  margin  subentire  to  ob- 
scurely crenate  with  teeth  0.1-0.2  mm  high,  ob- 
tuse to  rounded  at  the  base,  glabrous  above  and 
below,  venation  subpalmate.  Inflorescences  4-15 
cm  long,  erect,  racemes  or  spike-like,  rachis  ap- 
pressed puberulent,  flowers  distant  or  closely 
spaced,  bracts  2-6  mm  long,  1-3  mm  wide,  nar- 
rowly elliptic-oblong  to  narrowly  ovate-elliptic, 
sessile,  glabrous,  pedicels  2-4  mm  long  (-6  mm 
in  fruit),  glabrous  or  puberulent.  Flowers  usually 
puberulent  at  the  base,  calyx  lobes  4,  subequal, 
2-4  mm  long,  ca.  0.9  mm  wide,  lanceolate,  gla- 
brous or  minutely  puberulent;  corolla  ca.  4  mm 
long,  lobes  2-4  mm  wide,  pale  blue  to  blue-purple 
and  with  darker  lines;  styles  1.8-2.5  mm  long. 
Fruits  2.3-4  mm  long,  3-4.5  mm  wide,  obovoid 
with  slightly  or  moderately  depressed  apex  (ob- 
cordate), glabrous  or  minutely  puberulent  along 
the  distal  margins;  seeds  ovoid,  flattened  on  1 
face,  smooth. 

Plants  of  damp  open  or  partly  shaded  sites  in 
montane  forest  formations,  2400-3300  m  eleva- 
tion. Probably  flowering  throughout  the  year;  it  is 
common  along  the  Carretera  Interamericana  in  the 
Cordillera  de  Talamanca.  This  species,  a  native  of 
western  Eurasia,  is  now  widely  naturalized  in 
North  America  and  higher  elevations  in  Mexico 
and  Central  America. 

Veronica  serpyllifolia  is  recognized  by  its  nar- 
row unbranched  erect  stems,  small  subentire 
leaves,  short-pedicellate  flowers,  and  subequal  ob- 
long calyx  lobes.  This  species  is  the  second  most 
commonly  collected  species  of  Veronica  in  Costa 
Rica. 


68 


FIELDIANA:  BOTANY 


List  of  Accepted  Species  of  Scrophulariaceae 

Key:  END-CR  =  endemic  to  continental  Costa 
Rica;  END-CR&WP  =  endemic  to  Costa  Rica 
and  western  Panama;  END-WP  =  endemic  to 
western  Panama;  INTRO  =  introduced  weed; 
ORNAM  =  cultivated  ornamental;  ORNAM  & 
NAT  =  cultivated  and  naturalized;  ??  =  not  col- 
lected in  Costa  Rica  but  known  from  nearby  ar- 
eas. Total  number  of  species  covered  is  72;  the 
number  of  documented  native  species  is  48. 

Alectra  aspera  INTRO 
Alonsoa  meridionalis 
Angelonia  angustifolia  ORNAM 
Anisantherina  hispidula 
Antirrhinum  majus  ORNAM 

Bacopa  axillaris 
Bacopa  bacopoides  ?? 
Bacopa  egensis 
Bacopa  laxiflora 
Bacopa  monnieri 
Bacopa  monnierioides 
Bacopa  repens 
Bacopa  salzmannii 
Bacopa  sessiliflora 
Benjaminia  reflexa 
Buchnera  pusilla 
Buchnera  weberbaueri 

Calceolaria  irazuensis  END-CR 
Calceolaria  mexicana 
Calceolaria  microbe/aria 
Calceolaria  perfoliata 
Calceolaria  tripartita 
Capraria  biflora 
Castilleja  an'ensis 
Castilleja  irasuensis  END-CR 
Castilleja  lentil  END-CR 
Castilleja  quirosii  END-CR&WP 
Castilleja  talamancensis  END-CR 
Castilleja  tayloriorum  END-CR 
Cymhalaria  muralis  INTRO 

Darcya  costaricensis  END-CR 
Darcva  reliquiarum  END-WP 
Digitalis  purpurea  ORNAM  &  NAT 

Escobedia  grandiflora 
Hemichaena  fruticosa 

Lamarouxia  gutierrezii  END-CR&WP 

Lamarouxia  lanceolata 

Lamarouxia  viscosa 

Leucocarpus  perfoliatus 

Limosella  acaulis 

Linaria  canadensis  ?? 

Linaria  vulgaris  ORNAM 

Lindernia  Crustacea 

Lindernia  diffusa 

Lindernia  dubia 

Lophospernnim  eruhescens  ORNAM  &  NAT 

Maurandya  barclaiana  ORNAM 
Maurandya  scandens  ORNAM 


Mazus  pumila  INTRO 
Mecardonia  prociunbens 
Micranthemum  umhroxum  ?? 
Mimulus  glabratus  '.''.' 

Penstemon  gentianoitles  ORNAM 

Rus.\t'lia  equisctiformis  ORNAM  &  NAT 
Russelia  sa  rmt-n  to  MI 

Schistophragma  mt-.\icana 
Scoparia  an  nun 
Scoptiria  tiulci\ 
Sibthorpia  rcpena 
Stemodia  angulata 
Stemodid  (Inniiitifolia 
Stemodia  peduncularis 
Stemodia  verticilltita 

Tetranema  florilntnditm  END-CR 
Tetnnit-nui  gamboantun  END-CR 
Torcnia  fournieri  ORNAM 
Ton-nia  thouarsii  INTRO 

Veronica  arvenis  INTRO 
Veronica  peregrina  INTRO 
Veronica  persica  INTRO 
Veronica  polita  INTRO 
Veronica  serpyllifolia  INTRO 


SCHLEGELIACEAE  Reveal 

By  William  Burger  and  Kerry  Barringer 

REFERENCES — W.  D'Arcy,  Scrophulariaceae,  in 
Flora  of  Panama.  Ann.  Missouri  Bot.  Gard.  66: 
173-272,  1979.  R.  Olmstead  &  R  Reeves,  Evi- 
dence for  the  polyphyly  of  the  Scrophulariaceae 
based  on  chloroplast  rbcL  and  ndhF  sequences. 
Ann.  Missouri  Bot.  Gard.  82:  176-191,  1995.  J. 
Reveal,  Newly  required  suprageneric  names  in 
vascular  plants.  Phytologia  79:  68-76,  1996. 

Shrubs,  subshrubs  or  lianas,  erect  or  climbing, 
terrestrial  or  epiphytic,  autotrophic,  bisexual, 
stems  without  internal  phloem,  glabrous  or  with 
simple,  eglandular  or  glandular,  unicellular  or 
multiccllular  hairs,  nodes  lacking  intcrpetiolar 
lines  or  glandular  fields;  stipules  absent.  Leaves 
opposite  or  subopposite,  simple,  entire  (serrulate 
in  Synopsis),  usually  coriaceous  to  subcoriaceous, 
glabrous  or  puberulent,  pinnately  veined.  Inflo- 
rescences racemes,  cymes,  or  of  solitary  flowers 
in  leaf  axils,  sometimes  in  dense  fascicles,  brac- 
teoles  present,  pedicels  well  developed.  Flowers 
bisexual,  small  to  large,  often  showy,  calyx  irreg- 
ularly lobed,  5-lobed  or  split,  persistent  and  usu- 
ally accrescent  in  fruit;  corolla  tubular  to  funnel- 
form,  bilaterally  symmetric  or  almost  radially 


BURGER:  FLORA  COSTARICENSIS 


69 


symmetric,  5-Iobed,  the  tube  straight  (not  saccate 
or  spurred),  stamens  4,  borne  on  the  corolla  tube, 
filaments  tree,  alternating  with  the  corolla  lobes, 
anthers  2-thecous,  thecae  equal,  distinct,  a  stami- 
node  present,  disc  absent;  pistil  solitary,  ovary  su- 
perior. 2-locular,  ovules  many,  unitegmic,  borne 
on  2  large  axile  placentas,  the  style  1,  terminal, 
stigmas  simple.  Fruits  baccate,  indehiscent;  seeds 
many,  with  reticulate  exotesta,  the  embryo  small, 
straight. 

Schlegeliaceae  is  a  family  of  four  Neotropical 
genera  and  about  30  species.  The  genera  Gibson- 
iothamnus  and  Schlegelia  are  found  in  Costa  Rica, 
and  they  are  closely  related.  Until  recently  the 
genera  of  this  family  were  included  in  either  the 
Scrophulariaceae  or  Bignoniaceae  but  were  atyp- 


ical for  either  family.  Evidence  from  chloroplast 
DNA  sequences  (Olmstead  &  Reeves,  1995,  cited 
above)  indicates  that  the  family  is  not  closely  re- 
lated to  either  of  these  families  but  is  a  distinct 
group,  probably  most  closely  related  to  the  basal 
Verbenaceae,  such  as  Callicarpa.  In  recognition 
of  their  distinctiveness,  Reveal  (1996,  cited 
above)  established  the  family  Schlegeliaceae  for 
this  group.  Our  representatives  are  distinguished 
from  most  Bignoniaceae  by  their  simple  entire 
leaves  and  baccate  fruits.  The  family  also  differs 
from  many  bignons  in  lacking  tendrils,  interpetio- 
lar  lines,  and  interpetiolar  gland  fields  on  the 
stems.  The  Schlegeliaceae  are  distinguished  from 
most  Scrophulariaceae  by  their  woody  habit  and 
baccate-indehiscent  fruits. 


Key  to  the  Genera  of  Schlegeliaceae  in  Costa  Rica 

la.  Epiphytic  shrubs;  corollas  usually  reddish  to  purple  (less  often  white),  tubular  and  almost  radially 
symmetric;  staminodes  <  5  mm  long;  fruits  with  fleshy  covering;  leaves  usually  4-8  cm  long  .  .  . 
Gibsoniothamnus 

Ib.  Lianas  or  rarely  small  trees  or  epiphytic  shrubs;  corollas  white  to  pinkish  (in  Costa  Rica),  tubular- 
campanulate  and  usually  clearly  bilaterally  symmetric;  staminodes  usually  >  5  mm  long;  fruits  with 

a  hard  outer  covering;  larger  leaves  usually  >  10  cm  long  (except  in  5.  brachyantha) 

Schlegelia 


Gibsoniothamnus  L.  O.  Williams 

REFERENCES — W.  D'Arcy,  Gibsoniothamnus 
(Scrophulariaceae)  in  Flora  of  Panama,  Part  9. 
Ann.  Missouri  Bot.  Card.  66:  220-227.  1979.  A. 
Gentry,  Note  on  Gibsoniothamnus.  Fieldiana,  Bot. 
34:  55.  1971.  A.  Gentry,  Gibsoniothamnus  (Scro- 
phulariaceae) in  Panama.  Ann.  Missouri  Bot. 
Gard.  61:  533-537.  1974.  A.  Gentry,  New  or 
noteworthy  species  of  middle  American  Bigno- 
niaceae. Wrightia  7:  83-89.  1982.  P.  C.  Standley 
&  L.  O.  Williams,  Gibsoniothamnus  (Scrophular- 
iaceae) in  Flora  of  Guatemala.  Fieldiana,  Bot.  24: 
356-359.  1973.  L.  O.  Williams,  An  overlooked 
genus  of  the  Scrophulariaceae.  Fieldiana,  Bot.  32: 
211-214.  1970. 

Shrubs  or  small  trees,  epiphytic  or  rarely  ter- 
restrial, to  6  m  tall,  young  stems  terete  or  angular, 
glabrous  or  pubescent,  leaf  base  persistent  and  be- 
coming conspicuously  raised,  older  stems  with  a 
light-colored  bark.  Leaves  opposite  or  suboppos- 
ite,  equal  or  unequal  (anisophyllous)  at  a  node, 
clearly  articulated  at  the  base,  petioles  usually 
short,  merging  with  the  base  of  the  blade,  blades 
simple  and  entire,  often  elliptic,  coriaceous  or 


subcoriaceous  (rarely  chartaceous),  glabrous  or 
puberulent,  usually  with  glandular  pits  scattered 
on  the  underside,  domatia  often  present  in  the  ax- 
ils of  veins  and  midvein  below.  Inflorescences  of 
solitary  axillary  flowers  or  flowers  in  terminal 
cymes,  panicles,  racemes,  or  condensed  and  fas- 
ciculate, peduncles  present  or  absent,  pedicels 
elongate,  merging  with  the  calyx  and  often  the 
same  color  as  the  calyx,  usually  bracteate.  Flow- 
ers usually  colorful  throughout,  calyx  cupulate  or 
campanulate,  glabrous  or  puberulent,  lobes  usu- 
ally 5,  narrow  and  tooth-like  to  elongated  and  lat- 
erally compressed  or  winged,  persistent  and  often 
accrescent  in  fruit;  corolla  tubular  to  narrowly 
funnelform,  often  brightly  colored  in  purples  and 
reds,  radially  symmetric  or  slightly  2-lipped,  gla- 
brous externally  or  ciliolate  along  the  distal  edges, 
lobes  5,  rounded  and  subequal,  much  shorter  than 
the  tube,  tube  interior  with  hairs  near  the  stamen 
insertion;  stamens  4,  included,  equal  or  in  2  sub- 
equal  pairs,  inserted  above  the  base  of  the  tube, 
filaments  filiform,  bent  near  the  apex,  anthers  with 
a  broad  connective  and  2  separate  parallel  or 
slightly  divaricate  thecae,  staminode  1,  usually 
shorter  than  the  stamens;  disc  lacking;  ovary  2- 


70 


FIELDIANA:  BOTANY 


Gibsoniothamnus 
pterocalyx 


Schlegelia 
parvi flora 


Schlegelia 
fasti giata 


FIG.  9.     Schlcgcliaceae:  epiphytic  shrubs,  vines,  or  trees  with  berry-like  fruits;  species  of  Gibsoniothamnus  and 
Schlegelia. 


BURGER:  FLORA  COSTARICENSIS 


71 


locular,  ovules  many,  style  slender,  longer  than  the 
stamens,  stigma  capitate.  Fruits  fleshy  berries, 
globose  to  subglobose,  juicy  or  mucilaginous; 
seeds  many,  seed  coat  reticulate,  the  margins  or 
the  reticulae  bearing  elongate  filaments  that  are 
sticky  or  mucilaginous;  endosperm  absent  in  ma- 
ture seeds,  embryo  straight. 

Gibsoniothamnus  is  a  genus  of  12  species  na- 
tive to  southern  Mexico,  Central  America,  and  ad- 
jacent Colombia.  The  paucity  of  earlier  collec- 
tions resulted  in  D'Arcy's  broad  specific  concepts 


(1979),  but  additional  collecting  appears  to  justify 
the  recognition  of  a  larger  number  of  geographi- 
cally isolated  species.  The  domatia  vary  from 
tufts  of  hairs  in  vein  axils  to  well-defined  pits  or 
pockets  in  the  lower  leaf  surface;  they  may  be 
lacking  in  some  species.  Our  species  are  less  pu- 
bescent and  have  more  prominent  calyx  lobes 
than  does  G.  cornutus  (J.  D.  Smith)  A.  Gentry, 
which  is  restricted  to  Mexico  and  Guatemala  and 
probably  includes  G.  pithecobius  (Standl.  &  Stey- 
erm.)  L.  O.  Williams. 


Key  to  the  Species  of  Gibsoniothamnus 

la.  Flowers  usually  borne  in  distal  fascicles  of  5-25  flowers;  leaf  blades  elliptic-oblong  to  obovate,  to 
5  cm  wide  2 

Ib.  Flowers  1-3  in  the  axils  of  leaves  or  terminal;  leaf  blades  narrowly  elliptic  to  ovate  or  oblanceolate, 

to  3  cm  wide  3 

2a.  Leaf  blades  to  8  cm  long  and  to  3  cm  wide,  the  major  veins  deeply  impressed  above  when 

dried;  young  stems  usually  with  stiff  straight  hairs;  flowers  in  fascicles  of  5-1 1  

G.  epiphyticus 

2b.  Leaf  blades  to  15  cm  long  and  to  6  cm  wide,  the  major  veins  not  deeply  impressed  when  dried; 
young  stems  glabrous  or  with  few  hairs;  flower  in  fascicles  of  10-25  G.  parvifolius 

3a.  Corolla  red  or  purple;  sepal  lobes  and  veins  not  expanded  laterally  to  form  winged  margins,  calyx 
drying  blackish  or  yellowish;  1000-1500  m  elevation  in  evergreen  (rarely,  deciduous)  forests  of 
the  Pacific  slope G.  parvifolius 

3b.  Corolla  lavender,  rose,  or  white,  sepal  lobes  expanded  laterally  from  the  veins  and  somewhat 
winged,  often  drying  yellowish;  300-1400  m  elevation  in  evergreen  forests  on  the  Caribbean  slope 
and  along  the  Continental  Divide  4 

4a.  Corolla  lilac  or  rose;  calyx  lobes  only  slightly  expanded  along  the  veins,  leaves  to  70  mm  long; 
400-900  m  elevation  on  the  Caribbean  slope  G.  pterocalyx 

4b.  Corolla  white,  calyx  lobes  expanded  laterally  along  the  midveins  and  conspicuously  winged,  leaves 
to  25  mm  long;  1200-1400  m  elevation  on  Cerro  Colorado,  Panama  G.  stellatus 


Gibsoniothamnus  epiphyticus  (Standl.)  L.  O. 
Williams,  Fieldiana,  Bot.  34:  120.  1972.  Cler- 
odendrum  epiphyticum  Standl.,  Publ.  Field 
Mus.  Nat.  Hist.  Bot.  Ser.  22:  168.  1940.  Figure  9. 

Epiphytic  shrubs  to  ca.  1  m  tall,  leafy  branches 
2-4  mm  thick,  terete  to  slightly  angulate  below 
the  raised  leaf  bases,  with  stiff  straight  hairs  0.3- 
1  mm  long  but  glabrescent.  Leaves  opposite  or 
subopposite,  usually  of  2  different  sizes  at  each 
node  (anisophyllous),  petioles  4-9  mm  long,  0.8- 
1 .7  mm  thick,  puberulent  or  glabrous;  leaf  blades 
2-8  cm  long,  1.2-3  cm  wide,  elliptic  to  elliptic- 
oblong  or  ovate-elliptic,  apex  acuminate  to  acute 
or  rounded,  margin  entire  and  reflexed,  base  cu- 
neate  to  acute,  subcoriaceous,  lustrous  above,  gla- 
brous or  with  few  straight  hairs  to  1  mm  long, 
midvein  impressed  above,  2°  veins  2  or  3/side, 


domatia  in  vein  axils  beneath.  Inflorescences  of 
5-1 1  flowers  in  short  axillary  panicles  or  from  a 
fasciculate  base  near  the  apex  of  a  branchlet,  1° 
peduncles  to  7  mm  long,  bracts  2-4  mm  long, 
narrowly  triangular,  pedicels  15-20  mm  long, 
merging  gradually  with  the  calyx,  glabrous  or 
with  straight  hairs  to  0.8  mm  long.  Flowers  with 
calyx  8-1 1  mm  long,  3-4  mm  diam.,  glabrous  or 
slightly  puberulent  on  the  angles,  lobes  4-6  mm 
long,  1-1.5  mm  wide  at  the  base;  corolla  14-21 
mm  long,  3-4  mm  diam.,  tubular,  glabrous,  red- 
violet  or  purple,  lobes  1.5-2.5  mm  long;  stamens 
attached  3-4  mm  from  base  of  tube,  filaments  18- 
20  mm  long;  staminode  4-5  mm  long;  pistil  gla- 
brous, style  19-22  mm  long.  Mature  fruits  to  1 
cm  diam.,  becoming  white. 

Rarely  collected  plants  of  evergreen  montane 
cloud  forest  formations  of  the  Caribbean  slope, 


72 


FIELDIANA:  BOTANY 


600-1500  m  elevation.  Flowering  in  April-July. 
This  species  is  known  only  from  central  Costa 
Rica. 

Gibsoniothamnus  epiphyticus  is  recognized  by 
its  epiphytic  habit,  stiff  leaves  with  deeply  im- 
pressed venation,  fasciculate  flowers,  prominent 
winged  calyx  lobes,  and  red  or  purple  tubular  co- 
rollas. The  stiff  multicellular  hairs  are  distinctive 
when  present,  but  the  pubescence  is  often  decid- 
uous. This  species  is  similar  to  G.  pterocalyx  but 
differs  from  that  species  in  having  narrower,  sub- 
coriaceous  leaves  with  veins  impressed  above, 
red-violet  corollas,  and  shorter,  more  triangular 
calyx  lobes.  G.  mirificus  A.  Gentry  of  central  Pan- 
ama is  also  similar  but  that  species  has  solitary 
flowers,  longer  calyx  lobes,  and  thinner  leaves. 

Gibsoniothamnus  parvifolius  Barringer  (Novon 
9:  476.  1999).  Figure  9. 

Epiphytic  shrubs  to  2  m  tall,  leafy  stems  1-7 
mm  thick,  glabrous  to  sparsely  pubescent  with 
hairs  0.2-0.6  mm  long,  winged  or  angled  in  early 
stages  but  soon  terete  and  glabrescent.  Leaves 
equal  or  sometimes  strongly  unequal  at  a  node 
(with  the  smaller  leaves  more  rounded),  petioles 
2-15  mm  long,  0.4-2  mm  thick,  glabrous;  leaf 
blades  1.6-12  cm  long,  0.6-5  cm  wide,  blades 
lanceolate  to  narrowly  elliptic  or  obovate-elliptic, 
apex  acuminate,  margin  entire  and  reflexed  on 
drying,  base  acute  or  cuneate  and  decurrent  on  the 
petiole,  drying  subcoriaceous,  lustrous  and  dark 
olive-green  above,  glabrous  above,  glabrous  or 
with  thin  hairs  beneath,  2°  veins  usually  2-5/side, 
impressed  above,  pit  domatia  often  present  near 
the  vein  axils  beneath.  Inflorescences  of  solitary 
flowers  in  distal  leaf  axils  (2/node),  bracteoles  ca. 
1  mm  long,  linear,  pedicels  4-30  mm  long,  0.3- 
1.5  mm  thick,  glabrous  or  with  few  glandular 
hairs  in  early  stages,  bracteolate  at  base.  Flowers 
glabrous  externally,  calyx  campanulate,  tube  ca.  4 
mm  long  with  truncated  distal  margin,  lobes  4- 
10  mm  long,  0.5-1  mm  wide,  narrowly  triangular, 
glabrous,  spreading,  drying  dark;  corolla  18-38 
mm  long,  1.5-5  mm  diam.,  tubular,  reddish  violet 
to  wine-red  or  red,  lobes  2-4  mm  long,  upper 
lobes  broadly  ovate,  lateral  and  median  lobes 
ovate,  usually  more  darkly  colored  than  the  tube; 
filaments  15-26  mm  long,  staminode  1.7-5  mm 
long;  pistil  glabrous,  style  elongate,  ca.  18  mm 
long.  Fruits  becoming  10  mm  in  diam.,  globose 
or  oblate,  fleshy,  white  at  maturity. 

Plants  of  evergreen  cloud  forest  formations  in 
the  Chiriqui  highlands  and  in  the  Cordillera  de 


Guanacaste  and  Cordillera  de  Tilaran,  at  500- 
1500  m  elevation.  Flowering  in  February  and  Au- 
gust; fruiting  in  August.  This  species  is  known 
only  from  Costa  Rica  and  adjacent  Panama. 

Gibsoniothamnus  pan'ifolius  is  distinguished 
by  its  often  small  glabrous  leaves,  solitary  axil- 
lary flowers,  narrow  calyx  lobes,  slender  tubular 
purplish  or  reddish  corollas,  and  white  fleshy 
fruits.  The  specimens  from  northern  Costa  Rica 
tend  to  have  more  angled  stems  and  shorter  sta- 
minodia  than  the  Panamanian  specimens.  This 
species  can  be  distinguished  from  G.  epiphyticus 
of  central  Costa  Rica  by  its  small  obovate  or  lan- 
ceolate leaves,  its  calyx  lobes  more  than  5  mm 
long,  and  its  one-  or  two-flowered  inflorescences. 
The  larger-leaved  collections  resemble  G.  gran- 
diflorus  A.  Gentry  &  Barringer  of  central  Panama 
(see  Fig.  9).  Two  Costa  Rican  collections  were 
provisionally  placed  under  G.  grandiflorus.  but 
they  are  now  placed  under  an  expanded  circum- 
scription of  G.  parvifolius;  they  are  G.  Herrera 
3473  (MO)  and  J.  D.  Smith  6730  (us). 

Gibsoniothamnus  pterocalyx  A.  Gentry;  Ann. 
Missouri  Bot.  Gard.  61:  535.  1974.  Figure  9. 

Epiphytic  shrubs  to  2  m  tall,  leafy  stems  0.7- 
3  mm  thick,  glabrous,  at  first  slightly  angled  but 
soon  terete  and  pale  grayish.  Leaves  opposite  or 
subopposite,  unequal  or  subequal  at  a  node,  artic- 
ulated at  the  base,  petioles  2-13  mm  long,  0.4- 
0.8  mm  thick,  glabrous;  leaf  blades  1.7-7  cm 
long,  1.5-3.4  cm  wide,  narrowly  elliptic  to  nar- 
rowly obovate  or  oblanceolate,  apex  bluntly  acute 
or  rounded  and  slightly  emarginate,  margin  entire 
and  slightly  revolute,  base  acute  to  cuneate,  dry- 
ing subcoricaceous  to  chartaceous,  dark  olive 
green  and  lustrous  above,  glabrous,  2°  veins  2  or 
3/side,  strongly  ascending;  pit  domatia  often  pre- 
sent near  vein  axils  beneath.  Inflorescences  of 
solitary  axillary  flowers  or  few  flowers  in  con- 
densed axillary  panicles,  1°  peduncles  to  1  cm 
long,  bracts  2-3  mm  long,  narrowly  triangular, 
sparsely  villous,  pedicels  7-25  mm  long,  0.4-0.8 
mm  thick,  glabrous,  merging  gradually  with  the 
base  of  the  calyx.  Flowers  pale  lilac  or  rose,  gla- 
brous externally,  calyx  8-12  mm  long,  2-4  mm 
diam.,  calyx  lobes  5-9  mm  long,  0.4-1.3  mm 
wide,  linear  and  slightly  winged  at  the  base;  co- 
rolla 18-25  mm  long,  lilac  or  pink,  glabrous,  tube 
3-5  mm  diam.,  lobes  2-3  mm  long,  rounded;  fil- 
aments 1 5-20  mm  long,  staminode  5-6  mm  long, 
filiform;  ovary  glabrous,  style  16-19  mm  long. 


BURGER:  FLORA  COSTARICENSIS 


73 


Fruits  becoming  up  to  10  mm  diam.,  obovoid  to 
oblong  with  truncated  apex,  white. 

Infrequently  collected  plants  of  evergreen  low- 
er montane  rain  forest  formations  on  the  Carib- 
bean slope,  400-900  m  elevation  (intensive  col- 
lecting over  many  years  at  Monteverde  indicates 
that  the  upper  altitudinal  limit  of  900  m  is  real, 
and  not  an  artifact  of  insufficient  collecting). 
Flowering  in  April-July  and  October.  This  species 
ranges  along  the  Caribbean  escarpment  from 
northern  Costa  Rica  to  western  Panama. 

Gibsoniothamnus  pterocalyx  is  recognized  by 
its  epiphytic  habit,  glabrous  parts,  small  narrow 
lustrous  leaves,  usually  only  one  to  three  pink  or 
lilac  (white)  flowers  at  a  node,  and  prominent  se- 
pal lobes.  The  leaf  undersides  are  often  purplish, 
especially  along  the  veins.  This  species  is  super- 
ficially similar  to  G.  parvifolius  (q.v.). 

Gibsoniothamnus  stellatus  A.  Gentry  &  Bun  m- 
ger,  Novon  5:  121-122,  1995. 

Epiphytic  shrubs,  stems  angled  and  with  raised 
leaf  bases,  leafy  stems  ca.  2  mm  thick,  glabrous, 
becoming  grayish.  Leaves  of  the  same  node  un- 
equal or  subequal,  petioles  3-5  mm  long,  gla- 
brous, without  lateral  wings;  leaf  blades  5-25 
mm  long,  4-16  mm  wide  (larger  blades  15-25  X 
9-16  mm),  ovate-lanceolate  to  elliptic,  apex 
acute,  base  cuneate,  drying  coriaceous,  glabrous, 
punctate;  2°  veins  2  or  3/side,  domatia  sometimes 
present  near  the  base  beneath.  Inflorescences  of 
solitary  flowers  axillary  to  distal  leaves,  bracteate 
at  the  base,  pedicels  10-12  mm  long,  expanded 
distally,  glabrous.  Flowers  with  campanulate  pur- 
plish calyx,  glabrous,  tube  6-8  mm  long,  lobes  8- 
9  mm  long,  triangular  with  wings  3-4  mm  wide; 
corolla  20-25  mm  long,  4-5  mm  diam.,  tubular, 
white,  upper  and  lateral  lobes  3-4  mm  long  and 
wide,  ciliolate  along  the  edge;  filaments  17-18 
mm  long,  staminode  5-6  mm  long;  pistil  gla- 
brous. Fruits  not  seen  at  maturity. 

Gibsoniothamnus  stellatus  is  distinguished  by 
its  shrubby  epiphytic  habit,  small  leaves,  flowers 
with  winged  calyx  lobes  (giving  a  star-like  form), 
and  white  corollas.  This  species  is  known  only 
from  two  collections  from  1200-1400  m  elevation 
on  Cerro  Colorado  in  the  provinces  of  Bocas  del 
Toro  and  Chiriqui  in  western  Panama.  They  were 
flowering  in  April  and  July.  This  species  is  closely 


related  to  G.  alatus  of  eastern  Panama  and  adja- 
cent Choc6,  which  also  has  white  corollas  and 
winged  calyx  lobes  but  has  much  larger  leaves. 


Schlegelia  Miquel 

REFERENCES — A.  Gentry,  Schlegelia  (Bignoni- 
aceae)  in  Flora  of  Panama.  Ann.  Missouri  Bot. 
Card.  60:  923-930,  1973.  P.  Standley  &  L.  O. 
Williams,  Schlegelia  (Scrophulariaceae)  in  Flora 
of  Guatemala.  Fieldiana  Bot.  24,  pt.  9:  396-400, 
1973. 

Shrubs,  woody  vines,  lianas  or  small  trees,  ter- 
restrial or  epiphytic,  wood  without  anomalous 
vasculature,  stems  usually  terete;  pseudostipules 
small,  usually  appressed  against  the  branchlets. 
Leaves  opposite,  simple,  with  thick  terete  peti- 
oles, usually  becoming  articulated  at  the  base, 
blades  often  coriaceous,  margins  entire,  venation 
pinnate.  Inflorescences  terminal  or  axillary  pan- 
icles, fascicles  or  axillary  racemes,  peduncles  usu- 
ally woody,  often  with  conspicuous  small  bracts 
beneath  the  pedicels.  Flowers  with  a  subcoria- 
ceous  tubular  or  cupulate  calyx,  entire  or  splitting 
irregularly  into  small  lobes,  persisting  and  split- 
ting in  fruit;  corolla  campanulate  to  narrowly  tu- 
bular, white  to  pink  or  red  (yellow),  glabrous  ex- 
ternally but  glandular  lepidote  on  the  lobes  within, 
lobes  5,  rounded;  stamens  4,  in  2  unequal  pairs, 
borne  on  the  lower  half  of  the  corolla  tube,  in- 
cluded, anthers  glabrous,  thecae  divaricate,  a 
small  staminode  usually  present;  ovary  2-locular 
(sometimes  1-  locular  near  the  apex),  placenta 
central  on  the  septum  in  each  locule,  ovules  many, 
style  simple,  stigma  bifid  (trifid).  Fruits  berries 
with  hard  crustaceous  indehiscent  pericarp,  usu- 
ally globose;  seeds  narrow  and  angular,  wingless, 
small,  embedded  in  pulp. 

Schlegelia  is  a  genus  of  ca.  15  species,  ranging 
from  eastern  Mexico  and  the  West  Indies  to  Bra- 
zil. The  simple,  opposite,  usually  stiff  leaves, 
shrubby,  often  epiphytic  or  vining  habit,  usual 
lack  of  conspicuous  pubescence,  woody  inflores 
cence  branches,  calyx  tubes  with  poorly  devel- 
oped lobes,  and  round  berry-like  fruits  with  stiff 
rind  help  distinguish  this  genus.  Dr.  William 
D'Arcy  (MO)  is  currently  studying  this  genus. 


74 


FIELDIANA:  BOTANY 


Key  to  the  Species  of  Schlegelia 

la.  Corollas  usually  <  20  mm  long;  calyx  tube  4-6  mm  long  at  anthesis;  fruits  up  to  12  mm  diam.; 
leaves  usually  drying  stiffly  coriaceous  and  with  the  minor  venation  usually  not  evident  on  the 

upper  surface;  plants  often  epiphytic;  2-2100  m  elevation   2 

Ib.  Corollas  20-43  mm  long;  calyx  5-12  mm  long;  fruits  12-40  mm  diam.;  leaves  drying  stiffly 
chartaceous  to  subcoriaceous,  the  minor  veins  often  elevated  on  the  upper  surface;  plants  usually 

vines  or  lianas;  10-900(-1500)  m  elevation 3 

2a.  Corollas  mostly  8-1 1  mm  long;  leaf  blade  acute  to  obtuse  or  rounded  at  apex;  commonly  collected 

S.  parviflora 

2b.  Corollas  usually  15-20  mm  long;  leaf  blade  obtuse  to  rounded  at  apex;  rarely  collected   

S.  brachyantha 

3a.  Corolla  20-28  mm  long,  narrowly  tubular  with  short  (1-3  mm)  lobes;  fruits  12-16  mm  diam.; 
inflorescences  of  many  (>  15)  flowers  in  dense  woody  fascicles  or  panicles,  peduncles  1-20  cm 

long    S.  fastigiata 

3b.  Corolla  30-43  mm  long,  tubular-campanulate  with  prominent  (to  10  mm)  lobes;  fruits  30-40 
mm  diam.;  inflorescences  of  few  (1-5)  flowers  in  racemose  arrangements,  peduncles  usually  < 
1  cm  long S.  nicaraguensis 


Schlegelia  brachyantha  Griseb.,  Cat.  PI.  Cub. 
191.  1866. 

Lianas  or  vine-like  shrubs  to  6  m  high,  leafy 
stems  1-8  mm  thick,  glabrous  or  sparsely  puber- 
ulent  near  the  apex  and  on  the  new  growth,  drying 
grayish  or  dark,  terete  or  slightly  quadrangular. 
Leaves  with  petioles  3-9  mm  long,  1-2.5  mm 
thick,  becoming  articulated  and  thicker  at  the 
base,  glabrous  or  with  few  minute  (0.1-0.2  mm) 
hairs;  leaf  blades  4-12  cm  long,  1.5-8  cm  wide, 
broadly  elliptic  to  broadly  oblong  or  oblong-or- 
bicular, apex  bluntly  obtuse  to  rounded  or  short- 
acuminate,  base  obtuse  to  rounded  and  subtrun- 
cate,  drying  subcoriaceous  and  slightly  lustrous 
above  with  margin  often  recurved,  glabrous  above 
and  below,  punctate  beneath,  major  2°  veins  3-57 
side,  loop-connected  distally.  Inflorescences  ax- 
illary to  leaves  or  leafless  nodes,  apparently  ses- 
sile fascicles  with  few  (2-5)  flowers,  peduncles 
very  short  with  bracts  ca.  2  mm  long  on  congested 
nodes,  pedicels  2-8  mm  long,  glabrous  and  dry- 
ing black.  Flowers  not  seen  at  anthesis,  flower 
buds  3-5  mm  long,  with  few  minute  hairs,  drying 
dark;  corolla  15-20  mm  long,  white  or  pink,  5- 
6  mm  wide  at  throat,  lobes  ca.  5  mm  long.  Fruits 
6-10  mm  diam.,  green  turning  red. 

Rarely  collected  plants  of  evergreen  forest  for- 
mations, 20-1800  m  elevation.  In  Costa  Rica  it  is 
only  known  from  the  high  plateau  southeast  of 
Chatham  Bay,  Cocos  Island,  flowering  and  fruit- 
ing in  April  (R.  Foster  4125  F).  Also  known  from 
the  Chiriqui  highlands  in  Panama,  Venezuela,  and 
the  West  Indies. 


Schlegelia  brachyantha  is  distinguished  by  its 
smaller,  often  rounded  leaves,  few-flowered,  ses- 
sile inflorescences,  and  few  collections  from 
widely  separated  geographical  areas. 

Schlegelia  costaricensis  Stand!,  described  in 
the  Flora  of  Costa  Rica,  part  3  (Publ.  Field  Mus. 
Nat.  Hist.  Bot.  Ser.  18:  1128,  1938)  has  been 
transferred  to  the  Boraginaceae  as  Bourreria  cos- 
taricensis (Standl.)  A.  Gentry  (see  Phytologia  26: 
67-68,  1973). 

Schlegelia  fastigiata  Schery,  Ann.  Missouri  Bot. 
Gard.  29:  367.  1942.  Figure  9. 

Lianas  or  vines,  climbing  stems  ca.  15  mm 
diam.,  leafy  stems  1.8-6  mm  thick,  terete  or 
somewhat  flattened,  glabrous  (minutely  puberu- 
lent).  Leaves  with  petioles  3-8  mm  long,  1 .5-3.5 
mm  thick,  glabrous,  usually  drying  dark  and  ar- 
ticulated at  the  base;  leaf  blades  10-26(-33)  cm 
long,  3-9(-13)  cm  wide,  narrowly  elliptic-oblong 
to  narrowly  ovate-oblong,  apex  acuminate  or 
acute,  base  obtuse,  drying  stiffly  chartaceous  to 
subcoriaceous,  glabrous  above  and  below,  the  mi- 
nor venation  slightly  elevated  on  the  upper  sur- 
face (dried),  2°  veins  8-16/side.  Inflorescences 
congested  woody  panicles  borne  on  short  (1-2 
cm)  peduncles  or  longer  (8-14  cm)  leafless  stems, 
usually  with  more  than  15  flowers,  woody  pani- 
cles (fascicles)  2-8  cm  long,  2-10  cm  wide, 
woody  branches  short  and  grayish,  pedicels  4-8 
mm  long,  ca.  0.4  mm  thick,  minutely  puberulent, 
drying  black,  subtended  by  a  pair  of  small  (1  mm) 
bracts  on  a  slender  2°  peduncle  2-4  mm  long. 


BURGER:  FLORA  COSTARICENSIS 


75 


Flowers  with  truncated  calyx  tube  5-8  mm  long, 
cupular  with  entire  or  slightly  lobed  distal  margin, 
5-7  mm  diam.  at  apex,  purple  to  red-violet  or 
pink,  usually  glabrous,  splitting  in  fruit  and  to  14 
mm  wide;  corolla  20-28  mm  long,  tubular,  gla- 
brous externally,  4-6  mm  diam.,  4-8  mm  wide  at 
the  spreading  lobes,  white,  lobes  2-5  mm  long, 
rounded,  pink  to  pale  violet;  filaments  8-12  mm 
long,  anthers  1-1.5  mm  long,  divaricate,  ca.  2  mm 
wide;  ovary  lepidote  (Gentry,  1973b).  Fruits  12- 
1 6  mm  long,  subglobose,  yellow  to  orange  or  pur- 
ple, surface  somewhat  muriculate,  glabrous;  seeds 
ca.  2  X  0.5  mm,  narrowly  triangular  or  curved. 

Plants  of  the  evergreen  rain  forest  formations 
of  the  Caribbean  lowlands  and  slopes,  10-900  m 
elevation.  Probably  flowering  and  fruiting 
throughout  the  year,  but  our  collections  were  col- 
lected between  July  and  March.  The  species  rang- 
es from  Guatemala  to  Ecuador. 

Schlegelia  fasti giata  is  recognized  by  its  vining 
habit,  thinner  leaves  with  more  2°  veins  than  our 
other  species  of  Schlegelia,  flowers  borne  in 
densely  branched,  woody  fascicles  on  slender 
bracteate  pedicels,  calyx  cup  with  entire  margin, 
and  narrowly  tubular  white  corolla  with  pink 
lobes.  The  minor  venation  is  usually  clearly  ele- 
vated on  the  upper  leaf  surface  of  the  dried  leaves, 
a  helpful  characteristic  in  making  specific  deter- 
minations. This  species  was  incorrectly  placed  un- 
der S.  sulphurea  Diels  in  Flora  of  Panama,  mis- 
spelled as  5.  sulfurea  (Gentry,  1973b);  that  spe- 
cies has  yellow  corollas  and  is  endemic  to  eastern 
Ecuador. 

Schlegelia  nicaraguensis  Standl.,  Trop.  Woods 
16:  44.  1928.  5.  silvicola  L.  O.  Williams,  Fiel- 
diana,  Bot.  34:  126,  tab.  3.  1972. 

Lianas  or  woody  vines  (rarely  trees  or  epi- 
phytic shrubs),  climbing  stems  ca.  15  mm  diam., 
leafy  stems  2-6  mm  thick,  terete  or  4-angled,  mi- 
nutely puberulent  in  early  stages,  glabrescent, 
grayish  and  with  prominent  lenticels;  pseudosti- 
pules  to  3  mm  long.  Leaves  with  petioles  3-14 
mm  long,  1.5-4.2  mm  thick,  rounded  but  with  2 
adaxial  ridges,  glabrous,  drying  grayish  or  yel- 
lowish; leaf  blades  (4-)8-22  cm  long,  (2-)3.5- 
10  cm  wide,  elliptic  to  elliptic-oblong,  elliptic-ob- 
ovate  or  elliptic-ovate,  apex  acuminate  to  acute  or 
rounded,  base  obtuse  to  acute,  drying  subcoria- 
ceous  and  grayish,  glabrous  above  and  below  or 
with  a  few  hairs  along  the  midvein  beneath,  minor 
venation  flat  or  slightly  elevated  above,  2°  veins 
5-8/side.  Inflorescences  with  1-5  flowers,  ter- 


minal, axillary,  or  borne  on  older  stems,  race- 
mose, peduncles  6-10  mm  long  from  a  short  (1- 
6  mm)  woody  base,  rachis  with  small  (1-2  mm) 
bracts,  pedicels  8-20  mm  long,  ca.  0.8  mm  thick, 
minutely  (0.1-0.2  mm)  puberulent.  Flowers  with 
cupulate  or  campanulate  calyx  9-12  mm  long,  7- 
10  mm  diam.,  margin  entire  or  with  short  (1-2 
mm)  rounded  lobes,  glabrous  or  with  few  minute 
hairs  near  the  base,  purple  or  violet  with  white 
spots  in  life;  corolla  33-42  mm  long,  tubular- 
campanulate  and  bilabiate,  yellowish  white  or 
white,  with  rose  or  purplish  lobes,  tube  3-4  mm 
diam.  at  base  and  widening  to  the  mouth,  lobes 
6-10  mm  long,  with  rounded  apices;  stamens  16- 
23  mm  and  14-18  mm  long,  anthers  divaricate, 
thecae  ca.  3  mm  long,  staminode  3-5  mm  long; 
ovary  unilocular  near  the  apex.  Fruits  4-5  cm 
long,  3-4  cm  diam.,  obovoid  to  globose  with  nar- 
rowed base,  surface  lustrous  and  slightly  muri- 
culate, drying  brown;  seeds  ca.  3  mm  long,  0.8- 
1  mm  wide  in  the  center. 

Uncommon  vines  of  evergreen  rain  forest  for- 
mations along  the  Caribbean  slope,  100-600  m 
elevation  (to  1500  m  in  Nicaragua).  Probably 
fruiting  throughout  the  year  (Costa  Rican  flower- 
ing collections  have  been  made  only  in  October 
and  December-February).  This  species  ranges 
from  Veracruz,  Mexico,  to  Panama. 

Schlegelia  nicaraguensis  is  recognized  by  its 
woody  vining  habit,  subcoriaceous  leaves,  few- 
flowered  inflorescences,  large  bilabiate  corollas, 
and  large  fruits.  This  species  is  similar  to  5.  par- 
asitica  Sw.  of  Jamaica  (Gentry,  1973b). 

Schlegelia  parviflora  (Oerst.)  Monachino,  Phy- 
tologia  3:  103.  1949.  Dermatocalyx  parviflorus 
Oerst.,  Vidensk.  Meddel.  Dansk  Naturhis.  For- 
en.  Kjobenhavn  1855:  29.  1856.  S.  fuscata  A. 
Gentry,  Ann.  Missouri  Bot.  Gard.  60:  925.  1973 
(1974).  Figure  9. 

Epiphytic  shrubs  or  less  often  (in  Costa  Rica) 
woody  vines,  lianas,  or  small  trees,  stems  slightly 
quadrangular  or  terete,  leafy  stems  2-7  mm  thick, 
glabrous  or  sparsely  puberulent  at  the  shoot  apex; 
pseudostipules  2-3  mm  long.  Leaves  with  peti- 
oles 6-30  mm  long,  0.7-3  mm  thick,  becoming 
articulated  at  the  base  and  thickened,  glabrous; 
leaf  blades  (4-)6-18  cm  long,  (2-)3-13  cm  wide, 
broadly  elliptic  to  ovate-elliptic  or  ovate-oblong, 
apex  bluntly  acute  to  obtuse  or  rounded,  base  ob- 
tuse to  cuneate  and  often  slightly  decurrent  on  the 
petiole,  drying  coriaceous  and  yellowish  gray, 
glabrous  above  and  below,  minutely  punctate  be- 


76 


FIELDIANA:  BOTANY 


neath,  2°  veins  4-8/side,  minor  venation  usually 
obscure  above.  Inflorescences  axillary  to  leaves 
or  from  older  leafless  nodes,  fascicles  of  5-40 
flowers  in  racemose  or  paniculate  arrangements, 
2-5  cm  long  (to  8  cm  in  fruit),  1°  peduncles  1- 
10  mm  long,  1-2  mm  thick,  glabrous,  usually  dry- 
ing yellowish,  bracts  1-2  mm  long,  subulate,  ped- 
icels 1-4  mm  long  (measured  from  the  subtending 
bracteoles),  becoming  1.5  mm  thick  in  fruit. 
Flowers  with  calyx  4-6  mm  long,  3-4  mm  wide, 
cupulate  to  campanulate,  glabrous,  lobes  0.5-1.5 
mm  long,  rounded  distally;  corolla  8-1 1  mm 
long,  to  10  mm  wide  across  the  lobes,  white  with 
pink-tinged  lobes,  glabrous  externally,  lobes  2-4 
mm  long,  with  purple  lines  within;  filaments  ca. 
4  mm  long,  thecae  1.3  mm  long,  lilac;  style  ca.  6 
mm  long.  Fruits  8-12  mm  diam.,  globose  or 
somewhat  oblate,  becoming  purple  or  wine-red, 
surface  lustrous  and  slightly  muriculate,  subtend- 
ing calyx  to  10  mm  wide,  pale  green;  seeds  2-3 
mm  long,  0.5-1  mm  thick,  usually  wider  at  one 
end,  drying  black. 

Common  woody  plants  in  wet  evergreen  forest 
formations  on  both  the  Caribbean  and  Pacific 
slopes,  4-2200  m  elevation.  Probably  flowering 
and  fruiting  in  all  months,  but  with  lowland  plants 
usually  flowering  in  December-February  and 
highland  plants  with  peak  flowering  in  June-Au- 
gust. This  species  ranges  from  Belize  to  Brazil. 

Schlegelia  parviflora  is  recognized  by  its  usu- 
ally epiphytic  habit  (also  vines  or  trees),  stiffly 
coriaceous  leaves,  lack  of  pubescence,  fascicles  of 
flowers  on  few-branched  axes,  cupulate  or  cam- 
panulate calyx,  white  corolla  with  pink  and  purple 
coloring,  and  subglobose  purplish  fruits.  The  low- 
er elevation  collections  tend  to  have  larger  leaves, 
but  there  is  great  variation  in  leaf  size,  and  a  cline 
is  not  evident.  The  flowers  and  fruits  of  this  spe- 
cies are  distinctly  smaller  than  those  of  its  sym- 
patric  congeners.  A  very  few  Costa  Rican  collec- 
tions fit  the  description  of  S.  fuscata,  with  smaller 
flowers  drying  black  on  consistently  racemose 
axes.  However,  such  collections  seem  to  be  no 
more  than  extreme  variants  (perhaps  plants  of  ex- 
posed windy  sites)  in  a  species  that  exhibits  a 
broad  range  of  variation  in  leaf  and  inflorescence 
morphology. 


BIGNONIACEAE 

By  William  Burger  and  Alwyn  Gentry  (t) 

REFERENCES — A.  Gentry,  Bignoniaceae  in  Flora 
of  Panama.  Ann.  Missouri  Bot.  Gard.  60:  781- 


9771.  1973  (1974).  A.  Gentry,  Co-evolutionary 
patterns  in  Central  American  Bignoniaceae.  Ann. 
Missouri  Bot.  Gard.  61:  728-759.  1974.  A.  Gen- 
try, Bignoniaceae  of  southern  Central  America; 
distribution  and  ecological  specificity.  Biotropica 
8:  117-131.  1976.  A.  Gentry,  Bignoniaceae,  Part 
I  (Crescentieae  and  Tourrettieae).  Fl.  Neotrop. 
Monogr.  25(1):  1-130.  1980.  A.  Gentry,  Evolu- 
tionary patterns  in  Neotropical  Bignoniaceae. 
Mem.  New  York  Bot.  Gard.  55:  118-129.  1990. 
A.  Gentry,  Bignoniaceae — Part  II  (tribe  Teco- 
meae).  Fl.  Neotrop.  Monogr.  25(11):  1-370.  1992. 
P.  C.  Standley  &  L.  O.  Williams,  Bignoniaceae  in 
Flora  of  Guatemala.  Fieldiana,  Bot.  24(10):  153- 
232.  1974. 

Trees,  shrubs,  or  more  often  lianas  (rarely 
herbs  or  twining  herbs),  often  climbing  with  the 
aid  of  twisting  stems  or  tendrils,  the  tendrils  sim- 
ple or  distally  trifid  (with  sticky  pads  in  Mansoa 
parvifolia),  often  becoming  woody,  stems  terete 
or  angulate,  glabrous  or  pubescent,  often  with  in- 
terpetiolar  ridges  or  gland  fields  at  the  nodes;  stip- 
ules absent,  pseudostipules  present  or  absent. 
Leaves  opposite  or  whorled  (alternate  in  a  few 
genera),  simple,  2-foliolate,  3-foliolate,  pinnately 
compound  or  palmately  compound  (occasionally 
twice  compound  as  in  Jacaranda,  Pleonoioma, 
and  Tourrettia),  petiolate.  the  distal  leaflet  often 
replaced  by  a  tendril  in  climbing  species,  blades 
usually  entire,  sometimes  serrate  (not  lobed),  gla- 
brous to  densely  puberulent,  venation  palmate  or 
pinnate,  domatia  rarely  present.  Inflorescences 
terminal,  axillary,  or  cauliflorus,  cymose,  panic- 
ulate, racemose,  fasciculate,  or  of  solitary  flowers 
in  leaf  axils,  bracts  and  bracteoles  often  present, 
small  and  caducous,  pedicels  usually  well  devel- 
oped. Flowers  bisexual,  nearly  always  large  and 
showy,  calyx  united  and  tubular  to  campanulate, 
usually  with  5  lobes  or  teeth  (rarely  entire,  bila- 
biate, calyptrate,  or  spathe-like);  corolla  united 
and  funnelform  to  campanulate  with  a  basal  tube, 
usually  2-lipped  and  bilaterally  symmetrical  (rare- 
ly radially  symmetric),  the  5  lobes  imbricate  in 
bud  (rarely  valvate)  and  usually  rounded  distally; 
stamens  usually  4  (rarely  5  or  2),  alternate  with 
corolla  lobes,  filaments  usually  of  2  unequal  pairs, 
borne  on  the  lower  half  of  the  tube  and  free,  an- 
thers 2-thecous  (1-thecous),  free,  a  staminode  usu- 
ally present,  an  annular  disc  usually  present;  ova- 
ry superior,  2-locular  with  2  axile  placentas  or 
unilocular  with  2-4  intruded  parietal  placentas 
(rarely  4-locular  as  in  Tourrettia),  style  simple, 
stigma  2-lobed,  rounded.  Fruits  mostly  2-valved 


BURGER:  FLORA  COSTARICENSIS 


77 


capsules  with  septicidal  or  loculicidal  dehiscence 
(fruits  fleshy  or  hard  and  indehiscent  in  a  few  gen- 
era); seeds  mostly  flat  and  winged  in  capsular 
fruits,  often  embedded  in  pulp  or  mucilage  in  in- 
dehiscent  fruits,  lacking  endosperm,  cotyledons 
leaf-like. 

The  Bignoniaceae  are  a  family  of  about  112 
genera  with  over  800  species  in  tropical  and  tem- 
perate areas  of  the  world  but  with  the  greatest 
concentration  of  genera  and  species  (ca.  600)  in 
the  Neotropics.  The  family  is  often  easy  to  iden- 
tify because  of  its  woody  stems,  large  showy  five- 
lobed  and  two-lipped  corollas,  androecium  of  usu- 
ally four  functional  stamens  with  filaments  of  two 
lengths,  superior  ovary  with  many  ovules,  and 
usually  capsular  fruits  with  thin-winged  seeds. 
Gland  fields  are  sometimes  present  at  the  nodes, 
on  the  petioles,  or  at  the  base  of  the  leaf  blade. 
The  stems  are  woody  (except  in  Tourettia)  and 
often  climbing;  many  species  have  opposite  com- 
pound leaves  with  entire  margins.  Gibsoniotham- 
nus  and  Schlegelia,  often  assigned  to  Bignoni- 
aceae, have  now  been  placed  in  the  family  Schle- 
geliaceae  (q.v.).  On  the  basis  of  floral  morphology 
the  Bignoniaceae  were  thought  to  be  closely  re- 
lated to  the  Gesneriaceae,  Pedaliaceae,  and  Scro- 
phulariaceae.  However,  recent  DNA  studies  also 
suggest  relationships  with  Buddleiaceae  and  Ver- 
benaceae  (Olmstead  &  Reeves,  1995). 

This  is  publication  No.  2  in  the  Gentry  Invita- 
tion Series,  based  on  data  and  annotations  left  by 
the  late  Dr.  Alwyn  H.  Gentry,  who  died  in  an 
airplane  crash  in  Ecuador  on  3  August  1993.  The 
series  acknowledges  the  many  contributions  made 
by  Dr.  Gentry  to  our  understanding  of  the  Big- 
noniaceae. A  summary  of  his  life  and  a  complete 
bibliography  were  published  by  James  Miller 
(Ann.  Missouri  Bot.  Card.  83  pt.  4).  For  infor- 
mation on  the  computerized  databases  left  by 
Gentry,  contact  the  Missouri  Botanical  Garden.  It 
has  been  a  priviledge  to  work  on  Costa  Rica's 
Bignoniaceae  using  the  many  annotations  and 
publications  that  Alwyn  Gentry  left  us.  The  Big- 
noniaceae had  been  a  very  difficult  family  on 
which  to  work.  Many  genera  and  species  were 
described  from  material  that  included  only  flowers 
or  only  fruits.  The  resulting  number  of  monotypic 
genera  and  genera  with  disimilar  species  created 
considerable  confusion.  An  additional  problem 
was  that  tropical  trees,  and  especially  lianas,  had 
been  poorly  sampled  (Standley  &  Williams, 
1974).  Al  Gentry  stepped  into  this  situation  and 


attacked  the  problem  with  intelligence  and  vigor, 
both  in  the  field  and  in  herbaria.  He  made  order 
where  there  had  been  considerable  chaos,  and  we 
are  deeply  indebted  to  him  for  this  work.  His  ef- 
forts in  understanding  tropical  forests  and  their 
diversity,  together  with  his  work  on  behalf  of  con- 
servation, inspired  many  of  his  students  and  col- 
leagues. Al  Gentry's  untimely  death  cut  short 
what  was  already  a  rich  and  productive  botanical 
career. 

Thanks  to  Gentry's  work,  most  of  the  species 
of  New  World  Bignoniaceae  are  well  character- 
ized and  their  taxonomy  has  been  put  into  good 
order.  The  percentage  of  Costa  Rican  bignon  spe- 
cies that  range  from  Mexico  far  into  South  Amer- 
ica greatly  exceeds  that  for  most  plant  families  in 
our  flora.  There  appear  to  be  two  reasons  for  this. 
The  first  is  that  we  now  have  sound  biological 
species  concepts  recognizing  considerable  varia- 
tion within  each  binomial,  thanks  to  Al  Gentry's 
extensive  field  work.  The  second  and  more  fun- 
damental reason  is  related  to  the  thin  membra- 
nous-winged seeds,  which  have  proved  to  be  very 
successful  dispersal  agents  in  many  of  the  genera. 
As  Al  Gentry  pointed  out  (1983),  genera  and  spe- 
cies lacking  such  seeds  usually  have  much  more 
limited  geographical  ranges  (cf.  Amphitecna  and 
Parmentiera).  Interestingly,  although  a  few  gen- 
era of  Bignoniaceae  can  tolerate  the  cold  winters 
of  the  northeastern  United  States,  no  indigenous 
member  of  the  family  is  found  above  2300  m  el- 
evation in  Costa  Rica.  This  treatment  has  also 
benefited  from  comparison  with  a  draft  of  the 
family  for  the  Manual  Flora  of  Costa  Rica  project 
by  Quirico  Jimenez  and  J.  F.  Morales.  Our  inde- 
pendent work  on  these  plants  resulted  in  almost 
identical  treatments,  thanks  to  the  earlier  annota- 
tions by  Al  Gentry. 

The  Bignoniaceae  appear  to  be  a  modern  line- 
age in  which  recent  diversification  has  produced 
a  great  number  of  species  differing  in  a  variety  of 
minor  morphological  traits.  Some  of  these  minor 
variations  have  been  the  basis  for  erecting  what 
appeared  to  be  an  excessive  number  of  genera. 
Gentry  (1973a,  1979)  addressed  this  issue  and 
made  important  advances  in  creating  better  ge- 
neric concepts.  Although  further  "lumping" 
would  seem  to  be  desirable,  current  generic  con- 
cepts have  provided  a  useful  information  retrieval 
system  that  should  not  be  radically  altered  until 
the  evidence  for  doing  so  becomes  substantial. 


78 


FIELDIANA:  BOTANY 


Key  to  the  Genera  and  Unusual  Species  of  Bignoniaceae 

la.  Trees  and  shrubs,  stems  woody  and  erect  or  spreading,  tendrils  absent  [native  and  ornamental 

species]    2 

Ib.  Lianas  or  vines,  stems  woody  or  herbaceous,  stems  climbing  or  clambering,  tendrils  often  present 

16 

2a.  Leaves  pinnately  compound,  the  central  rachis  with  >  1  pair  of  lateral  leaflets  or  bipinnately 

compound  3 

2b.  Leaves  simple,  trifoliolate  or  palmately  compound  (with  all  leaflets  arising  from  the  apex  of 

the  petiole)    8 

3a.  Leaves  bipinnate  (twice  compound),  the  central  rachis  with  opposing  lateral  2°  rachises  on 
which  the  leaflets  are  borne;  flowers  lavender  to  bluish  purple,  staminode  larger  than  the 

stamens  and  resembling  a  style  [both  native  and  planted  ornamental  species]    

Jacaranda 

3b.  Leaves  pinnate,  the  central  rachis  bearing  the  usually  opposite  leaflets;  flowers  not  lavender 

or  bluish,  staminode  much  smaller  than  the  stamens,  not  resembling  a  style   4 

4a.  Corollas  3.5-6  cm  long,  yellow  or  orange  to  red-orange;  native  and  introduced  trees  .  .  5 
4b.  Corollas  6--12  cm  long,  yellow,  red  or  dark  purple  to  maroon;  introduced  ornamental  trees 

6 

5a.  Stamens  included;  flowers  usually  yellow;  leaflets  2-15  cm  long;  native  small  trees 

often  planted  for  ornament Tecoma  stans 

5b.  Stamens  exserted;  flowers  usually  orange-red;  leaflets  1-3.5  cm  long;  introduced  or- 
namental shrubs  with  clambering  branches    Tecoma  capensis 

6a.  Corollas  8-12  cm  long,  red  or  red-orange,  curved  and  opening  upward,  glabrous  externally; 
calyx  split  down  1  side  and  spathe-like;  commonly  planted  in  lower  elevation  evergreen 

areas Spathodea  campanulata 

6b.  Corollas  6-9  cm  long,  yellow  or  dark  purple  to  maroon,  not  usually  opening  upward, 
glabrous  or  puberulent  externally;  calyx  not  split  down  1  side  (not  spathe-like);  rarely 

planted  in  Central  America   7 

7a.  Corolla  dark  purple  or  maroon,  glabrous  externally;  leaves  7-9rfoliolate;  fruits  pendulous, 

ellipsoid-oblong   Kigelia  pinnata 

7b.  Corolla  yellow,  puberulent  externally;  leaves  5-7-foliolate;  fruits  not  ellipsoid  [plants  not 

recorded  from  Costa  Rica  and  not  included  in  text] Haplophragma  adenophyllum 

8a.  (from  2b)  Epiphytic  shrubs;  leaves  simple,  opposite,  usually  coriaceous;  corollas  radially  sym- 
metric or  only  slightly  2-lipped  (note  that  these  plants  have  now  been  transferred  to  Schlege- 

liaceae)    9 

8b.  Terrestrial  trees  (not  epiphytic);  corollas  usually  clearly  bilaterally  symmetric  and  2-lipped  [nei- 
ther red  nor  narrowly  tubular]    10 

9a.  Corollas  usually  red  or  purple,  narrowly  tubular  and  radially  symmetric;  fruit  with  fleshy 

covering;  leaves  usually  <  10  cm  long Gibsoniothamnus 

9b.  Flowers  white  or  pinkish,  tubular-campanulate;  fruit  with  hard  outer  covering;  leaves  to  18 

cm  long    •  Schlegelia 

lOa.  Leaves  opposite,  usually  2-9-foliolate;  corolla  thin  and  without  a  transverse  fold  on  the  lower 
(abaxial)  side,  white  to  yellow,  rose,  or  magenta;  flowers  usually  borne  in  racemose  panicles 

on  distal  stems;  fruits  dehiscent  capsules  with  thin  flat  overlapping  seeds 11 

lOb.  Leaves  fasciculate,  alternate  or  opposite,  simple  or  3-foliolate;  corolla  thick  and  stiff  with  a 
transverse  fold  on  the  lower  side,  white  to  greenish  white;  flowers  usually  1  to  few  and  borne 
on  thicker  branches  or  trunk;  fruits  indehiscent  with  fibrous-fleshy  or  hard  outer  shells,  seed 

angular  and  embedded  in  white  fleshy  pulp    14 

1  la.  Flowers  <  18  mm  long;  anthers  pilose;  fruits  spirally  twisted;  leaflets  7-9  [often  cuneate 

at  base] Godmannia 

lib.  Flowers  >  20  mm  long;  anthers  glabrous;  fruits  usually  straight;  leaflets  (l-)2-7  .  .   12 

12a.  Corolla  deeply  split  down  the  sides  and  strongly  2-lipped;  rarely  collected  small  trees  of 

the  Caribbean  slope  Tynanthus  macranthus 

BURGER:  FLORA  COSTARICENSIS  79 


12h.  Corolla  not  deeply  split  down  the  side,  slightly  2-lipped;  rare  and  common  species  of 

many  habitats 13 

13a.  Fruits  usually  held  erect  and  6-18  cm  long;  inflorescences  with  <  11  purple  flowers; 
leaves  simple  and  3-foliolate  on  the  same  branches;  small  trees  and  shrubs  of  seasonally 

very  dry  Guanacaste    Arrabidaea  costaricensis 

13b.  Fruits  usually  pendant  and  >  15  cm  long;  inflorescences  usually  with  >  10  pink  flowers 
or  the  corollas  yellow  or  white;  leaves  rarely  simple,  3-7-foliolate;  small  to  large  trees 

found  in  many  habitats  and  often  planted  as  ornamentals   Tabebuia 

14a.  (from  lOb)  Leaves  opposite;  3-foliolate  (often  cruciform);  calyx  split  on  1  side;  fruits  with 

fibrous  exocarp  (not  a  hard  shell)    Parmentiera 

14b.  Leaves  alternate  or  fasciculate;  simple  or  3-foliolate;  calyx  not  split  on  1  side;  fruits  with  a 

hard  exocarp  15 

15a.  Leaves  alternate;  ovules  on  2  parietal  placentas  or  axile  near  base;  seeds  large  (>  13  mm) 

Amphitecna 

15b.  Leaves  fasciculate;  ovules  on  4  parietal  placentas;  seeds  small  (<  8  mm)    Crescentia 

16a.  (from  Ib)  Leaves  simple;  tendrils  absent;  fruits  spherical  and  indehiscent,  seeds  angular  and  wing- 
less; stems  lacking  4-8  phloem  areas  in  cross-section;  with  1  axile  placenta  in  each  locule  (now 

transferred  to  Schlegeliaceae)    Schlegelia 

16b.  Leaves  usually  compound  (simple  leaves  sometimes  present  on  young  growth  or  the  base  of  new 

shoots);  tendrils  often  present;  fruits  usually  elongate  or  flattened,  dehiscing  to  release  flat,  usually 

winged  seeds;  stems  with  4-8  phloem  areas;  with  usually  2  axile  placentas  in  each  locule    .  .   17 

17a.  Leaflets  3-15  mm  long  on  young  climbing  stems,  becoming  30  mm  long  on  distal  stems  [4  leaflets/ 

node];  tendrils  ending  with  small  disc-like  adhesive  pads Mansoa  parvifolia 

17b.  Leaflets  not  so  small  on  young  climbing  stems,  becoming  more  than  3  cm  long  on  distal  stems; 

tendrils  not  ending  in  disc-like  pads 18 

18a.  Some  leaves  twice  compound,  some  petioles  bearing  2  or  3  petiolules  with  3  or  5  leaflets  each 

19 

1 8b.  Leaves  never  twice  compound,  simple  or  with  petioles  bearing  2  or  3  leaflets  [plants  with  woody 

stems;  inflorescences  never  with  sterile  and  fertile  dimorphic  flowers] 21 

19a.  Herbaceous  vines;  inflorescence  subspicate  with  2  different  kinds  of  flowers  (distal  flowers 
sterile);  corolla  greenish  to  purple;  fruits  ellipsoid  and  covered  with  prominent  hooked  spines 

Tourrettia  lappacea 

19b.  Woody  vines  and  lianas;  inflorescence  racemose  or  paniculate,  with  1  kind  of  flower;  fruits 

linear,  without  spines    20 

20a.  Corollas  white  or  yellowish  white,  glabrous  externally;  tendrils  tri-fid  at  the  apex 

Pleonotoma  variabilis 

20b.  Corollas  magenta  or  purple,  puberulent  externally;  tendrils  simple  [plants  not  known  from 

between  Belize  and  Colombia  and  not  included  in  the  text] Arrabidaea  inaequalis 

2 la.  Each  tendril  terminating  with  usually  3  sharp  stiff  claw-like  tips  or  hooks  [corolla  glabrous  exter- 
nally]     22 

21b.  Each  tendril  terminating  with  1-3  slender  tips,  not  hard  and  claw-like  or  hooked 24 

22a.  Corolla  deep  purple  or  magenta;  calyx  4-  or  5-lobed;  Golfo  Dulce  area  in  Central  America 

[capsules  usually  linear]    Parabignonia  steyermarkii 

22b.  Corolla  yellow;  calyx  subtruncate  to  spathe-like;  widely  ranging  in  Central  America  ...  23 
23a.  Capsules  linear,  1.5-2  cm  wide,  >  15  cm  long,  valves  not  splitting  in  half  (2/fruit);  thin 

flexible    Macfadyena 

23b.  Capsules  oblong,  2.5-4  cm  wide,  to  15  cm  long,  valves  splitting  longitudinally  in  half  at 

maturity  (4/fruit),  thick  woody Melloa  quadrivalvis 

24a.  Corolla  orange,  long-tubular  and  slightly  curved  (6-9  cm  long,  3-12  mm  diam.),  lobes  valvate  in 

bud;  planted  for  ornament  in  parks  and  gardens    Pyroslegia  venusta 

24b.  Corolla  not  orange,  without  a  long  narrow  slightly  curved  tube,  lobes  imbricate  in  bud;  planted 

for  ornament  and/or  native  wild  species 25 

25a.  Branchlets  6-sided,  hexagonal  in  cross-section,  with  6  prominent  longitudinal  ribs  along  the  young 
stems  [fruits  ellipsoid  to  oblong,  2.5-7  cm  wide,  surface  smooth  to  tuberculate]    26 

80  FIELDIANA:  BOTANY 


25b.  Branchlets  2-  or  4-sided  to  terete  or  subterete  in  cross-section,  with  0-4  longitudinal  ribs  along 

the  young  stems 28 

26a.  Calyx  double,  with  an  inner  and  outer  whorl  of  distal  lobes;  leaves  with  dendroid  (stellate 
or  branched)  hairs;  tendrils  usually  with  3  distal  tips;  corolla  strongly  2-lipped  with  the  upper 

and  lower  lobes  connate  at  anthesis Amphilophium 

26b.  Calyx  simple,  with  1  distal  whorl  of  lobes;  leaves  with  simple  hairs;  tendrils  with  3-15  distal 

divisions;  corolla  weakly  2-lipped,  lobes  free  and  reflexed  at  anthesis    27 

27a.  Corolla  white,  tube  strongly  curved  near  the  base;  fruit  oblong  with  a  covering  of  short 

spines;  widely  ranging    Pithecoctenium  crucigerum 

27b.  Corolla  rose-purple,  tube  not  curved;  fruits  without  spines;  not  known  from  between  central 

Nicaragua  and  Colombia  and  not  included  in  the  descriptions Distictis 

28a.  Leaves  with  minute  (0. 1  mm)  pellucid  flat  glands  on  both  surfaces;  branchlets  with  a  hollow  center; 
capsules  <  9  mm  wide,  linear  and  24-45  cm  long,  surface  minutely  puberulent  [corolla  puberulent 

externally,  white  or  greenish  yellow  with  purple  lobes) Stizophyllum 

28b.  Leaf  surfaces  without  pellucid-punctate  glands;  branchlets  without  a  hollow  center  (sometimes 
hollow  in  Paragonia);  capsule  >  10  mm  wide,  linear  to  oblong  or  rounded,  surfaces  glabrous  to 

puberulent,  smooth  to  spiny  or  muricate    29 

29a.  Leaves  with  branched  or  irregular  dendroid  hairs  (at  least  in  the  vein  axils  beneath);  capsules 
oblong  (3-10  cm  wide,  ca.  5-10  mm  thick),  valves  woody  and  flattened,  surface  smooth  [seeds 

thin  with  broad  lateral  wings] 30 

29b.  Leaves  with  slender  simple  or  flat  rounded-peltate  hairs  or  glabrous;  capsules  without  the  above 

combination  of  characters   31 

30a.  Interpetiolar  glandular  fields  absent  at  nodes;  calyx  >  20  mm  long;  corolla  yellow;  seeds  6- 

12  cm  wide,  wings  brown  and  opaque Callichlamys  latifolia 

30b.  Interpetiolar  gland  fields  present  at  the  nodes;  calyx  <  9  mm  long;  corolla  lavender  or  rose; 

seeds  3-5  cm  wide,  wings  translucent Xylophragma  seemannianum 

3 la.  Corolla  15-20  cm  long  with  long  narrow  tube  and  distal  rotate  lobes  (tubular-slaverform).  white 
and  puberulent  externally;  fruits  oblong-cylindric,  9-22  cm  long,  6-1 1  cm  wide  and  5-8  cm  thick 

Tanaecium  jaroba 

31b.  Corolla  <  15  cm  long  (mostly  tubular-campanulate)  and  without  the  combination  of  characters 

listed  above   32 

32a.  Lower  surface  of  leaf  with  gland  fields  in  axils  of  basal  veins  or  with  gland  field  at  apex  of  the 

petiole  [corolla  pink  to  magenta  or  white]   33 

32b.  Lower  surface  of  leaf  lacking  gland  fields,  apex  of  petiole  lacking  a  gland  field 35 

33a.  Leaf  axils  with  a  conical  structure  made  up  of  3  imbricate  series  of  scale-like  pseudostipules 
or  vegetative  parts  with  the  strong  odor  of  onion  or  garlic  [fruits  with  verrucose  or  smooth 

surface]    Mansoa 

33b.  Leaf  axils  lacking  a  conical  structure  of  imbricate  scale-like  pseudostipules  and  vegetative 

parts  lacking  an  onion-like  odor 34 

34a.  Native  wild  plants;  young  stems  lacking  conspicuous  pseudostipules  (rounded  leaf-like  pseu- 
dostipules present  in  C.  diversifolia);  corolla  pink  or  lavender  to  purple,  glabrous  or  puber- 
ulent externally;  fruits  oblong  or  linear Cydista 

34b.  Vines  planted  for  ornament  (rarely  collected  in  Costa  Rica);  young  stems  with  broadly  ovate 

leaf-like  pseudostipules;  corolla  magenta  to  violet,  glabrous  externally;  fruits  linear    

Saritea  magnified 

35a.  Corolla  strongly  bilabiate,  split  down  the  sides  beyond  the  middle,  usually  white;  rarely  collected 

Tynanthus 

35b.  Corolla  not  strongly  bilabiate,  not  split  down  the  sides  to  the  middle,  colors  various;  common  to 

rare  species    36 

36a.  Flowers  yellow  or  white  (sometimes  with  reddish  markings  in  Mussatia);  disc  present  beneath  the 

ovary;  valves  of  the  fruits  usually  woody,  >  25  mm  wide,  never  cchinate    37 

36b.  Flowers  rose  to  magenta  or  lavender  (white  in  species  of  Arrabidaea,  Cydista,  and  Lundia);  disc 
present  or  absent;  valves  of  the  fruits  not  woody  unless  echinate  or  tuberculate,  usually  <  25  mm 
wide  -41 

BURGER:  FLORA  COSTARICENSIS  81 


37a.  Corolla  <  20  mm  long,  calyx  <  2  mm  long,  widely  campanulate;  young  stems  strongly 
tetragonal  in  cross-section,  with  4  prominent  longitudinal  ridges Mussatia 

37h.  Corolla  >  25  mm  long,  calyx  >  4  mm  long,  cupular  to  tubular-campanulate;  young  stems 
terete  or  weakly  tetragonal  in  cross-section,  without  4  prominent  ridges 38 

38a.  Young  stems  with  interpetiolar  gland  fields  at  the  nodes;  fruits  tetragonal  (square)  in  cross- 
section,  ca.  2  cm  thick,  narrowly  oblong  [corolla  glabrous  externally]  

Ceratophytum  tetragonolobum 

38b.  Young  stems  lacking  interpetiolar  gland  fields;  fruits  flattened  and  broadly  oblong  to  elliptic 
or  suborbicular  39 

39a.  Corolla  glabrous  externally  or  with  minute  flat  peltate  hairs;  capsules  narrowed  at  the  base 
and  somewhat  stipitate,  flattened  and  elliptic  to  oblong  or  suborbicular  .  .  .  Anemopaegma 

39b.  Corolla  minutely  puberulent  externally;  capsules  not  stipitate,  flattened  and  elliptic  to  oblong 
40 

40a.  Inflorescences  10-20  cm  long;  calyx  entire;  tendrils  trifid  at  the  apex 

Distictella  magnoliifolia 

40b.  Inflorescence  2-6  cm  long;  calyx  usually  split  1-2  mm  along  2  sides;  tendrils  simple 

Adenocalymma  inundatum 

4 la.  (from  36b)  Pseudostipules  (cataphylls)  forming  small  bromeliad-like  clusters  in  distal  leaf  axils 

(especially  below  the  inflorescence);  fruits  ellipsoid  to  suborbicular  and  echinate,  5-9  cm  long 

with  spines  to  8  mm  long  [disc  absent,  venation  pinnate] Clytostoma  binatum 

41b.  Pseudostipules  not  forming  small  bromeliad-like  clusters  in  leaf  axils;  fruits  mostly  linear  and 

smooth  to  verrucose  42 

42a.  Calyx  split  down  1  side  and  spathe-like,  apex  thickened  and  curved;  fruit  surface  with  minute 

hairs  that  often  give  a  grayish  metallic  luster  [disc  absent;  fruits  linear-oblong  or  ellipsoid;  gland 

fields  absent  at  the  nodes] Phyrganocydia 

42b.  Calyx  not  split  down  1  side,  not  spathe-like  or  with  curved  thickened  tip;  fruit  surface  not  grayish 

lustrous 43 

43a.  Anthers  pubescent;  flower  buds  with  narrow  conical  tip,  the  apex  splitting  off  to  produce  an  entire 

margin  on  the  calyx  cup  at  anthesis  (also  splitting  as  growth  progresses)  [corollas  white  to  rose 

or  magenta,  minutely  puberulent  externally;  disc  absent;  fruits  20-60  cm  long,  1.5-2  cm  wide 

margin  and  midvein  prominent,  surface  densely  puberulent] Lundia 

43b.  Anthers  glabrous;  flower  buds  and  calyx  not  calyptrate  (sometimes  coming  off  as  a  conical  cap  in 

Paragonia} 44 

44a.  Tendrils  ending  in  3  short  tips;  corolla  dark  purple  to  deep  maroon,  glabrous  or  subglabrous 

externally;  fruits  (30-)50-110  cm  long,  linear,  surfaces  flat  and  glabrous  [disc  present] 

Martinella  obovata 

44b.  Tendrils  simple  or  bifid  at  apex;  corolla  whitish  or  rose  to  purple,  usually  minutely  puberulent 

externally;  fruits  <  60  cm  long,  surface  with  raised  edges  or  raised  midrib,  glabrous  or  puberulent 

45 

45a.  Calyx  >  15  mm  long,  2-lipped  or  lobed;  fruits  becoming  black,  with  tuberculate  or  verrucose 

surface  [disc  present] Arrabidaea  verrucosa 

45b.  Calyx  >  10  mm  long,  usually  truncate  at  the  apex;  fruits  not  becoming  dark  with  tuberculate  or 

verrucose  surface 46 

46a.  Disc  absent  beneath  the  ovary;  young  stems  terete  or  tetragonal,  lacking  interpetiolar  gland  fields; 

stems  with  8-16  phloem  areas  in  cross-section  Cydista 

46b.  Disc  present  beneath  the  ovary;  young  stems  terete,  often  with  interpetiolar  gland  fields;  stems 

with  4  phloem  areas  in  cross-section  47 

47a.  Capsule  valves  flat  and  smooth  (except  in  A.  verrucosa),  seeds  usually  with  translucent  wings 

differentiated  from  the  brown  center;  tendrils  simple  at  the  apex;  interpetiolar  gland  fields  often 

present;  leaflets  lacking  a  faint  sweet  odor  when  crushed;  deciduous  to  evergreen  wet  forests  .  .  . 

Arrabidaea 

47b.  Capsule  valves  convex  and  slightly  rough-surfaced,  seeds  uniformly  brownish;  tendrils  simple  or 

minutely  bifid  at  the  apex;  interpetiolar  gland  fields  rarely  present;  leaflets  with  faint  sweet  odor 

when  crushed;  evergreen  lowland  rain  forests    Paragonia  pyramidata 

82  FIELDIANA:  BOTANY 


Amphitecna 
isthmica 


A.   sessilifolia 

FIG.  10.     Bignoniaccae:  trees  with  simple  leaves,  fruit  a  calabash  or  pepo;  species  of  Amphitecna. 


BURGER:  FLORA  COSTARICENSIS 


83 


Parmentiera  macrophylla, 


FIG.  1 1.     Bignoniaccac:  trees  with  simple  or  compound  leaves,  fruit  a  calabash  or  pepo:  species  of  Crescentia  and 
Parmentiera. 


84 


FIELDIANA:  BOTANY 


'A  Tabebuia 

palustris 


Tynnanthus 
macranthus 


Godmania 
aesculi folia 


Arrabidea  costaricensis 

FIG.  12.     Bignoniaccac:  trees  and  shrubs  with  2-  to  7-foliolatc  leaves,  fruits  long  narrow  capsules. 


BURGER:  FLORA  COSTARICENSIS 


85 


Tabebuia 
chrysantha 


Tabebuia  impetiginosa 

FIG.  13.     Bignoniaceac:  trees  with  palmately  compound  leaves  and  showy  flowers;  species  of  Tabebuia. 


86 


FIELDIANA:  BOTANY 


Jacaranda 
i  mimosifolia 


isK  Jacaranda  copaia 


FIG.  14.     Bignoniaceae:  trees  with  bipinnately  compound  leaves  and  lavender  or  bluish  flowers;  species  of  Jaca- 
randa. 


BURGER:  FLORA  COSTARICENSIS 


87 


FIG.  15.     Bignoniaccac:  ornamental  trees  and  shrubs  and  an  ornamental  vine. 


88 


FIELDIANA:  BOTANY 


Mansoa 
parvifolia  "713 


Pleonotoma    l^variabilis 


Tanaecium  jaroba 


FIG.  16.     Bignoniaccac:  unusual  vines  with  twice -compound  leaves,  with  very  small  leaves,  or  with  a  very  long 
narrow  corolla  tube. 


BURGER:  FLORA  COSTARICENSIS 


89 


Parabignonia 
steyermarkl i 


Melloa  quadrivalvis 

FIG.  17.     Bignoniaccac:  vines  climbing  with  tendrils  ending  in  3  sharp  hard  claws. 


90 


FIELDIANA:  BOTANY 


&&OT  Pi  thecoctem  um 
crucigerum 


FIG.  18.     Bignoniaccac:  vines  with  cchinatc  or  prominently  tubcrculatc  fruits. 


BURGER:  FLORA  COSTARICENSIS 


91 


Ceratophytum 
tetragonolobum 


Cydlsta  potosina 


Callichlamys 
1  at i folia 


hyacinthina  f 

FIG.  19.     Bignoniaccae:  vines  with  broad  or  thick  elongate  fruits. 


92 


FIELDIANA:  BOTANY 


Anemopaegma 
chrysoleucum 


A.  orbiculatum 

FIG.  20.     Bignoniaccac:  vines  with  ellipsoid  or  rounded  fruits;  species  of  Anemopae^ma. 


BURGER:  FLORA  COSTARICENSIS 


93 


Adenocalymma 
inundatum 


Xylophragma 
seemanniana 


Amphilophium 
paniculatum 


magnoTri  fol  i  a 


FIG.  21.     Bignoniaceae:  vines  with  flattened  ellipsoid  or  oblong  fruits. 


94 


FIELDIANA:  BOTANY 


Phryganocydia  corymbosa 


Mansoa  standleyi   30 
cm 


FIG.  22.     Bignoniaccae:  vines  with  flattened  elongate  fruits  more  than  2  cm  wide. 


BURGER:  FLORA  COSTARICENSIS 


95 


Paragonla  ^pyrami  data  <^     Stizophyllum  riparium 

Fto.  23.     Bignoniaceae:  vines  with  very  long  linear  flattened  fruits. 


96 


FIELDIANA:  BOTANY 


Cydista      ^ttaequinoctialis 


Cydista  diversi folia 


Cydista  heterophy  1 1 a~*y 


lilacina 


FIG.  24.     Bignoniaccae:  vines  with  elongate  narrow  fruits;  species  of  Cydista. 


BURGER:  FLORA  COSTARICENSIS 


97 


Arrabidaea  candicans 


A.  chica  ^<^|         /^^^^m  \'^  K.  conjugata 

FIG.  25.     Bignoniaceac:  vines  with  elongate  flattened  narrow  fruits;  species  of  Arrabidea. 


98 


FIELDIANA:  BOTANY 


Arrabldaea  patellifera    Stizophyllum 

inaequilaterum 


Arrabidaea  mollisslma 

FIG.  26.     Bignoniaceae:  vines  with  elongate  narrow  fruits  and  usually  puberulcnt  leaves 


Amphilophium 
pannosum 


BURGER:  FLORA  COSTARICENSIS 


99 


Adenocalymma  Martius  ex  Meissner 

Lianas,  climbing  with  tendrils,  stem  terete, 
with  4  phloem  areas  in  cross-section,  glandular 
fields  absent  at  the  nodes;  pseudostipules  small, 
subulate.  Leaves  opposite,  3-foliolate  or  2-folio- 
late  with  terminal  tendril,  petiolate,  leaflets  with 
petiolules,  margins  entire,  venation  pinnate.  Inflo- 
rescences terminal  or  axillary,  narrow  racemes, 
flower  buds  subtended  by  caducous  bracts.  Flow- 
ers with  cupular  calyx,  5-lobed  and  2-lipped  or 
truncated,  usually  with  peltate  glands  near  the 
rim;  corolla  tubular-funnelform  to  tubular-cam- 
panulate,  yellow,  usually  puberulent  on  the  exte- 
rior; stamens  4,  filaments  of  2  lengths,  included, 
anthers  glabrous,  thecae  straight  and  divaricate  to 
slightly  divergent,  a  staminode  present;  disc  pul- 
vinate,  ovary  puberulent  or  with  peltate  hairs,  2- 
locular,  ovules  in  2  series  on  each  placenta.  Fruits 
oblong  woody  capsules,  valves  parallel  to  the  sep- 
tum, rounded  or  slightly  compressed,  median  vein 
not  elevated;  seeds  almost  wingless  or  with  2  lat- 
eral wings  poorly  differentiated  from  the  central 
body,  brownish  or  translucent  distally. 

Adenocalymma  is  a  Neotropical  genus  of  about 
36  species  ranging  from  Mexico  to  Argentina, 
with  most  of  the  species  in  eastern  Brazil.  One 
species  is  endemic  to  mangroves  in  eastern  Mex- 
ico, two  species  are  found  in  eastern  Panama- 
Choco,  and  the  following  wide-ranging  species 
occurs  in  Central  America. 

Adenocalymma  inundatum  Mart,  ex  DC.,  Prodi. 
9:  201.  1845.  Tabebuia  calderonii  Standl.,  J. 
Washington  Acad.  Sci.  14:  244.  1924.  A.  hin- 
tonii  Sandw.  Kew  Bull.  1936:  10.  1936.  A. 
calderonii  (Standl.)  Seibert,  Carnegie  Inst. 
Washington  Publ.  522:  428.  1940.  Figure  21. 

Lianas,  stems  1-15  cm  diam.,  tendrils  to  20  cm 
long,  leafy  stems  1 .7-8  mm  diam.,  essentially  gla- 
brous, terete,  drying  dark  with  whitish  lenticels 
but  becoming  grayish  in  age,  interpetiolar  line  of- 
ten present;  pseudostipules  2-3  mm  long.  Leaves 
with  petioles  2-8  cm  long,  0.8-2  mm  diam.,  gla- 
brous or  minutely  puberulent  at  the  apex,  petio- 
lules of  lateral  leaflets  7-27  mm  long,  often  thick- 
ened below  the  blade;  leaflet  blades  3.5-16  cm 
long,  2.5-7  cm  wide,  ovate  to  broadly  elliptic- 
ovate  or  elliptic-oblong,  apex  acuminate,  tips  to 
1 5  mm  long,  base  obtuse  or  slightly  rounded  and 
truncated,  drying  stiffly  chartaceous  and  lustrous 
above,  glabrous  except  for  few  minute  (0.05-0.1 
mm)  peltate  hairs  beneath,  2°  veins  5-8/side,  3° 


veins  often  elevated  on  the  dried  upper  surface. 
Inflorescences  2-8  cm  long,  peduncles  6-22  mm 
long,  minutely  papillate-puberulent,  bracts  6-12 
mm  long,  4-7  mm  wide,  early  caducous,  pedicels 
2-7  mm  long.  Flowers  with  calyx  4-8  mm  long, 
4-6  mm  diam.,  minutely  papillate-puberulent, 
lobes  1-2  mm  or  margin  irregularly  split;  corolla 
3.5-6.5  cm  long,  funnelform-campanulate,  yel- 
low, tube  5-18  mm  diam.,  minutely  puberulent 
with  crooked  hairs  externally,  lobes  10-20  mm 
long,  narrowed  below  and  rounded  distally,  be- 
coming reflexed;  filaments  15-20  and  10-16  mm 
long,  thecae  2-3  mm  long.  Fruits  9-18(-27)  cm 
long,  2-3.5  cm  wide,  13-22  mm  thick,  oblong, 
rounded  at  both  ends,  valves  convex  or  flattened, 
pale  grayish  and  rugulose-lenticellate;  seeds  16- 
19(-21)  mm  long,  44-64(-76)  mm  wide,  central 
area  17-20  mm  wide,  lateral  wings  membrana- 
ceous. 

Evergreen  lianas  in  deciduous  and  partly  decid- 
uous forests  of  the  Pacific  lowlands  (in  Costa 
Rica)  and  often  found  in  riverine  gallery  forests, 
2-400  m  elevation  (to  900  m  in  Chiapas).  Flow- 
ering primarily  March-August  in  Central  Ameri- 
ca, fruiting  December-February.  This  species 
ranges  from  Mexico  to  Brazil  and  occurs  on  Gre- 
nada. 

Adenocalymma  inundatum  is  recognized  by  its 
woody  climbing  habit  and  simple  tendrils,  oppo- 
site two-  or  three-foliolate  leaves,  short  inflores- 
ences,  papillate-puberulent  yellow  corollas,  nar- 
rowly oblong  woody  fruit,  and  thick-bodied, 
winged  seeds.  The  generally  glabrous  parts  and 
large  floral  bracts  that  leave  conspicuous  scars  are 
also  distinctive  characteristics.  The  floral  bracts 
have  the  same  texture  as  the  calyx  but  are  present 
for  only  a  short  time.  Gentry  used  the  name  A. 
apurense  (Kunth  in  H.B.K.)  Sandw.  for  this  ma- 
terial but  later  decided  that  A.  apurense  was  re- 
stricted to  South  America. 


Amphilophium  Kunth 

Lianas,  climbing  with  trifid  tendrils,  stems  te- 
rete, with  4  (5)  phloem  areas  in  cross-section, 
nodes  with  interpetiolar  ridges,  lacking  gland 
fields;  pseudostipules  leaf-like,  early  deciduous. 
Leaves  opposite,  3-foliolate  or  2-foliolate  and  of- 
ten with  a  tendril,  petiolate;  leaflets  with  entire 
margins  and  pinnate  venation.  Inflorescences  ter- 
minal, often  on  short  lateral  branches,  racemose 
panicles  with  short  lateral  branches,  bracts  sub- 
tending the  pedicels.  Flowers  with  a  cupulate- 


100 


FIELDIANA:  BOTANY 


campanulate  calyx  with  thick  inner  2-3-lobed  ca- 
lyx and- thinner  5-lobed  spreading  exterior;  corol- 
la tubular  and  bilabiate,  thick,  whitish  at  first  but 
becoming  purple  at  anthesis,  glabrous  or  puberu- 
lent  externally,  upper  lip  with  2  united  lobes,  low- 
er lip  with  3  united  lobes,  lobes  triangular;  sta- 
mens 4,  filaments  of  2  lengths,  anthers  glabrous, 
thecae  thick  and  divaricate,  staminode  present; 
disc  annular;  ovary  2-locular,  ovules  multiseriate 
in  each  locule.  Fruits  woody  capsules,  valves 
somewhat  compressed,  parallel  to  the  septum, 
smooth  to  tuberculate;  seeds  thin,  with  2  lateral 


membranous  wings  not  clearly  differentiated  from 
the  body  of  the  seed. 

Amphilophium  is  a  genus  of  about  eight  South 
American  species,  with  two  wider  ranging  species 
present  in  Costa  Rica.  Unusual  features  are  a  thick 
two-layered  calyx  and  the  stiff,  closed  corolla, 
split  almost  halfway  into  two  lobes  that  must  be 
forced  open  by  pollinators.  The  rough-surfaced 
fruits  and  corollas  that  are  white  but  turn  purple 
at  anthesis  are  also  unusual.  Distictis  laxiflora 
(DC.)  Greeenm.  of  Mexico  and  central  Nicaragua 
also  has  six-angled  stems,  but  that  species  has 
larger  corollas  and  smooth  elliptic  fruits. 


Key  to  the  Species  of  Amphilophium 

la.  Fruit  surface  muriculate;  pseudostipules  3-10  mm  long  and  caducous;  stems  and  petioles  sparsely 

to  densely  puberulent  with  simple  to  dendroid  hairs  0.05-0.4  mm  long  and  usually  drying  grayish 

to  yellowish  gray;  common  plants  in  deciduous,  partly  deciduous,  and  evergreen  forest  formations 

A.  paniculatum 

Ib.  Fruit  surface  verrucose;  pseudostipules  5-18  mm  long  and  often  persisting;  stems  and  petioles 
villous  with  simple  or  branched  hairs  1-3  mm  long  and  drying  yellowish  to  yellowish  brown; 
uncommon  plants  of  evergreen  rain  forest  formations A.  pannosum 


Amphilophium  paniculatum  (L.)  Kunth  in 
H.B.K.,  Nov.  Gen.  Sp.  3:  116.  1819.  Bignonia 
paniculata  L.,  Sp.  PI.  623.  1753.  A.  molle 
(Schldl.  &  Cham.)  Linnaea  5:  120.  1830.  A. 
paniculatum  var.  molle  (Schldl.  &  Cham.) 
Standl.,  Publ.  Field  Mus.  Nat.  Hist.  Bot.  Ser.  18: 
1114.  1938.  Figure  21. 

Lianas  to  ca.  15m  high,  stems  to  10  cm  diam., 
tendrils  to  17  cm  long,  leafy  stems  1.8-5  mm 
diam.,  with  6  prominent  longitudinal  ridges  (hex- 
agonal in  cross-section),  sparsely  to  densely  pu- 
berulent with  short  (0.1-0.4  mm)  scurfy-stellate 
and  minute  (0.05  mm)  peltate  hairs;  pseudosti- 
pules 3-10  mm  long,  usually  rounded,  caducous. 
Leaves  usually  2-foliolate,  petioles  3-7  cm  long, 
puberulent  in  longitudinal  lines,  petiolules  1-4.5 
cm  long;  leaflet  blades  3-16  cm  long,  2-11  cm 
wide,  broadly  ovate  to  ovate-oblong  or  ovate-el- 
liptic, apex  acuminate  to  caudate-acuminate,  base 
rounded  and  cordate  to  subcordate  or  truncate,  of- 
ten minutely  scurfy  puberulent  beneath,  larger 
(0.2  mm)  peltate  rounded  hairs  beneath  near  the 
base,  venation  subpalmate,  2°  veins  5-7/side,  dis- 
tal veins  strongly  ascending.  Inflorescences  ter- 
minal, often  on  short  lateral  shoots  and  apparently 
axillary,  7-30  cm  long,  racemiform  panicles  with 
short  lateral  branches,  1°  peduncle  ca.  3  cm  long. 


puberulent  like  the  stems,  bracts  caducous,  pedi- 
cels 1-9  mm  long.  Flowers  with  calyx  7-12  mm 
long,  5-7  mm  diam.,  cupulate.  minutely  puberu- 
lent, with  usually  reflexed  outer  lobes  3-4  mm 
long,  inner  calyx  2-  or  3-lobed;  corolla  20-35 
mm  long,  5-14  mm  diam.,  white  in  early  stages 
and  becoming  purple  at  anthesis,  glabrous  or  with 
few  hairs  externally,  lobes  11-20  mm  long,  nar- 
rowly triangular  and  not  opening;  filaments  ca.  17 
and  14  mm  long,  thecae  ca.  2  mm  long;  ovary 
puberulent,  style  ca.  24  mm  long.  Fruits  5-15  cm 
long,  2.8-5  cm  wide,  13-30  mm  thick,  oblong  or 
oblong-ellipsoid,  surface  muriculate  and  with  mi- 
nute peltate  hairs;  seeds  11-15  mm  long,  27-42 
mm  wide,  central  area  ca.  10  mm  wide,  brownish 
or  transparent  distally. 

Common  and  widespread  plants  of  both  sea- 
sonally deciduous  forests  and  evergreen  rain  for- 
est formations,  5-1800  m  elevation.  Flowering  in 
the  wet  season  and  early  dry  season  (Gentry, 
1973b),  but  Costa  Rican  collections  were  made 
primarily  in  May-July  and  October-January. 
Probably  fruiting  throughout  the  year.  This  spe- 
cies ranges  from  Veracruz,  Mexico,  to  northern 
Argentina. 

Amphilophium  paniculatum  is  recognized  by  its 
unusual  calyx  with  inner  and  outer  layers,  corollas 
that  are  thick  and  held  shut  at  anthesis,  and  the 


BURGER:  FLORA  COSTARICENSIS 


101 


woody  oblong  fruits  with  muriculate  surface. 
Stems  with  six  prominent  longitudinal  ridges, 
dendritic  hairs  (when  present),  and  ovate  blades 
with  palmate  venation  are  useful  vegetative  char- 
acteristics. The  thick  corolla  lobes  are  united  and 
the  two  lips  remain  connivent  until  forced  open 
by  large  bees.  The  flowers  are  full  size  and  white 
or  yellowish  a  day  before  anthesis;  they  then  be- 
come purple.  The  thick  two-layered  calyx  may  be 
protection  against  nectar-robbing  animals.  A  great 
range  of  variation  makes  separation  of  this  species 
into  two  varieties  based  on  pubescence  impracti- 
cal. Because  of  its  foliage  and  hexagonal  stems, 
this  species  can  be  confused  with  Pithecoctenium 
echinatum,  but  that  species  has  simple  hairs  and 
tendrils  with  as  many  as  15  distal  segments. 

Amphilophium  pannosum  (DC.)  Bur.  &  K. 
Schumann  in  Mart.,  Fl.  Bras.  8  (2):  209.  1896. 
Bignonia  pannosa  DC,  Prodr.  9:  148.  1845.  A. 
oxylophium  J.  D.  Smith,  Bot.  Gaz.  31:  119: 
1901.  Figure  26. 

Lianas  or  slender  vines  to  3  cm  diam.,  tendrils 
to  18  cm  long,  leafy  stems  1.5-5  mm  diam.,  with 
6  longitudinal  ridges,  densely  villous  with  hairs 
0.5-3.7  mm  long  yellowish  brown  when  dried; 
pseudostipules  5-18  mm  long,  equally  broad,  sub- 
sessile.  Leaves  usually  2-foliolate,  petioles  2-7 
cm  long,  1.3-2  mm  diam.,  densely  villous,  peti- 
olules  1-4  cm  long;  leaflet  blades  6-14  cm  long, 
3.5-10  cm  wide,  ovate  to  ovate-elliptic,  apex  acu- 
minate, base  rounded  and  truncate  to  cordate,  up- 
per surface  with  simple  hairs  0.5-2  mm  long,  low- 
er surface  densely  villous  with  branched  hairs  to 
2  mm  long,  venation  subpalmate,  2°  veins  4-6/ 
side.  Inflorescences  5-18  cm  long,  1°  peduncles 
2—4  cm  long,  densely  villous,  2°  peduncles  5-12 
mm  long,  bracts  1-1.5  mm,  linear  lanceolate,  ca- 
ducous, 2  bracteoles  present  below  the  calyx. 
Flowers  with  outer  calyx  8-14  mm  long,  densely 
villous,  lobes  to  6  mm  long,  rounded,  recurved  or 
rotate,  to  24  mm  wide;  corolla  22-30  mm  long, 
tubular,  6-9  mm  diam.,  at  first  yellowish  white  but 
turning  purple,  subglabrous  externally,  lip  1 1-14 
mm  long;  filaments  ca.  15  and  12  mm  long,  the- 
cae  ca.  2  mm  long;  style  ca.  2  cm  long.  Fruits  5- 
10  cm  long,  3-5.5  cm  wide,  elliptic-oblong,  com- 
pressed, surfaces  prominently  verrucose  with  tu- 
bercles ca.  2  mm  long,  a  central  vein  prominent 
or  not,  yellowish  pubescent;  seeds  ca.  12  X  48 
mm. 

Uncommon  plants  of  wet  evergreen  forest  for- 
mations on  both  the  Caribbean  and  southern  Pa- 


cific slopes,  5-1400  m  elevation  (but  note  that 
nearly  all  our  collections  come  from  the  southern 
Pacific  slope).  Flowering  in  January,  May,  and 
July-September.  The  species  ranges  from  central 
Costa  Rica  to  northern  Argentina. 

Amphilophium  pannosum  is  recognized  by  its 
climbing  habit,  six-angled  stems,  tendrils  with 
usually  trifid  tips,  densely  villous  yellow-brown 
pubescence,  rounded  pseudostipules,  usually  two- 
foliolate  leaves,  double-margined  calyx,  and  ver- 
rucose fruit.  Specimens  of  A.  pilosum  Standl. 
(1000-1400  m,  in  central  Honduras)  will  key  here 
because  of  similar  pubescence  and  often  retaining 
their  pseudostipules,  but  that  species  may  be  an 
unusual  variant  of  A.  paniculatum. 


Amphitecna  Miers 
Nomen  conservandum 

REFERENCE — A.  Gentry,  Amphitecna  in  Bigno- 
niaceae — Part  I  (Crescentieae  and  Tourrettieae). 
Flora  Neotropica  Monogr.  25(1):  1-130.  1980. 

Small  to  medium-sized  trees,  stems  without 
distinct  phloem  fields  in  cross-section,  glabrous, 
becoming  terete;  pseudostipules  and  glandular 
fields  absent  at  the  nodes.  Leaves  alternate  (sub- 
opposite),  simple,  sessile  to  short-petiolate,  blades 
entire,  subglabrous  or  glabrous  (domatia  absent), 
venation  pinnate.  Inflorescences  terminal,  axil- 
lary or  on  short-shoots  borne  on  older  branchlets 
or  trunks  (cauliflorus),  flowers  solitary  or  fascic- 
ulate, subtended  by  small  bracts,  pedicels  well  de- 
veloped and  drying  black,  with  or  without  brac- 
teoles, glabrous.  Flowers  drying  black,  calyx 
large,  cupular  or  tubular,  splitting  along  1  or  2 
sides  or  opening  irregularly,  usually  glabrous;  co- 
rolla tubular-campanulate  to  funnelform,  usually 
bent  upward  above  a  contracted  region  below  the 
middle  of  the  tube,  greenish  white  to  yellowish 
white,  thick-textured,  5-(4-)  lobed  and  slightly  2- 
lipped;  stamens  4,  slightly  exserted,  filaments 
equal  or  of  2  lengths,  anthers  glabrous,  thecae 
thick  and  somewhat  divergent;  a  staminode  pres- 
ent; ovary  1-  or  2-locular,  ovules  many  on  axile 
placentas  (when  2-locular)  or  on  2  parietal  pla- 
centas (where  unilocular).  Fruits  pendulous,  an 
indehiscent  hard-surfaced  and  thin-walled  pepo 
(calabash),  globose  to  ellipsoid,  usually  subtended 
by  a  circular  perianth  scar,  the  outer  shell  easily 
broken;  seeds  more  than  10  mm  diam.,  embedded 
in  a  soft  or  fleshy  white  pulp,  surfaces  smooth  and 
without  wings. 

Amphitecna  (formerly  called  Dendrosicus  and 


102 


FffiLDIANA:  BOTANY 


including  Enallagma)  is  a  genus  of  20  species 
ranging -from  Mexico,  Central  America,  and  the 
West  Indies  to  northern  South  America.  There  are 
two  centers  of  species  diversity:  southern  Mexico 
to  Belize  and  Guatemala,  and  Costa  Rica  to  Pan- 
ama. The  tree  habit,  lack  of  conspicuous  hairs, 
simple  large  alternate  leaves,  large  flowers  with 
thick  tissues  that  dry  black,  and  pepo-like  fruit 
with  rounded  seeds  are  unusual  traits  among  Big- 
noniaceae  and  are  characteristic  of  the  tribe  Cres- 
centieae.  The  perianth  usually  forms  a  circum- 
scissle  abscission  scar  around  the  base  of  the  fruit. 
The  corolla  often  has  an  abaxial  transverse  inden- 
tation just  distal  to  the  constricted  section  of  the 
corolla  tube,  where  the  distal  portion  of  the  wid- 
ening corolla  tube  bends  upward.  The  large  size, 


strong  odor  (usually  unpleasant),  large  amount  of 
nectar,  and  thick  texture  of  the  flowers  are  char- 
acteristic of  flowers  pollinated  by  bats.  The  flow- 
ers often  vary  significantly  in  size  within  a  single 
species;  leaf  sizes  and  shapes  also  vary  greatly 
within  species.  More  important,  herbarium  mate- 
rial suggests  that  there  may  be  populations  inter- 
mediate between  generally  recognized  species  in 
this  genus.  Whether  this  variation  is  due  to  hy- 
bridization, overly  narrow  species  concepts,  or  in- 
herent variability  is  difficult  to  determine  at  this 
time  because  many  species  are  represented  by 
only  a  few  collections.  Gentry  (1990)  interpreted 
the  greater  number  of  localized  endemic  species 
in  this  genus  as  being  a  result  of  mammal-dis- 
persed seeds,  in  contrast  to  genera  with  wind-dis- 
persed seeds. 


Key  to  the  Species  of  Amphitecna 

la.  Plants  of  ocean  shores  and  low-elevation  swamp  forests;  fruits  subglobose  and  rounded  distally; 
leaf  blades  often  rounded  distally,  coriaceous  A.  latifolia 

Ib.  Plants  of  well-drained  soils,  5-2200  m  elevation;  fruits  usually  oblong  to  ellipsoid  with  a  narrowed 
apex  (sometimes  globose  in  A.  gentryi);  leaf  blades  usually  acute  to  acuminate  at  the  apex,  char- 
taceous  to  subcoriaceous 2 

2a.  Flowers  borne  on  older  leafless  stems  and  trunks;  leaf  blades  to  50  cm  long  [calyx  20-34  mm  long, 
corolla  38-50  mm  long;  10-600  m  elevation] A.  kennedyi 

2b.  Flowers  terminal  on  leafy  shoots  or  on  short-shoots  near  the  leaves;  leaf  blades  to  30  cm  long  .  .  . 
3 

3a.  Flowers  usually  in  clusters  or  2-8,  pedicels  often  with  linear  bracteoles  1-4  mm  long  2-5  mm 
above  the  pedicel  base;  leaves  often  drying  dark;  ( 1 200-)  1 500-2200  m  elevation  [calyx  22-32  mm 
long,  corolla  33-53  mm  long;  fruits  13-17  cm  long]  A.  sessilifolia 

3b.  Flowers  usually  1  or  2,  pedicels  lacking  linear  bracteoles  near  the  base;  leaves  often  drying  brown 
or  grayish;  5-1400  m  elevation  4 

4a.  Calyx  22-28  mm  long,  corolla  44-60  mm  long;  fruits  10-14  cm  long;  5-800  m  elevation.  Pacific 
slope A.  isthmica 

4b.  Calyx  15-20  mm  long,  corolla  25-40  mm  long;  fruits  5-8  cm  long;  600-1400  m  elevation,  Carib- 
bean slope  A.  gentryi 


Amphitecna  gentryi  W.  Burger,  sp.  nov.  Figure 
10. 

Arbor  parva.  Folia  alternata,  glabra,  anguste 
obovata-elliptica  vel  elliptica-oblonga,  acuminata. 
Inflorescencia  terminalia,  floribus  1-2,  pedicelis 
glabris.  Calyx  15-20  mm  longus,  bipartitus,  lobis 
rotundatis,  glabris.  Corolla  22-28  mm  longa, 
campanulata,  alba.  Fructus  globosus  vel  ellipso- 
ideus,  seminibus  in  pulpa  inclusis.  TYPUS:  Costa 
Rica,  Puntarenas,  Monteverde,  in  lower  commu- 
nity, alt.  1350  m.  March  1981.  Haber  478  (holo- 
typus  MO  3162341,  isotypus  CR). 

Small  trees  5-10  m  tall,  trunks  to  18  cm  diam.. 


leafy  stems  2-6  mm  diam.,  terete,  glabrous,  yel- 
lowish or  pale  gray,  expanded  below  the  leaf  base. 
Leaves  alternate,  petioles  0-8  mm  long,  0.7-1.8 
mm  diam.,  glabrous,  often  with  lustrous  hard  yel- 
lowish tissue  at  the  base;  leaf  blades  6-16.5  cm 
long,  2-6.5  cm  wide,  elliptic-obovate,  obovate  or 
narrowly  elliptic-oblong,  apex  acute  to  caudate- 
acuminate,  gradually  narrowed  to  the  cuneate  or 
acute  base,  drying  chartaceous  and  grayish  or 
grayish  brown,  glabrous  above  and  below,  2° 
veins  7-12/side.  Inflorescences  terminal,  of  1 
flower  (rarely  2)  on  leafy  shoots  or  subterminal 
on  short  lateral  shoots,  subtended  by  hard  yellow- 
ish subulate  bracts  1-2  mm  long,  pedicels  30-42 


BURGER:  FLORA  COSTARICENSIS 


103 


mm  long,  glabrous,  sometimes  with  very  small  (1 
mm)  yellowish  bracteoles  1-5  mm  above  the 
base.  Flowers  with  calyx  15-20  mm  long,  ca.  10 
mm  diam.,  glabrous,  splitting  near  the  base  into  2 
broad  lobes,  rounded  distally;  corolla  25-40  mm 
long,  white  or  cream-green,  tube  6-1 1  mm  diam., 
papillate-puberulent  in  the  distal  half,  lobes  ca.  3 
X  10  mm;  filaments  20-23  mm  long,  thecae  ca. 
5  mm  long;  ovary  not  examined.  Fruits  5-8  cm 
long,  4-5  cm  diam.,  oblong  to  subglobose,  with 
a  slightly  narrowed  apex,  surface  smooth,  green 
drying  brown;  seeds  11-14  mm  long,  14-17  mm 
wide,  6-9  mm  thick. 

Plants  of  lower  montane  rain  forest  formations 
of  the  Caribbean  slope,  600-1400  m  elevation. 
Flowering  in  February-March  and  July;  fruiting 
in  March,  June-July,  and  September.  As  presently 
known,  this  species  ranges  from  Volcan  Cacao 
eastward  to  Braulio  Carillo  National  Park  in 
northern  and  central  Costa  Rica. 

Amphitecna  gentry!  is  recognized  by  its  small 
tree  habit,  smaller  subsessile  leaves  turning  gray- 
ish when  dried,  usually  solitary  terminal  flower, 
smaller  corollas,  and  globose  or  oblong  fruit. 
Specimens  placed  here  include  Gomez  et  al. 
21603,  Hammel  &  Chavarria  17533 A,  and  the 
following  collections  by  William  Haber  and  as- 
sociates: 478,  5841,  6673,  7473,  8235,  and  9272. 

Gentry  recognized  these  plants  as  distinct  and 
designated  these  plants  as  Amphitecna  sp.  aff.  A. 
donnell-smithii  (Sprague)  L.  O.  Williams,  a  low- 
land species  of  Belize  and  Guatemala  with  oblan- 
ceolate  leaves  of  similar  length.  This  species  also 
resembles  two  highland  species  of  northern  Cen- 
tral America  described  by  L.  O.  Williams:  A.  mol- 
inae,  of  Honduras,  and  A.  silvicola,  of  the  border 
between  Guatemala  and  Mexico.  More  important, 
it  should  be  noted  that  there  are  collections  that 
appear  to  be  intermediate  between  this  species 
and  A.  isthmica  at  elevations  of  300-600  m  (Her- 
rera  1966,  21 18),  implying  that  the  plants  placed 
here  could  also  be  considered  a  subspecific  high- 
land element  of  A.  isthmica.  Nevertheless,  the 
characteristics  used  in  the  key  seem  to  separate 
the  two  taxa  effectively,  and  only  further  collect- 
ing can  determine  whether  the  intermediates  are 
unusual  individuals  or  are  parts  of  a  cline. 

Amphitecna  isthmica  (A.  Gentry)  A.  Gentry, 
Taxon  25:  108.  1976.  Dendrosicos  isthmicus  A. 
Gentry,  Phytologia  26:  442.  1973.  Figure  10. 

Small  trees  5-10  m  tall,  trunks  to  15  cm  diam., 
leafy  stems  1.7-5  mm  diam.,  glabrous,  pale  yel- 


low or  grayish,  terete.  Leaves  subsessile  or  with 
petioles  to  10  mm  long,  1-2.5  mm  diam.,  gla- 
brous, often  with  hard  lustrous  yellowish  tissue  at 
the  base;  leaf  blades  6-22(-32)  cm  long,  2-8(-10) 
cm  wide,  narrowly  elliptic-obovate  to  narrowly  el- 
liptic-oblong, apex  acuminate,  gradually  narrowed 
to  the  cuneate  base,  drying  stiffly  chartaceous  and 
grayish,  glabrous,  2°  veins  8-17/side.  Inflores- 
cences terminal  (axillary),  usually  1  (2)  flower  on 
leafy  stems,  subtended  by  few  hard  subulate  yel- 
lowish bracts  1-2  mm  long,  pedicel  30-48  mm 
long,  0.8-1.7  mm  diam.,  glabrous,  without  brac- 
teoles. Flowers  drying  black  or  dark  brown,  calyx 
22-30  mm  long,  11-18  mm  wide,  splitting  to  near 
the  base  into  2  broad  obtuse  lobes,  glabrous;  co- 
rolla 44-60  mm  long,  30-40  mm  wide  at  the 
lobes,  greenish  white,  papillate  puberulent  in  the 
distal  half  externally,  tube  5-14  mm  diam.,  lobes 
3-8  mm  long,  with  reflexed  rim;  filaments  20-30 
mm  long,  thecae  4-5  mm  long;  ovary  bilocular  to 
above  the  middle.  Fruits  9-14  cm  long,  3-8  cm 
diam.,  ellipsoid-cylindric,  narrowed  at  the  ends, 
surfaces  smooth  and  punctate,  perianth  scar  thick- 
ened and  disc-like  at  base;  seeds  ca.  10-14  mm 
long,  4  mm  thick. 

Plants  of  wet  evergreen  rain  forest  formations 
on  the  Pacific  slopes,  5-800  m  elevation.  Flow- 
ering material  has  been  collected  in  February, 
May,  August,  and  October-December.  The  spe- 
cies ranges  from  northern  Costa  Rica  to  Colom- 
bia. 

Amphitecna  isthmica  is  recognized  by  its  low- 
land rain  forest  habitat,  small  tree  habit,  narrow 
alternate  leaves  that  usually  dry  grayish,  usually 
solitary  terminal  flowers  on  long  pedicels,  calyx 
splitting  into  two  rounded  lobes,  and  ellipsoid-cy- 
lindrica  fruits.  This  species  lacks  the  bracteoles  on 
the  pedicels  seen  in  A.  gentryi  and  A.  sessilifolia 
and  is  separated  from  A.  kennedyi  by  having  only 
terminal  flowers.  This  species  appears  to  be  re- 
stricted to  the  evergreen  Pacific  slope  in  Costa 
Rica,  which  helps  distinguish  it  from  the  closely 
related  A.  gentryi. 

Amphitecna  kennedyi  (A.  Gentry)  A.  Gentry, 
Taxon  25:  108.  1973.  Demdrosicos  kennedyi  A. 
Gentry,  Phytologia  26:  441.  1973.  Figure  10. 

Small  trees  and  shrubs  3-15  m  tall,  leafy  stems 
1.5-1 1  mm  diam.,  glabrous,  ridged  below  the  leaf 
base  but  becoming  terete  and  pale  grayish.  Leaves 
with  petioles  2-12  mm  long,  1-4  mm  diam.,  gla- 
brous, often  with  lustrous  hard  tissue  developed 
at  the  base;  leaf  blades  14-35(-50)  cm  long,  5- 


104 


FIELDIANA:  BOTANY 


13(-21)  cm  wide,  elliptic-obovate  to  narrowly  ob- 
ovate  or  oblanceolate,  apex  usually  short-acumi- 
nate, gradually  narrowed  to  the  cuneate  base,  dry- 
ing chartaceous  and  brownish  or  grayish,  glabrous 
above  and  below,  2°  veins  8-17(-22)/side,  often 
weakly  loop-connected  near  the  margin.  Inflores- 
cences cauliflorous,  1-3  flowers  on  short-shoots 
borne  below  leafy  nodes  on  branchlets  or  on  the 
surface  of  larger  stems,  subtended  by  stiff  grayish 
subulate  bracts  2-3  mm  long,  pedicels  25-45  mm 
long,  0.6-1.7  mm  diam.,  glabrous.  Flowers  dry- 
ing black,  calyx  20-34  mm  long,  9-20  mm  diam., 
split  to  near  the  base  to  form  2  lobes,  glabrous; 
corolla  38-56  mm  long,  tubular-campanulate,  1 3- 
19  mm  wide  at  the  mouth,  greenish  white  to  yel- 
lowish green,  with  peltate  or  papillate  hairs  on  the 
distal  half  externally,  lobes  forming  a  reflexed  bi- 
labiate rim;  filaments  24-28  mm  long,  thecae  4- 
5  mm  long;  ovary  lepidote,  1-locular  with  2  pa- 
rietal placentae.  Fruits  11-16  cm  long,  5-7  cm 
diam.,  ellipsoid,  apiculate  at  base  and  apex;  seeds 
13-15  mm  long,  embedded  in  pulp. 

Plants  of  evergreen  rain  forest  formations  of 
both  Caribbean  and  southern  Pacific  slopes  (in 
Costa  Rica),  10-600  m  elevation.  Flowering  col- 
lections were  made  in  January-April  and  July- 
September.  This  species  ranges  from  the  east  coast 
of  Honduras  to  central  Panama. 

Amphitecna  kennedyi  is  distinguished  by  its 
small  tree  habit,  flowers  borne  at  leafless  nodes  or 
on  the  older  stems,  corolla  with  poorly  developed 
lobes  forming  a  reflexed  rim,  ellipsoid  fruits,  and 
lowland  rain  forest  habitat.  The  lower  half  of  the 
corolla  often  has  a  smoother  texture  demarcated 
distally  by  a  transverse  ridge. 

Amphitecna  latifolia  (Miller)  A.  Gentry,  Taxon 
25:  108.  1976.  Crescentia  latifolia  Miller,  Card. 
Diet.  ed.  8:  306.  1768.  C.  obovata  Benth.,  Bot. 
Voy.  Sulphur  130.  1844.  Enallagma  latifolia 
(Miller)  Small,  Fl.  Miami  171.  1913.  -A.  obovata 
(Benth.)  L.  O.  Williams,  Fieldiana,  Bot.  36:  25. 
1973.  Dendrosicos  latifolius  (Miller)  A.  Gentry, 
Taxon  22:  644.  1973.  Figure  10. 

Small  trees  (2-)4-8(-12)  m  tall,  trunks  to  20 
cm  diam.,  branches  often  crooked,  leafy  stems 
2.5-12  cm  diam.,  terete,  glabrous,  often  becoming 
pale  gray,  lenticellate.  Leaves  alternate  or  subop- 
posite,  petioles  2-14  mm  long  but  poorly  differ- 
entiated from  the  leaf,  glabrous;  leaf  blades  10- 
28  cm  long,  4-14  cm  wide,  narrowly  or  broadly 
obovate  to  obovate-oblong  or  oblong,  apex  short- 
acuminate  to  rounded,  base  obtuse  to  cuneate  and 


slightly  decurrent  on  the  petiole,  drying  subcori- 
aceous  to  coriaceous,  glabrous  or  with  minute 
(0.05  mm)  peltate  hairs  beneath,  2°  veins  7-13/ 
side,  weakly  loop-connected  near  the  margin.  In- 
florescences terminal  or  below  leafless  nodes,  of 
1-3  flowers,  subtended  by  stiff  subulate  bracts  2- 
3  mm  long,  pedicels  (22-)40-63  mm  long,  gla- 
brous. Flowers  glabrous  externally,  drying  black, 
calyx  28-38  mm  long,  tube  7-15  mm  diam.,  split- 
ting along  the  lateral  sides  nearly  to  the  base  to 
form  2  divergent  lobes;  corolla  45-65  mm  long, 
tube  1 2-24  mm  diam.,  3-4  cm  wide  at  the  apex, 
tubular-campanulate,  greenish  white  or  pale  yel- 
lowish green,  lobes  5-15  mm  long;  filaments  ca. 
25  mm  long,  anthers  5-6  mm  long;  style  5-6  cm 
long.  Fruits  7-10  cm  long,  6-10  cm  wide,  glo- 
bose or  subglobose,  green,  surface  smooth  and 
hard,  with  thickened  disc-like  perianth  scar  at 
base;  seeds  13-16  mm  long,  14-17  mm  wide,  5- 
8  mm  thick,  surface  smooth. 

Small  trees  growing  just  inland  from  coastal 
mangroves  and  in  low-elevation  swamps  in  ev- 
ergreen and  deciduous  areas  of  both  the  Pacific 
and  Caribbean  coasts,  0-20(-200)  m  elevation. 
Flowering  and  fruiting  throughout  the  year  (more 
abundantly  in  the  wet  season).  This  species  ranges 
from  southern  Florida,  the  West  Indies,  Mexico, 
and  Central  America  to  Venezuela  and  Ecuador. 

Amphitecna  latifolia  is  recognized  by  the  tree 
habit,  stiff  simple  alternate  leaves,  glabrous  flow- 
ers drying  black  and  arising  directly  from  distal 
stems,  globose  indehiscent  fruit,  and  rounded 
seeds  embedded  in  pulp.  The  flowers  have  a  dis- 
agreeable odor.  Common  names  are  calabasillo  de 
playa,  jicarita,  jicaro  de  playa  and  swamp  cala- 
bash (Belize). 

Amphitecna  sessilifolia  (J.  D.  Smith)  L.  O.  Wil- 
liams, Fieldiana,  Bot.  36:  25.  April  1973.  Ta- 
bebuia  sessilifolia  J.  D.  Smith,  Bot.  Gaz.  25: 
156.  1898.  Neotuerckheimia  gonoclada  J.  D. 
Smith,  Bot.  Gaz.  47:  259.  1909.  Enallagma  ses- 
silifolia (J.  D.  Smith)  Standl.,  Publ.  Field  Mus. 
Nat.  Hist.,  Bot.  Ser.  18:  1120.  1938.  A.  sessili- 
folius  (J.  D.  Smith)  A.  Gentry,  Taxon  22:  646. 
Nov.  1973.  Figure  10. 

Trees  or  shrubs  to  12  m  tall,  trunks  7-25  cm 
diam.,  leafy  stems  4-1 1  mm  diam.,  glabrous,  with 
ridges  beneath  the  leaf  base,  pale  gray,  becoming 
terete.  Leaves  alternate,  petioles  2-10  mm  long, 
poorly  differentiated  from  the  blade,  glabrous; 
leaf  blades  10-35  cm  long,  2-11  cm  wide,  nar- 
rowly obovate  to  oblanceolate  or  narrowly  oblan- 


BURGER:  FLORA  COSTARICENSIS 


105 


ceolate,  apex  acute  to  acuminate,  gradually  nar- 
rowed to  the  cuneate  base,  drying  chartaceous, 
subglabrous  with  minute  (0.05  mm)  peltate  hairs, 
2°  veins  (8-)l  1-22/side,  often  arising  at  70-90° 
from  the  midvein.  Inflorescences  terminal  fasci- 
cles of  2-8  flowers,  basal  bracts  to  7  mm  long, 
pedicels  22-65  mm  long,  1-2  mm  diam.,  gla- 
brous, drying  black,  often  with  2  small  (1-4  mm) 
linear  bracteoles  2-5  mm  above  the  base.  Flowers 
with  calyx  inflated  before  anthesis,  22-32  mm 
long,  11-19  mm  diam.,  glabrous,  splitting  later- 
ally into  2  lobes,  margins  rounded;  corolla  33-53 
mm  long,  25-35  mm  wide  distally,  white  with 
greenish  tinge,  tube  6-13  mm  diam.,  bent  upward 
near  the  middle,  lobes  3-12  mm  long,  glabrous 
proximally  but  with  minute  (0. 1  mm)  peltate  hairs 
on  the  lobes  externally;  filaments  ca.  26  and  24 
mm  long,  thecae  ca.  5  mm  long.  Fruits  12-17  cm 
long,  6-9  cm  diam.,  subglobose  to  ellipsoid-ob- 
long with  narrowed  apex  to  1  cm  long,  surface 
smooth,  glabrous,  drying  dark;  seeds  15-20  mm 
long,  15-24  mm  wide. 

Understory  trees  within  evergreen  montane  rain 
forests,  1200-2200  m  elevation.  Flowering 
throughout  the  year.  This  species  ranges  from  the 
Cordillera  de  Tilaran  to  the  Chiriqui  highlands  of 
western  Panama. 

Amphitecna  sessilifolia  is  recognized  by  its  tree 
habit,  alternate  oblanceolate  leaves,  large  2-lobed 
calyx,  pale  green  corollas,  smooth  oblong-ellip- 
soid fruits,  and  restriction  to  montane  forests.  An- 
other important  distinction  is  that  the  thick  calyx 
splits  apart,  often  resulting  in  a  whitish  tissue  ex- 
posed along  the  edge.  A  number  of  specimens 
from  Monteverde  have  more  clearly  differentiated 
petioles  and  corolla  lobes  lacking  peltate  hairs  ex- 
ternally; these  have  been  annotated  A.  haberi  (an 
unpublished  name)  by  Gentry.  Because  they  are 
otherwise  identical  to  A.  sessilifolia  and  live  in 
the  same  kind  of  habitat,  it  seems  best  to  consider 
them  no  more  than  a  local  variant  of  A.  sessili- 


folia. Common  names  are  calabacero,  calabash, 
quacalillo,  jicarilla,  jicaro. 


Anemopaegma  Martius  ex  Meisner 
Nomen  conservandum 

Lianas  or  vines  (in  Central  America),  climbing 
with  simple  or  distally  trifid  tendrils,  stem  with  8 
(rarely  4)  phloem  areas  in  cross-section,  terete; 
nodes  usually  with  interpetiolar  lines,  gland  fields 
absent;  pseudostipules  leaf-like  to  small  or  absent. 
Leaves  opposite,  2-5-foliolate,  petiolate,  often 
bearing  a  tendril  coiled  near  the  tip,  blades  often 
drying  yellowish  or  grayish  green,  margins  entire, 
venation  pinnate.  Inflorescences  axillary  or  ter- 
minal, few-flowered  racemes  or  1  or  2  flowers, 
bracts  small,  pedicels  short.  Flowers  with  cupular 
or  campanulate  calyx,  truncated  and  subentire  or 
slightly  lobed,  often  with  glands  below  the  margin 
externally;  corolla  tubular-campanulate  or  funnel- 
form,  5-lobed  and  somewhat  2-lipped,  bright  yel- 
low to  pale  yellow  or  white,  externally  glabrous 
or  with  minute  glandular  peltate  hairs;  stamens  4, 
included,  filaments  of  2  lengths,  anthers  glabrous, 
thecae  straight  and  divaricate,  staminode  present; 
disc  pulviniform;  ovary  usually  stipitate,  ellipsoid, 
puberulent  or  with  peltate  hairs,  2-locular  with 
ovules  in  2-6  series  in  each  locule.  Fruits  woody 
or  coriaceous  capsules,  elliptic  to  orbicular,  valves 
strongly  flattened  or  rounded,  smooth;  seeds  flat, 
completely  surrounded  by  a  membranaceous  sub- 
orbicular  wing  or  wingless  and  with  larger  brown 
corky  seed  area  and  thinner  distal  area. 

Anemopaegma  is  a  genus  of  about  46  species 
ranging  from  Mexico  to  Brazil  and  Argentina; 
most  of  the  species  are  South  American.  The  great 
variation  found  within  some  species  has  resulted 
in  a  large  number  of  names,  and  it  is  often  diffi- 
cult to  demarcate  species.  Many  of  the  proposed 
species  probably  will  have  to  be  brought  together 
into  broader  species  concepts. 


Key  to  the  Species  of  Anemopaegma 

la.  Leaves  usually  with  4  or  5  leaflets A.  orbiculatum 

Ib.  Leaves  with  no  more  than  2  or  3  leaflets 2 

2a.  Pseudostipules  conspicuous  (4-9  mm  long);  tendrils  lacking  trifid  tips;  fruits  ellipsoid,  seeds  brown 

and  lacking  a  translucent  outer  wing  A.  chrysoleucum 

2b.  Pseudostipules  absent  or  minute;  tendrils  often  trifid  near  the  tip;  fruits  flattened  and  oblong,  seeds 

pale  yellowish  with  thin  translucent  circumferential  wing  (where  known)  3 

3a.  Leaves  usually  puberulent  beneath,  leaflet  margins  not  involute  [stems  terete,  lacking  corky  ridges; 

corolla  tube  subglabrous  externally] A.  puberulum 


106 


FIELDIANA:  BOTANY 


3b.  Leaves  subglabrous  beneath,  leaflet  margins  often  involute 4 

4a.  Corolla  tube  with  minute  peltate  hairs  externally;  stems  tetragonal  and  often  with  corky  ridges; 

seeds  not  known A.  santaritensis 

4b.  Corolla  tube  glabrous;  stems  subterete  and  longitudinally  striate;  seeds  suborbicular 

.  .  A.  chrysanthum 


Anemopaegma  chrysanthum  Dugand,  Caldasia 
4:  307.  1947. 

Lianas  or  slender  vines,  stems  to  2  cm  diam., 
tendrils  to  13  cm  long,  trifid  at  tip,  leafy  stems 
1.2-8  mm  diam.,  glabrous,  longitudinally  striate, 
grayish  and  becoming  lenticellate;  pseudostipules 
minute.  Leaves  foliolate,  petioles  8-50  mm  long, 
1-2.5  mm  thick,  glabrous,  petiolules  8-40  mm 
long;  leaflet  blades  5-17(-20)  cm  long,  3-9(-12) 
cm  wide,  broadly  ovate  to  elliptic,  apex  acute  or 
short-acuminate,  base  acute  to  obtuse,  drying  stiff- 
ly chartaceous  and  grayish  green,  subglabrous,  2° 
veins  3-7/side.  Inflorescences  short  axillary  ra- 
cemes with  2-8  flowers,  peduncles  ca.  10  mm 
long,  pedicels  10-15  mm  long.  Flowers  subgla- 
brous, calyx  6-9  mm  long,  4-6  mm  diam.  with 
distal  glands,  margin  subentire;  corolla  50-70 
mm  long,  yellow,  mouth  12-22  mm  wide,  lobes 
10-15  mm  long;  filaments  24-32  and  16-23  mm 
long,  thecae  4-5  mm  long.  Fruits  8-20  cm  long, 
5-8  cm  wide,  1  cm  thick,  oblong-elliptic,  flat- 
tened, surface  smooth,  yellow-brown,  stipe  1-2 
cm  long;  seeds  40-55  mm  diam.,  suborbicular. 

Plants  of  evergreen  forest  formations  of  the 
northern  cordilleras  in  Costa  Rica;  350-1000  m 
elevation.  Flowering  in  March-April.  This  species 
ranges  from  Mexico  to  Ecuador. 

Anemopaegma  chrysanthum  is  recognized  by 
its  three-tipped  tendrils,  two-foliolate  opposite 
leaves,  few-flowered  axillary  racemes,  glabrous 
yellow  corollas,  and  broad  flat  fruits  with  subor- 
bicular seeds.  Rarely  collected  in  Central  Ameri- 
ca, this  species  has  only  recently  been  found  in 
Costa  Rica. 

Anemopaegma  chrysoleucum  (Kunth  in  H.B.K.) 
Sandw.,  Lilloa  3:  459.  1938.  Bignonia  chryso- 
leucum Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3:  134. 
1819.  A.  puncticulatum  Pittier  &  Standl.,  J. 
Wash.  Acad.  Sci.  15:  461.  1925.  A.  macrocarpa 
Standl.,  Publ.  Field  Columb.  Mus.,  Bot.  Sen  4: 
262.  1929.  Figure  20. 

Lianas  or  slender  vines,  stems  to  2  cm  diam., 
tendrils  to  17  cm  long,  simple,  leafy  stems  1.2-5 
mm  diam.,  minutely  puberulent  at  the  nodes,  lon- 


gitudinally striate,  interpetiolar  lines  usually  well 
developed;  pseudostipules  4-9  mm  long,  broadly 
ovate.  Leaves  2-(3-)  foliolate,  petioles  8-34  mm 
long,  0.8-1.5  mm  diam.,  glabrous  or  minutely 
(0.1-0.2  mm)  puberulent,  petiolules  10-18  mm 
long;  leaflet  blades  5-13(-18)  cm  long,  2-6(-8) 
cm  wide,  narrowly  ovate-elliptic  to  elliptic,  apex 
acute  or  acuminate,  base  acute  to  obtuse,  drying 
chartaceous  and  yellowish  green  or  gray-green,  mi- 
nutely punctate,  glabrous  except  for  minute  hairs 
on  the  midvein  above,  2°  veins  5-8/side.  Inflores- 
cences of  1  or  2  axillary  flowers  (1-4/node)  or  a 
raceme  to  9  cm  long  with  ca.  5  flowers,  peduncles 
3-15  mm  long,  1  mm  diam.,  minutely  puberulent, 
pedicels  6-14  mm  long.  Flowers  with  calyx  8-1 1 
mm  long,  7-9  mm  diam.,  rounded  glands  and  mi- 
nute peltate  hairs  or  subglabrous,  margin  entire;  co- 
rolla 58-105  mm  long,  white  to  yellowish  white, 
glabrous  externally,  narrowed  base  of  tube  ca.  2 
cm  long,  mouth  1 2-20  mm  wide,  lobes  1 2-25  mm 
long,  17-24  mm  wide;  filaments  24-34  and  16-25 
mm  long,  thecae  4-5  mm  long.  Fruits  7-13  cm 
long,  3-5  cm  wide,  2  cm  thick,  ellipsoid,  surface 
smooth  and  lustrous,  yellow-brown,  stipe  ca.  1  cm 
long;  seeds  ca.  15  X  22  mm,  brown  throughout, 
margin  thin  but  not  translucent. 

Plants  of  river  edges  and  swampy  areas  in  low- 
land evergreen  rain  forest  formations  on  both  the 
Caribbean  and  Pacific  slopes,  0-50  m  elevation. 
Flowering  throughout  the  year.  This  species  rang- 
es from  Mexico  to  Venezuela  and  Peru. 

Anemopaegma  chrysoleucum  is  recognized  by 
its  slender  climbing  stems  with  simple  tendrils, 
usually  bifoliolate  opposite  leaves,  small  leaf-like 
pseudostipules,  few-flowered  axillary  inflores- 
cences, large  white  to  yellowish  corollas,  and  el- 
lipsoid fruits  with  flat  seeds  that  lack  thin  trans- 
lucent membranous  wings.  This  species  differs 
from  its  congeners  by  having  seeds  with  stiff 
cardboard-like  texture  that  are  dispersed  by  water. 

Anemopaegma  orbiculatum  (Jacq.)  DC.,  Prodr. 
9:  190.  1845.  Bignonia  orbiculata  Jacq.,  Sel. 
Stirp.  Am.  Hist.  184,  tab.  180,  fig.  79.  1763. 
Pithecoctenium  panamense  Benth.,  Bot.  voy. 
Sulphur  129.  1844.  Figure  20. 

Slender  lianas  to  5  m  high,  stems  to  25  mm 
diam.,  longitudinally  striate,  hollow,  tendrils  8-17 


BURGER.  FLORA  COSTARICENSIS 


107 


cm  long,  simple  or  with  trifid  arms  to  5  mm  long, 
leafy  stems  1.5-9  mm  diam.,  minutely  (0.1-0.2 
mm)  puberulent  with  thin  hairs,  an  interpetiolar 
line  present  or  absent;  pseudostipules  rarely  pres- 
ent. Leaves  4-  or  5-foliolate,  petioles  3-14  cm 
long,  1.2-2.2  mm  diam.,  minutely  puberulent, 
longitudinally  striate,  tendril  scar  ca.  2  mm  wide, 
petiolules  4-40(-55)  mm  long;  leaflet  blades  5- 
15(-22)  cm  long,  3-8(-9.5)  cm  wide,  ovate-ellip- 
tic to  ovate,  apex  acuminate,  base  obtuse  to 
rounded  and  subcordate,  drying  chartaceous  and 
greenish,  punctate  above,  minutely  and  sparsely 
puberulent  beneath,  2°  veins  5-7/side.  Inflores- 
cences axillary.  2-10  cm  long,  racemes  with  1- 
1 0  flowers  (rarely  with  a  cymose  lateral  branch  at 
the  first  node),  peduncles  3-30  mm  long,  ca.  1.2 
mm  diam.,  minutely  puberulent,  pedicels  6-12 
mm  long.  Flowers  with  cupular-campanulate  ca- 
lyx 6-10  mm  long,  5-9  mm  wide,  margin  sub- 
entire  or  with  5  irregular  minute  teeth,  minutely 
(0.1  mm)  puberulent;  corolla  33-70  mm  long, 
tube  14-20  mm  wide  at  the  mouth,  yellow  and 
often  with  the  lobes  paler,  glandular  puberulent 
externally  (more  densely  near  the  base),  lobes  8- 
18  mm  long,  the  lower  reflexed;  filaments  ca.  16 
and  23  mm  long,  thecae  ca.  4  mm  long,  divaricate 
at  180°.  Fruits  9-14  cm  long,  6-10  cm  wide,  ca. 
2  cm  thick  at  center,  suborbicular  to  broadly  el- 
liptic, with  a  basal  stipe  8-15  mm  long,  valves 
slightly  convex  or  flattened,  surface  smooth  and 
lustrous,  yellowish  brown;  seeds  30-38  mm  long, 
40-47  mm  wide,  elliptic-orbicular,  body  ca.  14 
mm  wide,  wings  transparent  near  the  periphery. 

Plants  of  forest  edges  in  evergreen  lowland  rain 
forest  formations  of  the  Caribbean  slope  and  Ni- 
coya  peninsula,  5-900  m.  Flowering  in  April- 
September.  Nearly  all  Costa  Rican  collections 
come  from  near  La  Selva  and  the  Rio  Sarapiqui- 
Rio  Frio  drainage  areas;  we  have  no  collections 
of  this  species  from  southern  Costa  Rica.  This 
species  ranges  from  El  Salvador  to  Colombia  and 
Venezuela. 

Anemopaegma  orbiculatum  is  recognized  by  its 
climbing  stems  with  simple  or  trifid  tendrils,  op- 
posite four-  or  five-foliolate  leaves,  subentire  ca- 
lyx cups,  bright  yellow  corollas,  and  suborbicular 
fruits  with  orbicular  winged  seeds.  This  is  the 
only  bignoniaceous  vine  in  Central  America  with 
consistently  five-foliolate  or  four-foliolate  leaves. 

Anemopaegma  puberulum  (Seibert)  Miranda, 
Anal.  Inst.  Biol.  Mexico  24:  93.  1953.  Chodan- 
thus  puberulum  Seibert,  Carnegie  Inst.  Wash- 
ington Publ.  527:  425.  1940.  Figure  20. 


Slender  lianas  to  2  cm  diam.,  tendrils  1 1-27 
cm  long  and  trifid  at  the  tip,  leafy  stems  2-6  mm 
diam.,  glabrous  or  puberulent  with  thin  straight 
hairs  0.1-0.3  mm  long,  longitudinaly  striate  when 
dry,  interpetiolar  ridge  usually  present;  pseudo- 
stipules  minute.  Leaves  2-foliolate  with  tendril  or 
tendril  scar,  petioles  18-40  mm  long,  1.5-2.3  mm 
diam.,  glabrous  or  minutely  puberulent,  petiolules 
10-45  mm  long,  1-2.2  mm  diam.;  leaflet  blades 
8-21  cm  long,  4-12  cm  wide,  ovate  to  ovate-el- 
liptic, apex  acuminate,  base  obtuse  to  truncate  or 
subcordate,  drying  stiffly  chartaceous,  usually  gla- 
brous above,  punctate,  minor  venation  raised 
above  (dried),  lower  surface  glabrous  to  pubes- 
cent with  straight  thin  hairs  to  0.1-0.4  mm  long, 
2°  veins  4-7/side,  3°  veins  prominent  on  both  sur- 
faces. Inflorescences  axillary,  2-8-flowered  ra- 
cemes to  6  cm  long,  peduncles  14-20  mm  long, 
ca.  1.5  mm  diam.,  glabrous  or  puberulent,  pedi- 
cels ca.  7  mm  long.  Flowers  with  cupular  calyx 
6-9  mm  long,  5-8  mm  diam.,  distal  margin  sub- 
entire  or  slightly  lobed,  puberulent,  glabrous  or 
with  few  minute  peltate  hairs;  corolla  50-71  mm 
long,  tubular  campanulate,  yellow,  glabrous  or  pa- 
pillate-puberulent  externally,  tube  12-21  mm 
diam.  at  mouth,  lobes  10-15  mm  long,  10-14  mm 
wide,  rounded,  filaments  ca.  14  and  20  mm  long, 
thecae  3-4  mm  long.  Fruits  10-21  cm  long  (in- 
cluding the  11-20  mm  stipe),  6-8  cm  wide,  ca. 
13  mm  thick,  oblong-elliptic,  valves  flattened, 
woody,  with  a  median  longitudinal  line,  surface 
lustrous,  yellowish  brown;  seeds  35-60  mm 
diam.,  suborbicular  with  thin  translucent  periph- 
eral wing,  body  14-18  mm  long. 

Uncommon  plants  of  evergreen  or  partly  decid- 
uous forest  formations  on  the  Caribean  and  south- 
ern Pacific  slopes  in  Costa  Rica,  5-800(-1200)  m 
elevation.  Probably  flowering  throughout  the  year 
(mostly  in  the  dry  season,  January-May).  This 
species  ranges  from  Veracruz,  Mexico,  to  Ecua- 
dor. 

Anemopaegma  puberulum  is  recognized  by  its 
vining  habit  with  trifid  tendrils,  opposite  two-fo- 
liolate  leaves,  few-flowered  axillary  inflorescenc- 
es, subentire  calyx  cups,  large  yellow  subglabrous 
corollas,  flattened  oblong-elliptic  woody  fruit,  and 
seeds  with  thin  suborbicular  peripheral  wings.  The 
glabrous  lustrous  upper  leaf  surface  with  elevated 
minor  venation,  lack  of  conspicuous  pseudostip- 
ules, and  fruits  with  a  longitudinal  line  down  the 
middle  also  help  distinguish  this  uncommon  spe- 
cies from  its  congeners.  Specimens  with  densely 
puberulent  stems  and  leaves  do  look  quite  differ- 
ent from  those  that  are  essentially  glabrous,  but 
they  seem  to  differ  in  no  other  way,  and  there  are 


108 


FffiLDIANA:  BOTANY 


intermediate  collections.  For  this  reason,  A.  chry- 
santhum  may  prove  to  be  a  synonym  of  A.  pub- 
erulum. 

Anemopaegma   santaritense   A.    Gentry,   Ann. 
Missouri  Bot.  Card.  58:  93.  1971.  Figure  20. 

Slender  lianas  climbing  with  trifid  (simple)  ten- 
drils 3-12  cm  long,  leafy  stems  1.5-4  mm  diam., 
glabrous  or  minutely  papillate-puberulent,  longi- 
tudinally ridged  or  with  longitudinal  corky  ridges, 
tetragonal,  interpetiolar  ridge  present  or  absent; 
pseudostipules  absent.  Leaves  2-foliolate,  with  a 
tendril  or  tendril  scar,  petioles  15-34  mm  long, 
0.9-1.3  mm  diam.,  glabrous  or  minutely  papillate- 
puberulent  along  the  upper  (adaxial)  surface,  pet- 
iolules  6-17  mm  long;  leaflet  blades  5-11  cm 
long,  2.5-6.5  cm  wide,  ovate  to  narrowly  ovate- 
elliptic  or  ovate-orbicular,  apex  acuminate  to  cau- 
date-acuminate, base  obtuse  to  somewhat  round- 
ed, drying  chartaceous  and  greenish,  glabrous  or 
with  minute  papillate  hairs  along  the  midvein  on 
both  surfaces,  2°  veins  3-5/side.  Inflorescences 
axillary,  flowers  1  or  2  or  2-8  on  a  raceme  to  9 
cm  long,  subglabrous,  bracts  0.5-1  mm  long,  ped- 
icels 5-14  mm  long.  Flowers  with  cupular  calyx 
6-10  mm  long,  5-8  mm  diam.,  glabrous  or  with 
few  minute  papillate  hairs,  margin  entire;  corolla 
52-75  mm  long,  tubular-campanulate,  17-20  mm 
diam.  at  mouth,  pale  yellow  or  white,  glabrous  or 
with  minute  peltate  hairs  externally,  lobes  8-12 
mm  long;  filaments  ca.  20  and  18  mm  long,  the- 
cae  3-4  mm  long;  ovary  stipitate,  ca.  2  mm  long. 
Fruits  not  known. 

Rarely  collected  plants  of  evergreen  rain  forest 
formations  on  the  Caribbean  slope  and  Golfo  Dul- 
ce,  0-1000  m  elevation.  Flowering  collections 
have  been  made  in  March-May  and  September  in 
Costa  Rica  and  Panama.  This  species  ranges  from 
eastern  Costa  Rica  to  Colombia. 

Anemopaegma  santaritensis  is  recognized  by 
its  climbing  habit  and  usually  trifid  tendrils,  te- 
tragonal stems,  opposite  two-foliolate  leaves,  few- 
flowered  axillary  inflorescences,  calyx  cups  with 
subentire  margins,  and  larger  yellow  or  white  co- 
rollas with  minute  peltate  hairs.  The  upper  leaf 
surfaces  are  dull  green  and  often  reflexed  along 
the  margins  when  dried. 


Arrabidaea  DeCandolle 

Lianas  climbing  with  simple  tendrils  (rarely 
small  trees  or  shrubs),  stems  terete,  with  4  phloem 
areas  in  cross-section,  interpetiolar  ridges  and 


gland  fields  often  present  at  the  nodes;  pseudo- 
stipules  usually  small  and  inconspicuous.  Leaves 
opposite,  petiolate,  3-foliolate  or  2-foliolate  with 
tendril  or  tendril  scar  (rarely  simple),  leaflets  pe- 
tiolulate,  margins  entire,  puberulent  to  subgla- 
brous, venation  pinnate  or  palmate,  domatia  rarely 
present.  Inflorescences  often  terminal  pyramidal 
thyrses  or  panicles,  also  axillary  or  from  leafless 
nodes,  usually  many-flowered,  bracts  minute,  ped- 
icels usually  puberulent.  Flowers  small  to  large, 
calyx  cupulate  to  tubular  or  campanulate  (rarely 
patelliform),  distally  truncated  and  entire  or  mi- 
nutely 5-denticulate,  puberulent;  corolla  tubular- 
campanulate  or  funnelform,  usually  somewhat  2- 
lipped,  pale  pink  to  purple  (white),  usually  dense- 
ly puberulent  externally,  lobes  5,  unequal,  round- 
ed at  the  apex;  stamens  4,  included,  filaments  of 
2  lengths,  anthers  glabrous,  thecae  usually  divar- 
icate, staminode  present;  disc  annular;  ovary  nar- 
rowly cylindrical,  with  peltate  hairs,  2-locular, 
ovules  2-seriate.  Fruits  long-linear  capsules,  sep- 
ticidally  dehiscent,  valves  flattened  parallel  to  the 
septum,  coriaceous  and  usually  with  a  raised  lon- 
gitudinal median  vein,  smooth  (rarely  tubercu- 
late);  seeds  much  wider  than  long  with  2  mem- 
branaceous  lateral  wings  (rarely  corky),  body  of 
the  seed  usually  differentiated  from  the  thin 
wings. 

Arrabidaea  is  a  genus  of  about  70  species  rang- 
ing from  Mexico  and  the  West  Indies  to  Argen- 
tina; the  majority  of  species  are  South  American. 
Arrabidaea  inaequalis  (DC.  ex  Spreng.)  K. 
Schum.  is  disjunct  between  Belize  and  Colombia 
but  should  be  searched  for  in  our  flora;  it  is  easily 
recognized  by  its  twice-compound  leaves. Two 
other  species  are  included  in  this  treatment  that 
have  not  been  collected  in  Costa  Rica:  A.  florida 
and  A.  pubescens  (q.v.).  Our  species  of  Arrabi- 
daea are  mostly  lianas  with  simple  tendrils  and 
two-  or  three-foliolate  leaves,  usually  having  pa- 
niculate inflorescences,  narrowly  cupular  calyx 
with  entire  or  minutely  dentate  margin,  purple  to 
lilac  or  rose  corollas  that  are  puberulent  external- 
ly, long  narrow  fruits  with  flattened  valves,  and 
flat  seeds,  often  with  thin  lateral  wings  transparent 
at  the  tips.  A  number  of  our  species  have  very 
short  flowering  periods,  which  may  account  for 
their  poor  representation  in  herbaria.  There  is 
great  variation  within  many  of  the  species,  which 
can  make  determination  of  specimens  lacking 
flowers  very  difficult.  These  plants  are  easily  con- 
fused with  species  of  Cydista  (lacking  a  disc  at 
the  base  of  the  ovary)  and  Lundia  (pubescent  an- 
thers). 


BURGER:  FLORA  COSTARICENSIS 


109 


Key  to  the  Species  of  Arrabidaea 

la.  Shrubs,  small  trees,  or  rarely  lianas,  tendrils  usually  absent;  gland  fields  absent;  simple  leaves  often 
present  beneath  3-foliolate  leaves  on  distal  stems;  inflorescences  short  racemes  with  <  10  flowers; 

fruits  often  held  erect  [plants  of  dry  deciduous  forests,  flowering  in  May-June]   

A.  costaricensis 

Ib.  Lianas,  tendrils  usually  present,  gland  field  present  or  absent;  simple  leaves  only  found  on  young 
plants  (except  in  A.  panel lifera);  inflorescences  usually  paniculate  with  >  10  flowers;  fruits  pen- 
dulous   2 

2a.  Calyx  14-23  mm  long,  tubular;  fruits  with  prominently  tuberculate  valves,  becoming  black  or  dark 
brown;  seeds  16-18  mm  long  [leaves  2-foliolate;  flowering  in  the  wet  season  in  evergreen  forest 
formations] A.  verrucosa 

2b.  Calyx  1-9  mm  long,  cupulate,  short-tubular  or  flattened  and  saucer-like;  fruits  with  smooth  or 
slightly  muricate  valves  becoming  yellowish  brown  to  dark  brown,  seeds  6-14  mm  long  3 

3a.  Calyx  1-4  mm  long,  flattened  and  saucer-like  (patelliform),  entire  and  lacking  teeth;  corolla  tube 
glabrous  externally  but  with  the  lobes  puberulent  and  whitish  [deciduous  forests,  flowering  in  July- 
October]  A.  patellifera 

3b.  Calyx  3-9  mm  long,  cupulate  to  short-tubular,  entire  or  with  minute  teeth;  corolla  tube  puberulent 
above  the  narrowed  base,  lobes  pink  to  purple  4 

4a.  Corolla  8-18  mm  long  [flowers  in  dense  distal  clusters  on  large  inflorescences;  leaves  usually 
subglabrous  and  drying  grayish;  interpetiolar  gland  fields  usually  absent;  not  known  to  occur  be- 
tween Nicaragua  and  central  Panama]  A.  florida 

4b.  Corolla  1 8-55  mm  long    5 

5a.  Leaves  usually  drying  reddish  brown  above  and  below,  usually  subglabrous  and  lustrous  with  minor 
venation  raised  above  and  below;  plants  of  wet  evergreen  forests  5-900  m  elevation  [flowering 
March-August]  A.  chica 

5b.  Leaves  not  drying  reddish  brown  and  lustrous  above  and  below,  usually  conspicuously  puberulent 
on  the  veins  or  on  the  lower  surface;  plants  of  deciduous  and  partly  deciduous  forest  formations 
(also  evergreen  forests  in  A.  candicans),  0-400  m  elevation  6 

6a.  Lower  leaf  surfaces  pale  grayish  with  a  dense  tomentum  of  minute  hairs;  flowering  in  July-Decem- 
ber [fruits  with  the  midvein  and  lateral  edges  usually  prominent;  central  area  of  seed  clearly  de- 
marked  from  the  translucent  wings]  7 

6b.  Lower  leaf  surface  not  pale  grayish  with  a  dense  tomentum,  hairs  0.1-1  mm  long;  flowering  in 

January-August 8 

7a.  Flowering  in  November-December;  seeds  24-34  mm  wide;  collected  in  Costa  Rica  in  partly 

deciduous  and  evergreen  forests  of  the  Pacific  slope    A.  candicans 

7b.  Flowering  in  July-September;  seeds  17-24  mm  wide;  not  known  from  Costa  Rica 

A.  pubescens 

8a.  Surface  of  fruit  puberulent  and  soft  to  the  touch;  calyx  4-6  mm  long  [margin  often  with  minute 
teeth  and  whitish];  flowering  in  December-early  March  A.  mollissima 

8b.  Surface  of  fruit  glabrous  or  minutely  puberulent,  not  soft  to  the  touch;  calyx  4-9  mm  long;  flowering 
in  April-August 9 

9a.  Calyx  cup  with  small  (0.3  mm)  distal  teeth;  stems  and  leaves  with  simple  and/or  branched  hairs; 

fruits  with  the  midvein  not  elevated;  mostly  flowering  in  April-June  (in  Costa  Rica) 

A.  corallina 

9b.  Calyx  cup  with  entire  distal  margin  (or  sometimes  split);  stems  and  leaves  with  simple  hairs;  fruits 
with  the  midvein  usually  elevated;  mostly  flowering  in  June-August A.  conjugate 

Arrabidaea  candicans  (L.  C.  Rich.)  DC.,  Prodr.  Lianas  to  7  cm  diam.,  climbing  with  simple 

9:  185.  1845.  Bignonia  candicans  L.  C.  Rich.,  tendrils    11-21    cm  long,   leafy  stems    1-6  mm 

Act.  Soc.  Hist.  Nat.  Paris  1:  110.  1792.  A.  pa-  diam.,   minutely   puberulent,   becoming  grayish, 

chycalyx  Sprague,  Bull.  Herb.  Boiss.,  ser.  2  6:  glandular  fields  conspicuous,  an  interpetiolar  line 

373.  1906.  A.  rhodothyrsus  Kra'nzl.,  Fedde  Re-  usually  present.  Leaves  with  petioles  45-97  mm 

pert.  17:  20.  1921.  Figure  25.  long,  1-2.3  mm  diam.,  minutely  puberulent,  pet- 

110  FIELDIANA:  BOTANY 


iolules  11-55  mm  long,  0.6-1.4  mm  diam.;  leaflet 
blades  4.5-20  cm  long,  3-12  cm  wide,  ovate  to 
broadly  ovate  or  ovate-elliptic,  apex  acute  to  acu- 
minate, base  obtuse  to  subtruncate  or  slightly 
rounded,  drying  chartaceous,  densely  and  minute- 
ly grayish  puberulent  beneath,  2°  veins  4-7/side, 
strongly  ascending.  Inflorescences  terminal  or  ax- 
illary to  distal  leaves,  20-40  cm  long,  to  30  cm 
wide,  a  pyramidal  thyrse,  peduncles  2-4  mm 
diam.,  densely  minutely  grayish  puberulent,  lat- 
eral branches  2-7  cm  long,  pedicels  1-3  mm  long. 
Flowers  with  calyx  3-5  mm  long,  2-4  mm  diam., 
tubular-conical,  densely  minutely  puberulent, 
margin  entire;  corolla  20-38  mm  long,  6-9  mm 
diam.  distally,  purple  or  rose  (white  within), 
densely  puberulent  externally,  lobes  5-1 1  mm 
long,  7-10  mm  wide,  rounded;  filaments  ca.  16 
and  13  mm  long,  thecae  2-2.5  mm  long.  Fruits 
12-35  cm  long,  7-12  mm  wide,  linear  to  linear- 
oblong,  valves  smooth,  midvein  and  margins 
slightly  elevated;  seeds  7-9  mm  long,  24-34  mm 
wide,  central  dark  area  14-18  mm  wide,  clearly 
differentiated  from  the  translucent  wings. 

Plants  of  evergreen  rain  forest  and  partly  de- 
ciduous forest  formations,  5-900  m  elevation. 
Flowering  November-December,  fruiting  Febru- 
ary-June. Uncommon  in  Costa  Rica,  it  is  known 
only  from  the  Pacific  slope,  from  Tacares  south- 
ward to  Golfo  Dulce.  This  species  ranges  dis- 
junctly  from  Mexico  to  Brazil. 

Arrabidaea  candicans  is  recognized  by  the 
climbing  habit,  simple  tendrils,  two-  or  three-fo- 
liolate  leaves,  large  inflorescences,  puberulent  ca- 
lyx with  entire  margin,  puberulent  rose  to  purple 
corollas,  long  linear  fruits,  and  transparent-winged 
seeds.  The  leaflets  with  grayish  lower  surface  and 
the  gland  fields  conspicuous  below  the  nodes  are 
useful  vegetative  distinctions. 

Arrabidaea  chica  (Humboldt  &  Bonpland)  Ver- 
lot,  Rev.  Hortic.  40:  154.  1868.  Bignonia  chica 
Humboldt  &  Bonpland,  PI.  Aequin.  1:  107,  pi. 
31.  1808.  Figure  25. 

Lianas  to  35  m  high,  stems  to  15  cm  diam., 
tendrils  to  12  cm  long,  leafy  stems  1.5-5  mm 
diam.,  glabrous  or  sparsely  and  minutely  (0.1 
mm)  puberulent,  longitudinally  striate,  nodal 
glandular  fields  visible  on  young  stems.  Leaves 
with  petioles  32-85  mm  long,  0.8-1.9  mm  diam., 
usually  glabrous,  petiolules  (4-)10-32(-46)  mm 
long;  leaflet  blades  (3-)5-16  cm  long,  2-8  cm 
wide,  narrowly  ovate-elliptic  to  elliptic-oblong  or 
elliptic,  acuminate  at  the  apex,  base  obtuse  or 


rounded,  drying  chartaceous  and  usually  reddish 
brown,  usually  glabrous  above  and  below  (in  Cen- 
tral America),  2°  veins  4-7/side,  minor  venation 
raised  on  both  surfaces  (dried).  Inflorescences 
terminal,  often  3-parted  at  a  branchlet  apex,  8-36 
cm  long,  peduncles  4-8  cm  long,  ca.  1.7  mm 
diam.,  very  sparsely  puberulent,  drying  dark, 
proximal  lateral  branches  to  7  cm  long,  pedicels 
4-7  mm  long.  Flowers  with  calyx  3-5  mm  long, 
2-3  mm  diam.,  cupulate,  minutely  papillate  pu- 
berulent, grayish  green,  distally  subentire  with 
small  (0.2  mm)  teeth;  corolla  18-34  mm -long, 
tube  6-1 1  mm  diam.,  lavender  to  purple  (white 
within),  densely  papillate  puberulent  externally, 
lobes  6-12  mm  long,  to  12  mm  wide;  filaments 
ca.  12  and  9  mm  long,  thecae  1.5-2  mm  long. 
Fruits  14-28  cm  long,  9-14  mm  wide,  linear- 
oblong,  valves  flat,  smooth  and  light  to  dark 
brown,  midvein  and  margins  slightly  raised;  seeds 
8-10  mm  long,  23-38  mm  wide,  wings  strongly 
differentiated  and  transparent. 

Uncommon  plants  of  wet  evergreen  rain  forest 
formations  on  both  Caribbean  and  Pacific  coasts 
(rarely  collected  in  moist  sites  in  seasonally  very 
dry  deciduous  formations),  5-900  m  elevation. 
Flowering  in  late  March-August.  This  species 
ranges  from  Mexico  to  Argentina. 

Arrabidaea  chica  is  recognized  by  its  climbing 
habit  with  simple  tendrils,  two-  or  three-foliolate 
leaves,  nearly  entire  cupular  calyx,  pale  purple  co- 
rollas densely  puberulent  externally,  and  narrow 
glabrous  fruits  with  winged  seeds.  The  leaves  are 
usually  glabrous  and  usually  become  reddish 
brown  and  lustrous  when  dried.  These  plants  have 
been  widely  used  as  a  source  of  red  dyes;  such  a 
use  has  been  reported  around  Santa  Ana  (Eche- 
verria  32  &  37  F).  Gentry  (1973b)  remarked  that 
puberulent  forms  of  this  species  intergraded  with 
A.  candicans,  but  in  Central  America  it  may  be 
better  to  assign  all  the  puberulent  forms  to  A.  can- 
dicans. 

Arrabidaea  conjugata  (Veil.)  Mart.,  Flora  24  (2), 
Beibl.  46.  1841.  Bignonia  conjugata  Veil.,  Fl. 
Flum.  245.  1825;  6:  tab.  18.  1827.  Figure  25. 

Lianas  climbing  to  the  tops  of  medium-size 
trees,  to  5  cm  diam.,  leafy  stems  1 .5-5  mm  diam., 
sparsely  very  minutely  puberulent,  glabrescent, 
gland  fields  often  conspicuous;  pseudostipules  ca. 
2  mm  long,  conical.  Leaves  with  petioles  2-13 
cm  long,  0.8-2  mm  diam.,  sparsely  minutely  pu- 
berulent, longitudinally  striate,  petiolules  8-55 
mm  long;  leaflet  blades  5-17(-21)  cm  long,  3.5- 


BURGER:  FLORA  COSTARICENSIS 


111 


10(-13)  cm  wide,  ovate-elliptic  to  ovate  or  ellip- 
tic-obovate,  apex  acuminate,  base  cuneate  to 
rounded,  drying  chartaceous  to  subcoriaceous, 
glabrous  or  with  minute  (0. 1  mm)  hairs  along  the 
veins  beneath  and  usually  with  longer  (0.2-0.5 
mm)  hairs  in  vein  axils  (domatia),  2°  veins  4-77 
side,  strongly  ascending  (subpalmate).  Inflores- 
cences terminal,  14-34  cm  long,  pyramidal  pan- 
icles with  3-5  pairs  of  lateral  branches,  peduncles 
to  10  cm  long,  2.3  mm  diam.,  pedicels  1-4  mm 
long,  densely  puberulent.  Flowers  with  tubular 
calyx  4-7  mm  long,  3-5  mm  diam.,  grayish  with 
a  dense  minute  (0. 1  mm)  puberulence,  margin  en- 
tire or  5-denticulate,  submarginal  glands  present; 
corolla  30-42  mm  long,  7-12  mm  wide  at  the 
mouth,  tubular-campanulate,  magenta  (white 
within),  puberulent  with  short  crooked  hairs,  lobes 
4-11  mm  long,  7-9  mm  wide;  filaments  12-16 
and  8-1 1  mm  long,  thecae  ca.  2  mm  long.  Fruits 
12-31  cm  long,  8-14  mm  wide,  linear-oblong, 
midvein  often  slightly  elevated,  surface  slightly 
muricate;  seeds  7-11  mm  long,  19-27  mm  wide, 
brown  throughout  except  for  transparent  distal  2- 
3  mm. 

Evergreen  lianas  of  the  seasonally  dry  decidu- 
ous forests  of  northwestern  Costa  Rica  (and  in 
partly  deciduous  forests  elsewhere),  5—150  m  el- 
evation. Flowering  in  June-September;  fruiting  in 
the  dry  season  (Gentry,  1973b).  The  short  flow- 
ering season  may  account  for  a  paucity  of  collec- 
tions. This  species  ranges  from  northwestern  Cos- 
ta Rica  to  Brazil. 

Arrabidaea  conjugata  is  recognized  by  its 
climbing  habit  with  simple  tendrils,  two-  or  three- 
foliolate  glabrate  leaves,  terminal  pyramidal  pan- 
icles, densely  puberulent  grayish  calyx  with  sub- 
marginal  glands,  puberulent  magenta  corolla,  and 
long  linear  flattened  fruit  with  two-winged  seeds. 
The  conspicuous  gland  fields  below  the  nodes, 
leaves  with  longer  hairs  in  vein  axils  beneath 
(when  present),  and  raised  minor  venation  also 
help  to  distinguish  this  species. 

Arrabidaea  corallina  (Jacq.)  Sandw.,  Kew  Bull. 
1953:  460.  1954.  Bignonia  corallina  Jacq., 
Fragm.  Bot.  37,  tab.  42,  fig.  1.  1800-1809.  B. 
glabrata  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3: 
137.  1819.  B.  obliqua  Kunth  in  H.B.K.,  Nov. 
Gen.  Sp.  3:  135.  1819.  Figure  25. 

Lianas  to  20  m  high,  stems  to  12  cm  diam., 
tendrils  to  15  cm  long,  leafy  stems  2-4  mm  diam., 
sparsely  to  densely  puberulent  with  simple  or 
branched  hairs  0.2-0.5  mm  long,  gland  fields  pre- 


sent or  absent  at  the  nodes.  Leaves  mostly  trifo- 
liolate,  petioles  2-9  cm  long,  0.7-2  mm  diam., 
usually  densely  puberulent  with  simple  or 
branched  hairs,  petiolules  3-38  mm  long;  leaflet 
blades  5-18  cm  long,  3-9(-12)  cm  wide,  ovate- 
elliptic  to  broadly  rounded-elliptic  or  elliptic-ob- 
ovate,  apex  short-acuminate,  base  narrowed  to 
rounded-truncate,  drying  thin  chartaceous,  green 
or  brown,  minutely  puberulent  on  the  veins  above, 
sparsely  to  densely  puberulent  beneath  with  sim- 
ple or  branched  hairs  0.2-0.8  mm  long,  2°  veins 
4-7/side.  Inflorescences  usually  short-branched 
(raceme-like)  panicles  from  the  axils  of  fallen 
leaves  or  terminal,  8-30  cm  long,  peduncles  2-24 
mm  long,  1.5-2  mm  diam.,  densely  puberulent, 
bracts  1-4  mm  long,  linear,  pedicels  1-12  mm 
long.  Flowers  with  cupulate  calyx  5-9  mm  long, 
4-7  mm  diam.,  sparsely  puberulent  with  multi- 
cellular  hairs  0.1-0.2  mm  long,  margin  subentire 
or  with  narrow  apiculate  teeth  0.3-1  mm  long 
(sometimes  split  and  appearing  lobed);  corolla 
27-46  mm  long,  8-12  mm  wide  at  the  mouth, 
magenta  to  lavender,  puberulent  externally,  lobes 
5-14  mm  long,  to  12  mm  wide;  filaments  ca.  15 
and  10  mm  long,  thecae  2.5-3  mm  long.  Fruits 
12-47  cm  long,  11-21  mm  wide,  valves  flat  and 
smooth  (minutely  pitted),  midvein  apparent  or 
not;  seeds  1 1-14  mm  long,  32-50  mm  wide,  cen- 
tral area  ca.  14  mm  wide,  thin  transparent  edges 
ca.  5  mm  wide. 

Rarely  collected  deciduous  lianas  of  seasonally 
dry  deciduous  forest  formations,  1-300  m  eleva- 
tion. Flowering  in  April-June  (July-November  in 
Panama;  Gentry,  1973b).  The  species  ranges  from 
Mexico  to  Argentina. 

Arrabidaea  corallina  is  recognized  by  its 
climbing  habit,  simple  tendrils,  two-  or  three-fo- 
liolate  leaves  with  usually  dense  pubescence  of 
branched  or  simple  hairs,  puberulent  magenta  or 
lavender  corollas,  long  narrow  fruits,  and  winged 
seeds.  The  calyx  tube  is  usually  sparsely  puberu- 
lent distally  and  of  thin  texture  with  venation  dry- 
ing darker,  and  narrow  teeth  (0.3  mm)  are  usually 
present  along  the  irregular  distal  margin. 

Arrabidaea  costaricensis  (Kranzl)  A.  Gentry, 
Brittonia  25:  231.  1973.  Saldanhaea  costaricen- 
sis Kranzl,  Fedde  Repert.  17:  124.  1921.  Arra- 
bidaea erecta  Miranda,  Anal.  Inst.  Biol.  Mex. 
24:  91.  1953.  Figure  12. 

Shrubs  or  small  trees  to  5  m  tall  (rarely  woody 
vines),  tendrils  absent  or  simple,  leafy  stems  2-5 
mm  diam.,  densely  puberulent  with  straight  or 


112 


FIELDIANA:  BOTANY 


curved  hairs  0.1-0.3  mm  long,  soon  glabrescent 
and  pale  grayish,  gland  fields  absent,  interpetiolar 
lines  present  or  absent.  Leaves  simple  to  3-fo- 
liolate,  petioles  3-55  mm  long,  0.8-1.7  mm 
diam.,  densely  puberulent,  petiolules  5-27  mm 
long  with  central  petiolule  much  longer  than  lat- 
erals; leaflet  blades  3-12  cm  long,  2.5-7.5  cm 
wide,  ovate-elliptic  to  broadly  ovate,  apex  acute 
or  short-acuminate,  base  narrowed  and  obtuse  to 
rounded,  drying  chartaceous  and  grayish  green  or 
brown,  smooth  or  slightly  scabrous  above.  With 
straight  hairs  ca.  0.2  mm  long,  lower  surface 
densely  puberulent,  the  soft  straight  hairs  to  0.3 
mm  long,  venation  subpalmate,  2°  veins  4-6/side, 
strongly  ascending.  Inflorescences  terminal  or  ax- 
illary at  leafless  nodes,  3-7  cm  long,  racemes  with 
fewer  than  10  flowers,  peduncle  5-14  mm  long, 
densely  puberulent,  bracts  caducous  or  to  4  mm 
long,  pedicels  2-5  mm  long.  Flowers  with  calyx 
3-5  mm  long,  3-4  mm  diam.,  cupular,  surface 
grayish  with  dense  minute  puberulence,  marginal 
teeth  5,  0.3-0.7  mm  long,  0.3-0.5  mm  wide  at 
base;  corolla  24-32  mm  long,  5-6  mm  wide  at 
the  mouth,  reddish  purple  (white),  densely  puber- 
ulent with  crooked  multicellular  hairs  externally. 
Fruits  8-21  cm  long,  6-14  mm  wide,  linear-ob- 
long valves  flattened,  with  prominent  midrib, 
brown,  smooth  or  with  pits  or  dots;  seeds  6-12 
mm  long,  18-26  mm  wide,  central  area  13-16 
mm  wide. 

Rarely  collected  shrubs  or  vines  of  seasonally 
very  dry  deciduous  forest  formations  in  northern 
Guanacaste,  10-300  m  elevation  (to  900  m  in 
Honduras).  Flowering  in  May-June;  fruiting  in 
June-January.  This  species  ranges  from  Mexico 
along  the  Pacific  slope  to  northwestern  Costa 
Rica. 

Arrabidaea  costaricensis  is  distinguished  by  its 
often  shrubby  habit  and  usual  lack  of  tendrils 
(sometimes  present),  grayish  indumentum  of 
straight  simple  hairs,  presence  of  both  simple  and 
three-foliolate  leaves,  few-flowered  inflorescenc- 
es, and  linear-oblong  fruits  with  two-winged 
seeds.  The  capsules  are  sometimes  held  upright, 
and  they  may  open  explosively.  The  simple  leaves 
are  usually  borne  near  the  base  of  a  shoot  or  at 
the  initiation  of  a  new  shoot,  with  the  following 
leaves  three-foliolate. 

Arrabidaea  florida  DC.,  Prodr.  9:  184.  1845.  A. 
panamensis  Sprague,  Bull.  Herb.  Boiss.  s£e.  2, 
6:  371.  1906. 


Lianas,  tendrils  to  8  cm  long  (often  lacking  on 
specimens),  leafy  stems  2-5  mm  diam.,  minutely 
(0.05  mm)  puberulent  and  lenticellate,  longitudi- 
nally striate,  interpetiolar  lines  usually  present, 
gland  fields  absent.  Leaves  2-  or  3-foliolate,  pet- 
ioles 10-60  mm  long,  0.8-1.5  mm  diam.,  minute- 
ly puberulent,  petiolules  5-24  mm  long;  leaflet 
blades  5-14  cm  long,  2.5-8  cm  wide,  ovate-ellip- 
tic to  ovate-oblong,  apex  acute  to  acuminate,  base 
acute  to  obtuse  or  rounded,  drying  stiffly  charta- 
ceous and  grayish  green  or  brown,  lustrous  and 
glabrous  above,  minutely  puberulent  beneath,  2° 
veins  4-7/side.  Inflorescences  terminal  or  axil- 
lary to  distal  leaves,  9-35  cm  long,  open  panicles 
to  30  cm  wide,  2°  peduncles  1 .5-7  cm  long,  distal 
branches  grayish  puberulent,  flowers  crowded  in 
distal  clusters,  subtended  by  narrow  bracts  1-4 
mm  long,  pedicels  0-3  mm  long.  Flower  buds 
narrowly  ellipsoid,  calyx  3-4  mm  long,  2-3  mm 
diam.,  minutely  puberulent  with  grayish  hairs, 
margin  entire  or  with  minute  teeth  0.1-0.3  mm 
long;  corolla  12-18  mm  long,  campanulate  with 
spreading  lobes,  pink  or  lavender  (white),  throat 
often  white,  5-7  mm  wide  at  the  mouth,  lobes  3- 
6  mm  long;  filaments  ca.  8  and  6  mm  long,  thecae 
1.5  mm  long.  Fruits  10-26  cm  long,  8-11  mm 
wide,  valves  flat,  brown  with  raised  midrib, 
smooth;  seeds  6-9  mm  long,  17-28(-36)  mm 
wide,  dark  central  area  11-14  mm  wide,  wings 
translucent. 

Rarely  collected  plants  (in  Costa  Rica)  of  moist 
forest  formations,  1-1 100  m  elevation.  Flowering 
in  July-October.  The  species  ranges  disjunctly 
from  Mexico  to  Bolivia  and  Brazil. 

Arrabidaea  florida  is  distinguished  by  its 
climbing  habit  with  simple  tendrils,  generally  gla- 
brous two-  or  three-foliolate  leaves,  large  inflo- 
rescences with  many  distal  clusters  of  small  flow- 
ers, pink  to  lavender  puberulent  corollas,  long 
narrow  fruit,  and  two-winged  seeds.  Sterile  plants 
of  this  species  resemble  Adenocalymma  inunda- 
tum  (cartilaginous  margins  on  leaflets)  and  Tyn- 
nanthus  croatianus  A.  Gentry  of  Panama  (tendrils 
with  trifid  tips). 

Arrabidaea  mollissima  (Kunth  in  H.B.K.)  Bu- 
reau &  K.  Shum.  in  Mart.,  Fl.  Bras.  8  (2):  46. 
1896.  Bignonia  mollissima  Kunth  in  H.B.K., 
Nov.  Gen.  Sp.  3:  133.  1819.  B.  littoralis  Kunth 
in  H.B.K.,  Nov.  Gen.  Sp.  3:  139.  1819.  A.  mol- 
licoma  Blake,  Cont.  Gray  Herb.  52:  92.  1917. 
A.  isthmica  Standl.,  J.  Wash.  Acad.  Sci.  18:  337. 
1925,  pro  parte.  Figure  26. 


BURGER:  FLORA  COSTARICENSIS 


113 


Lianas  (rarely  shrub-like),  stems  to  3  cm  diam., 
tendrils  to  12  cm  long,  leafy  stems  2-5  mm  diam., 
minutely  puberulent  with  thin  straight  hairs,  gla- 
brescent,  terete,  glandular  fields  rarely  present; 
pseudostipules  2-3  mm  long,  conical.  Leaves  de- 
ciduous in  dry  season,  2-  or  3-foliolate,  petioles 
25-80  mm  long,  1.1-2.8  mm  diam.,  densely  pu- 
berulent with  straight  simple  or  gland-tipped  hairs 
0.3-0.8  mm  long,  petiolules  6-35  mm  long;  leaf- 
let blades  4-13  cm  long,  2-10  cm  wide,  broadly 
ovate  to  ovate-orbicular,  apex  caudate-acuminate 
or  short-acuminate,  base  rounded  and  truncate  to 
subcordate,  drying  grayish  and  chartaceous,  upper 
surface  pubescent  with  straight  hairs  0.5-0.8  mm 
long,  lower  surface  densely  puberulent  with  soft 
hairs  0.3-0.7  mm  long,  venation  subpalmate,  2° 
veins  4-6/side.  Inflorescences  terminal  or  axil- 
lary to  leafless  nodes,  3-25  cm  long,  to  18  cm 
wide,  densely  puberulent  throughout,  flowers  in 
distal  groups  of  2-5,  pedicels  3-7  mm  long. 
Flowers  with  cupular  calyx  4-6  mm  long,  3-4 
mm  diam.,  densely  puberulent,  margin  subentire 
or  with  5  minute  (0.3  mm)  teeth;  corolla  26-54 
mm  long,  pink-lavender  to  red-purple  and  white 
in  the  throat  (rarely  white  throughout),  densely 
puberulent  externally,  tube  8-16  mm  diam.  at 
mouth,  lobes  6-12  mm  long;  filaments  ca.  16  and 
12  mm  long,  thecae  4-5  mm  long.  Fruits  15-28 
cm  long,  11-15  mm  wide,  valves  flat,  yellowish 
brown  with  minutely  puberulent  surface  (soft  to 
the  touch),  midvein  apparent  or  not;  seeds  11-13 
mm  long,  30-45  mm  wide,  central  area  ca.  12 
mm  wide,  distal  tranparent  area  of  wings  ca.  6 
mm  wide. 

Common  deciduous  lianas  in  seasonally  very 
dry  deciduous  forest  formations  and  open  vege- 
tation, 2-300  m  elevation.  Flowering  in  December 
to  early  March  (with  the  great  majority  of  flow- 
ering collections  made  in  January);  fruiting  in 
January-June.  This  species  ranges  from  Mexico, 
along  the  Pacific  Coast,  to  Colombia  and  Vene- 
zuela. 

Arrabidaea  mollissima  is  recognized  by  its 
densely  soft  pubescence,  broadly  ovate  or  round- 
ed leaf  blades,  flowering  when  leafless,  puberulent 
purple  or  pink  corollas,  long,  narrow,  softly  pu- 
berulent fruits,  and  two-winged  seeds  transparent 
at  the  lateral  margins.  Simple  tendrils  and  restric- 
tion to  areas  of  deciduous  forest  are  additional 
distinctions.  These  plants  can  be  very  striking 
when  in  full  flower,  after  the  leaves  have  been 
shed.  Compare  this  species  with  A.  corallina  (with 
glabrous  fruits). 


Arrabidaea  patellifera  (Schldl.)  Sandwith,  Kew 
Bull.  22:  413.  1968.  Bignonia  patellifera 
Schldl.,  Linnaea  8:  516.  1833.  Petastoma  pa- 
telliferum  (Schldl.)  Miers,  Proc.  R.  Hort.  Soc. 
3:  195.  1863.  P.  breviflorum  Standl.,  J.  Arnold 
Arbor.  11:  128.  1930.  Figure  26. 

Lianas,  to  over  10  m  high  and  5  cm  diam., 
tendrils  to  16  cm  long,  leafy  stems  2-8  mm  diam., 
densely  puberulent  with  curved  hairs  ca.  0.2  mm 
long  or  subglabrous,  terete,  an  interpetiolar  line 
often  present,  gland  fields  absent  at  the  node; 
pseudostipules  rarely  present  and  leaf-like,  round- 
ed. Leaves  simple  or  2-foliolate,  petioles  12-35 
mm  long  (shorter  in  compound  leaves),  1.2-2.3 
mm  diam.,  densely  puberulent,  petiolules  16-30 
mm  long;  leaflet  blades  (5-)6.5-14(-16)  cm  long, 
(3-)4-8(-12)  cm  wide,  ovate  to  ovate-oblong  or 
ovate-orbicular,  apex  acuminate,  base  narrowed  or 
rounded  and  truncate,  drying  chartaceous,  upper 
surface  with  scattered  straight  hairs  0.2-0.5  mm 
long,  puberulent  beneath  with  thin  straight  hairs 
ca.  0.4  mm  long,  venation  pinnate  to  palmate,  2° 
veins  3-6/side,  ascending.  Inflorescences  axillary 
or  often  terminal  and  3-parted,  6-28  cm  long,  to 
30  cm  wide  near  base,  peduncles  to  8  cm  long,  to 
4  mm  diam.,  densely  puberulent,  pedicels  4-7 
mm  long.  Flower  buds  white  apically,  calyx 
broadly  spreading  (saucer-like),  1-4  mm  long,  4- 
7  mm  wide,  margin  entire  and  undulate,  sparsely 
puberulent  externally;  corolla  23-40  mm  long, 
red-violet  to  magenta  or  lavender  (pink-white  in 
the  throat),  glabrous  externally  except  for  the 
lobes,  narrowed  (2  mm)  base  of  the  tube  6-8  mm 
long,  mouth  8-1 1  mm  wide,  lobes  5-1 1  mm  long, 
ca.  7  mm  wide;  filaments  ca.  12  and  9  mm  long, 
thecae  1.5-2  mm  long.  Fruits  1 1-39  cm  long,  9- 
14  mm  wide,  linear,  surface  of  valves  flat,  smooth, 
midvein  and  margins  slightly  raised;  seeds  8-10 
mm  long,  20-36  mm  wide,  central  area  1 1  mm 
wide,  wings  not  clearly  differentiated. 

Common  evergreen  lianas  in  seasonally  very 
dry  deciduous  and  partly  deciduous  forests  of  the 
Pacific  slope  (rarely  collected  in  evergreen  forests 
of  the  Caribbean  slope),  1-1200  m  elevation. 
Flowering  primarily  in  July-October  with  the  ma- 
jority of  collections  made  in  August;  fruiting  in 
January-March.  This  species  ranges  from  Mexico 
to  Brazil. 

Arrabidaea  patellifera  is  recognized  by  its 
climbing  habit  with  simple  tendrils,  simple  or  bi- 
foliolate  puberulent  leaves,  July-October  flower- 
ing period,  saucer-like  (patelliform)  calyx,  laven- 
der to  magenta  corollas,  and  linear  flattened  fruits 


114 


FIELDIANA:  BOTANY 


with  thin-winged  seeds.  The  frequent  presence  of 
simple  leaves,  strongly  ascending  secondary  veins 
in  a  subpalmate  pattern,  domatia,  and  the  occa- 
sional presence  of  rounded  leaf-like  little  pseu- 
dostipules  are  additional  distinctions.  The  flower 
buds  are  unusual  with  their  pubescent  distal  co- 
rolla lobes  whitish,  in  contrast  to  the  glabrous  ma- 
genta tube. 

Arrabidaea  pubescens  (L.)  A.  Gentry,  Brittonia 
25:  239.  1973.  Bignonia  pubescens  L.,  Sp.  PI., 
ed.  2,  2:  870.  1763.  Petastoma  pubescens  (L.) 
Miers,  Proc.  R.  Hort.  Soc.  3:  195.  1863.  A.  lun- 
dellii  Standl..  Publ.  Field  Mus.  Bot.  8:  48.  1930. 

Lianas  to  20  m  high  and  5  cm  diam.,  tendrils 
to  7  cm  long,  leafy  stems  2-7  mm  diam.,  grayish 
with  minute  (0.05-0. 1  mm)  appressed  hairs,  glan- 
dular fields  usually  present  at  the  nodes;  pseudo- 
stipules  small  or  absent.  Leaves  2-  or  3-foliolate. 
petioles  10-22  mm  long,  0.9-2.3  mm  diam.,  mi- 
nutely appressed  puberulent,  grayish,  petiolules 
12-23  mm  long;  leaflet  blades  4-13  cm  long, 
2.3-8  cm  wide,  ovate  to  ovate-elliptic  or  narrowly 
ovate,  apex  acuminate,  base  obtuse  or  rounded 
and  truncated,  drying  chartaceous  and  grayish  be- 
neath, upper  surface  subglabrous  or  with  minute 
(<  0.1  mm)  hairs,  lower  surface  with  minute  or 
longer  (0.4  mm)  hairs,  2°  veins  4-6  side.  Inflo- 
rescences to  35  cm  long,  open  terminal  or  axillary 
panicles,  peduncles  ca.  6  cm  long,  densely  ap- 
pressed puberulent,  pedicels  3-5  mm  long,  ca.  0.4 
mm  diam.,  grayish  puberulent.  Flowers  with  ca- 
lyx 3-5  mm  long,  2.5-4  mm  diam.,  tubular-cu- 
pulate,  margin  subentire,  grayish  puberulent;  co- 
rolla 18-33  mm  long,  funnelform-campanulate, 
lavender  to  lilac,  minutely  puberulent  externally, 
5-9  mm  wide  at  mouth,  lobes  8-10  mm  long; 
filaments  ca.  10  and  8  mm  long.  Fruits  (7-)  14- 
27  cm  long,  9-1 1  mm  wide,  linear-oblong, 
smooth  or  slightly  pitted,  midrib  and  margins 
raised;  seeds  6-9  mm  long,  17-32  mm  wide,  cen- 
tral area  5-6  mm  wide,  wings  transparent. 

Arrabidaea  pubescens  has  not  been  collected 
from  the  region  between  northeastern  Guatemala 
and  central  Panama,  but  it  may  occur  in  Costa 
Rica.  This  species  is  distinguished  by  the  simple 
tendrils,  nodes  with  gland  fields,  minute  dense  ap- 
pressed hairs  that  give  the  undersides  of  the  leaves 
and  petioles  a  grayish  appearance,  bi-  or  tri-fo- 
liolate  leaves,  and  linear  fruits.  The  smaller  lilac 
corollas  produced  in  July-September,  truncated 
grayish  calyx  tubes,  and  seeds  with  central  brown 
area  clearly  demarcated  from  the  transparent 


wings  are  other  noteworthy  characteristics.  This 
species  prefers  seasonally  dry  lowland  forests 
along  the  Caribbean  coast;  it  ranges  disjunctly 
from  Tamaulipas,  Mexico,  to  Brazil. 

Arrabidaea  verrucosa  (Standl.)  A.  Gentry,  Sel- 
byana  2:  43.  1977.  Adenocalymma  verrucosa 
Standl..  Publ.  Field  Columb.  Mus.,  Bot.  Ser.  4: 
323.  1929.  Martinella  verrucosa  (Standl.) 
Standl.,  Contrib.  Arnold  Arbor.  5:  138.  1933. 
Scobinaria  verrucosa  (Standl.)  Seibert,  Carne- 
gie Inst.  Washington  Publ.  522:  408.  1940.  Fig- 
ure 18. 

Lianas  to  35  m  high,  to  10  cm  diam.,  tendrils 
to  20  cm  long,  leafy  stems  2-7  mm  diam.,  sparse- 
ly and  minutely  (0.1  mm)  puberulent  (more  rarely 
with  hairs  to  1  mm  long),  gland  fields  and  inter- 
petiolar  lines  present  or  absent  at  nodes.  Leaves 
2-foliolate,  petioles  14-23(-55)  mm  long,  0.7-2.3 
mm  diam.,  minutely  papillate  puberulent,  petio- 
lules 12-28(-45)  mm  long;  leaflet  blades  6-17 
(-22)  cm  long,  3-9(-l  1 )  cm  wide,  ovate  to  ovate- 
elliptic,  narrowly  elliptic  or  elliptic-oblong,  apex 
acute  to  acuminate,  base  acute  to  slightly  rounded, 
drying  chartaceous  and  often  dark  brown,  upper 
surface  subglabrous  with  minute  hairs  on  the  mid- 
vein,  lower  surface  with  small  (0.2-0.5  mm) 
straight  hairs  or  branched  along  the  veins,  larger 
hairs  often  present  in  vein  axils  beneath  (domatia), 
venation  pinnate  or  subpalmate,  2°  veins  3-5/side, 
strongly  ascending.  Inflorescences  axillary,  5-15 
cm  long,  few-flowered  panicles,  peduncles  3-5 
cm  long,  sparsely  papillate-puberulent,  lenticel- 
late,  pedicels  6-14  mm  long,  ca.  0.8  mm  diam. 
Flowers  with  tubular-campanulate  calyx,  14-23 
mm  long,  4-1 1  mm  diam.,  glabrous  or  pubescent 
with  hairs  to  0.7  mm  long  near  the  base,  margin 
2-lipped  or  with  shallow  (1  mm)  sinuses  separat- 
ing the  rounded  lobes;  corolla  35-60(-79)  mm 
long,  tubular-campanulate,  lobes  magenta  to  pur- 
ple with  the  tube  white  or  pink,  puberulent  exter- 
nally, mouth  11-20  mm  wide,  lobes  8-18  mm 
long;  filaments  ca.  1 8  and  1 2  mm  long,  thecae  2- 
3  mm  long.  Fruits  ( 1 2-)22-42(-5 1 )  cm  long,  15- 
28  mm  wide,  linear-oblong,  valves  drying  dark 
and  verrucose-tubcrculate,  tubercles  0.3-2  mm 
high;  seeds  15-20  mm  long,  38-57  mm  wide, 
central  area  poorly  delimited. 

Plants  of  evergreen  rain  forest  formations  on 
both  the  Caribbean  and  Pacific  slopes,  1-700 
(-1300)  m  elevation.  Flowering  in  May-Novem- 
ber (with  the  great  majority  of  collections  in  July); 


BURGER:  FLORA  COSTARICENSIS 


115 


fruiting  throughout  the  year.  This  species  ranges 
from  Mexico  to  Bolivia. 

Arrabidaea  verrucosa  is  recognized  by  the 
climbing  habit  with  simple  tendrils,  bifoliolate 
leaves,  small  inflorescences,  long-tubular  calyx, 
puberulent  pink  or  white  and  purple  corollas,  and 
narrow  fruits  with  dark  rasp-like  verrucose-tuber- 
culate  valves.  The  wet-season  flowering  period 
and  wet  forest  habitats  are  also  characteristic.  This 
material  was  placed  under  Scobinaria  japurensis 
(DC.)  Sandwith  by  Gentry  in  Flora  of  Panama 
(1973b),  but  he  later  decided  that  that  name  be- 
longs to  a  species  restricted  to  Amazonian  South 
America  (see:  Selbyana  2:  45,  1977). 


Callichlamys  Miquel 

Lianas  or  shrub-like  when  young,  climbing 
with  simple  tendrils,  stems  terete,  with  4  phloem 
arms  in  cross-section,  an  interpetiolar  line  and 
glandular  fields  absent  at  the  node,  pseudostipules 
absent  (pseudostipule-like  structures  present  at  the 
base  of  axillary  shoots).  Leaves  opposite,  2-  or  3- 
foliolate,  terminal  leaflet  sometimes  replaced  by  a 
tendril,  petiolate,  leaflets  petiolulate,  margins  en- 
tire, venation  pinnate.  Inflorescences  axillary 
(terminal)  racemes,  with  2-12  mostly  opposite 
flowers,  peduncles  becoming  woody,  bracts  sub- 
tending the  pedicels  lanceolate.  Flowers  large,  ca- 
lyx tubular-inflated,  with  spongy  texture,  irregu- 
larly lobed  and  bilabiate;  corolla  tubular-campan- 
ulate,  yellow,  glabrous  externally,  lobes  broadly 
rounded;  stamens  4,  of  2  lengths,  anthers  gla- 
brous, thecae  divaricate,  staminode  absent;  ovary 
2-locular,  ovules  in  4-8  series  on  each  placenta, 
stigma  simple,  flattened.  Fruits  compressed  ob- 
long capsules,  valves  woody  and  smooth,  flat- 
tened parallel  to  the  septum;  seeds  large,  the  2 
thin  lateral  wings  poorly  differentiated  from  the 
central  area. 

Callichlamys  includes  a  single  distinctive  spe- 
cies ranging  from  Mexico  to  Brazil. 

Callichlamys  latifolia  (L.  C.  Rich.)  K.  Schum.  in 
Engler  &  Prantl.,  Nat.  Pflanzenfam.  4  (3b):  223. 
1894.  Bignonia  latifolia  L.  C.  Richard,  Act. 
Soc.  Hist.  Nat.  Paris  1:  110.  1792.  Tabebuia 
latifolia  (L.  C.  Rich.)  DC.,  Rev.  Bign.  (Bibl. 
Univ.  Geneve)  15.  1838.  T.  speciosa  Standl., 
Publ.  Field  Columb.  Mus.,  Bot.  Ser.  8:  49. 
1930.  C.  garnieri  Standl.  &  L.  O.  Williams, 
Ceiba  3:  130.  1952.  Figure  19. 


Lianas  to  over  30  m  high  and  8  cm  diam.  (rare- 
ly shrubs),  leafy  stems  3-9  mm  diam.,  minutely 
puberulent  at  first  but  soon  glabrescent  with  dark 
lenticels  (0.5-1.5  mm)  against  a  grayish  surface. 
Leaves  mostly  2-foliolate  (also  3-foliolate),  peti- 
oles 3-17  cm  long,  1.7-4.5  mm  diam.,  at  first 
minutely  puberulent  but  later  resembling  the 
stems,  petiolules  8-35  mm  long,  sulcate  above; 
leaflet  blades  7-24(-36)  cm  long,  4-14(-20)  cm 
wide,  broadly  elliptic  to  broadly  ovate-elliptic  or 
elliptic-oblong,  apex  acuminate  to  caudate-acu- 
minate, narrowed  tip  to  3  cm  long,  base  obtuse  to 
rounded,  drying  stiffly  chartaceous,  glabrescent 
on  both  surfaces  or  with  small  (0.4-0.5  mm) 
branched/stellate  hairs  beneath,  often  with  dense 
hairs  in  vein  axils  beneath  (domatia),  2°  veins  5- 
1  I/side.  Inflorescences  3-17  cm  long,  peduncles 
4-14  mm  long,  2-3  mm  diam.,  minutely  puber- 
ulent with  appressed  brownish  hairs,  bracts  to  5 
mm  long,  caducous,  pedicels  8-24  mm  long,  ca. 
1  mm  diam.,  drying  dark.  Flowers  with  calyx  28- 
48(-65)  mm  long,  8-30(-42)  mm  diam.,  tubular- 
inflated,  rounded  at  the  base,  greenish  white  to 
yellow,  lobes  12-20  mm  long;  corolla  5-9(-ll) 
cm  long,  bright  yellow,  tube  15-22(-28)  mm  wide 
near  the  mouth,  marked  with  reddish  lines  within, 
lobes  1.5-3  cm  long;  filaments  ca.  15-20  and  25- 
30  mm  long,  thecae  2-3  mm  long;  stigma  ca.  4 
X  1.8  mm.  Fruits  12-32  cm  long,  6-1 1  cm  wide, 
ca.  1  cm  thick,  oblong  or  elliptic-oblong  with 
rounded  ends,  flattened,  surface  smooth;  seeds 
2.5-4  cm  long,  6-1 1  cm  wide,  brownish. 

Plants  of  evergreen  rain  forest  formations  of 
both  Caribbean  and  Pacific  slopes  and  also  in  sea- 
sonally very  dry  deciduous  forest  areas,  5-600 
(-1000)  m  elevation.  Flowering  collections  have 
been  made  in  February-August  in  Costa  Rica 
(flowering  primarily  in  October-November  in 
Panama).  The  species  ranges  from  Veracruz, 
Mexico,  to  Brazil. 

Callichlamys  latifolia  is  recognized  by  its 
climbing  habit  with  simple  tendrils,  large  spongy 
calyx,  large  yellow  corolla,  and  flattened  oblong 
woody  fruits  with  large  brown  seeds.  The  grayish 
stems  with  dark  lenticels,  lack  of  gland  fields  and 
interpetiolar  lines  at  nodes,  and  larger  leaf  blades 
often  with  domatia  in  vein  axils  (or  with  branched 
hairs  beneath)  are  useful  in  identifying  sterile  ma- 
terial. Collections  from  dryer  forests  of  the  Pacific 
slope  tend  to  have  the  undersurface  of  the  leaflets 
covered  with  conspicuous  branched  hairs.  These 
collections  look  quite  different  from  those  of  the 
lowland  rain  forests  with  subglabrous  leaves.  The 
type  of  C.  garnieri  (Gamier  130  F,  from  the  Si- 


116 


FIELDIANA:  BOTANY 


erra  de  Managua,  Nicaragua)  has  leaves  with 
branched  hairs  and  unusually  large  (15  cm)  flow- 
ers subtended  by  reduced  leaves  on  a  terminal 
branchlet.  However,  the  fact  that  no  other  similar 
collection  has  been  seen  suggests  that  Gentry 
(1994)  was  correct  in  considering  it  an  aberrant 
individual  of  this  species. 


Ceratophytum  Pittier 

Lianas,  climbing  with  tendrils,  stems  terete, 
with  4  phloem  areas  in  cross-section,  nodes  with 
interpetiolar  ridges  and  glandular  fields;  pseudo- 
stipules  of  subulate  scales.  Leaves  opposite,  3- 
foliolate  or  2-foliolate  and  the  terminal  leaflet  re- 
placed by  a  tendril  or  tendril  scar,  leaflets  petiol- 
ulate,  margins  entire,  venation  pinnate,  webbed 
tissue  (domatia)  sometimes  present  in  vein  axils. 
Inflorescences  terminal,  corymbose  panicles  or 
condensed  fasicles,  with  1-20  flowers,  peduncles 
lepidote,  bracts  small,  pedicels  well  developed. 
Flowers  with  tubular  calyx,  coriaceous,  apex 
truncated,  with  short-linear  glandular  fields  near 
the  rim;  corolla  tubular-funnelform  and  slightly 
2-lipped,  yellow  to  yellowish  white,  minutely  pu- 
berulent  externally,  5-lobed;  stamens  4,  of  2 
lengths,  thecae  straight,  divaricate,  a  staminode 
present;  disc  cupular;  ovary  2-locular,  ovules  in 
6-8  series  on  each  placenta,  stigma  simple.  Fruits 
oblong-linear  capsules,  tetrangular  in  cross-sec- 
tion, expanded  and  rounded  at  the  base,  tapering 
to  the  apex,  valves  parallel  to  the  septum  (but  not 
flattened),  smooth;  seeds  with  2  lateral  papery 
wings,  brownish,  not  clearly  demarcated  from  the 
central  area. 

Ceratophytum  includes  a  single  species  ranging 
from  Mexico  and  the  Caribbean  to  Guyana  and 
Bolivia. 

Ceratophytum  tetragonolobum  (Jacq.)  Sprague 
&  Sandw.,  Kew  Bull.  1934:  222.  1935.  Bigno- 
nia  tetragonoloba  Jacq.,  Fragm.  Bot.  36,  tab. 
40,  fig.  2.  1800-1806.  Anemopaegma  toba- 
gense  Urb.,  Fedde  Repert.  14:  311.  1916.  Ad- 
enocalymma  heterophyllum  Si  ami  I..  Publ.  Field 
Columb.  Mus.,  Bot.  Ser.  8:  49.  1930,  not 
Kranzl.  C.  tobagense  (Urb.)  Sprague  &  Sandw., 
Kew  Bull.  1933:  322.  1933.  Ad.  standleyanum 
Lundell,  Carnegie  Inst.  Washington  Publ.  478. 
221.  1937,  based  on  Ad.  heterophyllum  Standl. 
Figure  19. 


Lianas  to  20  m  high,  stems  up  to  7  cm  diam.. 
tendrils  1-3  mm  diam.,  to  22  cm  long,  often  trifid, 
leafy  stems  1.3-7  mm  diam.,  subglabrous,  nodes 
with  gland  fields;  pseudostipules  1-3  mm.  Leaves 
2-  or  3-foliolate.  petioles  4-11  cm  long,  1.2-2 
mm  diam..  subglabrous  with  minute  peltate  hairs, 
petiolules  8-30  mm  long;  leaflet  blades  6-18  cm 
long,  3.5-9(-15)  cm  wide,  elliptic-oblong  to 
broadly  elliptic,  apex  short-acuminate  or  obtuse, 
base  obtuse  to  rounded,  drying  thin-chartaceous, 
with  scattered  minute  (0.1  mm)  peltate  hairs  be- 
neath, 2°  veins  3-7/side,  minor  venation  often 
prominent  above,  domatia  occasionally  present. 
Inflorescences  terminal,  1-8  cm  long,  with  2-20 
flowers,  peduncles  1-10  mm  long,  bracts  minute, 
pedicels  8-17  mm  long,  ca.  0.7  mm  diam.,  with 
minute  peltate  hairs.  Flowers  with  calyx  8-12 
mm  long,  5-9  mm  wide,  tubular  with  subentire 
rim,  appearing  glabrous  but  with  minute  peltate 
hairs  and  ciliolate  rim,  with  glands  near  the  mar- 
gin; corolla  5-8  cm  long,  tubular-funnelform  with 
tube  gradually  expanded  to  16-21  mm  wide  at  the 
mouth,  white  to  yellow,  with  a  dense  covering  of 
minute  glandular  hairs  drying  pale  yellowish 
brown,  lobes  15-22  mm  long,  rounded  or  obtuse; 
filaments  10-16  and  16-24  mm  long,  thecae  4  X 
1  mm;  ovary  ca.  5  mm  long,  with  peltate  hairs. 
Fruits  16-45  cm  long.  25-37  mm  wide,  15-26 
mm  thick,  linear-oblong,  narrowed  to  the  apex, 
yellowish  and  lenticellate;  seeds  8-13  mm  long, 
28-45  mm  wide,  lateral  wings  brown  or  gray,  not 
clearly  differentiated  from  central  area,  translu- 
cent. 

Uncommon  evergreen  climbers  of  deciduous, 
partly  deciduous,  or  wet  evergreen  lowland  rain 
forest  formations,  1-500  m  elevation.  Flowering 
in  January-June  and  October  in  Central  America; 
fruiting  throughout  the  year.  This  species  ranges 
from  Mexico  to  Bolivia. 

Ceratophytum  tetragonolobum  is  recognized  by 
its  climbing  habit,  three-tipped  tendrils,  bi-  or  tri- 
foliolatc  opposite  leaves,  compact  terminal  inflo- 
rescences, truncated  calyx  tube  with  distal  glands, 
white  or  yellowish  corolla  minutely  pubcrulent  on 
the  exterior,  thick  narrow  woody  fruits,  and  seeds 
with  poorly  differentiated  lateral  wings.  Young 
stems  with  prominent  glandular  fields  and  usually 
four  subulate  pseudostipules  at  the  nodes  and  oc- 
casional domatia  are  additional  characteristics. 


Clytostoma  Miers  ex  Bureau 

Lianas  climbing  with  simple  tendrils,  stems  4- 
angled  or  terete,  with  8  phloem  areas  in  cross- 


BURGER:  FLORA  COSTARICENSIS 


117 


section,  nodes  lacking  gland  fields;  pseudostipules 
usually  a  small  condensed  axis  of  subulate  cata- 
phylls.  Leaves  2-foliolate  (rarely  simple),  with  or 
without  a  terminal  simple  tendril,  petiolate,  mar- 
gins entire,  venation  pinnate.  Inflorescences  ter- 
minal or  axillary,  few-flowered  fascicles,  cymes, 
panicles,  or  the  flower  solitary,  subtended  by 
bracts  similar  to  the  cataphylls.  Flowers  with  cu- 
pular  or  campanulate  calyx  truncated  distally  and 
minutely  5-denticulate  or  with  5  linear  lobes, 
subglabrous  to  minutely  lepidote  or  puberulent; 
corolla  tubular-funnelform,  white  to  purple,  glan- 
dular-puberulent  or  with  minute  peltate  hairs  ex- 
ternally; stamens  4,  included,  filaments  of  2 
lengths,  anthers  glabrous,  thecae  straight  and  di- 
varicate, a  staminode  present;  disc  absent;  ovary 
2-locular,  ovules  2(-4)-seriate  in  each  locule. 


Fruits  ellipsoid  to  suborbicular  capsules,  valves 
convex  or  flattened  parallel  with  the  septum, 
woody  and  echinate  with  curved  spines;  seeds 
transverse-oblong,  corky,  flattened  but  not 
winged. 

Clytostoma  includes  nine  South  American  spe- 
cies and  one  that  ranges  from  Mexico  to  Brazil. 
Echinate  woody  fruits  and  flowers  lacking  a  nec- 
tariferous disc  are  distinctive  characteristics.  Cly- 
tostoma callistegioides  (Cham.)  Bureau  ex  Gri- 
sebach  of  southern  South  America  was  collected 
in  a  Costa  Rican  garden  in  1935  (Brenes  &  Ze- 
ledon  99;  Standley,  1938),  but  we  have  seen  no 
further  evidence  of  its  presence  in  Central  Amer- 
ica. It  differs  from  all  our  other  species  of  Big- 
noniaceae  in  Costa  Rica  by  having  prominent  lin- 
ear calyx  lobes  arising  from  a  truncated  calyx 
margin  (see  key  below). 


Key  to  the  Species  of  Clytostoma 

la.  Larger  leaf  blades  >  12  cm  long;  calyx  lobes  0.2-0.6  mm  long;  native    C.  binatum 

Ib.  Larger  leaf  blades  <  10  cm  long;  calyx  lobes  1-4  mm  long,  subulate  to  linear;  garden  ornamental 
(not  included  in  descriptions,  see  above)    C.  callistegioides 


Clytostoma  binatum  (Thunb.)  Sandw.,  Recueil 
Trav.  Bot.  Neerl.  34:  235.  1937.  Bignonia  bi- 
nata  Thunb.,  PI.  Bras.  3:  35.  1821.  Adenoca- 
lymma  ocositensis  J.  D.  Smith,  Bot.  Gaz.  18: 
209.  1893.  C.  isthmicum  Pittier,  Contrib.  U.S. 
Natl.  Herb.  18:  257,  pi.  106.  1917.  Petastoma 
ocositense  (J.  D.  Smith)  Kranzl.,  Fedde  Repert. 
17:  61.  1935.  Figure  18. 

Lianas  to  5  cm  diam.,  sometimes  forming 
clumps  1-2  m  high,  tendrils  to  12  cm  long,  leafy 
stems  2-7  mm  diam.,  rounded-tetrangular,  gla- 
brous or  very  minutely  (0.05  mm)  puberulent, 
lenticellate;  pseudostipules  forming  an  axillary 
cone  3-5  mm  long.  Leaves  2-foliolate,  petioles 
6-18(-31)  mm  long,  1-1.5  mm  diam.,  petiolules 
4-6(-21)  mm  long,  usually  glabrous;  leaflet 
blades  7-16(-19)  cm  long,  2-6(-8)  cm  wide,  el- 
liptic to  elliptic-oblong  or  oblong,  apex  acumi- 
nate, base  acute  to  cuneate,  drying  chartaceous 
and  with  minor  venation  raised  on  both  surfaces, 
usually  glabrous  on  both  surfaces,  2°  veins  5-87 
side.  Inflorescences  mostly  axillary,  0.4-8  cm 
long,  peduncle  subtended  by  subulate  bracts  (sim- 
ilar to  the  pseudostiples),  pedicels  6-32  mm  long, 
0.4-0.8  mm  diam.,  glabrous.  Flowers  with  cu- 
pular  calyx  4-8  mm  long,  3-5  mm  diam.,  gla- 
brous, with  many  longitudinal  veins  or  smooth, 


truncate  or  with  teeth  0.2-0.6  mm  long;  corolla 
45-80  mm  long,  lilac  to  blue-purple  (white), 
sparsely  and  minutely  (0.05  mm)  puberulent  ex- 
ternally, tube  12-24  mm  diam.  at  the  mouth, 
throat  white  internally,  lobes  6-16  mm  long;  fil- 
aments ca.  10-16  and  16-20  mm  long,  thecae  ca. 
3  mm  long;  ovary  covered  by  thick  rounded  hairs. 
Fruits  5-9  cm  long,  3-6  cm  wide,  broadly  elliptic 
to  suborbicular,  discoid-lenticular,  surfaces  of  the 
valves  covered  with  echinate  spines  to  8  mm  long, 
slender-tipped  and  slightly  curved  from  a  broad 
base;  seeds  12-19  mm  long,  19-24  mm  wide, 
suborbicular,  brown. 

Evergreen  or  deciduous  lianas  found  mostly  in 
swamp  forests,  river  edges,  and  seasonally  inun- 
dated areas  in  both  wet  evergreen  and  dry  decid- 
uous forest  formations,  1-200  m  elevation.  Flow- 
ering throughout  the  year  but  most  frequently  in 
September-November.  This  species  ranges  from 
Mexico  to  Brazil. 

Clytostoma  binatum  is  recognized  by  its  climb- 
ing habit  with  simple  tendrils,  bifoliolate  leaves, 
short  few-flowered  inflorescences,  essentially  gla- 
brous calyx  cups  with  minute  teeth  distally,  lilac 
to  blue-purple  (rarely  white)  corolla  sparsely  mi- 
nutely puberulent  on  the  exterior,  short  rounded 
fruit  covered  with  woody  echinate  projections, 


118 


FIELDIANA:  BOTANY 


and  wingless  seeds.  The  pseudostipule  resembles 
a  minute  bromeliad  plantlet  in  leaf  axils  and  is 
helpful  in  recognizing  this  species.  The  seeds  are 
apparently  adapted  to  water  dispersal;  conse- 
quently, the  plants  are  restricted  to  low  elevations. 


Crescentia  Linnaeus 

Small  to  medium-size  trees,  with  many  thick 
twisted  branches,  leafy  stems  becoming  terete 
with  enlarged  nodes,  gland  Melds  absent.  Leaves 
alternate  or  more  often  in  fascicles  of  3-9  on  con- 
densed short-shoots  in  the  axils  of  fallen  leaves, 
simple  or  3-foliolate  (1-  or  2-foliolate),  petioles  of 
compound  leaves  winged  and  leaf-like,  blades 
with  entire  margins,  subglabrous  or  sparsely  pu- 
berulent,  venation  pinnate.  Inflorescences  borne 
on  older  thick  branches  or  trunks,  peduncles  ab- 
sent, flowers  1-3  in  sessile  fascicles,  pedicels 
short  and  thick.  Flowers  with  large  tubular  calyx 
that  usually  splits  into  2  halves  at  maturity,  thick, 
glabrous  or  minutely  puberulent;  corolla  broadly 
campanulate  and  bilabiate,  narrowed  only  at  the 
base,  thick  textured,  usually  with  minute  peltate 
hairs  on  the  outer  surface,  tube  with  a  transverse 


fold  midway  across  the  lower  side,  distally  5- 
lobed  or  variously  erose  along  the  margin;  sta- 
mens 4,  subexserted,  filaments  of  2  lengths,  disc 
large,  staminode  present;  ovary  lepidote,  1-locular 
with  ovules  multiseriate  on  4  parietal  placentas, 
stigma  flat  and  expanded.  Fruits  large  pepos  (a 
gourd-like  calabash)  with  a  hard  corky  pericarp, 
subglobose  to  ovoid,  pulpy  within;  seeds  angular, 
without  wings,  less  than  10  mm  long,  embedded 
in  the  pulp. 

Crescentia  is  a  genus  of  six  species  in  Mexico, 
the  West  Indies,  Central  America,  and  (one  spe- 
cies) Amazonia.  The  genus  is  distinguished  by  the 
tree  habit,  mostly  fasciculate  leaves  on  alternate 
short-shoots,  few  large  thick-textured  cauliflorous 
flowers,  ovaries  with  a  single  locule  and  parietal 
placentation,  and  rounded  indehiscent  fruits  with 
seeds  embedded  in  pulp.  The  flowers  are  foul- 
smelling  and  adapted  for  bat  pollination.  The 
pulp-containing  fruits  are  dispersed  by  mammals. 
The  hard  outer  shell  of  the  fruits  has  been  used 
to  make  cups,  ladies,  and  bowls  (see  Standley  & 
Williams,  1974).  The  differentiation  of  the  two 
Central  American  species  is  not  complete,  and  hy- 
brids have  been  reported.  (See  Flora  Neotropica 
treatment;  Gentry,  1980.) 


Key  to  the  Species  of  Crescentia 

la.  Stems  with  3-foliolate  leaves,  simple  leaves  may  also  be  present;  fruits  up  to  10  cm  diam.  and 
globose;  wild  plants  (rarely  cultivated) C.  alata 

Ib.  Stems  lacking  3-foliolate  leaves,  all  the  leaves  simple;  fruits  >  13  cm  diam.  and  globose  to  ovoid; 
plants  widely  cultivated C.  cujete 


Crescentia  alata  Kunth  in  H.B.K.,  Nov.  Gen.  Sp. 
3:  158.  1819.  Parmentiera  alata  (Kunth  in 
H.B.K.)  Miers,  Trans.  Linn.  Soc.  26:  166.  1868. 
C.  ternata  Sesse  &  Moc.,  La  Naturaleza,  ser.  2. 
1  (append.);  94.  1889.  Figure  11. 

Trees  to  10  m  tall,  trunks  to  25  cm  diam.,  leafy 
stems  3-18  mm  diam.,  glabrous,  the  alternate 
nodes  becoming  enlarged  and  prominent  and  sub- 
tending the  condensed  leafy  short-shoots.  Leaves 
mostly  in  fascicles  of  3-9,  simple  or  3-foliolate 
(rarely  1-  or  2-foliolate),  simple  leaves  sessile, 
compound  leaves  with  winged  petioles  2.5-1 1  cm 
long,  4-11  mm  wide,  petiolules  absent;  leaf 
blades  4-15  cm  long,  5-35  mm  wide,  oblanceo- 
late  to  narrowly  obovate  (small  leaves  often 
rounded),  leaflet  blades  1.5-10  cm  long,  4-23  mm 
wide,  oblanceolate,  apex  rounded  or  emarginate. 


gradually  narrowed  to  the  acute  or  cuneate  base, 
drying  subcoriaceous  and  grayish  green,  glabrous. 
Inflorescences  of  1-3  flowers  borne  directly  on 
the  surfaces  of  larger  stems  and  trunks,  pedicels 
5-10  mm  long,  1.3  mm  diam.,  subtended  by  small 
(1.5  mm)  bracts,  glabrous  or  with  minute  peltate 
hairs.  Flowers  with  unpleasant  aroma,  calyx  14- 
28  mm  long,  9-14  mm  diam.,  usually  splitting 
into  2  subequal  halves,  glabrous  or  with  few  mi- 
nute peltate  hairs,  greenish;  corolla  4-6  cm  long, 
tube  narrowed  only  near  the  base,  22-30  mm 
diam..  greenish  purple  to  reddish  white,  surface 
glabrous  or  with  minute  peltate  glands,  5-lobed  or 
the  margin  irregularly  toothed;  filaments  ca.  30 
and  1 8  mm  long,  thecae  5-6  mm  long.  Fruits  7- 
10  cm  diam.,  globose,  greenish,  surface  smooth; 
seeds  6-7  mm  long,  7-9  mm  wide. 

Plants  of  seasonally  dry  deciduous  forest  for- 


BURGER:  FLORA  COSTARICENSIS 


119 


inations  of  the  Pacific  lowlands,  5-800  m  eleva- 
tion. Flowering  throughout  the  year  (collected 
mostly  in  January-September).  This  species  rang- 
es from  Mexico  along  the  Pacific  slope  to  north- 
western Costa  Rica  (introduced  in  Panama). 

Crescentia  alata  is  recognized  by  the  mix  of 
simple  and  trifoliolate  leaves  with  all  the  larger 
blades  being  oblanceolate  or  narrowly  obovate, 
the  few  flowers  coming  directly  from  the  trunk, 
thick  calyx  usually  splitting  in  two,  and  large  glo- 
bose gourd-like  fruits  borne  from  large  branches 
and  trunks.  The  trifoliolate  leaves,  with  sessile  ob- 
lanceolate leaflets  and  winged  petiole,  resemble  a 
Christian  cross  and  were  mentioned  in  early  Span- 
ish accounts  of  Mexico  (Standley  &  Williams, 
1974,  p.  187).  The  trees  are  noteworthy  for  their 
short  compact  form  and  dense  branching,  often 
contrasting  with  the  flat,  open,  grazed  land  around 
them.  The  dense  branching  is  often  the  habitat  for 
many  epiphytes.  There  may  be  hybridization  be- 
tween this  species  and  the  widely  planted  C.  cu- 
jete.  This  species  is  called  morro  or  jicaro,  but 
jicaro  is  also  the  name  of  the  following  species. 

Crescentia  cujete  L.,  Sp.  PI.  2:  626.  1753.  Figure 
11. 

Trees  to  10  m  tall,  with  dense  rounded  crowns, 
trunks  to  40  cm  diam.,  branches  often  crooked 
with  mostly  thick  branchlets,  leafy  stems  6-30 
mm  diam.,  glabrous,  nodes  becoming  thickened 
and  subtending  the  leafy  short-shoots.  Leaves 
usually  fasciculate  on  elevated  alternate  nodes,  1- 
15  and  varying  in  size  at  the  same  node,  unifoli- 
olate,  sessile,  glabrous;  leaf  blades  3-22(-26)  cm 
long,  l-6(-7.5)  cm  wide,  oblanceolate  to  narrow- 
ly obovate,  apex  acute  to  acuminate  or  rounded 
(emarginate),  gradually  narrowing  to  the  acute  or 
cuneate  base,  usually  drying  subcoriaceous  and 
pale  gray,  glabrous  or  with  minute  simple  or 
branched  hairs  along  the  midvein  beneath,  2° 
veins  5-14/side.  Inflorescences  of  1  or  2  flowers 
borne  on  older  branches  and  trunks,  pedicels  9- 
30  mm  long,  0.7-1.7  mm  diam.,  glabrous  or  with 
few  minute  peltate  hairs.  Flowers  with  calyx  18- 
34  mm  long,  splitting  into  2  subequal  lips  12-24 
mm  wide;  corolla  55-70  mm  long,  dull  white  to 
yellowish  white,  with  purple  lines,  tube  18-25 
mm  diam.,  lobes  triangular  with  narrow  tip,  sub- 
glabrous  or  with  minute  peltate  hairs  externally; 
filaments  ca.  30  and  10  mm  long,  anthers  6-9  mm 
long,  slightly  divergent.  Fruits  13-25  cm  diam. 
or  to  30  cm  long,  subglobose  or  ovoid,  surface 


smooth,  subglabrous;  seeds  7-8  mm  long,  4-6 
mm  wide. 

Widely  cultivated  plants  in  regions  of  both  de- 
ciduous and  wet  evergreen  forest  formations;  1- 
1200  m  elevation.  Flowering  irregularly  through- 
out the  year.  This  species  may  have  been  indige- 
nous to  southeastern  Mexico,  but  it  is  now  culti- 
vated throughout  tropical  America,  and  its 
original  range  is  unknown. 

Crescentia  cujete  is  recognized  by  its  fascicles 
of  narrow  simple  leaves  from  alternate  thickened 
nodes,  oblanceolate  blades,  few  flowers  borne  di- 
rectly on  larger  stems  and  trunks,  large  calyx  usu- 
ally splitting  into  two,  large  thick  white  to  yellow- 
ish corolla,  and  larger  globose  gourd-like  fruit 
(calabash).  The  hard  shell  of  the  pericarp  is  often 
used  as  a  cup,  ladle,  or  other  kitchen  utensil;  the 
round,  dried,  empty  fruits  are  often  incised  or 
painted  for  ornament.  The  pulp  of  the  fruit  has 
been  used  as  a  purgative,  according  to  Standley 
(1938),  who  reported  that  the  oval  fruits  are  called 
guacales  and  the  globose  ones  jicaras.  The  wood 
is  hard  but  not  durable.  Jicaro,  calabacero,  cala- 
bash, and  wild  calabash  are  common  names. 


Cydista  Miers 

REFERENCE — W.  D.  Hauk,  A  review  of  the  ge- 
nus Cydista.  Ann.  Missouri  Bot.  Gard.  84:  815- 
840.  1997  (1998). 

Lianas  climbing  with  simple  tendrils,  stems 
terete  to  tetrangular,  with  8-16  phloem  areas  in 
cross-section,  nodes  usually  with  interpetiolar 
lines,  gland  fields  absent  at  the  nodes;  pseudos- 
tipules  absent  or  leaf-like.  Leaves  opposite,  sim- 
ple or  2-(4-)  foliolate,  petiolate,  margins  entire, 
venation  pinnate,  glands  or  gland  fields  often 
present  in  basal  vein  axils  beneath.  Inflores- 
cences terminal  or  axillary,  cymose  racemes  or 
panicles  (thyrses),  often  with  fewer  than  15  flow- 
ers, bracts  minute  or  absent,  flowers  pedicellate. 
Flowers  with  cupular  or  campanulate  calyx,  gla- 
brous to  glandular  puberulent  or  with  minute  pel- 
tate hairs,  glands  present  or  absent  distally,  mar- 
gin truncate  and  entire  to  5-lobed  or  irregularly 
split;  corolla  funnelform  to  tubular-campanulate, 
2-lipped,  lavender  to  purple  or  white,  usually 
with  minute  glandular  or  peltate  hairs  externally, 
with  2  upper  and  3  lower  lobes,  broadly  rounded; 
stamens  4,  included,  filaments  of  2  lengths,  an- 
thers glabrous,  thecae  divaricate;  disc  absent; 
ovary  with  glandular  or  peltate  hairs  on  the  sur- 


120 


FIELDIANA:  BOTANY 


face,  2-locular,  ovules  in  2  series  in  each  locule, 
stigma  2-lobed.  Fruits  linear  to  narrowly  oblong 
woody  capsules,  valves  2,  flattened  parallel  to 
the  septum,  septicidally  dehiscent,  surface  usu- 
ally smooth,  glabrate  to  puberulent;  seeds  flat 
and  transversely  oblong,  the  central  body  of  the 
seed  not  clearly  differentiated  from  the  2  lateral 
wings. 

Cydista  is  a  Neotropical  genus  of  six  species, 
ranging  from  Mexico  and  the  West  Indies  to  Bra- 
zil and  Bolivia.  Five  species  have  been  found  in 


Costa  Rica,  but  two  are  known  from  only  single 
collections.  The  mix  of  simple  and  bifoliolate 
leaves,  presence  of  glands  or  gland  fields  in  basal 
vein  axils  (on  underside  of  leaf),  narrow  capsules, 
and  brown  two-winged  seeds  with  central  area  not 
clearly  differentiated  help  distinguish  the  genus. 
The  absence  of  a  disc  is  consistent  with  reports 
that  the  flowers  do  not  produce  nectar  and  deceive 
pollinators  by  multiple  "big-bang"  flowering  of  a 
few  days.  Compare  species  of  Arrabidaea  and 
Clytostoma. 


Key  to  the  Species  of  Cydista 

la.  Fruits  linear  or  linear-oblong,  10-25  mm  wide,  to  45  cm  long;  commonly  encountered  plants  in 

areas  of  dry  deciduous,  partly  deciduous,  or  (less  often)  evergreen  forests    2 

Ib.  Fruits  narrowly  oblong,  27-43  mm  wide,  to  24  cm  long;  more  rarely  encountered  plants  of  evergreen 

or  partly  deciduous  forests    4 

2a.  Stems  with  persisting  rounded  pseudostipules  3-15  mm  long  [leaf- venation  mostly  palmate; 

flowering  mostly  in  June-August;  fruits  with  thin  flat  valves]   C.  diversifolia 

2b.  Pseudostipules  usually  absent  or  not  broadly  ovate 3 

3a.  Plants  usually  flowering  in  April-May  while  leafless,  in  dry  deciduous  forest  formations;  calyx 
usually  split  into  2  or  3  lobes;  fruits  with  2  longitudinal  ridges  on  the  flat  valve  surface;  leaves 
simple  or  2-foliolate,  leaf  venation  mostly  palmate-subpalmate,  lower  leaf  surfaces  never  dense- 
ly hirtellous C.  heterophylla 

3b.  Plants  flowering  throughout  the  year  with  leaves  fully  expanded,  in  both  deciduous  and  ever- 
green habitats;  calyx  usually  entire  distally;  fruits  smooth  and  flat  on  the  valve  surfaces;  leaves 
usually  only  2-foliolate,  venation  pinnate  to  subpalmate;  leaf  surfaces  densely  hirtellous  to 

pilose  or  subglabrous  and  lustrous C.  aequinoclialis 

4a.  Valves  of  fruit  with  smooth  surface;  lower  leaf  surface  without  glands  or  gland  fields  in  basal  vein 

angles;  leaflet  blades  mostly  2-5  cm  wide;  Mexico  to  Costa  Rica  C.  potosina 

4b.  Valves  of  fruit  with  longitudinally  wrinkled  surface;  lower  leaf  surface  usually  with  glands  in  basal 
vein  angles;  leaflet  blades  mostly  5-15  cm  wide;  Costa  Rica  to  Bolivia    C.  lilacina 


Cydista  aequinoctialis  (L.)  Miers,  Proc.  R.  Hort. 
Soc.  3:  191.  1863.  Bignonia  aequinoctialis  L., 
Sp.  PI.  2:  623.  1753.  B.  sarmentosa  Bertol.,  Fl. 
Guatimal.  25.  1840.  B.  sarmentosa  var.  hirtella 
Benth.,  Bot.  voy.  Sulphur  128.  1845.  C.  sar- 
mentosa (Bertol.)  Miers,  Proc.  R.  Hort.  Soc.  3: 
192.  1863.  Levya  nicaraguensis  Bureau  ex 
Baillon,  Hist.  PI.  10:  29.  1888.  Arrabidaea  gua- 
temalensis  K.  Schum.  &  Loes.,  Bot.  Jahrb.  Syst. 
23:  129.  1896.  Arrabidaea  pseudochica 
Kra'nzl.,  Repert.  Spec.  Nov.  Regni  Veg.  17:  19. 
1921.  Anemopaegma  tonduzianum  Kra'nzl., 
Fedde  Repert.  17:  116.  1921.  Cydista  pubescens 
Blake,  Contrib.  U.S.  Natl.  Herb.  24:  23.  1922. 
Arrabidaea  isthmica  Standl.,  J.  Wash.  Acad. 
Sci.  15:  461.  1925,  pro  parte  (leaves).  C.  ae- 
quinoctialis var.  hirtella  (Benth.)  A.  Gentry, 
Ann.  Missouri  Bot.  Card.  60:  838.  1973.  Figure 
24. 


Lianas  to  over  20  m  high,  to  8  cm  diam.,  ten- 
drils 4-21  cm  long,  leafy  stems  2-7  cm  diam., 
glabrous  to  sparsely  or  densely  pilose  with 
straight  hairs  to  1  mm  long  (in  var.  hirtella), 
young  stems  tetragonal;  pseudostipules  usually  in- 
conspicuous (leaf-like  to  4  mm  long),  caducous. 
Leaves  simple  or  2-foliolate,  petioles  1 1-47  mm 
long,  1 .2-2  mm  diam.,  petiolules  9-40  mm  long, 
glabrous  to  sparsely  or  densely  pilose  (var.  hirtel- 
la); leaflet  blades  6-16  cm  long,  3-10  cm  wide, 
ovate-elliptic  to  elliptic,  apex  acuminate,  base  ob- 
tuse or  rounded  and  subtruncate,  drying  stiffly 
chartaceous,  glabrous  to  sparsely  puberulent 
above,  minutely  puberulent  on  the  veins  to  dense- 
ly pubescent  beneath  (in  var.  hirtella),  2°  veins  4- 
7/side,  basal  veins  often  strongly  ascending  (sub- 
palmate).  Inflorescences  terminal  or  axillary, 
short  (3-15  cm)  panicles  of  3-15  flowers  (flowers 


BURGER:  FLORA  COSTARICENSIS 


121 


rarely  solitary),  peduncles  1.5-10  cm  long, 
sparsely  to  densely  puberulent,  pedicels  5-25  mm 
long.  Flowers  with  calyx  4-9  mm  long,  4-7  mm 
wide,  cupular,  margin  entire  or  with  minute  teeth, 
surface  glabrous  to  densely  puberulent  (in  var. 
hirtella),  sometimes  with  gland  fields  near  the  dis- 
tal margin;  corolla  (25-)35-75  mm  long,  funnel- 
form-campanulate,  magenta  to  pink  or  white, 
throat  often  yellowish  or  with  purplish  lines  with- 
in, minutely  lepidote  externally,  tube  (6-)  11 -22 
mm  diam.  at  the  mouth,  lobes  12-24  mm  long, 
14-25  mm  wide;  filaments  11-18  and  8-13  mm 
long,  thecae  3-5  mm  long.  Fruits  21-45  cm  long, 
17-24  mm  wide,  linear-oblong,  ends  rounded, 
valves  flat  with  slightly  raised  submarginal  ridges, 
drying  dark  or  blackish,  smooth;  seeds  (8-)  12-20 
mm  long,  (26-)41-70  mm  wide,  pale  brown 
throughout,  sometimes  transparent  at  the  tips. 

Common  evergreen  lianas  of  lowland  wet  ev- 
ergreen forest  areas,  partly  deciduous  forests,  and 
seasonally  very  dry  deciduous  formations,  0-800 
m  elevation.  Flowering  throughout  the  year  but 
with  a  peak  in  April-May;  fruiting  mostly  in  the 
dry  season  (January-May).  This  species  ranges 
from  Mexico  and  the  West  Indies  to  Brazil,  Bo- 
livia, and  Paraguay. 

Cydista  aequinoctialis  is  recognized  by  the 
climbing  habit  with  simple  tendrils,  opposite  sim- 
ple or  bifoliolate  leaves,  short,  few-flowered  inflo- 
rescences, usually  entire  calyx  cups,  pink  to  ma- 
genta or  white  corollas  minutely  lepidote  on  the 
exterior,  long  narrow  flat  fruits,  and  seeds  with 
uniform  brownish  coloring.  These  plants  are  usu- 
ally leafless  when  flowering  and  fruiting  (Gentry, 
1973a).  An  important  identifying  characteristic  is 
that  gland  fields  are  sometimes  present  in  the  axils 
of  secondary  veins  on  the  lower  leaf  surface.  This 
species  is  extremely  variable  in  leaflet  form,  flow- 
er size,  and  vesture;  the  more  densely  pubescent 
individuals  (formerly  called  C.  sarmentosa  or  C. 
pubescens)  have  been  designated  var.  hirtella 
(Bentham)  A.  Gentry.  The  more  puberulent  vari- 
ety is  usually  encountered  in  dry  vegetation,  and 
the  glabrous  variety  is  more  common  in  evergreen 
formations. 

Cydista  diversifolia  (Kunth  in  H.B.K.)  Miers, 
Proc.  R.  Hort.  Soc.  3:  192.  1863.  Bignonia  di- 
versifolia Kunth  in  H.B.K.,  Nov.  Gen.  Sp. 
quart,  ed.  3:  133,  folio  ed.  3:  104.  1819.  Pleon- 
otoma  diversifolium  (Kunth  in  H.B.K.)  Bureau 
&  K.  Schum.  in  Mart.,  Fl.  Bras.  8(2):  274. 
1897.  Figure  24. 


Lianas  to  5  cm  diam,  to  20  m  high,  tendrils 
11-21  cm  long,  leafy  stems  2-7  mm  diam., 
strongly  tetragonal  with  4  longitudinal  ridges, 
subglabrous  or  sparsely  pubescent  with  thin  hairs 
to  0.4  mm  long;  pseudostipules  3-15  mm  long, 
leaf-like,  rounded-ovate.  Leaves  2-foliolate  or 
sometimes  simple,  petioles  18-45  mm  long,  ca.  1 
mm  diam.,  surface  similar  to  stems,  petiolules  10- 
31  mm  long;  leaflet  blades  4-13  cm  long,  2.5-8 
cm  wide,  ovate  to  broadly  ovate,  apex  acuminate 
with  tip  to  13  mm  long,  base  rounded  and  sub- 
cordate  to  shallow-cordate,  drying  chartaceous, 
upper  surface  subglabrous,  lower  surface  glabrous 
to  sparsely  pubescent  with  thin  hairs  ca.  0.4  mm 
long,  venation  palmate  with  3  major  veins  or  sub- 
palmate  with  3  or  4  2°  veins/side,  gland  fields  of- 
ten present  in  the  basal  vein  axils  beneath.  Inflo- 
rescences terminal  or  axillary,  6-20  cm  long, 
panicles  with  4-25  flowers,  peduncles  to  3-9  cm 
long,  ca.  1.5  mm  diam.,  similar  to  stems,  lateral 
branches  8-25  mm  long,  bracts  0.4-1  mm  long, 
pedicels  4-12  mm  long.  Flowers  with  calyx  cups 
3.5-5  mm  long,  ca.  4  mm  diam.,  entire  or  split 
along  the  margin,  with  minute  peltate  hairs  or  thin 
hairs  along  the  margin;  corolla  22-46  mm  long, 
funnelform-campanulate,  magenta  to  lavender  or 
bluish  purple,  subglabrous  externally,  tube  8-16 
mm  diam.  at  the  mouth,  throat  white  with  purple 
stripes  within,  lobes  7-22  mm  long,  broadly 
rounded;  filaments  ca.  16  and  13  mm  long,  thecae 
2-2.5  mm  long.  Fruits  27-41  cm  long,  10-16 
mm  wide,  linear,  valves  flat  and  smooth,  dark 
brown;  seeds  10-13  mm  long,  28-57  mm  wide, 
pale  yellowish  brown  throughout,  wings  thin. 

Common  evergreen  vines  in  seasonally  dry  de- 
ciduous formations  (rarely  in  evergreen  forests), 
5-500  m  elevation.  Flowering  throughout  the  year 
but  most  often  collected  in  June-August;  fruiting 
in  November-March.  This  species  ranges  from 
Mexico  and  the  West  Indies  to  Colombia  and  Ven- 
ezuela. 

Cydista  diversifolia  is  recognized  by  its  climb- 
ing habit  with  simple  tendrils,  usually  bifoliolate 
leaves  with  rounded  subcordate  base  and  palmate 
venation,  open-paniculate  inflorescences,  wet-sea- 
son flowering,  bright  lavender  to  bluish  purple  co- 
rollas, long  narrow  flattened  fruits,  and  uniformly 
colored  thin  seeds.  The  persisting  rounded  pseu- 
dostipules (four  per  node),  square  stems  with 
prominent  ridges  and  hollow  in  early  stages,  and 
gland  fields  in  the  basal  vein  axils  are  additional 
vegetative  features. 

Cydista  heterophylla  Seibert,  Carnegie  Inst. 
Wash.  Publ.  522:  417.  1940.  Bignonia  lepidota 


122 


FIELDIANA:  BOTANY 


Seem.,  Bot.  voy.  Herald  179.  1854,  non  Kunth 
in  H.B.K.  Figure  24. 

Lianas  to  10  m  high,  to  10  cm  diam.,  tendrils 
to  18  cm  long,  leafy  stems  1.8-5  mm  diam.,  te- 
rete, glabrescent,  nodes  with  or  without  interpe- 
tiolar  lines;  pseudostipules  1-2  mm  long,  conical. 
Leaves  simple  or  2-foliolate  (with  or  without  ten- 
drils), petioles  15-50  mm  long,  glabrous  or 
sparsely  puberulent  with  minute  peltate  hairs,  lon- 
gitudinally striate,  petiolules  5-50  mm  long;  leaf- 
let blades  5-14(-17)  cm  long,  3-9(-l  1)  cm  wide, 
broadly  to  narrowly  ovate,  apex  bluntly  obtuse  to 
short-acuminate,  base  rounded  and  truncate  to 
subcordate,  drying  thin-chartaceous,  subglabrous 
on  both  surfaces,  venation  palmate  with  3  major 
veins  or  subpalmate,  2°  veins  3-5/side,  glandular 
fields  usually  present  in  basal  vein  axils  beneath. 
Inflorescences  mostly  axillary  to  fallen  leaves, 
1.5-7(-14)  cm  long,  racemose,  peduncles  8-14 
mm  long,  0.7-1.8  mm  diam.,  densely  covered 
with  peltate  (lepidote)  hairs,  pedicels  6-13  mm 
long,  vesture  like  the  peduncle.  Flowers  with  ca- 
lyx 3.5-6  mm  long,  4-5  mm  diam.,  densely  cov- 
ered with  peltate  hairs  (glands  sometimes  pres- 
ent), margin  entire  or  split  into  several  lobes;  co- 
rolla 47-66  mm  long,  funnelform-campanulate, 
lavender  to  purple-magenta,  glabrous  or  with  mi- 
nute peltate  hairs  externally,  tube  11-17  mm  wide 
at  the  mouth,  lobes  9-20  mm  long,  to  22  mm 
wide;  filaments  ca.  18  and  12  mm  long,  thecae  4- 
5  mm  long.  Fruits  18-33  cm  long,  14-25  mm 
wide,  linear-oblong,  with  2  raised  longitudinal 
ridges  and  raised  edges  on  each  flattened  valve, 
smooth;  seeds  8-13  mm  long,  34-64  mm  long, 
wings  with  transparent  tips. 

Deciduous  lianas  of  the  seasonally  very  dry  de- 
ciduous forest  formations  and  drier  areas  within 
evergreen  formations,  5-300  m  elevation  (to  1000 
m  in  Honduras).  Flowering  in  April-May,  fruiting 
in  November-March.  This  species  ranges  from 
Mexico  to  northern  Colombia. 

Cydista  heterophylla  is  recognized  by  its  climb- 
ing habit  with  simple  tendrils,  the  simple  or  bi- 
foliolate  opposite  leaves,  short  racemose  inflores- 
cences with  vesture  of  minute  peltate  hairs,  lav- 
ender or  purple  corollas,  long  narrow  fruit  with 
two  central  longitudinal  ridges  and  raised  edges, 
and  winged  seeds  with  transparent  distal  tips. 
Stem  tips  of  young  plants  often  have  four  leaves 
(two  pairs  of  simple  leaves).  The  gland  fields  in 
the  axils  of  basal  veins  may  be  difficult  to  see. 
The  short  flowering  period  when  leaves  are  usu- 
ally absent  and  restriction  to  deciduous  forest  ar- 
eas are  additional  characteristics. 


Cydista  lilacina  A.  Gentry,  Mem.  New  York  Bot. 
Card.  29:  277.  1978.  Figure  24. 

Lianas  climbing  with  simple  tendrils  (not 
seen),  leafy  stems  ca.  4  mm  diam.,  terete,  gla- 
brous or  with  few  minute  hairs,  drying  black; 
pseudostipules  1-2  mm  long.  Leaves  opposite  or 
subopposite,  2-foliolate  or  simple,  petioles  17-50 
mm  long,  1.4-3.3  mm  diam.,  subglabrous,  drying 
dark,  petiolules  9-23  mm  long;  leaflet  blades  5- 
19(-27)  cm  long,  3-13(-17)  cm  wide,  ovate  to 
ovate-oblong,  apex  obtuse  with  a  small  acutp  tip, 
base  obtuse  to  rounded  and  often  unequal,  drying 
stiffly  chartaceous  and  dark  brown,  subglabrous 
above  and  below,  venation  pinnate,  2°  veins  4-67 
side,  glands  usually  present  at  the  basal  vein  axils 
beneath.  Inflorescences  terminal,  ca.  23  cm  long, 
panicles  with  well-separated  opposite  lateral 
branches  to  4  cm  long,  peduncle  4-9  cm  long,  2- 
3  mm  diam.,  glabrous,  drying  dark,  flowers  in  dis- 
tal groups  of  2  or  3,  pedicels  5-14  mm  long. 
Flowers  with  calyx  4-7  mm  long,  cupular  with  5 
prominent  (0.5-1.2  mm)  teeth  or  entire,  subgla- 
brous but  with  rounded  discoid  flat  glands;  co- 
rolla 40-55  mm  long,  tubular-funnelform,  lav- 
ender with  a  white  throat,  tube  10-15  mm  diam. 
at  mouth,  glabrous  or  appressed  puberulent  exter- 
nally, lobes  to  20  mm  long;  filaments  ca.  20  and 
13  mm  long,  thecae  3-4  mm  long;  ovules  4-se- 
riate.  Fruits  14-32  cm  long,  3-4  cm  wide,  ob- 
long, valves  flattened,  margins  rounded,  midvein 
flat,  surface  wrinkled;  seeds  15-21  mm  long,  35- 
58  mm  wide,  brownish. 

Plants  of  lowland  evergreen  rain  forest  forma- 
tions of  the  Pacific  slope,  10-100  m  elevation.  A 
collection  from  Reserva  Biol6gica  Carara,  flow- 
ering in  early  February  (Ziiniga  90),  is  the  only 
record  known  from  Central  America.  Otherwise, 
this  species  ranges  from  northern  South  America 
to  Brazil  and  Bolivia. 

Cydista  lilacina  is  recognized  by  its  climbing 
habit,  simple  or  bifoliolate  leaves,  terminal  pani- 
cle, calyx  with  round  flat  glands,  and  oblong  fruits 
with  flattened  wrinkled  surface.  The  tendency  to 
dry  dark  (especially  stems  and  inflorescence 
axes),  general  lack  of  puberulence,  and  lowland 
wet  forest  habitat  are  additional  characteristics. 

Cydista  potosina  (Schum.  &  Loes.)  Loes.,  Re- 
pert.  Spec.  Nov.  Regni  Veg.  16:  209.  1919.  Ar- 
rabidaea  potosina  Schum.  &  Loes.,  Bull.  Herb. 
Boissier  3:  618.  Clytostoma  mayanum  Si  ami  I.. 
Carnegie  Inst.  Wash.  Publ.  461.  86.  1935.  Fig- 
ure 19. 


BURGER:  FLORA  COSTARICENSIS 


123 


Lianas,  tendrils  to  15  cm  long,  slender,  leafy 
stems  1.7-5  mm  diam.,  glabrous  or  with  minute 
peltate  hairs,  tetragonal  with  4  prominent  longi- 
tudinal ridges,  interpetiolar  ridge  usually  present; 
pseudostipules  linear  or  triangular,  1-8  mm. 
Leaves  2-foliolate,  petioles  12-37  mm  long,  1- 
1.5  mm  diam.,  sparsely  puberulent,  striate,  petio- 
lules  9-40  mm  long;  leaflet  blades  4.5-15  cm 
long,  2-8  cm  wide,  ovate  to  ovate-elliptic  or 
ovate-oblonge,  apex  acute  to  acuminate,  base  ob- 
tuse or  rounded,  drying  chartaceous  and  brown, 
upper  surface  glabrous,  lower  surface  glabrous  or 
with  short  (0.2-0.4  mm)  thin  hairs  along  the 
veins,  venation  pinnate,  2°  veins  3-6/side.  Inflo- 
rescences axillary  or  terminal,  2-10  cm  long,  ra- 
cemes with  2-9  flowers,  peduncles  5-85  mm 
long,  glabrous  or  with  minute  (0.05  mm)  peltate 
hairs,  pedicels  5-18  mm  long.  Flowers  with  calyx 
5-7  mm  long,  4-6  mm  diam.,  narrow-cupular, 
drying  dark,  subglabrous,  sometimes  with  round- 
ed flat  glands  distally,  margin  entire  with  5  tri- 
angular minute  lobes  or  irregular  with  pale  edge; 
corolla  43-57  mm  long,  funnelform-campanulate, 
lavender  or  white,  with  minute  glandular  hairs  ex- 
ternally, tube  1 1-16  mm  diam.  at  the  mouth,  lobes 
9-16  mm  long,  broadly  rounded;  filaments  ca.  17 
and  20  mm  long,  thecae  ca.  3  mm  long.  Fruits 
17-24  cm  long,  27-43  mm  wide,  narrowly  ob- 
long, narrowed  at  the  apex,  valves  flat  with  the 
margins  elevated,  midvein  flat  or  elevated,  surface 
smooth  and  dark;  seeds  14-23  mm  long,  45-70 
mm  wide,  brownish  but  transparent  at  the  wing 
tips. 

Plants  of  wet  lowland  evergreen  and  partly  de- 
ciduous forest  formations  on  both  Caribbean  and 
Pacific  slopes,  5-400  m  elevation  (to  800  m  in 
northern  Central  America).  Flowering  in  May- 
August.  This  species  ranges  from  Veracruz,  Mex- 
ico, to  Costa  Rica  but  is  rarely  collected  along  the 
Pacific  slope  of  Central  America. 

Cydista  potosina  is  recognized  by  its  climbing 
habit  with  simple  tendrils,  bifoliolate  leaves, 
short,  few-flowered  racemes,  lavender  or  white 
corollas,  and  narrowly  oblong  fruits  with  winged 
seeds.  The  lack  of  gland  fields  in  the  axils  of  basal 
veins  (lower  leaf  surface),  consistently  pinnate  ve- 
nation, and  elevated  ridges  on  leafy  stems  help 
distinguish  this  species  from  some  of  its  conge- 
ners. 

Dendrosicus  species  are  now  placed  in  Amphi- 
tecna. 


Distictella  Kuntze 

Lianas  (shrubs),  climbing  with  distally  coiled 
and  trifid  tendrils,  stems  with  4  phloem  areas  in 
cross-section,  branchlets  terete,  nodes  lacking  in- 
terpetiolar lines  and  glandular  fields;  pseudostip- 
ules short  and  inconspicuous.  Leaves  opposite, 
petiolate,  3-foliolate  or  2-foliolate  with  terminal 
tendril,  petiolules  well  developed,  margins  entire, 
venation  pinnate,  domatia  absent,  gland  fields  of- 
ten present  in  the  basal  vein  axils  beneath.  Inflo- 
rescences terminal  or  axillary,  racemes  or  race- 
mose panicles  with  short  lateral  branches,  rachis 
minutely  puberulent,  bracts  small  and  caducous. 
Flowers  with  cupulate  calyx,  minutely  puberu- 
lent, margin  truncated  and  subentire,  glandular 
fields  often  present  near  the  edge;  corolla  tubular- 
campanulate,  white,  minutely  and  densely  puber- 
ulent externally,  lobes  5;  stamens  4,  of  2  lengths, 
anthers  glabrous,  thecae  straight  and  divaricate,  a 
staminode  present;  disc  annular;  ovary  2-locular, 
ovules  in  4-8  series  on  each  placenta,  stigma  sim- 
ple. Fruits  laterally  compressed  or  biconvex  cap- 
sules, the  2  valves  flattened  parallel  to  the  septum, 
thick  and  woody;  seeds  flat,  woody  or  with  thin 
wings  on  2  lateral  sides  and  distally,  usually 
brown. 

Distictella  is  a  genus  of  13  South  American 
species  and  one  species  that  ranges  from  Costa 
Rica  to  Brazil. 

Distictella  magnoliifolia  (Kunth  in  H.B.K.) 
Sandw.,  Lilloa  3:  460.  1938.  Bignonia  magno- 
liaefolia  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3: 
136.  1819.  Figure  21. 

Lianas  to  30  m  high  and  8  cm  diam.,  tendrils 
to  18  cm  long,  0.6-2.3  mm  diam.,  leafy  stems  3- 
8  mm  diam.,  terete,  minutely  (0.05-0.1  mm)  pu- 
berulent with  reddish  brown  hairs.  Leaves  2-fo- 
liolate, petioles  8-55  mm  long,  1.3-2.7  diam., 
petiolules  6-23  mm  long,  often  thickened  below 
the  blade,  minutely  puberulent;  leaf  blades  8-27 
cm  long,  4-12  cm  wide,  elliptic  to  elliptic-oblong 
or  ovate-elliptic,  apex  short-acuminate,  base  ob- 
tuse to  cuneate,  drying  chartaceous,  glabrous 
above,  minutely  puberulent  on  the  veins  beneath, 
2°  veins  5-8/side,  glands  often  present  in  basal 
vein  axils.  Inflorescences  terminal  or  axillary, 
10-28  cm  long,  racemose  panicles,  rachis  densely 
minutely  puberulent,  brownish,  2°  peduncles  (lat- 
eral branches)  6-15  mm  long,  1.5-2  mm  diam., 
bracts  2-4  mm  long,  pedicels  4-8  mm  long. 
Flowers  with  calyx  9-13  mm  long,  8-11  diam., 


124 


FIELDIANA:  BOTANY 


cupular  with  entire  margin,  minutely  papillate  pu- 
berulent,  longitudinal  gland  fields  2-3  mm  long 
often  present  distally;  corolla  42-65(-73)  mm 
long,  30-45  mm  wide  at  the  lobes,  white  with 
yellow  throat,  densely  minutely  glandular  puber- 
ulent  externally,  tube  10-17  mm  diam.  above  the 
narrowed  (3  mm)  basal  portion,  lobes  10-18  mm 
long,  rounded  distally  and  narrowed  near  the  base; 
filaments  ca.  22  and  16  mm  long.  Fruits  7.5-23 
cm  long,  32-55  mm  wide,  15-20  mm  thick,  nar- 
rowly oblong  to  elliptic-oblong,  valves  woody, 
surface  smooth;  seeds  19-24  mm  long,  28-38 
mm  wide,  oblong  to  suborbicular,  brown,  body  of 
seed  14-17  mm  wide,  not  differentiated. 

Rarely  collected  plants  of  lowland  Caribbean 
evergreen  rain  forest  formations  ca.  100  m  ele- 
vation, fruiting  in  November.  The  species  ranges 
from  northeastern  Costa  Rica  (Morales  et  al.  3212 
from  La  Selva)  to  Peru  and  Brazil. 

Distictella  magnoliifolia  is  recognized  by  its 
climbing  habit  with  distally  trifid  tendrils,  oppo- 
site bifoliolate  leaves,  terminal  raceme-like  inflo- 
rescences, minutely  puberulent  calyx  cups  with 
entire  margins  and  distinct  glandular  areas,  large 
white  corollas,  woody  oblong  fruit,  and  flat  brown 
seeds  in  which  the  opaque  wings  extend  around 
three  sides  of  the  central  area.  The  above  descrip- 
tion is  based  on  South  American  material. 

Gibsoniothamnus  is  now  placed  in  the  Schle- 
geliaceae  (q.v.). 


Godmania  Hemsley 

Small  to  medium-size  trees,  bark  smooth  to 
longitudinally  ridged,  interpetiolar  line  and  glan- 
dular field  absent  at  the  nodes.  Leaves  opposite, 
long-petiolate,  palmately  5-9  foliolate,  basal  leaf- 
lets smaller  than  distal,  leaflets  petiolulate,  mar- 
gins entire  (serrate  on  juvenile  shoots),  venation 
pinnate.  Inflorescences  terminal,  compact  many- 
branched  corymbose  panicles  with  many  flowers, 
bracts  small,  puberulent,  pedicels  well  developed. 
Flowers  with  small  (<  2  mm)  broadly  campanu- 
late  calyx,  subentire  or  with  5  small  lobes;  corolla 
urceolate-campanulate  and  slightly  2-lipped,  yel- 
low and  brownish,  puberulent  externally,  lower 
lip  3-lobed,  lobes  valvate  in  bud  and  triangular; 
stamens  4,  of  2  lengths,  included,  filaments  and 
anthers  pubescent,  thecae  divaricate,  a  staminode 
present;  disc  annular-pulvinate;  ovary  minutely 
puberulent,  2-locular,  ovules  multiseriate  in  each 
locule.  Fruits  linear-cylindric  capsules,  twisted, 


dehiscing  loculicidally  (perpendicular  to  the  sep- 
tum), valves  coriaceous;  seeds  with  2  lateral 
membranaceous  wings. 

Godmania  is  a  Neotropical  genus  of  two  spe- 
cies characterized  by  the  smaller  yellowish  flow- 
ers with  valvate  lobes,  twisted  linear  fruits,  and 
palmately  compound  opposite  leaves.  A  second 
species  occurs  in  the  caaiinga  formations  of  Bra- 
zil. The  genus  is  a  member  of  tribe  Tecomeae  and 
related  to  Tabebuia  (Gentry,  1992). 

Godmania  aesculifolia  (Kunth  in  H.B.K.)  Stand). 
in  Standl.  &  Calder6n,  Lista  Prelim.  PI.  El  Sal- 
vador 200.  1925.  Bignonia  aesculifolia  Kunth 
HI  H.B.K.,  Nov.  Gen.  Sp.  3:  140.  1819.  Tecoma 
fuscata  Mocino  ex  DC.,  Prodr.  9:  221.  1845. 
Cybislax  macrocarpa  Benth.  in  Benth.  & 
Hook.,  Gen.  PI.  2:  1043.  1876.  G.  macrocarpa 
(Benth.)  Hemsl.,  Diag.  PI.  Nov.  Mex.  2:  35. 
1879.  Figure  12. 

Trees  or  shrubs,  3-7(-20)  m  tall,  leafy  stems 
3-8  mm  diam.,  minutely  (0.1-0.2  mm)  puberu- 
lent, brown,  terete,  becoming  grayish  and  lenti- 
cellate.  Leaves  1 1-38  cm  long,  with  5,  7,  or  9 
leaflets,  petioles  4.5-18  cm  long,  2-3  mm  diam., 
petiolules  2-24  mm  long,  usually  densely  puber- 
ulent; leaflet  blades  2.5-16  cm  long,  1.5-7  cm 
wide,  elliptic-obovate,  obovate,  narrowly  ellip- 
tic-oblong or  oblanceolate,  apex  short  caudate- 
acuminate  to  gradually  acuminate,  margin  entire 
(with  teeth  on  juvenile  shoots),  base  acute  to  cu- 
neate,  drying  stiffly  chartaceous,  sparsely  to 
densely  puberulent  with  thin  straight  hairs  0.1- 
0.3  mm  long  beneath,  punctate  on  both  surfaces, 
2°  veins  6-13/side,  loop-connected  distally.  In- 
florescences terminal,  5-10  cm  long,  6-10  cm 
wide,  often  a  flat-topped  panicle,  1°  peduncle  2- 
9  mm  long,  2°  peduncles  12-15  mm  long,  dense- 
ly puberulent,  bracts  ca.  0.7  mm  long,  pedicels 
2-9  mm  long,  ca.  0.5  mm  diam.  Flowers  with 
calyx  1-2  mm  long,  2-3  mm  wide,  broadly  ob- 
conic  or  campanulate,  minutely  puberulent,  sub- 
entire  or  5-lobed  (lobes  ca.  0.2  mm  and  obscure); 
corolla  12-16  mm  long,  urceolate  to  campanu- 
late and  slightly  2-lipped,  yellow  with  red-brown 
marking  on  the  upper  lobes,  tube  5-7  mm  diam. 
for  most  of  its  length,  densely  minutely  puberu- 
lent externally,  lobes  1-5  mm  long;  filaments  ca. 
9  and  7  mm  long,  thecae  less  than  1  mm  long; 
style  ca.  9  mm  long.  Fruits  28-60(-100)  cm 
long,  10-14  mm  wide,  8-14  mm  thick,  long-lin- 
ear, somewhat  flattened  and  becoming  coiled  or 
twisted,  with  ca.  6  prominent  longitudinal  ridg- 


BURGER:  FLORA  COSTARICENSIS 


125 


es  on  each  face;  seeds  7-11  mm  long,  50-135 
mm  wide,  central  area  well  differentiated,  6-8 
mm  long,  10-12  mm  wide,  wings  translucent. 

Infrequent  deciduous  trees  of  deciduous  and 
partly  deciduous  forests  of  the  Meseta  Central 
and  Pacific  slope,  5-1200  m  elevation.  Flower- 
ing in  February-July;  fruiting  in  November- 
March.  This  species  ranges  from  Mexico  to  Bo- 
livia. 

Godmania  aesculifolia  is  recognized  by  the 
tree  habit,  opposite  palmately  compound  leaves 
with  three  to  nine  leaflets,  the  short,  many- 
branched  terminal  inflorescences  with  small 
crowded  flowers,  short  calyx,  and  campanulate 
or  urceolate  yellow  corolla.  The  long  linear  spi- 
rally coiling  fruits  and  seeds  with  clearly  differ- 
entiated translucent  membranaceous  lateral 
wings  are  additional  characteristics.  The  branch- 
es often  have  an  unpleasant  odor  when  broken. 


Jacaranda  Jussieu 

Trees  (shrubs  in  Brazil),  stems  quadrangular  or 
terete,  nodes  without  gland  fields  or  interpetiolar 
lines;  pseudostipules  absent.  Leaves  opposite, 
usually  bipinnate  (pinnate  to  simple  in  Brazilian 
spp.),  often  large,  petiolate,  lateral  pinnae  oppo- 
site or  alternate,  leaflets  sessile  or  subsessile,  often 
strongly  asymmetric  with  1  side  broader  than  the 
other,  margin  entire  or  with  distal  teeth,  base  usu- 
ally asymmetric,  glabrous  or  puberulent,  venation 
pinnate.  Inflorescences  terminal,  axillary  or  from 
older  leafless  nodes,  panicles  or  thyrses  (racemes) 
with  many  flowers,  pedunculate,  lateral  branches 


usually  opposite  and  not  crowded,  pedicels  sub- 
tended by  enlarged  ebracteate  nodes.  Flowers 
showy,  calyx  broadly  cupulate  to  tubular-campan- 
ulate,  short,  5-lobed  to  denticulate  or  subentire; 
corolla  tubular-campanulate  or  campanulate-fun- 
nelform  above  a  narrowed  base,  slightly  bilabiate, 
blue  or  lavender  to  magenta  (white),  densely  pu- 
berulent to  glabrous  externally,  the  tube  contract- 
ed near  the  base;  stamens  4,  filaments  of  2 
lengths,  anthers  glabrous,  unequal  or  with  only  1 
theca,  staminode  usually  exceeding  the  stamens 
with  glandular  hairs  along  its  length  and  at  the 
apex,  pollen  3-colpate;  disc  pulviniform,  ovary 
glabrous  or  puberulent,  2-locular,  ovules  on  ca.  8 
series  in  each  locule.  Fruits  capsules,  oblong  or 
rounded  in  outline,  strongly  flattened  perpendic- 
ular to  the  septum  and  dehiscing  parallel  to  the 
plane  of  compression,  surfaces  glabrous  or  lepi- 
dote;  seeds  thin,  flat,  with  translucent  or  brown 
circumferential  wings  (wider  in  width  than  in 
length),  seed  area  and  membranaceous  wing  dif- 
ferentiated or  not. 

Jacaranda  is  a  genus  of  49  species  ranging 
from  Mexico  to  Argentina  (Gentry  &  Morawetz 
in  Gentry,  1992),  belonging  to  the  tribe  Teco- 
meae.  The  beauty  of  the  trees  when  in  flower  has 
made  a  few  of  the  species  ornamental  favorites, 
now  planted  throughout  the  tropics  and  subtrop- 
ics.  The  genus  stands  out  within  the  family  be- 
cause of  its  tree  habit,  opposite  bipinnate  leaves 
with  usually  many  small  leaflets,  bluish  or  lav- 
ender flowers,  and  flat,  rounded  woody  fruits.  The 
long  staminode,  often  adorned  with  glandular 
hairs,  is  another  important  generic  distinction  (it 
can  be  mistaken  for  a  style). 


Key  to  the  Species  of  Jacaranda 

la.  Calyx  4-7  mm  long  and  tubular;  leaflets  20-80  mm  long,  6-25  mm  wide  [corolla  densely  puberulent 
externally;  fruits  with  flat  rounded  margin;  trees  to  45  m  tall;  wet  forests  of  both  the  Caribbean  and 
Pacific  slopes]  J.  copaia 

Ib.  Calyx  1-3  mm  long  and  cupular  or  saucer-like;  leaflets  6-24  mm  long,  1-10  mm  wide    2 

2a.  Corolla  mostly  glabrous  externally;  leaflets  3.5-10  mm  wide;  fruits  5-12  cm  long,  oblong  or  rounded 
with  strongly  undulating  lateral  margins;  growing  wild  in  evergreen  forests  of  the  Pacific  slope  .  . 
J.  caucana 

2b.  Corolla  minutely  puberulent  externally;  leaflets  1-4  mm  wide;  fruits  4-7  cm  long,  suborbicular 
with  flattened  or  slightly  undulating  margins,  widely  planted  for  ornament  in  evergreen  and  partly 
deciduous  areas  J.  mimosifolia 


126 


FIELDIANA:  BOTANY 


Jacaranda  caucana  Pittier,  Contrib.  U.S.  Natl. 
Herb.  18:  258.  1917.  J.  ficifolia  D.  Don,  sec 
Seem.,  Bot.  Voy.  Herald  181.  154,  non  Don.  J. 
trianae  Kranzl.,  Fedde  Repert.  17:  226.  1921. 
J.  caucana  ssp.  sandwithiana  A.  Gentry,  Ann. 
Missouri  Bot.  Card.  60:  858.  1973  (1974).  Fig- 
ure 14. 

Trees  8-28  m  tall,  trunks  up  to  1  m  diam., 
leafy  stems  3-12  mm  diam.,  glabrous  or  with  few 
minute  peltate  hairs,  young  stems  drying  dark 
with  narrow  whitish  lenticels,  older  stems  gray. 
Leaves  bipinnate,  20-45  cm  long,  petioles  4-8 
cm  long,  1.8-4  mm  diam.,  sparsely  minutely  pu- 
berulent,  rachis  sulcate  above,  with  lateral  wings 
ca.  0.5  mm  wide,  lateral  pinnae  7-18  pairs,  4-17 
cm  long,  petiolules  3-10  mm  long;  leaflet  blades 
8-28  mm  long,  3.5-10  mm  wide  (terminal  leaflets 
to  37  X  16  mm),  asymmetrically  oblong-rhombic, 
apex  obtuse  with  a  small  apiculate  tip,  base  with 
narrowly  cuneate  and  broadly  obtuse  sides,  drying 
chartaceous,  glabrous  above  with  major  veins  im- 
pressed, grayish  beneath  with  thin  hairs  0.2-0.4 
mm  long  2°  veins  4-7/side.  Inflorescences  usu- 
ally borne  at  distal  leafless  nodes,  6-15  cm  long, 
paniculate  with  3-30  flowers,  peduncles  3-25  mm 
long,  0.7-2  mm  diam.,  glabrous  or  sparsely  pu- 
berulent,  drying  dark,  pedicels  0.5-3  mm  long 
(ca.  6  mm  including  subtending  node).  Flowers 
with  calyx  ca.  1 .5-3  mm  long,  1 .8-4  mm  wide  at 
the  apex,  broadly  campanulate,  sparsely  papillate 
puberulent,  bluntly  5-toothed;  corolla  35-48  mm 
long,  tubular-campanulate  and  often  bent  almost 
90°  near  the  base,  purple-blue,  subglabrous  exter- 
nally, tube  10-17  mm  diam.  near  the  mouth,  lobes 
ca.  10  X  12  mm;  filaments  ca.  14  and  13  mm 
long,  thecae  1.5-2  mm  long,  staminode  22-25 
mm  long  with  hairs  to  1  mm  long.  Fruits  6-12 
cm  long,  4-7  cm  wide,  oblong-elliptic  (narrowed 
at  base  and  apex),  10-17  mm  thick,  margins  thin- 
ner and  undulating  (2  lobes/side);  seeds  8-19  mm 
long,  23-42  mm  wide,  wings  brown  or  slightly 
translucent,  not  clearly  differentiated. 

Trees  of  evergreen  forest  formations  of  the 
southern  Pacific  slope  (in  Costa  Rica),  5-1000  m 
elevation.  Flowering  in  March-May;  fruiting  in 
November-March.  The  species  ranges  from 
southwestern  Costa  Rica  to  Venezuela. 

Jacaranda  caucana  is  recognized  by  its  tree 
habit,  opposite  bipinnate  leaves  with  small  asym- 
metric leaflets,  flowers  with  small,  widely  flaring 
calyx  cups,  purple-blue  corollas  mostly  glabrous 
on  the  exterior,  and  the  flat  woody  fruits  with  un- 
dulating margin.  Costa  Rican  material  is  placed  in 


subspecies  sandwithiana,  having  shorter  or  un- 
developed calyx  lobes  and  a  corolla  tube  that  is 
glandular-pilose  externally.  Four  subspecies  were 
recognized  by  Gentry  and  Morawetz  (Gentry, 
1992). 

Jacaranda  copaia  (Aublet)  D.  Don,  Edinburg 
Phil.  J.  9:  267.  1823.  Bignonia  copaia  Aublet. 
Hist.  PI.  Guiane  Fr.  2:  650.  tab.  262,  fig.  1,  tab. 
265.  1775.  J.  spectabilis  Mart,  ex  DC.,  Prodr. 
9:  229.  1845.  J.  superba  Pittier.  Bol.  Soc.  Ve- 
nez.  Ci.  Nat.  6:  19.  1940.  J.  copaia  ssp.  spec- 
tabilis (Mart,  ex  DC.)  A.  Gentry,  Rhodora  79: 
441.  1977.  Figure  14. 

Trees  to  45  m  tall,  trunks  to  ca.  50  cm  diam., 
with  few  upright  branches  forming  a  plume-like 
large-leaved  crown  when  young,  leafy  stems  4- 

12  mm  diam.,  glabrous,  subtetragonal,  lenticel- 
late.  Leaves  30-160  cm  long,  to  60  cm  wide,  pet- 
ioles 4-29  cm  long,  2.5-10  mm  diam.,  minutely 
papillate   puberulent  or  muricate,   terete,  rachis 
narrowly  sulcate  above  (without  wings),  pinnae 
7-35  cm  long,  petiolules  0-3  mm  long;  leaflet 
blades  20-80  mm  long,  6-18(-25)  mm  wide, 
asymmetrically   rhomboid-elliptic   to  elliptic-ob- 
long, apex  acuminate,  base  asymmetric  with  nar- 
rowly cuneate  and  obtuse  sides,  drying  dark  and 
chartaceous,  upper  surface  with  minute  hairs  on 
the  midvein,  sparsely  minutely  puberulent  on  the 
veins  beneath,  2°  veins  3-7/side.  Inflorescences 
to  48  cm  long,  open  pyramidal  panicles  with  lat- 
eral branches  to  30  cm  long  (sometimes  with  3 
terminal  thyrses),  peduncles  to  8  cm  long,  3-5 
mm  diam.,  minutely  puberulent  or  with  peltate 
hairs.  Flowers  with  calyx  tube  4-7  mm  long,  3- 
5  mm  diam,  densely  minutely  (0.1  mm)  puberu- 
lent, irregularly  lobed  (0.3  mm)  or  subentire;  co- 
rolla 23-50  mm  long,  tubular-campanulate  above 
the  narrowed  (8-10  mm)  base,  blue  or  lavender 
(white  in  the  throat),  densely  minutely  puberulent 
externally,  tube  8-14  mm  diam.  distally,  lobes  3- 

13  mm  long;  filaments  ca.  12  and  9  mm  long, 
thecae  ca.  2  mm  long,  staminode  24-27  mm  long, 
pubescent,  bifurcate  at  tip.  Fruits  4.5-12  cm  long, 
3-6  cm  wide,  5-8  mm  thick,  oblong-rounded,  sur- 
face smooth  but  minutely  muricate;  seeds  11-14 
mm  long,  21-30  mm  wide,  seed  ca.  5  mm  diam., 
wings  translucent  with  brown  rays. 

Trees  of  rain  forests  on  both  Caribbean  and  Pa- 
cific slopes,  10-600  m  elevation.  Flowering  in 
February- April;  fruiting  in  July-November.  This 
species  ranges  from  southern  Mexico  to  Brazil 
and  Bolivia. 


BURGER:  FLORA  COSTARICENSIS 


127 


Jacaranda  copaia  is  distinguished  by  the  tall 
tree  habit,  opposite  bipinnate  leaves,  strongly 
asymmetric  leaflets,  tubular  calyx,  bluish  lavender 
corolla  densely  puberulent  externally,  flat  oblong 
woody  fruit,  and  seeds  with  brown  rays  in  the 
otherwise  transparent  wings.  When  mature,  this  is 
one  of  the  most  beautiful  trees  of  southern  Central 
America  (Standley,  1938).  As  a  young  tree,  this 
species  has  a  slender  unbranched  trunk  with  as- 
cending distal  branches  forming  a  terminal  tuft 
with  very  large  bipinnate  leaves.  The  mimosoid 
legume  Schizolobium  parahybum  (Veil.)  Blake  is 
quite  similar  in  this  regard;  both  species  are  vi- 
sually striking  and  often  seen  in  lowland  rain  for- 
est secondary  growth  (Gentry,  1973b).  Gallinazo 
is  a  common  name. 

Jacaranda  mimosifolia  D.  Don,  Bot.  Reg.  8:  tab. 
631.  1822.  J.  ovalifolia  R.  Br.,  Bot.  Mag.  tab. 
2327.  1822.  Figure  14. 

Trees,  usually  small  to  medium  size  (8-15  m 
tall),  stems  4-9  mm  diam.,  usually  glabrous,  te- 
rete, with  elongate  lenticels.  Leaves  13-34  cm 
long,  petioles  to  4  cm  long,  1.8-2.6  mm  diam., 
lateral  pinnae  3-10  cm  long,  with  narrow  wings 
0.3-0.5  mm  wide,  with  13-41  opposite  or  alter- 
nate, subsessile  leaflets;  leaflet  blades  5-14  mm 
long,  1-4  mm  wide  (terminal  leaflets  to  22  X  6 
mm),  elliptic  to  elliptic-obovate,  apex  acute  to 
acuminate,  slightly  asymmetric  at  the  obtuse/cu- 
neate  base,  glabrous  above,  glabrous  or  puberu- 
lent along  the  edge  and  major  veins  beneath,  2° 
veins  3-7/side.  Inflorescences  terminal,  9-38  cm 
long,  open  panicles  with  well-separated  opposite 
branches,  peduncles  34-88  mm  long,  1-4  mm 
diam.,  pedicels  1-2  mm  long  (to  subtending 
node),  flowers  usually  in  distal  cymes.  Flowers 
with  calyx  cup  1-2  mm  long,  ca.  2  mm  wide, 
sparsely  minutely  puberulent,  margin  with  5  tri- 
angular or  acute  teeth  0.2-1  mm  long;  corolla 
30-45  mm  long,  tubular-campanulate,  bluish  lav- 
ender or  lavender,  densely  puberulent  near  the 
base  with  hairs  0.1-0.3  mm  long,  less  densely  pu- 
berulent distally,  tube  9-13  mm  diam.  near  the 
mouth,  lobes  3-8  mm  long;  filaments  ca.  16  and 
13  mm  long,  thecae  ca.  2  mm  long,  staminode 
20-25  mm  long.  Fruits  30-70  mm  long,  37-60 
mm  wide,  12-20  mm  diam.,  orbicular  to  oblong- 
rounded,  lenticular  with  thinner  flattened  edges, 
surface  smooth  and  slightly  muricate;  seeds  9-13 
mm  long,  11-19  mm  wide,  seed  ca.  7  X  6  mm, 
clearly  differentiated  from  the  transparent  circum- 
ferential wing. 

Jacaranda  mimosifolia,  native  to  northern  Ar- 


gentina and  adjacent  Bolivia,  is  widely  planted 
throughout  the  tropics  and  subtropics  as  an  orna- 
mental tree.  In  Central  America  it  is  usually 
grown  between  500  and  1400  m  elevation,  flow- 
ering in  January-July,  with  peak  flowering  in 
April-May.  Standley  and  Williams  (1974)  de- 
scribed the  colorful  effect  of  these  showy  trees  in 
Guatemala.  The  very  small  leaflets  on  opposite 
bipinnate  leaves,  bluish  lavender  flowers  bloom- 
ing in  the  dry  season,  and  hard,  rounded,  lens- 
shaped  fruits  help  distinguish  this  species.  Jaca- 
randa and  Jacaranda  are  the  Spanish  and  English 
names. 


Kigelia  DeCandolle 

Medium-size  trees  up  to  20  m  tall,  with  broad 
spreading  crowns  and  inflorescences  hanging 
from  the  lower  branches.  Leaves  opposite,  peti- 
olate,  imparipinnate  with  3-1 1  leaflets,  the  ter- 
minal leaflet  larger  than  the  laterals  and  borne  on 
a  longer  petiolule,  margin  entire  to  dentate,  ve- 
nation pinnate.  Inflorescences  panicles  with  few 
short  lateral  branches  from  an  elongate  central  ra- 
chis,  long-pedunculate  and  pendulous,  bracts 
small,  pedicels  well  developed.  Flowers  large,  ca- 
lyx campanulate  or  cupular,  splitting  into  2-5 
lobes,  coriaceous,  usually  glabrous;  corolla  cam- 
panulate-funnelform  and  distally  curved,  glabrous 
on  the  exterior,  5-lobed  and  somewhat  2-lipped, 
deep  purple  to  dark  wine-red  (orange-red);  sta- 
mens 4,  of  2  lengths,  subexserted  at  the  mouth  of 
the  corolla,  thecae  slightly  divergent;  disc  annular; 
ovary  lepidote,  1-locular  with  2  parietal  placentas 
or  bilocular  in  the  lower  half,  style  short,  stigma 
simple.  Fruits  indehiscent,  pendulous,  cylindric- 
oblong,  fibrous-woody;  seeds  many,  without 
wings. 

Kigelia  is  now  believed  to  comprise  a  single 
very  variable  species,  native  to  savannas  and  for- 
ests in  central  tropical  Africa.  (In  the  past  some 
authors  recognized  as  many  as  ten  species.)  These 
trees  are  occasionally  grown  as  unusual  exotics  in 
tropical  parks  and  gardens. 

Kigelia  pinnata  (Jacq.)  DC.,  Rev.  Bign.  (Bibl. 
Univ.  Geneve)  24.  1838.  Crescentia  pinnata 
Jacq.,  Collect.  3:  203,  tab.  18.  1791.  K.  africana 
DC.,  Fl.  Nigrit.  463.  1849. 

Trees  6-12  m  (rarely  20  m)  tall,  twigs  often 
hollow,  leafy  stems  6-14  mm  diam.,  glabrous  or 
minutely  puberulent,  glandular  fields  absent  at  the 
nodes.  Leaves  to  50  cm  long,  with  usually  7  or  9 


128 


FIELDIANA:  BOTANY 


leaflets,  petioles  6-14  cm  long,  ca.  3  cm  diam., 
petiolules  of  lateral  leaflets  1-4  mm  long;  leaf 
blades  (2-)5-16  cm  long,  (1.5-)3-7  cm  wide,  el- 
liptic-oblong to  elliptic-obovate,  apex  obtuse  to 
short-acuminate,  margin  entire  or  dentate  distally, 
base  asymmetric  in  lateral  leaflets  (rounded/cu- 
neate),  drying  stiffly  chartaceous,  glabrous  or  with 
minute  (0.1  mm)  straight  hairs,  2°  veins  7-1  I/side. 
Inflorescences  terminal  and  pendulous,  to  40  cm 
long,  peduncles  ca.  20  cm  long,  3  mm  diam.,  with 
lateral  branches  to  9  cm  long  and  2.5  mm  diam., 
pedicels  10-14  mm  long.  Flowers  held  horizon- 
tally, calyx  2-3  cm  long,  12-20  mm  wide  at  the 
apex,  truncated  at  the  base,  lobes  to  12  mm  long, 
coriaceous;  corolla  6-10  cm  long,  tube  ca.  14  mm 
diam.  at  base,  narrowing  to  1 1  mm  and  then  open- 
ing to  4-6  cm  wide,  dark  maroon  or  wine-red; 
anthers  ca.  8  mm  long;  ovary  10-11  mm  long,  3 
mm  diam.  Fruits  25-50(-80)  cm  long,  5-12  cm 
diam.,  tubular-oblong,  pendulous,  glabrous  on  the 
exterior. 

Kigelia  pinnata  is  an  African  tree  rarely  en- 
countered in  Neotropical  parks  and  gardens  (our 
only  Costa  Rican  collections  come  from  Golfito). 
The  pendulous  raceme-like  inflorescences  with 
large  dark  wine-red  or  maroon  flowers  held  hor- 
izontally and  the  large  pendulous  sausage-like 
fruits  make  this  a  unique  species.  These  plants  are 
adapted  for  bat  pollination,  with  inflorescences 
long-pendulous  beneath  the  lower  branches,  and 
night-blooming  flowers  with  thick  tissues  and  foul 
odor. 


Lundia  DeCandolle 
Nomen  conservandum 

Lianas  climbing  with  tendrils  (simple  or  trifid 
near  the  tip),  stems  with  4  phloem  areas  in  cross- 


section,  branchlets  terete,  nodes  often  wider  than 
the  stems,  with  interpetiolar  gland  fields;  pseu- 
dostipules  small  or  absent.  Leaves  opposite,  2-  or 
3-foliolate.  petiolate,  blades  usually  somewhat 
rounded  near  the  base,  margin  entire,  venation 
palmate  or  subpalmate,  tufts  of  hairs  (domatia)  of- 
ten present  in  the  proximal  vein  axils  beneath.  In- 
florescences axillary  or  terminal,  usually  less  than 
20  cm  long,  paniculate,  pedunculate,  bracts  mi- 
nute or  absent,  flowers  pedicellate.  Flowers  with 
calyptrate-conical  calyx  buds,  calyx  margin  entire 
or  splitting,  puberulent;  corolla  tubular-funnel- 
form,  white  to  rose  or  magenta,  puberulent  exter- 
nally; stamens  4,  included,  filaments  of  2  lengths, 
anthers  puberulent,  thecae  divaricate  and  straight, 
staminode  small;  disc  absent;  ovary  densely  pu- 
berulent, 2-locular,  ovules  many  in  2-6  series  in 
each  locule.  style  puberulent  or  glabrous.  Fruits 
capsules,  linear  and  smooth,  valves  flattened  par- 
allel to  the  septum,  surface  densely  puberulent, 
median  vein  and  margins  usually  elevated,  de- 
hiscing septicidally;  seeds  flat,  thin,  transverse-ob- 
long, with  2  lateral  membranaceous  wings  trans- 
parent at  the  tips. 

Lundia  is  a  genus  of  12  species  ranging  from 
Mexico  to  Brazil  and  Bolivia.  The  genus  is  dis- 
tinguished by  its  unusual  calyx,  externally  puber- 
ulent white  to  rose  or  purple  corollas,  stamens 
with  divaricate  puberulent  thecae  (often  in  a 
straight  line),  and  long  narrow  fruits  with  densely 
puberulent  surface.  The  early  flower  buds  are  tu- 
bular with  narrowly  conical  tip  that  is  deciduous 
to  produce  a  calyx  with  an  entire  margin,  which 
then  may  split  as  growth  continues.  The  calyx  re- 
mains obconic  at  the  base  as  opposed  to  the 
rounded  base  in  Paragonia  pyramidata,  which 
may  also  have  calyptrate  calyces.  Costa  Rican 
species  of  Lundia  are  also  vegetatively  similar  to 
species  of  Arrabidaea  and  Cydista. 


Key  to  the  Species  of  Lundia 

la.  Flowers  white;  style  glabrous  except  at  the  base;  calyx  drying  yellowish,  entire  or  sometimes  split; 

occasional  plants  of  evergreen  and  partly  deciduous  forests   L  corymbifera 

Ib.  Flowers  pink  or  magenta;  style  puberulent  near  the  tip  or  along  its  length;  calyx  drying  brownish 

and  often  split  to  form  2  lips;  rarely  collected  in  evergreen  forests    L.  puberula 


Lundia  corymbifera  (Vahl)  Sandw.,  Recueil 
Trav.  Bot.  Ne~erl.  34:  229.  1937.  Bignonia  cor- 
ymbifera Vahl,  Eclog.  Am.  2:  45,  pi.  17.  1798. 
B.  umbrosa  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3: 
138.  1819.  L.  valenzuelae  Dugand,  Mutisia  10: 
7.  1952.  Figure  23. 


Lianas  to  3  cm  diam.,  tendrils  4-14  cm  long, 
leafy  stems  1 .7-6  mm  diam.,  minutely  puberulent 
with  thin  hairs  0.1-0.2  mm  long,  longitudinally 
ridged  (striate),  gland  fields  prominent;  pseudo- 
stipules  conical.  Leaves  2-  or  3-foliolate,  petioles 
8-64  mm  long,  0.8-1.7  mm  diam.,  minutely  pu- 


BURGER:  FLORA  COSTARICENSIS 


129 


berulent,  petiolules  7-42  mm  long;  leaflet  blades 
5-14  cm  long,  2-9  cm  wide,  ovate-elliptic  to 
broadly  ovate,  apex  caudate-acuminate  to  long- 
acuminate  (tips  ca.  15  mm  long),  base  rounded 
and  truncate  to  subcordate,  slightly  or  clearly 
asymmetric,  drying  thin-chartaceous  and  brown, 
minutely  puberulent  on  the  midvein  above,  with 
whitish  hairs  0.1-0.3  mm  long  beneath  and  larger 
hairs  sometimes  present  in  the  vein  axils  (doma- 
tia),  venation  palmate  or  subpalmate,  2°  veins  3- 
7/side.  Inflorescences  axillary  or  terminal,  5-10 
cm  long,  peduncles  3-4.5  cm  long,  ca.  1  mm 
diam.,  densely  puberulent  with  short  (0.1-0.2 
mm)  yellowish  hairs,  pedicels  5-8  mm  long. 
Flowers  with  calyx  3-6  mm  long,  3-4  mm  diam., 
cupulate-campanulate  to  obconic,  margin  entire  or 
split  on  1  or  2  sides,  minutely  puberulent;  corolla 
26-43  mm  long,  tubular-funnelform,  white, 
densely  minutely  puberulent  externally,  with  yel- 
low ridges  in  the  throat,  lobes  4-12  mm  long; 
filaments  ca.  14  and  9  mm  long,  thecae  2-3  mm 
long,  divaricate.  Fruits  28-60  cm  long,  15-20 
mm  wide,  valves  thick,  with  raised  midrib  and 
edges,  densely  minutely  velutinous;  seeds  6-14 
mm  long,  19-40  mm  wide,  central  area  ca.  10 
mm  wide,  lateral  wings  with  darker  interior  and 
clear  distal  areas. 

Vines  often  found  along  stream  edges  in  ever- 
green and  partly  deciduous  forest  formations  of 
the  Pacific  slope,  10—900  m  elevation.  Flowering 
in  June-December;  fruiting  in  February-March. 
This  species  ranges  from  central  Costa  Rica  to 
Brazil  and  Bolivia. 

Lundia  corymbifera  is  distinguished  by  its  vin- 
ing  habit  with  simple  tendrils,  opposite  bi-  or  tri- 
foliolate  leaves,  leaflets  rounded  at  the  base,  short 
inflorescences,  unusual  calyx  buds,  pubescent  an- 
thers, white  corolla  puberulent  on  the  exterior,  and 
long  linear  fruits  with  two-winged  seeds.  The 
flower  buds  are  ellipsoid  and  often  have  an  apic- 
ulate  apex  that  usually  dehisces  as  a  cap,  leaving 
an  entire  margin. 

Lundia  puberula  Pittier,  Contrib.  U.S.  Natl. 
Herb.  18:  258.  15  Sept.  1917.  L  dicheilocalyx 
Blake,  Contrib.  Gray  Herb.  52:  94.  28  Sept. 
1917.  L.  schumanniana  Kranzl.,  Fedde  Repert. 
17:  120.  1921.  L.  colombiana  Dugand,  Caldasia 
4:  236.  1946. 

Lianas  to  5  cm  diam.,  tendrils  5-15  cm  long, 
leafy  stems  1.6-6  mm  diam.,  minutely  (0.1  mm) 
puberulent,  longitudinally  ridged  or  striate,  pseu- 
dostipules  inconspicuous.  Leaves  2-  or  3-folio- 


late,  petioles  2.4-6  cm  long,  petiolules  10-20  mm 
long,  0.6-1  mm  diam.,  minutely  puberulent;  leaf 
blades  6-12(-16)  cm  long,  3.5-7(-9)  cm  wide, 
ovate  to  ovate-oblong,  apex  long-acuminate  (tips 
ca.  2  cm  long),  base  rounded  and  truncate  to  cor- 
date, slightly  asymmetric,  drying  thin-chartaceous 
and  brown,  sparsely  puberulent  above,  minutely 
puberulent  beneath  and  often  with  longer  (0.2-0.6 
mm)  hairs  in  the  vein  axils  (domatia),  venation 
palmate  or  subpalmate,  2°  veins  3-5/side.  Inflo- 
rescences axillary  or  terminal,  3-12  cm  long,  pe- 
duncles 15-55  mm  long,  1-2  mm  diam.,  minutely 
puberulent,  bracts  ca.  1  mm  long,  pedicels  3—8 
mm  long,  ca.  0.6  mm  diam.  Flowers  with  calyx 
4-8  mm  long,  3-4  mm  diam,  tubular,  entire  or 
split  on  the  margin,  minutely  puberulent  with 
short  (0.1  mm)  hairs;  corolla  30-58  mm  long, 
tubular-campanulate  or  funnelform,  slightly  2- 
lipped,  pale  pink  to  magenta,  densely  minutely 
puberulent  externally,  lobes  8-21  mm  long;  fila- 
ments 14-20  and  9-13  mm  long,  thecae  2-3  mm 
long.  Fruits  long  linear  capsules,  23-48  cm  long, 
15-17  mm  wide,  valves  flat  with  raised  midvein, 
surface  velutinous;  seeds  9-12  mm  long,  33-35 
mm  wide,  wings  brown  to  transparent  distally. 

Rarely  collected  vines  in  lowland  evergreen 
rain  forests  of  the  Caribbean  and  Pacific  slopes, 
0-900  m  elevation.  Flowering  throughout  the 
year.  This  species  ranges  from  Mexico  and  Belize 
to  Peru. 

Lundia  puberula  is  recognized  by  its  vining 
habit  with  simple  tendrils,  the  opposite  bi-  or  tri- 
foliolate  leaves  with  long-acuminate  tips  and 
rounded  base,  the  entire  or  split  calyx,  pubescent 
anthers,  pink  to  magenta  corollas  minutely  puber- 
ulent externally,  and  long  slender  fruits.  The  usual 
presence  of  tufted  hairs  in  vein  axils,  gland  fields 
at  nodes,  and  lowland  rain  forest  habitat  are  ad- 
ditional distinctions.  This  species  is  easy  to  con- 
fuse with  some  species  of  Arrabidaea  and  Cydis- 
ta. 


Macfadyena  A.  DeCandolle 

Lianas,  tendrils  with  hardened  curved  trifid 
tips,  stems  terete,  cross-section  with  ca.  8  phloem 
areas  and  fissured  xylem,  roots  with  swollen  tu- 
bers, interpetiolar  gland  fields  present  or  absent; 
pseudostipules  small,  lanceolate  to  ovate.  Leaves 
evergreen  or  deciduous,  opposite,  2-foliolate,  pet- 
iolate,  often  with  a  tendril,  margin  entire,  venation 
pinnate.  Inflorescences  axillary  or  terminal  on 
short-shoots,  1  or  2  or  3-15  on  short  cymes  or 


130 


FIELDIANA:  BOTANY 


reduced  panicles,  pedicels  subtended  by  nodes. 
Flowers  with  tubular  or  campanulate  calyx,  lobed 
or  variously  split  to  spathe-like,  usually  thin-tex- 
tured; corolla  tubular-campanulate  or  tubular-fun- 
nelform,  2-lipped,  bright  yellow,  glabrous  exter- 
nally, with  5  rounded  lobes;  stamens  4,  filaments 
of  2  lengths,  anthers  glabrous,  thecae  straight,  di- 
varicate, staminode  present;  disc  annular-pulvi- 
nate,  ovary  with  minute  peltate  hairs  to  puberulent 
or  glabrate,  locules  2,  ovules  in  2-4  series  in  each 
locule.  Fruits  elongate  linear  capsules,  valves 
parallel  to  the  septum,  flat,  smooth,  midvein 
slightly  raised;  seeds  flat,  transversely  oblong. 


central  area  not  clearly  differentiated  from  the  2 
thin  lateral  wings. 

Macfadyena  is  a  genus  of  four  species  ranging 
from  Mexico  and  the  West  Indies  to  Argentina. 
The  plants  are  dimorphic,  with  the  leaf  blades  of 
young  plants  usually  smaller  and  more  lanceolate 
than  the  blades  of  more  mature  stems.  In  addition, 
these  plants  are  unusual  in  that  they  arc  able  to 
climb  up  tree  trunks  using  their  short,  three- 
clawed  tendrils  as  grappling  hooks;  advcntitous 
roots  also  aid  in  adhering  to  their  support.  The 
flowers  can  vary  greatly  in  size,  even  on  the  same 
vine. 


Key  to  the  Species  of  Macfadyena 

la.  Calyx  usually  split  on  1  side  (spathe-like)  with  apical  tip  curved  backward;  seeds  20-25  mm  wide, 
brown  throughout,  fruits  to  30  cm  long;  pseudostipules  subulate-lanceolate;  plants  of  lowland 
swamps  below  100  m  elevation  in  evergreen  forest  formations  M.  uncata 

Ib.  Calyx  subentire  or  variously  lobed  or  split  (not  spathe-like);  seeds  40-65  mm  wide,  translucent  near 
the  tips,  fruits  to  130  cm  long;  pseudostipules  often  ovate  and  striate;  plants  of  well-drained  sites 

in  deciduous  and  partly  deciduous  or  evergreen  forest  formations,  10-1200  m  elevation  

M.  unguis-cati 


Macfadyena  uncata  (Andr.)  Sprague  &  Sandw., 
Recueil  Trav.  Bot.  Neerl.  34:  215.  1937.  Big- 
nonia  uncata  Andr.,  Bot.  Repos.  tab.  530.  1808. 
B.  uncinata  G.  Meyer,  Prim.  Fl.  Essequeb.  210. 
1818.  M.  uncinata  (G.  Meyer)  A.  DC.,  Prodr. 
180.  1845.  M.  guatemalensis  Blake,  Contrib. 
U.S.  Natl.  Herb.  24:  24.  1922.  Figure  17. 

Lianas  to  30  m  high,  tendrils  with  3  stiff 
curved  distal  tips,  leafy  twigs  1-4  mm  diam.,  mi- 
nutely puberulent  or  glabrous,  terete,  gland  fields 
usually  visible  on  young  stems;  pseudostipules  ca. 
2  mm  long,  subulate-lanceolate.  Leaves  with  pet- 
ioles 8-36  mm  long,  1-1.7  mm  diam.,  glabrous 
or  sparsely  puberulent  with  thin  hairs  ca.  0.2  mm 
long,  petiolules  4-27  mm  long;  leaflet  blades  5- 
20  cm  long,  2-9  cm  wide,  narrowly  ovate  to  ovate 
or  ovate-elliptic,  apex  acuminate,  often  with  a 
long  (2  cm)  narrow  tip,  base  obtuse  or  cuneate 
(rounded),  drying  chartaceous  and  often  dark, 
subglabrous  or  with  very  minute  hairs  above,  mi- 
nutely puberulent  and  with  appressed  peltate  hairs 
beneath,  2°  veins  5-8/side.  Inflorescences  axil- 
lary to  foliage  leaves,  flowers  1-3  in  reduced 
cymes  or  panicles,  peduncles  0-9  mm  long,  ca.  1 
mm  diam.,  glabrous,  drying  black,  bracts  1-4  mm 
long,  lanceolate,  caducous,  pedicels  ca.  10  mm 


long.  Flowers  with  calyx  15-27  mm  long,  6-14 
mm  diam.,  glabrous  or  with  few  minute  peltate 
hairs,  usually  split  on  1  side  (spathaceous)  with 
the  narrowed  apex  recurved;  corolla  40-85  mm 
long,  tubular-funnelform,  yellow,  glabrous  exter- 
nally, 8-18  mm  diam.  at  the  mouth,  lobes  7-17 
mm  long;  filaments  ca.  23  and  18  mm  long,  the- 
cae 2.5  mm  long.  Fruits  16-30  cm  long.  15-19 
mm  wide,  valves  drying  dark  brown;  seeds  12- 
14  mm  long,  20-25  mm  wide,  transversely  ob- 
long-rectangular with  short  lateral  wings,  dark 
brown  throughout. 

Plants  of  swampy  sites  in  evergreen  lowland 
forest  formations  of  both  Caribbean  and  Pacific 
slopes,  1-110  m  elevation.  Probably  flowering 
throughout  the  year.  This  species,  often  found  in 
mangrove  formations,  ranges  from  Mexico  to 
Brazil. 

Macfadyena  uncata  is  recognized  by  its  climb- 
ing habit  with  three-clawed  tendrils,  bifoliolate 
leaves  with  acuminate  blades,  few-flowered  axil- 
lary inflorescences,  spathe-like  calyx,  yellow  co- 
rollas glabrous  externally,  and  narrow  fruits  with 
short-winged  seeds.  It  is  also  unusual  in  being  re- 
stricted to  swampy  habitats  at  low  elevations. 

Macfadyena  unguis-cati  (L.)  A.  Gentry,  Britton- 
ia  25:  236.  1973.  Bignonia  unguis-cati  L.,  Sp. 


BURGER:  FLORA  COSTARICENSIS 


131 


PI.  2:  623.  1753.  Doxantha  unguis-cati  (L.) 
Miers  emend.  Rehder,  Mitt.  Deutsch,  Dendrol. 
Gesel.  1913:  262.  1913.  B.  dasyonyx  Blake, 
Contrib.  Gray  Herb.  52:  93.  1917.  D.  dasyonyx 
(Blake)  Blake,  J.  Bot.  61:  192.  1923.  Figure  17. 

Lianas  to  30  m  high,  to  7  cm  diam.,  tendrils 
2-5  cm  long  with  3  stiff  curved  terminal  "claws" 
5-14  mm  long,  leafy  stems  1.3-4  mm  diam.,  gla- 
brescent  and  grayish,  gland  fields  present  or  ab- 
sent at  nodes;  pseudostipules  lanceolate  to  ovate, 
often  with  parallel  longitudinal  ridges  (striate). 
Leaves  dimorphic  (young  vines  vs.  mature 
stems),  2-foliolate,  petioles  1 1-27(-47)  mm  long, 
0.8-1.5  mm  diam.,  glabrous  or  sparsely  and  mi- 
nutely puberulent,  petiolules  4-16(-25)  mm  long; 
leaf  blades  5-16  cm  long,  1-7  cm  wide,  narrowly 
ovate-elliptic  to  narrowly  ovate  or  elliptic,  apex 
acuminate  with  narrowed  tip  to  2  cm  long,  base 
rounded  to  acute,  drying  thinly  chartaceous,  most- 
ly glabrous  except  for  minute  puberulence  on  the 
midvein  and  with  peltate  hairs  beneath,  2°  veins 
4-8/side.  Inflorescences  terminal  on  short-shoots 
or  axillary  to  leafless  nodes,  panicles  of  up  to  15 
flowers,  peduncles  8-28  mm  long,  usually  gla- 
brous, pedicels  8-10  mm  long.  Flowers  with  ca- 
lyx 6-15  mm  long,  7-12  mm  wide,  cupular  to 
campanulate,  thin-textured,  distal  margin  irregu- 
larly split  or  with  lobes  0.5-2  mm  long,  glabrous 
or  with  few  peltate  hairs;  corolla  42-80  mm  long, 
tubular  campanulate  or  funnelform,  deep  yellow 
or  yellow-orange,  glabrous  externally,  tube  12-18 
mm  wide  at  mouth,  lobes  9-18  mm  long;  fila- 
ments ca.  20  and  14  mm  long,  thecae  ca.  3  mm 
long,  staminode  rarely  forming  a  fifth  stamen. 
Fruits  24-145  cm  long,  11-19  mm  wide,  linear, 
tapering  at  both  ends,  valves  smooth,  with  minute 
peltate  hairs,  drying  dark  brown;  seeds  9-14  mm 
long,  40-65  mm  wide,  mostly  brown  with  lateral 
tips  transparent. 

Deciduous  lianas  of  seasonally  dry  deciduous, 
partly  deciduous,  and  (less  often)  evergreen  forest 
formations,  10-900(-1300)  m  elevation.  Flower- 
ing primarily  in  February-June.  This  species 
ranges  from  Mexico  and  the  West  Indies  to  Ar- 
gentina. 

Macfadyena  unguis-cati  is  recognized  by  its 
climbing  habit  with  short  distally  three-clawed 
tendrils,  opposite  bifoliolate  leaves,  inflorescences 
usually  flowering  before  the  new  leaves  are  fully 
flushed,  thin  calyx  with  irregular  margin,  deep 
yellow  corollas  glabrous  externally,  long-linear 
flattened  fruits,  and  two-winged  seeds.  The  un- 


usual dimorphism  and  climbing  habit  (see  discus- 
sion under  genus)  are  additional  distinctions.  The 
leaves  on  very  young  plants  may  be  only  1  cm 
long  and  rounded  (Standley,  1938).  Gentry 
(1973b)  reported  that  the  seedlings  of  this  species 
can  persist  for  extended  periods  in  drier  areas. 
Una  de  gato. 


Mansoa  DeCandolle 

Lianas  climbing  with  distally  trifid  tendrils  or 
simple  tendrils  with  a  terminal  peltate  disc,  stems 
terete  or  somewhat  quadrangular,  with  4-8  phlo- 
em areas  in  cross-section,  often  with  conspicuous 
glandular  fields  at  the  nodes;  pseudostipules  well 
developed  to  inconspicuous.  Leaves  opposite,  2- 
or  3-foliolate,  glabrous  or  puberulent,  petioles 
sometimes  with  a  glandular  field  at  the  apex,  mar- 
gin entire,  venation  pinnate  or  subpalmate.  Inflo- 
rescences axillary  to  distal  leaves  or  leafless 
nodes  (terminal),  panicles,  racemes  or  small  cor- 
ymbs, usually  few-flowered,  pedicels  subtended 
by  minute  bracts  or  ebracteate.  Flowers  with  cu- 
pulate  or  tubular-campanulate  calyx,  5-lobed  or 
the  margin  truncated,  often  with  plate-shaped 
glands  on  the  distal  surface;  corolla  narrowly  tu- 
bular at  the  base,  distally  tubular-funnelform  to 
tubular-campanulate,  white  to  reddish  purple,  pu- 
berulent externally  (tube  may  be  glabrous);  sta- 
mens 4,  filaments  of  2  lengths,  anthers  glabrous 
or  villous,  thecae  curved  or  straight,  divaricate,  a 
staminode  present;  disc  annular-pulvinate;  ovary 
cylindric,  lepidote  to  papillate,  2-locular,  ovules  in 
2-4  series  in  each  locule.  Fruits  linear-oblong 
capsules,  valves  parallel  to  the  septum,  woody, 
thin  and  flat  to  thick  and  terete,  surface  smooth 
or  with  elevated  spines;  seeds  flat  with  2  thin  lat- 
eral distally  membranaceous  wings  or  corky  and 
lacking  lateral  wings. 

Mansoa  is  a  genus  of  15  Neotropical  species, 
mostly  South  American.  This  generic  concept 
now  includes  a  number  of  species  formerly  placed 
in  Pachyptera  and  Pseudocalymma  (Gentry, 
1973b;  Standley  &  Williams,  1974).  Five  distinc- 
tive species  are  placed  here:  one  with  very  small 
leaves  (M.  parvifolia),  two  whose  vegetative  parts 
smell  like  onions  (M.  hymenaea  and  M.  standle- 
yi),  one  with  verrucose-echinate  fruits  (M.  verru- 
cifera),  and  one  with  large  white  corollas  (M.  ker- 
ere).  Except  for  M.  hymenaea,  the  species  of  this 
genus  have  rarely  been  collected  in  Costa  Rica. 


132 


FIELDIANA:  BOTANY 


Key  to  the  Species  of  Mansoa 

la.  Leaflets  <  30  mm  long;  tendrils  <  3  cm  long,  with  flat  disc-like  pad  at  the  apex    

M.  parvifolia 

Ib.  Leaflets  >  40  mm  long;  tendrils  usually  >  5  cm  long,  usually  ending  in  3  slender  tips  (disc  absent) 

2 

2a.  Fruit  surfaces  covered  with  spine-like  tubercles  1-5  mm  long;  gland  fields  absent  at  the  nodes; 
calyx  with  well-defined  lobes  to  4  mm  long  [leaves  without  a  garlic-like  or  onion-like  odor)    .... 

M.  verrucifera 

2b.  Fruits  with  flat  smooth  surfaces;  gland  fields  usually  present  at  the  nodes;  calyx  with  truncated  or 

irregular  margins  or  rarely  with  lobes  to  2  mm  long 3 

3a.  Vegetative  parts  lacking  the  odor  of  onion;  pseudostipules  usually  in  a  vertical  series  of  3  in 
each  leaf  axil,  acute;  leaves  mostly  3-foliolate;  inflorescences  racemose;  corolla  pubcruleht  on 

tube  and  lobes  externally,  white  (in  Central  America)    M.  kerere 

3b.  Vegetative  parts  with  the  odor  of  onion  or  garlic  when  crushed;  pseudostipules  usually  incon- 
spicuous, obtuse;  leaves  2-foliolate;  inflorescences  panicles  or  racemes,  corolla  puberulent  only 

on  the  lobes  externally,  reddish  purple    4 

4a.  Calyx  4-7  mm  long,  green  at  maturity;  fruits  15-25  mm  wide,  with  midvein  raised  on  surface  of 

the  valves;  plants  of  deciduous  and  partly  deciduous  forests M.  hymenaea 

4b.  Calyx  9-21  mm  long,  green  to  purplish  at  maturity;  fruits  with  the  midvein  obscure  on  the  valve 
surface;  plants  mostly  in  evergreen  forest  formations    M.  siandleyi 


Mansoa  hymenaea  (DC.)  A.  Gentry,  Ann.  Mis- 
souri Bot.  Card.  66:  782.  1979.  Bignonia  hy- 
menaea DC.,  Prodr.  9:  158.  1845.  Adenocalym- 
ma  macrocarpum  J.  D.  Smith,  Bot.  Gaz.,  40:  9. 
1915.  A.  ciliolatum  Blake,  Contrib.  Gray  Herb. 
52:  90.  1917.  A.  hosmeca  Pittier,  Contrib.  U.S. 
Natl.  Herb.  18:  256.  1917.  Petastoma  tondu- 
zianum  Kra'nzl.,  Fedde  Repert.  17:  56.  1921. 
Figure  22. 

Lianas  to  5  cm  diam.,  tendrils  10-15  cm  long, 
leafy  stems  0.8-7  mm  diam.,  subtetragonal,  gla- 
brous (new  shoots  sparsely  minutely  puberulent), 
becoming  pale  yellowish  brown  or  gray;  pseudo- 
stipules  ca.  3  mm  long,  ovoid,  striate.  Leaves  2- 
foliolate,  petioles  6-38  mm  long,  0.7-2  mm 
diam.,  glabrous  or  very  minutely  (0.05  mm)  pu- 
berulent, petiolules  8-26  mm  long,  thickened  at 
the  blade;  leaflet  blades  4-14  cm  long,  2-8  cm 
wide,  ovate  to  ovate-oblong  or  suborbicular,  apex 
short-acuminate  to  bluntly  obtuse,  base  obtuse  to 
rounded-truncate  (subcordate),  drying  stiffly  char- 
taceous  and  grayish  or  green,  glabrous  on  both 
surfaces,  2°  veins  3-10/side.  Inflorescences  axil- 
lary or  terminal,  4-10  cm  long,  panicles,  pedun- 
cles 1-5  cm  long,  ca.  2  mm  diam.,  pedicels  5-20 
mm  long,  sparsely  to  densely  puberulent  with  thin 
straight  hairs.  Flowers  with  calyx  4-7  mm  long, 
4-6  mm  diam.,  cupulate,  puberulent  to  subgla- 
brous,  margin  subentire  or  minutely  5-lobed,  cil- 
iolate;  corolla  37-65  mm  long,  tubular-campan- 


ulate  (slightly  2-lipped),  white  to  lavender  or  dark 
pink,  glabrous  or  with  the  lobes  puberulent  exter- 
nally, tube  8-14  mm  diam.,  lobes  9-14  mm  long; 
filaments  ca.  18  and  13  mm  long,  thecae  ca.  3 
mm  long.  Fruits  15-34  cm  long,  15-23  mm  wide, 
linear-oblong,  flat,  midvein  raised,  surface 
smooth;  seeds  11-15  mm  long.  34-62  mm  wide, 
center  ca.  12  mm  diam.,  not  clearly  differentiated, 
wings  translucent. 

Common  plants  of  seasonally  dry  deciduous 
and  partly  deciduous  forests  of  the  Pacific  slope. 
10-1300  m  elevation.  Flowering  primarily  in  De- 
cember-March. This  species  ranges  from  northern 
Mexico  to  southeastern  Brazil. 

Mansoa  hymenaea  is  recognized  by  its  climb- 
ing habit  with  trifid  tendrils,  opposite  bifoliolate 
broadly  ovate  leaves,  short  inflorescences,  white 
to  rose  or  lilac  corollas  puberulent  distally.  flat, 
narrow  fruits  tapering  at  both  ends,  and  thin  two- 
winged  seeds.  The  strong  odor  of  garlic  or  onions 
when  vegetative  parts  are  crushed  is  a  very  dis- 
tinctive feature.  Specimens  placed  here  were 
called  Pseudocalymma  sagotii  (Bur.  &  K. 
Schum.)  Sandwith  by  Williams  in  Flora  of  Gua- 
temala (Standley  &  Williams,  1974).  Ajillo  and 
pedo  de  padre  are  common  names. 

Mansoa  kerere  (Aublet)  A.  Gentry,  Ann.  Mis- 
souri Bot.  Gard.  66:  783.  1979.  Bignonia  kerere 
Aublet,  Hist.  PI.  Guiane  Fr.  2:  644,  tab.  260. 
1775.  Tanaecium  zetekii  Stand!.,  Contrib.  Ar- 


BURGER:  FLORA  COSTARICENSIS 


133 


nold  Arbor.  5:  140,  tab.  19.  1933.  Pachyptera 
kerere  (Aublet)  Sandw.,  Recueil  Trav.  Bot. 
Nteerl.  34:  219.  1937.  Figure  22. 

Lianas  to  5  cm  diam.,  tendrils  6-16  cm  long, 
leafy  stems  1.8-6  mm  diam.,  somewhat  quadran- 
gular in  cross-section,  glabrous  or  minutely  pu- 
berulent  with  thin  straight  hairs  ca.  0.1  mm  long, 
gland  fields  often  conspicuous  with  sunken  pits; 
pseudostipules  2-6  mm  long,  often  3/axil  and  the 
more  abaxial  progressively  shorter,  lanceolate. 
Leaves  2-  or  3-foliolate,  petioles  10-85  mm  long, 
0.8-2.2  mm  diam.,  minutely  puberulent,  longitu- 
dinally striate,  with  glandular  pits  at  the  apex,  pet- 
iolules  12-61  mm  long;  leaflet  blades  8-18(-24) 
cm  long,  3-9(-ll)  cm  wide,  ovate  to  ovate-ob- 
long or  narrowly  ovate-elliptic,  apex  acuminate  to 
acute  (rarely  rounded  or  emarginate),  base  obtuse 
to  rounded  and  narrowly  subcordate,  drying  thinly 
chartaceous  and  brown,  surfaces  subglabrous  with 
minute  hairs  along  the  major  veins  on  both  sur- 
faces, 2°  veins  5-9/side.  Inflorescences  terminal 
or  axillary,  1 .5-4  cm  long,  racemes,  axis  2-3  mm 
diam.,  stem-like  with  many  nodes  and  reflexed 
bracts  1-3  mm  long,  pedicels  8-14  mm  long. 
Flowers  with  calyx  9-1 1  mm  long,  5-8  mm 
diam.,  deep-cupular,  minutely  papillate-puberu- 
lent  to  subglabrous  with  peltate  hairs,  rounded 
glands  present  on  distal  half,  margin  with  few 
short  (2  mm)  broad  (3  mm)  lobes  or  irregular; 
corolla  4-7  cm  long,  narrowly  tubular-funnel- 
form,  white  or  dull  white  (yellowish  within),  gla- 
brous proximally  or  minutely  puberulent  exter- 
nally, tube  3-9  mm  diam.,  lobes  5-15  mm  long; 
filaments  ca.  20  and  15  mm  long,  thecae  2-4  mm 
long.  Fruits  (10-)22-42  cm  long,  17-40  mm 
wide,  narrowly  oblong  with  narrowed  apex  and 
base,  flat  or  somewhat  rounded,  midvein  elevated 
or  obscure,  surface  smooth  and  with  round  glands; 
seeds  18-24  mm  long,  28-38  mm  wide,  with 
corky  texture,  wings  absent. 

Rarely  collected  plants  of  wet  evergreen  forests 
on  both  the  Caribbean  and  Pacific  lowlands,  0- 
150  m  elevation.  These  plants  are  usually  found 
along  stream  margins,  lake  shores,  and  coastal 
marshes  (Gentry,  1973b).  This  species  has  been 
collected  in  the  Rio  Sixaola  (Talamanca  Valley) 
and  Golfo  Dulce  areas  in  Costa  Rica.  Probably 
flowering  throughout  the  year.  This  species  ranges 
from  Belize  to  Brazilian  Amazonia. 

Mansoa  kerere  is  distinguished  by  its  climbing 
habit  with  three-tipped  tendrils,  opposite  two-  or 
three-foliolate  leaves,  short  racemose  inflores- 
cences, long  white  puberulent  corollas,  pubescent 


anthers,  and  narrowly  oblong  fruits.  Deeply  pitted 
gland  fields  at  the  apex  of  the  petiole  and  the 
three-seriate  pseudostipules  in  leaf  axils  are  useful 
vegetative  distinctions.  The  seeds  may  be  corky 
(the  typical  variety  dispersed  by  water)  or  thin  and 
winged,  e.g.,  var.  incarnata  (Aublet)  A.  Gentry. 
The  Central  American  collections  all  have  white 
corollas,  but  some  South  American  populations 
have  red  flowers  (var.  erythraea  Dugand). 

Mansoa  parvifolia  (A.  Gentry)  A.  Gentry,  Ann. 
Missouri  Bot.  Gard.  66:  783.  1979.  Pachyptera 
parvifolia  A.  Gentry,  Phytologia  26:  448.  1973. 
Figure  16. 

Woody  or  herbaceous  vines  with  stems  to  1  cm 
diam.,  climbing  with  short  (2-5  mm)  tendrils 
bearing  a  flat  rounded  terminal  disc  1-5  mm  wide, 
leafy  stems  0.2-0.7  mm  diam.,  glabrous  or  sparse- 
ly puberulent  with  hairs  ca.  0.1  mm  long;  pseu- 
dostipules ca.  0.3  mm  long,  conical.  Leaves  2- 
foliolate  and  often  with  a  tendril,  petioles  l-3(-8) 
mm  long,  glabrous  abaxially  and  sparsely  to 
densely  minutely  puberulent  on  the  adaxial  sur- 
face, petiolules  0.5-2(-7)  mm  long;  leaflet  blades 
(3-)6-18(-30)  mm  long,  2-9(-17)  mm  wide,  el- 
liptic-oblong to  oblong  or  oblong-obovate,  apex 
obtuse  (rounded),  base  asymmetric  with  cuneate 
and  slightly  rounded  sides,  drying  chartaceous 
and  grayish  beneath,  surfaces  with  short  (0. 1  mm) 
thin  straight  hairs,  2°  veins  2-4/side,  often  ob- 
scure, loop-connected.  Inflorescences  axillary  at 
leafless  nodes,  short,  2-  or  3-flowered,  bracts  ab- 
sent, peduncle  5-6  mm  long,  pedicels  1-1.5  mm 
long,  subglabrous  or  with  minute  peltate  hairs. 
Flowers  with  calyx  4-6  mm  long,  5-8  mm  wide, 
campanulate,  truncated  or  slightly  2-lipped,  sub- 
glabrous;  corolla  35-50  mm  long,  tubular-funnel- 
form,  white  or  pale  purple,  minutely  puberulent 
externally,  tube  8-12  mm  diam.  at  the  mouth, 
lobes  11-13  mm  long;  filaments  16-25  and  11- 
18  mm  long,  thecae  2-4  mm  long.  Fruits  23-43 
cm  long,  20-25  mm  wide,  linear-oblong,  surface 
smooth  (midvein  obscure),  pale  brown;  seeds  not 
seen. 

Rarely  collected  plants  of  evergreen  lowland 
rain  forest  formations,  10-700  m  elevation.  Only 
sterile  trunk-climbing  specimens  with  very  small 
leaves  have  been  collected  in  Central  America. 
This  species  ranges  from  Honduras  to  Bolivia. 

Mansoa  parvifolia  is  one  of  the  most  distinctive 
species  of  Bignoniaceae.  The  slender  stems  cling- 
ing to  tree  trunks  by  means  of  short  tendrils  end- 
ing in  a  peltate  disc  and  the  very  small  opposite 


134 


FIELDIANA:  BOTANY 


bifoliolate  leaves  are  unique  among  the  species  of 
the  family.  The  few-flowered  inflorescences  and 
puberulent  white  or  rose  corollas  are  additional 
characteristics  but  are  rarely  seen.  The  leaves  be- 
come 20-30  mm  long  on  distal  flowering  stems, 
but  these  stems  have  not  been  collected  in  Central 
America.  The  slender  climbing  stems  with  four 
small  leaflets  at  each  node  are  reminiscent  of 
some  Peperomia  species  with  whorled  leaves. 

Mansoa  standleyi  (Steyermark)  A.  Gentry,  Ann. 
Missouri  Hot.  Card.  66:  783.  1979.  Pseudoca- 
lymma  standleyi  Steyermark,  Publ.  Field  Mus. 
Nat.  Hist.,  Bot.  Sen  23:  235.  1947.  Pachyptera 
standleyi  (Steyermark)  A.  Gentry,  Brittonia  25: 
236.  1973.  Ps.  alliaceum  var.  macrocalyx 
Sandw.,  Kew  Bull.  1953:  468.  1954. 

Lianas  to  8  cm  diam.,  tendrils  10-14  cm  long 
with  3  distal  arms  to  2  cm  long,  vegetative  parts 
smelling  like  garlic,  leafy  stems  2.5-8  mm  diam., 
glabrous,  longitudinally  striate,  nodes  with  a  few 
punctate  glands;  pseudostipules  small,  conical. 
Leaves  2-foliolate,  petioles  9-48  mm  long,  1.7- 
2.7  mm  diam.,  glabrous,  petiolules  7-25  mm  long, 
thickened  at  the  apex;  leaflet  blades  10-20(-24) 
cm  long,  6- 11  (-14)  cm  wide,  elliptic-ovate  to 
broadly  ovate,  apex  acute  to  short-acuminate,  base 
obtuse  or  rounded,  drying  green  to  olive  green 
and  chartaceous  (subcoriaceous  in  life),  surfaces 
glabrous,  with  gland  fields  in  the  axils  of  basal 
veins  beneath,  2°  veins  6-9/side.  Inflorescences 
axillary  or  terminal,  to  45  cm  long,  pyramidal 
panicles  or  reduced  to  racemose,  usually  with 
short  (1-3  cm)  opposite  branches,  peduncles  to  18 
cm  long,  glabrous,  bracts  8-19  mm  long,  6-11 
mm  wide,  bright  red,  caducous,  pedicels  8-20  mm 
long.  Flowers  with  cupulate  calyx  (9-)  17-21  mm 
long,  (8-)  15-1 8  mm  wide,  rounded  at  the  base, 
glabrous,  margin  with  5  undulate  lobes  or  sub- 
entire,  thin-papery  in  life  and  green  to  red-violet; 
corolla  (38-)66-85  mm  long,  tubular-campanu- 
late,  rose-red  to  violet  on  lobes  and  upper  tube, 
base  of  tube  yellow,  glabrous  externally,  lobes  6- 
27  mm  long,  apically  obtuse;  filaments  1 1-22  and 
22-30  mm  long,  thecae  3-4  mm  long,  linear. 
Fruits  35-75  cm  long,  26-38  mm  wide,  elongate- 
linear,  acute  at  both  ends,  flat  (midvein  obscure), 
pale  brown;  seeds  20-28  mm  long,  75-110  mm 
wide,  central  area  poorly  differentiated,  wings 
whitish,  thin. 

Uncommon  plants  of  evergreen  forest  forma- 
tions on  both  the  Caribbean  and  Pacific  slopes, 
50-1100  m  elevation  (1300-1400,  m  elevation  in 


Guatemala).  Probably  flowering  throughout  the 
year.  This  species  ranges  from  Mexico  to  Peru  and 
Brazil. 

Mansoa  standleyi  is  recognized  by  its  climbing 
habit  with  distally  trifid  tendrils,  mostly  glabrous 
parts,  opposite  bifoliolate  leaves  with  large  leaf- 
lets, large  cupulate  calyces,  large  pinkish  corollas, 
and  long  linear  fruits  with  large  winged  seeds. 
The  plants  are  also  distinctive  because  of  the  on- 
ion-like odor  of  vegetative  parts  and  flowers  and 
the  gland  fields  in  basal  vein  axils  of  the  lower 
leaf  surface.  This  species  was  treated  under  Pa- 
chyptera in  Flora  of  Panama  and  as  Pseudoca- 
lymma  sagotii  var.  macrocalyx  (Sandw.)  L.  O. 
Williams  in  Flora  of  Guatemala  (Standley  &  Wil- 
liams, 1974,  p.  216).  The  type  collection  (Stey- 
ermark 33533  F)  cites  bejuco  de  ajo  as  a  common 
name  in  Guatemala. 

Mansoa  verrucifera  (Schldl.)  A.  Gentry,  Ann. 
Missouri  Bot.  Gard.  63:  62.  1976.  Bignonia  ver- 
rucifera Schldl.,  Linnaea  26:  655.  1853.  Aden- 
ocalymma  verruciferum  (Schldl.)  Miers,  Ann. 
Mag.  Nat.  Hist.,  ser  3,  7:  393.  1861.  A.  fissum 
Loes.,  Verh.  Bot.  Vereins  Prov.  Brandenberg 
65:  102.  1923.  A.  seleri  Loes.,  Verh.  Bot.  Ver- 
eins Prov.  Brandenberg  65:  101.  1923.  Ono- 
hualcoa  seleri  (Loes.)  Lundell,  Contrib.  Univ. 
Michigan  Herb.  7:  52.  1942.  O.  fissa  (Loes.) 
Sandw.,  Kew  Bull.  1946:  88.  1947.  A.  perezii 
Standl.  &  L.  O.  Williams,  Ceiba  3:  61.  1952. 
O.  verrucifera  (Schldl.)  A.  Gentry,  Ann.  Mis- 
souri Bot.  Gard.  60:  885.  1973  (1974).  Figure 
18. 

Lianas  to  20  m  high,  to  over  8  cm  diam.,  ten- 
drils to  15  cm  long,  leafy  stems  1.3-9  mm  diam., 
terete  or  slightly  quadrangular,  subglabrous  or 
sparsely  and  minutely  puberulent  with  peltate  or 
straight  hairs  ca.  0. 1  mm  long,  V-shaped  interpe- 
tiolar  ridges  often  present,  gland  fields  absent; 
pseudostipules  inconspicuous.  Leaves  3-  or  2-fo- 
liolate, petioles  2-9  cm  long,  1.3-2.7  mm  diam., 
subglabrous,  longitudinally  striate,  lateral  petio- 
lules 8-35  mm  long,  terminal  petiolule  3-7  cm 
long;  leaflet  blades  6-14(-17)  cm  long,  2.5-9(-l  1) 
cm  wide,  ovate-elliptic  to  ovate-triangular  or 
ovate-oblong,  apex  acute  to  acuminate,  base  ob- 
tuse to  rounded,  drying  chartaceous,  upper  surface 
glabrous  except  for  the  midvein,  lower  surface 
subglabrous,  often  with  gland  fields  in  proximal 
vein  axils  beneath,  2°  veins  5-7/side,  basal  sec- 
ondary veins  strongly  ascending.  Inflorescences 
axillary  to  leaves  or  fallen  leaves,  3-12  cm  long, 


BURGER:  FLORA  COSTARICENSIS 


135 


racemose  or  narrowly  paniculate  (thyrse)  with  3- 
15  flowers,  peduncle  puberulent  and  lenticellate, 
pedicels  4-7  mm  long,  ca.  1  mm  diam.,  sparsely 
to  densely  puberulent  with  hairs  0.1-0.2  mm  long. 
Flowers  with  calyx  8-12  mm  long,  4-6  mm 
diam.,  narrowly  cupuiar,  surface  sparsely  to 
densely  minutely  puberulent,  with  5  longitudinal 
ridges  terminating  in  apical  teeth  0.5-3  mm  long; 
corolla  47-82  mm  long,  tubular-campanulate,  li- 
lac to  purple,  puberulent  with  thin  curved  hairs 
ca.  0.2  mm  long  externally,  lobes  13-35  mm  long, 
rounded  distally,  throat  white;  filaments  23-24 
and  11-16  mm  long,  thecae  ca.  4  mm  long.  Fruits 
13-30  cm  long,  17-33  mm  wide,  linear-oblong, 
valves  flat  to  rounded  and  covered  with  tubercles 
1-5  mm  long  from  a  base  1-4  mm  wide;  seeds 
12-17  mm  long,  37-58(-80)  mm  wide,  wings  dis- 
tally translucent,  central  area  poorly  differentiat- 
ed. 

Rarely  collected  plants  of  evergreen  and  partly 
deciduous  forest  formations  along  the  Pacific  slope 
in  Costa  Rica,  10-1 100  m  elevation.  Flowering  in 
January  and  April  (March-June  from  Mexico  to 
Nicaragua);  fruiting  in  December.  The  species 
ranges  from  Mexico  to  Bolivia  and  Guyana. 

Mansoa  verrucifera  is  recognized  by  its  climb- 
ing habit  with  tendrils  that  are  distally  trifid,  op- 
posite two-  or  three-foliolate  leaves  with  gland 
fields  in  vein  axils  beneath,  short  axillary  inflo- 
rescences, prominently  five-dentate  calyx  lobes, 
large  purple  or  rose  corollas  minutely  puberulent 
externally,  and  unusual  woody  fruits  with  verru- 
cose-tuberculate  surface.  The  longer  petiolules  of 
terminal  leaflets  (in  three-foliolate  leaves),  strong- 
ly ascending  basal  secondary  veins,  and  stems 
lacking  well-developed  interpetiolar  gland  fields 
or  pseudostipules  are  additional  distinctions.  This 
species  was  treated  as  Adenocalymma  fissum  in 
Flora  of  Guatemala  (Standley  &  Williams,  1974) 
and  as  Onohualcoa  verrucifera  in  Flora  of  Pan- 
ama (Gentry,  1973b). 


Marti  nella  Bail  Ion 

Lianas  climbing  with  distally  trifid  tendrils, 
stems  terete,  cross-sections  of  stems  with  large 
pith  and  4  phloem  areas,  nodes  with  an  interpe- 
tiolar ridge,  without  glandular  fields;  pseudo- 
stipules  absent.  Leaves  opposite,  2-foliolate,  pet- 
iolate,  often  bearing  a  distally  trifid  tendril,  mar- 
gin entire,  glabrous  or  with  short  glandular  hairs, 
venation  pinnate.  Inflorescences  axillary  ra- 
cemes, pedunculate,  bracts  minute  or  absent,  ped- 


icels prominent.  Flowers  with  calyx  closed  and 
conical  in  bud,  tubular-campanulate,  opening  ir- 
regularly with  4-  or  5-lobed  margin  or  splitting 
and  2-lipped;  corolla  tubular-campanulate  or  fun- 
nelform,  dark  wine-red,  tube  with  a  narrow  base, 
glabrous  or  with  minute  peltate  hairs  externally; 
stamens  4,  included,  filaments  of  2  lengths,  an- 
thers glabrous,  thecae  straight,  divaricate,  a  small 
staminode  present;  ovary  linear-cylindric,  2-loc- 
ular,  ovules  in  4  (2)  series  in  each  locule.  Fruits 
long  linear  capsules,  valves  parallel  to  the  septum, 
thin  and  flat,  smooth,  midvein  obscure,  margins 
slightly  thickened;  seeds  thin,  transversely  oblong 
with  2  poorly  differentiated  lateral  membrana- 
ceous  wings. 

Martinella  is  a  genus  of  two  species,  one  South 
American  and  one  ranging  from  Mexico  to  Brazil 
and  Bolivia.  The  very  long  fruits  and  unusual  col- 
or of  the  corolla  help  distinguish  our  representa- 
tive of  this  genus. 

Martinella  obovata  (Kunth  in  H.B.K.)  Bureau  & 
K.  Schum.  in  Mart.,  Fl.  Bras.  8(2):  161,  pi.  84. 
1896.  Spathodea  obovata  Kunth  in  H.B.K. , 
Nov.  Gen.  Sp.  3:  147.  1819.  Bignonia  obovata 
(Kunth  in  H.B.K.)  Spreng.,  Syst.  Veg.  2:  830. 
1825.  Figure  23. 

Lianas  to  over  25  m  high,  to  7  cm  diam.,  ten- 
drils 5-25  cm  long  with  tips  2-24  mm  long,  leafy 
stems  1 .7-9  mm  diam.,  glabrous  or  minutely  glan- 
dular puberulent,  a  raised  interpetiolar  ridge  usu- 
ally present  at  nodes.  Leaves  2-foliolate,  petioles 
2-7  cm  long,  1-2.3  mm  diam.,  petiolules  12-32(- 
66)  mm  long,  glabrous  or  very  minutely  (0.05 
mm)  puberulent;  leaflet  blades  6-16(-21)  cm 
long,  2.5-8(-15)  cm  wide,  ovate-elliptic  to  ovate- 
oblong  or  broadly  ovate,  apex  short-acuminate  or 
acuminate  with  narrow  tip,  base  obtuse  to  rounded 
(subcordate)  and  slightly  asymmetric,  drying 
chartaceous  and  often  dark,  glabrous  on  both  sur- 
faces with  very  minute  hairs  along  the  major 
veins  beneath,  flat  glands  often  present  along  the 
midvein  beneath,  2°  veins  3-7/side.  Inflorescenc- 
es axillary,  5-15  cm  long,  racemes  with  1-21 
flowers,  peduncles  16-26  mm  long,  ca.  1.5  mm 
diam.,  glabrous  or  very  minutely  puberulent, 
bracts  less  than  1  mm  long,  pedicels  6-14  mm 
long.  Flowers  with  calyx  11-16(-21)  mm  long, 
6-9(-12)  mm  diam.,  tubular  or  tubular-campanu- 
late, margin  irregularly  split  or  2-4-lobed,  surface 
glabrous  with  many  parallel  longitudinal  veins  or 
minutely  puberulent;  corolla  48-65  mm  long, 
campanulate  above  a  narrowed  (3-5  mm  diam.) 


136 


FIELDIANA:  BOTANY 


tubular  base,  dark  reddish  purple  or  maroon,  8- 
21  mm  wide  at  the  mouth,  lobes  6-15  mm  long, 
glabrous  or  with  minute  glandular  peltate  hairs 
externally;  filaments  15-17  and  11-14  mm  long, 
thecae  2.5-3  mm  long.  Fruits  31-130  cm  long, 
14-22  mm  wide,  2-4  mm  thick,  valves  flat  with 
smooth  surface,  acute  at  the  ends,  midveins  slight- 
ly elevated  or  obscure,  margins  thickened;  seeds 
12-15  mm  long,  (30-H3-60  mm  wide,  central 
area  ca.  17  mm  wide,  wings  membranaceous,  pale 
brownish. 

Uncommon  plants  of  moist  evergreen  forest 
formations  on  both  Caribbean  and  Pacific  slopes, 
20-1600  m  elevation.  Flowering  in  February-July 
in  Costa  Rica.  This  species  ranges  from  Mexico 
to  Brazil  and  Bolivia. 

Martinella  obovata  is  recognized  by  its  climb- 
ing habit  with  distally  trifid  tendrils,  opposite  two- 
foliolate  leaves  often  with  long  petioles,  axillary 
few-flowered  racemes,  tubular  calyx  with  irregu- 
lar margins,  deep  purple  or  maroon  corollas,  and 
long  thin  linear  fruits  with  smooth  flat  valves.  In 
addition,  the  petioles  and  petiolules  are  often 
twisted,  and  the  leaflets  are  lustrous  above  in  life. 
The  fruits  can  be  more  than  1  m  long. 


Melloa  Bureau 

Lianas,  tendrils  with  3  terminal  claw-like  arms, 
stems  terete,  cross-section  with  irregular  phloem 
areas,  interpetiolar  glandular  fields  present  or  ab- 
sent; pseudostipules  ovate  to  subulate.  Leaves  op- 
posite, 2-foliolate,  petiolate,  often  with  a  tendril, 
margins  entire,  glabrous,  venation  pinnate.  Inflo- 
rescences terminal  or  axillary,  panicles  or  few- 
flowered  cymes,  foliaceous  bracts  present  and  ear- 
ly caducous,  pedicels  prominent.  Flowers  with  tu- 
bular-campanulate  thin-textured  calyx,  distally  ir- 
regularly lobed  or  spathaceous  with  recurved  tip; 
corolla  tubular-funnelform,  yellow,  glabrous  ex- 
ternally; stamens  4,  filaments  of  2  lengths,  anthers 
glabrous,  thecae  straight,  divaricate,  a  staminode 
present;  ovary  flattened-ovoid,  2-locular,  ovules 
multiseriate  in  each  locule.  Fruits  capsules,  ellip- 
soid-oblong and  slightly  flattened,  valves  parallel 
to  the  septum,  thick  and  woody,  splitting  along 
the  middle  at  maturity;  seeds  thin,  transversely 
oblong  with  2  lateral  wings,  seed  body  well  dif- 
ferentiated from  the  membranaceous  wings. 

Melloa  has  a  single  species,  ranging  from  Mex- 
ico to  Argentina. 


Melloa  quadrivalvis  (Jacq.)  A.  Gentry,  Brittonia 
25:  237.  1973.  Bignonia  quadrivalvis  Jacq., 
Fragm.  Bot.  37,  tab.  40,  fig.  3.  1800-1809.  Fig- 
ure 17. 

Lianas  to  over  20  m  high,  tendrils  with  3  stiff- 
ened curved  tips  11-18  mm  long,  leafy  stems  1.3- 
6  mm  diam.,  glabrous,  longitudinally  striate,  in- 
terpetiolar ridge  usually  present;  pseudostipules 
1.5-3.5  mm  long,  subulate.  Leaves  with  petioles 
1 1-55  mm  long,  1-2  mm  diam.,  glabrous,  striate, 
petiolules  6-20  mm  long;  leaflet  blades  4-1 1  (-15) 
cm  long,  2-5(-7)  cm  wide,  ovate-elliptic  to  ovate- 
lanceolate  or  elliptic,  apex  acute  to  acuminate, 
base  obtuse  or  slightly  rounded  and  truncate,  dry- 
ing chartaceous,  glabrous  above  and  below,  2° 
veins  3-6/side.  Inflorescences  axillary  or  termi- 
nal (often  on  lateral  short  shoots),  paniculate  or 
dichotomous  cymes,  mostly  3-9  flowered,  flowers 
usually  in  distal  triads  (rarely  solitary),  peduncles 
13-36  mm  long,  0.8-1.3  mm  diam..  glabrous  and 
drying  dark,  with  caducous  bracts  7-21  mm  long 
and  3-6  mm  wide,  pedicels  16-32  mm  long. 
Flowers  with  tubular-campanulate  calyx.  10-21 
mm  long,  9-13  mm  diam.,  surface  glabrous  (rare- 
ly puberulent),  without  glands,  margin  split  and 
spathaceous  or  irregularly  lobed;  corolla  35-75 
mm  long,  tubular-funnelform,  bright  yellow,  gla- 
brous or  subglabrous  externally  (lobes  ciliolate), 
tube  8-15  mm  diam.,  lobes  10-18  mm  long;  fil- 
aments 21-28  and  14-18  mm  long,  thecae  4-5 
mm  long;  disc  1.5-2  mm  high  (subtended  by  a 
broad  thin  layer  and  appearing  double).  Fruits  8- 
15  cm  long,  25-52  mm  wide,  15-20  mm  thick, 
ellipsoid-oblong  with  narrowed  ends,  valves  5-8 
mm  thick,  each  splitting  longitudinally  into  2 
equal  parts,  surface  dark  brown  with  many  pale 
lenticel-like  spots;  seeds  10-16  mm  long,  25-52 
mm  wide,  central  area  9-12  mm  wide  and  clearly 
differentiated  from  the  transparent  wings. 

Uncommon  lianas  of  seasonally  dry  deciduous 
or  partly  deciduous  (less  often  evergreen)  forest 
formations  on  both  Caribbean  and  Pacific  slopes, 
10-1000  m  elevation.  Flowering  in  February-Au- 
gust; fruiting  throughout  the  year.  This  species 
ranges  from  northeastern  Mexico  to  Argentina. 

Melloa  quadrivalvis  is  recognized  by  its  climb- 
ing habit  with  three-clawed  tendrils,  opposite  two- 
foliolate  glabrous  leaves,  few-flowered  inflores- 
cences, bright  yellow  corollas,  and  thick  woody 
ellipsoid-oblong  fruits  that  split  into  four  parts. 
The  large  floral  bracts  would  be  a  useful  feature 
for  determination,  but  they  are  quickly  caducous 
and  rarely  collected.  These  plants  are  similar  to 


BURGER:  FLORA  COSTARICENSIS 


137 


those  of  Adenocalymma  innundatum  (q.v.),  but 
their  fruits  are  very  different. 


Mussatia  Bureau  ex  Baillon 

Lianas,  climbing  with  distally  coiled  simple 
tendrils,  branchlets  quadrangular,  with  8  (16) 
phloem  areas  in  cross-section,  nodes  without  in- 
terpetiolar  gland  fields;  pseudostipules  large  and 
resembling  small  leaves.  Leaves  opposite,  petio- 
late,  2-foliolate,  often  with  a  terminal  tendril,  ten- 
dril with  1  tip,  leaflets  petiolulate,  margins  entire, 
venation  pinnate,  domatia  present.  Inflorescences 
terminal,  panicles  with  opposite  branching,  bracts 
inconspicuous,  flowers  pedicellate.  Flowers  with 
short  calyx  forming  a  shallow  cup,  margin  entire 
to  lobed  or  split;  corolla  funnelform  and  2-lipped, 
yellow  with  red  or  brownish  stripes  within,  mi- 
nutely glandular  with  peltate  hairs  externally;  sta- 
mens 4,  included,  filaments  of  2  lengths,  anthers 
glabrous,  thecae  short,  divergent  or  divaricate; 
disc  cupular,  ovary  2-locular,  ovules  in  4-6  series 
in  each  locule,  stigma  simple.  Fruits  oblong  to 
linear-oblong  capsules,  valves  parallel  to  the  sep- 
tum, woody,  flat  or  slightly  convex,  surface  tu- 
berculate  to  muriculate;  seeds  thin,  with  2  poorly 
differentiated  lateral  wings. 

Mussatia  is  a  genus  of  two  species;  the  other 
species  is  found  in  eastern  Amazonia  and  Guiana. 

Mussatia  hyacinthina  (Standl.)  Sandw.,  Recueil 
Trav.  Bot.  Neerl.  34:  218.  1937.  Tynanthus  hy- 
acinthinus  Standl.,  Carnegie  Inst.  Wash.  Publ. 
461:  87.  1935.  Figure  19. 

Lianas  to  30  m  high,  stems  to  18  cm  diam., 
tendrils  12-23  cm  long,  leafy  stems  2.5-8  mm 
diam.,  strongly  quadrangular  with  4  raised  longi- 
tudinal ridged  edges,  minutely  (0.1  mm)  puberu- 
lent  but  soon  glabrescent  and  pale  gray;  pseudo- 
stipules  usually  4/node,  3-18  mm  long,  ovate  or 
rounded.  Leaves  with  petioles  4-9(-12)  cm  long, 
1.5-2.5  mm  diam.,  subglabrous  or  minutely  pu- 
berulent,  petiolules  1.4-4.5(-6)  cm  long;  leaflet 
blades  7-20(-26)  cm  long,  4-13(-17)  cm  wide, 
broadly  ovate  to  ovate-elliptic  or  ovate-suborbic- 
ular,  apex  acute  to  acuminate,  base  obtuse  or 
rounded  and  truncate,  drying  chartaceous,  gla- 
brous above,  minutely  papillate-puberulent  on  the 
veins  or  subglabrous  beneath,  with  longer  hairs  in 
vein  axils  (domatia),  2°  veins  4-7/side,  minor  ve- 
nation slightly  raised  above  (dried).  Inflorescenc- 
es terminal  (often  on  short  lateral  shoots),  5-16 


cm  long,  peduncles  3-40  mm  long,  ca.  1.5  mm 
diam.,  with  minute  (0.1  mm)  peltate  hairs,  dark 
brown,  2°  peduncles  ca.  1  cm  long,  bracts  0.5-1 
mm  long,  pedicels  2-5  mm  long  (above  bract 
scars).  Flowers  with  broadly  campanulate  calyx 
1.2-2.3  mm  long,  3-3.5  mm  wide,  margin  entire, 
glabrous;  corolla  15-24  mm  long,  funnelform  and 
bilabiate,  white  with  maroon  tips,  becoming  yel- 
lowish brown  or  reddish  brown,  minutely  papil- 
late-puberulent, tube  2.5-7  mm  diam.,  lobes  5-10 
mm  long;  filaments  ca.  8-12  and  6-7  mm  long; 
ovary  3  mm  long,  with  minute  peltate  hairs. 
Fruits  17-37  cm  long,  4-6  cm  wide,  ca.  2  cm 
thick,  valves  obtuse  at  both  ends,  surface  muricate 
with  rounded  projections  ca.  1  mm  high,  brown; 
seeds  20-43  mm  long,  50-120  mm  wide,  central 
area  20-40  mm  wide,  wings  membranaceous, 
pale  brown. 

Uncommon  plants  of  evergreen  rain  forest  for- 
mations on  both  the  Caribbean  and  Pacific  slopes, 
50-900  m  elevation.  Flowering  in  February-May 
in  Central  America.  This  species  ranges  from 
eastern  Mexico  to  Brazil. 

Mussatia  hyacinthina  is  recognized  by  the 
woody  climbing  habit,  two-foliolate  leaves,  small 
flowers,  short  calyx  cups  with  entire  margins,  bi- 
labiate white  or  yellowish  brown  corollas,  long 
thick  muricate  woody  fruits,  and  large  winged 
seeds.  Leaves  with  hairs  in  vein  axils  beneath 
(domatia),  square  stems  with  four  prominent  ridg- 
es, rounded  pseudostipules,  and  tendrils  with  sim- 
ple tips  are  useful  vegetative  distinctions.  Com- 
pare this  species  with  Cydista  diversifolia,  which 
has  similar  stems  and  pseudostipules.  The  fruits 
are  similar  to  those  of  Callichlamys,  where  the 
valves  have  a  smoother  surface. 

The  genera  Onohualcoa  and  Pachyptera  are 
now  considered  to  be  synonyms  of  Mansoa  (q.v.). 


Parabignonia  Bureau  ex  K.  Schumann 

Lianas  climbing  with  tendrils  with  3  stiff 
curved  distal  "claws,"  stems  terete,  with  4  phlo- 
em areas  in  cross-section,  interpetiolar  gland 
fields  present;  pseudostipules  small  and  ovate. 
Leaves  opposite,  2-foliolate,  petiolate,  blades  en- 
tire, usually  glabrous,  venation  pinnate  or  subpal- 
mate.  Inflorescences  axillary,  very  short  and  few- 
flowered,  usually  solitary  racemes,  bracts  short 
and  narrow,  pedicels  subglabrous  or  minutely  pu- 
berulent.  Flowers  with  tubular-campanulate  ca- 
lyx, 4-  or  5-lobed,  rounded  or  acute  at  the  apex; 


138 


FIELDIANA:  BOTANY 


corolla  funnelform-campanulate  and  somewhat  2- 
lipped,  glabrous  externally  or  with  minute  pub- 
erulence  on  the  lobes,  lobes  5,  rounded;  stamens 
4,  included,  filaments  of  2  lengths,  anthers  gla- 
brous, thecae  straight,  divergent,  staminode  pre- 
sent; ovary  oblong,  with  minute  peltate  hairs,  2- 
locular,  ovules  2-seriate  in  each  locule.  Fruits 
capsules,  narrowly  linear,  valves  parallel  to  the 
septum  but  dehiscing  perpendicular  to  it;  seeds 
thin,  transversely  oblong,  with  2  membranaceous 
lateral  wings. 

A  genus  of  two  species  (more  if  Paradolichan- 
dra  is  included),  one  ranging  from  southern  Costa 
Rica  to  Venezuela  and  the  other  in  southern  Bra- 
zil. Recently  collected  material  (1992-1993)  is  the 
first  record  of  the  genus  in  Central  America. 

Parabignonia  steyermarkii  Sandw.,  Bol.  Soc. 
Venez.  Cienc.  Nat.  26:  446.  1966.  Figure  17. 

Lianas  climbing  to  over  25  m  high,  stems  to 
19  cm  diam.,  tendrils  5-12  cm  long,  with  3 
curved  tips,  leafy  stems  2-6  mm  diam.,  glabrous 
(in  ours)  or  with  short  thin  hairs,  terete,  becoming 
brown  with  elliptic  lenticels  ca.  1  mm  long;  in- 
terpetiolar  gland  fields  inconspicuous.  Leaves  2- 
foliolate,  petioles  10-28  mm  long,  1.2-1.7  mm 
diam.,  petiolules  4-1 1  mm  long,  glabrous,  drying 
dark;  leaflet  blades  4-11  cm  long,  2-6.5  cm 
wide,  ovate-elliptic,  ovate,  ovate-oblong,  or  ellip- 
tic-oblong, apex  acute  or  short-acuminate,  base 
obtuse  to  slightly  rounded,  drying  stiffly  charta- 
ceous  and  dark,  glabrous  above  and  below,  2° 
veins  4-7/side,  basal  pair  strongly  ascending 
(subtripli veined).  Inflorescences  axillary,  1-5- 
flowered  racemes  or  corymbs  or  flowers  solitary, 
1-3  cm  long,  peduncle  1-5  mm  long,  glabrous, 
drying  dark,  bracts  1.3-2.5  mm  long,  linear-lan- 
ceolate or  subulate,  pedicels  6-18  mm  long,  gla- 
brous or  very  minutely  (0.03  mm)  puberulent. 
Flowers  with  calyx  6-12(-22)  mm  long,  4-7(-14) 
mm  wide,  cupulate  or  campanulate,  subglabrous, 
drying  dark,  lobes  2-4(-7)  mm  long,  rounded  to 
obtuse;  corolla  35-62  mm  long,  funnelform-cam- 
panulate, deep  magenta  (in  ours)  to  rose-purple  or 
pale  lavender,  10-15  mm  wide  at  the  mouth,  tube 
glabrous  but  the  lobes  minutely  papillate  puber- 
ulent externally,  lobes  10-17  mm  long;  filaments 
17-24  and  12-16  mm  long,  thecae  2.5-3.5  mm 
long.  Fruits  not  seen  at  maturity,  young  fruits  ca. 
10  cm  long,  4-5  mm  diam.,  valves  glabrous; 
seeds  unknown. 

Rarely  collected  climbers  of  lowland  evergreen 
rain  forest  formations  in  the  Golfo  Dulce  area  at 


300-450  m  elevation  (300-1200  m  in  Venezuela). 
Collected  with  flowers  in  January  (Aguilar  817, 
Gentry  el  al.  78682)  at  Rancho  Quemada  in  the 
Reserva  Forestal  Golfo  Dulce.  This  species  is 
known  only  from  southwestern  Costa  Rica  and 
Venezuela. 

Parabignonia  steyermarkii  is  recognized  by  its 
climbing  habit  with  distally  3-clawed  tendrils,  op- 
posite 2-foliolate  leaves,  short  few-flowered  axil- 
lary inflorescences,  4-  or  5-lobed  calyx,  and  ma- 
genta corollas  with  externally  glabrous  tube  and 
minutely  puberulent  lobes.  Costa  Rican  collec- 
tions seem  to  have  broader  thinner  leaves  than 
those  from  Venezuela.  In  addition,  the  calyx  is 
smaller  in  Costa  Rican  material  (Venezuelan  mea- 
surements are  in  parenthesis  in  the  description 
above). 


Paragonia  Bureau 

REFERENCE — W.  D.  Hauk.  A  review  of  the  ge- 
nus Paragonia  (Bignoniaceae).  Ann.  Missouri 
Bot.  Card.  85:  460.  1998. 

Lianas  climbing  with  the  aid  of  minutely  sim- 
ple, bifid  or  trifid  tendrils,  stems  terete,  with  4 
phloem  areas  in  cross-section,  becoming  lenticel- 
late,  glabrate  to  lepidote  or  puberulent,  nodes 
lacking  gland  fields;  pseudostipules  conical  with 
broad  base  and  narrow  tip,  usually  curved  in  to- 
ward the  stem.  Leaves  opposite,  2-foliolate,  pet- 
ioles usually  with  gland  fields  near  the  apex, 
blades  entire,  glabrate  to  densely  puberulent,  pin- 
nately  veined.  Inflorescences  terminal  or  axillary, 
paniculate,  often  large  and  many-flowered,  mi- 
nutely bracteate,  pedicellate.  Flowers  with  cupu- 
late calyx,  with  minute  peltate  hairs,  distally  trun- 
cate or  irregularly  lobed  or  split,  ciliolate;  corolla 
tubular-campanulate,  rose  red  to  lilac  or  magenta, 
densely  and  minutely  puberulent  with  crooked 
moniliform  hairs;  stamens  4,  filaments  of  2 
lengths,  anthers  glabrous,  thecae  straight,  divari- 
cate; disc  prominent,  ovary  narrowly  cylindric, 
locules  2,  ovules  in  2  series  in  each  locule.  Fruits 
woody  capsules,  linear  to  linear-oblong,  valves 
flattened  parallel  to  the  septum,  surface  smooth  to 
tuberculatc  (in  ours),  midvein  slightly  elevated  or 
not  apparent;  seeds  thin,  transversely  oblong  with 
2  lateral  wings,  brown,  the  central  area  not  dif- 
ferentiated from  the  wings. 

Paragonia  is  a  genus  of  two  species,  one  re- 
stricted to  eastern  Brazil. 


BURGER:  FLORA  COSTARICENSIS 


139 


Paragonia  pyramidata  (L.  C.  Rich.)  Bureau,  Vi- 
densk.  Meddel.  Dansk.  Naturhist.  Foren.  Kjob- 
enhavn  1893:  104.  \S94.Bignoniapyramidata 
L.  C.  Rich.,  Actes  Soc.  Hist.  Nat.  Paris  1:  110. 
1792.  B.  sinclairii  Benth.,  Bot.  Voy.  Sulphur 
129.  1844.  Tabebuia  pyramidata  (L.  C.  Rich.) 
DC.,  Prodr.  9:  214.  1845.  Arrabidaea  dichasia 
J.  D.  Smith,  Bot.  Gaz.  20:  6.  1895.  Figure  23. 

Lianas,  stems  to  5  cm  diam.,  often  with  thick- 
ened nodes,  tendrils  5-17  cm  long,  distally  simple 
or  bifid  (rarely  trifid),  leafy  stems  2-6  mm  diam., 
glabrous  or  less  often  minutely  puberulent,  inter- 
petiolar  ridge  often  present;  pseudostipules  3-7 
mm  long,  sharply  acute.  Leaves  2-foliolate,  peti- 
oles 9-45  mm  long,  1.4-3  mm  diam.,  subgla- 
brous  to  minutely  puberulent,  often  with  glands 
near  the  apex,  petiolules  7-35(-46)  mm  long,  sul- 
cate  above;  leaflet  blades  6-19(-25  cm)  long,  3- 
9(-13)  cm  wide,  elliptic-oblong  to  elliptic  or 
broadly  ovate-oblong,  apex  acute  to  short-acumi- 
nate or  rounded,  base  obtuse  to  slightly  rounded, 
drying  stiffly  chartaceous,  glabrous  above,  gla- 
brous beneath  or  minutely  puberulent  on  the  mid- 
vein  beneath,  with  minute  (0.05  mm)  punctate  de- 
pressions beneath,  2°  veins  4-8/side.  Inflores- 
cences terminal,  10-30  cm  long,  peduncles  3-11 
cm  long,  ca.  2  mm  diam.,  minutely  puberulent, 
pedicels  4-9  mm  long.  Flowers  with  cupular  ca- 
lyx 4-7  mm  long,  4-8  mm  diam.,  subglabrous  or 
minutely  puberulent,  margin  entire  to  split  or  with 
minute  (0.3  mm)  teeth;  corolla  35-65  mm  long, 
tubular-campanulate,  pink  to  lavender  or  magenta 
(white  at  the  base),  densely  puberulent  with 
crooked  hairs  0.1-0.3  mm  long  externally,  12-22 
mm  wide  at  the  mouth;  filaments  ca.  18  and  13- 
16  mm  long,  thecae  2-2.5  mm  long.  Fruits  (10- 
)32-61  cm  long,  10-17  mm  wide,  5-8  mm  thick, 
narrowed  at  base  and  apex,  valves  slightly  rough 
to  the  touch  with  muricate  surface  of  small  pro- 
jections; seeds  8-12  mm  long,  21-43  mm  wide, 
oblong  with  blunt  tips,  uniformly  grayish  brown 
to  dark  brown.  Common  plants  of  evergreen  low- 
land rain  forest  formations  on  both  Caribbean  and 
Pacific  slopes,  5-800  m  elevation.  They  are  found 
on  well-drained  slopes  and  in  poorly  drained 
swamps.  Flowering  occasionally  throughout  the 
year  but  with  the  majority  of  flowering  in  Janu- 
ary-May. The  species  ranges  from  Mexico  to 
Brazil  and  Bolivia. 

Paragonia  pyramidata  is  recognized  by  its 
climbing  habit  with  simple  or  bifid  tendrils,  op- 
posite two-foliolate  leaves  with  gland  field  at  pet- 
iole apex,  many-flowered  terminal  panicles,  cu- 


pular calyx  with  irregular  margin,  pink  to  magenta 
corollas  densely  puberulent  on  the  exterior,  long 
linear  fruits  with  flat  muricate  surfaces,  and  thin, 
uniformly  brown,  blunt-ended,  two-winged  seeds. 
The  pseudostipules  and  sweet  smell  when  leaves 
are  crushed  are  useful  vegetative  features.  The  im- 
mature calyx  has  a  conical  apex  that  sometimes 
splits  off  in  a  calyptrate  fashion  as  in  Lundia 
(q.v.).  Compare  also  the  vegetatively  similar  Cer- 
atophytum  tetragonolobium,  which  differs  in  hav- 
ing trifid  tendrils  and  interpetiolar  gland  fields. 

Parmentiera  DeCandolle 

REFERENCE — A.  Gentry,  Parmentiera  in  Big- 
noniaceae — part  I  (Crescentiae  and  Tourrettieae). 
Flora  Neotropica  Monogr.  25(1):  1-130.  1980. 


Trees  or  shrubs,  stems  terete,  leaves  subtended 
by  simple  spines  or  hard  smooth  shelf-like  tissue, 
an  interpetiolar  line  or  ridge  absent;  pseudostip- 
ules absent.  Leaves  opposite  or  subopposite  (oc- 
casionally alternate),  sometimes  fasciculate,  usu- 
ally 3-  or  5-foliolate  (rarely  simple  or  2-  or  4- 
foliolate),  petioles  sometimes  with  narrow  lateral 
wings,  leaflets  mostly  glabrous,  entire  or  with  few 
distal  teeth,  venation  pinnate,  domatia  usually 
present  in  vein  axils  beneath.  Inflorescences  ter- 
minal or  often  on  leafless  short-shoots  or  cauliflo- 
rous  on  trunk  and  older  stems,  with  1-3  flowers, 
bracts  minute,  pedicels  drying  black.  Flowers 
drying  black,  calyx  tubular,  entire  distally  but  split 
abaxially  and  spathe-like,  subglabrous;  corolla 
white  to  greenish  white,  tubular-campanulate  and 
often  bent  in  the  middle  (with  a  transverse  ridge 
across  the  lower  side)  of  the  throat,  2-lipped  with 
2  upper  lobes  partly  united,  thick-textured,  gla- 
brous externally;  stamens  4,  subexserted  near  the 
mouth,  filaments  subequal,  thecae  thick,  straight, 
somewhat  divergent,  a  staminode  present;  disc  an- 
nular; ovary  cylindric,  2-locular,  ovules  multiser- 
iate.  Fruits  fleshy  and  indehiscent,  elongate  ob- 
long to  linear,  terete  or  with  prominent  longitu- 
dinal ridges,  surface  smooth,  outer  cortex  firm  and 
fleshy;  seeds  borne  on  a  fibrous-fleshy  central 
core,  small,  flattened,  with  or  without  a  narrow 
circumferential  wing. 

Parmentiera  is  a  genus  of  ten  species  ranging 
from  Mexico  to  Panama,  with  one  in  Colombia. 
The  genus  is  unusual  because  of  its  glabrous  flow- 
ers that  dry  black,  tubular  calyx  split  down  one 
side,  large  white  or  greenish  white  two-lipped  co- 
rollas with  thick  texture  that  usually  have  a  trans- 


140 


FIELDIANA:  BOTANY 


verse  fold  at  the  bottom  of  the  tube,  and  the  elon- 
gate fleshy  fruits.  The  usually  opposite  three-  or 
five-foliolate  leaves,  blades  with  domatia  beneath, 
and  hard  spines  or  hard  tissue  at  leaf  bases  are 
additional  distinguishing  features.  All  of  the  spe- 


cies are  probably  pollinated  by  bats,  with  seeds 
adapted  to  dispersal  by  mammals.  Our  introduced 
species  may  resemble  Crescentia  alata,  but  that 
species  has  leaves  in  alternate  fascicles  and  the 
petioles  are  broadly  winged. 


Key  to  the  Species  of  Parmentiera 

la.  Native  trees  found  in  evergreen  forest  formations;  leaves  usually  3-foliolate  or  5-foliolate,  petioles 
lacking  narrow  green  margins,  larger  leaf  blades  3-17  cm  long 2 

Ib.  Introduced  trees  planted  in  gardens  and  parks  (rarely  collected);  leaves  mostly  3-foliolate.  petioles 

usually  with  narrow  green  margins  or  wings,  larger  leaf  blades  2-9  cm  long  4 

2a.  Leaves  usually  5-foliolate,  largest  blades  to  9  cm  long;  600-1500  m  elevation  in  the  Cordillera 

de  Guanacaste  and  near  Tilaran P.  valerii 

2b.  Leaves  usually  3-foliolate,  largest  blades  to  17  cm  long;  wet  forests  of  the  Caribbean  slope  10- 

750  m  3 

3a.  Corollas  ca.  25  mm  long;  fruits  ca.  15  mm  diam.;  rarely  collected  P.  dressleri 

3b.  Corollas  50-75  mm  long;  fruits  ca.  30-60  mm  diam.;  more  commonly  collected  

P.  macrophylla 

4a.  Stems  usually  with  2  small  hard  spines  subtending  each  node;  flowers  terminal  or  in  axils  of  distal 
leaves;  fruits  >  30  mm  diam.  in  life;  petioles  with  very  narrow  margins  P.  aculeata 

4b.  Stems  without  spines;  flowers  borne  on  trunks  and  older  branches;  fruits  <  25  mm  diam.  in  life; 
petioles  often  with  winged  margins  P.  cereifera 


Parmentiera  aculeata  (Kunth  in  H.B.K.)  Seem., 
Bot.  Voy.  Herald  183.  1854.  Crescentia  aculea- 
ta Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3:  158. 
1819.  C.  edulis  Moc.  ex  DC.,  Prodr.  9:  244. 
1845,  non  C.  edulis  Desv.  P.  edulis  DC.,  Prodr. 
9:  244.  1845.  Figure  11. 

Shrubs  to  medium-size  trees  2-6  m  tall,  leafy 
stems  1-10  mm  diam.,  glabrous,  terete,  leaf  bases 
subtended  by  a  forward-pointing  sharp-tipped 
conical  spine  3-8  mm  long,  with  smooth  glossy 
surface.  Leaves  opposite  or  occasionally  alter- 
nate, 3-foliolate  or  simple,  2-4/node,  petioles  1 1- 
47  mm  long,  0.6-1.8  mm  wide,  with  narrow 
wings  0.1-0.5  mm  wide,  petiolules  0-15  mm 
long;  leaflet  blades  25-90  mm  long  (distal  leaf- 
lets), 6-40  mm  wide,  elliptic  to  rhombic-elliptic, 
apex  obtuse,  base  cuneate  and  decurrent  on  the 
petiole,  drying  chartaceous,  glabrous  above,  with 
depressions  and  minute  hairs  in  the  vein  axils  be- 
neath (domatia),  2°  veins  3-7/side.  Inflorescences 
of  1-4  terminal  flowers  or  1  or  2  flowers  in  axils 
of  near-terminal  leaves  or  fasciculate  on  older 
stems,  pedicels  12-21  mm  long,  0.7-1  mm  diam., 
with  few  minute  peltate  trichomes,  drying  dark. 
Flowers  with  calyx  25-35  mm  long,  spathe-like 
or  split  more  than  once,  glabrous,  drying  dark; 
corolla  4-6.5  cm  long,  campanulate,  white  or 


greenish  with  purple  lines,  glabrous  externally, 
tube  20-25  mm  wide  at  the  mouth;  thecae  8-9 
mm.  Fruits  9-17  cm  long,  to  3  cm  diam.  in  life 
(12-20  mm  diam.  when  dried),  linear-cylindric 
with  narrowed  tips,  usually  slightly  curved,  with 
thick  longitudinal  ridges. 

Pannentiera  aculeata  is  recognized  by  the  tree 
habit  (usually  with  a  short  trunk  and  dense 
crown),  nodes  with  short  spines,  mostly  opposite 
and  trifoliolate  leaves,  spathe-like  calyx,  white  or 
greenish  corollas,  and  slightly  curved  succulent 
fruits  that  are  edible.  This  species  has  not  been 
collected  in  Costa  Rica,  but  Standley  (1938)  re- 
ported it  to  be  cultivated  in  the  hot  lowlands.  Its 
natural  range  is  Mexico  to  Honduras.  Common 
names  in  northern  Central  America  are  cuajilote, 
pepino  de  arbol,  and  cow  okra. 

Parmentiera  cereifera  Seem,  in  Hook.,  J.  Bot. 
3:  302.  1841. 

Small  trees  to  7  mm  tall,  trunks  to  20  cm  diam., 
with  strongly  ascending  main  branches  from  near 
the  ground  and  open  crown,  leafy  stems  1.2-6 
mm  diam.,  glabrous,  terete,  usually  with  hard 
shelf-like  tissue  below  the  leaf  base.  Leaves  op- 
posite, often  with  a  smaller  leaf  in  the  axil  of  a 
larger  (4/node),  3-foliolate,  petioles  l-4(-6)  cm 


BURGER:  FLORA  COSTARICENSIS 


141 


long,  to  3  mm  wide  with  narrow  greenish  wings, 
petiolules  not  clearly  differentiated;  leaflet  blades 
0.5_7(-9.5)  cm  long,  0.5-3(-4)  cm  wide,  elliptic 
to  elliptic-rhombic,  terminal  leaflets  larger  than 
the  laterals,  apex  acute  to  acuminate,  base  acute 
and  decurrent  on  the  petiole,  glabrous  above,  with 
small  hairs  in  vein  axils  beneath  (domatia),  2° 
veins  3-6/side.  Inflorescences  of  solitary  flowers 
or  2  or  3  flowers  on  short-shoots  borne  on  trunks 
and  larger  branches,  peduncles  (to  bracts)  ca.  3 
mm  long,  pedicels  9-16  mm  long,  0.7-1  mm 
diam.,  glabrous  or  with  minute  peltate  trichomes, 
drying  black.  Flowers  with  calyx  2-5  cm  long, 
9-25  mm  diam.,  split  and  spathe-like,  glabrous; 
corolla  37-64  mm  long,  tubular-campanulate  and 
with  a  transverse  fold  on  the  lower  side,  white, 
glabrous  externally,  tube  18-29  mm  wide  at  the 
mouth,  lobes  ca.  10  mm  long;  filaments  26-35 
mm  long.  Fruits  30-54  cm  long,  1-2.5  cm  diam., 
linear-cylindric  and  slightly  curved,  yellow,  gla- 
brous or  with  few  minute  peltate  trichomes;  seeds 
3-4  mm  long,  3-4  mm  wide,  with  a  narrow  mu- 
cilaginous wing. 

Parmentiera  cereifera,  originally  described 
from  the  Caribbean  slope  of  central  Panama,  is 
not  known  to  occur  in  Costa  Rica,  but  it  is  often 
planted  as  a  botanical  curiosity.  The  short  trunk 
with  spreading  branches,  opposite  or  fasciculate 
trifoliolate  leaves,  cauliflorous  flowers,  white  co- 
rolla, and  pendulous  yellow  linear  fleshy  fruits 
make  the  species  quite  distinctive. 

Parmentiera  dressier!  A.  Gentry,  Wrightia  7:  85. 
1982. 

Small  trees  2-3  m  tall,  stems  terete  to  suban- 
gulate,  without  spines  or  prominent  lenticels. 
Leaves  opposite  or  fasciculate,  3-foliolate,  lateral 
petiolules  poorly  distinguished  from  the  base  of 
the  blade,  petioles  3-8  cm  long,  glabrous,  petio- 
lules of  terminal  leaflet  ca.  1-2  cm  long;  leaflet 
blades  5-13  cm  long,  3.5-5.5  cm  wide,  elliptic  to 
elliptic-ovate,  apex  acute  to  acuminate,  base  cu- 
neate,  glabrous  except  for  tufts  of  hairs  in  the  vein 
axils  below  (domatia).  Inflorescences  cauliflorous 
or  ramiflorous,  usually  of  solitary  flowers  borne 
on  pedicels  less  than  1  cm  long,  glabrous.  Flow- 
ers with  spathaceous  calyx  16-17  mm  long,  lep- 
idote  near  the  base,  glabrous  distally,  bluntly 
acute;  corolla  ca.  25  mm  long,  broadly  tubular- 
funnelform,  white,  20  mm  wide  at  the  mouth, 
lobes  ca.  8  mm  long,  glabrous;  stamens  and  pistil 
not  examined.  Fruits  23-24  cm  long,  ca.  15  mm 
wide,  cylindric,  not  ridged. 


Plants  of  rain  forest  formations  in  northern 
Costa  Rica  and  the  Caribbean  lowlands.  Flower- 
ing in  August;  fruiting  in  March  (Panama).  The 
species  ranges  from  northern  Costa  Rica  to  central 
Panama. 

Parmentiera  dressleri  is  recognized  by  its  tree 
habit,  trifoliolate  leaves,  solitary  flowers  with  spa- 
thaceous calyx,  and  cylindric  fruits.  This  species 
was  at  first  thought  to  be  a  variant  of  P.  macro- 
phylla,  but  the  much  smaller  flowers  and  much 
narrower  fruits  indicate  that  the  plants  placed  here 
are  distinct.  We  have  not  seen  Costa  Rican  ma- 
terial and  base  this  report  on  the  original  descrip- 
tion and  the  Manual  Flora  manuscript  of  J.  F.  Mo- 
rales and  Q.  Jimenez  (1997). 

Parmentiera  macrophylla  Standl.,  Publ.  Field 
Columb.  Mus.  Hot.  Ser.  4:  263.  1929.  Figure 
11. 

Small  or  medium-size  trees  3-12(-25)  m  tall, 
trunk  unbranched  for  much  of  its  length,  to  30  cm 
diam.,  leafy  stems  1.8-4  mm  diam.,  glabrous,  be- 
coming pale  gray  and  terete,  petioles  often  sub- 
tended by  smooth  thickened  shelf-like  tissue. 
Leaves  opposite  or  subopposite,  3-foliolate  (rare- 
ly 2-4-foliolate),  petioles  2-10  cm  long,  0.8-1.5 
mm  diam.,  glabrous,  sulcate  above,  lateral  petio- 
lules 2-9  mm  long  and  merging  with  base  of 
blade,  terminal  petiolules  8-32  mm  long;  leaflet 
blades  (2-)6-17  cm  long,  (l-)2-6  cm  wide  (lat- 
eral blades  smaller  than  the  terminal),  elliptic  to 
elliptic-obovate  or  elliptic-rhombic,  apex  acumi- 
nate or  caudate-acuminate,  base  narrowly  cuneate 
and  decurrent  on  the  petiole,  drying  thin-charta- 
ceous,  subglabrous  above,  with  minute  sessile  pel- 
tate trichomes  beneath  and  tufts  of  hairs  (or 
webbed  tissue)  in  the  vein  axils  (domatia),  2° 
veins  4-7/side.  Inflorescences  cauliflorous,  1  or 
2  flowers  borne  on  trunk  and  larger  branches,  ped- 
icels 1-2  cm  long,  glabrous  or  with  minute  (0.05 
mm)  hairs,  drying  dark.  Flowers  with  tubular  ca- 
lyx 28-48  mm  long,  8-13  mm  diam.,  spathe-like 
(split  abaxially),  green  or  purplish,  glabrous;  co- 
rolla 50-75  mm  long,  trumpet-shaped  to  tubular 
campanulate,  white  or  the  tube  greenish  white, 
glabrous  externally,  with  a  saccate  bulge  in  the 
floor  of  the  throat,  tube  10-14  mm  diam.,  in  the 
proximal  half,  mouth  to  24  mm  wide;  filaments 
subequal,  32-38  mm  long,  thecae  5-6  mm  long, 
1-2  mm  wide.  Fruits  20-47  cm  long,  3-6  cm 
diam.  in  life,  narrowly  cylindric  with  longitudinal 
ribs  (8-sided  in  cross-section),  yellowish  and 
fleshy  (shrinking  on  drying);  seeds  ca.  6-9  mm 


142 


FIELDIANA:  BOTANY 


long,  5-6  mm  wide,  flat,  obovoid  with  rounded 
emarginate  (obcordate)  apex,  brown. 

Trees  of  the  shaded  Caribbean  rain  forest  inte- 
rior, 10-750  m  elevation.  Flowering  irregularly 
throughout  the  year  (more  often  in  the  wet  sea- 
son). This  species  ranges  from  southeastern  Nic- 
aragua along  the  Caribbean  slope  to  central  Pan- 
ama. 

Parmentiera  macrophylla  is  recognized  by  its 
tree  habit,  usually  opposite  trifoliolate  leaves,  cau- 
liflorous  flowers,  spathe-like  calyx,  thick  white 
corolla,  and  narrow  fleshy  fruits.  This  species  may 
flower  when  only  3  m  tall  with  a  stem  only  2.5 
cm  diam.  (Gentry,  1973b).  It  is  similar  to  P.  trun- 
ciflora  Standl.  &  L.  O.  Williams,  but  the  latter 
species  has  flowers  borne  on  prominent  (1-2  cm) 
woody  short-shoots  and  grows  at  higher  eleva- 
tions in  the  mountains  west  of  Jinotega,  Nicara- 
gua. 

Parmentiera  valerii  Standl.,  J.  Wash.  Acad.  Sci. 
17:  16.  1927.  Figure  11. 

Small  to  medium-size  trees  5-20  m  tall,  trunk 
to  1  m  diam.,  leafy  stems  1.2-5  mm  diam.,  gla- 
brous, pale  grayish  with  conspicuous  lenticels. 
Leaves  opposite  or  subopposite,  usually  5-folio- 
late,  less  often  3-7-foliolate,  petioles  18-70  mm 
long,  0.8-1.5  mm  diam.,  glabrous,  petiolules  0- 
1 2  mm  long  and  intergrading  into  the  base  of  the 
blade,  glabrous;  leaflet  blades  (1.2-)2-9  cm  long, 
(6-)8-34  mm  wide,  elliptic  to  narrowly  elliptic  or 
elliptic-obovate,  apex  acute  to  short-acuminate, 
base  acute  to  cuneate  and  decurrent  on  the  petiole, 
drying  thin  chartaceous,  glabrous  above  or  with 
the  midvein  minutely  puberulent,  glabrous  be- 
neath but  with  crooked  hairs  in  depressions  in  the 
vein  axils  (domatia),  2°  veins  3-6/side.  Inflores- 
cences of  few  fasciculate  cauliflorous  flowers  on 
lower  part  of  the  trunk,  pedicels  1 5-25  mm  long, 
glabrous,  drying  dark.  Flowers  with  tubular  calyx 
32-40  mm  long,  12-17  mm  diam..  glabrous, 
spathe-like;  corolla  60-75  mm  long,  tubular-cam- 
panulate,  white  or  greenish  white,  glabrous  exter- 
nally. Fruits  12-35  cm  long,  2-4  cm  diam.  in  life 
(1.2-3.5  cm  dried),  glabrous  (purplish  to  green- 
violet  in  life);  seeds  5-7  mm  long,  7-8  mm  wide. 


thin-lenticular,  obovate  with  obtuse  base  and  cor- 
date apex. 

Plants  of  the  moist  evergreen  lower  montane 
forest  understory,  600-1500  m  elevation.  Flow- 
ering throughout  the  year.  This  species  has  been 
collected  only  on  the  Cordillera  de  Guanacaste 
and  near  TilarSn  in  northwestern  Costa  Rica. 

Pannentiera  valerii  is  recognized  by  its  tree 
habit,  opposite  usually  pentafoliolate  leaves,  few 
cauliflorous  flowers,  spathe-like  calyx,  thick  white 
corrolla,  and  pendulous  cylindric  fleshy  fruits. 
The  limited  geographical  area  and  narrow  altitu- 
dinal  range  are  also  noteworthy.  Jicaro  danto  and 
pepino  de  danta  are  common  names. 


Phryganocydia  Martius  ex  Bureau 

Lianas  and  herbaceous  vines,  climbing  with 
simple  (distally  unbranched)  tendrils,  stems  most- 
ly terete,  cross-section  with  4  or  8  phloem  areas, 
interpetiolar  gland  fields  absent;  pseudostipules 
absent  or  early  caducous.  Leaves  opposite,  simple 
or  2-foliolate,  petiolate,  blades  glabrous  or  with 
few  minute  peltate  scales,  venation  pinnate.  Inflo- 
rescences terminal  or  axillary,  panicles  with  few 
flowers  on  bifurcating  axes,  or  of  1  or  2  flowers, 
subglabrous  with  few  peltate  scales.  Flowers  with 
tubular-funnelform  calyx  split  more  than  60%  and 
spathe-like,  tip  often  thickened  and  reflexed;  co- 
rolla tubular-funnel  form,  lavender  to  magenta, 
subglabrous  with  few  minute  peltate  scales  exter- 
nally, 2-lipped  and  5-lobed;  stamens  4,  filaments 
of  2  lengths,  anthers  glabrous,  thecae  straight,  di- 
varicate, staminode  present;  disc  absent;  ovary  2- 
locular.  ovules  in  2  series  in  each  locule.  Fruits 
linear-oblong  flattened  capsules,  valves  parallel  to 
the  septum,  surface  smooth  but  with  a  dense  cov- 
ering of  minute  lepidote  trichomes;  seeds  2- 
winged  with  thin  lateral  wings  or  corky-ovoid 
without  wings. 

Phryganocydia  is  unusual  because  of  its  spa- 
thaceous  calyx  with  thickend  tip  and  the  fruit  sur- 
face with  a  dense  covering  of  minute  hairs.  Our 
species  are  mostly  glabrous  or  with  minute  (0.05- 
0.1  mm)  sessile  peltate  scales. 


Key  to  the  Species  of  Phryganocydia 

la.  Fruits  narrowly  oblong  to  linear,  <  28  mm  wide,  to  60  cm  long,  surface  grayish;  seeds  thin  with 
lateral  membranaceous  wings;  leaves  consistently  2-foliolate;  plants  of  well-drained  sites    . 

P.  corymbosa 


BURGER:  FLORA  COSTARICENSIS 


143 


Ib.  Fruits  broadly  ovate,  >  35  mm  wide,  to  8  cm  long,  surface  yellowish  to  dark  brown;  seeds  thick 

and  corky,  lacking  thin  wings;  leaves  simple  or  2-foliolate;  plants  of  mangroves  and  lowland  swamps 

P.  phellosperma 


Phryganocydia  corymbosa  (Vent.)  Bureau  ex  K. 
Schum.  in  Engler  &  Prantl,  Naturl.  Pflanzen- 
fam.  4(3b):  224,  fig.  89H.  1894.  Spathodea  cor- 
ymbosa Vent.,  Choix,  tab.  40.  1807;  Mem. 
Math.  Phys.  Inst.  Natl.  France  1:  1-20.  1807. 
5.  laurifolia  Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3: 
114.  1819.  5.  orinocensis  Kunth  in  H.B.K., 
Nov.  Gen.  Sp.  3:  147.  1819.  Figure  22. 

Lianas  or  herbaceous  vines,  1-25  m  high,  to  5 
cm  diam.,  tendrils  7-16  cm  long,  leafy  stems  2- 
6  mm  diam.,  sparsely  to  densely  covered  with  flat 
peltate  scales  0.05-0.1  mm  diam.,  terete,  with  8 
phloem  areas  in  cross-section,  interpetiolar  ridge 
usually  present.  Leaves  2-foliolate  (simple  on 
young  stems),  petioles  8-33(-70)  mm  long,  1.2- 
2  mm  diam.,  vesture  similar  to  the  stem,  petiolules 
6-40  mm  long;  leaf  blades  5-15(-22)  cm  long, 
3-9(-ll)  cm  wide,  ovate-elliptic  to  ovate-oblong 
or  oblong-rounded,  apex  acute  to  short-acuminate 
or  rounded,  base  obtuse  to  rounded,  stiffly  char- 
taceous  to  subcoriaceous,  glabrous  on  both  sur- 
faces, 2°  veins  3-7/side.  Inflorescences  terminal 
or  axillary,  3-15  cm  long,  flowers  usually  cy- 
mose,  peduncles  to  5  cm  long,  pedicels  2-3  mm 
long  (node  to  calyx  base)  and  subtended  by  an 
internode  10-30  mm  long,  drying  black.  Flowers 
with  calyx  26-37(-48)  mm  long,  3-14  mm  diam., 
gradually  expanding  in  width,  with  a  straight  or 
curved  terminal  projection  3-9  mm  long,  split 
down  1  side  with  entire  margins,  surface  with  few 
peltate  scales;  corolla  55-85(-97)  mm  long,  tu- 
bular-funnelform,  rose-lilac  to  purplish  (often 
with  a  magenta  area  beneath  the  upper  2  lobes), 
subglabrous  externally,  9-24  mm  wide  at  the 
mouth,  throat  white  with  purple  lines,  lobes  1-3 
cm  long;  filaments  18-21  and  10-13  mm  long, 
thecae  3.5-4.5  mm  long.  Fruits  13-53  cm  long, 
16-26  mm  wide,  narrowly  oblong  to  linear-ob- 
long, flat  with  smooth  surface,  grayish  with  dense 
covering  of  minute  peltate  scales,  midvein  slightly 
elevated;  seeds  13-21  mm  long,  27-73  mm  wide, 
pale  brown  throughout,  wings  membranaceous. 

Plants  of  lowland  evergreen  or  partly  deciduous 
forest  formations,  10-400  m  elevation.  Flowering 
in  May-December  in  the  Golfo  Dulce  area  (not 
known  elsewhere  in  Costa  Rica);  flowering 
throughout  the  year  in  Panama.  The  species  is 
common  in  central  and  eastern  Panama  but  has 


only  been  collected  on  the  Burica  Peninsula  in 
western  Panama.  The  species  ranges  from  south- 
western Costa  Rica  to  Brazil. 

Phryganocydia  corymbosa  is  recognized  by  its 
climbing  habit  with  simple  tendrils,  opposite  bi- 
foliolate  leaves,  few-flowered  dichotomously 
branched  inflorescences,  spathe-like  calyx  with 
terminal  appendage,  large  pinkish  corollas,  and 
elongate  flattened  fruits  differing  greatly  in  length 
and  with  grayish  crystalline  surface.  The  leaflet 
blades  are  sometimes  nearly  as  broad  as  long.  An 
interesting  feature  of  this  species  is  that  some 
plants  flower  while  only  1  m  high  in  open  road- 
side vegetation.  Vegetatively,  this  species  is  very 
similar  to  Cydista  aequinoctialis,  but  the  fruit  of 
that  species  lacks  the  surface  trichomes  found  in 
P.  corymbosa. 

Phryganocydia  phellosperma  (Hemsl.)  Sandw., 
Kew  Bull.  1940:  302.  1941.  Macfadyena  phel- 
losperma Hemsl.,  Biol.  Central  Am.  Bot.  2: 
492.  1892. 

Lianas  to  2.5  cm  diam.,  tendrils  8-15  cm  long, 
stem  with  4  phloem  areas  in  cross-section,  leafy 
stems  2-4  mm  diam.,  terete,  reddish  brown  and 
with  scattered  minute  peltate  scales,  lacking  con- 
spicuous lenticels;  pseudostipules  absent.  Leaves 
2-foliolate  or  simple,  petioles  8-42  mm  long,  1- 
2.4  mm  diam.,  subglabrous,  petiolules  8-32  mm 
long,  sulcate  above;  leaf  blades  5-13  cm  long,  3— 
8  cm  wide,  ovate  to  ovate-triangular,  ovate-oblong 
or  oblong-elliptic,  apex  short-  to  long-acuminate, 
base  obtuse  to  rounded  and  often  truncate,  drying 
chartaceous  and  grayish,  glabrous  above,  glabrous 
or  with  few  minute  (0.05  mm)  peltate  scales  be- 
neath, 2°  veins  4-8/side.  Inflorescences  terminal, 
usually  of  2  (1)  flowers,  peduncles  ca.  10  mm 
long,  2°  peduncles  to  4  cm  long,  subglabrous,  ped- 
icel 2-3  mm  long,  often  terminated  by  a  pair  of 
linear-elliptic  bracts  6-8  mm  long.  Flowers  with 
calyx  18-27  mm  long,  4-10  mm  diam.,  terminat- 
ed by  a  thick  tip  2-3  mm  long,  split  along  side, 
subglabrous;  corolla  44-65  mm  long,  tubular- 
funnelform,  lavender  with  a  white  throat,  glabrous 
or  with  few  minute  hairs  externally,  8-20  mm 
wide  at  the  mouth,  lobes  12-25  mm  long;  fila- 
ments 13-15  and  8-1 1  mm  long.  Fruits  4-8  cm 
long,  35-67  mm  wide,  ca.  6  mm  thick,  broadly 


144 


FIELDIANA:  BOTANY 


ovate  to  ovate-suborbicular,  surface  dark  brown 
and  minutely  papillate  puberulent  (smooth  to  the 
touch);  seeds  20-27  mm  long,  22-35  mm  wide, 
thick,  corky,  and  wingless. 

Rarely  collected  plants  of  mangrove  swamps 
and  wet  muddy  sites  along  the  Pacific  coast,  0- 
20  m  elevation.  Flowering  mostly  in  the  wet  sea- 
son. The  species  ranges  from  southwestern  Costa 
Rica  to  Colombia. 

Phryganocydia  phellosperma  is  recognized  by 
its  climbing  habit  with  simple  tendrils,  opposite 
bifoliolate  or  simple  leaves,  glabrous  leaflets 
rounded  at  the  base,  usually  two-flowered  termi- 
nal inflorescences,  spathe-like  calyx,  pink  or  ma- 
genta corollas,  flattened  broadly  ovate  fruits,  and 
corky  seeds  lacking  thin  lateral  wings.  The  seeds 
of  this  species  are  probably  dispersed  by  water, 
consistent  with  its  restriction  to  mangroves  and 
low  wet  sites. 


Pithecoctenium  Martius  ex  Meisner 

Lianas,  tendrils  trifid  with  0-5  further  distal 
divisions  (sometimes  with  adhesive  pads),  stems 
strongly  6-angled,  with  4  phloem  areas  in  cross- 
section,  interpetiolar  lines  often  present,  gland 
fields  absent;  pseudostipules  narrowly  spatulate  or 
lanceolate,  caducous.  Leaves  opposite,  2-  or  3- 
foliolate  (occasionally  simple),  petiolate,  margin 
entire,  venation  palmate  to  pinnate.  Inflorescen- 
ces terminal  (or  terminal  on  axillary  short-shoots), 
racemes  or  racemose  panicles,  bracteate,  flowers 
pedicellate.  Flowers  with  cupulate  calyx,  distally 
truncated  or  minutely  5-toothed,  densely  puberu- 
lent, without  glands;  corolla  tubular-campanulate, 
relatively  thick,  puberulent  externally,  2-lipped,  5- 
lobed;  stamens  4,  filaments  of  2  lengths,  anthers 
glabrous,  thecae  straight,  divaricate,  a  small  stam- 
inode  present;  ovary  2-locular,  ovules  multiseriate 
in  each  locule.  Fruits  thick  woody  capsules, 
valves  parallel  with  the  septum  and  somewhat 
compressed,  covered  with  short  woody  spines; 
seeds  thin,  with  membranaceous  wing  around  3 
sides,  central  body  clearly  differentiated  from  the 
wings. 

Pithecoctenium  is  a  Neotropical  genus  of  four 
species,  one  wide-ranging  and  the  others  found  in 
Argentina  and  eastern  Brazil. 

Pithecoctenium  crucigerum  (L.)  A.  Gentry,  Tax- 
on  24:  121.  1975.  Bignonia  crucigera  L.,  Sp. 
PI.  2:  624.  1753.  B.  echinata  Jacq.,  Enum.  PI. 
Carib.  25.  1760.  B.  tiliaefolia  Kunth  in  H.B.K., 


Nov.   Gen.   Sp.   3:.  136.    1819.   P.   echinatnm 
(Jacq.)  Baill.,  Hist.  PI.  10:  8.  1888.  Figure  18. 

Lianas  to  ca.  10  cm  ilium.,  to  over  20  m  high, 
tendrils  5-20  cm  long,  with  3-15  slender  divi- 
sions (sometimes  terminating  in  a  disc  that  be- 
comes attached  to  bark),  leafy  stems  1.5-5  mm 
ilium.,  with  minute  peltate  scales  and  with  or 
without  slender  hairs  ca.  0.2  mm  long,  with  6 
prominent  longitudinal  ridges;  pseudostipules  8- 
15  mm  long,  1-2  mm  diam.  Leaves  opposite,  2- 
or  3-foliolate  (simple),  petioles  14-72  mm  long, 
1-2  mm  diam.,  subglabrous  or  with  thin  hairs  ca. 
0.2  mm  long,  petiolules  28-80  mm  long,  with  lon- 
gitudinal ridges;  leaf  blades  (4-)6-18  cm  long, 
(2.5-)4-13  cm  wide,  ovate  to  broadly  ovate  or 
ovate-oblong,  apex  acuminate,  base  rounded  and 
truncate  to  cordate,  drying  thin  chartaceous,  both 
surfaces  with  lustrous  peltate  scales  0.05-0. 1  mm 
diam.,  simple  hairs  0.2  mm  long  present  or  absent, 
venation  palmate  or  subpalmate,  2°  veins  4-67 
side,  strongly  ascending,  with  disc-like  glands  at 
the  base  beneath.  Inflorescences  3-19  cm  long, 
3-15  flowered  racemes  or  racemose  panicles,  pe- 
duncle 1.5-2.5  mm  thick,  resembling  the  stems, 
lateral  branches  7-12  mm  long,  bracts  1-6  mm 
long,  pedicels  1-2  mm  long.  Flowers  with  calyx 
7-11  mm  long,  8-10  mm  wide,  densely  puberu- 
lent and  lepidote,  margin  entire  or  minutely  5- 
dentate,  eglandular;  corolla  32-62  mm  long,  tu- 
bular-campanulate but  often  bent  at  almost  90° 
near  the  base,  white  or  yellowish  white,  densely 
puberulent  externally  with  minute  hairs  (drying 
yellowish),  lobes  6-17  mm  long;  filaments  17-21 
and  12-17  mm  long,  thecae  3-4  mm  long.  Fruits 
12-23(-31)  cm  long,  50-65(-75)  mm  wide,  23- 
38  mm  thick,  ellipsoid-oblong  with  somewhat 
flattened  sides,  surface  densely  tuberculate  with 
woody  projections  1-3  mm  high,  1.5-2.5  mm 
diam.  at  the  base,  yellowish  brown;  seeds  24- 
36(-41)  mm  long,  45-88(-95)  mm  wide,  central 
area  18-28  mm  wide,  wings  transparent. 

Common  lianas  of  deciduous,  partly  deciduous, 
and  wet  evergreen  forest  formations  on  both  Ca- 
ribbean and  Pacific  slopes,  1-1100  m  elevation. 
Flowering  primarily  in  May-June;  fruiting  mostly 
in  December-February.  This  species  ranges  from 
Mexico  to  Argentina. 

Pithecoctenium  crucigerum  is  recognized  by  its 
climbing  habit  with  multifid  tendrils,  opposite  bi- 
or  trifoliolate  leaves  (occasionally  simple),  race- 
mose inflorescences,  usually  entire  densely  pu- 
berulent calyx  cups,  densely  puberulent  white  co- 
rollas sharply  bent  near  the  middle,  echinate-tu- 


BURGER:  FLORA  COSTARICENSIS 


145 


berculate  woody  fruits,  and  thin  seeds  with  large 
thin  transparent  wings  on  three  sides.  The  hex- 
agonal twigs,  elongate  pseudostipules,  multi- 
branched  tendrils  (sometimes  terminating  in 
discs),  subpalmate  leaf  venation,  and  minute  shiny 
peltate  scales  on  leaves  are  useful  vegetative  at- 
tributes. Vegetatively,  these  plants  resemble  Am- 
philophium,  but  those  species  have  branched  hairs 
and  trifid  tendrils  and  lack  the  elongate  pseudos- 
tipules of  P.  crucigerum.  The  seeds  are  often 
called  mariposas  and  palomitas  in  Central  Amer- 
ica (Standley,  1938).  Batefta,  cucharilla,  peine  de 
mico,  and  peina  de  mono  are  common  names. 


Pleonotoma  Miers 

Lianas  and  vines,  climbing  with  tendrils  usu- 
ally 3-parted  at  the  apex,  stems  hollow  at  the  cen- 
ter, with  4  radiating  phloem  arms  in  cross-section 
and  quadrangular,  an  interpetiolar  line  usually  pre- 
sent at  the  node;  pseudostipules  leaf-like,  tubular, 
or  absent.  Leaves  opposite,  bi-  or  triternate  to  tri- 
pinnate,  the  terminal  petiolule  of  the  first  division 
often  replaced  by  a  tendril,  margins  entire,  vena- 
tion pinnate,  domatia  sometimes  present.  Inflo- 
rescences axillary  or  terminal  racemes,  bracts 
small,  lateral  bracteoles  sometimes  present,  pedi- 
cels well  developed.  Flowers  with  calyx  cupular, 
truncated  at  the  apex  and  subentire  or  with  5  mi- 
nute teeth;  corolla  tubular-funnelform,  white  to 
yellowish,  usually  glabrous  along  the  tube  and  pu- 
berulent  distally;  stamens  4,  of  2  lengths,  includ- 
ed, anthers  glabrous,  thecae  straight,  divaricate, 
pollen  grains  3-colpate,  a  staminode  present;  disc 
cupular-pulvinate;  ovary  oblong,  puberulent  or 
lepidote,  ovules  many  in  2  series  on  the  central 
placentae.  Fruits  capsule,  linear  to  narrowly  ob- 
long, valves  flat,  paralleling  the  septum,  smooth; 
seeds  thin  and  with  2  membranaceous  lateral 
wings. 

Pleonotoma  is  a  genus  of  14  species;  all  are 
South  American  except  for  one,  which  ranges 
northward  into  Central  America. 

Pleonotoma  variabilis  (Jacq.)  Miers,  Proc.  R. 
Hort.  Soc.  N.S.  3:  184.  1863.  Bignonia  varia- 
blilis  Jacq.,  Hort.  Schoenb.  2:  45,  tab.  212. 
1797.  Figure  16. 

Vines  or  lianas,  climbing  by  tendrils  (3-15  cm 
long)  with  3  short  distal  arms  5-20  mm  long,  larg- 
er stems  ca.  3  cm  diam.,  leafy  stems  2-7  mm 
diam.,  with  4  prominent  longitudinal  ridge,  subgl- 


abrous;  pseudostipules  2-5  mm  long.  Leaves  op- 
posite, bicompound  with  2-  or  3-trifoliate  parts, 
1°  petioles  1-7.5  cm  long,  1-2  mm  diam.,  gla- 
brous, 2°  petioles  2-6  cm  long,  longitudinally  stri- 
ate,  glabrous,  petiolules  7-40  mm  long  with  those 
of  central  leaflet  often  twice  as  long  as  laterals; 
leaflet  blades  3-18  cm  long,  1-9  cm  wide,  elliptic 
to  elliptic-ovate  or  elliptic-obovate,  apex  acute  to 
acuminate,  base  obtuse  to  rounded  (lateral  leaflets 
often  asymmetric  at  the  base),  drying  chartaceous, 
glabrous  except  for  short  (0.2-0.5  mm)  whitish 
hairs  along  the  midvein  and  vein  axils  beneath 
(domatia),  2°  veins  4-7/side.  Inflorescences  usu- 
ally terminal  racemes,  peduncle  and  rachis  2-8 
cm  long,  with  3-12  flowers,  pedicels  6-14  mm 
long,  0.6-1.4  mm  diam.,  drying  dark,  with  minute 
peltate  glands  or  subglabrous.  Flowers  with  cu- 
pular calyx  5-9  mm  long,  5-7  mm  diam.,  mostly 
glabrous,  margin  subentire  with  teeth  ca.  0.2  mm 
long  and  a  ciliolate  or  glabrous  edge;  corolla  50- 
90  mm  long,  tubular-funnelform,  white  or  yellow- 
ish white,  tube  10-20  mm  wide  at  the  mouth,  gla- 
brous along  the  tube  and  minutely  glandular  lep- 
idote distally,  lobes  14-22  mm  long,  rounded; 
longer  stamens  ca.  27  mm  long,  thecae  ca.  4  mm 
long,  divaricate;  ovary  ca.  4  X  1.5  mm.  Fruits 
18-35  cm  long,  1.5-2.7  cm  wide,  3-7  mm  thick, 
linear  to  linear-oblong,  surface  flat  and  slightly 
muricate,  drying  dark  and  lustrous;  seeds  1-1.5 
cm  long,  3-5  cm  wide,  central  part  ca.  12  mm 
wide,  wings  translucent  at  the  tips. 

Uncommon  evergreen  lianas  of  deciduous, 
partly  deciduous,  and  evergreen  forest  formations, 
10-800  m  elevation.  Known  from  the  Caribbean 
lowlands  of  Nicaragua  but  as  yet  collected  only 
on  the  Pacific  slope  in  Costa  Rica.  Flowering  in 
February-May;  fruiting  in  December-April.  The 
species  ranges  from  Guatemala  to  Trinidad  and 
the  Amazon  Basin. 

Pleonotoma  variabilis  is  recognized  by  its  vin- 
ing  habit  with  trifid  tendrils,  the  strongly  four-an- 
gled hollow  stems,  the  twice-compound  opposite 
leaves  (each  of  which  have  two  or  three  trifoliate 
parts),  short  racemes,  truncated  calyx  cup,  and 
white  or  yellowish  corolla.  The  tendrils  are  usu- 
ally borne  from  a  1°  petiole  that  also  bears  two 
trifoliolate  2°  petioles.  This  species  and  the  her- 
baceous Tourrettia  lappacea  are  the  only  Costa 
Rican  climbing  bignons  having  twice-compound 
leaves. 


Podranea  Sprague 

Scandent  shrubs  or  climbing  vines,  tendrils  ab- 
sent, stems  terete,  gland  fields  absent  at  the  nodes; 


146 


FIELDIANA:  BOTANY 


pseudostipules  not  developed.  Leaves  opposite, 
imparipinnate  with  opposite  lateral  leaflets,  blades 
entire  or  distally  serrate,  venation  pinnate.  Inflo- 
rescences terminal,  solitary,  paniculate  with  op- 
posite lateral  branches,  pedicels  subtended  by  ca- 
ducous bracts.  Flowers  large,  calyx  tubular  or 
campanulate  (slightly  inflated),  glabrous,  with  5 
prominent  lobes  shorter  than  the  tube;  corolla  tu- 
bular-funnelform  or  tubular-campanulate,  laven- 
der, slightly  2-lipped,  glabrous  externally,  the  5 
lobes  rounded;  stamens  4,  included,  filaments  of 
2  lengths,  anthers  glabrous,  thecae  straight,  par- 
allel and  later  divergent,  staminode  small;  disc 
saucer-shaped,  thick;  ovary  ovoid,  2-locular, 
ovules  ca.  6-seriate  in  each  locule,  stigma  of  2  flat 
ovate  lobes.  Fruits  elongated  linear  capsules, 
valves  leathery,  compressed  parallel  to  the  sep- 
tum, dehiscing  perendicular  to  the  septum;  seeds 
flat,  2-winged. 

Podranea  is  a  monotypic  African  genus  of  east- 
tropical  and  southern  Africa,  now  a  popular  gar- 
den ornamental  in  the  tropics.  The  name  Podra- 
nea is  an  anagram  of  Pandorea,  a  similar  and 
closely  related  genus  of  five  species  native  to  Ma- 
laysia, Australia,  and  the  western  Pacific  (with 
two  species  widely  cultivated  ornamental  vines). 
Pandorea  differs  in  having  pod-like  oblong  cap- 
sules with  woody  valves,  and  a  noninflated  calyx. 

Podranea  ricasoliana  (Tanfani)  Sprague  in  Thi- 
selton-Dyer,  Fl.  Cap.  4  (2):  450.  1904.  Tecoma 
ricasoliana  Tanfani,  Bull.  Soc.  Tosc.  Ortic. 
1887:  17.  1887.  Pandorea  ricasoliana  (Tanfani) 
Baill.,  Hist.  PI.  10:  40.  1888.  Figure  15. 

Scandent  shrubs  and  vines,  tendrils  absent, 
leafy  stems  1-5  mm  diam.,  glabrous,  becoming 
pale  gray  in  age  with  rounded  lenticels,  interpe- 
tiolar  lines  usually  present.  Leaves  6-25  cm  long, 
with  usually  5-13  leaflets,  petioles  16-60  mm 
long,  0.6-1.5  mm  diam.,  glabrous,  petiolules  2- 
12  mm  long;  leaflet  blades  (12-)20-55  mm  long, 
8-25  mm  wide,  lanceolate  to  ovate-lanceolate  or 
ovate-elliptic,  apex  acuminate,  margin  with  3-9 
teeth/side,  base  somewhat  asymmetric  (rounded 
and  cuneate)  on  lateral  leaflets,  glabrous,  2°  veins 
3-5/side.  Inflorescences  terminal,  6-18  cm  long, 
paniculate  with  short  opposite  lateral  branches, 
glabrous,  bracts  2-4  mm  long,  subulate,  pedicels 
4-14  mm  long.  Flowers  with  calyx  14-20  mm 
long,  6-9  mm  diam.,  whitish  to  pale  lavender  in 
life,  with  few  round  glands  on  the  distal  surface, 
with  5  apiculate  lobes  3-6  mm  long;  corolla  5-8 
cm  long,  tubular-campanulate,  pale  lavender  to 


pink  with  purple  areas  near  the  2  adaxial  lobes 
(white  inside),  glabrous  externally  but  with  dense 
crooked  hairs  in  the  sinuses  of  the  lobes;  filaments 
18-20  and  14-16  mm  long,  thecae  3-4  mm  long. 
Fruits  25-30  cm  long,  linear  and  terete  (rarely 
produced  in  cultivation). 

Podranea  ricasoliana,  native  of  southern  Afri- 
ca, is  now  widely  cultivated  as  a  garden  orna- 
mental. It  is  recognized  by  its  clambering  habit 
(without  tendrils),  opposite  pinnately  compound 
leaves,  narrow  leaflets  with  serrate  margins, 
slightly  inflated  calyx  with  prominent  lobes,  and 
large  pink  corollas,  often  with  tufts  of  hairs  in  the 
sinuses  of  the  lobes.  It  flowers  throughout  the  year 
and  does  well  from  near  sea  level  to  2000  m  el- 
evation when  grown  on  a  wall  or  trellis  in  open 
sunny  locations.  Called  bombilla,  linda,  and  m(r- 
ame  linda  in  Guatemala. 


Pyrostegia  Presl 

Vines  or  lianas,  climbing  with  the  aid  of  coiling 
petiole-borne  tendrils  usually  trifid  near  the  tip, 
stems  with  longitudinal  ribs,  lacking  interpetiolar 
gland  fields;  pseudostipules  small  or  inconspicu- 
ous. Leaves  opposite  or  subopposite,  petiolate,  2- 
or  3-foliolate,  petiolules  present,  margins  entire, 
vention  pinnate.  Inflorescences  terminal  or  axil- 
lary panicles,  often  much  branched,  pedicels  sub- 
tended by  small  paired  bracts.  Flowers  with  cu- 
pular  calyx,  minutely  glandular  lepidote  or  subgl- 
abrous,  distal  margin  truncated  with  5  small  teeth; 
corolla  tubular  or  narrowly  funnelform,  orange  or 
reddish  orange,  with  5  lobes,  valvate  in  bud;  sta- 
mens 4,  exserted,  anthers  glabrous,  thecae 
straight,  parallel  or  slightly  divergent;  disc  annu- 
lar; ovary  narrowly  tubular,  stigma  bifid.  Fruits 
dry  capsules,  valves  parallel  to  septum,  smooth 
and  leathery,  with  an  indistinct  median  vein;  seeds 
thin,  2-winged  and  transverse-oblong,  wings 
smooth  and  with  hyaline  margins. 

A  genus  of  four  species,  originally  native  to 
tropical  South  America.  One  species  is  now  wide- 
ly planted  for  its  showy  orange  flowers. 

Pyrostegia  venusta  (Ker-Gawler)  Miers,  Proc.  R. 
Hort.  Soc.  N.S.  3:  188.  1863.  Bignonia  venusta 
Ker-Gawler,  Bot.  Reg.  3:  5.  1818.  B.  ignea 
Veil.,  Fl.  Flumin.  244.  1825.  P.  ignea  (Veil.)  K. 
B.  Presl.,  Bot.  Bemerk.  93:  1843.  Figure  15. 

Vines,  climbing  with  tendrils  to  15  cm  long, 
leafy  stems  2-5  mm  diam.,  glabrous  or  with  short 


BURGER:  FLORA  COSTARICENSIS 


147 


(0.2-0.5  mm)  hairs  near  the  nodes,  with  6  or  8 
longitudinal  ribs.  Leaves  3-foliolate  or  2-foliolate 
and  often  with  a  tendril,  petioles  7-35  mm  long, 
0.8-1.8  mm  diam.,  with  short  hairs  along  the  ad- 
axial  side,  petiolules  4-18  mm  long  (distal  leaflet 
with  longer  petiolules);  leaflet  blades  3-9  cm 
long,  1.5-6  cm  wide,  ovate-elliptic  to  ovate-ob- 
long, apex  acute  to  caudate-acuminate,  base  ob- 
tuse to  slightly  rounded  and  truncate  or  subcor- 
date,  drying  chartaceous,  glabrous,  minutely 
punctate  beneath,  2°  veins  3-5/side.  Inflorescenc- 
es dense  panicles  usually  terminal  on  short  shoots, 
often  pendulous  with  8-25  flowers,  rachis  to  9  cm 
long,  borne  on  woody  peduncles  5-50  mm  long, 
1-2.5  mm  diam.,  glabrous  or  puberulent  along  1 
side,  pedicels  12-20  mm  long,  glabrous.  Flowers 
with  calyx  4-5  mm  long,  4-6  mm  wide,  puber- 
ulent and  glandular  lepidote  or  subglabrous,  distal 
margin  minutely  ciliolate,  subentire  or  with  mi- 
nute (0.3  mm)  lobes  (teeth);  corolla  6-9  cm  long, 
2-3  mm  diam.  near  the  base,  8-12  mm  wide  at 
the  mouth,  orange,  tubular  to  narrowly  tubular- 
funnelform,  straight  or  slightly  curved,  lobes  6- 
18  mm  long,  oblong  with  blunt  apex,  becoming 
reflexed,  whitish  puberulent  along  the  margin;  sta- 
mens slightly  exserted,  filaments  3-5  cm  long, 
thecae  ca.  4  mm  long.  Fruits  20-30  cm  long,  10- 
16  mm  wide;  seeds  1.2-1.6  mm  long,  3-4  cm 
wide. 

Pyrostegia  venusta  is  recognized  by  its  climb- 
ing habit  with  distally  trifid  tendrils,  opposite  2- 
or  3-foliolate  leaves,  long  narrowly  tubular  orange 
corollas,  and  exserted  stamens.  These  plants  are 
often  seen  climbing  over  garden  walls,  with  their 
clusters  of  brilliant  orange  flowers  making  a  fine 
display.  The  corollas  are  usually  slightly  curved, 
held  horizontally  when  in  anthesis,  and  drooping 
afterward.  Native  to  Brazil,  Bolivia,  and  Para- 
guay, this  species  is  now  planted  as  an  ornamental 
throughout  the  tropics.  In  Central  America  it  is 
grown  from  near  sea  level  to  1500  m  elevation. 
This  species  has  been  called  chorro,  chorro  de 
oro,  San  Carlos,  and  Triquitraque  in  Central 
America. 


Saritea  Dugand 

Lianas  climbing  with  simple  tendrils,  young 
stems  terete  and  without  interpetiolar  gland  fields; 
pseudostipules  conspicuous  and  leaf  like  (ca.  5- 
25  mm  long).  Leaves  opposite,  short-petiolate,  2- 
foliolate,  leaf  blades  5-12  cm  long,  3-7  cm  wide, 
cuneate  at  the  base,  subglabrous,  basal  2°  veins 


strongly  ascending,  usually  with  a  gland  field  in 
the  axil  of  proximal  veins  beneath.  Inflorescences 
usually  terminal  with  1-9  flowers,  peduncles  and 
pedicels  drying  dark,  subglabrous  with  few  mi- 
nute lepidote  hairs,  flowers  sometimes  in  distal 
triads.  Flowers  with  tubular  subglabrous  calyx  6- 
10  mm  long,  margin  truncate;  corolla  6-9  cm 
long,  tubular-campanulate,  magenta,  glabrous  ex- 
ternally; stamens  4,  filaments  ca.  2.8  and  2.1  mm 
long,  anthers  glabrous  and  straight,  staminode 
small;  disc  small;  ovules  2-seriate  (or  appearing 
1 -seriate)  in  each  locule.  Fruits  linear  capsules, 
valves  flattened  parallel  to  the  septum;  seeds  thin, 
with  2  membranous  lateral  wings. 

Saritea  magnifica  (Sprague  ex  van  Steenis)  Du- 
gand, native  to  Colombia  and  Ecuador,  is  the  only 
species  of  this  genus  and  is  widely  planted  as  an 
ornamental.  This  taxon  is  closely  related  to  Cy- 
dista  and  Phyrganocydia  and  also  to  Arrabidaea 
(Gentry,  1973b).  Although  not  recorded  from 
Costa  Rica,  this  species  may  be  expected  in  gar- 
dens. The  bifoliolate  leaves  with  cuneate-based 
leaflets,  gland  fields  in  vein  axils  beneath,  con- 
spicuous pseudostipules,  and  large  magenta  co- 
rollas make  it  easy  to  identify. 

Schlegelia  is  now  placed  in  the  family  Schle- 
geliaceae  (q.v.). 

Scobinaria  is  now  considered  to  be  part  of  Ar- 
rabidaea. See  Gentry's  discussion  (Selbyana  2: 
43-45.  1977). 


Spathodea  Beauvois 

Trees  with  well-developed  trunks,  nodes  lack- 
ing interpetiolar  lines  or  glands;  pseudostipules 
absent.  Leaves  opposite  or  3/node,  petiolate,  im- 
paripinnately  compound,  lateral  leaflets  opposite, 
venation  pinnate.  Inflorescences  terminal  and 
erect,  compact  racemes  and  somewhat  corymb- 
like  with  the  lower  pedicels  longer  than  the  distal. 
Flowers  large,  calyx  spathe-like  and  curved  with 
a  narrowed  apex,  opening  from  beneath  (abaxi- 
ally);  corolla  with  a  narrow  cylindric  base  and 
large  curved-campanulate  throat,  deep  red  to  or- 
ange-red, glabrous  on  the  exterior,  5-lobed;  sta- 
mens 4,  anthers  with  slender  divaricate  thecae; 
ovary  2-locular,  narrowly  oblong,  ovules  multis- 
eriate  in  each  locule,  stigma  flat  and  bifid.  Fruits 
narrowly  oblong-ellipsoid  capsules,  dehiscence 
perpendicular  to  the  septum  along  1  side;  seeds 


148 


FIELDIANA:  BOTANY 


flat  with  broad  thin  hyaline  wing  around  the  entire 
circumference. 

The  genus  Spathodea,  member  of  the  tribe  Te- 
comeae,  contains  a  single  species  native  to  tropi- 
cal Africa.  A  densely  puberulent  variant  of  East 
Africa  is  sometimes  segregated  as  5.  nilotica 
Seem.  The  compact  terminal  inflorescences  with 
large  red  corollas  and  spathaceous  calyx  are  dis- 
tinctive. 

Spathodea  campanulata  Beauvois,  Fl.  Oware  1: 
47,  tab.  27.  1805.  Figure  15. 

Trees  to  25  m  tall,  trunks  to  40  cm  diam.,  bark 
somewhat  rough  and  scaling  near  the  base,  leafy 
stems  3-12  mm  diam.,  minutely  appressed  puber- 
ulent, lenticellate.  Leaves  19-40  cm  long,  with 
7-13(-17)  leaflets,  petioles  to  20  cm  long,  petio- 
lules  1-3  mm  long  (terminal  leaflet  with  petiolule 
to  15  mm  long),  ca.  1  mm  diam.,  puberulent;  leaf 
blades  5-14  cm  long,  2-6  cm  wide,  elliptic  to 
ovate-elliptic  or  ovate-oblong,  apex  acute  to  acu- 
minate, base  acute  to  obtuse  or  rounded  on  1  side 
and  asymmetric,  puberulent  on  the  veins  beneath 
to  densely  puberulent  throughout,  with  a  gland  at 
the  base  of  the  midvein  abaxially,  2°  veins  5- 107 
side.  Inflorescences  compact  terminal  corymb- 
like  racemes  with  a  closely  placed  spiral  of  6-35 
flowers,  ca.  7  X  15  cm  (not  including  corollas), 
pedicels  to  6  cm  long,  distal  pedicels  shorter  than 
the  proximal  and  giving  a  flat-topped  effect. 
Flowers  with  calyx  2-6  cm  long,  split  on  1  side, 
recurved  and  acuminate,  densely  sericeous,  yel- 
lowish brown;  corolla  6-15  cm  long,  5-9  cm 
wide  at  the  apex,  widely  campanulate  and  curved 
from  a  short-tubular  base,  deep  red  to  orange-red, 
lobes  2-3  cm  long,  obtuse,  distal  edge  of  the  co- 
rolla crinkled  and  yellowish.  Fruits  17-27  cm 
long,  3-7  cm  wide,  1-2  cm  thick,  narrowly  ellip- 
soid-oblong; seeds  ca.  15  X  20  mm,  with  thin 
membranaceous  wing. 

Spathodea  campanulata  is  recognized  by  its 
tree  habit,  opposite  pinnately  compound  leaves, 
compact  terminal  inflorescences,  curved  spatha- 
ceous calyx,  and  very  large  curved-campanulate 
red  corollas.  This  is  a  very  striking  ornamental 
tree  when  in  full  flower,  with  the  huge  corollas 
often  in  a  whorl-like  configuration.  The  distal  un- 
opened flower  buds  often  form  a  tight  cluster  on 
which  birds  can  perch  while  taking  nectar  from 
the  upwardly  open  corollas.  Native  to  evergreen 


forests  of  tropical  central  Africa,  the  species  is 
now  cultivated  throughout  the  tropics.  In  Costa 
Rica  it  has  been  grown  successfully  in  evergreen 
and  partly  deciduous  areas  below  1500  m  eleva- 
tion. African  tulip  tree,  llama  del  bosque,  and  tn- 
lipdn  are  common  names. 


Sti/(>ph\  limn  Miers 

Lianas  or  vines,  climbing  with  simple  or  trifid 
tendrils,  stems  with  4  phloem  areas  in  cros's-sec- 
tion,  terete,  longitudinally  striate,  hollow  in  the 
center,  nodes  without  gland  fields;  pseudostipules 
spatulate  or  inconspicuous,  caducous.  Leaves  op- 
posite, 2-  or  3-foliolate,  petiolate,  blades  with 
conspicuous  pellucid-lustrous  round  sessile  glands 
on  the  lower  surface,  venation  palmate  or  subpal- 
mate.  Inflorescences  axillary  to  distal  (sometimes 
early  developing)  leaves,  few-flowered  racemes, 
peduncles  short,  bracts  well  developed,  pedicels 
puberulent.  Flowers  with  cupulate  or  campanulate 
calyx,  often  slightly  inflated,  5-lobed  or  irregular- 
ly split  (2-lipped),  densely  puberulent.  thin-tex- 
tured; corolla  campanulate-funnelform,  slightly 
2-lipped,  white  or  yellowish  white,  puberulent  and 
with  minute  glandular  hairs  externally,  the  lobes 
often  tinged  with  pink,  lobes  rounded;  stamens  4. 
filaments  of  2  lengths,  anthers  glabrous  or  puber- 
ulent along  the  lines  of  dehiscence,  thecae 
straight,  divaricate,  staminode  present;  disc  thick; 
ovary  linear-tetragonal,  2-locular,  ovules  2-seriate 
in  each  locule.  Fruits  linear  capsules,  valves  flat- 
tened parallel  to  the  septum,  slightly  convex,  de- 
hiscence septicidal,  surface  puberulent;  seeds 
transversely  oblong,  thin  and  flat,  with  2  translu- 
cent lateral  wings  weakly  differentiated  from  the 
central  area. 

A  Neotropical  genus  of  three  species,  one  of 
which  ranges  from  Mexico  to  Amazonia.  The  ge- 
nus is  unusual  because  of  the  pellucid  dots  on  the 
lower  leaf  surfaces,  the  hollow  stems,  and  the 
very  slender  fruits,  which  may  resemble  stems. 
Central  American  material  of  our  two  species  is 
difficult  to  separate  on  the  basis  of  herbarium  ma- 
terial. Both  species  exhibit  a  wide  range  of  vari- 
ability as  regards  density  of  pubescence,  size  of 
leaves,  and  size  and  shape  of  calyx  and  corolla. 
Study  in  the  field  is  the  basis  for  determing  their 
status  as  separate  species  (Gentry,  1 973b,  p.  936). 


BURGER:  FLORA  COSTARICENSIS 


149 


Key  to  the  Species  of  Stizophyllum 

la.  Branchlets  and  petioles  yellowish  brown  to  reddish  brown  or  dark  brown  with  hairs  0.5-2  mm  long; 

tendrils  with  3-branched  tips;  calyx  10-18  mm  long;  fruit  surface  with  longer  (to  1  mm)  hairs  .  .  . 

5.  inaequilaterum 

Ib.  Branchlets  and  petioles  grayish  or  pale  brownish  with  short  (0.1-0.5  mm)  hairs;  tendrils  simple 

with  unbranched  tip  (rarely  2-branched);  calyx  7-12  mm  long,  chartaceous;  fruit  surface  with  minute 

(0. 1-0.3  mm)  hairs    S.  riparium 


Stizophyllum  inaequilaterum  Bureau  &  K. 
Schum.  in  Mart.,  Fl.  Bras.  8(2):  221.  1896.  Big- 
nonia  inaequilatera  Poeppig  ex  Bureau  &  K. 
Schum.  in  Mart.,  Fl.  Bras.  8(2):  222,  1896,  pro 
syn.,  non  Poeppig  ex  Bureau,  Adansonia  8:  289. 
1868.  Figure  26. 

Lianas  and  vines,  to  3  cm  diam.,  tendrils  8-18 
cm,  with  3  short  (6-20  mm  long)  distal  parts, 
leafy  stems  2-7  mm  diam.,  densely  villous  with 
yellowish  brown  hairs  0.5-2  mm  long,  older 
stems  glabrescent;  pseudostipules  to  8  mm  long. 
Leaves  2-  or  3-foliolate,  petioles  22-68(-150) 
mm  long,  densely  villous,  petiolules  9-28(-60) 
mm  long;  leaflet  blades  7-13(-19)  cm  long,  4- 
7(-14)  cm  wide,  ovate  to  oblong  or  elliptic-ob- 
long, apex  acute  to  acuminate,  margin  entire  or 
denticulate,  base  of  lateral  leaflets  asymmetric 
with  a  rounded  and  a  cuneate  side,  drying  char- 
taceous, upper  surface  with  slender  curved  hairs 
0.5-1  mm  long,  lower  surface  more  densely  vil- 
lous, venation  pinnate  or  subpalmate,  2°  veins  3- 
6/side.  Inflorescences  a  few-flowered  terminal 
fascicle  or  contracted  raceme  on  a  short  axillary 
shoot,  peduncles  ca.  1  cm  long,  bracts  conspicu- 
ous, linear,  pedicels  8-24  mm  long,  densely  vil- 
lous with  reddish  brown  hairs.  Flowers  with  calyx 
10-18  mm  long,  7-10  mm  diam.,  inflated-cam- 
panulate,  thin-chartaceous,  villous  with  hairs  to 
1 .5  mm  long,  margin  irregular  or  with  lobes  to  4 
mm  long;  corolla  5-7  cm  long,  tubular-funnel- 
form,  greenish  yellow  with  pink  lobes  (purple), 
tube  puberulent  externally,  lobes  8-1 1  mm  long; 
1 1 km ic ntx  20-21  and  15-16  mm  long,  thecae  2 
mm  long.  Fruits  24-35  cm  long,  4-6  mm  wide, 
3-4  mm  thick,  linear,  densely  brownish  villous; 
seeds  not  seen. 

Rarely  collected  plants  of  lowland  evergreen 
rain  forest  formations  on  the  Caribbean  slope,  0- 
500  m  elevation.  Flowering  in  March-May.  The 
species  ranges  from  eastern  Nicaragua  to  Peru. 

Stizophyllum  inaequilaterum  is  recognized  by 
its  climbing  habit  with  trifid  tendrils,  dense  pu- 
bescence of  reddish  brown  hairs,  opposite  bi-  or 


trifoliolate  leaves,  short  few-flowered  inflores- 
cences, thin  villous  calyx  with  irregular  margins, 
puberulent  corollas  with  pink  lobes,  and  slender 
long-linear  pubescent  fruits.  The  dense  pubes- 
cence of  longer  (0.5-2  mm)  hairs  and  distally  tri- 
fid tendrils  (rarely  well  preserved  in  herbarium 
material)  help  distinguish  this  species  from  its 
more  common  congener. 

Stizophyllum  riparium  (Kunth  in  II  BK 
Sandw.,  Lilloa  3:  462.  1938.  Bignonia  riparia 
Kunth  in  H.B.K.,  Nov.  Gen.  Sp.  3:  138.  1819. 
Adenocalymma  flos-ardeae  Pittier,  Contrib. 
U.S.  Natl.  Herb.  18:  256.  1917.  A.  punctifolium 
Blake,  Contrib.  U.S.  Natl.  Herb.  24:  22.  1922. 
5.  flos-ardeae  (Pittier)  Sandw.,  Recueil  Trav. 
Bot.  Neerl.  34:  212.  1937.  5.  punctifolium 
(Blake)  Sandw.,  Recueil  Trav.  Bot.  Need.  34: 
212.  1937.  Figure  23. 

Lianas  and  vines  to  5  cm  diam.,  tendrils  7-15 
cm  long,  apex  simple  or  rarely  with  reduced  sec- 
ond tip,  leafy  stems  1.3-6  mm  diam.,  sparsely  to 
densely  puberulent  with  hairs  to  0.7  mm  long,  te- 
rete; pseudostipules  3-8  mm  long,  strap-shaped. 
Leaves  2-  or  3-foliolate,  petioles  12-110  mm 
long,  ca.  1  mm  diam.,  puberulent,  lateral  petio- 
lules 7-37  mm  long  (terminal  to  70  mm);  leaflet 
blades  3.5-1 7(-20)  cm  long,  2-9.5(-12)  cm  wide, 
ovate-elliptic,  ovate,  or  elliptic-oblong  to  narrow- 
ly ovate-oblong,  apex  acute  to  short-acuminate, 
margin  entire  or  with  1-7  obtuse  teeth/side  in  ear- 
ly stages,  base  rounded  and  truncate  to  subcor- 
date,  drying  thinly  chartaceous,  sparsely  puberu- 
lent with  slender  hairs  0.2-0.6  mm  long  above, 
more  densely  puberulent  beneath,  with  pellucid 
dots  0.1-0.2  mm  diam.  beneath,  venation  subpal- 
mate, 2°  veins  4-6/side.  Inflorescences  terminal 
fascicles  or  axillary  few-flowered  racemes,  pe- 
duncles 1-5  cm  long,  puberulent  like  the  stems, 
pedicels  4-7  mm  long.  Flowers  with  calyx  7-12 
mm  long,  4-9  mm  diam.,  deeply  cupular  or  tu- 
bular-funnelform,  densely  minutely  puberulent, 
margin  5-lobed  or  irregular,  glandular  on  lobes; 


150 


FIELDIANA:  BOTANY 


corolla  32-53  mm  long,  tubular-funnelform  to  tu- 
bular-campanulate,  white  to  pale  yellowish  white, 
minutely  papillate  puberulent,  lobes  5-8  mm 
long;  filaments  15-18  and  1 1-12  mm  long,  thecae 
2-2.5  mm  long.  Fruits  24-45  cm  long,  3-8  mm 
wide,  3.5-5  mm  thick,  linear  and  narrowed  at  the 
ends,  straight  or  more  often  curved,  lenticular  in 
cross-section,  surface  densely  minutely  puberu- 
lent, grayish;  seeds  4-6  mm  long,  14-26  mm 
wide,  transversely  oblong,  wings  translucent. 

Climbers  in  lowland  evergreen  rain  forest  for- 
mations, 0-300  m  elevation.  Flowering  mostly  in 
June-November  (but  rarely  seen  with  flowers  and 
rare  in  herbaria).  This  species  ranges  from  eastern 
Mexico  to  the  Amazon  basin. 

Stizophyllum  riparium  is  recognized  by  its 
climbing  habit  with  simple  tendrils,  hollow  pu- 
berulent stems,  opposite  bi-  or  trifoliolate  leaves, 
blades  with  pellucid  dots  beneath,  white  or  yel- 
lowish corollas  puberulent  externally,  and  the 
very  narrow  linear  usually  curved  fruits.  Leaflets 
on  juvenile  growth  may  have  conspicuous  broad- 
based  teeth.  The  differences  between  this  and  the 
preceding  species  are  often  very  difficult  to  dis- 
cern in  herbarium  material. 


Tabebuia  Gomes  ex  De  Candolle 

REFERENCES — A.  Gentry,  A  revision  of  Tabe- 
buia (Bignoniaceae)  in  Central  America.  Brittonia 
22:  246-264.  1970.  A.  Gentry,  Bignoniaceae— 
Part  II  (tribe  Tecomeae).  Flora  Neotropica,  Mon- 
ogr.  25(11).  1992. 

Small  to  large  trees  or  shrubs,  wood  usually 
dense  and  fine-grained,  branchlets  terete,  nodes 
lacking  interpetiolar  gland  fields;  pseudostipules 
absent.  Leaves  opposite,  palmately  3-7(9-)-fo- 
liolate  or  less  often  simple  or  1  -foliolate,  petiolate, 


basal  leaflets  usually  smaller  and  with  shorter  pet- 
iolules  than  the  more  terminal,  venation  pinnate. 
Inflorescences  terminal,  short,  open  or  congested 
panicles,  few-flowered  racemes  or  fascicles  of  1 
to  few  flowers,  bracts  subtending  the  pedicels; 
some  species  flowering  when  the  tree  is  leafless. 
Flowers  with  tubular  to  cupular  or  campanulate 
calyx,  margin  2-5-lobed,  irregular  or  truncate, 
surface  with  flat  rounded  appressed,  stellate  or 
straight  hairs;  corolla  tubular-funnelform  to  tu- 
bular-campanulate,  white,  yellow,  rose,  purple,  or 
red,  glabrous  or  puberulent  externally,  lobes  5. 
rounded;  stamens  4,  exserted  or  included,  fila- 
ments of  2  lengths,  anthers  glabrous,  thecae 
straight,  divaricate,  staminode  present;  disc  cush- 
ion-shaped, ovary  oblong,  2-locular,  ovules  in  2 
to  many  series  in  each  locule.  Fruits  capsules, 
linear-cylindric  to  oblong-cylindric,  subterete, 
valves  dehiscing  approximately  perpendicular  to 
the  septum,  surface  smooth  to  verrucose-muricate, 
glabrous  to  variously  puberulent;  seeds  thin, 
transversely  oblong  with  2  thin  lateral  wings 
(thicker,  corky,  and  without  wings  in  T.  palustris 
et  al.). 

Tabebuia,  a  member  of  tribe  Tecomeae,  is  a 
Neotropical  genus  of  100  species,  with  many  spe- 
cies in  South  America  and  the  West  Indies.  In 
Central  America  most  of  the  species  can  be  dis- 
tinguished by  the  tree  habit,  opposite  usually  pal- 
mately compound  leaves,  compact  but  showy  ter- 
minal inflorescences  (often  in  flower  when  the 
leaves  are  absent),  large  corollas,  and  the  linear 
to  oblong  fruits  that  are  rounded  in  cross-section 
and  split  at  right  angles  to  the  septum.  In  our  spe- 
cies, the  corolla  may  be  white,  yellow,  or  rose  to 
purple.  When  in  full  flower,  these  are  some  of  the 
most  beautiful  trees  in  Central  America.  The 
wood  of  this  genus  is  a  commercially  important 
tropical  hardwood,  and  the  bark  contains  phar- 
macologically active  compounds  (Gentry,  1992). 


Key  to  the  Species  of  Tabebuia 

la.  Seeds  corky,  without  thin  transparent  lateral  wings;  fruits  to  1 1  cm  long;  corollas  white  with  yel- 
lowish markings  within;  shrubs  and  small  trees  of  mangroves  and  river  deltas  along  the  Pacific 
coast;  leaves  simple  or  3-foliolate,  leaflets  mostly  narrowly  lanceolate T.  palustris 

Ib.  Seeds  thin  with  transparent  lateral  wings;  fruits  usually  >  15  cm  long;  corolla  yellow  to  rose  or 

purple  (white  with  pink  lobes  in  T.  rosea);  medium  to  large  trees,  not  found  in  mangrove  formations 

(occasionally  in  lowland  riversides);  leaves  usually  3-,  5-,  or  7-foliolate,  leaflets  rarely  lanceolate 

2 

2a.  Corollas  rose,  purple,  or  white,  marked  with  pink;  leaflets  and  stems  without  stellate  hairs;  fruits 
glabrous  or  with  flat  rounded  appressed  peltate  (lepidote)  hairs  or  minutely  papillate,  the  surfaces 
smooth  .  


BURGER:  FLORA  COSTARICENSIS 


151 


2b.  Corollas  bright  yellow;  leaflets  with  stellate  hairs  (sometimes  only  in  vein  axils  beneath  and  difficult 
to  see);  fruits  stellate-pubescent  to  minutely  villous  or  with  an  undulate  surface  if  with  minute 

rounded  appressed  (lepidote)  hairs    4 

3a.  Surface  of  leaflets  with  minute  flat  rounded  (lepidote)  hairs;  corolla  glabrous  externally;  calyx 
with  minute  flat  rounded  hairs;  capsule  with  minute  papillate  or  appressed  peltate  hairs;  native 
in  dry  deciduous  to  wet  evergreen  rain  forest  formations  and  planted  as  ornamental,  0-1500  m 

elevation   T.  rosea 

3b.  Surface  of  leaflets  with  straight  hairs,  at  least  in  the  vein  axils  beneath;  corolla  puberulent 
externally;  calyx  with  minute  straight  or  scurfy  hairs;  capsule  mostly  glabrous;  in  deciduous 

and  partly  deciduous  forest,  5-300  m  elevation    T.  impetiginosa 

4a.  Leaflets  with  minute  stellate  hairs  only  in  the  vein  axils  beneath,  leaves  often  7-foliolate;  fruits 
with  slightly  undulate  surface  and  appressed  rounded  flat  (lepidote)  hairs  [trees  of  lowland  rain 

forests]    T.  quayacan 

4b.  Leaflets  with  stellate  hairs  scattered  on  the  surface  beneath  or  along  the  veins,  leaves  rarely  7- 

foliolate;  fruit  with  few  to  many  minute  stellate  or  tomentulous  hairs 5 

5a.  Leaves  puberulent  mostly  along  veins  and  petioles;  calyx  without  long  simple  hairs,  lobes 
becoming  reflexed;  corolla  lobes  with  dark  interconnecting  veins  to  the  distal  margin;  fruits 

minutely  stellate-puberulent;  trees  of  evergreen  rain  forest  formations  (in  Costa  Rica) 

T.  chrysantha 

5b.  Leaves  densely  puberulent  over  the  entire  surface  beneath;  calyx  yellowsh  pubescent  with  thin 
straight  hairs  2-7  mm  long,  lobes  erect;  corolla  lobes  with  venation  not  so  clearly  reaching  the 

margin;  fruits  tomentulous;  trees  of  deciduous  and  partly  deciduous  forest  formations 

.    T.  ochracea 


Tabebuia  chrysantha  (Jacq.)  G.  Nichols.,  111. 
Diet.  Gard.  4:  1.  1887.  Bignonia  chrysantha 
Jacq.,  Hort.  Schoenb.  2:  45,  tab.  211.  1797.  Te- 
coma  chrysantha  (Jacq.)  DC.,  Prodr.  9:  221. 
1845.  Tec.  evenia  J.  D.  Smith,  Bot.  Gaz.  20:  8. 
1895,  pro  parte  (fls.).  Tec.  palmeri  Kranzl.,  Fed- 
de  Repert.  17:  220.  1921.  Tab.  chrysantha  ssp. 
pluvicola  A.  Gentry,  Phytologia  35:  190.  1977. 
Figure  13. 

Trees  to  30  m  tall,  trunks  to  1  m  diam.,  bark 
light  gray  and  smooth  with  few  widely  spaced 
long  vertical  furrows,  leafy  stems  3-9  mm  diam., 
terete,  densely  puberulent  with  stellate  hairs  0.1- 
0.3  mm  long,  glabrescent,  becoming  pale  gray. 
Leaves  usually  5-foliolate  (3-,  4-,  7-foliolate), 
petioles  (5-)8-19(-28)  cm  long,  1.5-3.5  mm 
diam.,  densely  stellate  puberulent,  petiolules  1-9 
cm  long  (basal  laterals  8-20  mm),  with  node-like 
area  below  the  blade;  leaflet  blades  (4-)7-21(-26) 
cm  long,  (2-)3-12  cm  wide  (proximal  lateral  leaf- 
lets smaller  than  the  distal),  elliptic  to  broadly  el- 
liptic or  elliptic-obovate,  apex  acuminate  to  cau- 
date-acuminate, margin  entire  or  sometimes  blunt- 
ly serrate,  base  obtuse  to  rounded  and  slightly 
truncate,  drying  thinly  to  stiffly  chartaceous, 
subglabrous  to  sparsely  stellate  puberulent  above, 
more  densely  stellate-puberulent  beneath,  peltate 
trichomes  0.02-0.05  mm  diam.  often  conspicuous 


beneath,  2°  veins  6-12/side.  Inflorescences  ter- 
minal, condensed  (almost  fasciculate)  panicles 
with  7-25  flowers,  peduncles  4-20  mm  long, 
densely  stellate  pubescent,  pedicels  4-20  mm 
long,  1.2-2  mm  diam.  Flowers  with  calyx  8-14 
(-19)  mm  long,  7-10(-13)  mm  diam.,  cupular- 
funnelform  or  campanulate,  densely  pubescent 
with  hairs  0.3-0.8  mm  long,  lobes  2-3  mm  long, 
broadly  triangular;  corolla  52-95  mm  long,  tu- 
bular-funnelform,  bright  yellow  with  narrow  red- 
dish lines  within,  glabrous  externally,  lobes  9-28 
mm  long;  filaments  16-22  and  10-15  mm  long, 
thecae  2-3  mm  long.  Fruits  20-50(-90)  cm  long, 
14-24  mm  wide,  linear-cylindric,  narrowed  at  the 
ends,  surfaces  smooth  and  minutely  stellate  pu- 
berulent; seeds  6-12  mm  long,  14-38  mm  wide, 
wings  transparent,  well  differentiated. 

Trees  of  wet  evergreen  lowland  rain  forest  for- 
mations on  both  the  Caribbean  and  Pacific  slopes 
in  Costa  Rica  and  Panama,  20-600  m  elevation 
(also  occurring  in  much  drier  habitats  in  its  range 
from  Mexico  to  Nicaragua  and  in  Venezuela). 
Flowering  primarily  in  January-March,  occasion- 
ally at  other  times.  The  species  ranges  from  Mex- 
ico, disjunctly,  to  Peru. 

Tabebuia  chrysantha  is  recognized  by  its  large 
tree  habit,  small  stellate  hairs  on  many  parts,  op- 
posite palmately  5-foliolate  leaves,  compact  ter- 
minal inflorescences,  large  brilliant  yellow  exter- 


152 


FIELDIANA:  BOTANY 


nally  glabrous  corollas,  and  long  linear-cylindric 
fruits  with  transparent-winged  seeds.  The  leaflets 
are  sometimes  serrate.  The  trees  make  a  brilliant 
show  when  flowering,  in  part  because  they  usually 
flower  when  all  the  leaves  have  fallen.  Common 
names  are  corteza  and  cones  amarilla. 

Our  collections  are  placed  in  subspecies  pluvi- 
cola  A.  Gentry,  which  ranges  from  northeastern 
Costa  Rica  to  northern  Venezuela  and  Ecuador. 
Subspecies  chrysantha  is  found  from  Mexico  to 
Nicaragua  and  disjunctly  in  Amazonian  Peru.  The 
two  subspecies  can  be  separated  by  the  following 
key. 


la.  Trees  usually  10-20  m  tall,  usually  found  in 
seasonally  dry  deciduous  or  partly  deciduous 
forests  below  1 200  m  elevation;  calyx  <  1 0  mm 
long,  densely  rufescent  with  stellate  to  curved- 
barbate  hairs  ca.  1  mm  long;  fruits  15-50  X 
0.8-2  cm,  persistently  stellate-tomentose,  often 
striate  or  with  rough  surface.  .  .  .  subsp.  chry- 
santha 

Ib.  Trees  to  30  m  tall,  usually  found  in  lowland 
and  lower  montane  evergreen  forest  formations, 
0-1500  m;  calyx  >  10  mm  long,  rufescent  with 
short  stellate  hairs  ca.  0.5  mm  long;  fruits  30- 
80  X  1 .5-2.4  cm,  nearly  glabrescent  and  smooth. 
subsp.  pluvicola 


Tabebuia  guayacan  (Seem.)  Hemsley,  Biol. 
Centr.  Am.  Bot.  2:  495.  1882.  Tecoma  quaya- 
can  Seem.,  Bot.  Voy.  Herald,  180.  1854. 

Trees  to  50  m  tall,  trunks  to  1.7  m  diam.,  bark 
gray  or  brown  and  ridged  with  well-developed 
vertical  furrows,  leafy  stems  2-8  mm  diam., 
subglabrous  or  with  few  minute  hairs,  becoming 
pale  grayish,  terete.  Leaves  5-  or  7-foliolate,  pet- 
ioles, 8-23  cm  long,  1.5-3  mm  diam.,  sparsely 
puberulent  with  scurfy  hairs  or  glabrous,  petio- 
lules  1-7  cm  long  (basal  lateral  petiolules  1-3  cm 
long),  geniculate  and  drying  dark  below  the  blade; 
leaflet  blades  5-20(-30)  cm  long,  3-1 1(- 15)  cm 
wide  (terminal  larger  than  laterals),  ovate-elliptic 
to  elliptic  or  lanceolate,  apex  acute  to  acuminate, 
margin  entire  (serrate  in  juveniles),  base  rounded 
to  obtuse  (lateral  leaflets  usually  asymmetric), 
drying  thin-chartaceous,  subglabrous  or  minutely 
puberulent  on  the  veins  beneath  and  with  short 
stellate  hairs  in  vein  axils  (domatia),  lower  surface 
with  minute  appressed  peltate  hairs,  2°  veins  6-97 
side.  Inflorescences  compact  panicles  with  flow- 


ers in  diads  or  triads  and  opening  at  the  same 
time,  peduncles  6-14  mm  long,  pedicels  10-18 
mm  long,  0.6-0.9  mm  diam.,  sparsely  puberulent 
with  scurfy  hairs  0.1-0.3  mm  long.  Flowers  with 
calyx  9-15  mm  long,  4-9  mm  diam.,  campanu- 
late,  with  minute  scurfy  scattered  hairs,  margin 
split  or  with  2-5  rounded/obtuse  lobes;  corolla 
63-105  mm  long,  tubular- funnelform,  yellow  with 
brown  lines  within,  glabrous  externally,  13-22 
mm  wide  at  the  mouth,  lobes  22-35  mm  long; 
filaments  15-21  and  11-16  mm  long,  thecae  2-3 
mm  long.  Fruits  30-60  cm  long,  12-28  mm 
wide,  linear,  surface  subglabrous  to  thick  stellate- 
puberulent;  seeds  6-1 1  mm  long,  23-40  mm 
wide,  wings  transparent  and  clearly  differentiated. 

Rarely  collected  trees  of  lowland  evergreen  rain 
forest  formations,  5-600  m  elevation.  Flowering 
in  February-March  (March-May  elsewhere  in 
Central  America);  seeds  are  released  in  May-July. 
The  species  ranges  from  Mexico  to  Peruvian 
Amazonia. 

Tabebuia  quayacan  is  recognized  by  its  tree 
habit,  opposite  palmately  penta-  or  septafoliolate 
subglabrous  leaves,  usually  narrow  leaflets,  com- 
pact terminal  inflorescences,  large  yellow  corol- 
las, and  long  linear  cylindric  fruits.  The  leaves  are 
often  almost  glabrous,  except  for  the  minute 
round  flat  peltate  trichomes  and  the  stellate  hairs 
in  vein  axils  beneath.  The  hard  wood  has  been 
used  as  uprights  in  home  construction,  as  axles, 
and  for  tool  handles  (Englesing  181,  Nicaragua). 
This  species  is  very  closely  related  to  T.  serrati- 
folia  (Vahl)  Nichols.,  a  wide-ranging  species  of 
South  America  (Gentry,  1992).  Common  names 
are  cartes  or  cortez. 

Tabebuia  impetiginosa  (Mart,  ex  DC.)  Standl., 
Publ.  Field  Mus.  Nat.  Hist.,  Bot.  Ser.  11:  176. 
1936.  Tecoma  impetiginosa  Mart,  ex  DC., 
Prodr.  9:  218.  1845.  Tab.  palmeri  Rose,  Con- 
trib.  U.S.  Natl.  Herb.  1:  109.  tab.  1 1.  1891.  Tab. 
nicaraguensis  Blake,  Contrib.  Gray  Herb.  52: 
95.  1917.  Tab.  dugandii  Standl.,  Trop.  Woods 
36:  17.  1933.  Figure  13. 

Trees  to  30  m  tall,  trunks  to  70  cm  diam.,  bark 
slightly  furrowed,  leafy  stems  3-8  mm  diam.,  gla- 
brous or  sparsely  puberulent  with  minute  (0.05 
mm)  hairs,  terete,  longitudinally  striate,  becoming 
smooth  and  grayish.  Leaves  3-  or  5-foliolate  (in 
Central  America),  petioles  4-14  cm  long,  1.3-2.5 
mm  diam.,  sparsely  and  minutely  puberulent,  lon- 
gitudinally striate,  petiolules  7-50  mm  long  (basal 
laterals  to  15  mm);  leaflet  blades  (3-)  11 -19  cm 


BURGER:  FLORA  COSTARICENSIS 


153 


long,  (2-)4-8  cm  wide  (basal  leaflets  often  much 
smaller  than  distal),  elliptic  to  elliptic-oblong  or 
narrowly  ovate-elliptic,  apex  short-  to  long-acu- 
minate, margin  serrate  in  juvenile  plants,  base 
acute  to  obtuse  or  rounded,  drying  chartaceous, 
glabrous  above,  lower  surface  subglabrous  or  with 
short  (0.4  mm)  thin  hairs  along  the  midvein,  and 
with  tufts  of  hairs  in  the  vein  axils  (domatia),  2° 
veins  9-12/side.  Inflorescences  terminal,  com- 
pact-paniculate (congested-fasciculate),  peduncles 
2-12  mm  long,  1-1.7  mm  diam.,  densely  yellow- 
ish pubescent  with  thick  stellate  hairs,  pedicels  2- 
6  mm  long.  Flowers  with  calyx  5-8  mm  long,  3- 
6  mm  diam.,  tubular-campanulate,  sparsely  to 
densely  yellowish  stellate  puberulent,  margin 
truncate  or  slightly  5-lobed;  corolla  35-75  mm 
long,  tubular-funnelform,  purple  to  red-purple 
(yellow  within),  sparsely  to  densely  minutely  pu- 
berulent externally,  15-25  mm  wide  at  the  mouth, 
lobes  10-17  mm  long;  filaments  16-23  and  10- 
16  mm  long,  thecae  2.5-3.5  mm  long.  Fruits  15- 
56  cm  long,  12-25  mm  wide,  10-13  mm  thick, 
surface  subglabrous  and  drying  dark,  with  few 
longitudinal  ridges;  seeds  9-13  mm  long,  30- 
48(-80)  mm  wide,  central  area  17-18  mm  wide, 
wings  translucent  and  clearly  differentiated. 

Deciduous  trees  of  seasonally  very  dry  decid- 
uous or  partly  deciduous  (rarely  in  lowland  ev- 
ergreen) forest  formations,  10-300  m  elevation 
(to  1400  m  elsewhere).  Flowering  in  November- 
February;  fruiting  in  January-February.  This  spe- 
cies ranges  from  Mexico  to  Argentina. 

Tabebuia  impetiginosa  is  recognized  by  its  tree 
habit,  opposite  palmately  tri-  or  pentafoliolate 
leaves,  compact  terminal  inflorescences,  external- 
ly puberulent  purple  corollas,  long  linear  fruits, 
and  seeds  with  transparent  wings.  The  leaflets  are 
mostly  glabrous.  Gentry  (1973b,  1992)  discussed 
the  regional  differentiation  and  nomenclatural  as- 
pects of  this  species.  Common  names  are  cortez 
negro  and  roble  macho.  Compare  T.  rosea 

Tabebuia  ochracea  (Cham.)  Standl.,  Field  Mus. 
Nat.  Hist.  Bot.  Ser.  11:  176.  1936.  Tecoma 
ochracea  Cham.,  Linnaea  7:  653.  1832.  Tab. 
chrysantha  (Jacq.)  Nichols.,  Diet.  Gard.  4:1. 
1897,  sensu  Sandw.,  non  Jacq.  Tab.  neochry- 
santha  A.  Gentry,  Brittonia  22:  260.  1970.  Tab. 
ochracea  subsp.  neochyrsantha  (A.  Gentry)  A. 
Gentry,  Ann.  Missouri  Bot.  Gard.  60:  948. 
1973.  Figure  13. 

Trees  8-25  m  tall,  bark  with  flat-surfaced  gray 
ridges  alternating  with  darker  furrows,  leafy  stems 


3-7  mm  diam.,  densely  stellate  pubescent  with 
hairs  0.2-0.5  mm  long,  soon  glabrescent  and  te- 
rete. Leaves  5-foliolate,  petioles  4-18  cm  long, 
1.3-2.7  mm  diam.,  densely  stellate  pubescent, 
petiolules  5-58  mm  long,  lateral  basal  petiolules 
3-15  mm  long;  leaflet  blades  4-22  cm  long  (bas- 
al laterals  2-11  cm),  2-14  cm  wide,  oblong-ob- 
ovate  or  elliptic-obovate  to  broadly  elliptic,  apex 
acuminate,  margin  serrate  in  juvenile  plants,  base 
obtuse  to  rounded  and  truncate,  drying  charta- 
ceous, densely  stellate  puberulent  beneath  with 
hairs  ca.  0.5  mm  wide,  2°  veins  6-12/side.  Inflo- 
rescences terminal,  condensed  subcapitate  clus- 
ters of  2-  and  3-flowered  groups  or  contracted 
panicles,  distal  stems  often  terminating  in  a 
rounded  pubescent  inflorescence  "bud"  1-2  cm 
diam.,  peduncles  and  pedicels  not  usually  visible 
in  the  dense  pubescence  of  yellowish  (dried)  hairs 
to  1.5  mm  long.  Flowers  sweet-scented,  calyx  8- 
13  mm  long,  4-8  mm  diam.,  campanulate,  dense- 
ly yellowish  (dried)  tomentulous  with  straight 
hairs  to  2(-7)  mm  long  (stellate  at  base),  margin 
with  5  lobes  1-2  mm  long;  corolla  36-75(-83) 
mm  long,  tubular-funnelform,  yellow  with  red 
lines  within,  glabrous  externally  except  near  the 
lobes,  lobes  10-25  mm  long;  filaments  15-20  and 
9-15  mm  long,  thecae  1.5-2.5  mm  long.  Fruits 
1 1-38  cm  long,  10-17  mm  wide,  5-10  mm  thick, 
linear-cylindric,  surface  densely  tomentulous  with 
hairs  0.3-2  mm  long  (sometimes  subglabrous  and 
with  prominent  longitudinal  ridges);  seeds  6-12 
mm  long,  17-88  mm  wide,  wings  transparent, 
well  differentiated. 

Common  deciduous  trees  of  seasonally  dry  de- 
ciduous and  partly  deciduous  forest  formations  of 
the  Pacific  slope,  5-900  m  elevation.  Flowering 
in  January-June;  fruiting  in  February-June.  This 
species  is  represented  in  our  area  by  subspecies 
neochrysantha,  which  ranges  from  Guatemala  to 
northwestern  Venezuela;  the  other  subspecies  are 
South  American. 

Tabebuia  ochracea  subspecies  neochrysantha 
is  recognized  by  its  tree  habit,  opposite  palmately 
pentafoliolate  leaves,  compact  inflorescences  (ear- 
ly stages  form  round  pubescent  "buds"  at  the  tips 
of  distal  stems),  bright  yellow  corollas  mostly  gla- 
brous externally,  long  cylindric  fruits,  and  seeds 
with  transparent  wings.  The  leaflets  are  densely 
stellate  tomentulose  beneath.  This  species  is  sim- 
ilar to  T.  chrysantha  of  evergreen  forests;  both  are 
called  cones  amarilla  and  cortez. 

Tabebuia  palustris  Hemsley,  Biol.  Centr.  Am. 
Bot.  2:  495.  1882.  Figure  12. 


154 


FIELDIANA:  BOTANY 


Shrubs  or  small  trees  1 .5-4  m  tall,  trunks  to  5 
cm  diam.,  often  twisted  and  sometimes  forming 
dense  colonies,  leafy  stems  2-8  mm  diam.,  terete, 
surface  with  flat  rounded  trichomes.  Leaves  sim- 
ple or  3-foliolate  (2-foliolate),  petioles  24-100 
mm  long,  1.5-2.5  mm  diam.,  with  appressed  pel- 
tate hairs  but  often  appearing  glabrous,  often  with 
gland  fields  along  the  proximal  half,  petiolules  of 
distal  leaves  12-35  mm  long;  leaflet  blades  6-19 
cm  long,  1.4-5  cm  wide,  narrowly  lanceolate  to 
narrowly  elliptic  or  elliptic-oblong,  apex  acute, 
base  acute  to  cuneate,  drying  yellowish  and  sub- 
coriaceous,  glabrous  above,  densely  lepidote  with 
minute  (0. 1  mm)  appressed  rounded  trichomes  be- 
neath, 2°  veins  8-13/side.  Inflorescences  terminal 
on  short  shoots  or  terminal  in  a  distal  branch-di- 
chotomy, flowers  1-5,  often  in  diads  or  triads,  pe- 
duncle 8-25  mm  long,  bracts  2-8  mm  long,  linear, 
pedicels  9-11  mm  long,  with  minute  appressed 
rounded  hairs.  Flowers  with  calyx  11-17  mm 
long,  6-8  mm  diam.,  cupulate-funnelform,  drying 
dark,  surface  with  flat  rounded  hairs  ca.  0.1  mm 
diam.,  margin  cleft  into  2-4  lobes  1-5  mm  long; 
corolla  40-75  mm  long,  tubular-funnelform, 
white  with  yellow  lines  within,  glabrous  external- 
ly, 9-18  mm  wide  at  mouth,  lobes  8-23  mm  long; 
filaments  18-26  and  14-19  mm  long,  thecae  3 
mm  long.  Fruits  5-1 1  cm  long,  14-26  mm  diam., 
oblong-cylindric  with  a  narrow  tip,  surface 
smooth,  covered  by  minute  peltate  scales;  seeds 
14-18  mm  long,  18-22  mm  wide,  suborbicular 
with  corky  texture,  wings  absent. 

Shrubs  of  tidal  mangrove  formations  and  sea- 
side marshes  along  river  estuaries,  0-10  m  ele- 
vation. Flowering  throughout  the  year.  The  north- 
ernmost collection  (Croat  663  from  Playa  Coco, 
Guanacaste)  may  be  an  outlier.  The  estuary  of  the 
Rio  Tempisque  may  be  the  northernmost  area  in 
which  this  species  is  common;  it  is  common 
around  Golfo  Dulce.  This  species  ranges  along  the 
Pacific  Coast  from  Costa  Rica  to  northwestern  Ec- 
uador. 

Tabebuia  palusiris  is  recognized  by  its  Pacific 
coastal  habitats,  short  shrubby  (small  tree)  habit, 
opposite  usually  trifoliolate  leaves  with  narrow 
leaflets,  few  white  flowers,  and  cylindrical  fruit 
with  corky  wingless  seeds.  The  habitat,  gland 
fields  on  petioles,  narrow  leaf  blades,  and  corky 
seeds  easily  distinguish  this  species  from  all  its 
congeners  in  Costa  Rica. 

Tabebuia  rosea  (Bertol.)  DC..  Prodr.  9:  215. 
1845.  Tecoma  rosea  Bertol.,  Fl.  Guatimal.  25. 
1840.  Tec.  mexicana  Mart,  ex  DC.,  Prodr.  9: 


218.  1845.  Sparattosperma  rosea  (Bertol.) 
Miers,  Proc.  R.  Hort.  Soc.  3:  99.  1863.  Tub. 
mexicana  (Mart,  ex  DC.)  Hemsl.,  Biol.  Centr. 
Am.  Bot.  2:  495.  1882.  Tab.  pentaphylla  (L.) 
Hemsl.,  Biol.  Centr.  Am.  Bot.  2:  495.  1882,  non 
Bignonia  pentaphylla  L.  Couralia  rosea  (Ber- 
tol.) J.  D.  Smith,  Bot.  Gaz.  20:  9.  1895.  Tec. 
evenia  J.  D.  Smith,  Bot.  Gaz.  20:  8.  1895,  pro 
parte.  Figure  13. 

Trees  5-30  m  tall,  to  1  m  diam.,  bark  dark  gray 
and  rough  with  a  distinct  pattern  of  vertical  ridg- 
es, leafy  stems  3-12  mm  diam.,  with  minute  pel- 
tate scales,  glabrescent,  becoming  terete.  Leaves 
5-foliolate,  basal  leaflets  often  smaller  than  distal, 
petioles  6-17(-32)  cm  long,  2-3.4  mm  diam..  lep- 
idote, petiolules  of  distal  leaflets  2-7(-ll)  cm 
long;  leaflet  blades  5-18(-35)  cm  long,  3-10 
(-18)  cm  wide  (distal  leaflets  larger),  elliptic  to 
elliptic-oblong,  apex  acute  to  acuminate,  margin 
entire,  base  cuneate  or  rounded,  drying  stiffly 
chartaceous  and  grayish,  both  surfaces  with  flat 
appressed  rounded  hairs  0.05-0.1  mm  diam.  (lep- 
idote), gland  fields  sometimes  present  along  the 
midvein  near  the  base  beneath,  2°  veins  7-12/side. 
Inflorescences  2-20  cm  long,  short  or  condensed 
panicles  often  with  dichotomous  branching,  pe- 
duncles 8-25  mm  long,  lepidote,  bracts  1-2  mm 
long,  pedicels  4-20  mm  long,  with  minute  peltate 
scales.  Flowers  with  calyx  11-21  mm  long,  6-1 1 
mm  diam.,  cupular-funnelform,  surface  densely 
minutely  papillate  puberulent  or  lepidote,  margin 
with  2  or  3  rounded  or  obtuse  lobes  separated  by 
irregular  sinuses  1-3  mm  deep;  corolla  53-97 
mm  long,  tubular-funnelform,  pink  to  purple  or 
white  with  pinkish  lobes,  often  yellowish  within, 
glabrous  externally,  lobes  18-26  mm  long;  fila- 
ments 14-20  and  10-15  mm  long,  thecae  2.5-3.5 
mm  long.  Fruits  21-38  cm  long,  10-18  mm 
diam.,  linear-cylindric,  densely  covered  with  mi- 
nute peltate  or  papillate  hairs,  apex  narrowly  acu- 
minate; seeds  8-12  mm  long,  28-45  mm  wide, 
wings  transparent  and  sharply  differentiated. 

Trees  of  deciduous,  partly  deciduous,  and  low- 
land evergreen  rain  forest  formations  on  both  Ca- 
ribbean and  Pacific  coasts,  0-1100  m  elevation 
(planted  at  elevations  up  to  1500  m).  Flowering 
primarily  in  January-March;  fruits  usually  dehisc- 
ing in  April-May.  This  species  ranges  from  Mex- 
ico to  Venezuela  and  western  Ecuador. 

Tabebuia  rosea  is  recognized  by  its  tree  habit, 
opposite  palmately  pentafoliolate  leaves,  tubular 
and  irregularly  lobed  calyx,  large  pink  or  white 
and  pink  corollas,  linear-cylindric  fruits,  and  seeds 


BURGER:  FLORA  COSTARICENSIS 


155 


with  thin  transparent  wings.  The  leaf  surfaces  ap- 
pear glabrous  but  have  a  dense  covering  of  minute 
appressed  peltate  hairs;  some  leaves  have  gland 
fields  along  the  base  of  the  midvein  beneath.  Un- 
like T.  chrysantha,  the  leaflets  of  this  species  are 
always  entire.  This  species  is  one  of  the  most  im- 
portant commercial  sources  of  fine  quality  hard- 
wood; it  is  used  in  making  furniture  and  interior 
trim  and  for  other  purposes.  The  tree  flowers  with 
or  without  its  foliage  and  is  one  of  Central  Amer- 
ica's most  beautiful  native  species,  often  planted 
for  ornament.  Common  names  are  roble,  roble  de 
bianco,  and  roble  de  sabana. 


Tanaecium  Swartz 

Lianas  climbing  with  simple  tendrils,  stems  te- 
rete, with  8  or  16  phloem  areas  in  cross-section, 
interpetiolar  gland  fields  present  or  absent;  pseu- 
dostipules  small  or  absent.  Leaves  opposite,  2-  or 
3-foliolate,  petiolate,  blades  often  with  a  cyanide 
odor  when  crushed,  venation  pinnate.  Inflores- 
cences axillary  or  terminal,  panicles  or  racemose 
panicles,  or  solitary  in  distal  leaf  axils.  Flowers 
with  cupulate  calyx,  margin  entire  or  with  5  mi- 
nute teeth,  often  with  flat  rounded  glands  on  the 
distal  surface;  corolla  with  a  very  long  tube  and 
salverform  distally,  white,  externally  glabrous  or 
puberulent,  lobes  5;  stamens  4,  exserted  or  partly 
exserted,  filaments  of  2  lengths,  anthers  glabrous, 
thecae  straight  or  curved,  somewhat  divergent, 
staminode  present;  ovary  oblong,  lepidote,  2-loc- 
ular,  ovules  multiseriate  in  each  locule.  Fruits 
capsules,  oblong  to  ellipsoid-cylindric,  valves 
parallel  to  the  septum,  thick,  woody,  convex, 
smooth;  seeds  lacking  wings  or  with  2  lateral 
poorly  differentiated  membranaceous  wings. 

Tanaecium  is  a  genus  of  about  six  species  rang- 
ing from  Costa  Rica  and  the  West  Indies  to  Brazil. 

Tanaecium  jaroba  Sw.,  Prodr.  92.  1788.  T.  al- 
biflora  DC.,  Prodr.  9:  245.  1845.  Figure  16. 

Lianas  climbing  with  tendrils  5-22  cm  long, 
leafy  stems  2.5-7  mm  diam.,  glabrous,  terete, 
nodes  with  10-15  round  flat  glands  between  the 
leaf  bases;  pseudostipules  ca.  3  mm  long  or  ab- 
sent. Leaves  2-  or  3-foliolate,  petioles  4-9  cm 
long,  1.5-2.3  mm  diam.,  glabrous  or  puberulent, 
petiolules  7-37  mm  long  (shorter  on  lateral  leaf- 
lets); leaflet  blades  6.5-16  cm  long,  3-9  cm  wide, 
elliptic-oblong  to  elliptic  or  ovate-elliptic,  apex 
acute  to  short-acuminate,  base  obtuse  to  rounded 
and  somewhat  truncate,  glabrous  above,  glabrous 


or  with  curled  thin  hairs  to  0.6  mm  long  beneath, 
2°  veins  5-9/side.  Inflorescences  terminal  or  ax- 
illary, solitary  flowers  in  distal  leaf  axils  or  short 
few-flowered  racemes  on  leafless  stem-tips,  ped- 
icels 7-24  mm  long,  1-2  mm  diam.,  glabrous, 
drying  black.  Flowers  with  calyx  9-16  mm  long, 
5-10  mm  diam,  deeply  cupulate,  subglabrous, 
with  flat  circular  glands  distally,  margin  subentire 
with  minute  (0.3  mm)  teeth;  corolla  1 1-24  cm 
long,  long-tubular-salverform,  white,  minutely  pu- 
berulent externally,  tube  3.5-5  mm  diam.  for  most 
of  its  length,  ca.  10  mm  wide  at  the  throat,  lobes 
17-27  mm  long;  filaments  ca.  28  and  15  mm  long, 
thecae  5-6  mm  long.  Fruits  9-22  cm  long,  5—1 1 
cm  wide,  4-8  cm  thick,  ellipsoid-oblong  to  ob- 
long, pale  brown,  subglabrous;  seeds  25-33  mm 
long,  30-32  mm  wide,  woody,  angular  with  thin 
edge  on  3  sides. 

Rarely  collected  plants  of  evergreen  lowland 
rain  forest  formations  along  the  Caribbean  coastal 
plain.  Collected  in  flower  in  late  May  (Pittier 
3631)  and  early  October  (Gomez-Laurito  12913) 
in  Costa  Rica.  This  species  ranges  from  south- 
eastern Nicaragua  and  Jamaica  to  Venezuela  and 
Peru. 

Tanaecium  jaroba  is  recognized  by  its  climbing 
habit  with  simple  tendrils,  opposite  bi-  or  trifoli- 
olate  leaves,  few-flowered  inflorescences,  very 
long  slender  white  corollas,  and  rounded  woody 
fruits  with  woody  seeds.  No  other  species  of  Big- 
noniaceae  in  Central  America  has  such  long  and 
narrow  corolla  tubes.  The  flowers  have  a  sweet 
odor  (like  Hedychium,  Zingiberacae)  and  open  at 
night.  They  are  probably  pollinated  by  sphingid 
moths  with  very  long  tongues.  Little  Central 
American  material  of  this  species  has  been  seen; 
the  description  is  based  largely  on  South  Ameri- 
can material  and  Gentry's  (1973b)  description. 


Tecoma  Jussieu 

Shrubs  or  small  trees,  stems  terete,  interpetio- 
lar lines  weakly  differentiated  at  the  node  (glan- 
dular fields  absent).  Leaves  opposite,  simple,  3- 
foliolate  or  pinnately  compound  with  opposite  lat- 
eral leaflets  and  a  terminal  leaflet,  petiolate,  blades 
very  variable  in  form,  margins  serrate,  venation 
pinnate.  Inflorescences  terminal  racemes  or  ra- 
cemose panicles,  bracts  small  and  inconspicuous, 
flowers  pedicellate.  Flowers  showy,  calyx  cupu- 
late, glands  often  present  on  the  distal  surface, 
margin  with  5  deltoid  lobes;  corolla  tubular-cam- 
panulate  to  tubular-funnelform,  radially  symmet- 


156 


FIELDIANA:  BOTANY 


ric,  glabrous  externally,  yellow  to  orange,  5- 
lobed;  stamens  4,  exserted  or  included,  filaments 
of  2  lengths,  anthers  glabrous  or  puberulent,  the- 
cae  divergent,  pollen  3-colpate,  a  staminode  pres- 
ent; disc  cupular;  ovary  lepidote,  2-locular,  ovules 
in  2  series  in  each  locule,  stigma  flat.  Fruits  long 
linear  capsules,  compressed  parallel  to  the  septum 
but  dehiscing  perpendicular  to  the  septum,  valves 
smooth  and  glabrous;  seeds  thin,  with  2  lateral 


membranaceous  wings  sharply  differentiated  from 
the  central  area  of  the  seed. 

Tecoma  is  a  genus  of  14  species,  with  two  in 
Africa  and  12  in  the  New  World  (ranging  from 
the  southernmost  United  States  to  Argentina). 
Gentry  (1992)  placed  Tecomaria  under  Tecoma. 
One  species  is  both  native  and  planted  in  gardens 
in  Central  America;  the  other  is  an  introduced  or- 
namental. 


Key  to  the  Species  of  Tecoma 

la.  Corollas  yellow,  tubular-campanulate;  leaflet  blades  3-14  cm  long;  fruits  9-25  cm  long;  wild  and 
Planted  T.  stans 

Ib.  Corollas  orange  to  red-orange,  tubular;  leaflet  blades  1-4  cm  long;  fruits  5-12  cm  long;  planted 
ornamental T  capensis 


Tecoma  capensis  (Thunb.)  Lindley,  Bot.  Reg.  13: 
tab.  1117.  1827.  Bignonia  capensis  Thunb., 
Prodr.  105.  1800.  Tecomaria  capensis  (Thunb.) 
Spach.  Hist.  Nat.  Veg.  Phan.  9:  137.  1840.  Fig- 
ure 15. 

Shrubs  or  scandent  subshrubs  1-2  m  tall,  leafy 
stems  1.5-4  mm  diam.,  usually  terete,  minutely 
(0.05-0.1  mm)  puberulent.  Leaves  4-14  cm  long, 
pinnate  with  7  or  9  (11)  leaflets,  petioles  10-18 
mm  long,  0.5-1.2  mm  diam.,  rachis  sulcate  above, 
minutely  puberulent;  leaflet  blades  8-37  mm 
long,  8-16  mm  wide,  broadly  ovate  to  ovate- 
rhombic  or  broadly  ovate-elliptic,  apex  acute  to 
obtuse,  distal  margin  with  4-7  teeth/cm,  base 
broadly  cuneate  or  rounded,  mostly  glabrous  but 
with  thin  white  hairs  in  vein  axils  beneath  (dom- 
atia),  2°  veins  4-7/side.  Inflorescences  7-15  cm 
long,  racemes  with  10-40  flowers,  with  short  (4- 
9  mm)  lateral  branches,  bracts  3-14  mm  long, 
mostly  linear,  pedicels  6-12  mm  long.  Flowers 
with  calyx  4-8  mm  long,  3-4  mm  diam.,  sub- 
glabrous,  lobes  0.5-1  mm  long,  triangular  with  an 
apiculate  apex;  corolla  4-8  cm  long,  tubular,  or- 
ange to  orange-red,  glabrous  externally,  tube  2-3 
mm  diam.  at  base,  8-1 1  mm  diam.  at  mouth, 
lobes  9-14  mm  long;  thecae  3  mm  long.  Fruits 
rarely  produced  in  Central  America,  5-12  cm 
long,  6-1 1  mm  wide;  seeds  5-6  mm  long,  18-22 
mm  wide. 

Tecoma  capensis  is  planted  as  an  ornamental  in 
the  tropics  and  subtropics;  its  vigorous  growth 
and  scandent  branches  make  it  useful  in  hedges. 
The  opposite  dark  green  leaves  with  7-11  small 
serrate  leaflets  and  brilliant  curved-tubular  or- 
ange-red corollas  with  exserted  anthers  make 


these  plants  easy  to  identify.  Not  often  seen  in 
Central  America,  these  plants  are  likely  to  grow 
best  at  middle  (1000-2000  m)  elevations.  The  ge- 
nus Tecomaria  with  two  African  species  is  now 
considered  part  of  Tecoma  (Gentry,  1992).  Julia 
and  cape  honeysuckle  are  common  names. 

Tecoma  stans  (L.)  Juss.  ex  Humboldt  in  H.B.K.. 
Nov.  Gen.  Sp.  3:  144.  1819.  Bignonia  stans  L., 
Sp.  PI.  ed.  2,  2:  871.  1763.  Figure  15. 

Shrubs  or  small  trees  l-6(-12)  m  tall,  with 
many  branches,  trunk  up  to  25  cm  diam.  with 
ridged  dark  brown  bark,  leafy  stems  2-5  mm 
diam.,  glabrous,  drying  pale  yellowish  gray. 
Leaves  8-22(-28)  cm  long,  usually  with  7,  9,  or 
11  (1-5)  leaflets,  petioles  25-90  mm  long,  0.8- 
1.7  mm  diam.,  glabrous  or  puberulent  at  base  of 
leaflets,  sulcate  above,  petiolules  of  lateral  leaflets 
0-2  mm  long,  terminal  petiolules  6-18  mm  long; 
leaflet  blades  3-14  cm  long,  1-6  cm  wide  (ter- 
minal leaflet  larger  than  laterals),  elliptic-ovate  to 
narrowly  elliptic-oblong  or  lanceolate,  apex  acu- 
minate, margin  with  3  or  4  prominent  teeth/cm, 
base  cuneate  and  slightly  decurrent  on  the  petiole, 
lateral  leaflets  asymmetric  at  base,  drying  thin- 
chartaceous,  glabrous  (in  ours),  2°  veins  6-10/ 
side.  Inflorescences  5-18  cm  long,  racemose  or 
paniculate  with  8-20  flowers,  peduncles  1-3  cm 
long,  glabrous,  bracts  ca.  1  mm  long,  triangular, 
pedicels  3-8  mm  long,  glabrous.  Flowers  with  ca- 
lyx 4-7  mm  long,  2.3-4  mm  diam.,  cupulate,  gla- 
brous, with  submarginal  glands,  lobes  0.7-2  mm 
long,  acute  or  apiculate;  corolla  3.5-6  cm  long, 
to  4  cm  wide  at  the  rotate  lobes,  tubular-campan- 
ulate from  a  narrow  (2-3  mm  diam.)  base,  yellow, 


BURGER:  FLORA  COSTARICENSIS 


157 


12-20  mm  diam.  at  the  mouth,  lobes  ca.  15  X  18 
mm,  rounded;  filaments  ca.  16  and  21  mm  long, 
thecae  3-4  mm  long,  pubescent.  Fruits  9-22 
(-26)  cm  long,  5-8  mm  wide,  linear,  cylindric  or 
slightly  flattened  perpendicular  to  the  septum,  nar- 
rowed at  base  and  apex,  surfaces  glabrous,  pale 
brown;  seeds  4-6.5  mm  long,  17-28  mm  wide, 
wings  translucent,  central  area  ca.  4  X  7  mm. 

Plants  of  deciduous  and  partly  deciduous  forest 
(or  escaped  from  gardens  in  evergreen  areas)  of 
the  Pacific  slope  and  central  highlands,  10- 
1300(-1800)  m  elevation.  Flowering  primarily  in 
November-March;  fruiting  throughout  the  year. 
This  species,  originally  ranging  from  southern- 
most Arizona  and  Florida  to  Argentina,  is  widely 
planted  as  an  ornamental  in  the  tropics  and  sub- 
tropics. 

Tecoma  stans  is  recognized  by  its  opposite  im- 
paripinnately  compound  leaves  with  serrate  leaf- 
lets, large  bright  yellow  corollas  that  are  only 
slightly  bilaterally  symmetric  (obscurely  2- 
lipped),  and  long  narrow  seed  pods  with  2-winged 
seeds.  This  species  is  rather  uniform  in  morphol- 
ogy in  southern  Central  America.  In  Mexico  and 
among  cultivated  forms  there  is  extraordinary  var- 
iation, with  some  plants  having  simple  linear  (25 
X  1  cm)  leaves  and  others  with  the  blades  densely 
hirtellous  beneath.  This  great  range  of  variation 
has  produced  many  specific  and  varietal  names 
(see  synonomy  of  Gentry,  1992).  An  often  strag- 
gly growth  habit  prevents  this  species  from  being 
a  more  popular  ornamental.  Candelillo,  carbon- 
cillo,  and  vainillo  are  common  names. 


Tourrettia  Fougeroux 

Herbaceous  annual  vines,  climbing  by  means 
of  tendrils  with  3  coiled  branches  and  dichoto- 
mous  tips  (transformed  leaflets),  stems  tetragon- 
ous  with  4  longitudinal  ridges,  an  interpetiolar 
line  or  ridge  usually  present  at  the  node,  glandular 
fields  absent;  pseudostipules  absent.  Leaves  op- 
posite, twice  divided  with  3  2°  petioles  each  bear- 
ing 3-5  leaflets  or  with  2  2°  petioles  bearing  3-5 
leaflets  and  a  distally  branched  tendril;  blades 
with  strongly  serrate  margin,  lateral  leaflets  asym- 
metric and  sometimes  divided,  thin,  venation  pin- 
nate. Inflorescences  terminal  spike-like  racemes, 
sparsely  to  densely  puberulent  with  thin  gland- 
tipped  hairs,  bracts  linear,  pedicels  short,  the  up- 
per flowers  usually  sterile  and  deciduous  (with  ca- 
lyx and  corolla  not  clearly  separated).  Flowers  of 
two  kinds  with  the  distal  usually  sterile  and  short- 
er than  the  lower  (proximal)  fertile  flowers,  calyx 


tubular  near  the  base  and  separating  into  2  lobes 
distally  (upper  and  lower)  in  fertile  flowers,  de- 
ciduous during  anthesis;  corolla  tubular  and  2- 
lipped,  slightly  longer  than  the  calyx,  orange  to 
red,  puberulent  distally;  stamens  4,  of  2  lengths, 
included,  anthers  glabrous,  thecae  divaricate, 
staminode  absent;  disc  a  thin  cup;  ovary  ovoid,  4- 
locular,  ovules  many  on  uniseriate  placentas,  sur- 
face short-echinate,  style  slender,  stigma  simple. 
Fruits  slightly  woody  capsules  covered  with 
straight  and  uncinate  spines,  dehiscing  septicidal- 
ly  into  2  valves  united  near  the  base;  seeds  small, 
oblong,  with  a  thin  narrow  wing. 

Tourrettia  contains  a  single  species,  ranging 
from  the  highlands  of  Guatemala  to  Argentina. 
The  unusual  fruits  are  reminiscent  of  the  family 
Pedaliaceae,  but  the  compound  leaves,  vining  ten- 
drils, and  winged  seeds  are  characteristics  of  the 
Bignoniaceae. 

Tourrettia  lappacea  (L'Her.)  Willd.,  Sp.  PI.  ed. 
4,  3:  263.  1801.  Dombeya  lappacea  L'Her., 
Stirp.  Nov.  33,  tab  17.  1785.  T.  volubilis  J.  F. 
Gmel.,  Syst.  Nat.  2:  940.  1791.  Figure  16. 

Herbaceous  vines  to  3  m  high,  climbing  with 
slender  (0.3-0.7  mm)  coiled  tendrils  3-10  cm 
long,  leafy  sterns  0.8-5  mm  diam.,  with  2  or  4 
prominent  longitudinal  ridges,  subglabrous  or 
sparsely  puberulent  with  thin  multicellular  hairs 
0.2-0.5  mm  long  (especially  at  the  nodes).  Leaves 
opposite,  to  25  cm  long,  1°  petiole  3-5  cm  long, 
1.5-3.3  mm  diam.,  usually  glabrous,  2°  petioles 
2-6  cm  long  bearing  3-5  leaflets,  petiolules  of 
central  leaflet  8-18  mm  long;  leaflet  blades  0.8- 
10  cm  long,  the  central  leaflet  2-8(-10)  cm  long, 
1.3-5  cm  wide,  ovate  to  ovate-elliptic,  apex  acute, 
margin  strongly  serrate  with  teeth  0.3-3  mm  high 
(4-9  teeth/cm),  drying  thin  and  dull  greenish,  gla- 
brous above  and  below,  2°  veins  6-13/side.  Inflo- 
rescences to  20  cm  long,  with  both  sterile  distal 
flowers  (5-10  mm  long)  and  fertile  (to  2  cm  long) 
usually  proximal  fertile  flowers,  peduncles  3-12 
cm  long,  subglabrous  to  densely  pubescent  with 
gland-tipped  hairs,  pedicels  0.8-3.5  mm  long,  ca. 
0.3  mm  diam.,  with  a  linear  bract  to  4  mm  long 
from  near  the  base.  Flowers  dimorphic,  distal  ca- 
lyx 4-7  mm  long  and  reddish,  fertile  calyx  11- 
15  mm  long,  greenish,  lobes  slightly  longer  than 
the  tube,  subglabrous  to  densely  glandular  puber- 
ulent; corolla  of  fertile  flowers  12-24  mm  long, 
3-4  mm  diam.  near  the  base,  greenish  to  orange, 
rose  red  or  deep  scarlet  red,  puberulent  except 
near  the  base,  upper  lobe  8-12  mm  long,  hooded 
(galeate);  filaments  8-9  and  9-12  mm  long,  an- 


158 


FIELDLANA:  BOTANY 


ther  thecae  2-3  mm  long;  style  to  16  mm  long. 
Fruits  3-5  cm  long,  body  of  the  fruit  ca.  3  X  1 .4 
cm,  ellipsoid,  surface  covered  with  longer  (7-14 
mm)  uncinate  reddish  spines  and  shorter  (1-3 
mm)  acute  spines,  drying  dark  brown;  seeds  6-7 
mm  long,  4-5  mm  wide,  flat. 

Plants  of  moist  open  sites  and  forest  edges  in 
evergreen  lower  montane  forest  formations, 
(600-)1100-2100  m  elevation.  Flowering  and 
fruiting  throughout  the  year.  This  species  ranges 
from  Mexico  to  Venezuela,  Peru,  and  northwest- 
ern Argentina. 

Tourrettia  lappacea  is  a  very  unusual  herba- 
ceous vine  with  slender  divided  coiling  tendrils, 
twice-compound  opposite  leaves,  strongly  serrate 
thin  blades,  and  colorful  spike-like  inflorescences 
usually  bearing  flowers  of  two  different  forms. 
The  sterile  (usually  terminal)  flowers  are  shorter 
and  have  the  calyx  and  corolla  often  united,  while 
the  larger  fertile  flowers  have  the  calyx  clearly 
differentiated  and  deciduous.  In  addition,  the  two- 
valved  fruits  covered  with  both  shorter  straight 
spines  and  longer  hooked  spines  are  distinctive. 
Because  of  the  slender  tendril-bearing  stems  and 
spiny  fruits,  these  plants  are  sometimes  mistaken 
for  Cucurbitaceae  (see  Gentry,  1980). 


Tynanthus  Miers 

Lianas  climbing  with  the  aid  of  simple  or  trifld 
tendrils  (rarely  small  trees?),  stems  subterete  to 
quadrangular  with  4  phloem  areas  in  cross-sec- 


tion, often  with  an  interpetiolar  ridge  at  the  nodes 
but  interpetiolar  gland  fields  lacking;  pseudostip- 
ules  lacking  or  leaf-like.  Leaves  opposite.  2-  or 
3-foliolate,  often  with  the  terminal  leaflet  replaced 
by  a  simple  or  distally  trifid  tendril,  blades  entire, 
venation  pinnate.  Inflorescences  terminal  or  ax- 
illary, usually  small  (4-12  cm)  open  or  condensed 
panicles  or  dichasia,  pedicels  densely  puberulent. 
Flowers  with  small  cupular  or  campanulate  calyx, 
puberulent,  apex  subtruncate  with  5  denticulate 
lobes  or  setae,  persisting;  corolla  funnelform  and 
somewhat  curved,  bilabiate  and  split  to  near  the 
middle,  white,  puberulent  on  the  exterior,  with  4 
or  5  lobes  (upper  lobe  emarginate  or  2-lobed);  sta- 
mens 4,  of  2  lengths,  included  or  slightly  exserted. 
anthers  glabrous,  thecae  divaricate,  staminode 
present;  disc  very  small;  ovary  conical,  densely 
puberulent,  2-locular,  ovules  in  2-4  series  on  the 
placenta.  Fruits  long-linear  capsules,  slightly  flat- 
tened with  valves  parallel  to  the  septum,  flat  and 
smooth  with  a  slightly  or  distinctly  raised 
(winged)  margin,  dehiscing  septicidally;  seeds 
flat,  with  2  lateral  hyaline  wings. 

Tynanthus  is  a  genus  of  about  1 2  species  rang- 
ing disjunctly  from  southern  Mexico  and  the  West 
Indies  to  Brazil  and  Bolivia.  The  genus  has  a 
number  of  species  with  unusually  small  flowers 
for  Bignoniaceae;  for  example,  the  corollas  of  T. 
guatemalensis  J.  D.  Smith  are  only  6-9  mm  long. 
Our  species,  based  on  only  two  collections,  is  the 
only  representative  of  the  genus  known  from  be- 
tween Guatemala  and  Central  Panama;  for  com- 
parison we  include  a  key  to  the  three  species  of 
the  genus  known  in  Central  America. 


Key  to  the  Central  American  Species  of  Tynanthus 

la.  Corollas  25-30  mm  long;  small  trees  (?)  without  tendrils;  lower  leaf  surfaces  grayish  with  a  dense 
covering  of  minute  scurfy  hairs;  known  only  from  the  Caribbean  slope  of  Costa  Rica  at  100-800 
m  elevation  T.  macron  thus 

Ib.  Corollas  6-20  mm  long;  lianas  climbing  with  tendrils;  lower  leaf  surfaces  greenish,  glabrous  or 
with  minute  hairs  along  the  veins;  Panama  or  Mexico  to  Guatemala  and  not  included  in  the  de- 
scriptions   2 

2a.  Corollas  6-9  mm  long;  leaves  mostly  elliptic  to  ovate-elliptic;  ranging  from  central  Mexico  to  Belize 
and  Guatemala T.  guatemalensis 

2b.  Corollas  12-20  mm  long;  leaves  mostly  ovate  to  broadly  ovate;  central  to  eastern  Panama 

T.  croatianus 


Tynanthus  macranthus  L.  O.  Williams,  Fieldi- 
ana  Bot.  31:  250.  1967.  Figure  12. 

Small  trees  3  m  tall  (in  the  type)  but  also  lia- 
nas, leafy  stems  1.5-5  mm  diam.,  densely  grayish 


puberulent  with  minute  (ca.  0.1  mm)  rounded, 
peltate  or  scurfy  hairs,  terete,  node  with  a  slightly 
elevated  interpetiolar  ridge.  Leaves  2-foliolatc, 
petioles  8-17  mm  long,  ca.  1  mm  diam.,  densely 
puberulent  like  the  stems,  petiolules  6-1 1  mm 


BURGER:  FLORA  COSTARICENSIS 


159 


long,  sulcate  above;  leaf  blades  4-12  cm  long,  2- 
6  cm  wide,  elliptic-oblong  to  oblong,  apex  acute 
to  abruptly  short-acuminate,  base  asymmetric 
with  narrowed  and  slightly  rounded  sides,  drying 
stiffly  chartaceous  and  brown  above,  upper  sur- 
face minutely  papillate  puberulent  or  subglabrous, 
lower  surface  grayish  to  brownish  and  densely  to 
sparsely  puberulent  with  whitish  hairs  0.05-0.1 
mm  long,  2°  veins  3-6/side.  Inflorescences  axil- 
lary, 1-3  cm  long,  few-flowered  racemes  or  di- 
chasia,  peduncle  10-16  mm  long,  1  mm  diam., 
densely  puberulent,  bracts  1-2  mm  long,  cadu- 
cous, pedicels  3-6  mm  long.  Flowers  with  calyx 
6-9  mm  long,  4-5  mm  diam.,  tubular-campanu- 
late,  densely  pale  brownish  puberulent  (dried), 
margin  subentire  with  5  minute  (0.3  mm)  teeth; 
corolla  22-23  mm  long,  tubular-funnelform,  2- 
lipped  and  split  along  the  sides  to  within  5-8  mm 
of  the  base,  white,  densely  pale  yellowish  brown 
puberulent  on  the  exterior  when  dried,  tube  2-4 
mm  diam.  near  the  base,  5-8  mm  diam.  at  the 
mouth  (lower  lip  becoming  bent  downward  and 
recurved  in  Barringer  et  al.  2671),  lobes  5-12 
mm  long,  obtuse  at  the  apex;  thecae  ca.  2  mm 
long,  the  pair  U-shaped  when  dried;  ovary  mi- 
nutely puberulent.  Fruits  unknown. 

Rarely  collected  plants  of  wet  evergreen  rain 
forest  formations  of  the  Caribbean  slope,  100-800 
m  elevation.  Flowering  in  late  April  and  early 
July.  Endemic  to  Costa  Rica  and  northwestern 
Panama. 

Tynanthus  macranthus  is  recognized  by  its  op- 
posite bifoliolate  leaves  with  blades  usually  gray- 
ish puberulent  beneath,  the  short  axillary  racemes, 
the  densely  puberulous  calyx  with  minutely  den- 
tate margin,  and  the  densely  puberulous  white  co- 
rollas deeply  split  along  the  two  sides.  Although 
the  type  (Lent  42)  was  said  to  be  a  tree,  other 
collections  appear  to  be  lianas.  The  flowers  of  this 
species,  while  modest  in  size  for  the  family,  are 
large  for  the  genus.  No  other  bignon  in  our  flora 
has  a  corolla  tube  so  deeply  incised. 


Xylophragma  Sprague 

Lianas  climbing  with  simple  tendrils  (undivid- 
ed distally),  stems  terete  or  subquadrangular,  with 
4  phloem  areas  in  cross-section,  interpetiolar 
glandular  fields  present;  pseudostipules  short, 
acute.  Leaves  opposite,  3-foliolate  or  2-foliolate 
with  the  terminal  leaflet  sometimes  replaced  by  a 
tendril,  pubescence  of  small  simple  or  branched 
(dendritic)  hairs,  venation  pinnate.  Inflorescences 


racemes  or  panicles  with  racemose  branches,  usu- 
ally axillary  on  older  branchlets,  bracts  present, 
pedicels  often  with  paired  bracteoles.  Flowers 
with  cupular  calyx,  margin  with  5  short  teeth,  pu- 
bescent with  stellate  or  branched  hairs;  corolla 
tubular-campanulate,  slightly  bilabiate,  rose  to 
lavender  or  purple,  puberulent  on  the  exterior;  sta- 
mens 4,  of  2  lengths,  anthers  glabrous,  thecae 
straight  and  divaricate,  a  staminode  present;  disc 
cupular;  ovary  ovoid,  2-locular,  ovules  6-8  series 
on  each  placenta,  stigma  simple.  Fruits  capsules, 
valves  woody,  flat,  and  parallel  to  the  septum,  me- 
dian vein  not  evident,  smooth  or  with  raised  glan- 
dular areas;  seeds  thin  with  2  lateral  membrana- 
ceous  wings  clearly  delineated  from  the  central 
seed  area. 

Xylophragma  is  a  genus  of  four  species  cen- 
tered in  South  America,  with  one  species  ranging 
northward  as  far  as  Mexico. 

Xylophragma  seemannianum  (Kuntze)  Sandw., 
Kew  Bull.  1953:  469.  1954.  Saldanhaea  see- 
manniana  Kuntze,  Rev.  Gen.  PI.  2:  480.  1891. 
Distictis  rovirosana  J.  D.  Smith,  Bot.  Gaz.  20: 
7.  1895.  Adenocalymma  cocleense  Pittier,  Con- 
trib.  U.S.  Natl.  Herb.  18:  255.  1917.  Figure  21. 

Lianas  stems  to  10  cm  diam.,  tendrils  to  20  cm 
long,  leafy  stems  2-6  mm  diam.,  at  first  puberu- 
lent with  branched  or  stellate  hairs  ca.  0.2  mm 
long,  glabrescent,  terete,  developing  whitish  len- 
ticels,  gland  fields  often  present  at  the  nodes; 
pseudostipules  1-3  mm  long,  conical.  Leaves  3- 
foliolate  or  2-foliolate  with  a  tendril,  petioles  4- 
16(-20)  cm  long,  0.7-2.2  mm  diam.,  minutely  pu- 
berulent with  straight  or  crooked  hairs  0.1-0.3 
mm  long,  petiolules  6-40(-70)  mm  long,  petio- 
lules  of  central  leaf  often  twice  the  length  of  lat- 
eral petiolules;  leaflet  blades  (4-)6-16(-23)  cm 
long,  (2.5-)4-10(-13)  cm  wide,  ovate,  broadly  el- 
liptic, obovate  or  slightly  rhombic,  apex  acute  to 
acuminate,  base  obtuse  to  slightly  rounded  and 
truncated,  drying  chartaceous,  minutely  puberu- 
lent on  the  veins  above,  usually  puberulent  be- 
neath with  branched  hairs  0.2-0.5  mm  long  be- 
neath, 2°  veins  5-7/side.  Inflorescences  terminal 
on  axillary  short-shoots,  racemes  or  short  pani- 
cles, peduncles  5-20  mm  long,  ca.  1  mm  diam., 
densely  stellate  puberulent,  bracts  3-6  mm  long, 
linear  to  linear-lanceolate,  pedicels  5-15  mm 
long.  Flowers  with  calyx  4-8  mm  long,  3-5  mm 
diam.,  densely  papillate  or  stellate  puberulent, 
teeth  (lobes)  0.2-1  mm  long,  ca.  0.5  mm  wide; 
corolla  (2.5-)3-6  cm  long,  tubular-funnelform, 


160 


FIELDIANA:  BOTANY 


lavender  to  rose  and  minutely  puberulent  on  the 
exterior,  yellow  and  white  within,  tube  1.5-2  mm 
diam.  near  the  base,  10-18  mm  wide  at  the 
mouth,  lobes  to  22  X  16  mm  and  acute;  filaments 
ca.  9  and  14  mm  long,  thecae  2.3-3.5  mm  long; 
ovary  ca.  3  mm  long,  lepidote.  Fruits  5-16  cm 
long,  3-5  cm  wide,  ca.  5  mm  thick,  oblong-round- 
ed, valves  flat  and  smooth;  seeds  14-22 
(-29)  mm  long,  28-52  mm  wide,  central  dark  area 
17-24  mm  wide. 

Common  deciduous  lianas  of  seasonally  dry 
deciduous  and  partly  deciduous  forest  formations 
(rarely  collected  in  evergreen  forest  formations), 
1-500  m  elevation.  Flowering  in  March-May 
when  the  plants  are  usually  leafless;  fruits  mature 
and  open  in  the  following  year's  dry  season.  This 
species  ranges  from  Veracruz,  Mexico,  to  north- 
ern Brazil. 

Xylophragma  seemannianum  is  recognized  by 
its  climbing  habit  with  simple  woody  tendrils 
(rarely  shrubs),  the  usually  trifoliolate  leaves, 
short  inflorescences  usually  blooming  when  the 
leaves  are  absent,  showy  lavender  or  magenta  co- 
rollas, and  oblong  flattened  fruits  with  two- 
winged  seeds.  The  gland  fields  at  older  nodes, 
larger  leaves  with  symmetric  lateral  leaflets,  and 
pubescence  of  small  branched  hairs  (dendroid,  of- 
ten appearing  farinose)  are  also  helpful  in  recog- 
nizing this  species. 

List  of  Accepted  Species  of  Bignoniaceae 

Key:  END-CR  =  endemic  to  continental  Costa 
Rica;  END-CR&WP  =  endemic  to  Costa  Rica 
and  western  Panama;  END-WP  =  endemic  to 
western  Panama;  INTRO  =  introduced  weed; 
ORNAM  =  cultivated  ornamental;  ORNAM  & 
NAT  =  cultivated  and  naturalized;  ??  =  not  col- 
lected in  Costa  Rica  but  known  from  nearby  ar- 
eas. Total  number  of  species  covered  is  79;  the 
number  of  documented  native  species  is  69.  One 
new  species,  Amphitecna  gentryi,  is  described 
here. 

Adenocalymma  inundatum 

Amphilophium  paniculatum 

Amphilophium  pannosum 

Amphitecna  gentryi  END-CR  &  SP.  NOV. 

Amphitecna  isthmica 

Amphitecna  kennedyi 

Amphitecna  latifolia 

Amphitecna  sessilifolia  END-CR  &  WP 

Anemopaegma  chrysanthum 

Anemopaegma  chrysoleucum 

Anemopaegma  orbiculatum 

Anemopaegma  puberulum 


Anemopaegma  santaritense 
Arrabidaea  candicans 
Arrabidaea  chica 
Arrabidaea  conjugata 
Arrabidaea  corallina 
A  rrabidaea  costaricensis 
Arrabidaea  florida 
A  rrabidaea  mollissimu 
Arrabidaea  patellifera 
Arrabidaea  pubescens  ?? 
Arrabidaea  verrucosa 

Callichlamyx  latifolia 
Ceratophytum  tetragonolobum 
Clytostoma  binatum 
Crescentia  alata 
Crescentia  cujete 
Cydista  aequinoctialis 
Cydista  diversifolia 
Cydista  heterophylla 
Cydista  lilacina 
Cydista  potosina 

Distictella  magnoliifolia 
Godmania  aesculifolia 

Jacaranda  caucana 

Jacaranda  copaia 

Jacaranda  mimosifolia  ORNAM 

Kigelia  pinnata  ORNAM 

Lundia  corymbifera 
Lundia  puberula 

Macfadyena  uncata 
Macfadyena  unguis-cati 
Mansoa  hymenaea 
Mansoa  kerere 
Mansoa  pan'ifolia 
Mansoa  standleyi 
Mansoa  verrucifera 
Martinella  obovata 
Melloa  quadrivalvis 
Mussatia  hyacinthina 

Parabignonia  steyermarkii 
Paragonia  pyramidata 
Parmentiera  aculeata  ORNAM 
Parmentiera  cereifera  ORNAM 
Parmentiera  dressleri 
Parmentiera  macropltylla 
Parmentiera  valerii  END-CR 
Phyrganocydia  <  -orymhoxa 
Phryganocydia  phellosperma 
Pithecoctenium  crucigerum 
Pleonotoma  variabilis 
Podranea  ricolaaiana  ORNAM 
Pyrostegia  venusta  ORNAM 

Spathodea  campanulata  ORNAM 
Stizophyllum  inaequilaterum 
Sti-ophyllum  riparium 

Tabebuia  chrysantha 
Tahebuia  guayacan 
Tabebuia  impetiginosa 
Tabebuia  ochracea 
Tabebuia  palustris 
Tabebuia  rosea 


BURGER:  FLORA  COSTARICENSIS 


161 


Tanned  urn  jaroba 

Tecoma  capensis  ORNAM 

Tecomu  stems  ORNAM  &  native 

Tourretliei  lappacea 

Tynunllnis  meicreinthtis  END-CR  &  WP 

Xyliiphreignui  seemannianum 


PEDALIACEAE 

By  William  Burger 

Herbs  or  more  rarely  shrubs  or  small  trees, 
stems  erect,  often  with  mucilaginous  glands  that 
become  slimy  when  wet;  stipules  absent.  Leaves 
opposite  or  the  distal  alternate,  petiolate,  simple, 
entire  or  lobed,  pinnately  or  palmately  veined.  In- 
florescences of  usually  solitary  axillary  flowers, 
sometimes  cymose,  usually  with  glands  at  the 
base  of  the  pedicel.  Flowers  bisexual,  bilaterally 
symmetric,  sepals  5,  united  at  the  base  or  free; 
corolla  tubular  to  campanulate  and  slightly  2- 
lipped,  5-lobed,  imbricate  in  bud;  stamens  4,  fil- 
aments of  2  lengths,  anthers  dorsifixed,  parallel  or 
separate,  a  staminode  usually  present;  disc  present 
on  1  side  of  the  ovary;  ovary  superior  (rarely  in- 
ferior), 2-  or  4-locular,  placentation  axile,  ovules 
few  to  many,  style  slender,  stigma  2-lobed.  Fruits 
capsules  or  indehiscent,  often  woody  and  spiny  or 
with  projections;  seeds  few  to  many,  smooth,  en- 
dosperm thin. 

Pedaliaceae  is  an  Old  World  family  of  about  1 5 
genera  and  50  species.  Closely  related  to  Scro- 
phulariaceae,  this  family  is  distinguished  by  its 
slime  cells  and  unusual  fruits.  Some  authors  have 
enlarged  this  family  to  include  Martyniaceae 
(Cronquist,  1981),  but  that  family  has  parietal  pla- 
centation and  is  restricted  to  the  New  World.  The 
only  representative  of  this  family  likely  to  be  seen 
in  Central  America  is  Sesamum  orientate,  the 
source  of  sesame  seed. 


Sesamum  Linnaeus 

Annual  or  perennial  herbs,  stems  erect,  usually 
with  mucilaginous  sap  or  glands.  Leaves  opposite 
or  alternate,  petiolate,  blades  with  pinnate  vena- 
tion. Inflorescences  of  usually  solitary  flowers  in 
leaf  axils  or  sometimes  in  few-flowered  cymes, 
pedicels  subtended  by  lateral  glands.  Flowers 
with  5-parted  calyx;  corolla  tube  oblique  at  the 
base  and  slightly  expanded  (gibbous),  slightly  2- 


lipped  with  5  spreading  lobes;  stamens  4  (rarely 
2),  inserted  near  the  base  of  the  corolla  tube,  an- 
thers divergent  (sagittate);  ovary  superior  or  part- 
ly inferior,  2-locular  with  many  ovules.  Fruits 
capsules,  oblong,  somewhat  4-angled  in  cross- 
section,  loculicidally  dehiscent. 

Sesamum  is  a  tropical  genus  of  about  12  spe- 
cies, native  to  Africa  and  Asia.  The  following  spe- 
cies has  been  widely  cultivated  in  tropical  regions 
for  its  oil-containing  seeds. 

Sesamum  orientate  L.,  Sp.  PI.  634.  1753.  S.  in- 
dicum  L.,  Sp.  PI.  634.  1753.  Figure  27. 

Erect  annual  herbs  to  ca.  1 .5  m  tall,  stems  sim- 
ple or  branched,  leafy  stems  1-5  mm  diam.,  with 
thin  whitish  hairs  0.2-1.5  mm  long;  bracts  may 
resemble  stipules  at  flowering  nodes.  Leaves 
mostly  alternate,  petioles  6-35(-80)  mm  long, 
0.5-1. 3(-2)  mm  diam.,  puberulent  with  thin  hairs 
ca.  0.5  mm  long;  leaf  blades  3-15  cm  long,  0.7- 
9  cm  wide,  varying  from  broadly  ovate-triangular 
(or  deeply  3-lobed  in  basal  leaves)  to  lanceolate 
or  narrowly  oblong,  apex  acute  to  obtuse,  margin 
entire  (sometimes  with  rounded  lobes),  base  acute 
to  obtuse,  drying  chartaceous,  sparsely  puberulent 
with  thin  hairs  on  both  surfaces,  2°  veins  5-6/side 
and  strongly  ascending.  Inflorescences  of  solitary 
axillary  flowers,  bracts  to  4  mm  long,  linear,  ped- 
icels 1-2  mm  long  (to  5  mm  in  fruit),  with  a  pair 
of  lateral  sessile  round  glands  (0.7  mm  diam.)  at 
the  base,  puberulent.  Flowers  with  calyx  4.5-6.5 
mm  long,  sepals  united  only  at  the  base,  ca.  5  mm 
long,  narrowly  triangular,  acute,  with  thin  white 
hairs;  corolla  20-30  mm  long,  white  to  pink  or 
lavender,  sparsely  puberulent  with  thin  hairs  0.3- 
1 .5  mm  long,  lobes  2-7  mm  long,  broadly  round- 
ed. Fruits  17-30  mm  long,  8-10  mm  wide,  ob- 
long-rectangular, base  rounded,  apex  acute  or 
apiculate,  with  a  central  longitudinal  sulcus  on  the 
nondehiscent  sides,  pale  brown,  puberulent;  seeds 
3  X  1 .8  X  1  mm,  ovate-lenticular,  lustrous  white. 

Cultivated  or  occasionally  escaped  plants  in 
seasonally  dry  deciduous  areas,  100-800  m  ele- 
vation in  Central  America.  Flowering  in  June-De- 
cember; fruiting  in  July-January.  Rarely  encoun- 
tered in  Costa  Rica,  this  species  is  naturalized  in 
parts  of  Guatemala  and  Honduras.  Cultivated 
since  ancient  times  in  southern  and  western  Asia, 
it  has  been  introduced  throughout  the  tropics. 

Sesamum  orientale  is  recognized  by  its  solitary 
axillary  flowers  subtended  by  glands  at  the  base 
of  the  pedicels,  bent  campanulate  white  or  pink 
corollas,  and  the  distinctive  oblong  capsules. 


162 


FIELDIANA:  BOTANY 


Redrawn 

froK 

S.Troyo 


Tetranema  florlbundum 

FIG.  27.     Unusual  herbs  of  the  Martyniaccae,  Pedaliaccac,  and  Scrophulariaccae  families. 


BURGER:  FLORA  COSTARICENSIS 


163 


Ajonjoli  and  sesame  are  common  names.  The  oil 
expressed  from  the  seeds  is  used  much  like  olive 
oil,  and  the  seeds  are  used  on  bread  or  in  making 
sweets. 


MARTYNIACEAE 

By  William  Burger 

Herbs,  usually  with  viscid  gland-tipped  multi- 
cellular  hairs;  stipules  absent.  Leaves  alternate  or 
opposite,  simple,  petiolate,  leaf  blades  entire  to 
undulate,  dentate  or  somewhat  lobed.  Inflores- 
cences terminal  or  axillary,  racemose,  flowers 
pedicellate,  often  subtended  by  1  or  2  large  brac- 
teoles,  sometimes  becoming  thick  and  fleshy  in 
age.  Flowers  bisexual,  calyx  5-parted  or  5-lobed 
and  split  down  1  side,  glandular  puberulent  exter- 
nally; corolla  bilaterally  symmetric  and  somewhat 
2-lipped,  tubular  at  base  and  expanded  to  cam- 
panulate  or  funnelform  distally,  5-lobed  with  the 
2  upper  lobes  exterior  in  bud;  stamens  4  or  2, 
filaments  equal  or  of  2  lengths,  arising  from  the 
base  of  the  expanded  corolla  tube,  anthers  of  each 
stamen  pair  at  first  coherent,  2-thecous,  stami- 
nodes  1-3;  disc  annular;  ovary  superior,  unilocu- 
lar  but  with  2  intruding  parietal  placentas  and  ap- 
pearing 2-  or  4-locular  by  intrusion  of  the  placen- 
tas, ovules  few  to  many,  style  slender,  stigma  2- 
lobed.  Fruits  woody  capsules,  usually  with  distal 
recurved  horns,  exocarp  fleshy  and  deciduous,  en- 
docarp  hard  and  woody;  seeds  few  to  many, 
sculptured,  black,  compressed  or  oblong. 

This  family  of  three  genera  is  restricted  to  the 
New  World,  but  some  authors  place  it  in  the  Pe- 
daliaceae  (Cronquist,  1981).  The  Martyniaceae 
differ  from  the  Pedaliaceae  in  having  intruding 
parietal  placentae  and  lacking  glands  at  the  base 
of  the  pedicels,  and  they  do  not  become  slimy 
when  wet.  The  monotypic  genus  Martynia  is  rare- 
ly collected  in  Costa  Rica.  A  second  genus  and 
species,  Proboscidia  triloba  (Cham.  &  Schldl.) 
Decne.,  is  found  in  northern  Central  America. 
That  genus  is  distinguished  by  having  a  spathe- 
like  calyx,  four  fertile  stamens,  and  distal  horns 
of  the  capsule  usually  surpassing  the  body  of  the 
fruit  in  size. 


Martynia  Linnaeus 

Erect  annual  herbs,  puberulent  to  villous  with 
thin   multicellular  gland-tipped   viscid   hairs  on 


most  parts.  Leaves  opposite  or  alternate  distally, 
petiolate,  blade  broadly  ovate  with  palmate  ve- 
nation. Inflorescences  terminal  or  axillary,  short- 
racemose,  viscid  puberulent,  flowers  subtended  by 
2  large  bracts.  Flowers  with  deeply  5-parted  ca- 
lyx, sepals  unequal,  thin-textured;  corolla  cam- 
panulate  from  a  short  tubular  base,  somewhat  2- 
lipped  and  oblique,  with  5  broadly  rounded  lobes; 
stamens  2,  included,  filaments  borne  from  the 
apex  of  the  short  narrow  basal  tube,  anthers 
strongly  divaricate  (180°)  and  coherent  on  their 
sides,  staminodes  2  or  3;  ovary  with  2  intruding 
parietal  placentae,  stigma  2-lobed.  Fruits  a 
horned  capsule,  exocarp  glandular  pubescent  but 
deciduous  and  exposing  the  hard  woody  exocarp, 
adaxial  and  abaxial  sides  of  the  exocarp  each  with 
4  longitudinal  ribs  separated  by  deep  sulci,  the 
hard  recurved  distal  horns  shorter  than  the  body 
of  the  fruit. 

Martynia  annua  L.,  Sp.  PI.  618.  1753,  non  L. 
Syst.  Nat.  1113.  1759.  M.  diandra  Gloxin,  Obs. 
Bot.  14.  1785.  M.  angulosa  Lam.,  Encycl. 
Meth.  Bot.  2:  112.  1786.  Carpoceras  angulata 
A.  Rich.,  Bull.  Sci.  Nat.  Geol.  21:  98.  1830. 
Disteira  angulosa  Raf.,  Fl.  Telluriana:  68.  1881. 
Vatkea  diandra  O.  Hoffm.,  Verh.  Bot.  Bran- 
denb.  73:  45.  1881;  Linnaea  43:  554.  1882.  Fig- 
ure 27. 

Annual  erect  herbs  0.3-1.5  m  tall,  stems 
sparsely  to  densely  viscid  pubescent  with  hairs  ca. 
0.5  mm  long.  Leaves  with  petioles  4-14(-40)  cm 
long,  1.4-5  mm  diam.,  villous  with  gland-tipped 
hairs  0.3-1  mm  long;  leaf  blades  7-24(-40)  cm 
long,  5-21  (-32)  cm  wide,  ovate  to  ovate-trian- 
gular (sometimes  weakly  3-  or  5-lobed),  apex  ob- 
tuse, margin  with  widely  triangular  or  obtuse 
lobes  and  more  numerous  minute  (0.5  mm)  teeth, 
base  cordate  to  subcordate,  drying  membrana- 
ceous,  upper  surface  with  scattered  short  hairs, 
lower  surface  with  hairs  and  pellucid  dots,  vena- 
tion palmate,  2°  veins  3-4/side.  Inflorescences  3- 
6  cm  long,  terminal  or  axillary  to  distal  leaves, 
with  5-15(-20)  flowers  bracts  12-23  mm  long, 
broadly  elliptic  and  somewhat  similar  to  the  se- 
pals, pedicles  12-28  mm  long,  slender,  glandular 
pubescent.  Flowers  with  5  unequal  sepals  16-20 
mm  long,  4-7  mm  wide,  oblong  to  broadly  ellip- 
tic or  oblanceolate,  obtuse  or  rounded  at  the  apex, 
ciliolate  on  the  edge;  corolla  40-55  mm  long, 
white  or  pinkish  white  with  rounded  purple  spots 
on  the  sepal  lobes  within  and  yellow  area  on  the 
floor  of  the  throat,  lobes  broadly  rounded  distally, 


164 


FIELDIANA:  BOTANY 


the  2  smaller  (adaxial)  lobes  held  erect,  the  broad 
abaxial  lobe  forming  the  lower  lip;  anthers  ca.  8 
mm  wide,  filaments  ca.  12  mm  long;  style  ca.  30 
mm  long.  Fruits  3-4  cm  long,  1.6-2  cm  wide, 
slightly  curved  and  boat-shaped  (curved  ellip- 
soid), the  distal  ends  with  hard  recurved  sharp- 
tipped  horns  ca.  1  cm  long,  woody  endocarp  with 
deep  sulci  and  drying  black. 

Plants  of  seasonally  dry  deciduous  forest  areas 
5-300  m  elevation  (to  1000  m  in  Honduras,  to 
2400  m  in  Guatemala).  Flowering  in  July-August; 
fruiting  in  August-February.  Although  common 
in  central  Honduras  and  along  the  shore  of  Lake 
Nicaragua  near  Granada,  this  species  has  been 
collected  only  a  few  times  in  Guanacaste  Prov- 
ince. The  species  ranges  from  Mexico  to  north- 
eastern Costa  Rica. 

Martynia  annua  is  recognized  by  its  short-lived 
herbaceous  habit,  viscid  pubescence,  broad  pal- 
mately  veined  leaves,  racemose  inflorescences, 
five  large  subequal  sepals,  and  whitish  slightly 
two  lipped  corolla  with  purple  spots.  The  mature 
woody  fruits  are  especially  distinctive  with  their 
two  distal  recurved  sharp-tipped  horns.  The  some- 
what coherent  anthers  are  reminiscent  of  those 
seen  in  Gesneriaceae.  Unas  de  diablo  is  a  com- 
mon name. 


OROBANCHACEAE 

By  Luis  D.  Gomez  P.  and  William  Burger 

REFERENCES — L.  D.  G6mez,  Notes  on  the  bi- 
ology of  Central  American  Orobanchaceae.  Bre- 
nesia  17:  389-396.  1980.  G.  B.  von  Mannagetta, 
Orobanchaceae,  in  A.  Engler,  Pflanzenreich  IV, 
261:  1-348.  1930.  J.  Thieret,  The  Genera  of  Oro- 
banchaceae in  the  Southeastern  United  States.  J. 
Arnold  Arbor.  52:  404-434.  1971. 


Annual  or  perennial  herbs,  all  parts  lacking 
chlorophyll,  parasitic  on  the  roots  of  other  plants, 
arising  from  a  thickened  base,  stems  simple  or 
few-branched,  thin  to  thick,  usually  succulent, 
glabrous  to  pubescent,  hairs  simple  or  glandular; 
stipules  absent.  Leaves  alternate  and  simple,  re- 
duced to  sessile  scales,  without  chlorophyll,  mar- 
gins entire.  Inflorescences  simple  or  less  often 
branched,  flowers  in  racemes  or  spikes  (rarely  sol- 
itary and  terminal),  flowers  solitary  in  the  axils  of 
bracts,  sessile  or  pedicellate,  bracteoles  usually 
present.  Flowers  bisexual,  calyx  radially  or  bilat- 
erally symmetric,  tubular  or  campanulate  with  2- 
5  unequal  lobes  or  teeth,  sometimes  split  on  1  or 
2  sides,  open  or  valvate  in  bud;  corolla  tubular  to 
campanulate  and  usually  2-lipped,  4-  or  5-lobed. 
tube  curved  or  straight,  upper  lip  2-lobed  or  en- 
tire; stamens  4,  included  or  exserted,  alternate 
with  the  corolla  lobes,  filaments  2  pairs  of  unequal 
length,  borne  on  the  proximal  half  of  the  corolla 
tube,  free,  anthers  free,  dorsifixed,  2  thecous  with 
1  or  2  thecae  fertile,  free  or  coherent,  opening  by 
a  longitudinal  slit,  a  staminode  present  or  absent; 
ovary  superior,  1-locular  with  4  (2-6)  intruded  pa- 
rietal placentae,  ovules  many  and  anatropous, 
style  simple,  stigma  simple  or  2-4-lobed.  Fruits 
capsules,  dehiscing  loculicidally  into  2  or  3 
valves;  seeds  many,  small,  with  ornamented  testa, 
embryo  undifferentiated. 

A  family  of  ca.  .17  genera  and  200  species 
(Mabberley,  1987),  in  temperate  and  subtropical 
climates  throughout  the  northern  hemisphere,  but 
with  the  greatest  number  of  species  in  the  Old 
World.  The  lack  of  chlorophyll,  usually  simple  or 
few-branched  stems,  tubular  two-lipped  corollas, 
unilocular  ovary  with  intruding  parietal  placentae, 
and  capsular  fruits  make  these  plants  easy  to  iden- 
tify. The  aboveground  parts  of  these  plants  are 
short-lived  and  are  easily  missed  by  collectors. 
One  genus  is  native  to  Central  America  and  one 
may  be  found  as  an  introduced  weed.  This  family 
is  closely  related  to  Scrophulariaceae. 


Key  to  the  Genera  of  Orobanchaceae 

la.  Scales  overlapping  on  the  lower  stem;  corolla  glabrous  externally;  anthers  slightly  exserted;  parasites 
in  high-elevation  oak  forests •  Conopholis 

Ib.  Scales  not  overlapping  on  the  lower  stems;  corolla  puberulent  externally;  anthers  included;  rarely 
collected  parasites  at  higher  elevations  in  open  fields  Orobanche 


BURGER:  FLORA  COSTARICENSIS 


165 


Conopholis  Wallroth 

REFERENCE — R.  Haynes,  A  monograph  of  the 
genus  Conopholis  (Orobanchaceae).  Sida  4:  246- 
264.  1971. 

Herbs  with  short  erect  unbranched  stems,  par- 
asitic on  oak  (Quercus)  roots,  aerial  parts  of  1  or 
several  annual  flowering  stems  that  are  covered 
by  imbricate  scales  near  the  base,  stems  arising 
from  a  gall-like  perennial  base,  simple  or  rarely 
branched,  glabrous  to  densely  glandular-pubes- 
cent. Leaves  represented  by  broad-based  sessile 
scales  that  are  imbricate  at  the  base  and  alternate 
distally,  yellowish  white  at  first  and  becoming 
brown,  intergrading  with  the  very  similar  bracts. 
Inflorescences  spike-like  racemes  with  thick  axis, 
flowers  separate  or  crowded,  bracts  exceeding  the 
calyx  in  length,  sessile  or  short-pedicellate,  1  or 
2  small  bracteoles  arising  from  the  base  of  the 
calyx.  Flowers  with  tubular  or  spathe-like  calyx, 
cleft  on  one  side,  irregularly  2-,  4-,  or  5-lobed  or 
dentate  on  the  distal  margin,  divisions  acute  or 
rounded;  corolla  tubular  and  2-lipped,  yellowish 
white,  the  upper  lip  curved  or  straight,  the  lower 
lip  with  (1  or  2)  3  subequal  spreading  lobes;  sta- 
mens 4,  exserted,  anthers  with  spurs,  thecae 
slightly  divergent,  glabrous  or  sparsely  pilose; 
ovary  unilocular  with  4  parietal  placentas,  stigma 
discoidal.  Fruits  2-valved  capsules,  usually  en- 
closed by  the  persisting  perianth,  dehiscing  irreg- 
ularly; seeds  oval  to  angular,  testa  reticulate. 

Conopholis  is  a  North  and  Central  American 
genus  of  two  species,  according  to  Haynes  (1971, 
reference  above).  A  brief  overview,  however, 
makes  it  appear  that  the  collections  of  Conopholis 
could  also  be  interpreted  as  representing  either  a 
single  variable  species  or  several  species.  The 
main  characteristics  used  by  Haynes  to  distinguish 
the  two  species,  apart  from  geographic  separation, 
are  quite  variable,  even  within  a  single  popula- 
tion. Earlier,  Standley  (1938)  placed  the  Costa  Ri- 
can  material  under  the  name  C.  americana  (L.) 
Wallroth.  The  genus  is  distinctive  because  of  its 
lack  of  green  pigmentation,  parasitic  association 
with  oak  trees,  short  thick  succulent  spike-like 
stems  with  imbricate  bract-like  scales,  many  dis- 
tally crowded  flowers,  and  pale  yellowish  tubular 
two-lipped  corollas. 

Conopholis  alpina  Liebmann,  Forh.  Skand.  Na- 
turf.  Mode  4:  184.  1847.  C.  sylvatica  Lieb- 
mann, Forh.  Skand.  Naturf.  Mode  4:  185.  C. 
mexicana  Gray  ex  Watson,  Proc.  Am.  Acad. 


Arts  Sci.  18:  131.  1883.  C.  panamensis  Wood- 
son,  Ann.  Missouri  Bot.  Gard.  25:  835.  1935. 
C.  alpina  var.  mexicana  (Gray  ex  Watson) 
Haynes,  Sida  4:  255.  1971.  Figure  28. 

Parasitic  herbs,  stems  6-24(-30)  cm  tall,  4-12 
mm  diam.,  glabrous,  proximal  scales  imbricate, 
thickened  at  the  base,  glabrous.  Leaves  repre- 
sented by  sessile  scales  6-18(-21)  mm  long,  4- 
7(-l  1)  mm  wide,  ovate-lanceolate  to  narrowly  ob- 
long-lanceolate, widest  just  above  the  base,  apex 
acute  to  acuminate,  margins  entire,  drying  sub- 
coriaceous,  surfaces  glabrous,  often  dark  brown 
distally  (in  life),  drying  uniformly  yellowish 
brown.  Inflorescences  2-18  cm  long,  2-4  cm 
wide,  bracts  8-20  mm  long,  separate  or  overlap- 
ping, usually  concealing  the  calyx,  pedicels  0-3 
mm  long,  bracteoles  2-4  mm  long,  ca.  1.5  mm 
wide,  ovate-elliptic,  on  the  lateral  sides  of  the  ca- 
lyx or  absent.  Flowers  with  calyx  8-12  mm  long, 
3-4  mm  diam.,  ovoid-tubular,  usually  split  down 
the  anterior  side,  lobes  2-5,  acute  to  rounded;  co- 
rolla 7-15  mm  long,  curved  downward,  white  to 
greenish  white  (yellowish  white  when  dried),  gla- 
brous externally;  filaments  9-15  mm  long,  anthers 
1.5-2.5  mm  long,  with  basal  appendages  ca.  0.4 
mm  long,  usually  glabrous.  Fruits  8-18  mm  long, 
6-1 1  mm  diam.,  ovoid-rounded,  drying  dark, 
style  often  persisting;  seeds  0.6-1  mm  long,  with 
rounded  edges,  surface  yellowish  to  dark  brown 
and  lustrous,  minutely  reticulate  (20X). 

Plants  of  the  forest  floor  in  montane  oak  forests, 
2000-2800  m  elevation.  Flowering  in  December- 
April;  fruiting  in  February-July.  Costa  Rican  col- 
lections come  from  the  Cordillera  de  Talamanca 
and  Volcan  Irazu.  This  species,  as  circumscribed 
by  Haynes,  ranges  from  the  southwestern  United 
States  to  northern  Panama. 

Conopholis  alpina  is  recognized  by  its  lack  of 
green  pigment,  restriction  to  high-elevation  oak 
forests,  short  thick  unbranched  stems,  curved  tu- 
bular yellowish  corollas,  exserted  stamens,  and 
many-seeded  capsules.  The  seeds  germinate  only 
near  young  oak  roots,  and  the  flowers  are  polli- 
nated by  bumblebees  (Gomez,  1980).  Haynes 
placed  our  material  in  variety  alpina,  which  rang- 
es from  southern  Mexico  to  Panama.  However, 
the  senior  author  has  seen  these  plants  in  a  num- 
ber of  areas  ranging  from  Chiapas  to  Panama  and 
believes  that  there  is  a  discontinuity  of  morphol- 
ogy in  the  Guatemala-Honduras  region.  Thus,  it 
is  possible  that  the  plants  of  southern  Central 
America  are  deserving  of  taxonomic  recognition 
as  C.  panamensis. 


166 


FIELDIANA:  BOTANY 


FIG.  28.     Orobanchaccac:  Conopholis  alpina.  A,  Calyx  in  lateral  view.  B,  Scpaloid  hractcolc.  C,  A  different  calyx 
in  side  view.  D,  Ventral  view  of  stamen.  E,  Lateral  view  of  stamen.  F,  Placcntation.  G,  Inflorescence  and  base. 


BURGER:  FLORA  COSTARICENSIS 


167 


Orobanche  Linnaeus 

Herbs  parasitic  on  roots,  lacking  chlorophyll, 
stems  simple  or  branched,  usually  succulent,  usu- 
ally covered  with  minute  gland-tipped  hairs. 
Leaves  reduced  to  alternate  sessile  scales,  sepa- 
rate along  the  stem,  yellowish  to  brown  or  purple, 
margins  entire.  Inflorescences  terminal  spikes  or 
racemes,  bracts  conspicuous  and  similar  to  the 
scales,  bracteoles  present  or  absent.  Flowers  with 
calyx  cupulate  to  campanulate,  2-5-lobed  or  di- 
vided to  the  base  adaxially  and  abaxially,  lobes 
equal  or  unequal;  corolla  tubular,  often  with  mi- 
nute gland-tipped  hairs,  2-lipped,  4-  or  5-lobed, 
upper  lip  entire  or  2-lobed;  stamens  4,  included, 
filaments  borne  near  the  base  of  the  tube,  anthers 
2-thecous,  thecae  parallel;  disc  not  apparent;  ova- 
ry 1-locular,  ovules  many  on  4  parietal  placentas, 
stigma  2-lobed.  Fruits  capsules,  oblong,  enclosed 
within  the  persisting  corolla;  seeds  ellipsoid, 
coarsely  reticulate. 

Orobanche  is  a  genus  of  about  150  species, 
mostly  north  temperate,  and  is  especially  speciose 
in  Europe.  The  following  species  has  become  a 
serious  agricultural  problem  in  some  parts  of  the 
world. 


Orobanche  minor  J.  E.  Smith,  English  Bot.  6: 
422,  tab.  422.  1797. 

Herbs  with  single  or  several  clustered  stems  8- 
80  cm  tall,  the  stems  rarely  with  distal  branches, 
4-10  mm  diam.,  pale  rose-yellow  to  brownish, 
surface  with  glandular  hairs  0.2-1  mm  long. 
Leaves  represented  by  alternate  simple  scales  1- 
3  cm  long,  sessile,  broadly  ovate,  acute,  venation 
parallel.  Inflorescences  spicate,  bracts  9-12  mm 
long,  ca.  4  mm  wide,  sessile,  difficult  to  distin- 
guish from  the  sepals  in  dried  material,  pedicels 
0-2(-5)  mm  long,  glandular  puberulent,  bracte- 
oles absent.  Flowers  with  calyx  usually  split  to 
the  base  abaxially  and  adaxially,  each  portion 
with  2  long  narrow  acuminate  lobes;  corolla  12- 
18  mm  long,  slightly  purplish  with  purple  veins 
but  usually  yellowish  at  the  base  and  lower  lip, 
sparsely  puberulent  externally,  tube  ca.  4  mm 
diam.;  filaments  ca.  6  and  7  mm  long. 

Orobanche  minor  is  not  native  to  the  Americas 
and  is  rarely  collected  in  the  American  tropics. 
The  senior  author  observed  this  species  near  Pru- 
sia,  at  1500  m  elevation  on  the  slopes  of  Vocan 
Irazu,  Cartago  province,  in  November  1979  (G6- 
mez  7140  CR,  usj),  where  it  was  parasitizing  a 


species  of  clover  (Trifolium)  and  two  species  of 
grasses.  This  adventive  population  probably  came 
from  introduced  grass  or  forage  (G6mez,  1980). 
These  plants  are  distinctive  because  of  their  lack 
of  green  pigmentation,  erect  unbranched  stems 
with  gland-tipped  hairs,  scale-like  leaves  similar 
to  the  bracts,  split  calyx,  and  tubular  purple  and 
yellowish  corollas.  Although  associated  with  a  va- 
riety of  herbs  and  small  shrubs,  this  species  is 
often  found  parasitizing  plants  of  the  family  Fa- 
baceae.  In  tropical  areas  they  are  unlikely  to  be 
found  below  1000  m  elevation. 


Literature  Cited 

BURGER,  W.,  AND  Q.  JIMENEZ.  1994.  A  new  species  of 
Psychotria  subgenus  Psychotria  (Rubiaceae)  from 
Costa  Rica.  Novon,  4:  206-208. 

CANNE,  J.  1980.  Seed  morphology  in  Aureolaria  and  70- 
manthera  (Scrophulariaceae)  (abstract).  Botanical  So- 
ciety of  America  Miscellaneous  Series  Publication 
158:  20. 

CRONQUIST,  A.  1981.  An  Integrated  System  of  Classifi- 
cation of  Flowering  Plants.  Columbia  Univ.  Press, 
New  York,  1262  pp. 

D'ARCY,  W.  G.  1979.  Scrophulariaceae,  in  Flora  of  Pan- 
ama. Annals  of  the  Missouri  Botanical  Garden,  66: 

173-272. 

GENTRY,  A.  H.  1973a.  Generic  delimitations  of  Central 
American  Bignoniaceae.  Brittonia,  25:  226-242. 

.  1973b  (1974).  Bignoniaceae,  in  Flora  of  Pana- 
ma. Annals  of  the  Missouri  Botanical  Garden,  60: 
781-977. 

.  1979.  Additional  generic  mergers  in  Bignoni- 
aceae. Annals  of  the  Missouri  Botanical  Garden,  66: 
778-787. 

.  1980.  Bignoniaceae — Part  I  (Crescentieae  and 

Tourettieae).  Flora  Neotropica,  25(1):  1-130. 

.  1983.  Dispersal  and  distribution  of  Bignoni- 
aceae. Sonderbd.  Naturwissenschaftlicher  Verein 
Hamburg,  7:  303-314. 

— .  1990.  Evolutionary  patterns  in  Neotropical  Big- 
noniaceae. Memoirs  of  the  New  York  Botanical  Gar- 
den, 55:  119-129. 

— .  1992.  Bignoniaceae — Part  II  (Tribe  Tecomeae). 
Flora  Neotropica,  25(11):  1-370. 

-.  1994.  Bignoniaceae.  Flora  of  Nicaragua  project. 


Missouri  Botanical  Garden  (unpublished  manuscript). 

GOMEZ,  L.  D.  1980.  Notes  on  the  biology  of  Central 
American  Orobanchaceae.  Brcnesia,  17:  389-396. 

MABBERLEY,  D.  J.  1987.  The  Plant  Book.  Cambridge 
Univ.  Press,  New  York,  706  pp. 

MORALES,  J.  F,  AND  Q.  JIMENEZ.  1997.  Bignoniaceae. 
Flora  of  Costa  Rica  project,  Missouri  Botanical  Gar- 
den (unpublished  manuscript). 

OLMSTEAD,  R.,  AND  P.  REEVES.  1995.  Evidence  for  the 
polyphyly  of  the  Scrophulariaceae  based  on  chloro- 


168 


FffiLDIANA:  BOTANY 


plast  rbcL  and  ndhF  sequences.  Annals  of  the  Mis- 
souri Botanical  Garden,  82:  176-191. 

REVEAL,  J.  L.  1995.  Newly  required  supragcneric  names 
in  vascular  plants.  Phytologia,  79:  68-76. 

MINOD,  M.  1918.  Contribution  a  I'ctudc  du  genre  Ste- 
modia  et  du  groupe  dcs  Stemodiees  in  Amcrique.  Bul- 
letin do  Socie'te  Botaniquc  Geneve,  send  II,  10:  155- 
252. 

STANDLEY,  P.  C.  1938.  Flora  of  Costa  Rica.  Publications 
of  the  Field  Museum  of  Natural  History,  Botanical 
Series,  18(3). 

STANDLEY,  P.  C.  AND  L.  O.  WILLIAMS.  1973.  Scrophular- 


iaccac,  in  Flora  of  Guatemala.  Ficldiana:  Botany,  24: 
9(4):  319-416. 

-.    1974.  Bignoniaccac,  in  Flora  of  Guatemala. 


Fieldiana:  Botany,  24(10):  153-232. 

SUTTON,  D..  AND  R.  HAMPSHIRE.  1995.  Scrophulariaccac. 
Flora  of  Nicaragua  project,  Missouri  Botanical  Garden 
(unpublished  manuscript). 

WAGSTAFF,  S.  J.,  AND  R.  G.  OI.MSTKAD.  1997.  Phylogeny 
of  Labiatae  and  Vcrbenaceae  inferred  from  r/>cL  se- 
quences. Systematic  Botany,  22:  165-179. 

WILLIAMS,  L.  O.  1970.  An  overlooked  genus  of  the  Scro- 
phulariaccac. Ficldiana:  Botany,  32:  211-214. 

— .  1972.  Scrophulariaccac,  in  Tropical  American 
plants,  XII.  Fieldiana:  Botany.  34(8):  118-132. 


BURGER:  FLORA  COSTARICENSIS 


169 


Index 

The  index  includes  all  accepted  names  (in  roman  type),  synonyms  (italics),  common  English  names 
(roman)  and  vernacular  Spanish  names  (italics).  The  text  includes  only  one  new  species  (boldface), 
Amphitecna  gentryi. 


Adenocalymma  100 
Adenocalymma  apurcnsc  101 
Adenocalymma  calderonii  100 
Adenocalymma  ci liola turn  \  33 
Adenocalymma  cocleense  160 
Adenocalymma  fissum  135,  136 
Adenocalymma  flos-ardeae  1 50 
Adenocalymma  heterophyllum  \  \  1 
Adenocalymma  hintonii  100 
Adenocalymma  hosmeca  133 
Adenocalymma  inundatum  100 
Adenocalymma  macrocarpum  133 
Adenocalymma  ocositensis  1 1 8 
Adenocalymma  perezii  135 
Adenocalymma  punctifolium  1 50 
Adenocalymma  seleri  135 
Adenocalymma  standleyanum  1 1 7 
Adenocalymma  veruciferum  135 
Adenocalymma  verrucosa  1 1 5 
African  tulip  tree  149 
Agalinis  5 
Agalinis  albida  5 
Agalinis  hispidula  17 
Agalinis  peduncularis  5 
ajillo  133 
ajonjoli  164 
Alectra  14 
Alectra  aspera  14 
Alectra  brasiliensis  14 
Alectra  fluminensis  14 
Alectra  melatnpyroides  14 
Alonsoa  15 

Alonsoa  meridionalis  15 
Alonsoa  warscewiczii  19 
Amphilophium  100 
Amphilophium  molle  101 
Amphilophium  oxylophium  102 
Amphilophium  paniculatum  101 

var.  molle  101 

Amphilophium  pannosum  102 
Amphilophium  pilosum  102 
Amphitecna  102 
Amphitecna  donnell-smithii  104 
Amphitecna  gentryi  103 
Amphitecna  haberi  106 
Amphitecna  isthmica  104 
Amphitecna  kennedyi  104 
Amphitecna  latifolia  105 
Amphitecna  molinae  104 
Amphitecna  obovata  105 
Amphitecna  sessilifolia  105 
Amphitecna  silvicola  104 
Anemopaegma  106 
Anemopaegma  chrysanthum  107, 

109 

Anemopaegma  chrysoleucum  107 
Anemopaegma  macrocarpa  107 
Anemopaegma  orbiculatum  107 


Anemopaegma  pubcrulum  108 
Anemopaegma  puncticulatum  1 07 
Anemopaegma  santaritense  109 
Anemopaegma  tobagense  117 
Anemopaegma  tonduzianum  121 
angelon  17 
Angelonia  16 
Angelonia  angustifolia  16 
Angelonia  ciliaris  16 
Anisantherina  5,  17 
Anisantherina  hispidula  17 
Anonymous  umbrosa  53 
Antirrhinum  18 
Antirrhinum  majus  18 
Antirrhinum  canadensis  45 
Antirrhinum  cymbalaria  36 
Antirrhinum  linaria  46 
Antirrhinum  vulgaris  117 
Antirrhinum  muralis  87 
Arrabidaea  109 
Arrabidaea  candicans  1 10 
Arrabidaea  chica  1 1 1 
Arrabidaea  conjugata  1 1 1 
Arrabidaea  corallina  1 12 
Arrabidaea  costaricensis  1 1 2 
Arrabidaea  dichasia  140 
Arrabidaea  e recta  1 12 
Arrabidaea  florida  1 1 3 
Arrabidaea  guatemalensis  121 
Arrabidaea  inaequalis  109 
Arrabidaea  isthmica  113,  121 
Arrabidaea  lundellii  1 14 
Arrabidaea  mollicoma  113 
Arrabidaea  mollissima  113 
Arrabidaea  pacy calyx  1 10 
Arrabidaea  panamensis  113 
Arrabidaea  pattelifera  1 14 
Arrabidaea  potosina  123 
Arrabidaea  pseudochica  121 
Arrabidaea  pubescens  1 1 5 
Arrabidaea  rhodothyrsus  \  10 
Arrabidaea  verrucosa  1 1 5 
Asarina  erubescens  49 
Asarina  scandens  50 


Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 
Bacopa 


18 

auriculata  21 
axillaris  20 
bacopoides  20 
bracteolata  20 
curtipes  23 
decumbens  21 
cgcnsis  21 
humilis  23 
lacertosa  21 
laxiflora  21 
limosa  22 
monnieri  22 


Bacopa  monnierioides  22 
Bacopa  myriophylloides  25 
Bacopa  naias  25 
Bacopa  pan'iflora  22 
Bacopa  procumbens  52 
Bacopa  ranaria  22 
Bacopa  reflexa  25 
Bacopa  rcpens  23 
Bacopa  salzmannii  23 
Bacopa  sessiliflora  24 
Bacopa  valerii  23 
Bacopa  violacea  23 
bateita  146 
bees,  Centris  17 
bejuco  de  ajo  135 
Benjaminia  24 
Benjaminia  reflexa  25 
Benjaminia  utriculariaeformis  24 
Bignonia  aeqinoctialis  121 
Bignonia  aesculifolia  125 
Bignonia  binata  118 
Bignonia  candicans  110 
Bignonia  capensis  157 
Bignonia  chica  \  1 1 
Bignonia  chrysantha  152 
Bignonia  chrysoleucum  107 
Bignonia  conjugata  111 
Bignonia  copaia  127 
Bignonia  corallina  1 1 2 
Bignonia  corymbifera  129 
Bignonia  crucigera  145 
Bignonia  dasyonyx  132 
Bignonia  diversifolia  122 
Bignonia  echinata  145 
Bignonia  glabrata  112 
Bignonia  hymenaea  133 
Bignonia  ignea  147 
Bignonia  inaequilatera  150 
Bignonia  kerere  133 
Bignonia  latifolia  \  16 
Bignonia  lepidota  122 
Bignonia  litoralis  113 
Bignonia  magnoliaefolia  124 
Bignonia  mollicoma  113 
Bignonia  mollissima  113 
Bignonia  obliqua  112 
Bignonia  obovata  136 
Bignonia  orbiculata  107 
Bignonia  paniculata  101 
Bignonia  pannosa  102 
Bignonia  patellifera  \  1 4 
Bignonia  pubescens  \  14 
Bignonia  pyramidata  140 
Bignonia  quadrivalvis  137 
Bignonia  riparia  150 
Bignonia  sarmentosa  121 
Bignonia  sinclairii  140 
Bignonia  stans  157 


170 


FIELDIANA:  BOTANY 


Bignonia  tetragonoloba  \  \  1 
Bignonia  tiliaefolia  145 
Bignonia  umbrosa  129 
Bignonia  uncata  131 
Bignonia  uncinata  131 
Bignonia  ungiiis-cati  1 3 1 
Bignonia  variabilis  146 
Bignonia  venusta  147 
Bignonia  verrucifera  135 
Bignoniaccac  77 

list  of  species  161 
boca  de  dragon  18 
boca  de  la  vieja  17 
boca  de  ledn  1 8 
bombilla  147 
baton  de  oro  28 
Bourreria  costaricensis  75 
Buchnera  25 
Buchnera  grandiflora  97 
Buchnera  leiantha  26 
Buchnera  longifolia  26 
Buchnera  major  26 
Buchnera  mexicana  26 
Buchnera  palustris  27 
Buchncra  pusilla  26 
Buchnera  tinctoria  26 
Buchnera  weberbaueri  26 

Cabomba  25 
Caconapea  auriculata  21 
Caconapea  axillaris  34 
Caconapea  conferta  24 
Caconapea  paniflora  22 
Caconapea  sessiliflora  24 
calabacero  106,  120 
calabash  106,  120 
calabash,  swamp  105 
calabasillo  de  playa  1 05 
Calceolaria  27 
Calceolaria  irazucnsis  28 
Calceolaria  costaricensis  28 
Calceolaria  heterophylla  30 
Calceolaria  mexicana  28 
Calceolaria  microbefaria  29 
Calceolaria  perfoliata  30 
Calceolaria  sciadephora  30 
Calceolaria  storkii  29 
Calceolaria  trachelifolia  28 
Calceolaria  trilobata  30 
Calceolaria  tripartita  30 
Calceolaria  urticina  28 
Callichlamys  116 
Callichlamys  garneri  1 16 
Callichlamys  latifolia  1 16 
candelillo  158 
carhoncillo  158 
Capraria  31 
Capraria  bi  flora  3 1 

var.  pilosa  3 1 
Capraria  Crustacea  47 
Capraria  durantifolia  61 
Capraria  gratioloides  48 
Carpoceras  angulata  164 
Castilleja  32 
Castilleja  agrestis  33 
Castilleja  arvensis  33 


Castilleja  aurantiaca  34 
Castilleja  hicolor  34 
Castilleja  chiriquiensis  34 
Castilleja  communis  33 
Castilleja  irasuensis  33 
Castilleja  lentil  34 
Castilleja  quirosii  34 
Caxtilleja  seibertii  34 
Castilleja  talamancensis  35 
Castilleja  tayloriorum  35 
Ccratophy  turn  1 1 7 
Ceratophytum  tciragonolobum  117 
Ceratophytunt  tobagense  I  1 7 
Chelone  gentianoides  54 
Chodanthus  puberulum  108 
chorro  148 
chorro  de  oro  148 
Clerodendrwn  epiphyticum  72 
Clytostoma  1 1 7 
Clytostoma  binatum  1 1 8 
Clytostoma  callistegioidcs  1 1 8 
C  'lyiostoma  islhmicum  1 1 8 
Clytostoma  mayanum  123 
Coliseum  Ivy  88 
coloring  agents,  dyes  39 
Conobea  alata  43 
Conobea  pusilla  57 
Conopholis  165 
Conopholis  alpina  166 

var.  mexicana  166 
Conopholis  americana  166 
Conopholis  mexicana  166 
Conopholis  panamensis  1 66 
Conopholis  sylvatica  166 
coral  56 
coralillo  56 
corrimiento  62 
cones  152 

carles  amarilla  153,  154 
coneza  153,  154 
cortez  negro  154 
Couralia  rosea  155 
cow  okra  141 
Crcscentia  1 19 
Crescentia  aculeata  141 
Crescentia  alata  1 19 
Crcscentia  cujcte  120 
Crescentia  edulis  141 
Crescentia  latifolia  105 
Crescentia  ohovata  105 
Crescentia  pinnata  1 28 
Crcscentia  ternata  119 
cucharilla  146 
culantrillo  58 
(\bistax  macrocarpa  125 
Cydista  120 
Cydista  aequinoctalis  1 2 1 

var.  hirtella  121 
Cydista  divcrsifolia  122 
Cydista  heterophylla  122 
Cydista  lilacina  123 
Cydista  potosina  123 
Cvdista  puhescens  1 2 1 
Cvdista  sarmentosa  121 
Cymbalaria  36 


Cymbularia  muralis  36 

Daixsa  36 

Darcya  costaricensis  37 
Darcya  reliquiarum  37 
Dtndrosicos  i\ihiuicn\  l()4 
Dfiulrosicos  kennedyi  104 
Dendrosicos  latijoliu\  105 
Dermatocalyx  par\-ijioms  76 
i/i.U/M/  39 
Digitalis  38 
Digitalis  purpurea  38 
Distictella  124 
Distictclla  inagnoliilblia  124 
Distictis  laxiflora  101 
Distictis  rovirasana  160 
Disteira  angulo\a  164 
Dodartieae  100 
domatia  196 
Dombeya  lappacea  158 
Doxantha  dasyonyx  132 
Doxantha  unguis-cati  132 

Enallagma  latifolia  105 
Enallagma  sexsil (folia  1 05 
Erinus  procumbens  52 
Erinus  verticillatus  62 
escoba  amarga  58 
Escobedia  39 
Escobedia  curialis  39 
Escobedia  grandiflora  39 
Escobedia  lacvis  39 
Escobedia  long/flora  39 
Escobedia  reticulata  39 
Escobedia  scabrifolia  39 
escoheta  58 
escobilla  amarga  58 

Fagelia  perfoliata  30 
figure  scales  5 
y/or  </<'  //u/fV>  35 
flowering  phenology  120 
foxglove  39 

gallinazo  128 
gallito  34 
gallitos  28 
Gentry,  A.  H.  v.  78 
Gerardia  5. 
Gerardia  hispidula  17 
Gibsoniothamnus  70 
Gibsoniolhamnus  alatus  74 
Gibsoniothamnus  cornutus  72 
Gibsoniothamnus  cpiphyticus  72 
Gibsoniothamnus  grandiflorus  73 
Gibsoniothamnus  mirilicus  73 
Gibsoniothamnus  parvifolius  73 
Gibsoniothamnus  pithccobius  72 
Gibsoniothamnus  ptcrocalyx  73 
Gibsoniothamnus  stcllatus  74 
Globifcra  umbrostim  53 
Glossotylis  aspcra  14 
GcxJmania  125 
Gcxlmannia  acsculifolia  125 
Godmannia  macrocarpa  1 25 
Gratiola  anagallidea  48 


INDEX 


171 


Gratiola  dubia  48 
Gratiola  inaequalis  48 
Gratiola  repens  23 
guacales  120 

Hebe  66 
Hcmichacna  40 
Hemichaena  fruticosa  40 
Hemimcridae  18 
Herpestis  auriculata  2 1 
Herpestis  axillaris  34 
Herpestis  bacopoides  20 
Herpestis  caparioides  52 
Herpestis  ciliata  23 
Herpestis  laxiflora  21 
Herpestis  peduncularis  52 
Herpestis  ranaria  22 
Herpestis  reflexa  24 
Herpestis  repens  23 
Herpestis  salzmannii  23 
Herpestis  sessiliftora  24 
hierba  santa  62 
Hydranthelium  egense  21 

Ilysanthes  dubia  48 
Ilysanthes  gratioloides  48 

jacaranda,  jacaranda  \  28 
Jacaranda  126 
Jacaranda  caucana  127 

ssp.  sandwithiana  127 
Jacaranda  copaia  127 

ssp.  spectabilis  127 
Jacaranda  ficifolia  127 
Jacaranda  mimosifolia  128 
Jacaranda  ovalifolia  128 
Jacaranda  spectabilis  127 
Jacaranda  superba  127 
Jacaranda  trianae  127 
jfcaras  120 
jicarita  105 
jf'/caro  106,  119,  120 
jicaro  danto  143 
jicaro  de  playa  105 
Jovellana  60 
>//a  157 

Kenilworth  Ivy  88 
Kigelia  128 
Kigelia  africana  128 
Kigelia  pinnata  128 

Lamourouxia  41 
Lamourouxia  gutierrezii  42 
Lamourouxia  lanceolata  42 
Lamourouxia  longiflora  42 
Lamourouxia  longifolia 

var.  lanceolata  42 
Lamourouxia  rhinanthifolia  43 
Lamourouxia  smithii  43 
Lamourouxia  veijensis  42 
Lamourouxia  viscosa  42 
Lamourouxia  scabra  42 
Leucocarpus  43 
Leucocarpus  alatus  43 
Leucocarpus  fruiticosus  40 


Leucocarpus  pcrfoliatus  43 

Levya  nicaraguensis  121 

Limnophila  costaricensis  25 
forma  aquatica  25 
forma  semiterrestris  25 

Limosclla  44 

Limosclla  acaulis  44 

Limosella  americana  44 

Limosella  australis  45 

Linaria  45 

Linaria  canadensis  45 

Linaria  texana  45 

Linaria  vulgaris  46 

Linaria  cymbalaria  36 

///M/fl  393 

Lindernia  46 

Lindernia  anagallidea  48 

Lindernia  Crustacea  47 

Lindernia  dianthera  52 

Lindernia  diffusa  47 

Lindernia  dubia  48 

Lindernia  gratioloides  48 

Lindernia  inaequalis  48 

Lindernia  japonica  5 1 

Lindernia  microcalyx  48 

Lindernia  thouarsii  65 

Lindernia  verticillata  62 

llama  del  bosque  149 

lluvia  de  coral  56 

Lobelia  pumila  5 1 

Lophospermum  48 

Lophospermum  erubescens  49 

Lundia  129 

Lundia  colombiana  130 

Lundia  corymbifera  129 

Lundia  dicheilocalyx  130 

Lundia  puberula  130 

Lundia  schumanniana  130 

Lundia  valenzuelae  129 

Lysimachia  monnieri  22 

Macfadyena  130 
Macfadyena  guatemalensis  1 3 1 
Macfadyena  phellosperma  144 
Macfadyena  uncata  131 
Macfadyena  uncinata  131 
Macfadyena  unguis-cati  131 
Macuillamia  limosa  23 
Macuillamia  repens  23 
Manga  de  la  Senora  39 
Mansoa  132 
Mansoa  hymenaea  133 
Mansoa  kerere  133 

var.  erythraea  134 

var.  incarnata  134 
Mansoa  parvifolia  134 
Mansoa  standleyi  135 
Mansoa  verrucifera  135 
mariposas  146 
Marline!  la  136 
Martinella  obovata  136 
Martinella  verrucosa  1 1 5 
Martyniaceae  164 
Martynia  164 
Martynia  angulosa  164 
Martynia  annua  164 


Martynia  diandra  164 
Maurandya  49 
Maurandya  barclaiana  50 
Maurandya  erubescens  49 
Maurandya  scandens  50 
Maurandya  semperflorens  50 
Mazus  51 

Mazus  japonicus  51 
Mazus  pumilus  51 
Mazus  rugosus  51 
Mecardonia  51 

Mecardonia  montevidensis  52 
Mecardonia  procumbens  52 
medicinal  properties  32,  39,  151 
Melasma  melampyroides  14 
Mella  laxiflora  21 
Melloa  137 

Melloa  quadrivalvis  137 
Micalia  grandiflora  97 
Micranthemum  52 
Micranthemum  pilosum  53 
Micranthemum  standleyi  53 
Micranthemum  umbrosum  53 
Mimulus  53 
Mimulus  glabratus  53 
Mimulus  perfoliatus  43 
mi'rame  Undo  147 
monkey  flowers  53 
Monniera  axillaris  34 
Monniera  reflexa  50 
Monocardia  humilis  23 
Monocardia  violacea  23 
morro  1 1 9 
Mussatia  138 
Mussatia  hyacinthina  138 
Myoporaceae  31 

Naiadothrix  longipes  25 
Neotuerckheimia  gonoclada  105 
Nortenia  thouarsii  65 

Onohualcoa  138 
Onohualcoa  fissa  135 
Onohualcoa  seleri  135 
Onohualcoa  verrucifera  135,  136 
Orobanchaceae  5,  165 
Orobanche  168 
Orobanche  minor  168 

Pachyptera  132,  138 
Pachyptera  kerere  134 
Pachytera  parvifolia  \  34 
Pachyptera  standleyi  135 
palomitas  146 
Pandorea  146 
Pandorea  ricasoliana  147 
Parabignonia  138 
Parabignonia  steyermarkii  139 
Paradolichandra  139 
Paragonia  139 
Paragonia  pyramidata  140 
Parmentiera  140 
Parmentiera  aculeata  141 
Parmentiera  alata  1 19 
Parmentiera  cerifera  141 
Parmentiera  dressleri  142 


172 


FIELDIANA:  BOTANY 


Parmentiera  edulis  141 
Parmentiera  macrophylla  142 
Parmentiera  trunciflora  143 
Parmentiera  valerii  143 
Pedaliaceae  162 
Pedicularis  meUonpyroides  14 
pedo  de  padre  \  33 
peina  de  mico  146 
peina  de  mono  146 
Peloria  118 
Pennywort  88 
Penstemon  54 
Penstcmon  gentianoidcs  54 
Penstemon  skutchii  54 
pepino  de  danto  143 
Petastoma  breviflorum  1 14 
Petastoma  ocositense  1 1 8 
Petastoma  patellifera  1 14 
Petastoma  pubescens  1 14 
Petastoma  tonduzianum  133 
Phryganocydia  143 
Phryganocydia  corymbosa  144 
Phryganocydia  phellospcrma  144 
Pithecoctenium  145 
Pithecoctenium  crucigerum  145 
Pithecoctenium  echinatum  145 
Pithecoctenium  panamensis  107 
Pleonotoma  146 
Pleonotoma  diversifolia  122 
Pleonotoma  variabilis  146 
Podranea  146 
Podranea  ricolasiana  147 
pollinators  23 
Porodittia  60 
porto  hello  17 
Proboscidia  triloba  164 
Pseudocalymma  alliaceum 

var.  macrocalyx  135 
Pseudocalymma  sagotii  133 

var.  macrocalyx  135 
Pseudocalymma  standleyi  135 
Pyrostegia  147 
Pyrostegia  ignea  147 
Pyrostegia  venusta  147 
Pyxidaria  Crustacea  47 
Pyxidaria  diffusa  47 

quacalillo  106 

Quinquelobulus  utriculariaeoides 
25 

Ranaria  monnierioides  22 
Reichardia  scandens  50 
Rhodochiton  49 
roble  155 

roble  de  bianco  1 55 
roble  de  sabana  155 
roble  macho  154 
Russelia  55 

Russelia  colombiana  56 
Russelia  equisetiformis  55 
Russelia  flavoviridis  56 
Russelia  juncea  55 
Russelia  oxyphylla  56 
Russelia  sarmentosa  56 
Russelia  tabacensis  56 


Saldanhaea  costaricensis  1 1 2 
Saldanhaea  seemanniana  160 
San  Carlos  148 
Saritea  148 
Saritca  magnifica  148 
Schistophragma  56 
Schistophragma  mcxicana  57 
Schistophragma  pusilla  57 
Schlcgcliaceae  69 
Schlegclia  74 
Schlegclia  brachyantha  75 
Schlegelia  costaricensis  75 
Schlegclia  fastigiata  75 
Schlegelia  fuscata  76 
Schlegclia  nicaraguacnsis  76 
Schlegelia  parasitica  76 
Schlegclia  parviflora  76 
Schlegelia  silvicola  76 
Schlegclia  sulfurea  76 
Scobinaria  148 
Scobinaria  japurensis  115 
Scobinaria  verrucosa  \  \  5 
Scoparia  57 
Scoparia  annua  58 
Scoparia  dulcis  58 
Scrophularia  fluminensis  14 
Scrophularia  meridionalis  15 
Scrophularia  procumbens  23 
Scrophulariaceae  1,  2 

list  of  species  69 
sesame  162 
Sesamum  162 
Sesamum  indicum  162 
Sesamum  orientalc  162 
Sibthorpia  58 
Sibthorpia  pichinensis  59 
Sibthorpia  rcpens  59 
Sibthorpia  triandra  59 
slipper  flowers  27 
snapdragon  18 
Sparattosperma  rosea  155 
Spathodea  148 
Spathodea  campanulata  149 
Spathodea  corymbosa  144 
Spathodea  laurifolia  144 
Spathodea  nilotica  148 
Spathodea  obovata  136 
Spathodea  orinocensis  144 
Stemodia  59 
Stemodia  angulata  60 
Stemodia  arenaria  62 
Stemodia  costaricensis  37 
Stemodia  durantifolia  61 
Stemodia  jorullcnsis  61 
Stemodia  jorullensis 

ssp.  re  plans  60 
Stemodia  pan-ijiora  62 
Stemodia  pcduncularis  61 
Stemodia  reliquiarum  37 
Stemodia  siliquosa  57 
Stemodia  verticillata  62 
Stemodiacra  angulata  60 
Stemodiacra  peduncularis  6 1 
Stizophyllum  149 
Stizophyllum  flos-ardeae  150 
Stizophyllum  inacquilatcrum  150 


Stizophyllum  punctifolium  150 
Stizophyllum  riparium  150 
swamp  calabash  105 
sweet  broom  58 

Tabebuia  151 
Tabehuia  calderonii  100 
Tabebuia  chrysantha  152.  154 
ssp.  chrysantha  152 
ssp.  pluvicola  152.  153 
Tabebuia  dugandii  153 
Tabebuia  guayacan  153 
Tabebuia  impctiginosa  153 
Tabebuia  latifolia  1 1 6 
Tahebuia  me.\icana  155 
Tabebuia  neochrysantha  154 
Tabebuia  nicaraguensis  153 
Tabebuia  ochracca  154 

ssp.  neochrysantha  154 
Tabebuia  palmeri  153 
Tabebuia  palustris  154 
Tabebuia  pentaphylla  1 55 
Tahebuia  pyramidata  140 
Tabebuia  rosea  155 
Tabebuia  scrratifolia  153 
Tabebuia  sessilifolia  105 
Tabebuia  sinclairii  139 
Tabebuia  speciosa  1  16 
Tanaecium  156 
Tanaecium  albiflora  156 
Tanaecium  jaroba  1 56 
Tanaecium  zetekii  133 
Tccoma  156 
Tccoma  capensis  157 
Tecoma  chrysantha  152 
Tecoma  evenia  152,  155 
Tecoma  fuscata  1 25 
Tecoma  guayacan  153 
Tecoma  impetiginosa  153 
Tecoma  mexicana  155 
Tecoma  ochracea  154 
Tecoma  palmeri  1 52,  4 1 2 
Tecoma  ricasoliana  147 
Tecoma  rosea  155 
Tecoma  stans  157 
Tecomaria  156 
Tecomaria  capensis  157 
Tclrancma  62 
Tctranema  floribundum  63 
Tctrancma  gamboanum  63 
Tctranema  mcgaphyllum  63 
Tctrancma  roscum  172 
Torcnia  64 
Torcnia  asiatica  65 
Torenia  Crustacea  47 
Torcnia  fournicri  64 
Torcnia  thouarsii  65 
Tourrcttia  158 
Tourrcttia  lappacca  158 
Tourrettia  volubilis  158 
triquitraque  148 
tulipdn  149 
Tynanthus  159 
Tynanthus  guatcmalcnsis  159 
Tynanthus  hyacinthinus  I3H 
Tynanlhus  macranthus  159 


INDEX 


173 


una  de  gato  \  32  Veronica  65                                                Veronica  serpyllifolia  68 

unas  de  diablo  165  Veronica  arvensis  66                                     van  humifusa  68 

Uroskinnera  63  Veronica  crenulata  68                               Veronica  tenella  68 

Usteria  scandens  50  Veronica  didyma  68                                  Veronica  xalapensis  67 

Utricularia  25  Veronica  humifusa  68 

Veronica  pcregrina  67 

.   .„     .  co  t           •   en                                    Wil  ichia  repens  59 

vainillo  1 58  van  xalapensis  67 

Vandelia  Crustacea  47  Veronica  persica  67 

Vandelia  diffusa  47  Veronica  polita  68                                     Xylophragma  160 

Vatkea  diandra  164  Veronica  rotundifolia  67                           Xylophragma  seemannianum  160 


174  FIELDIANA:  BOTANY 


UNIVERSITY  OF  ILLINOIS-URBANA