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ISSN  0968-0446 


Bulletin  of 

The  Natural  History 

Museum 


THE  NATURAL 
HISTORY  MUSEUM 


PRESENTED 
GENERAL  LIBRARY 


Botany  Series 


THE 

NATURAL 
HISTORY 
MUSEUM 


VOLUME  28         NUMBER  1         25  JUNE  1998 


The  Bulletin  of  The  Natural  History  Museum  (formerly:  Bulletin  of  the  British  Museum 
(Natural  History)  ),  instituted  in  1949,  is  issued  in  four  scientific  series,  Botany, 
Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology. 

*vl    *       * 

The  Botany  Series  is  edited  in  the  Museum's  Department  of  Botany 
Keeper  of  Botany:  Dr  S.  Blackmore 

Editor  of  Bulletin:  Ms  M.J.  Short 


•'•" 
Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and  ever- 

growing collections  of  the  Museum,  both  by  the  scientific  staff  and  by  specialists  from  elsewhere 
who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are  works  of  reference  that  will  remain 
indispensable  for  years  to  come.  All  papers  submitted  for  publication  are  subjected  to  external  peer  review  for 
acceptance. 

A  volume  contains  about  160  pages,  made  up  by  two  numbers,  published  in  the  Spring  and  Autumn. 
Subscriptions  may  be  placed  for  one  or  more  of  the  series  on  an  annual  basis.  Individual  numbers  and  back 
numbers  can  be  purchased  and  a  Bulletin  catalogue,  by  series,  is  available.  Orders  and  enquiries  should  be 
sent  to: 

Intercept  Ltd. 

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months  for  the  UK,  and  9  months  for  the  rest  of  the  world. 


World  List  abbreviation:  Bull.  nat.  Hist.  Mus.  Lond.  (Bot.) 
©  The  Natural  History  Museum,  1998 


Botany  Series 
ISSN  0968-0446  Vol.  28,  No.  1,  pp.  1-66 

The  Natural  History  Museum 

Cromwell  Road 

London  SW7  5BD  Issued  25  June  1998 

Typeset  by  Ann  Buchan  (Typesetters),  Middlesex 

Printed  in  Great  Britain  by  Henry  Ling  Ltd.,  at  the  Dorset  Press,  Dorchester,  Dorset 


Bull.  nut.  Hist.  Mus.  Land.  (Bot.)  28(1):  1-16 


Issued  25  June  1998 


Morphology  and  ecology  of  seedlings,  fruits 
and  seeds  of  Panama:  Vochysiaceae 


THE  NATURAL 
HISTORY 


NANCY  C.  GARWOOD     ^ 

Department  of  Botany,  The  Natural  History  Museum,  Cromwell  Road,  London  SW7  5BD 


CONTENTS 


18  JUN  1998 

pnr^cu*rgn> 

GENERAL  LIBRARY 


Introduction 

Methods 

Vochysiaceae 

Vochysia  ferruginea  Mart 

Vochysia  guatemalensis  Donn.  Sm 

Vochysia  sp 

Key  to  known  seedlings  of  Vochysia  in  Panama  and  Central  America  . 
Discussion  ... 


Fruit 
Seed 
Embryo 


Seedling  .................................................................................................................................................................................  10 

Phylogeny  ..............................................................................................................................................................................  12 

Ecology  ..................................................................................................................................................................................  14 

References  ...  ...  15 


SYNOPSIS  Seedlings,  saplings,  fruits  and  seeds  of  Vochysia  ferruginea  (Vochysiaceae)  are  described,  and  ecological  data  on 
these  stages  summarized,  as  part  of  the  ongoing  Seedling  Flora  Project  centred  on  Barro  Colorado  Island,  Panama.  Seedlings  of 
two  other  species  reported  from  Panama  are  also  described.  The  morphology  of  these  stages  is  compared  to  that  of  other  taxa  in 
the  family.  The  potential  use  of  seed  and  seedling  characters  in  phylogenetic  studies  within  the  family  is  discussed.  Special 
attention  is  drawn  to  unusual  two-parted  cotyledonary  stipules,  ontogenetic  changes  in  leaf  hair  morphology,  and  complex  ptyxis 
and  vernation  of  the  embryonic  cotyledons. 


INTRODUCTION 


The  aim  of  the  Seedling  Flora  Project,  initiated  in  1985,  is  to 
produce  an  illustrated  flora  of  seedlings,  saplings,  fruits  and  seeds 
for  the  neotropical  forest  centred  on  Barro  Colorado  Island,  Panama 
(see  Garwood  &  Humphries,  1993;Garwood,  1994,  1995,  in  prep.). 
This  is  the  account  of  Vochysia  ferruginea  Mart.  (Vochysiaceae).  It 
is  the  most  abundant  Vochysia  in  Panama,  common  in  the  younger 
forests  of  Barro  Colorado  Island  and  Central  Panama;  four  other 
species  are  known,  but  only  from  a  few  collections  (Robyns,  1967a, 
b\  Croat,  1978;  D'Arcy,  1987).  Ranging  from  Central  America 
through  the  Amazon  basin,  V.  ferruginea  is  also  the  most  widespread 
species  in  the  genus,  in  contrast  to  most  species  which  have  rela- 
tively limited  distributions  (Stafleu,  1948).  Because  of  its  fast 
growth  rate  and  valuable  timber,  V.  ferruginea  has  been  identified  as 
a  potentially  important  forestry  species  for  use  in  plantations  or 
managed  secondary  forests  in  the  neotropics  (Finegan,  1992;  Condit 
et  al.,  1993;  Butterfield  &  Fisher,  1994). 

In  this  account,  I  describe  the  seedlings,  saplings,  fruits  and  seeds 
of  V.  ferruginea,  and  the  seedlings  of  two  other  species  known  from 
Panama,  then  compare  these  stages  to  those  of  other  species  in  the 
family  to  identify  characters  of  potential  phylogenetic  importance. 
Fruit  and  seed  characters  have  been  important  in  delimiting  the 

©  The  Natural  History  Museum,  1998 


tribes  and  genera  of  Vochysiaceae;  subgeneric  classifications  and 
keys  to  species,  however,  have  been  based  almost  solely  on  floral 
and  vegetative  characters  (Warming,  1875;  Stafleu,  1948-1954; 
Keay  &  Stafleu,  1953).  There  have  been  few  comparative  studies  of 
the  gross  morphology  or  anatomy  of  fruits  and  seeds  (Corner,  1976; 
Boesewinkel  &  Venturelli,  1987).  Seedlings  of  few  species  have 
been  described  in  detail  (delAmo,  1979;  Ricardi  et  al.,  1987;  Flores, 
19930,  b). 


METHODS 

Standard  methods  for  germinating  seeds,  producing  voucher  speci- 
mens for  each  seed  collection,  growing  and  harvesting  seedlings, 
collecting  unvouchered  seedlings  (wildlings)  from  the  forest,  and 
manipulating  the  descriptive  database  were  developed  for  the  Seed- 
ling Flora  Project  (Garwood,  1994,  1995)  and  were  used  for  this 
account.  The  detailed  descriptions  follow  the  standard  format  devel- 
oped for  the  Project.  Descriptive  terms  have  been  standardized  for 
use  across  all  families  in  the  Project,  but  generally  follow  common 
use.  See  de  Vogel  (1980)  for  definitions  of  seedling-specific  terms 
and  Corner  (1976)  and  Boesewinkel  &  Bouman  (1984)  for  seed- 
specific  terms.  To  facilitate  identifying  isolated  seeds,  the  base  of  the 


N.C.  GARWOOD 


seed  is  arbitrarily  defined  as  the  part  nearest  the  hilum,  rather  than 
using  a  reference  point  within  the  fruit.  Leaf  nodes  of  seedlings  are 
numbered  sequentially  from  the  first  leaf-bearing  node  above  the 
cotyledons.  The  following  less  conventional  terms  are  used:  for 
length  of  pubescence  or  size  of  other  surface  features  -  minute  (<  0. 1 
mm),  short  (0. 1  -0.5  mm),  long  (0.5- 1 .0  mm),  very  long  (>  1 .0  mm); 
for  relative  thickness  of  the  seed  coat  or  pericarp  layers  compared  to 
total  width  of  the  structure  -  very  thick  (>  25%  width),  thick  (10- 
25%),  thin  (1-10%),  and  very  thin  (<  1%). 


VOCHYSIACEAE 

HABIT  AND  DISTRIBUTION.  Trees  or  shrubs,  rarely  subshrubs. 
Neotropical  except  for  one  very  small  African  genus;  6-8  genera  and 
c.  200  species,  3  genera  and  8  species  from  Panama,  1  genus  and 
species  known  from  Barro  Colorado  Island. 

TAXONOMIC  REFERENCES.  Stafleu  ( 1 948),  Keay  &  Stafleu  ( 1 953), 
Robyns  (1967a,  b),  Cronquist  (1981),  D'Arcy  (1987),  Kawasaki  (in 
press). 

Vochysia  Aubl. 

HABIT  AND  DISTRIBUTION.  Trees,  shrubs  or  rarely  subshrubs. 
Neotropical;  c.  100  species,  5  species  in  Panama,  1  species  known 
from  Barro  Colorado  Island. 

Vochysia  ferruginea  Mart,  in  Mart.  &  Zucc.,  Nov.  Gen.  sp.  pi.  1: 

151,1.92(1824). 
Fig.  1. 

HABIT  AND  DISTRIBUTION.  Trees,  6-35  m  tall,  of  moist  to  wet 
lowland  and  premontane  forests,  usually  in  secondary  forest.  Nica- 
ragua to  Peru  and  Brazil. 

COLLECTIONS.  Panama.  Barro  Colorado  Island:  Garwood  1 630A 
(F,  BM,  PMA,  adult  vouchers),  Garwood  1630B-F  (seedlings), 
Garwood  2005  (seedlings),  Foster  s.n.  (seedlings  with  fruit  voucher); 
50  seedlings  to  37  cm  tall  examined. 

Fruits 

Infructescences  terminal  and  upper-axillary  thyrses,  of  1-  to  5-flow- 
ered  helicoid  cymose  branches,  usually  with  <  1  fruit  maturing  per 
branch.  Rachis  densely  short-pubescent,  with  hairs  2-branched,  ± 
sessile  to  short-stalked,  appressed  to  ascending,  straight  to  sinuous; 
lateral  branches  (including  pedicels)  8-15  mm  long,  slightly  to 
moderately  pubescent. Receptacles  undeveloped.  Stipes  absent.  Fruits 
dry  capsules,  from  superior  ovaries.  Capsules  ±  woody,  loculicidally 
dehiscent;  completely  septate;  17-26  x  8-12  x  8-12  mm;  straight; 
oblong  to  obovate  in  outline,  3-lobed  to  3-cleft  in  t.s.,  not  com- 
pressed; margins  entire;  base  rounded  to  abruptly  truncate;  apex 
rounded  to  slightly  retuse,  with  stylar  scar  to  1  mm  wide;  surface 
dark  brown  to  black-brown,  dull,  slightly  to  prominently  warty, 
often  irregularly  clear-  to  white-scaly,  glabrous.  Seeds  1-3,  1  per 


locule,  intermediary,  separated  by  mesocarp  and  endocarp,  com- 
pressed laterally;  long  axis  of  seed  radially  parallel  to  long  axis  of 
fruit.  Placentae  axile.  Funicle  absent.  Valves  3,  dehiscing  from  apex, 
slightly  spreading  apart  then  each  folding  back,  remaining  firmly 
attached  at  base.  Fruit  wall  thick,  0.4-1.5  mm  wide,  thickest  cen- 
trally, 3-layered.  Exocarp  very  thin,  =  0.05  mm  wide,  ±  hard-papery, 
dark  brown  to  black-brown.  Mesocarp  thick,  0.2-1.3  mm  wide, 
thickest  centrally  between  locules,  hard-spongy  to  ±  woody,  with  2 
large  longitudinal  resin-filled  canals  near  axis  between  each  pair  of 
locules  and  several  smaller  chambers  along  locules  (resin  drying 
red-brown  and  glassy),  light  to  medium  brown.  Endocarp  thin,  ~  0. 1 
mm  wide,  crustaceous,  cream  to  light  brown;  inner  surface  smooth, 
minutely  transversely  fibrous,  glabrous,  glossy,  medium 
brown-orange.  When  fruit  unripe,  exocarp  grey-green. 

Seeds 

Seeds  20-25  x  5-6  x  1-2  mm;  hilar-anatropous;  1 -winged;  trans- 
versely half-obovate  in  outline,  compressed  and  elliptic  in  t.s.  in 
seminiferous  area,  planar  in  wing;  margins  constricted  distally  in 
seminiferous  area  on  hilar  side,  constricted  below  wing  on  opposite 
side;  base  widely  truncate;  apex  widely  rounded;  outer  surface  ± 
smooth  to  slightly  rough,  minutely  rough  under  hairs  at  higher 
magnification,  densely  long-  to  very  long-pubescent,  with  hairs 
sinuous,  appressed,  light  to  medium  brown-orange;  glossy,  medium 
brown  to  brown-orange,  dark  brown  under  hairs;  not  exfoliating. 
Wing  lateral,  fibrous,  ~  15-17  x  5-6  mm,  ±  half-elliptic  in  outline, 
planar,  asymmetrically  rounded  at  apex,  entire  along  margins,  with 
curved  ridge  slightly  raised,  ±  flattened,  extending  from  base  to  3/4 
length  of  wing  and  forming  thickened  straight  margin;  formed  from 
very  long,  interwoven,  testal  hairs.  Seminiferous  area  lateral,  ~  10- 
1 1  x  4-5  mm,  ±  elliptic  in  outline,  compressed,  darker  than  wings, 
with  short  fringe  of  interwoven  hairs  on  side  opposite  wing.  Aril 
absent.  Sarcotesta  absent.  Hilum  basal,  along  straight  side,  exposed, 
flush,  light  brown,  lighter  than  testa,  linear,  7-10  mm  long.  Micro- 
pyle  subbasal  to  marginal  (visible  internally  at  tip  of  radicle,  possibly 
continuing  through  thickened  ridge  of  wing  and  terminating  along 
straight  side  past  hilum),  concealed  externally.  Lens  absent.  Storage 
reserves  absent.  Embryo  large,  =  10-11  mm  long,  filling  seminifer- 
ous area,  straight,  ±  oblong  or  ellipsoid,  compressed;  axis 
well-developed,  with  long  4-winged  hypocotyl  separated  from  very 
short  radicle  by  annular  ridge  at  collet,  half-exposed;  cotyledons 
well-developed,  thin,  with  compressed  obliquely  supervolute  ptyxis 
and  convolute  (sensu  imbricate)  vernation,  with  wider  side  of  each 
cotyledon  angled  down  and  partially  enclosing  hypocotyl,  with 
shape  obscured  by  folding  and  base  hidden  within  folded  blade; 
plumule  rudimentary.  Vascular  bundle  half-lateral  on  hilar  side, 
unbranched,  thin,  with  tip  exposed  at  end  of  hilum  and  fragments  of 
the  funicular  bundle  often  exposed  in  hilum;  passing  through  short 
raphe,  terminating  marginally  at  tip  of  seminiferous  area.  Seed  coat 
very  thin,  ±  papery,  undifferentiated  at  maturity.  Testa  very  thin,  ± 
thicker  under  hilum  and  around  micropyle,  ±  papery,  1 -layered, 
medium  to  dark  brown,  densely  long-pubescent  (see  above);  inner 
surface  irregular  and  minutely  rough,  glabrous,  glossy,  dark  brown. 
Tegmen  absent. 


Fig.  1     Vochysia  ferruginea  Mart.  Fruit:  A,  undehisced  fruit;  B,  dehisced  capsule;  C,  t.s.  Seed:  D,  seed,  lateral  view;  E,  embryo  in  seminiferous  region, 
with  seed  coat  partially  removed;  F,  embryonic  cotyledon  and  radicle,  with  one  side  of  cotyledon  unfolded  and  second  cotyledon  removed  to  show 
primary  veins,  radicle,  and  angle  of  folding;  G,  t.s.  through  seminiferous  region  at  level  marked  by  arrow  in  E-F,  view  toward  radicle  and  wing.  Seedling: 
H,  seedling;  J,  cotyledon;  K,  cotyledonary  stipules,  hypocotyl,  and  petiole  ridges;  L-M,  cotyledonary  stipules  after  epicotyl  expands;  N,  first  leaf;  O, 
stipules  at  first  node.  Sapling:  P,  young  sapling;  Q,  branched  sapling;  R,  leaf  at  node  15;  S,  stipules  at  about  node  20.  Scale  bars:  black  or  black  and 
white,  1  mm  units;  white  only,  1  cm  units.  Symbols:  ac,  annulate  collet;  ab,  abaxial  surface  of  cotyledon;  ad,  adaxial  surface  of  cotyledon;  ^-c,, 
cotyledons  1  and  2;  en,  endocarp;  ex,  exocarp;  m,  midvein  of  cotyledon;  ms,  mesocarp;  r,  radicle;  re,  resin  canals;  sd,  seed;  so,  outline  of  seed  (dotted 
line);  ts,  testa;  f,-f4,  folds  of  cotyledon. 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


N.C.  GARWOOD 


Seedlings  and  saplings 

Seedlings  phanero-epigeal  with  photosynthetic  cotyledons. 

DEVELOPMENT.  Radicle  emerges  from  hilar  side  of  seed  near 
wing.  Hypocotyl  elongates  into  loop,  then  straightens,  often  carry- 
ing seed  upward.  Cotyledons  emerge  from  seed  and  fully  expand 
before  first  leaves  are  initiated.  Shoot  rest  period  is  short  during  the 
cotyledon  stage,  about  1  month  in  sun  and  3  months  in  shade.  Seed 
reserves  are  exhausted  by  the  end  of  this  period,  as  albino  seedlings 
die  at  the  cotyledon  stage  without  initiating  any  leaves.  Growth  is 
continuous,  with  a  pair  of  leaves  initiated  about  every  month  in  sun 
and  every  2-3  months  in  shade,  although  shade-produced  leaves  are 
smaller. 

ROOTS.  Primary-  root  30-50  x  0.6-0.8  mm,  circular  in  t.s.,  slightly 
sinuous,  cream  to  white,  becoming  medium  orange-brown;  narrow 
annular  ridge  encircles  axis  at  collet,  becoming  indistinct  with  age; 
surface  texture  and  colour  between  hypocotyl  and  root  change 
conspicuously  and  abruptly  at  collet.  Secondary  roots  to  0.2  mm 
wide,  sinuous,  very  sparsely  scattered  along  primary  root.  Root 
hairs  appear  abruptly  and  densely  on  primary  root  just  below  collet. 
Sapling  root  system:  taproot  thick,  long-tapering;  secondary  roots  ± 
thin  and  common  along  primary  root;  tertiary  and  higher  order  roots 
very  fine  and  numerous. 

STEMS.  Hypocotyl  15-67  x  0.25-0.50  mm,  ±  square  in  t.s.,  be- 
coming circular  in  t.s.  and  hollow  with  age;  4-ridged  to  4-winged, 
each  wing  to  0.5  mm  tall,  continuing  at  apex  into  lower  margin  of 
cotyledon  petiole  and  terminating  at  base  abruptly  above  collet; 
glabrous;  medium  yellow-green  to  green,  becoming  dark  brown  or 
red-brown.  Epicotyl  3-12  x  0.3-0.5  mm,  ±  square  in  t.s.,  becoming 
circular  with  age,  4-winged,  each  0.3-1.0  mm  tall  and  terminating 
apically  in  stipules,  glabrous  or  sparsely  short-pubescent,  medium 
green  to  yellow-green.  Hairs  short,  T-shaped  (ratio  of  arm  lengths  = 
1 : 1 ),  ±  sessile,  appressed,  straight,  and  light  brown  to  orange-brown. 
Cataphylls  absent.  Internodes  1-10,  2-15  mm  long;  later  internodes 
to  34  mm;  similar  to  epicotyl  except  later  internodes  becoming 
densely  short-  to  long-pubescent,  with  hairs  long,  unequally 
2-branched  (arm  length  ratio  to  8:1),  short-stalked,  ascending,  and 
curved  or  sinuous  or  coiled.  Older  stems  dark  brown,  with  wings 
often  red-brown,  hollow,  often  sparsely  pubescent  with  age;  wood 
cream-coloured.  Lenticels  appearing  on  stems  >  2  mm  wide,  sparse 
on  stems  2—4  mm  wide;  mostly  0.5  mm  long,  ±  round  to  elliptic, 
light  brown,  lighter  on  dark  stems.  Buds:  Cotyledonary  buds  hidden 
in  axils  and  inconspicuous  above  cotyledon  scars,  rarely  protruding 
to  ==  0.5  mm  in  older  seedlings,  moderately  short-pubescent.  Lateral 
buds  usually  slightly  protruding  out  of  axils,  to  1  mm  long  at  later 
nodes,  moderately  pubescent  at  early  nodes  to  densely  pubescent  at 
later  nodes,  with  accessory  bud  above  sylleptic  branches  at  later 
nodes.  Terminal  buds  naked,  partially  covered  by  stipules,  moder- 
ately short-pubescent  at  early  nodes  to  densely  long-pubescent  at 
later  nodes,  with  hairs  light  brown  to  orange-brown. 

COTYLEDONS.  Two,  opposite,  foliaceous,  simple,  petiolate,  stipu- 
late, emergent  from  seed,  spreading  apart,  persistent  at  least  until 
node  3  matures,  falling  sometime  before  node  9  matures  (between  9 
and  20  months).  Blades  10-15  x  13-23  mm;  straight,  flat,  depressed 
obovate,  with  right  half  (viewed  from  the  lower  abaxial  side)  usually 
slightly  larger,  2-lobed  to  2-cleft;  apex  of  lobes  obtuse  to  broadly 
obtuse;  base  emarginate  or  cordate,  often  asymmetrical  with  right 
lobe  slightly  larger,  or  half-rounded/half-cordate;  margins  entire. 
Blades  glabrous;  green  above,  lighter  below.  Venation  palmate, 
3-nerved;  acro-brochidodromous,  basal  and  perfect.  Primary  veins 
3;  moderate-sized;  impressed  to  slightly  raised  above,  moderately 
raised  below;  central  primary  vein  forked,  with  straight  stem  '/2-3/4 


distance  to  apex  and  branches  slightly  recurved;  lateral  primary 
veins  basal,  =  3/4  length  of  blade,  straight  to  curved,  diverging  at 
narrowly  acute  angle,  looped,  joining  branches  of  central  vein  to 
form  large  ±  triangular  to  obovate  intercostal  area,  the  pair  of 
intercostal  areas  usually  spanning  about  60-70%  of  the  width  of  the 
cotyledons.  Secondary  veins  moderate-sized  basally  along  lateral 
primaries  becoming  fine  apically;  16-24,  2-4  along  midvein,  6-10 
along  lateral  primaries,  6-10  along  branches  of  midvein;  mostly 
marginal,  alternate  along  midvein;  'A>-3/4  length  of  blade  (mostly  'A>- 
'/4  along  branches  of  midvein),  longest  basally;  mostly  curved  or 
arched  along  lateral  primaries,  hooked  along  branches  of  midvein, 
curved  to  zigzag  along  midvein,  rarely  scythe-shaped;  diverging  at 
narrowly  acute  angles  (<  45°)  basally  on  lateral  primaries  to  perpen- 
dicular angles  along  branches  of  midvein;  looped  marginally, 
sometimes  percurrent  in  intercostal  area,  mostly  obtusely  joined; 
slightly  raised  above,  slightly  raised  below.  Inter-secondary  veins 
absent.  Tertiary  veins  reticulate  within  intercostal  area,  looped  in 
marginal  area.  Petioles  0.5-1 .5  mm  long,  transversely  oblong  in  t.s., 
straight  to  slightly  curved,  2-ridged  above,  with  ridges  continuing 
into  lateral  primary  veins,  4-winged  along  margin,  with  lower  wings 
continuing  into  hypocotyl  wings  and  upper  wings  terminating  in 
fleshy  knobs  near  centre  of  hypocotyl,  glabrous.  Stipules  usually 
fused  and  interpetiolar,  sometimes  separate  and  paired;  composed  of 
two  structures,  both  variable  in  shape  and  development.  The  lower 
part  usually  a  ±  conical  projection  between  the  petioles  at  the  base  of 
the  upper  marginal  ridge  of  each  petiole,  sometimes  with  two  or 
three  separate  projections  or  these  absent,  mostly  0.125-0.25  mm 
long,  usually  perpendicular  to  the  stem,  sometimes  gland-like 
apically.  The  upper  part  usually  a  flattened  interpetiolar  scale-like 
outgrowth  just  above  the  petioles,  initially  folded  over  the  terminal 
bud,  becoming  erect  and  visible  as  the  epicotyl  expands,  0.25-0.5 
mm  tall  and  0.5-0.75  mm  wide,  ±  transversely  oblong  to  depressed 
ovate,  usually  2-  to  5-lobed  or  -cleft,  but  sometimes  divided  into  two 
separate  parts  or  unlobed,  the  apex  of  lobes  gland-like  when  young; 
persistent,  papery,  green  becoming  light  brown. 

LEAVES.  Opposite,  decussate  on  trunk  axis,  distichous  on  branches; 
simple;  short-petiolate,  stipulate.  Leaf  blades  17-32  x  3-9  mm  at 
first  node;  20-67  x  5-20  mm  at  nodes  2-10;  to  142  x  49  mm  at  later 
nodes;  65-125  x  25-37  mm  on  branches;  straight;  narrowly  elliptic 
at  nodes  1-3,  narrowly  elliptic  or  narrowly  obovate  at  later  nodes, 
elliptic  or  narrowly  elliptic  or  narrowly  obovate  or  linear-obovate  on 
branches;  apex  narrowly  obtuse  or  acuminate  at  nodes  1-10,  acumi- 
nate to  long-acuminate  at  later  nodes  and  on  branches,  mucronate  at 
most  nodes;  base  mostly  attenuate  at  early  nodes,  sometimes  nar- 
rowly cuneate  at  later  nodes,  mostly  acute  on  branches;  margins 
entire,  sometimes  undulate.  Blades  green  above,  lighter  below;  flat 
between  secondary  veins  above  when  fresh;  papery;  densely  minutely 
pusticulate  below  (from  internal  crystals),  but  weakly  so  at  nodes  1- 
2;  densely  minutely  pusticulate  above,  with  pustules  usually  white 
on  larger  leaves;  slightly  to  moderately  short-  to  long-pubescent  on 
blade  and  veins  below  at  early  nodes  becoming  densely  long-  to  very 
long-pubescent  only  on  the  midvein  at  later  nodes;  glabrous  on  blade 
above,  slightly  to  moderately  ±  short-pubescent  on  midvein  at  early 
nodes  to  densely  long-pubescent  at  later  nodes;  slightly  to  moder- 
ately short-  to  long-pubescent  along  margin,  often  also  densely 
minute-pubescent  when  young  at  later  nodes;  pubescence  light 
brown  to  orange-brown.  Hairs  on  midvein  above  simple,  ±  erect, 
straight  to  slightly  curved,  mostly  0.5-0.75  mm  long  on  first  node,  to 
1-2  mm  long  on  later  nodes;  those  on  blade  and  veins  below 
T-shaped  (ratio  of  arm  lengths  ~  1:1),  ±  sessile,  appressed  and 
straight  at  early  nodes,  mostly  0.25-0.75  mm  long,  becoming  pro- 
gressively more  unequally  2-branched  (ratio  of  arm  lengths  mostly 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


<  4:1  on  blades  but  to  8:1  on  midvein),  short-stalked,  ascending, 
curved  or  sinuous  or  coiled  at  later  nodes;  those  on  margins  T-shaped 
or  2-branched  (as  above)  and,  at  later  nodes,  also  minute,  simple,  ± 
erect  and  sinuous.  Young  leaves  conduplicate,  spreading  open  before 
half-grown,  densely  pubescent,  light  green,  rigid,  erect  to  horizon- 
tal. Venation  pinnate;  brochidodromous.  Midvein  moderate-sized; 
straight;  flush  to  slightly  raised  above,  impressed  at  later  nodes  and 
when  fresh,  prominently  raised  below.  Secondary  veins  ±  fine;  14— 
17  on  leaves  at  first  node,  15-30  at  nodes  2-10,  to  30  at  later  nodes 
and  on  branches;  alternate;  mostly  '/6-'/3  length  of  blade,  longest 
centrally;  mostly  hooked  or  arched,  sometimes  also  slightly  zigzag; 
diverging  at  moderately  acute  angles  (45"-65°);  looped,  sometimes 
diffusely  looped  basally  on  larger  leaves,  obtusely  joined;  slightly 
raised  above,  slightly  raised  below.  Inter-secondary  veins  infre- 
quent. Tertiary  veins  reticulate  within  intercostal  area,  looped  in 
marginal  area.  Petioles  0.5^.0  mm  long,  <  2  mm  long  at  nodes  1- 
1 0;  ±  circular  in  t.s.,  straight  to  slightly  curved;  shallowly  channelled, 
with  erect  narrow  margins  above;  sparsely  to  moderately  pubescent. 
Stipules  paired;  0.5-1 .0  mm  long  at  nodes  1-6,  to  3  mm  long  at  later 
nodes  and  on  branches;  simple,  triangular  to  narrowly  triangular; 
persistent,  ±  papery;  densely  pubescent,  pale  against  dark  older 
stems;  the  inner  margin  and  base  continuing  to  and  perhaps  above 
the  petiole;  the  outer  margin  and  base  usually  terminating  abruptly, 
but  sometimes  forming  a  low  interpetiolar  ridge  with  the  stipule  of 
the  opposite  leaf  as  the  stem  thickens  at  early  but  not  later  nodes. 

ARCHITECTURE.  Trunk  axis  monopodial  to  at  least  37  cm  tall 
(based  on  one  individual).  Initial  axis  orthotropic.  Branches 
plagiotropic  but  ±  ascending,  with  leaves  distichous  through  twist- 
ing of  internodes;  sylleptic;  alternate  and  perpendicular  at  two 
consecutive  nodes  (semi-decussate);  first  produced  3 1-34  cm  above 
the  collet  (at  about  nodes  17-18).  First  branch  internode  60-75  mm 
long,  much  longer  than  superadjacent  trunk  internode  (10-30  mm). 
First  branch  leaf  half-  to  full-size  of  later  branch  leaves.  Single  or 
paired  sylleptic  secondary  branches  were  produced  on  the  primary 
branches  24-29  cm  from  the  trunk.  Leaves  held  horizontal.  Several 
factors  indicate  that  growth  of  the  trunk  axis  above  the  branches 
stopped  or  slowed  considerably  while  the  branches  were  develop- 
ing, so  that  overall  growth  of  the  trunk  was  rhythmic  (see  Fig.  1Q): 
the  trunk  axis  was  very  short  (about  4  cm  long)  compared  to  the 
length  of  the  branches  (28-38  cm  long);  it  was  non-woody  and  very 
thin  compared  to  the  woody  internodes  on  the  trunk  below  the 
branches  and  on  the  lower  parts  of  the  branches;  and  it  had  matured 
only  one  pair  of  leaves  compared  to  7-8  pairs  on  the  branches .  At  the 
time  of  harvest,  the  trunk  axis  had  re-initiated  growth,  although  the 
primary  and  secondary  branches  were  still  elongating.  Rhythmic 
growth,  if  substantiated,  would  suggest  Massart's  model. 

Ecology 

PHENOLOGY.  In  Panama,  Vochysiaferruginea  usually  flowers  from 
late  March  to  early  July,  and  sometimes  to  a  lesser  extent  in 
September-October,  and  its  fruits  mature  August-October  (Croat, 
1978).  Ripe  fruits  were  dehiscing  and  releasing  mature  seeds  in 
October  (Garwood  1630A).  Elsewhere  in  Central  America,  it  prima- 
rily flowers  in  April-June  and  October-December  and  fruits 
July-September  and  December-January  (Croat,  1978;Flores,  19936; 
Moreira  &  Arnaez,  1994). 

DISPERSAL.  When  ripe,  valves  of  the  dark  brown  capsules  open 
and  fold  back,  exposing  the  glossy  bright  orange-brown  endocarp 
and  releasing  the  seeds.  Seeds  are  wind-dispersed. 

VIABILITY.  Seeds  remain  viable  for  2-3  months  when  stored  at 
ambient  temperature  (24-26°C)  and  moisture  levels  if  well-aerated 


(Flores,  1993ft).  Flores  (1993ft)  considers  the  seeds  orthodox,  al- 
though the  initial  moisture  content  of  the  seeds  (23%)  is  rather  high. 

GERMINATION.  Seeds  germinate  rapidly  (13  days)  at  moderate 
percentages  (39-50%)  in  both  sun  and  shade  in  the  growing  house 
(this  study).  Other  studies  also  report  rapid  germination  (8-14  days) 
in  the  forest  or  growing  house,  but  higher  germination  percentages 
(80-95%)  (Flores,  1993ft;  Moreira  &  Arnaez,  1994). 

ESTABLISHMENT  AND  GROWTH.  The  earliest  stages  of  natural  seed- 
ling establishment  and  growth  have  not  been  studied,  although 
Flores  (1993ft)  noted  that  germinating  seeds  and  young  seedlings 
suffered  high  predation  from  ants  and  other  herbivores.  Boucher  et 
al.  (1994)  reported  exceptional  seedling  densities  and  sapling  growth 
after  Hurricane  Joan  in  Nicaragua  in  October  1989,  which  killed  all 
adults  in  the  population.  Four  months  after  the  hurricane,  there  were 
94  500  seedlings  and  saplings  (0.1-1.5  m  tall)  per  hectare,  but  few 
young  trees  (3.2-10  cm  DBH).  The  extremely  open  canopy,  created 
by  the  hurricane  through  defoliating  and  felling  trees,  favoured 
survivorship  and  growth  of  these  juveniles,  such  that  young  trees 
were  abundant  four  years  later  and  were  growing  on  average  at  about 
1 .4  cm  DBH/year.  Under  the  less  open  canopy  of  Barro  Colorado 
Island,  not  recently  if  ever  disturbed  by  hurricanes,  similar-sized 
individuals  (1-9.9  cm  DBH)  grew  on  average  only  0.45  cm  DBH/ 
year,  but  larger  individuals  (>  10  cm  DBH)  grew  more  rapidly,  1.3- 
1.7  cm  DBH/year  (Condit  et  al.,  1993). 

The  population  dynamics  and  appropriate  management  strategies 
for  this  species  may  differ  greatly  in  hurricane-prone  areas  such  as 
Nicaragua  compared  to  hurricane-free  areas  such  as  Panama.  Where 
hurricanes  are  common  in  the  Caribbean,  they  occur  mostly  August- 
October  during  the  period  Vochysia  ferruginea  is  maturing  and 
dispersing  its  seeds.  A  hurricane  occurring  late  in  the  season  might 
blow  the  wind-dispersed  seeds  over  large  areas  of  disturbed  forest, 
expanding  the  population.  An  earlier  hurricane  might  kill  adult  trees 
before  seeds  could  be  matured  and  dispersed,  preventing  regenera- 
tion from  seed  and  eliminating  the  local  population,  unless  most 
regeneration  comes  from  established  shade-tolerant  juveniles.  It  is 
not  yet  known  whether  seedling  recruitment  after  disturbance  is 
primarily  from  recently  germinated  seedlings  from  the  current 
year's  seed  crop  or  older  shade-tolerant  juveniles  surviving  from 
previous  years. 

Vochysia  guatemalensis  Donn.  Sm.  in  Bot.  Gaz.  12:  131,  pi.  22 

(1887). 
Vochysia  hondurensis  Sprague  in  Bull.  Misc.  Inform.,  Kew  1922: 

183(1922). 
Fig.  2A-D. 

HABIT  AND  DISTRIBUTION.  Trees,  30-55  m  tall.  Veracruz,  Mexico, 
to  Panama. 

COLLECTIONS.  Belize.  Cayo  District,  Chiquibul:  Palmar,  Garwood 
403 1  (BM,  seedlings  with  seed  voucher);  New  Maria,  Monro  867  to 
871,  Monro,  Howe  &  King  M4162X,  M7231X,  M7340X  (seed- 
lings) and  M6002X  (sapling);  28  seedlings  to  17  cm  tall  examined, 
sapling  height  about  2  m. 

TAXONOMIC  NOTE.  The  seeds  associated  with  Garwood  4031 
unmistakenly  identify  the  seedlings  to  genus.  If  one  accepts  the 
synonymy  of  Vochysia  guatemalensis  and  V.  hondurensis  (Standley 
&  Steyermark,  1949;  Gaos,  1978;  Flores,  1993a),  then  only  one 
species  has  been  reported  from  Belize  (Stafleu,  1948;  Standley  & 
Steyermark,  1949).  That  the  mature  leaves  and  stems  are  nearly 
glabrous  supports  the  identification  of  these  collections  as  V. 
guatemalensis. 


N.C.  GARWOOD 


Fig.  2     Vochysia  guatemalensis  Donn.  Sm.  Seedling:  A,  cotyledon;  B,  first  leaf;  C-D,  cotyledonary  stipules.  Vochysia  sp.  Seedling:  E-F,  cotyledons;  G, 
cotyledonary  stipules;  E,  Hampshire  &  Whitefoord  269;  F-G,  Garwood  135.  Scale  bars:  black,  1  mm;  white,  1  cm. 


Seedlings  and  saplings 

Seedlings  phanero-epigeal  with  photosynthetic  cotyledons. 

ROOTS.  Primary  root  30-70  x  0.8-1 .2  mm,  circular  in  t.s.,  slightly 
sinuous,  cream-coloured;  narrow  annular  ridge  encircles  axis  at 
collet,  becoming  indistinct  with  age.  Secondary  roots  sinuous, 
scattered  along  primary  root. 

STEMS.  Hypocotyl  50-85  x  1  mm,  square  to  rectangular  in  t.s., 
becoming  circular  with  age;  4-ridged  or  4-winged,  each  wing  <  0.25 
mm  tall,  continuing  at  apex  into  lower  margin  of  cotyledon  petiole; 
glabrous;  green.  Epicotyl  5-22  x  1  mm,  ±  square  in  t.s.  when  young, 
becoming  circular  with  age;  4-winged,  each  wing<  0.25  mm  tall  and 
terminating  apically  in  a  stipule;  glabrous;  green.  Cataphylls  absent. 
Internodes  1-16, 3-5  mm  long;  4-winged,  each  wing  about  0.25  mm 
tall  and  terminating  apically  in  a  stipule;  glabrous;  green.  Older 
stems  light  brown  to  grey-brown,  hollow.  Buds:  Cotyledonary  buds 
hidden  in  axils,  sparsely  pubescent.  Lateral  buds  protruding  out  of 
axils,  <  0.5  mm  long  at  nodes  1-15,  sparsely  short-pubescent  at  first 
node,  becoming  densely  pubescent  at  later  nodes.  Terminal  buds 
naked,  sparsely  pubescent  at  first  node,  becoming  moderately  pu- 
bescent at  later  nodes.  Hairs  on  all  buds  clear  to  white,  mostly  <  0.25 
mm  long. 

COTYLEDONS.  Two,  opposite,  foliaceous,  simple,  petiolate,  stipu- 
late, emergent  from  seed,  spreading  apart,  persistent  until  at  least 
leaves  at  nodes  2-3  mature.  Blades  15-26  x  25-37  mm;  straight, 
flat,  ±  depressed  obovate,  with  right  half  (seen  from  lower  or  abaxial 
side)  usually  larger  at  both  base  and  apex;  apex  truncate  to  emarginate, 
often  obliquely  so,  rarely  ±  rounded;  base  cordate,  rounded,  to  ± 


truncate,  often  obliquely  so;  margins  entire.  Venation  palmate,  3- 
nerved;  acro-brochidodromous,  basal  and  perfect.  Primary  veins  3, 
±  fine,  slightly  raised  above,  moderately  raised  below;  central 
primary  vein  forked,  with  straight  stem  '/2  to  3A  distance  to  apex  and 
branches  slightly  recurved;  lateral  primary  veins  basal,  about  3A 
length  of  blade,  mostly  curved,  diverging  at  narrowly  acute  angle, 
looped,  joining  branches  of  central  vein  and  forming  ±  small  obovate 
or  triangular  intercostal  area  or  rarely  branching  before  joining  the 
central  vein  and  forming  a  ±  indistinct  intercostal  area,  the  pair  of 
intercostal  areas,  when  distinct,  usually  spanning  about  30-50%  of 
the  width  of  the  cotyledon.  Secondary  veins  moderate-sized  basally 
along  lateral  primaries  becoming  fine  apically;  12-16,  0-1  along 
midvein,  8-10  along  lateral  primaries,  4-5  along  branches  of  midvein; 
mostly  marginal;  to  2/3  length  of  blade,  longest  basally;  mostly 
arched  or  hooked;  diverging  at  narrowly  acute  angles  basally  on 
lateral  primaries  to  perpendicular  angles  along  branches  of  midvein; 
looped  marginally,  acutely  to  obtusely  joined;  flush  above,  moder- 
ately raised  below.  Petioles  2-3  mm  long,  transversely  oblong  in  t.s., 
2-ridged  above,  with  ridges  continuing  into  lateral  primary  veins;  4- 
ridged  along  margins,  the  lower  ridges  continuing  down  into  the 
ridges  or  wings  on  the  hypocotyl  and  the  upper  ridges  terminating  at 
the  stipules;  glabrous.  Stipules  separate  and  paired  or  sometimes 
fused  and  interpetiolar,  composed  of  two  structures,  both  variable  in 
shape  and  development.  The  lower  part  a  pair  of  ±  conical  projec- 
tions between  the  petioles  near  the  base  of  the  upper  marginal  ridge 
of  each  petiole,  one  or  both  sometimes  absent,  mostly  0.25-0.75  mm 
long,  usually  perpendicular  to  the  stem,  sometimes  gland-like 
apically.  The  upper  part  usually  a  pair  of  flattened  scale-like 
outgrowths  between  and  just  above  the  petioles,  initially  folded  over 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


the  terminal  bud  but  becoming  erect  and  visible  as  the  epicotyl 
expands,  each  ±  ovate  to  rhombic,  0.5-0.75  mm  tall  and  wide,  the 
apex  often  3-lobed  and  gland-like,  the  pair  sometimes  fused  to- 
gether, then  0.5-0.75  x  0.75-1.5  mm  tall  and  wide  and  the  apex 
multilobed  or  irregular. 

LEAVES.  Opposite,  decussate  on  seedling  trunk  axis,  sometimes  3- 
whorled  on  sapling  trunk  axis;  simple,  short-petiolate;  stipulate. 
Leaf  blades  30-69  x  1 1-19  mm  at  first  node;  30-82  x  13-17  mm  at 
nodes  2-15,  to  195  x  52  mm  on  saplings;  straight;  narrowly  obovate 
to  narrowly  elliptic  at  nodes  1-15  and  on  sapling;  apex  acuminate  or 
long-acuminate,  rarely  acute  at  first  node;  bases  cuneate  to  narrowly 
acute;  margins  entire.  Blades  green,  papery  at  early  nodes  becoming 
subcoriaceous  on  larger  saplings;  sometimes  moderately  pusticulate 
above  and  below  on  later  leaves,  the  pustules  pale  above;  the  blade 
nearly  glabrous  when  fully  expanded,  the  midvein  below  often 
sparsely  short-pubescent,  but  moderately  to  densely  pubescent  on 
the  blade  and  midvein  below  and  along  margins  when  expanding 
leaves  of  seedlings  are  <  10  mm  long.  Hairs  simple,  basifixed,  <  0.25 
mm  long,  clear  to  white.  Venation  pinnate,  brochidodromous.  Midvein 
moderate-sized;  straight;  flush  to  slightly  raised  above,  prominently 
raised  below.  Secondary  veins  thin,  14-22  at  first  node,  to  22  at  later 
nodes;  alternate;  mostly  '/4  to  V3  length  of  blade,  longest  centrally; 
mostly  hooked  or  arched,  diverging  at  a  moderately  acute  angle; 
looped,  obtusely  joined;  slightly  raised  above  and  below. 
Intersecondary  veins  infrequent.  Tertiary  veins  reticulate  in  inter- 
costal area,  looped  in  marginal  area.  Petioles  1  mm  long  at  first  node, 
to  2  mm  long  at  nodes  2-15,  to  7  mm  on  saplings;  shallowly 
channelled,  with  erect  narrow  margins  above.  Stipules  paired;  1 .0- 
1.2  mm  long  at  first  node,  to  1.7  mm  long  at  nodes  2-15;  simple, 
narrowly  triangular;  the  inner  margin  and  base  continuing  above  the 
petiole  and  apparently  terminating  near  the  lateral  bud;  the  outer 
margin  and  base  usually  terminating  abruptly,  but  sometimes  form- 
ing a  low  interpetiolar  ridge  with  the  stipule  of  the  opposite  leaf  as 
the  stem  thickens  at  some  early  nodes;  persistent;  often  sparsely 
short-pubescent  toward  the  apex. 

Vochysia  sp. 

Fig.  2E-G. 

COLLECTIONS.  Panama.  Colon  Province:  Santa  Rita  Ridge, 
Garwood  135  (seedlings  with  seed  fragment);  Chiriqui  Province: 
Fortuna,  Hampshire  &  Whitefoord  269  (BM,  seedlings);  15  seed- 
lings to  9  cm  tall  examined. 

TAXONOMIC  NOTE.  Specific  identification  is  not  yet  possible,  as 
Garwood  135  had  only  a  seed  fragment  attached  and  Hampshire  & 
Whitefoord  269  lacked  seeds  altogether.  In  addition  to  Vochysia 
ferruginea,  V.  allenii  Standl.  &  L.O.  Williams,  V.  jefensis  A. 
Robyns,  and  V.  speciosa  Warm,  have  been  reported  from  eastern 
Panama  and  V.  hondurensis  (-  V.  guatemalensis)  from  western 
Panama  (D'Arcy,  1987).  As  immature  leaves  of  both  collections 
have  T-shaped  hairs  >  0.25  mm  long  on  the  lower  surfaces  (see 
below),  it  is  not  V.  guatemalensis,  which  has  simple  basifixed  hairs 
<  0.25  mm  long  (see  above).  The  T-shaped  hairs  on  the  first  leaves 
ally  it  with  V.  ferruginea,  but  the  cotyledons  are  about  1.5-2  times 
larger.  Thus,  the  Chiriqui  collection  is  a  new  species  record  for 
western  Panama.  As  the  cotyledons  from  the  Chiriqui  collection 
are  consistently  larger  than  those  from  the  Colon  collection  (Fig. 
3),  two  species  might  be  represented.  Given  their  similarity  and  the 
paucity  of  material,  they  are  described  together  below,  with  the  few 
differences  noted. 

Seedlings 

Seedlings  phanero-epigeal  with  photosynthetic  cotyledons. 


ROOTS.  Primary  root  45-70  mm  long,  1  mm  wide,  circular  in  t.s., 
slightly  sinuous,  medium  brown. 

STEMS.  Hypocotyl  50-86  x  =  1  mm  long  and  wide,  ±  square  to 
rectangular  in  t.s.,  4-ridged  to  4-winged,  each  wing  <  0.25  mm  tall 
and  continuing  at  apex  into  lower  margin  of  petiole;  glabrous. 
Epicotyl  (immature)  4-ridged  or  4-winged,  each  wing  <  0.25  mm 
tall;  glabrous.  Buds:  Cotyledonary  buds  hidden  in  axils  (Garwood 
1 35)  or  0.75  mm  tall  and  densely  pubescent  (Hampshire  &  Whitefoord 
269). 

COTYLEDONS.  Two,  opposite,  foliaceous,  simple,  petiolate,  stipu- 
late, emergent  from  seed,  spreading  apart,  persistent.  Blades  14-21 
x  22-40  mm  (14-17  x  25-32  mm  in  Garwood  135,  19-21  x  34-^0 
mm  in  Hampshire  &  Whitefoord  269);  straight,  flat,  depressed 
obovate,  with  right  side  (viewed  from  lower  or  abaxial  surface) 
usually  larger  than  or  sometimes  equal  to  left  side;  apex  truncate, 
emarginate,  or  cordate,  sometimes  obliquely  so;  base  rounded, 
truncate  or  emarginate,  sometimes  unequal;  margins  entire.  Blades 
green,  glabrous.  Venation  palmate,  3-nerved;  acro-brochidodromous, 
basal  and  perfect.  Primary  veins  3,  moderate-sized;  central  primary 
vein  forked,  with  straight  stem2/3  to3A  distance  to  apex  and  branches 
recurved;  lateral  primary  veins  basal,  about  3/4  length  of  blade, 
mostly  curved,  diverging  at  narrowly  acute  angle,  looped,  joining 
branches  of  central  vein  to  form  ±  small  obovate  or  triangular 
intercostal  area,  the  pair  of  intercostal  areas  usually  spanning  about 
40-50%  of  the  width  of  the  cotyledon.  Secondary  veins  moderate- 
sized  basally  becoming  fine  apically;  10-16,  0-2  along  midvein, 
6-9  along  lateral  primaries,  3-5  along  branches  of  midvein;  mostly 
marginal;  '/3  to  nearly  full  length  of  blade,  longest  basally;  mostly 
arched  to  hooked,  diverging  at  narrowly  acute  angles  basally  to 
perpendicularly  apically;  looped  marginally,  acutely  to  perpendicu- 
larly joined;  flush  above,  moderately  raised  below.  Tertiary  veins 
reticulate  within  intercostal  areas,  looped  in  marginal  areas.  Petioles 
about  2  mm  long,  transversely  oblong  in  t.s.,  2-ridged  above,  with 
ridges  continuing  into  the  lateral  primary  veins;  4-ridged  along 
margin,  with  lower  ridges  continuing  into  hypocotyl  ridges  and 
upper  ridges  terminating  in  the  stipules;  glabrous.  Stipules  usually 
fused  and  interpetiolar,  sometimes  separate  and  paired;  composed  of 
two  structures,  both  variable  in  shape  and  development.  The  lower 
part  a  pair  of  ±  conical  projections  between  the  petioles  near  the  base 
of  the  upper  marginal  ridge  of  each  petiole,  sometimes  absent  or 
bifid,  mostly  0.25-0.5  mm  long,  usually  perpendicular  to  the  stem, 
sometimes  gland-like  apically.  The  upper  part  usually  a  flattened 
scale-like  outgrowth  between  and  just  above  the  petioles,  initially 
folded  over  the  terminal  bud  but  becoming  erect  and  visible  as  the 
epicotyl  expands,  ±  transversely  oblong  to  depressed  ovate,  0.5- 
0.75  x  0.75-1.0  mm  tall,  3-  to  5-lobed  and  gland-like  at  the  apex, 
sometimes  divided. 

LEAVES  (only  immature  leaves  at  first  node  present).  Opposite, 
decussate  on  trunk  axis;  simple,  short-petiolate.  Blades  moderately 
pubescent  below  and  along  margins  when  expanding  blades  are  <  5 
mm  long,  sparsely  pubescent  when  half-expanded.  Hairs  0.25- 
0.625  mm  long,  T-shaped,  sessile,  pale  orange-browa  Venation 
pinnate,  brochidodromous. 

Key  to  known  seedlings  of  Vochysia  in  Panama  and 
Central  America 

1 .      T-shaped  hairs  present  on  lower  surface  of  young  leaves  at  first  and  early 
nodes 

2.  Cotyledons  13-23  mm  wide  and  10-15  mm  long 


V.  ferruginea 


N.C.  GARWOOD 


2.  Cotyledons  22-40  mm  wide  and  14-21  mm  long V.  sp. 

Simple  basifixed  hairs  present  on  lower  surface  of  young  leaves  at  first 
and  early  nodes  or  hairs  absent;  cotyledons  25-37  mm  wide  and  1 5-26 
mm  long V.  guatemalensis 


DISCUSSION 

Vochysiaceae  is  a  moderate-size,  primarily  neotropical  family  di- 
vided into  two  tribes.  Vochysieae  includes  Vochysia  (100  species), 
Callisthene  (12  species),  Salvertia  (1  species),  and  Qualea  sensu 
lato  (77  species,  including  Ruizterania);  Erismeae  includes  Erisma 
(16  species)  and  Erismadelphus  (2  species),  the  only  African  taxa 
(Stafleu,  1948-1954;  Keay  &  Stafleu,  1953;  Marcano-Berti,  1969; 
Kawasaki,  in  press).  The  monotypic  Euphronia  (=  Lightia),  some- 
times included  in  Vochysiaceae  (Lleras,  1976;  Cronquist,  1981)  or 
segregated  as  the  Euphroniaceae  (Marcano-Berti,  1989,  1990),  is 
not  closely  related  to  Vochysiaceae  according  to  recent  molecular 
studies  (Litt  et  al.,  1995)  and  will  not  be  further  discussed. 

Fruit 

There  are  two  basic  types  of  fruit  in  Vochysiaceae  (Warming,  1 875; 
Stafleu,  1948- 1954;  Keay  &  Stafleu,  1953).  The  dehiscent  trilocular 
capsules  of  Vochysieae,  derived  from  superior  ovaries,  contain 
winged  seeds.  The  indehiscent  unilocular  fruits  of  Erismeae,  derived 
from  inferior  to  semi-inferior  ovaries,  have  persistent  enlarged 
winged  calyx  lobes.  Within  Vochysieae,  fruits  of  Callisthene  have  a 
thick  persistent  central  column  and  a  fragile  exfoliating  exocarp  not 
found  in  the  other  genera.  Several  of  the  12  ovules  per  locule  mature 
into  seeds  in  Qualea  (Stafleu,  1953);  one  or  more  of  the  few  ovules 
per  locule  mature  in  Callisthene,  (Stafleu,  1952).  In  Vochysia  and 
Salvertia,  however,  only  one  of  the  two  ovules  in  each  locule  usually 
develops  into  a  mature  seed  (Warming,  1875;  Stafleu  1948;  Standley 
&Steyermark,  1949;Robyns,  1967a;Gaos,  1978;Moreira&Arnaez, 
1994;  this  study),  although  two  mature  according  to  Flores  (1993a, 
b). 

Fruits  of  Vochysia  ferruginea  are  typical  of  the  genus  (Stafleu, 
1948),  being  3-locular  loculicidal  capsules,  oblong  or  obovate  in 
outline  and  3-angled.  Variation  in  size,  shape,  and  surface  texture  of 
fruits  in  Vochysia  is  not  well-documented:  fruits  of  many  species  are 
still  unknown  (Warming,  1875;  Stafleu,  1948;  but  see  van  Roosmalen, 
1985).  The  gross  morphology  of  fruits  has  been  little  studied.  Stafleu 
( 1 948)  described  the  pericarp  of  the  genus  as  consisting  of  a  black  or 
dark  bluish  exocarp  'strongly  adhering  to  the  shining  golden-yellow 
endocarp' ,  an  observation  repeated  by  later  authors  (Robyns,  1 967a; 
van  Roosmalen,  1985),  but  Flores  (1993a,  b)  noted  the  presence  of 
a  well-developed  mesocarp  in  both  V.  ferruginea  and  V?  guatemalensis. 
In  V.  ferruginea,  there  is  a  brown  to  black  exocarp,  a  light  to  medium 
brown  spongy  to  woody  mesocarp,  thickest  near  the  centre  of  the 
fruit,  and  a  pale  crustaceous  endocarp  of  even  thickness  around  each 
locule  (Fig.  1C).  The  mesocarp  is  penetrated  by  longitudinal  resin 
canals  adjacent  to  the  locules  (Fig.  1C)  and  is  also  well-vascularized 
(Flores  1993a,  b).  In  mature  fruit,  the  endocarp  is  as  well-developed 
around  empty  locules  as  around  those  filled  with  mature  seeds, 
indicating  that  development  of  each  locule  is  not  dependent  on  the 
growth  of  fertilized  ovules  within  it.  The  locules  complete  develop- 
ment before  the  seeds  and  determine  the  shape  of  the  seed  wing 
(Boesewinkel  &  Venturelli,  1987). 

Seed 

Seeds  are  winged  in  the  Vochysieae.  The  wings  are  large  and 
unilateral  in  Vochysia,  Qualea,  and  Salvertia,  but  small  and  circum- 


ferential in  Callisthene,  and  are  formed  from  compacted  testal  hairs 
in  Vochysia  and  Salvertia,  but  from  extensions  of  the  testa  in  Qualea 
and  Callisthene  (Warming,  1875;  Stafleu,  1948-1953;  Boesewinkel 
&  Venturelli,  1987).  The  seeds  of  V.  ferruginea  are  typical  of  the 
genus  (Warming,  1875).  The  wing  is  on  the  anti-hilar  side  of  the 
seed,  and  the  micropyle  extends  along  the  seminiferous  area  and 
terminates  near  the  hilum,  as  in  V.  pygmaea  Bong,  and  Salvertia 
(Boesewinkel  &  Venturelli,  1987).  In  contrast,  the  wing  of  Qualea  is 
on  the  hilar  side  of  the  seed,  such  that  the  micropyle  extends  along 
the  anti-hilar  margin  of  the  wing  and  opens  at  the  apex  of  the  wing 
away  from  the  hilum  (Boesewinkel  &  Venturelli,  1987).  The  final 
shape  of  the  wing  in  Vochysia  is  determined  by  the  shape  of  the 
locule:  the  unicellular  hairs  grow  upward  from  the  testa  to  fill  and 
take  their  shape  from  the  empty  locule  (Boesewinkel  &  Venturelli, 
1987).  Hence,  much  of  the  variation  in  size  and  shape  of  seeds 
among  species  in  Vochysia  probably  arises  from  differences  in  the 
size  and  shape  of  the  mature  locules. 

There  has  been  little  work  on  seed  anatomy  of  Vochysiaceae 
(Corner,  1976;  Boesewinkel  &  Venturelli,  1987).  Stafleu  (1948, 
1 953)  described  the  testa  of  Vochysia  as  brown,  thin,  and  chartaceous, 
and  the  tegmen  as  very  thin,  paper  white,  and  adhering  to  the  testa, 
and  the  testa  of  Qualea  as  chartaceous.  The  mature  seed  coat  of  V. 
pygmaea,  however,  consists  only  of  the  outer  layers  of  the  testa, 
because  the  inner  layers  of  the  testa  are  crushed  and  the  tegmen 
resorbed  during  development  (Boesewinkel  &  Venturelli,  1987). 
The  seed  coat  also  appeared  as  one  undifferentiated  layer  at  maturity 
in  V.  ferruginea  (this  study),  but  Flores  (1993a,  b)  described  a  dark 
exotesta  distinct  from  the  dead  cell  layers  and  air  spaces  of  the 
mesotesta  and  endotesta  and  remnants  of  the  tegmen  in  V.  ferruginea 
and  V.  guatemalensis.  In  contrast,  the  mature  seed  coat  of  Qualea 
densiflora  Warm,  and  Callisthene  retains  both  the  inner  layers  of  the 
testa  and  tegmen  (Boesewinkel  &  Venturelli,  1987).  The  seed  coat  of 
Erisma  and  Erismadelphus  is  thick  and  contains  a  large  branched 
vascular  bundle  (Boesewinkel  &  Venturelli,  1987).  The  vascular 
bundle  of  V.  ferruginea,  hitherto  undescribed  in  Vochysieae,  is 
unbranched  and  terminates  at  the  tip  of  the  seminiferous  area.  The 
hilum  of  V.  ferruginea  (7-10  mm  long)  is  longer  than  in  V.  pygmaea 
(3-4  mm  long),  and  stretches  over  a  greater  proportion  of  the  length 
of  the  winged  seed  (35-40%  versus  about  20%).  Due  to  the  long 
hilum,  the  seed  type  of  V.  ferruginea  is  described  as  hilar-anatropous. 

Endosperm  is  absent  in  mature  seeds  of  Vochysiaceae  (Bentham 
&  Hooker,  1867;  Baillon,  1874;  Warming,  1875;  Petersen,  1896; 
Stafleu,  1948-1953;  Standley  &  Steyermark,  1949;  Robyns,  1967a; 
Gaos,  1978;  Boesewinkel  &  Venturelli,  1987;  Flores,  1993o,  b). 
None  was  seen  in  Vochysia  ferruginea.  Occasional  reports  that 
endosperm  is  rarely  well-developed  (Cronquist,  1981;  Niembro 
Rocas,  1989)  do  not  appear  well-founded.  A  thin,  whitish  aril, 
adhering  to  the  testa,  was  reported  in  V.  guatemalensis  (Gaos,  1978) 
and  in  the  family  (Niembro  Rocas,  1989):  this  appears  to  be  a 
mistranslation  of  Stafleu's  (1948)  description  of  the  tegmen  (see 
above). 

Embryo 

In  all  genera,  the  embryo  of  the  mature  seed  is  straight  and  the 
plumule  and  radicle  poorly  developed  (Warming,  1875;  Stafleu, 
1948-1954).  In  Vochysia  ferruginea,  the  embryonic  axis  is  well- 
developed;  the  hypocotyl  is  relatively  long  compared  to  the  poorly 
developed  radicle,  compressed,  4-winged,  and  annulate  at  the  collet. 
The  winged  hypocotyl  and  annulate  collet  persist  into  the  seedling 
stage,  confirming  the  identification  of  the  structures  in  the  seed.  In 
Vochysieae,  the  embryonic  cotyledons  are  generally  thin,  folia- 
ceous,  and  folded  in  various  ways  (see  below),  whereas,  in  Erismeae, 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


they  are  thick,  fleshy,  and  not  or  scarcely  folded  (Baillon,  1874; 
Warming,  1875;  Boesewinkel  &  Venturelli,  1987;  Kawasaki,  in 
press).  The  cotyledons  of  Erismadelphm  are  plano-convex  and 
slightly  convolute  at  the  margins  (Boesewinkel  &  Venturelli,  1987), 
while  those  of  Erisma  are  narrowly  elongate  and  semi-cylindrical 
(Baillon,  1874),  plano-convex  (Boesewinkel  &  Venturelli,  1987),  or 
flat  (Stafleu,  1954). 

The  embryonic  cotyledons  have  been  described  as  unequal  in 
Vochysia  (Warming,  1875;  Stafleu,  1948)  and  Salvertia  (Stafleu, 
1 948),  although  no  comparable  information  was  given  for  Qualea  or 
Callisthene.  Unequal  could  mean  either  that  each  cotyledon  is 
asymmetrical  or  that  the  two  cotyledons  are  heterocotylar,  differing 
in  shape  or  size.  Cotyledons  are  asymmetric  in  two  species  of 
Vochysia  and  one  species  of  Qualea  but  symmetric  in  one  species  of 
Vochysia  and  Salvertia  (Table  1 ),  based  on  the  illustrations  of  the 
unfolded  embryo  in  Warming  ( 1 875).  In  V.ferruginea,  each  cotyle- 
don is  usually  asymmetric,  with  the  right  side  (viewed  from  the 
lower  abaxial  surface)  larger,  at  least  basally.  The  base  of  the  smaller 
side  tends  to  be  rounded,  and  that  of  the  larger  side  cordate,  as  in  the 
expanded  cotyledon  of  the  seedling  (Fig.  U).  The  embryonic  coty- 
ledons of  V.  ferruginea,  when  unfolded,  and  the  fully  expanded 
cotyledons  of  the  seedling  (Fig.  1 H,  P),  are  isocotylar.  The  larger 
side  of  one  cotyledon  opposes  the  smaller  side  of  the  other  on  the 
embryo  and  seedling,  hence,  the  two  cotyledons  are  identical  in 
shape,  not  mirror-images. 

Most  monographic  and  floristic  works  indicate  that  the  embry- 
onic cotyledons  of  Vochysiaceae  are  folded,  but  give  insufficient 
detail  to  characterize  the  complex  pattern  of  folding.  It  is  usually 
unclear  which  component  of  folding  is  described:  ptyxis,  the  folding 
of  an  individual  organ  such  as  a  leaf  in  the  bud  or  cotyledon  in  the 
seed,  or  vernation,  the  relationship  of  the  immature  organs  to  each 
other  (Bell,  1991).  Some  terms,  such  as  convolute,  are  used  for  both 
ptyxis  and  vernation  but  have  different  meanings  (Bell,  1991). 
Definitions  of  terms  vary  considerably  among  authors. 

In  Vochysia  ferruginea,  each  cotyledon  is  folded  four  times 
around  the  longitudinal  axis  of  the  embryo  (Fig.  1E-G).  The  folds 
(f,-f4)  are  parallel  to  the  radicle  but  not  parallel  to  the  midvein  of 
the  cotyledon  (Fig.  IF).  The  margin  of  the  larger  side  of  the 
cotyledon  (the  right  side  as  viewed  from  the  lower  abaxial  surface) 
is  on  the  exterior  of  the  embryo,  while  the  margin  of  the  smaller 
side  is  in  the  interior.  The  folded  smaller  side  is  completely  en- 
closed within  the  folded  larger  side.  The  inner  margin  is  narrowly 
bent  back  with  the  adaxial  surfaces  appressed  at  the  first  fold  (f,). 
Because  of  the  oblique  folding  of  the  cotyledon,  the  base  of  the 
larger  side  of  each  asymmetric  cotyledon  is  folded  around  the 
hypocotyl,  nearly  obscuring  it  (Fig.  IF).  Except  for  the  com- 
pressed nature  of  the  embryo  and  distinct  lines  of  folding,  ptyxis  is 
essentially  supervolute  (often  called  convolute)  but  oblique.  It  is 
not  conduplicate,  although  the  cotyledon  appears  to  be  folded 
around  the  midvein  in  cross-sections  through  the  middle  of  the 
embryo  (Fig.  1G).  Vernation  is  essentially  convolute  sensu  Bell 
(1991),  although  unusually  compressed.  Each  cotyledon  partially 
overlaps  the  other,  with  inner  margins  opposite  near  the  centre  of 
the  embryo  and  outer  margins  opposite  on  the  exterior  (Fig.  1G). 
Confusingly,  vernation  is  not  convolute  sensu  Radford  et  al. 
(1974),  because  one  rolled  or  supervolute  cotyledon  is  not  com- 
pletely enclosed  within  a  second  rolled  cotyledon. 

Convolute  vernation  and  supervolute  ptyxis  are  also  found  in 
Vochysia  pygmaea,  V.  tucanorum  Mart.,  and  Salvertia  convallari- 
dorea  A.  St.-Hil.,  as  seen  in  transverse  sections  of  the  embryo 
(Warming,  1875:  figs  16-11,  17-11;  Boesewinkel  &  Venturelli,  1987: 
fig.  10B).  The  embryos  are  less  compressed  and  angular  than  in  V. 
ferruginea,  and  the  inner  margins  of  the  cotyledons  overlap  rather 


than  being  folded  back  individually.  Embryos  of  V.  guatemalensis,  V. 
obscura  Warm.,  V.  tucanorum,  and  5.  convallaridorea  (Warming, 
1875:  figs  13,  1611,  1711;  Flores,  1993a:  fig.  9)  are  externally  similar 
to  that  of  V.  ferruginea,  and  the  position  of  the  outer  cotyledon 
margin  is  also  consistent  with  convolute  vernation.  Ptyxis  is  prob- 
ably oblique  is  most  species  because  the  hypocotyl  is  hidden,  as  in  V. 
ferruginea,  but  may  differ  in  V.  obscura,  given  that  the  hypocotyl  is 
well-exposed  in  spite  of  the  deeply  lobed  cotyledon  base.  Although 
folding  appears  somewhat  angular  in  section  because  the  seed  is 
compressed,  cotyledons  are  not  plicate  in  the  sense  of  fan-like.  Thus, 
there  is  one  basic  pattern  of  vernation  and  ptyxis  for  cotyledons  of 
Vochysia  and  Salvertia,  with  minor  variations,  whether  cotyledons 
are  asymmetric  or  not. 

Embryonic  cotyledons  of  the  genus  Vochysia  have  previously 
been  described  as  convolute  or  rolled  (Bentham  &  Hooker,  1867; 
Petersen,  1896;  Flores,  \993a,  b),  spirally  convolute  (Baillon,  1874), 
obliquely  spirally  convolute  (Warming,  1875),  spirally  folded 
(Boesewinkel  &  Venturelli,  1987),  and  spirolobal  (Stafleu,  1948). 
The  cotyledons  of  the  species  discussed  above  are  not  spirolobal,  a 
term  now  used  for  bent  embryos  with  incumbent  once-folded  coty- 
ledons (Duke,  1969;Radfordetal.,  1974). The  remaining  descriptions 
are  not  inconsistent  with  convolute  vernation  and/or  obliquely 
supervolute  (=  convolute)  ptyxis,  but  do  not  completely  characterize 
ptyxis  and  vernation. 

In  Qualea,  embryonic  cotyledons  have  been  described  as  convo- 
lute or  rolled  (Petersen,  1896;  Boesewinkel  &  Venturelli,  1987)  or 
convolute  and  plicate  (Warming,  1875;  Stafleu,  1953).  The  cotyle- 
dons of  Q.  grandiflora  Mart.  (Warming,  1875:  fig.  8-II)  are  more 
complexly  folded  than  in  Vochysia,  although  the  complete  pattern 
cannot  be  resolved  from  the  embryo  and  its  cross-section.  Supervolute 
ptyxis  and  convolute  vernation,  as  in  Vochysia,  can  be  eliminated, 
however,  and  folding  may  be  irregular  but  is  not  fan-like.  The 
exposed  margins  of  the  cotyledon  are  very  uneven,  almost  lobed.  In 
contrast,  the  exposed  cotyledon  margins  of  Q.  multiflora  Mart. 
(Warming,  1875:  fig.  7-II)  are  unlobed  and  parallel,  similar  to 
Vochysia  except  that  they  are  close  together  on  the  exterior  of  the 
embryo.  There  appears  to  have  been  some  folding  around  the 
longitudinal  axis  through  the  radicle,  but  the  specific  pattern  is  again 
unknown.  In  both  species  of  Qualea,  the  folded  cotyledons  nearly 
obscure  the  embryonic  axis.  Thus,  ptyxis  and  vernation  appear  very 
different  in  these  two  Qualea  species  compared  to  the  homogeneous 
pattern  observed  in  Vochysia. 

Embryonic  cotyledons  in  Callisthene  have  been  described  as 
plicate  and/or  convolute  (Warming,  1875;  Petersen,  1896;  Stafleu, 
1952;  Boesewinkel  &  Venturelli,  1987).  In  C.  major  Mart.  (Warm- 
ing, 1875:  fig.  3-II),  the  embryo  is  straight  and  compressed.  Each 
cotyledon  is  irregularly  folded,  appearing  more  wrinkled  (corru- 
gate) than  plicate  on  the  surface;  each  margin  is  involute,  folding 
back  with  the  adaxial  surfaces  touching.  The  folded  cotyledons  are 
pushed  together  for  most  of  their  length  and  width,  but  the  folds  are 
only  rarely  synchronous  between  the  two.  The  involute  margin  of  the 
first  cotyledon  is  exposed  at  the  edge  of  the  embryo  and  overlaps  the 
involute  but  hidden  margin  of  the  second,  whereas  the  exposed 
involute  margin  of  the  second  cotyledon  abuts,  rather  than  overlaps, 
the  hidden  margin  of  the  first.  No  standard  terms  describe  this 
pattern  of  folding,  but  corrugate-involute  ptyxis  and  semi-convolute 
or  valvate-convolute  vernation  might  be  used.  In  contrast  to  other 
genera  in  Vochysieae,  the  embryonic  axis  is  fully  exposed  and  the 
embryo  appears  almost  spathulate,  except  for  the  wrinkled  slightly 
overlapping  cotyledons. 

Embryos  of  Vochysia,  Qualea,  and  Callisthene  are  rich  in  lipids 
and  poor  in  starch,  those  of  Erisma  are  poor  in  lipids  and  starch,  and 
those  of  Erismadelphus  are  rich  in  lipids  and  starch  (Boesewinkel  & 


10 

Venturelli,  1987).  The  seeds  of  V.  hondurensis  are  rich  in  protein,  as 
well  as  lipids  (Blanche  et  al.,  1991). 

Seedling 

There  is  little  published  information  on  seedlings  and  saplings  of 
Vochysiaceae.  Only  Vochysia  duquei  Pilg.,  V.  guatemalensis,  and  V. 
ferruginea  (del  Amo,  1979;  Ricardi  et  al.,  1987;  Flores,  19930.  b; 
this  study)  have  been  described  in  detail.  Limited  data  are  available 
for  ten  additional  species  and  some  information  can  be  inferred  from 
the  embryos  of  five  species  (Tables  1-2).  In  Erismeae,  cotyledons  of 
Erismadelphus  are  cryptocotylar,  hypogeal,  and  food-storing.  Em- 
bryonic cotyledons  ofErisma  are  also  fleshy  and  food-storing,  but  it 
is  not  known  whether  these  are  cryptocotylar  or  phanerocotylar,  or 
epigeal  or  hypogeal,  after  germination.  No  other  information  on 
seedlings  is  available  for  Erismeae. 

In  Vochysieae,  cotyledons  are  all  phanerocotylar,  epigeal,  and 
foliaceous  (Table  1 ).  Where  known,  they  persist  after  the  first  leaves 
are  produced,  and  presumably  continue  to  photosynthesize.  In 
Vochysia  thyrsoidea  Pohl,  they  are  the  major  photosynthetic  organ 


20- 


10- 


15        20 
Length  (mm) 


Fig.  3     Relationship  of  cotyledon  length  and  width  in  Vochysiaceae. 
Vochysia  species:  circles,  V.  ferruginea  (Panama:  Garwood  1630; 
Garwood  2005;  Foster  s.n.;  Costa  Rica:  Flores,  19936);  squares,  V. 
guatemalensis  (Belize:  Garwood  4031;  Monro  868-871,  Monro  et  al. 
M4162X;  del  Amo,  1979;  Costa  Rica:  Flores,  1993a);  diamonds,  V.  sp. 
(Panama:  Hampshire  &  Whitefoord  269);  octagons,  V.  sp.  (Panama: 
Garwood  135);  inverted  triangles,  V  duquei  (Venezuela:  Ricardi  et  al., 
1987);  open  triangle,  V.  surinamensis  (Guyana:  Polak,  1992);  triangle 
with  black  circle,  V.  sp.  (French  Guiana:  Oldeman  &  Fundter,  1986); 
triangle  with  white  circle,  V  thyrsoidea  (Brazil:  Rizzini,  1965).  Other 
genera  (stars):  C,  Callisthene  major  (Brazil:  Labouriau  et  al.,  1964);  Q, 
Qualea  grandiflora  (Brazil:  Thomazini  et  al.,  1973).  Shading  of 
symbols  indicates  country  of  origin:  black,  Panama;  cross-hatching, 
Belize;  horizontal  hatching,  Costa  Rica;  vertical  hatching,  Mexico; 
white.  South  America.  Dashed  lines  represent  width  to  length  ratios  of 
2.0  (upper  line)  and  1.3  (lower  line).  Data  are  measurements  of  all 
seedlings  in  each  collection,  values  cited  in  references,  or  my  estimates 
from  illustrations  in  references  where  values  are  not  given.  Overlapping 
points  have  been  off-set  by  0.5-1  .0  mm  to  improve  readability. 


N.C.  GARWOOD 

for  many  months  (Rizzini,  1965).  Cotyledons  range  from  about  1  to 
3  cm  long  and  1  to  5  cm  wide  (Fig.  3).  They  are  smallest  in 
Callisthene  major  and  largest  in  an  unidentified  species  of  Vochysia 
(Fig.  3).  Within  Vochysia,  those  of  V.  ferruginea  are  the  smallest  (Fig. 
3).  Cotyledons  are  mostly  1.3  to  2.0  times  wider  than  long  in 
Vochysia,  Salvertia,  and  Callisthene,  but  only  0.8  to  1.3  times  wider 
than  long  in  Qualea,  and  broadest  above  the  middle  in  Vochysia  and 
Salvertia.  They  are  broadest  around  the  middle  in  Callisthene  and 
Qualea  (Table  1;  Fig.  3).  They  are  asymmetric  or  symmetric  in 
Vochysia  (Fig.  U  and  2 A,  E-F),  as  in  the  embryos,  but  this  may  be 
variable  within  some  species  or  collections  (Table  l).The  apex  of  Q. 
grandiflora  was  short-acuminate,  that  of  C.  major  ±  obcordate,  and 
those  of  Vochysia  truncate,  rounded,  retuse  or  rarely  obcordate  (Fig. 
U  and  2A,  E-F;  Table  1 ).  Venation  can  only  be  assessed  in  Vochysia, 
but  the  pattern  is  unusual.  The  midvein  forks  toward  the  apex, 
whether  the  apex  is  rounded  or  lobed,  and  two  strong  lateral  prima- 
ries diverge  from  the  base:  in  most  species,  the  lateral  primaries  join 
with  the  recurved  branches  of  the  midvein  to  form  triangular  to 
obovate  intercostal  areas  (Fig.  U  and  2A,  E-F;  Table  1).  Smaller 
veins  form  one  or  more  concentric  series  of  loops  around  the  central 
intercostal  areas. 

The  cotyledonary  stipules  in  the  three  species  of  Vochysia  de- 
scribed in  this  study  are  unusual  two-parted  structures  (Fig.  1K-M; 
Fig.  2C-D,  G).  The  lower  part  usually  consists  of  one  to  three  narrow 
projections  between  the  petioles,  usually  perpendicular  to  the 
hypocotyl,  situated  at  the  base  of  the  upper  (adaxial)  surface  of  the 
petiole  where  the  upper  ridge  or  wing  of  the  petiole  terminates.  The 
upper  part  consists  of  a  flattened  interpetiolar  scale-like  outgrowth 
just  above  the  petioles,  initially  folded  over  the  terminal  bud  but 
becoming  erect  and  visible  as  the  epicotyl  expands,  completely  or 
partially  divided  or  torn  into  two  to  five  parts  but  rarely  unlobed.  The 
apex  of  the  lower  and/or  upper  parts  often  appears  gland-like. 
Neither  part  appears  to  be  fused  over  the  petioles,  and  the  axillary 
buds  are  often  visible  above  the  cotyledonary  petiole.  Although 
there  was  some  intraspecific  variation,  the  stipules  of  V.  ferruginea 
usually  had  one  lower  projection  and  a  lobed  upper  part  and  those  of 
V.  guatemalensis  two  lower  projections  and  two  separate  upper 
parts.  The  reduced  stipules  of  V.  guatemalensis  have  also  been 
described  as  2-3  trichomes  of  a  glandular  nature  (Flores,  1993a),  but 
the  thick  3-lobed  pair  of  structures  illustrated  (fig.  7e)  appear  rather 
large  and  fleshy  for  trichomes  and  may  be  a  different  form  of  the 
variable  structures  described  in  this  study.  Two-parted  cotyledonary 
stipules  similar  to  those  of  Vochysia  have  not  been  seen  in  seedlings 
of  more  than  600  dicotyledonous  species  in  88  families  surveyed 
from  Panama  (Garwood,  in  prep.),  although  the  fleshy  cotyledonary 
stipules  of  Lafoensia  (Lythraceae)  are  similar  to  the  lower  stipular 
parts  of  Vochysia. 

Leaves  are  opposite  at  the  early  nodes  in  Vochysia,  Qualea,  and 
Callisthene  (Table  2).  In  Vochysia,  the  pair  of  leaves  at  the  first  node 
is  decussate  with  respect  to  the  cotyledons  and  leaves  at  the  second 
node  (Table  2).  In  Q.  grandiflora,  the  first  pair  of  fully  developed 
leaves  is  parallel  to  the  cotyledons  but  decussate  to  the  second  pair 
of  leaves  (Table  2);  a  pair  of  'prophylls'  between  the  cotyledon  and 
first  pair  of  leaves  (Thomazini  et  al.,  1973)  might  be  vestiges  of  the 
'missing'  pair  of  decussate  leaves  or  homologous  to  the  interpetiolar 
cotyledonary  stipules  of  Vochysia.  Leaves  remain  opposite  into  the 
adult  stage  in  V.  ferruginea,  but  become  3-  then  4-whorled  in 
V.  guatemalensis  sometime  in  the  sapling  stage.  The  initial  axis  of  C. 
major  appears  to  be  plagiotropic,  with  opposite  distichous  leaves 
smaller  than  the  cotyledons,  but  this  will  need  confirmation  because 
it  is  difficult  to  interpret  details  in  the  published  photograph  of  the 
seedlings  (Labouriau  et  al.,  1964:  fig.  26). 

Stem  characters  are  known  only  from  Vochysia.  The  collet,  the 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


11 


Table  1  Cotyledon  characters  in  seedlings  and  embryos  of  Vochysiaceae. 


Genus 
Species 

Source 

Emer- 
gence 

Posi- 
tion 

Devel- 

Persistence 

Width:Length 
Ratio 

Shape 

Base 

Apex  Symmetry     Venation      Stipules 

opment    node       months 

Vochysia 

V.  densiflora  Spruce 

18* 

- 

- 

F 

- 

1.7 

2 

5-7 

8 

A 

V  duquei  Pilg. 

15 

P 

E 

Fr 

2              >1 

1.6-1.9 

2-3,6 

4-6 

3-5 

S                7,2               + 

V.ferruginea 

6 

P 

E 

F 

3-8            >9 

1.3-1.9 

2 

3-6 

6-7 

A                1-3            f+f,g 

Mart. 

10 

- 

E 

- 

- 

- 

_ 

_ 

_ 

_                  _                _ 

4b 

P 

E 

F 

2 

1.6 

2 

6 

5 

8^4                 7                 f 

V.  guatemalensis 

6 

P 

E 

F 

3 

1.2-1.8 

2 

3-6 

3-5 

A                1-3            s+s,g 

Donn.  Sm. 

10 

- 

E 

- 

-              _ 

_ 

_ 

_ 

_ 

_                  _                _ 

4a 

P 

E 

F 

/ 

1.2-1.3 

2 

6 

4 

A                7,4              s,g 

1,5 

P 

E 

F 

6 

1.5 

2 

4-5 

4 

A                 1,2 

V.  obscura  Warm. 

18* 

- 

- 

F 

-              - 

1.4 

2 

7 

4-5 

S 

V  surinamensis  Stafleu 

14 

P 

E 

n 

6 

2.7 

1 

— 

3-4 

—                  —                _ 

V.  thyrsoidea  Pohl 

13 

- 

E 

- 

=  12 

- 

- 

- 

- 

_                  _                _ 

16 

P 

E 

Fr 

1             >12 

=  7-2 

7 

2 

4 

_                  _                _ 

V.  tucanorum  Mart. 

18* 

- 

- 

F 

-              - 

7.6 

2 

6-7 

3^t 

A 

V.  sp.  (Brazil) 

2 

P 

E 

F 

-              - 

- 

1 

_ 

- 

—                  _                _ 

V.  sp.  (Fr.  Guiana) 

11 

P 

E 

F 

- 

2.7 

2 

4 

5 

5                2,5 

V.  sp.  (Panama) 

6 

P 

E 

F 

1 

1.4-2.1 

2 

3-5 

4-6 

S,A               1-3            s+f,g 

Salvertia 

S.  convallaridorea 

18* 

- 

- 

F 

_              - 

1.6 

2 

2-3 

4-5 

S 

A.  St.-Hil. 

Qualea 

Q.  grandiflora 

12 

P 

E 

F 

2              >6 

- 

- 

- 

- 

_                  _                _ 

Mart. 

3 

P 

E 

F 

_               4 

- 

- 

— 

_ 

_                  _                _ 

17 

P 

E 

F 

2              >6 

1.1 

5 

3^f 

7 

Al                                    P 

Q.  multiflora  Mart. 

18* 

- 

- 

F 

- 

0.8-1.3 

4 

3 

9 

A 

Callisthene 

C.  major  Mart. 

8 

P 

E 

Fl 

1-3? 

1.2-1.5 

3 

- 

5-6 

_                   _                 _ 

18* 

- 

- 

F 

- 

1.2-1.6 

- 

- 

- 

-                   -                 - 

Erisma 

E.  spp. 

7* 

- 

- 

R 

- 

- 

- 

- 

- 

- 

Erismadelphus 

E.  exsul  Mildbr. 

9 

C 

H 

R 

- 

- 

- 

- 

- 

_                   _                 _ 

E.  sp. 

9 

C 

H 

R 

- 

- 

- 

- 

- 

Source:   1,  del  Amo  1979;  2,  Duarte  1979;  3,  Felippe  &  Dale  1990;  4a,b,  Flores  1993a,  b;  5,  Gaos  1978;  6,  Garwood,  this  study;  7,  Kawasaki,  in  press;  8.  Labouriau  et  al.  1964: 

9,  Miquel  1987;  10,  Moreira  &  Arnaez  1994;  11,  Oldeman  &  Fundter,  1986;  12,  Paulilo  et  al.  1993;  13,  Paviani  &  Haridasan  1988;  14,  Polak  1992;  15,  Ricardi  et  al.  1987;  16, 

Rizzini  1965;  17,  Thomazini  et  al.  1973;  18,  Warming  1875.   *,  all  characters  based  only  on  the  embryonic  cotyledon  in  the  mature  seed. 

Emergence:  P,  phanerocotylar;  C,  cryptocotylar. 

Position:  E,  epigeal;  H,  hypogeal. 

Development:  F,  foliaceous;  R,  fleshy,  presumably  with  stored  reserves;  r,  somewhat  fleshy  or  succulent,  with  reserves  or  stored  water. 

Persistence:  measured  as  i)  the  last  recorded  leaf  node  to  produce  leaves  while  the  cotyledons  are  still  present  (e.g.  node=3.  cotyledons  still  present  when  leaves  produced  at 

node  3)  or  ii)  the  length  of  time  [months]  that  cotyledons  persist  after  germination.  These  measures  underestimate  persistence  when  only  the  youngest  seedling  stages  have 

been  studied. 

Width:Length  Ratio:  width  divided  by  length. 

Shape:  1,  obtriangular  to  shallowly  obtriangular,  including  'triangular';  2,  depressed  obovate;  3,  transversely  oblong;  4,  very  widely  obovate;  5,  transversely  widely  elliptic  to 

transversely  widely  oblong;  6  transversely  reniform,  rectangular-reniform. 

Base  &  apex:  1,  short-acuminate;  2,  acute;  3,  rounded;  4,  truncate;  5,  retuse  to  emarginate;  6,  cordate,  obcordate,  2-lobed;  7,  2-cleft  or  -parted;  8,  3-lobed;  9,  irregular.  When 

blade  is  asymmetric,  the  base  and/or  apex  may  be  unequal  on  the  two  sides  (e.g.  half-cordate,  half-rounded)  or  oblique  (e.g.  obliquely  truncate). 

Symmetry  of  blade:  A,  asymmetrical;  S,  symmetrical. 

Venation:  1,  branches  of  forked  central  primary  vein  join  lateral  primaries  forming  a  pair  of  obovate  to  triangular  intercostal  areas;  2,  palmate  or  3-nerved;  3,  basal  acro- 

brochidodromous;  4,  basal  actinodromous;  5,  branches  of  central  primary  vein  forked;  lateral  primaries  each  form  a  closed  loop  apically,  terminating  near  but  not  joined 

directly  to  the  tips  of  the  central  vein. 

Stipules:  s,  usually  separate;  f,  usually  fused  and  interpetiolar;  f+f,  f+s,  and  s+s,  two  structures  (fused  or  separate)  present  at  cotyledonary  node  (first  is  lower  structure,  second 

is  upper  structure);  g,  gland-like;  +,  present;  P  'prophylls'  present  above  cotyledons. 

-,  Information  not  available. 

Characters  in  italics  were  inferred  or  estimated  from  illustrations  in  the  citations,  not  from  the  text  of  the  original  author.  Where  authors  identified  the  cotyledons  as  foliaceous, 

it  was  assumed  that  they  were  also  phanero-epigeal. 


junction  of  the  radicle  and  hypocotyl,  is  annulate  in  at  least  three 
species  (Table  2;  Fig.  IF).  The  hypocotyl,  epicotyl,  and  later 
internodes  are  usually  square,  with  ridges  or  wings  on  each  corner 
(Table  2).  Ridges  on  the  hypocotyl  continue  without  interruption 
into  the  ridges  on  the  lower  margins  of  the  cotyledonary  petioles  in 
V.ferruginea  and  V.  guatemalensis  (Figs  IK,  2G;  Flores,  1993a,  b). 
In  contrast,  the  wings  or  ridges  on  the  epicotyl  and  later  internodes 


terminate  in  small  narrowly  triangular  to  linear  stipules  at  the  top  of 
each  internode  (Fig.  1O,  S;  Table  2),  with  the  inner  margin  and  base 
of  the  stipules  apparently  extending  above  the  petiole  but  not 
attached  to  it.  The  winged  stems  do  not  originate  directly  from 
decurrent  leaf  bases,  as  sometimes  suggested,  because  the  attenuate 
base  of  the  blade  narrows  into  a  medial  pair  of  short  thin  ridges  on 
the  upper  (adaxial)  surface  of  the  petiole,  interrupting  the  lamina 


12 


N.C.  GARWOOD 


Table  2  Root,  stem,  and  leaf  characters  in  seedlings  of  Vochysiaceae. 


Genus 
Species 

Source 

Collet 

Hypocotyl 

Epicotyl 

Internodes 

Phyllotaxis 

Leaf 
stipules 

Nodes  1-2 

Later  nodes 

Vochysia 

V  duquei 

15 

a 

s 

s 

0 

-,v 

+ 

V.  ferruginea 

6 

a 

s,w 

s,w 

s,w                         o,d 

o,d 

+,s 

4b 

c 

s 

- 

o,d 

o,d 

- 

V.  guatemalensis 

6 

a 

s,w 

s,w 

s,w                        o,d 

o,w 

+,s 

4a 

c 

s,w 

s,w 

o,d 

o,w 

- 

1,5 

c 

c,w 

s,w 

s,w                         o,d 

0 

+,i 

V.  surinamensis 

14 

- 

- 

- 

S,W                                       0 

o,v 

- 

V.  thyrsoidea 

13 

c 

t 

- 

- 

- 

- 

V.  sp.  (Panama) 

6 

- 

s,w 

w 

0 

- 

- 

Qualea 

Q.  grandiflora 

17 

- 

- 

- 

o,p 

- 

— 

Callisthene 

C.  major 

8 

- 

o 

a? 

Sources  and  authorities  for  names:  see  Table  1. 

Collet  (junction  of  hypocotyl  and  primary  root):  a,  annulate  ridge  present;  c,  constriction,  abrupt  change,  or  other  feature  present,  but  nature  of  structure  not  discernable  from 

illustration. 

Hypocotyl.  Epicotyl,  and  Internode:  cross-sectional  shape  of  hypocotyl,  epicotyl,  and  early  internodes  (about  first  ten),  respectively:  s,  square  to  rectangular;  c,  circular;  t, 

tuberous  and  circular,  at  least  when  older;  w,  4  wings  or  ridges  present. 

Phyllotaxis: 

Nodes  1-2:  o,  leaves  at  first  node  opposite;  d,  pair  of  leaves  at  first  node  decussate  to  both  cotyledons  and  pair  of  leaves  at  second  node;  p,  pair  of  leaves  at  first  node  parallel  to 

cotyledons  and  decussate  to  pair  of  leaves  at  second  node. 

Later  nodes:  o,  leaves  opposite  and  decussate  (at  least  to  node  5,  sometimes  reported  up  to  node  20);  d,  leaves  opposite  and  decussate  on  larger  seedlings,  saplings,  and  adults; 

w,  leaves  whorled  on  larger  seedlings  or  saplings  (and  adults),  with  three  or  four  leaves  per  node,  size  of  juveniles  at  time  of  transition  from  opposite  to  whorled  leaves 

unknown;  v,  juvenile  leaves  unknown,  but  adult  leaves  whorled;  a,  axis  becoming  plagiotropic  with  pairs  of  leaves  distichous. 

Leaf  stipules:  +,  small,  narrowly  triangular  to  linear  stipules  present  from  the  first  node;  i,  stipules  intrapetiolar;  s,  stipules  separate. 

-,  Information  not  available. 

Characters  in  italics  were  inferred  or  estimated  from  illustrations  in  the  citations,  not  from  the  text  of  the  original  author. 


between  the  blade  and  wings.  It  is  unclear,  however,  whether  the 
petiolar  ridges  are  joined  to  the  lateral  extensions  of  the  stipules. 

Striking  ontogenetic  changes  are  seen  in  hair  morphology  in 
Vochysia  ferruginea  (this  study).  The  epicotyl  and  first  leaves  are 
only  sparsely  pubescent  below,  but  stems  and  leaves  at  higher 
nodes  become  progressively  more  densely  pubescent.  Hairs  on 
early  internodes  and  lower  leaf  surfaces  are  nearly  sessile, 
appressed,  and  T-shaped,  with  straight  arms  of  equal  length;  those 
on  later  leaves  and  stems  are  short-stalked,  ascending  to  erect,  2- 
branched,  with  the  longer  arm  curved  to  coiled  and  about  eight 
times  longer  than  the  shorter  arm.  Hairs  on  the  adult  leaves  are 
similar,  except  the  branches  are  even  more  unequal  in  length 
(Garwood,  pers.  obs.)-  On  seedlings  to  adults  of  V.  ferruginea, 
hairs  on  the  midrib  above  are  always  simple.  In  Vochysia  sp. 
(Garwood  135  and  Hampshire  &  Whitefoord  269),  hairs  on  the 
epicotyl  and  eophylls  were  similar  to  those  on  V  ferruginea.  In 
contrast,  hairs  on  the  young  expanding  leaves  of  V  guatemalensis 
were  simple  and  basifixed,  but  these  were  usually  sparse  or  absent 
on  mature  leaves  (Flores,  1993a;  this  study).  All  parts  of  V  duquei 
are  glabrous  (Ricardi  et  al.,  1987). 

Seedlings  from  the  Brazilian  cerrado  have  a  number  of  special 
morphological  adaptations  to  the  dry  cerrado  environment  (Labouriau 
et  al.,  1964;  Rizzini,  1965;  Paviani  &  Haridasan,  1988).  Vochysia 
thyrsoidea  has  ±  fleshy  succulent  cotyledons,  a  thick  watery 
hypocotyl  which  is  mostly  buried,  minute  eophylls  which  remain 
hidden  between  the  cotyledons,  greater  growth  of  the  taproot  rather 
than  shoot  in  the  first  few  months,  and  a  swollen  subterranean  organ 
which  forms  from  the  hypocotyl  and  upper  part  of  the  taproot  in  the 
first  two  years.  Qualea  grandiflora  also  forms  a  swollen  under- 
ground organ. 


Phylogeny 

There  have  been  no  phylogenetic  analyses  of  relationships  within 
Vochysiaceae.  Since  the  completion  of  Stafleu's  ( 1 948- 1 954)  mono- 
graph more  than  40  years  ago,  the  numbers  of  collections  and 
species  described  have  increased,  but  major  systematic  work  has 
been  limited  to  elevating  one  section  of  Qualea  to  Ruizterania 
(Marcano-Berti,  1969)  and  monographing  Erisma  (Kawasaki,  in 
press).  Fruit  and  seed  characters  have  long  been  important  in  delim- 
iting genera  within  the  family,  but  the  utility  of  seedling  characters 
has  been  untested  because  seedlings  have  not  been  available  for 
study.  The  potential  of  the  new  seedling  and  seed  characters  de- 
scribed above  to  contribute  to  phylogenetic  and  systematic  studies  is 
discussed  here.  If  such  traits  look  promising,  data  on  more  species 
should  be  collected. 

Phanero-epigeal  seedlings  are  reported  only  from  Vochysieae, 
and  crypto-hypogeal  seedlings  from  Erismeae,  following  the  current 
tribal  division.  In  Vochysieae,  shape  characters  of  the  foliaceous 
cotyledon  fell  into  three  groups  corresponding  to  Qualea,  Callis- 
thene, and  Vochysia/Salvertia,  although  some  variation  occurred  in 
all  groups.  Symmetry  of  the  cotyledons  varied  most  in  Vochysia,  and 
vernation  and  ptyxis  of  the  embryonic  cotyledons  inQualea.  Several 
traits  generally  uncommon  in  dicotyledon  seedlings  occur  together 
in  Vochysia,  including  unusual  cotyledonary  stipules  and 
cotyledonary  venation,  annulate  collet,  and  winged  hypocotyls  and 
early  internodes,  but  these  have  not  been  assessed  in  many  species  or 
other  genera.  Supra-cotyledonary  prophylls,  also  unusual  in  dicoty- 
ledon seedlings,  were  reported  in  one  species  of  Qualea.  The 
existence  of  considerable  variation  in  cotyledon  characters,  and  the 
presence  of  many  unusual  seedling  characters  in  this  small  sample  of 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


13 


about  20  species  (Tables  1-2),  suggests  further  study  of  other  taxa 
would  be  profitable. 

Ontogenetic  changes  from  the  seedling  to  adult  stages  can  also  be 
important  phylogenetic  characters.  In  Vochysia,  leaves  of  adults  are 
either  opposite  or  3-  or  4-verticillate,  but  the  first  seedling  leaves  of 
all  species  described  are  opposite  (Table  2).  In  V.  guatemalensis, 
leaves  remain  opposite  at  early  seedling  nodes,  but  gradually  be- 
come 3-  then  4-verticillate  in  larger  saplings  (Flores,  1993o). 
Although  presence  of  opposite  seedling  leaves  in  all  taxa  would  not 
be  phylogenetically  informative,  the  timing  of  the  switch  from 
opposite  to  verticillate  leaves  may  provide  a  useful  character.  Adult 
leaves  are  also  verticillate  in  Salvertia  and  some  species  of  Erisma, 
but  opposite  in  other  taxa  (Stafleu,  1948-1954). 

Pronounced  ontogenetic  changes  in  hair  morphology  were  ob- 
served in  Vochysia  ferruginea,  but  not  in  V.  guatemalensis.  Because 
studies  of  ontogenetic  changes  of  trichomes  on  leaves  of  seedlings 
and  adults  have  helped  identify  groups  of  species  within  the  large 
and  complex  genera  of  Solanum  and  Rhododendron  (Seithe  1979, 
1980;  Seithe  &  Anderson,  1982),  further  study  would  be  warranted. 
Unfortunately,  the  morphology  of  hairs  in  Vochysia  is  poorly  known. 
They  have  been  described  as  simple  or  sometimes  unicellular  and  2- 
branched  (Warming,  1875;  Metcalfe  &  Chalk,  1950);  hairs  have 
arms  of  equal  length  and  short  stalks  in  at  least  one  species 
(Shanmukha  Rao  &  Vijayalakshmi  Sarma,  1992).  Stafleu  (1952) 
indicated  that  simple  and  biramose  hairs  occurred  in  Vochysieae,  but 
did  not  specifically  describe  those  of  Vochysia  (Stafleu,  1948), 
although  frequent  references  to  'peculiar  indumentum'  or  'charac- 
teristic indumentum'  suggests  that  their  morphology  is  unusual. 

An  examination  of  adult  specimens  of  Vochysia  at  BM  indicates 
that  species  with  2-branched  or  simple  hairs  are  both  common 
(Table  3).  The  distribution  of  simple  and  branched  hairs  is  clearly  not 
random  with  respect  to  the  sections  of  Vochysia  delimited  by  Stafleu 
(1948).  Branched  hairs  are  especially  abundant  in  subsection 
Ferrugineae,  which  contains  V.  ferruginea.  There  was  considerable 
variation  among  species  in  the  morphology  of  the  branched  hairs  of 
the  adult  stage,  ranging  from  nearly  sessile  appressed  T-shaped  hairs 
with  straight  branches  of  equal  length,  to  long-stalked  erect  2- 
branched  hairs  with  curled  arms  of  equal  or  unequal  length  (Garwood, 
pers.  obs.).  In  some  species,  the  smaller  arm  was  so  reduced  in 
length  that  the  hair  could  easily  be  misinterpreted  as  simple,  sug- 
gesting that  at  least  some  of  the  simple  hairs  in  Vochysia  might  be 
derived  from  much-reduced  branched  hairs,  as  occurs  in  Malpighi- 


aceae  (Anderson,  1990).  The  range  of  hair  morphology  from  the 
seedling  to  adult  stages  in  V.  ferruginea  spans  nearly  the  entire  range 
of  branched  hairs  seen  in  the  genus.  Therefore,  examining  the 
ontogeny  of  branched  hairs  might  reveal  similarities  in  the  early 
stages  that  would  be  phylogenetically  informative  in  comparison 
with  other  species  of  Vochysia,  or  other  genera.  Hairs  are  also  simple 
or  2-branched  in  Qualea  (Stafleu,  1953),  but  stellate  in  Erisma 
(Warming,  1875;  Stafleu,  1954),  and  simple  in  Callisthene  (Stafleu, 
1952),  Salvertia  (Garwood,  pers.  obs.),  and  Erismadelphus 
(Garwood,  pers.  obs.). 

Seed,  fruit,  and  seedling  characters  also  bear  on  the  higher  order 
relationships  of  the  family.  Vochysiaceae  has  usually  been  included 
in  the  Polygalales  with  Polygalaceae,  Trigoniaceae,  Malpighiaceae, 
Xanthophyllaceae,  Tremandaceae,  and  Krameriaceae  (Dahlgren, 
1980;Takhtajan,  1980;  Cronquist,  1981).  Boesewinkel  &  Venturelli 
(1987)  used  anatomy  of  the  seed  coat  to  reassess  the  position  of  the 
family  in  Polygalales.  They  concluded  that  only  Polygalaceae  and 
Xanthophyllaceae  should  be  retained  in  the  Polygalales,  and  that  the 
relationship  of  Vochysiaceae  and  the  remaining  families  needed 
further  study.  Reviewing  numerous  floral  and  vegetative  characters 
in  Malpighiaceae,  Anderson  (1990)  decided  that  Malpighiaceae  was 
most  similar  to  Vochysiaceae  and  Trigoniaceae  and  least  similar  to 
Polygalaceae  and  Krameriaceae.  The  presence  of  2-branched  uni- 
cellular hairs  in  Vochysiaceae,  Trigoniaceae,  and  Malpighiaceae, 
and  their  absence  in  Polygalaceae,  also  supports  this  grouping 
(Shanmukha  Rao  &  Vijayalakshmi  Sarma,  1992). 

Recent  molecular  analyses  of  the  angiosperms  using  the 
chloroplast  gene  rbcL  also  supported  the  break-up  of  the  Polygalales, 
but  not  the  grouping  of  Vochysiaceae  with  Malpighiaceae  and 
Trigoniaceae  (Chase  et  al.,  1993).  Instead,  Vochysiaceae,  repre- 
sented by  Qualea,  fell  within  the  Myrtales.  A  more  detailed  study  of 
Myrtales  using  the  same  gene  confirmed  the  Myrtalean  affinities  of 
Vochysiaceae,  represented  by  Vochysia,  Erisma,  and  Qualea  (Conti 
et  al.,  1996).  Vochysiaceae  was  sister  group  to  the  Myrtaceae  and 
these,  with  Heteropyxidaceae  and  Psiloxylacecae,  formed  the  sister 
clade  to  that  containing  Melastomataceae,  Memecylaceae,  and  other 
families.  Work  with  chloroplast  gene  matK  also  supports  placement 
of  Vochysiaceae  in  Myrtales,  but  it  falls  between  Myrtaceae  (with 
Heteropyxidaceae  and  Psiloxylacecae)  and  Melastomataceae  (with 
Memecylaceae)  (P.  Wilson,  pers.  comm.). 

Previous  systematic  treatments  based  on  morphology  have  never 
included  Vochysiaceae  in  the  Myrtales  (Conti  et  al.,  1996),  regard- 


Table  3      Distribution  of  simple  and  branched  hairs  in  Vochysia. 


Section 


Subsection 


Number  of  species 


Described 

K,,  ^'i:,!!  .11  /IG/1Q\ 

Examined 

Type  of  hair 

by  Matleu  (iy4o) 

for  study 

Simple 

Branched 

Unknown 

Vochysiella 

Decorticanthes 

10 

6 

6 

0 

0 

Calophylloideae 

10 

7 

6 

0 

1 

Ciliantha 

Micranthae 

11 

9 

8 

0 

1 

Lutescentes 

23 

13 

6 

4 

3 

Discolores 

3 

2 

0 

2 

0 

Chrysophyllae 

1 

1 

1 

0 

0 

Megalanthae 

5 

3 

3 

0 

0 

Ferrugineae 

31 

22 

1 

18 

3 

Pachyantha 

- 

3 

0 

— 

— 

~ 

Type  of  hair:  simple,  only  simple  unbranched  hairs  present  on  leaves,  stems,  and  reproductive  parts;  branched,  some  or  all  of  hairs  present  branched,  including  T-shaped  hairs; 
unknown,  specimens  glabrous  or  hairs  too  small  or  dense  to  characterize.  Leaves,  stems,  and  inflorescences  of  one  to  several  specimens  of  each  species  at  BM  monographed  by 
Stafleu  (1948)  were  examined  under  a  40x  dissecting  microscope. 


14 

ing  this  relationship  as  so  remote  that  the  presence  of  two  of  the 
most  definitive  Myrtalean  characters,  vestured  pits  in  the  second- 
ary xylem  and  bicollateral  vascular  bundles  in  the  primary  stem, 
was  not  considered  significant  (van  Vliet  &  Baas,  1984).  In  retro- 
spect, many  vegetative  characters  of  Vochysiaceae  are  equally  at 
home  in  Myrtales  (sensu  Dahlgren  &  Thorne,  1984)  as  in 
Polygalales,  including  opposite  or  verticillate  leaves  with  entire 
margins,  brochidodromous  venation,  and  stipules.  The  highly 
zygomorphic  flowers  with  one  fertile  stamen  are,  however,  unu- 
sual in  the  order  (Conti  et  al.,  1996).  When  a  thorough 
morphological  analysis  of  the  relationship  of  Vochysiaceae  within 
Myrtales  is  undertaken,  several  seedling  and  seed  characters  de- 
scribed for  Vochysiaceae  deserve  further  study. 

Vochysiaceae  share  other  vegetative  characters  with  Myrtales, 
including  the  occasional  presence  of  2-branched  unicellular  or 
tufted  hairs  in  Combretaceae,  Lythraceae,  and  Myrtaceae 
(Metcalfe  &  Chalk,  1950).  Cotyledonary  stipules,  generally  un- 
common in  dicotyledons,  occur  as  rudimentary  glandular 
structures  in  some  Lythraceae,  Onagraceae,  and  Myrtaceae,  and 
as  2-  to  3-lobed  structures  or  glandular  processes  near  the  basal 
margins  of  the  blade  in  other  groups  of  Myrtaceae  (Weberling, 
1984).  The  latter  type  might  be  homologous  to  the  unusual 
cotyledonary  stipules  of  Vochysia  (Flores,  1993«;  this  study), 
which  are  composed  of  an  upper  lobed  part  and  lower  possibly 
glandular  part. 

Wings  are  well-developed  on  adult  stems  in  some  Lythraceae 
and  Penaeaceae,  and  often  develop  auricles  at  the  apex,  which  are 
distinct  from  the  rudimentary  stipules  found  closer  to  the  petioles 
(Weberling,  1984).  In  Lythraceae,  auricled  wings  are  already 
present  at  the  seedling  stage  in  Lafoensia  (Garwood,  in  prep.). 
Wings  are  well-developed  on  seedlings  of  Vochysia,  although  usu- 
ally reduced  to  ribs  on  adults  (Stafleu,  1948;  Garwood,  pers. 
obs.).  As  the  small  stipules  of  Vochysia  occur  at  the  apex  of  the 
wings,  they  may  be  homologous  to  the  auricles  at  the  apex  of  the 
wings  of  these  Myrtales,  rather  than  the  small,  often  dissected, 
stipules  characteristic  of  Myrtales  (Weberling,  1984).  If  so, 
Vochysia  may  lack  true  stipules.  Herbarium  specimens  of  V. 
ferrugineae  and  V.  guatemalensis  seedlings  were  carefully  exam- 
ined for  the  presence  of  dissected  or  rudimentary  stipules  over  the 
petioles,  but  none  were  seen  (Garwood,  pers.  obs.).  The  occa- 
sional presence  of  a  minute  shrivelled  structure,  however,  suggests 
that  actively  growing  seedlings  should  be  examined  to  verify 
whether  minute  caducous  stipules  are  present  but  hidden  by  the 
small  erect  expanding  leaves.  In  seedlings  of  Myrtaceae,  auricles 
are  lacking  on  the  winged  epicotyl  and  later  internodes  of  Psidium 
and  Calycolpus  and  on  the  winged  hypocotyl  of  Myrcia,  and 
wings  are  lacking  in  Eugenia,  but  all  have  rudimentary  gland-like 
stipules  (Garwood,  in  prep.). 

Seeds  of  Vochysiaceae  and  Myrtales  generally  lack  endosperm 
and  arils  and  have  straight  or  slightly  curved  embryos  (Cronquist, 
1981;  Dahlgren  &  Thorne,  1984).  A  fibrous  exotegmen  and  an 
endotestal  crystal  layer  are  found  in  Qualea  and  some  Lythraceae, 
Combretaceae,  and  Onagraceae,  but  the  exotegmen  lacks  fibres  and 
is  crushed  in  Vochysia  and  some  Melastomataceae  and  Myrtaceae 
(Corner,  1976;  Dahlgren  &Thorne,  1984;  Boesewinkel  &Venturelli, 
1987).  The  embryonic  cotyledons  are  variously  folded  in 
Vochysiaceae  and  many  Myrtales,  with  folding  frequently  described 
as  'spiral'  in  Vochysia  as  well  as  Combretaceae  and  Oliniaceae 
(Dahlgren  &  Thorne,  1984).  As  brief  descriptions  of  vernation  and 
ptyxis  in  Vochysiaceae  are  generally  incomplete  or  incorrect  (see 
above),  further  detailed  studies  should  be  carried  out  throughout  the 
order. 


N.C.  GARWOOD 


Ecology 


Seed  germination  has  been  the  most  frequently  studied  aspect  of  the 
seed  and  seedling  ecology  of  Vochysiaceae.  Germination  is  usually 
rapid,  occurring  1  week  to  3  months  after  seed  dispersal  or  sowing  in 
species  of  Vochysia,  Salvertia,  and  Qualea  from  both  cerrado  and 
rainforest  (Rizzini,  1965;  Thomazini  et  al.,  1973;  Macedo,  1977; 
Duarte,  1979;  Sabatier,  1985;  Paviani  &  Haridasan,  1988;  de  Godoy 
&  Felippe,  1992a;  Moreira  &  Arnaez,  1994;  Flores  1993a,  b;  this 
study;  M.  Bustamante,  pers.  comm.;  P.E.  Oliveira,  pers.  comm.;  P. 
Wilson,  pers.  comm.).  Chemical  or  mechanical  scarification  of  the 
seed  coat,  or  its  complete  removal,  may  hasten  germination  in  Q. 
grandiflora  (Felippe,  1990;  Felippe  &  Dale,  1990),  but  is  not 
required  (M.  Bustamante,  pers.  comm.;  P.E.  Oliveira,  pers.  comm.; 
P.  Wilson,  pers.  comm.).  Germination  is  not  inhibited  by  dark  or 
shaded  conditions  in  Q.  retusa  Spruce  ex  Warm.,  Q.  grandiflora,  or 
V.ferruginea  (Macedo,  1977;  Felippe,  1990;  this  study),  although  it 
can  be  somewhat  depressed  in  direct  sunlight  (Flores,  19936). 
Recently  dispersed  seeds  of  V.  maxima  Ducke  buried  under  the  leaf 
litter  either  lost  viability  or  had  germinated  within  2  months  (Viana, 
1990).  Thus,  there  is  no  indication  of  any  type  of  seed  dormancy  in 
the  family,  and  seeds  should  form  transient,  rather  than  persistent, 
seed  banks  under  natural  conditions. 

Under  artificial  conditions,  seeds  of  Qualea  cordata  Spreng., 
Vochysia  guatemalensis,  and  V.ferruginea  retained  high  viability  for 
at  least  2-3  months  when  stored  at  room  temperature  and  humidity 
(de  Godoy  &  Felippe,  1992a;  Flores,  1993a,  b),  in  spite  of  a 
relatively  high  initial  moisture  content  in  Vochysia  (23-32%,  fresh 
weight  basis).  Reducing  the  moisture  content  extended  viability  of 
the  apparently  orthodox  seeds,  although  they  were  intolerant  of 
chilling  (Flores,  \993a).  The  potential  to  store  seed  should  promote 
the  ongoing  silvicultural  development  of  the  family. 

Seeds  of  all  species  are  wind-dispersed,  as  indicated  by  the 
morphology  of  seeds  of  Vochysieae  and  indehiscent  fruits  of 
Erismeae,  except  Erisma  calcaratum  (Link)  Warm,  from  the  Ama- 
zonian varzea,  which  is  water-dispersed  (Stafleu,  1954).  In  tropical 
forests,  seeds  of  Vochysia  maxima  are  dispersed  near  the  parent, 
mostly  <  30  m  from  the  trunk  (Viana,  1990),  seedlings  of  V.  maxima 
and  V.  ferruginea  persist  in  the  shaded  understorey  (Viana,  1990; 
Garwood,  pers.  obs.),  but  growth  is  fast  only  in  treefall  gaps  and 
larger  openings  (Viana,  1990;  Boucher  et  al.,  1994).  In  plantations, 
seedlings  of  V.  guatemalensis  and  V.ferruginea  can  reach  2-3  m  tall 
within  three  years  (Flores  1993a,  b).  Thus,  tall  canopy  Vochysia 
appear  to  be  gap  opportunists,  regenerating  from  a  persistent  seed- 
ling bank  following  disturbances  throughout  the  year.  Regeneration 
of  Qualea  may  follow  a  different  pattern.  Following  a  severe  hurri- 
cane, most  of  the  severely  defoliated  adults  of  Q.  paraensis  Ducke 
survived  but  there  was  no  regeneration  from  seedlings  or  saplings;  in 
contrast,  all  the  equally  abundant  and  severely  defoliated  adults  of  V. 
ferruginea  died,  but  there  was  extensive  regeneration  from  seedlings 
and  saplings  (Boucher  et  al.,  1994).  If  this  pattern  is  typical,  species 
of  Vochysia  may  be  more  amenable  to  sustainable  management  or 
plantation  development  than  Qualea. 

In  the  drier  and  more  open  cerrado,  seedlings  of  Qualea 
grandiflora,  Q.  cordata,  and  Vochysia  thyrsoidea  appear  to  grow 
inherently  slowly,  with  neither  extra  moisture  nor  nutrients  pro- 
moting rapid  growth  (Rizzini,  1965;  de  Godoy  &  Felippe,  19926; 
Paulilo  et  al.,  1993;  Felippe  &  Dale,  1990).  Although  the  basis  for 
the  slow  growth  is  not  understood,  many  species  develop  special 
morphological  adaptations  to  the  cerrado  environment,  such  as 
large  subterranean  storage  organs  and  thick  succulent  cotyledons 
(Labouriau  et  al.,  1964;  Rizzini,  1965;  Paviani  &  Haridasan, 
1988). 


SEEDLINGS,  FRUITS  AND  SEEDS  OF  PANAMA:  VOCHYSIACEAE 


15 


ACKNOWLEDGEMENTS.  I  am  especially  grateful  to  Margaret  Tebbs  for 
illustrations,  Simon  Thornton-Wood  for  cross-sectional  diagrams  of  seeds, 
Ruth  Dempsey  for  composition  of  the  figures,  and  Eduardo  Sierra  for 
technical  assistance  in  Panama.  I  also  thank  C.J.  Humphries  for  encourage- 
ment and  advice;  S.  Knapp,  D.M.  Williams  and  an  anonymous  reviewer  for 
helpful  comments  on  the  manuscript;  M.  Beasley  for  bibliographic  assist- 
ance; INRENARE  for  permission  to  work  in  Panama;  M.L.  Kawasaki,  M. 
Bustamante,  P.E.  Oliveira,  and  P.O.  Wilson  for  sharing  work  in  press, 
unpublished  data,  and  observations;  and  the  many  people  on  BCI  and  at  STRI 
who  helped  the  Project  in  various  ways  but  are  too  numerous  to  thank  here 
individually.  The  Seedling  Flora  Project  has  been  generously  supported  by 
the  National  Science  Foundation  (BSR-85-17395),  the  Natural  Environment 
Research  Council  (GR3-695 1 ),  the  Smithsonian  Tropical  Research  Institute, 
The  Natural  History  Museum,  and  the  Linnean  Society  of  London. 


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Bull.  not.  Hist.  Mus.  Land.  (Bot.)  28(1):  17^0 


Issued  25  June  1998 


A  revision  of  the  genus  Mandragora 
(Solanaceae) 


STEFAN  UNGRICHT*  SANDRA  KNAPP  AND  JOHN  R.  PRESS 

Department  of  Botany,  Tne~Natural  History  Museum,  Cromwell  Road,  London  SW7  5BD 
*  Present  address:  Waldmatt  6,  CH-5242  Birr,  Switzerland 


CONTENTS 


Introduction 

Mythological  and  medicinal  history 

Taxonomic  history 

Materials  and  methods 

Material  examined 

Taxonomic  concepts 

Morphometrics 

Cladistics 

Results  and  discussion 

Species  delimitations  using  morphometric  analyses 

Phylogeny 

Biogeography 

Taxonomic  treatment 

Mandragora  L 

Key  to  the  species  of  Mandragora 

1.  Mandragora  officinarum  L 

2.  Mandragora  turcomanica  Mizg 

3.  Mandragora  caulescens  C.B.  Clarke 

References 

Exsiccatae 

Taxonomic  index  ... 


17 
18 
18 
19 
19 
20 
21 
22 
22 
22 
26 
26 
29 
29 
30 
30 
33 
34 
36 
38 
40 


SYNOPSIS.  The  Old  World  genus  Mandragora  L.  (Solanaceae)  is  revised  for  the  first  time  across  its  entire  geographical  range. 
The  introduction  reviews  the  extensive  mythological  and  medicinal  as  well  as  the  taxonomic  history  of  the  genus.  On 
morphological  and  phenological  grounds  three  geographically  widely  disjunct  species  can  be  distinguished:  the  Mediterranean 
M.  officinarum  L.,  the  narrowly  local  Turkmenian  endemic  M.  turcomanica  Mizg.  and  the  Sino-Himalayan  M  caulescens  C.B. 
Clarke.  The  generic  monophyly  of  Mandragora  L.  as  traditionally  circumscribed  is  supported  by  cladistic  analysis  of 
morphological  data.  The  ecological  and  historical  phytogeography  of  the  genus  is  discussed  and  alternative  biogeographical 
scenarios  are  evaluated.  Finally,  a  concise  taxonomic  treatment  of  the  taxa  is  provided,  based  on  the  evidence  of  the  preceeding 
analyses. 


INTRODUCTION 


The  nightshade  family  (Solanaceae)  is  a  cosmopolitan  but  predomi- 
nantly tropical  group  and  is  especially  well  represented  in  the 
Neotropics.  The  family  is  of  considerable  economic  importance. 
Food  plants  include  potato  (Solanum  tuberosum  L.),  tomato  (Sola- 
num  lycopersicum  L.  (=Lycopersicon  esculentum  Mill.)),  pepper 
(Capsicum  annuum  L.)  and  aubergine  (Solanum  melongena  L.j. 
Another  species  of  great  commercial  value  is  tobacco  (Nicotiana 
tabacum  LJ.  The  family  also  contains  ornamental  plants  such  as 
garden  petunias  (species  of  Petunia  Juss.)  and  the  Chinese  lantern 
(Phy  sails  alkekengi  L.j.  Many  members  of  the  family  are  poisonous, 
but  several  such  as  deadly  nightshade  (Atropa  belladonna  L.), 
henbane  (Hyoscyamus  niger  L.)  and  thorn-apple  (Datura  stramo- 
nium L.)  provide  drugs  traditionally  used  in  medicine.  The  subjects 
of  this  study,  the  mandrakes  (Mandragora  L.),  belong  to  this  latter 
group  of  pharmaceutically  interesting  Solanaceae. 


The  long  history  of  mythology  and  medicinal  use  of  the  mandrake 
combined  with  the  variable  morphology  and  phenology  have  led  to 
considerable  confusion  in  the  classification  of  Mandragora.  The 
nomenclatural  situation  is,  considering  the  small  size  of  the  genus, 
amazingly  chaotic.  In  particular  the  Mediterranean  Mandragora  is 
burdened  with  synonyms,  and  some  authors  (Mill,  in  litt.)  have 
expressed  doubts  about  the  placement  of  the  Himalayan  mandrake 
in  Mandragora  at  all.  This  study  represents  the  first  taxonomic 
revision  of  the  genus  over  its  entire  range.  We  address  the  following 
main  questions  in  this  study.  Firstly,  which  of  the  described  Man- 
dragora taxa  are  justifiable  if  the  whole  range  of  the  morphological 
variation  is  considered?  Secondly,  is  the  genus  as  currently  delim- 
ited (including  the  Himalayan  mandrake,  M.  caulescens) 
monophyletic?  Finally,  does  the  phylogeny,  combined  with  the 
distribution  patterns  of  the  taxa,  support  a  particular  historical 
biogeographical  scenario  and  do  the  ecological  requirements  of  the 
defined  taxa  impinge  upon  this? 


©  The  Natural  History  Museum,  1998 


18 

Mythological  and  medicinal  history 

The  origin  of  the  myth  surrounding  the  mandrake  root  is  hard  to 
trace.  It  is  clear,  however,  that  the  mandrake  was  a  focus  of  supersti- 
tion and  belief  in  magic  and  demons  for  many  centuries.  The 
sometimes  vaguely  humanoid  shape  of  the  root  and  its  potent, 
psychotropically  active  alkaloids  are  probably  responsible  for  the 
long  history  of  myths.  However,  the  properties  ascribed  to  the  plant 
as  a  whole,  and  especially  to  its  root,  are  highly  ambiguous  and  the 
first  references  in  Ancient  Egypt  and  in  the  Bible  remain  controver- 
sial. Ornaments  on  the  casket  and  throne  from  the  tomb  of 
Tutankhamun  seem  to  depict  mandrakes  (Hepper,  1990;  Roland, 
199 1 ).  Mandragora  is  not  known  to  be  native  in  Egypt  but  may  have 
been  cultivated  there.  The  Biblical  'Dudaim'  is  often  said  to  be 
identical  with  Mandragora  (Moldenke  &  Moldenke,  1952;  Fein- 
brun-Dothan,  1978;  Fleisher  &  Fleisher,  1994).  Dudaim  is  mentioned 
in  the  Song  of  Solomon  (7:  13)  and  in  another  passage  from  the  Old 
Testament  (Genesis  30:  14-18),  according  to  which  it  was  used  in 
relation  to  fertility.  The  'Moly '  of  Homer  in  his  Odyssey  is  presumed 
to  be  the  mandrake,  as  is  the  'Baraa'  of  Flavius  Josephus'  Jewish 
Wars,  where  it  is  described  as  shining  in  the  dark  (sic)  and  is 
attributed  exorcistic  powers.  Pliny  in  his  Natural  History  and 
Dioscorides  in  De  Materia  Medica  wrote  that  the  mandrake  was 
known  as  'Circaeon'  after  Circe,  the  mythical  sorceress  who  turned 
men  into  sexually  supercharged  swine  (Marzell,  1975;  Roland  1991; 
Fleisher  &  Fleisher,  1 994).  The  Ancient  Greeks  and  Romans  consid- 
ered the  mandrake  so  potent  and  valuable  as  a  narcotic  and  restorative 
plant  that  the  collectors  of  its  roots  had  to  obey  rigid  ceremonial 
rules  which  were  described  by  Theophrastus  and  Pliny  (Randolph, 
1905). 

In  Medieval  Europe  the  plant  was  known  as  'Alraune'  in  Germany 
and  'Main  de  gloire'  in  France.  Carved  to  emphasize  the  anthropo- 
morphic male  or  female  character  of  the  root,  it  was  popular  as 
aphrodisiac,  charm  against  sterility  and  love  talisman  alike.  But  the 
plant  was  soon  connected  with  witchcraft;  for  example,  Joan  of  Arc, 
who  was  tortured  and  burnt  at  the  stake  in  143 1  as  a  heretic,  was  also 
accused  of  the  witches'  crime  of  possessing  a  mandrake  (Thompson, 
1934).  It  was  believed  that  the  plant  grew  under  gallows  and  that  the 
root  owed  its  human  shape  to  urine  or  sperm  of  unjustly  hanged  men. 
The  gathering  of  the  root  was  considered  to  be  highly  dangerous  and 
dogs  had  to  perform  the  supposedly  lethal  procedure  of  pulling  out 
the  plants,  whereupon  the  roots  would  shriek  so  'that  living  mortals 
hearing  them  run  mad':  Shakespeare,  Romeo  and  Juliet  Act  IV 
(Randolph,  1905;  Moldenke  &  Moldenke,  1952;  Marzell,  1975; 
Roland,  1991;  Talalaj  et  al,  1992;  Fleisher  &  Fleisher,  1994). 
Perhaps  the  most  interesting  aspect  of  the  mythology  of  the  man- 
drake is  convergent  myths  in  the  eastern  area  of  the  genus  in  the 
disjunct  region  of  Central  Asia,  the  Himalayas  and  China  where 
Mandragora  has  a  similarly  long  history  of  application  and  tradi- 
tional use.  For  instance  in  Sikkim,  the  roots  of  Mandragora 
caulescens,  called  'Lakshmana',  were  used  in  magical  rites  and  in 
folk  medicine  (Mehra,  1979);  and  in  Central  Asia  the  mandrake, 
called  'Khaoma-soma',  was  considered  a  holy  plant  (Khlopin, 
1979). 

The  mythology  of  the  mandrake  and  its  use  in  traditional  medi- 
cine are  hard  to  keep  apart.  In  a  wide  range  of  applications  it  was 
used  as  an  aphrodisiac,  hypnotic,  emetic  and  purgative,  sedative  and 
narcotic,  for  example  as  a  pain-killer  at  childbirth  as  well  as  to 
relieve  crucifixion  agony  (Duke,  1985;  Talalaj  et  al.,  1992);  in  fact, 
according  to  Heiser  (1969),  it  has  been  questioned  that  Christ  died 
on  the  Cross  at  all,  suggesting  that  the  vinegar  supplied  to  him  had 
contained  the  drug,  inducing  merely  a  deathlike  trance.  The  second- 
ary metabolites  of  major  pharmaceutical  importance  are  chiefly 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 

tropane  alkaloids  and  their  esters.  Staub  (1962),  Hegnauer  (1973), 
Romeike  (1978),  Evans  (1979)  and  Jackson  &  Berry  (1979)  list 
alkaloids  of  Mandragora  and  other  Solanaceae.  Tropane  alkaloids 
occur  mainly  in  the  Solanaceae,  but  they  are  not  entirely  restricted  to 
this  family;  other  tropane  alkaloids  of  importance  include  cocaine, 
found  in  the  family  Erythroxylaceae.  However,  esters  with  tropic  or 
related  acids  of  alkamines  derived  from  tropane  do  seem  to  be 
restricted  to  the  Solanaceae  (Romeike,  1978). 

The  most  important  alkaloid  components  of  Mandragora  are: 
atropine,  apoatropine,  belladonnine,  cuscohygrine,  hyoscyamine, 
hyoscyamine-N-oxide,  norhyoscyamine,  scopolamine,  3oc-tigloyl- 
oxytropane,  3oc,6p-ditigloyloxytropane,  and  6p-hydroxyhyoscy- 
amine.  The  root,  stem,  leaves,  fruits  and  seeds  of  Mandragora  all 
contain  a  different  alkaloid  profile.  However,  the  root  is  considered 
to  be  the  site  of  synthesis  of  the  tropane  alkaloids  (Romeike,  1978) 
and  is  the  most  poisonous  part  of  Mandragora,  due  to  the  high 
concentration  therein.  The  total  content  of  alkaloids  in  the  root  is 
around  0.4%  (Kessler,  1951;  Maugini,  1959;  Duke,  1985),  hyo- 
scyamine being  the  most  important  constituent  (Duke,  1985).  In 
contrast,  the  berries  are  said  to  be  sweet  and  edible  in  moderation 
(Viney,  1994).  The  synthetic  pathways  leading  to  the  alkaloids  in 
different  taxa  of  the  Solanaceae  have  been  proposed  to  be  homolo- 
gous to  each  other  (Tetenyi,  1987)  and  to  be  a  linear  or  spiralling 
phyletic  sequence.  Tropane  alkaloids  are  effective  as  analgesics, 
anaesthetics,  antispasmodics,  and  are  used  to  increase  the  circula- 
tion, reduce  secretions  and  dilate  pupils.  Scopolamine  is  a  hypnotic 
(Duke,  1985)  and  achieved  a  notorious  reputation  as  a  'truth-drug'  in 
police  and  secret  service  investigations  (Heiser,  1969).  'Quishen',  a 
Chinese  herbal  medicine  consisting  of  dried  roots  of  M.  caulescens, 
is  used  in  Yunnan  and  Xizang  as  a  ginseng  substitute  for  all  kinds  of 
physical  weaknesses  (Xiao  &  He,  1983).  In  allopathic  western 
medicine,  however,  the  mandrake  has  largely  lost  its  importance  as 
valued  source  of  alkaloids,  although  the  plant  itself  is  still  sur- 
rounded by  myth. 

Taxonomic  history 

The  classification  of  Mandragora 

MEDITERRANEAN  PLANTS.  As  mentioned  above,  considering  the 
small  size  of  the  genus,  there  has  been  a  fair  amount  of  confusion  in 
respect  to  the  classification  of  Mandragora.  The  long  taxonomic 
history,  especially  of  the  Mediterranean  Mandragora  species,  led  to 
a  large  number  of  often  imprecisely  defined  species  and  subspecies 
or  varieties.  As  a  consequence,  there  is  a  plethora  of  names,  and  their 
somewhat  indiscriminate  use  renders  the  meaning  and  reliability  of 
identifications  of  specimens  in  herbaria,  as  well  as  in  the  literature, 
highly  problematic.  Here  we  present  the  taxonomic  history  of  the 
Mediterranean  part  of  the  genus  in  periods,  with  examples  of  major 
taxonomic  works  and  their  treatment  of  the  genus,  in  order  to  outline 
the  main  taxonomic  trends  through  the  centuries. 

Pre-Linnaean:  The  Ancients  distinguished  two  species  that  they 
called  the  female  or  black  mandrake  (Mandragorasfcemina),  and  the 
male  or  white  mandrake  (Mandragoras  mas).  These  plants  were 
described  by  both  Dioscorides  and  Pliny  (cf.  Randolph,  1905  for 
translations),  and  this  division  was  widely  adopted,  for  instance  by 
Parkinson  (1629)  and  by  Jean  Bauhin  et  al.  (1651)  as  late  as  the 
seventeenth  century.  The  male  mandrake  corresponds  to  a  spring 
flowering  entity;  the  female  mandrake  represents  an  autumn  flower- 
ing one.  Caspar  Bauhin  (1623)  and  Tournefort  (1719),  on  the  other 
hand,  each  recognized  three  'species'.  Both  authors  listed  Man- 
dragora fructu  rotundo  and  M.  /lore  subcaeruleo,  purpurascente. 


MANDRAGORA 


19 


Bauhin's  third  species  was  M.fructu  pyri  while  Tournefort's  was  M. 
flore  subcaeruleo,  foliis  minoribus,  fructu  globoso). 

Linnaeus:  In  hisHortus cliff ortianus,  Linnaeus  ( 1 738)  considered 
these  variants  as  mere  forms  of  a  single  species.  In  the  first  edition  of 
Species  plantarum  (1753),  he  accordingly  recognized  only  one 
species  that  he  named  Mandragora  officinarum.  Later,  however,  he 
changed  his  mind  with  respect  to  the  distinctness  of  this  taxon  and 
placed  it  within  his  genus  Atropa,  calling  it  Atropa  mandragora 
(Linnaeus,  1759,  1762). 

Post-Linnaean:  Linnaeus's  idea  of  a  single  species  did  not  last 
very  long.  The  nineteenth  and  the  first  half  of  the  twentieth  centuries 
were  characterized  by  an  increasingly  elaborate  division  of  the 
genus  (Table  1 ).  Heldreich  ( 1 886)  additionally  described  an  infertile 
hybrid  between  two  vernal  entities:  Mandragora  vernalis  Bertol. 
and  M.  haussknechtii  Heldr.,  called  M.  x  hybrida  Haussk.  &  Heldr. 

Post-1950:  In  recent  decades,  however,  there  has  been  a  trend 
towards  reversing  this  ever  greater  subdivision  of  the  genus.  In  fact, 
since  the  1950s  there  have  been  a  number  of  proposals  to  re-adopt 
the  two  traditional,  pre-Linnaean  taxa  only  (Tercinet,  1950;  Hawkes, 
19726;  Jackson  &  Berry,  1979).  This  viewpoint  was  adopted  in 
Flora  Europaea  (Hawkes,  1972a),  using  the  names  Mandragora 
officinarum  L.  for  a  vernal  entity  and  M.  autumnalis  Bertol.  for  an 
autumnal  one.  However,  M.  officinarum  L.  might  be  considered  an 
ambiguous  name  since  it  was  formerly  used  for  an  autumnal  entity 
as  well  (Greuter  &  Rechinger,  1967;  see  Table  1).  Hawkes  (19726) 
rejected  this  reservation  on  the  grounds  of  the  priority,  the  wide  use 
and  the  generic  type  status  of  the  Linnaean  epithet  officinarum. 
Knapp  in  Jarvis  et  al.  (1993)  finally  lectotypified  M.  officinarum  L. 
with  a  specimen  in  the  Bursar  herbarium,  to  which  no  phenological 
data  are  attached. 

SlNO-HiMALAYAN  PLANTS.  In  contrast  to  the  Mediterranean  plants, 
the  classification  of  the  disjunct  Central  and  East  Asian  part  of  the 
genus  was  never  as  contentious  and  did  not  arouse  much  comment, 
in  part  reflecting  its  far  younger  taxonomic  history.  Mandragora 
caulescens  was  described  by  C.B.  Clarke  (1883)  in  Hooker's  The 
flora  of  British  India  (Hooker  fil.  and  Thomson  labelled  the  speci- 
mens 'Scopolia  humilis '  but  this  name  was  never  published).  Anisodus 
caulescens  (C.B.  Clarke)  Diels  and  A.  mariae  Pascher,  as  well  as 
Mairella  yunnanensis  H.  Lev.,  were  placed  in  synonymy  with 
Mandragora  caulescens  in  the  Flora  of  China  (Zhang  et  al.,  1994). 
Grubov  (1970)  described  a  new  species  from  Xizang,  China  which 
he  named  Mandragora  tibetica,  and  Kuang  Ko-zen  &  Lu  An-ming 
(1978)  described  a  new  species  found  in  Qinghai  and  Xizang  which 
they  named  Mandragora  chinghaiensis.  In  the  Flora  of  China 
(Zhang  et  al.,  1994)  the  latter  two  taxa  were  provisionally  put  into 
synonymy  with  M.  caulescens  until  a  more  detailed  revision  could 

Table  1     Post-Linnaean  classifications  of  the  increasingly  subdivided 
Mediterranean  taxa  of  Mandragora. 


Classification  type        Vernal  species 


Autumnal  species 


1  VERNAL,  1  AUTUMNAL  TAXON 
Sprengel  (1825)        M.  vernalis  Bertol. 

1  VERNAL,  2  AUTUMNAL  TAXA 

Bertoloni  ( 1 835),      M.  vernalis  Bertol. 
Dunal  (1852) 

2  VERNAL,  2  AUTUMNAL  TAXA 

Heldreich  ( 1 886)      M.  vernalis  Bertol. 

M.  haussknechtii  Heldr. 

3  VERNAL,  2  AUTUMNAL  TAXA 

Vierhapper  (1915)    M.  mas  Garsault 

M.  hispanica  Vierh. 
M.  haussknechtii  Heldr. 


M.  autumnalis  Bertol. 

M.  officinarum  L. 
M.  microcarpa  Bertol. 

M.  autumnalis  Spreng. 
M.  microcarpa  Bertol. 

M.  autumnalis  Bertol. 
M.  foemina  Garsault 


establish  their  distinctiveness.  Mandragora  shebbeard  C.E.C. 
Fischer,  described  from  a  single  specimen  from  Xizang  (Fischer, 
1934),  was  transferred  to  the  monotypic  solanaceous  genus 
Przewalskia  by  Grubov  (1970).  One  attempt  to  subdivide  Man- 
dragora caulescens  was  made  by  Grierson  &  Long  (1978)  who  split 
it  into  four  subspecies  (subsp.  caulescens,  subsp.  flavida  Grierson 
&  Long,  subsp.  purpurascens  Grierson  &  Long  and  subsp.  brevi- 
calyx  Grierson  &  Long)  differentiated  on  the  basis  of  corolla  colour 
and  various  morphometric  characters,  especially  of  flower  parts. 

TURKMENIAN  PLANTS.  Finally,  Mandragora  turcomanica  Mizg. 
was  described  in  1942  from  a  small  area  (the  Shevlan  'pocket', 
Mizgireva,  1955)  near  Kara  Kala  in  the  southwestern  Kopet  Dag  in 
Turkmenistan.  Only  a  few  collections  of  this  restricted  endemic 
exist,  and  these  are  all  to  be  found  in  herbaria  of  the  former  Soviet 
Union.  Its  status  as  a  distinct  taxon  has  never  really  been  questioned, 
but  only  because  so  few  specimens  exist  for  comparison.  Mizgireva 
( 1 955)  worked  for  1 2  years  on  the  detailed  anatomy  and  morphology 
of  this  species  and  her  revision  remains  the  best  source  for  data  on 
characters  and  their  variation. 

The  position  of  Mandragora  within  the  Solanaceae 

More  than  200  years  after  the  original  outline  by  Jussieu  (1789),  the 
higher  level  classification  of  Solanaceae  is  far  from  being  settled, 
and  the  placement  of  Mandragora  in  the  various  proposed  classifi- 
cations is  just  as  controversial.  In  the  following  review,  the  most 
widely  followed  classifications  of  the  Solanaceae  are  presented 
together  with  the  positions  of  Mandragora  and  the  other  genera  used 
in  our  cladistic  analysis:  Anisodus  Link,  Atropa  L.,  Lycium  L.  and 
Nicotiana  L. 

In  his  artificial  sexual  system,  Linnaeus  (1753)  placed  Man- 
dragora within  his  group  Pentandria  Monogyna,  together  with 
Atropa,  Lycium  and  Nicotiana.  Jussieu  (1789)  defined  the  'order' 
Solaneae  with  Mandragora  in  group  II  (fructus  baccatus)  alongside 
Atropa  and  Lycium.  Nicotiana  on  the  other  hand  was  placed  in  group 
I  (fructus  capsularis).  In  the  nineteenth  century,  classifications  be- 
came more  elaborate  and  tribes  and  subtribes  were  established  in 
order  to  approximate  a  more  'natural'  classification.  In  Table  2,  the 
three  most  widely  adopted  classifications  of  the  Solanaceae  in  this 
period  (Dunal,  1852;Bentham,  1 876;  Wettstein,  1895)  are  presented 
together  with  the  more  recent  one  of  Baehni  (1946). 

The  last  few  decades  have  seen  the  analysis  of  new  types  of  data, 
such  as  DNA  sequences  (Olmstead  &  Sweere,  1994)  and  secondary 
plant  metabolites  (Tetenyi,  1987),  as  well  as  application  of  cladistic 
methodology  to  morphological  data  (Hoare  &  Knapp,  1997).  This 
has  led  to  new  hypotheses  concerning  relationships  within  the 
Solanaceae,  as  well  as  the  placement  of  Mandragora.  However, 
opinion  still  differs  widely  on  whether  Mandragora  should  be 
placed  within  the  tribe  Hyoscyameae  (Hoare  &  Knapp,  1997;  = 
Atropeae  sensu  Tetenyi,  1987)  or  in  an  isolated,  basal  monogeneric 
tribe  Mandragoreae  (Olmstead  &  Sweere,  1994;  Hunziker,  1995; 
Olmstead  et  al.,  in  press). 


MATERIALS  AND  METHODS 


Material  examined 

This  revision  is  based  exclusively  on  herbarium  specimens.  The 
material  examined  includes  the  Mandragora  specimens  in  the  her- 
baria of  The  Natural  History  Museum  of  London  (BM)  and  Kew  (K) 
together  with  specimens  borrowed  from  the  following  herbaria: 


20  S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 

Table  2    Tribal  classification  of  the  Solanaceae  and  the  position  of  Mandragora  and  the  outgroup  genera  Anisodus,  Atropa,  Lycium  and  Nicotiana. 


Author 

Classification 

Genera  of  this  study 

Dunal(1852) 

Nolaneae 

Nolanineae 

Grabowskieae 

Solaneae 

Triguereae 

Solaneae                                        Solanineae 

Atropineae 

Mandragora 

Atropa 

Lycineae 

Lycium 

Datureae 

Hyoscyameae 

Anisodus  (Scopolia) 

Nicotianeae 

Nicotiana 

Retzieae 

Fabianeae 

Metternichieae 

Cestreae 

Bentham(1876) 

Solaneae 

Atropeae 

Mandragora 

Atropa 

Lycium 

Hyoscyameae 

Anisodus  (Scopolia) 

Cestrineae 

Nicotiana 

Salpiglossidae 

Wettstein(1895) 

Nicandreae 

Solaneae 

Lyciinae 

Atropa 

Lycium 

Hyoscyaminae 

Anisodus  (Scopolia) 

Solaninae 

Mandragorinae 

Mandragora 

Daturae 

Cestreae 

Cestrinae 

Goetzeinae 

Nicotianinae 

Nicotiana 

Salpiglossideae 

Baehni(1946) 

Solaneae 

Solaninae 

Sarachinae 

Margaranthinae 

Physalidinae 

lochrominae 

Goetzeinae 

Discopodiinae 

Atropeae 

Atropinae 

Mandragora 

Atropa 

Lycium 

Markeinae 

Hyoscyaminae 

Anisodus  (Scopolia) 

Parabouchetiinae 

Anthocercideae 

Nicotianeae 

Nicotianinae 

Nicotiana 

Daturinae 

Nicandrinae 

Salpiglossideae 

Berkeley  (UC,  JEPS),  Edinburgh  (E),  Kunming  (KUN),  Palermo 
(PAL)  and  Reading  (RNG).  The  collections  of  the  Museum  National 
d'Histoire  Naturelle  in  Paris  (P)  were  examined  on  site  during  a 
research  visit.  Regrettably  it  was  not  possible  to  obtain  any  speci- 
mens, including  the  types,  of  Mandragora  turcomanica  and  M. 
tibetica  from  St.  Petersburg  (LE)  or  M.  chinghaiensis  from  Xining 
(HNWP). 

Material  with  the  same  collection  number  or  specimens  collected 
at  the  same  locality  and  at  the  same  date  were  unified  and  treated  as 
one  sample.  Unlocalized  material  or  specimens  with  illegible, 
doutbful  or  very  imprecise  collection  locality  and  collector  data 
were  excluded,  cultivated  material  being  the  exception.  Only  collec- 
tions with  precise  locality  data  are  listed  in  the  taxonomic  account  of 
the  species.  A  complete  listing  of  the  specimens  examined  can  be 
found  in  the  Exsiccatae. 


Taxonomic  concepts 

The  adopted  taxonomic  methodology  in  this  study  of  herbarium 
material  is  necessarily  largely  pattern-oriented.  If  process-based 
terms  like  'population'  or  'hybrid'  are  used,  these  properties  are  only 
informally  applied,  i.e.  inferred  rather  than  rigidly  defined,  let  alone 
experimentally  circumscribed. 

The  adopted  species  concept  is  operational,  i.e.  the  ability  to 
distinguish  between  sets  of  specimens  serves  as  the  criterion  for  the 
delimitation  of  the  terminal  taxa  (species).  Cladistic  analysis  results 
in  an  estimate  of  these  relationship  between  the  species  and  the 
delimitation  of  higher  level  taxa,  i.e.  monophyletic  groups  and  their 
hierarchical  relationships.  The  requirement  for  a  species  description 
is  thus  at  least  one  distinguishing  qualitative,  or  discontinuous 
quantitative,  diagnostic  character  state  (monothetic  species)  or  a 
unique  set  of  character  states  (polythetic  species)  (cf.  Nelson  & 


MANDRAGORA 

Platnick,  1 981;  Nixon  &  Wheeler,  1990).  Morphological  and  pheno- 
logical  variation  plus  ecological  and  geographical  information  are 
the  data  that  were  collected  from  herbarium  specimens  and  their 
labels  respectively. 

Morphometrics 

Specimen  sampling 

The  inherent  weakness  of  the  traditional  herbarium  record  of  a  taxon 
for  taxonomic  studies  is  well  known  (Du  Rietz,  1930;  Anderson  & 
Turrill,  1935;  Anderson,  1941).  For  species  that  are  well  represented 
in  herbaria,  one  can  obtain  a  more  or  less  accurate  record  of  the 
general  range  of  variation.  Seldom  or  never,  however,  is  it  possible 
to  calculate  frequencies  of  the  different  variants  and  the  resolution  of 
the  classification  is  limited  by  the  lack  of  a  random  sample. 

Plant  variation  also  occurs  on  different  hierarchical  levels  (cf. 
Bateman,  1989),  which  can  become  confused  and  overlap.  In  Man- 
dragora  the  confounding  of  within-group  and  between-group 
variation  is  most  pronounced  in  the  Sino-Himalayan  complex,  due 
to  the  secondary  growth  of  the  plants  during  the  late  flowering  and 
the  fruiting  period  (ontogenetic  variation,  see  Fig.  9,  also  Hoare  & 
Knapp,  1997:  fig.  5d).  Combining  all  samples  to  a  pooled  matrix 
easily  leads  to  a  confusion  of  the  two  sources  of  variation  which  can 
seriously  compromise  interpretation  of  many  applications  of 
morphometric  analyses  (Gibson  et  al,  1984).  Avoiding  within- 
group  variation,  for  example  by  calculating  the  mean  or  only 
considering  the  largest  or  smallest  example  of  a  particular  character 
is  not  a  solution,  but  merely  reduces  the  total  amount  of  variation 
encountered  in  a  taxon,  leading  ultimately  to  a  simplistic  taxon 
delimitation.  In  the  taxonomic  process,  characters  and  their  variabil- 
ity necessarily  have  priority  over  all  other  considerations  since  taxa 
are  attributed  to  characters — not  the  other  way  around  (characters  all 
attributed  to  individuals).  It  follows  that  the  basal  unit  of  investiga- 
tion, the  specimen,  must  be  evaluated  in  respect  to  its  variability. 
Therefore,  maximal  as  well  as  minimal  values  of  the  flowering  part 
measurements  of  each  specimen  were  assessed,  rather  than  solely 
mean  or  maximal  values. 

Character  sampling 

A  genus  with  a  long  taxonomic  history  is  unlikely  to  yield  any  'new' 
gross  morphological  characters  which  distinguish  species.  This  is 
especially  true  when  working  with  inadequate  herbarium  material 
(i.e.  incomplete  with  much  missing  data  and/or  very  old  and  often  in 
poor  condition).  The  emphasis  in  this  study  was  thus  much  more  on 
evaluating  the  traditional  and  obvious  characters  in  respect  to  their 
ability  consistently  to  delimit  proposed  taxa.  The  analysis  was 
therefore  carried  out  using  the  following  data. 

GEOGRAPHY.  The  entire  distributional  range  of  Mandragora  was 
divided  into  20  geographic  areas.  These  are  (1)  Cultivated  speci- 
mens, typically  from  botanic  gardens  north  of  the  Alps,  (2)  North 
Africa,  (3)  Iberia,  (4)  northern  Italy  and  Dalmatia,  (5)  peninsular 
Italy  with  Sardinia  and  Sicily,  (6)  mainland  Greece,  (7)  Aegean 
islands,  (8)  Asia  Minor,  (9)  Cyprus,  (10)  Levant,  (11)  Kopet  Dag, 
(12)  Nepal,  (13)  Sikkim,  (14)  Bhutan,  (15)  Arunchal  Pradesch 
(Assam),  (16)  north  Myanmar  (Burma),  (17)  Yunnan,  (18)  Sichuan, 
(19)  Qinghai  and  (20)  Xizang. 

PHENOLOGY.  Flowering  and  fruiting  dates  were  recorded,  result- 
ing in  the  cumulative  reproductive  period. 

MORPHOLOGY.  Corolla  length,  calyx  length,  fruit  size,  fruiting 
calyx  length,  leaf  length,  leaf  length/width  ratio.  The  corolla  colour 


21 

of  herbarium  material  is  not  reliably  determinable  and  was  thus  only 
recorded  if  it  was  stated  on  the  labels.  The  shape  of  the  berry 
(globose  or  ovoid)  was  excluded  as  a  character  since  it  is  not 
accessible  through  pressed  herbarium  material  and  is  only  rarely 
stated  on  the  labels.  Berry  shape  in  the  Solanaceae  is  quite  variable 
and  this  variation  is  occasionally  a  developmental  sequence  (Bohs, 
1994). 

Data  analysis 

Univariate  and  multivariate  analyses  were  applied  as  a  descriptive, 
explorative  tool  to  study  the  joint  relationships  of  the  morphological, 
phenological  and  geographical  variables  of  the  specimens  at  hand. 
As  mentioned  above,  since  the  examined  specimens  are  not  a 
random  sample,  the  result  of  inferential  data  analysis  with  signifi- 
cance testing  would  be  invalid.  A  cluster  analysis  or  a  cladistic 
analysis  of  the  data  at  population  level,  on  the  other  hand,  was 
rejected  on  the  grounds  that  these  methods  assume  hierarchical 
relationships,  an  assumption  that  is  not  justified.  Phenotypic  charac- 
ters may  be  distributed  clinally  or  in  a  reticulate  manner,  but  are 
unlikely  to  be  strictly  hierarchical  (Crisp  &  Weston,  1993).  If 
hierarchical  structure  exists,  it  would  suggest  that  the  populations 
have  already  undergone  speciation  and  have  to  be  treated  as  terminal 
taxa  in  the  phylogenetic  analysis.  The  explorative  data  analysis  was 
performed  on  SYSTAT  for  the  Macintosh  Version  5.2  (Wilkinson, 
1992). 

UNIVARIATE  ANALYSES.  Phenological  data  are  circular  (Fisher, 
1993).  The  reproductive  period  was  thus  analysed  in  a  polar  coordi- 
nate system.  In  order  to  dampen  peak  values,  which  might  merely 
reflect  a  particularly  high  collecting  effort,  the  collection  proportion 
of  each  month  was  converted  to  its  logarithm. 

MULTIVARIATE  ANALYSES.  Ordination  techniques  reduce  a  data 
matrix  of  distances  (similarities)  among  the  samples  to  one  or  a  few 
dimensions.  These  types  of  techniques  seem  to  be  most  appropriate 
under  a  nonhierarchical  model  of  infraspecific  variation.  Principal 
component  analysis  (PCA)  is  the  method  most  commonly  applied  in 
multivariate  systematics  studies  (James  &  McCulloch,  1990).  It 
aims  to  show  the  eventual  existence  of  distinguishable  groups  with 
discontinuous  variation  without  a  priori  postulates  of  groups  such  as 
populations.  Since  data  sets  with  missing  data  cannot  be  processed 
using  PCA,  only  the  most  commonly  represented  characters  of  the 
herbarium  specimens  (see  above  and  Table  3)  were  analysed.  Fortu- 
nately, these  are  also  the  characters  that  are  most  commonly  applied 
to  differentiate  between  the  traditionally  recognized  Mediterranean 
taxa.  In  constrast  to  many  other  analyses  of  specimen-based  data, 
within-collection  variation  was  accounted  for  by  recording  and 
analysing  both  minimal  and  maximal  values  of  the  flowering  part 
lengths.  Only  fully  developed  and  open  flowers  were  considered  and 
measured.  The  length/width  ratio  of  the  leaves  was  not  included  in 
the  PCA  analysis  on  the  grounds  that  ratios  are  often  not  linearly 
related  to  the  length  measurements  (Miles  &  Ricklefs,  1 984).  All  the 
variables  have  been  standardized  (normalized)  by  the  standard 


Table  3     Variables  used  in  the  principal  component  analysis  of  the 
Mediterranean  complex. 


Length  measurements 

Variable  1 

Corolla  length  max. 

mm 

Variable  2 

Corolla  length  min. 

mm 

Variable  3 

Calyx  length  max. 

mm 

Variable  4 

Calyx  length  min. 

mm 

Variable  5 

Leaf  length  max. 

cm 

Phenology 

Variable  6 

Flowering  time 

month 

22 

deviation  to  obtain  a  variance  of  one  and  a  mean  of  zero  by 
performing  the  analysis  on  a  correlation  matrix. 

Due  to  a  lack  of  available  specimens  from  Asia  of  the  taxa 
Mandragora  caulescens  subsp.  brevicalyx,  M.  chinghaiensis,  M. 
tibetica  and  M.  turcomanica,  the  discussion  of  variability  among 
these  taxa  (and  its  implications  on  their  delimitation)  has  to  remain 
qualitative. 

Cladistics 

Cladistic  analysis  is  a  parsimony-based  search  for  hierarchical 
arrangements  or  patterns  of  terminal  units  (in  most  cases  species). 
The  premises  are  accordingly  that  there  is  a  hierarchical  structure  of 
the  taxa  and  therefore  of  the  characters  used  to  define  them,  and  that 
the  level  of  homoplasy  in  the  chosen  characters  is  low  enough  not  to 
infer  convergent  groups.  As  a  strictly  numerical  procedure,  there  is 
no  need  for  additional  models  or  assumptions  relating  to  biological 
processes,  such  as  reproduction  or  descent  (Davis  &  Nixon,  1992). 
The  resulting  cladogram  therefore  represents  a  synapomorphy 
scheme,  merely  suggesting  closeness  of  relationship  due  to  recency 
of  shared  common  ancestry.  It  might  be  congruent  with  the  actual 
phylogenetic  tree,  featuring  the  ancestors  and  descendants;  how- 
ever, there  are  many  possible  phylogenetic  trees  compatible  with 
one  cladogram  (Eldredge,  1979).  The  cladogram  was  rooted  by 
means  of  outgroup  comparison. 

A  phylogeny  of  the  tribe  Hyoscyameae  based  mainly  on  morpho- 
logical characters  was  recently  published  by  Hoare  &  Knapp  (1997) 
suggesting  that  Mandragora,  as  traditionally  defined,  is  mono- 
phyletic.  To  test  the  monophyly  of  Mandragora  as  a  whole,  we 
included  in  this  analysis  material  of  species  representing  two  other 
genera  of  the  Hyoscyameae,  Anisodus  luridus  Link  and  Atropa 
belladonna  (see  Table  4).  The  three  recognized  terminal  taxa  of 
Mandragora  represent  the  ingroup  (M.  caulescens,  M.  officinarum, 

Table  4     Taxa  used  in  the  cladistic  analysis. 

Nicotiana  glauca  Graham 
Lycium  chinense  Mill. 
Anisodus  luridus  Link 
Atropa  belladonna  L. 
Mandragora  officinarum  L. 
M.  turcomanica  Mizg. 
M.  caulescens  C.B.  Clarke 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 
Table  6     Data  matrix  used  in  the  HENNIG86  analysis 


Taxon 


Character 


1  I 
1234567890  1 


Nicotiana  glauca 
Lycium  chinense 
Anisodus  luridus 
Atropa  belladonna 
Mandragora  officinarum 
M.  turcomanica 
M.  caulescens 


00200010000 
0020000001  0 
11201  111110 
10201  10001  1 
11011  10001  0 
11011  10101  ? 
11111  10111  1 


M.  turcomanica).  The  outgroup,  Nicotiana  glauca  Graham,  was 
chosen  based  upon  previous  cladistic  analyses  of  chloroplast  DNA 
data  sets  (Olmstead  &  Palmer,  1992;  Olmstead  &  Sweere,  1994; 
Olmstead  et  al.,  in  press).  Lycium  chinense  Mill,  was  included  as  an 
additional  taxon  (but  not  defined  as  an  outgroup,  see  Tables  2  and  4). 

Although  acknowledging  that  there  is  no  such  thing  as  an  a  priori 
'perfect  cladistic  character' (Thiele,  1993),  the  suggestion  of  Chappill 
(1989)  and  Stevens  (1991)  not  to  include  purely  morphometric, 
quantitative  data  in  the  cladistic  analysis  was  followed.  This  limita- 
tion is  feasible  in  view  of  the  minimal  size  of  the  ingroup  and  the 
already  established  analysis  of  an  extensive  character  set  including 
quantitative  character  states  (Hoare  &  Knapp,  1997).  The  characters 
used  in  the  present  work  (Table  5)  and  their  states  are  largely  self- 
explanatory.  The  data  matrix  used  in  the  analysis  is  presented  in 
Table  6. 

Cladistic  analyses  were  undertaken  withHENNlG86  (Farris,  1988) 
using  the  ie  option  (implicit  enumeration)  with  all  characters 
unordered,  thus  avoiding  any  character  state  change  assumptions. 
The  generated  cladogram  statistics  include  length  (L),  the  number  of 
character  state  changes  on  the  tree,  the  ensemble  consistency  index 
(CI)  which  provides  a  measure  of  character  fit  on  the  cladogram 
(amount  of  homoplasy),  and  the  ensemble  retention  index  (RI) 
which  expresses  the  fraction  of  similarities  on  the  cladogram  inter- 
preted as  synapomorphy  (Farris,  1989).  Both  of  these  indices  have 
an  optimal  value  of  1 . 


RESULTS  AND  DISCUSSION 


Table  5     Character  set  used  in  the  cladistic  analysis. 

Habit 

1 .  Woody  plant  (shrub  or  tree)  0;  herb  1 
Root 

2.  Tap-root:  enlarged  tap-root  absent  0;  present  1 
Stem 

3.  Axis:  condensed  0;  first  condensed,  later  elongated  1 ;  elongated  2 
Leaves 

4.  At  anthesis:  leaves  clustered  in  a  rosette  0;  leaves  alternate  1 
Inflorescence 

5.  Type:  numerous  flowers  in  panicle  0;  l(-3)  axillary  flowers  1 
Corolla 

6.  Shape:  tubular-infundibuliform  0;  campanulate-urceolate  1 
Fruit 

7.  Type:  berry  0;  capsule  16 
Fruiting  calyx 

8.  Not  enlarged  in  fruit  0;  markedly  enlarged  in  fruit  1 

9.  Without  prominent  ribs  0;  with  prominent  ribs  1 
Seeds 

10.  Embryo:  not  curved  0;  curved  circularly  or  in  a  spiral  1 

1 1 .  Testal  cells  deep  0;  shallow  1 


Species  delimitation  using  morphometric  analyses 

For  the  Mediterranean  taxa  a  number  of  specific  hypotheses  were 
tested  using  the  above  described  statistical  analyses. 

Univariate  analysis  of  reproductive  period. — Are  there  two 
distinct  vernal  and  autumnal  taxa  ? 

By  adopting  the  two  species  Mandragora  vernalis  and  M.  autumn- 
alis,  Sprengel  ( 1 825)  argued  against  Linnaeus's  unified  Mandragora 
officinarum  concept.  Even  in  later,  more  elaborate  classifications  the 
division  into  these  two  major  groups  remained  important  (Table  1) 
since  it  was  obvious  that  two  groups  with  widely  differing  flowering 
periods  would  hardly  be  able  to  interbreed  due  to  the  seasonal 
isolation.  So,  is  there  evidence  in  the  compiled  Mediterranean  data 
set  that  there  are  two  phenologically  distinct  species?  Figure  1 
shows  the  distribution  of  the  collecting  dates  (in  flower  or  in  fruit)  as 
an  indicator  of  the  reproductive  period. 

It  is  evident  that  there  is  actually  only  one,  very  extensive  period 
of  reproductive  activity  from  autumn  to  spring.  In  fact  only  during 
the  hottest  summer  months  does  the  cycle  break.  If  conditions  are 


MANDRAGORA 


23 


Mar 


Jun 


Feb 


Jan 


Dec 


Sep 


Nov 


Fig.  1     Relative  distribution  of  the  vegetative  period  of  the  Mediterranean 
complex  (Mandragora  officinaruiri)  in  the  course  of  the  year 
(logarithimized  monthly  fractions:  for  analysis  see  text). 

favourable,  the  same  individual  can  flower  twice  a  year,  in  autumn 
and  then  again  in  spring,  as  can  be  seen  from  annotated  herbarium 
material,  especially  from  cultivated  material  in  botanical  gardens. 
The  key  factor  is  probably  sufficient  rainfall;  for  instance,  sudden 
and  widespread  flowering  after  exceptionally  heavy  autumn  rains 
was  recorded  from  Tunisia  (label  data  from  Tomkinson  72).  A  more 
detailed  discussion  pertaining  to  the  influence  of  the  climate  and  the 
summer  dormancy  is  given  in  the  section  on  ecological  biogeo- 
graphy  below. 

Multivariate  analysis  of  morphometric  characters.  -  Are 
there  distinct  clusters  within  the  Mediterranean  mandrakes 
that  are  recognizable  from  gross  morphology? 

The  different  eigenvector  values  of  the  PCA  (Table  8)  indicate  the 
relative  contribution  of  the  variables.  The  first  three  axes  (see  Table 
7)  explain  92.2%  of  the  variation  if  length  measurements  are  consid- 
ered only  (Fig.  2a),  and  87. 1%  of  the  variation  if  the  analysis  was  run 
including  the  phenological  variable  of  flowering  time  (Fig.  2b).  In 
both  cases  it  is  obvious  that  no  clear-cut  clusters  can  be  distin- 
guished. Cryptic  species  may  exist  within  the  Mediterranean  region 
but  are  likely  to  be  revealed  only  if  extensive,  field-based  studies 
over  the  entire  range,  as  well  as  transplant  experiments,  are  under- 
taken (Clausen  et  al,  1940).  The  information  currently  available 
does  not  justify  recognition  of  more  than  a  single  species  of  Mediter- 
ranean mandrake. 

Other  characters  traditionally  applied  in  keys  but  not  analysed 
quantitatively  here  include  the  following: 

Corolla  colour.  Hawkes  (1972b)  considered  the  elaborate  split- 
ting of  the  vernal  and  autumnal  groups  as  unjustified  and  recognized 
only  the  white-flowered  vernal  plants  from  northern  Italy  and  Dal- 
matia  as  distinct  from  the  remaining  Mediterranean  group.  Restricted 
to  only  a  small  part  of  the  total  Mediterranean  range,  the  white- 
flowered  plant  has  often  been  considered  as  the  most  distinctive 
group  (Heldreich,  1886;  Vierhapper,  1915).  The  diagnostic  value  of 
this  character  which  has  been  extensively  used  in  past  and  current 


Table  7     Principal  components  (PC)  of  the  Mediterranean  complex.  A: 
Length  measurements  only.  B:  Length  measurements  and  flowering 
period  combined. 


Component 


Eigenvalue 


%  of  variance  explained 


A     PC  1 

3.248 

64.97 

PC  2 

0.977 

19.54 

PC  3 

0.383 

7.66 

B     PC  1 

3.328 

55.47 

PC2 

1.410 

23.50 

PC  3 

0.490 

8.17 

Table  8     Eigenvectors  of  the  three  most  important  axes  (component 
loadings). 


Variable 


PC  1 


PC  2 


PC  3 


A 

1 

Corolla  length  max. 

0.885 

0.023 

0.131 

2 

Corolla  length  min. 

0.875 

0.011 

0.435 

3 

Calyx  length  max. 

0.898 

0.114 

-0.391 

4 

Calyx  length  min. 

0.924 

0.065 

-0.147 

5 

Leaf  length  max. 

-0.197 

0.979 

0.047 

B 

1 

Corolla  length  max. 

0.882 

0.067 

-0.160 

2 

Corolla  length  min. 

0.851 

0.189 

0.042 

3 

Calyx  length  max. 

0.897 

0.123 

0.066 

4 

Calyx  length  min. 

0.905 

0.156 

0.175 

5 

Leaf  length  max. 

-0.181 

0.865 

-0.453 

6 

Flowering  time 

-0.413 

0.763 

0.471 

keys  is  minimal,  however,  if  one  considers  that  flowers  are  very 
variable  in  colour  even  within  individuals  (Jury  et  al.  12238  from 
Morocco,  Jury  &  Jury  13111  from  Spain). 

Fruiting  calyx  length:  The  relative  length  of  the  accrescent  fruit- 
ing calyx  to  the  berry  (longer:  M.  autumnalis  or  shorter:  M. 
officinarum)  has  also  been  proposed  as  a  diagnostic  character 
(Hawkes,  19726).  Again,  it  seems  that  within-group  variation  is  just 
as  high,  considering  for  instance  a  specimen  housed  in  P  of  a 
cultivated  plant  which  features  two  berries  that  are  clearly  shorter 
than  the  calyx,  but  three  others  which  are  longer. 

Leaf  shape:  This  is  perhaps  the  most  variable  character  of  all.  One 
can  get  at  least  an  idea  of  the  variability  if  the  length  to  width  ratios 
of  mature  leaves  are  compared.  The  variability  within  an  individual 
ranges  from  1.7:1-4:1  (in  Reverchon  s.n.)  to  4.6:1-10.5:1  (in 
Welwitsch  159).  The  total  variability  within  the  whole  set  of  exam- 
ined specimens  was  found  to  be  between  1.5:1-10.5:1. 

Any  distinction  between  Mandragora  officinarum  and  M. 
autumnalis  or  any  other  splitting  of  the  Mediterranean  Mandragora 
on  morphological  or  phenological  grounds  would  be  intentionally 
giving  formal  status  to  divergent  ends  of  clines.  Thus  overlapping 
intermediates  would  be  difficult  if  not  impossible  to  assign  to  either 
of  the  taxa  in  any  other  way  than  arbitrarily.  Extensive  overlap  and 
putative  hybrids  must  be  considered  if  aiming  at  expressing  the 
whole  range  of  variability.  This  confirms  Meikle's  (1985)  doubts 
concerning  the  validity  of  the  different  proposed  Mediterranean 
species.  Although  certain  plants  of  particular  populations  are  quite 
different  in  appearance  from  some  plants  of  other  populations,  when 
the  total  pattern  of  variation  over  the  entire  range  is  considered,  no 
diagnosable  taxa  within  the  Mediterranean  complex  of  Mandragora 
are  discernable.  Further  morphological  support  for  the  unification  of 
the  Mediterranean  Mandragora  comes  from  the  anatomy  of  the  tap 
roots  which  has  been  shown  to  be  identical  in  vernal  and  autumnal 
taxa  and  in  their  chemical  composition  which  is  also  similar  (Jackson 
&  Berry,  1979). 

For  Mandragora  turcomanica,  a  statistically  valid  morphometric 
analysis  is  not  feasible  with  the  available  literature  data  only.  The 


24 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


/.• 
•/  • 

•• 


I 


•2 


f  1 


•2-10  1 

FACTOR(2) 


•1  0  1 

FACTORO) 


-3 


I 


T  I  T 


J I I I 


-2 


-1 


0  1  2  3  -4  -3  -2  -1  0  1  2  3 

FACTORC2)  FACTOR(3) 

Fig.  2     Principal  component  analysis  of  the  Mediterranean  complex  (Mandragora  officinarum).  Top  pair:  a.  Bottom  pair  b.  Left:  PC  1,2.  Right:  PC  2,3. 


range  of  the  Turkmenian  plants  is  so  small  that  variability  at  popula- 
tion level  is  considered  equivalent  with  the  variability  over  the  whole 
range.  Mizgireva  ( 1 955)  meticulously  documented  teratogenic  forms 
of  flowers  and  fruits  of  M.  turcomanica,  but  did  not  cite  field 
collected  vouchers  for  these  forms.  She  also  documented  huge 
variability  in  the  shape  and  size  of  tap  roots,  and  described  the 
growth  of  the  plant  in  detail,  from  seed  germination  to  fruiting. 
Mandragora  turcomanica  is  accepted  as  a  distinct  species  on  the 
grounds  of  the  evidence  given  in  Mizgireva  (1942,  1955,  1978)  and 
Lincevskij  (1955).  The  whole  plant  is  much  larger  than  M. 
officinarum.  The  lower  leaves  (±  90  x  60  cm)  are  especially  large 
resulting  in  a  rosette  with  a  diameter  of  150-180  cm.  Mandragora 
turcomanica  blooms  in  both  the  autumn  and  the  spring  (Mizgireva, 


1955,  see  taxonomic  treatment),  with  different  individuals  in  the 
population  exhibiting  one  or  the  other  flowering  time.  Geographi- 
cally the  species  is  isolated,  occuring  only  in  a  narrowly  restricted 
range,  and  within  that  range  it  is  only  found  in  localized  regions 
(Mizgireva,  1955).  It  is  also  widely  disjunct,  being  some  1500  km 
from  the  closest  Mediterranean  Mandragora  population  in  the  Near 
East  and  about  2500  km  from  the  nearest  Himalayan  population  in 
western  Nepal. 

In  common  with  the  Mediterranean  mandrake,  the  Himalayan 
plants  make  up  a  complex  with  considerable  variation  in  size,  shape 
and  colour  of  the  flowers  and  leaves.  The  following  questions  are 
addressed: 


MANDRAGORA 

Is  the  recognition  of  the  four  infraspecific  taxa  sensu 
Grierson  &  Long  within  Mandragora  caulescens  sensible? 

After  checking  herbarium  material  from  almost  the  entire  range  of 
Mandragora  caulescens  (north-western  Sichuan  and  southern 
Qinghai  excluded),  Grierson  &  Long  (1978)  were  well  aware  of  the 
extreme  variability  of  the  morphological  characters  they  used  to 
describe  their  four  subspecies.  The  characters  used  were  corolla 
colour,  calyx  and  corolla  lengths  and  overall  plant  size.  As  with 
Mandragora  officinarum,  flower  colour,  according  to  observations 
of  the  fresh  material  given  on  herbarium  labels,  is  variable  within 
individuals  as  well  as  within  local  populations  (e.g.  Alden  et  al. 
1 185:  'flowers  yellow  to  purplish-brown',  Long  et  al.  266:  'corolla 
yellow,  purple  at  base,  inside  and  out',  and  Polunin  631:  'petals 
yellow  with  purple  veins').  Colours  probably  merely  reflect  to  what 
extent  the  yellow  pigments  are  masked  by  purple  ones.  Grierson  & 
Long  (1978)  were  unable  to  find  any  diagnostic  morphometric 
discontinuities.  Their  proposed  classification  is  based  on  the  rule 
that  due  to  its  type  status,  the  holotype  of  M.  caulescens,  collected  in 
Sikkim  and  representing  an  intermediate  between  the  extremes  of 
the  'typical'  small,  yellow-flowered  plants  and  the  large,  purple- 
flowered  ones,  must  retain  the  name  M.  caulescens  (and  must  thus 
be  given  the  subspecific  epithet  caulescens).  They  therefore  split  the 
complex  not  into  two  but  into  three  subspecies  representing  the  two 
extreme  forms  (subsp.  flavida  and  subsp.  purpurascens)  and  the 
intermediates  (subsp.  caulescens),  and  additionally  described  a 
fourth  subspecies  with  a  shorter  calyx  (subsp.  brevicalyx)  based  on 
only  two  collections  from  Xizang.  As  a  consequence,  their  classifi- 
cation is  based  on  non-diagnostic,  largely  overlapping  characters, 
rendering  M.  caulescens  subsp.  caulescens  as  a  'basket'  taxon  for 
any  intermediates.  Indeed,  it  might  be  hypothesized  that  there  are  at 
least  two  different  species  present  within  this  complex  with  frequent 


25 

introgression  of  the  sympatric  populations.  However,  the  many 
putative  'hybrids'  linking  them  and  resulting  in  an  extensive  overlap 
of  their  features  make  them  unsuitable  as  terminal  taxa  for  the 
cladistic  analysis.  Furthermore,  a  restrictive  taxonomic  concept  as 
described  earlier  does  not  allow  for  the  recognition  of  infraspecific 
taxa  if  their  characters  and  their  ranges  are  extensively  overlapping. 

Are  Mandragora  tibetica  and  M.  chinghaiensis  distinct 
species  or  are  they  both  conspecific  with  M.  caulescens? 

As  in  the  case  of  the  Turkmenian  mandrake,  the  unavailability  of  the 
types  and  a  reasonable  number  of  specimens  renders  the  data 
somewhat  meagre,  so  the  original  descriptions  by  Grubov  ( 1 970)  for 
Mandragora  tibetica  and  by  Kuang  &  Lu  ( 1 978)  for  M.  chinghaiensis 
must  provide  much  of  the  necessary  information.  These  descrip- 
tions, together  with  their  accompanying  keys,  which  distinguish  the 
newly  described  species  from  M.  caulescens,  recognize  the  small, 
yellow-flowered  plants  as  distinct  from  the  larger,  purple-flowered 
ones.  Both  new  taxa  are  therefore  covered  by  the  range  of  variability 
given  in  Grierson  &  Long's  ( 1978)  subdivision  of  M.  caulescens  into 
four  subspecies,  which  are  defined  only  by  overlapping  differential 
characters  and,  as  such,  are  not  acceptable  as  distinct  taxa.  Indeed,  a 
specimen  in  P,  Polunin  et  al.  878,  is  annotated  asM.  chinghaiensis  by 
Lu  An-ming  and  is  also  an  isotype  of  M.  caulescens  subsp.  flavida. 
Some  doubts  must  remain  concerning  M.  tibetica  and  M.  caulescens 
subsp.  brevicalyx  that  together  may  represent  a  single  distinct  taxon 
with  a  very  short  calyx.  An  unusual  specimen  from  western  China 
(Wilson  4195  (K),  no  precise  locality  given)  bears  a  single  flower 
with  a  conspicuously  narrow  corolla.  A  note  by  Grierson  and  Long 
on  the  herbarium  sheet  suggests  that  they  considered  this  specimen 
not  conspecific  or  even  congeneric  with  M.  caulescens.  Without 
further  material,  however,  proposing  a  new  taxon  would  be  just  as 


Fig.  3     The  single  most  parsimonious  cladogram  from  the  HENNIG86  analysis.  The  characters  are  discussed  in  the  text,  and  characters  states  are  shown  in 
Table  5.  For  characters  marked  on  the  branches  of  the  cladogram:  single  lines  indicate  unambiguous  synapomorphies,  parallel  lines  parallelisms  and 
crosses  reversals  (homoplasy). 


26 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


speculative  as  disregarding  the  flower  as  a  mere  artefact  of  prepara- 
tion. The  synonymy  presented  in  the  taxonomic  treatment  here, 
especially  in  respect  to  the  undercollected  plants  from  central  and 
northeastern  Xizang,  is  therefore  provisional  and  subject  to  confir- 
mation once  more  material  of  both  flowering  and  fruiting  plants 
becomes  available.  Only  then  will  it  be  possible  to  judge  whether  or 
not  there  exist  distinct  taxa  not  included  within  the  range  of  variation 
of  M.  caulescens. 

Mandragora  is  a  genus  with  pronounced  variability,  both  within 
and  between  individuals  and  therefore  necessarily  within  the  pro- 
posed polymorphic  species.  Furthermore,  discordant  variation  in  the 
applied  gross  morphological  characters  impedes  a  workable  subdi- 
vision based  on  a  whole  set  of  differential  characters,  rather  than  just 
one  (apparently  non-existent)  diagnostic  character. 

Phylogeny 

The  cladistic  analysis  resulted  in  a  single  most  parsimonious 
cladogram  of  length  =  16,  CI  =  75  and  RI  =  73  (Fig.  3).  The  genus 
Mandragora  defined  in  the  traditional  sense  is  monophyletic  in  our 
analysis:  the  species  share  synapomorphies  in  characters  3  (a  con- 
densed axis),  4  (leaves  clustered  in  a  rosette)  and  9  (fruiting  calyx 
with  prominent  ribs).  If  the  inaperturate  pollen  known  from  M. 
caulescens  and  M.  officinarum  (see  Hoare  &  Knapp,  1997  for 
details)  is  also  found  in  M.  turcomanica,  this  too  would  be  a 
synapomorphy  of  the  genus. 

Mandragora  officinarum  and  M.  turcomanica  are  more  closely 
related  to  each  other  than  to  M.  caulescens.  They  share  reversals  in 
characters  3  (a  condensed  axis)  and  9  (a  fruiting  calyx  without 
prominent  ribs),  but  no  unambiguous  synapomorphies.  The  peculiar 
life-cycle,  i.e.  the  mode  of  development  of  the  leaves  and  flowers,  is 
evidence  for  a  shared  common  ancestry  of  the  Mediterranean- 
Turkmenian  clade.  Mizgireva  (1955)  also  suggested  this  in  her 
revision  of  the  Turkmenian  species.  In  order  to  determine  more 
clearly  the  relationships  among  the  three  species  of  Mandragora,  a 
different,  probably  field-based  set  of  characters  is  necessary.  Here, 
however,  we  are  concerned  more  with  the  delimitation  of  terminal 
taxa,  especially  among  the  Mediterranean  plants,  and  whether  or  not 
the  genus  as  currently  defined  is  monophyletic. 

A  comparison  between  alternative  hypotheses  concerning  the 
position  of  Mandragora  based  on  morphological,  chemical  and 


molecular  (chloroplast  DNA)  characters  is  given  in  Figure  4.  The 
most  pronounced  discrepancy  concerning  the  position  of  Man- 
dragora is  that  the  molecular  data  set  (Fig.  4a)  deviates  from  the 
morphological  (Fig.  4c)  and  chemical  ones  (Fig.  4b)  by  attributing 
this  genus  a  very  independent  position  due  to  extremely  divergent 
and  autapomorphic  cpDNA  sequences.  This  is  reflected  in  the 
proposal  of  a  monogeneric  tribe  Mandragoreae  (Olmstead  et  al.,  in 
press).  Morphological  and  chemical  evidence,  on  the  other  hand, 
suggests  a  placement  within  the  tribe  Hyoscyameae  (Hoare  & 
Knapp,  1997;  =  AtropeaesenswTetenyi,  1987).  The  present  study  is 
far  too  limited  in  its  scope  to  allow  any  conclusions  concerning 
higher  level  classification  within  the  Solanaceae.  However,  it  is  clear 
that  our  limited  data  set  supports  the  inclusion  of  Mandragora 
within  the  Hyoscyameae  (synapomorphies  in  characters  1,  5  and  6) 
but,  as  mentioned  above,  a  much  larger  and  more  field-based  data  set 
will  be  needed  to  address  this  problem.  Mandragora  is  an  anoma- 
lous genus  in  the  family,  both  morphologically  and  in  terms  of 
molecules,  but  this  should  not  prevent  us  from  eventually  identifying 
its  true  affinities  and  relatives  given  adequate  data. 

Biogeography 

The  biogeography  of  an  exclusively  Laurasian  genus  within  the 
Solanaceae  is  of  interest  because  the  family  is  largely  Gondwanan  in 
distribution.  It  is  useful  to  approach  biogeography  from  two  differ- 
ent angles.  Ecological  biogeography  considers  a  short  temporal 
scale  in  evaluating  ecological  factors  relevant  for  present-day  distri- 
butions of  taxa.  Historical  biogeography,  on  the  other  hand,  is 
concerned  with  the  change  of  distributions  and  the  potential  causes 
thereof  on  a  long  temporal  scale.  We  attempt  here  to  summarize 
some  of  the  most  important  data  with  respect  to  the  geographical 
distribution  of  Mandragora,  both  now  and  in  the  past,  and  pinpoint 
major  gaps  in  knowledge. 

Ecological  biogeography 

The  present  approximate  distribution  of  the  genus  is  given  in  Figure 
5.  Apart  from  the  herbarium  material  examined,  the  sources  pre- 
sented in  Table  9  were  used.  Most  comprehensive  are  the  distribution 
data  in  Heldreich  (1886)  and  Vierhapper  (1915).  Mandragora  does 
not  appear  to  penetrate  into  the  Arabian  Peninsula,  being  absent 


ABC 

Fig.  4     Differing  hypotheses  of  the  relationships  of  Mandragora  within  the  Solanaceae.  a:  Olmstead  et  al.,  in  press,  b:  Tetenyi,  1987,  c:  Hoare  &  Knapp,  1997. 


MANDRAGORA 


27 


20 


Fig.  5     Distribution  of  the  genus  Mandragom.  1)  M.  officinarum,  2)  M.  turcomanica,  and  3)  M.  caulescens. 


even  in  the  most  northern  parts  of  Saudi  Arabia  (Migahid,  1989)  as 
well  as  from  Mesopotamia  (Rechinger,  1964;  Mouterde,  1979).  The 
genus  was  naturalized  in  Malta  in  the  nineteenth  century  but  is  now 
presumably  extinct  there  (Borg,  1927;  Maugini,  1959).  Mandra- 
gora  does  not  appear  in  works  covering  France  and  Corsica  (Fournier, 
1990),  Albania  (Demiri,  1983),  Egypt  (Tackholm,  1974;  Boulos, 
1995)  or  Libya  (Siddiqi,  1978)  and  is  presumed  absent  from  these 
areas.  Likewise,  the  genus  is  absent  from  the  Macaronesian  archi- 
pelagos of  the  Canaries,  Madeira  and  the  Azores  (not  recorded  either 
by  Hansen  &  Sunding,  1993;  or  Press  &  Short,  1994). 

Mandragora  turcomanica  is  found  only  in  the  Kopet  Dag  range  of 
southern  Turkmenistan.  It  has  not  yet  been  reported  from  the  Iranian 
part  of  these  mountains  (Rechinger,  1972).  In  eastern  Asia,  Man- 
dragora (M.  caulescens)  occurs  in  the  central  and  eastern  Himalaya 

Table  9     Sources  used  for  determining  the  overall  distribution  of  the 
genus  Mandragora. 


Geographical  area 


Reference 


Tunisia 
Algeria 
Morocco 
Portugal 
Spain  (incl. 

Balearic  Is.) 
Italy  (incl.  Sicily, 

Sardinia) 

Former  Yugoslavia 
Greece  (incl.  Aegean  Is.) 
Turkey 
Cyprus 
Syria 

Lebanon 

Israel 

Jordan 

Turkmenistan 

Nepal 

India  (Sikkim, 

Arunachal  Pradesh) 
Bhutan 
Myanmar 
China 


Bouquet  (1952),  Pettier- Alapetite  (1981) 

Quezel&  Santa  (1963) 

Jahandiez  &  Maire  (1934) 

Amaral  Franco  (1984) 

Lazaro  e  Ibiza  (1907),  Bonafe  Barcelo  (1980), 

Valdesetal.  (1987) 
Maugini  (1959),  Pignatti  (1982), 

Cellinese  et  al.  (1994) 
Hayek  &Markgraf(  1931) 
Halacsy  (1902),  Hayek  &  Markgraf  (1931) 
Baytop(1978) 
Meikle  (1 985),  Viney  ( 1994) 
Post  (1933),  Feinbrun-Dothan  (1978), 

Mouterde  (1979) 
Post  (1933),  Feinbrun-Dothan  (1978), 

Mouterde  (1979) 

Post  (1933),  Feinbrun-Dothan  (1978) 
Post  (1933),  Feinbrun-Dothan  (1978) 
Pojarkova  (1955),  Kurbanov  ( 1 994) 
Grierson&  Long  (1978) 
Hara  (1966),  Grierson  &  Long  (1978),  Deb(1979) 

Hara  (1971),  Grierson  &  Long  (1978) 
Grierson  &  Long  (1978) 
Grierson  &  Long  (1978),  Zhang  et  al.  (1994). 
Zhang  &Lu(  1994) 


and  southwestern  China,  but  there  are  no  records  from  the  western 
extensions  of  these  mountain  ranges  (Karakorum,  Pamir,  Hindu 
Kush  and  Tien  Shan). 

Apart  from  historical  factors,  dealt  with  below,  the  principal 
factor  determining  the  geographical  distribution  of  taxa  on  a  large 
scale  is  typically  climate.  Although  one  might  disagree  on  the 
relative  importance  of  means  and  extreme  values  of  various  climatic 
variables,  the  prime  importance  of  climate  in  general  has  long  been 
accepted  (Cain,  1944). 

A  convenient  way  of  depicting  and  comparing  different  climates 
is  by  means  of  standardized  diagrams.  The  purpose  of  the  so-called 
ecological  climate-diagrams  is  to  portray  the  climate  of  a  locality  in 
a  readily  comparable  way  and  visually  to  present  the  most  important 
climatic  factors  affecting  the  growth  of  plants  (Walter  et  al.,  1975). 
By  choosing  a  scale  at  which  10°C  correspond  to  20  mm  precipita- 
tion, a  relatively  wet  (dark)  and  relatively  dry  (light)  season  can  be 
identified  on  the  diagrams  (Fig.  6).  These  climate  diagrams  make  it 
evident  that  the  range  of  Mandragora  comprises  not  only  quantita- 
tively different  climates  but  qualitatively  different  climate  types. 
The  amount  and  especially  the  main  season  of  rainfall  is  particularly 
significant. 

The  Mediterranean  climate  is  characterized  by  mild,  rainy  winters 
and  hot,  dry  summers.  Although  the  total  amount  of  rainfall  for  other 
locations  for  Mandragora  officinarum  may  differ  somewhat  from 
that  presented  here  (Iraklio,  on  the  northern  coast  of  Crete,  Fig.  6), 
the  general  aspect  of  a  winter  rainfall  regime  remains  the  same  all 
over  the  Mediterranean  basin,  even  extending  towards  Afghanistan 
and  the  Hindu  Kush.  The  typical  vegetation  consists  of  adapted 
sclerophyllous  vegetation,  analogous  to  that  found  in  the  same 
climate  type  in  California,  Chile,  the  Cape  region  of  South  Africa 
and  southwestern  Australia. 

The  Kopet  Dag  is  a  Transcaspian  mountain  range  surrounded  by 
the  Irano-Turanian  desert.  The  climate  there  represents  a  continen- 
tal, dry  version  of  the  Mediterranean  type  with  much  lower  rainfall 
(Fig.  6:  Kisil-Arvat))  and  an  extended  summer  arid  period.  During 
this  dry  summer,  Mandragora  turcomanica  exhibits  a  long  period  of 
xerothermic  dormancy  (Kurbanov,  1994).  Mizgireva  (1955)  sug- 
gested that  M.  turcomanica  is  much  more  cold-tolerant  than  its 
mediterranean  congeners.  The  typical  vegetation  in  the  Kopet  Dag 
consists  of  broadleafed  deciduous  forest  in  the  moister  microhabitats 
and  open  shrubland  in  drier  areas. 

The  eastern  part  of  the  Mandragora  range  in  the  Sino-Himalayan 


28 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


Irakiio  27  m  alt. 


Gangtok  1725m  alt. 


IN 
so 
M 
4* 
20 
0 


Kisil-Arvat  101  m  alt. 


100 

m 
M 

40 
20 
0 


30 
20 
10 
0 


Gyantse  3996  m  alt. 


A 


Fig.  6    Climate  diagrams  of  four  representative  weather  stations  in  three 
different  climate  regimes.  Given  are  the  rainfall  and  temperature  curves, 
resulting  in  wet  (dark  shading)  and  dry  (light  shading)  seasons.  Based 
on  Walter  et  al.  1975. 

region  is  characterized  by  a  completely  different  rainfall  regime 
altogether.  The  influence  of  the  monsoon  renders  not  the  winter  but 
the  summer  the  season  of  the  highest  rainfall.  The  amount  of 
precipitation  during  the  monsoon  period,  however,  does  vary  enor- 
mously. The  Himalayan  chain  functions  as  a  rain  screen  resulting  in 
high  rainfall  on  the  southern  slope  of  the  mountain  range  (Fig.  6: 
Gangtok)  as  well  as  in  the  southwestern  Chinese  mountains  of 
Yunnan  and  Sichuan,  and  much  lower  rainfall  in  the  inner  regions  of 
Xizang  (Fig.  6:  Gyantse).  Even  at  high  altitudes,  winters  are  very 
mild  in  the  high  rainfall  areas  (see  climate-diagram  of  Gangtok, 
about  1700  m  above  sea  level)  and  snow  is  scarce.  Generally 
speaking,  the  eastern  Himalaya  and  southeastern  Xizang,  Sichuan 
and  Yunnan  enjoy  a  much  more  humid  and  milder  climate  than  the 
western  Himalaya  and  central  Xizang.  The  locality  data  of  the 
examined  specimens  suggest  that  the  distributional  boundary  of  the 
Himalayan  mandrake  towards  the  central  Xizang  highland  lies 
within  the  200-400  mm  annual  precipitation  strip  north  of  the  main 
ranges.  Due  to  the  immense  altitudinal  gradient,  the  vegetation  is 
strikingly  zonal  and  ranges  from  subtropical  forests  to  alpine  scrub 
and  meadows.  The  upper  forest  limit  lies  between  about  3600  and 
4200  m,  the  upper  tree  limit  between  4200  and  4500  m.  The  belt  in 


between  is  characterized  mainly  by  a  succession  of  various  Rhodo- 
dendron species.  This  is  the  typical  habitat  of  Mandragora  caule- 
scens.  The  permanent  snow  line  lies  at  about  4800  m  in  the  south  and 
5800  m  in  the  north  towards  central  Xizang  (Walter,  1962). 

Other  factors  affecting  plant  distribution,  albeit  on  a  smaller  and 
less  important  scale,  include  soil  type  and  competition.  Man dragora 
officinarum  especially  grows  in  ruderal  habitats,  on  clayey  and 
nitrogen-rich  soils.  So  far,  it  is  not  clear  whether  M.  caulescens 
occurs  on  acidic  (granitic)  or  base-rich  (limestone)  soils  in  the 
subalpine  and  alpine  zone  of  the  Himalaya.  It  cannot  be  ruled  out 
that  there  might  be  two  different,  vicariant  ecological  groups  as  is 
often  the  case  in  the  Alps,  for  example  in  Rhododendron  (Walter, 
1966).  The  type  of  climate  and  soil  typically  only  indirectly  deter- 
mine the  occurrence  of  a  species  in  a  certain  locality,  in  that  they 
affect  the  fitness  of  that  species  in  relation  to  others  (Walter,  1977). 
Both  Mandragora  officinarum  and  M.  turcomanica  seem  more  or 
less  restricted  to  low-competition  localities,  representing  typical  r- 
species  (good  colonizers  but  poor  competitors). 

Historical  biogeography 

The  atemporal  cladogram  of  a  group  together  with  historical  evi- 
dence, namely  paleoclimatic  and  paleogeographical  data  allow  for 
the  formulation  of  historical  scenarios.  A  scenario  is  a  model  of  the 
basic  features  (including  space  and  time)  of  the  evolutionary  history 
of  a  group  which  can  serve  to  stimulate  directions  of  further  investi- 
gation. However,  the  essentially  speculative  character  of  such  a 
model  renders  it  more  like  a  tool  than  a  serious  scientific  statement 
(Eldredge,  1979)  and  ad  hoc  assumptions  are  inevitable  (Humphries 
&  Parenti,  1986).  In  attempting  to  construct  scenarios  for  the 
historical  biogeography  of  any  group,  plate  movement,  eustasy 
(worldwide  changes  of  sea  level)  and  climate  changes  are  consid- 
ered the  major  factors  to  be  considered  (Hallam,  1981).  In  the  case 
of  the  Solanaceae,  virtually  no  fossil  record  is  available  (PaleoNet 
Listserver,  http://www.ucmp.berkeley.edu/Paleonet/),  thus  histori- 
cal hypotheses  concerning  Mandragora  are  inevitably  founded  on 
extant  taxa  only. 

The  Mediterranean  region  presents  a  very  complex  tectonic  jig- 
saw puzzle,  hard  to  resolve  adequately  (Hallam,  1981).  It  seems 
clear,  however,  that  during  most  of  the  Tertiary,  summer  was  the 
rainy  season  and  only  before  the  Pleistocene  was  there  a  shift 
towards  the  current  pattern  of  rainfall  in  winter.  The  plants  had  to 
adapt  and  reduce  activity  during  the  dry  summer  months  (Walter, 
1977).  The  northern  Mediterranean  localities  for  Mandragora  in 
northern  Italy  and  on  the  Dalmatian  coast  are  very  patchy,  resulting 
in  a  highly  fragmented  range  there,  and  the  precise  status  of  these 
populations  is  uncertain.  We  have  been  unable  to  discover  any 
modern  collections  from  these  regions  (Cellinese  et  al.,  1994)  and 
some,  or  all,  of  the  populations  may  no  longer  exist.  Vierhapper 
(1915)  maintained  that  these  plants  are  Tertiary  relics,  the  scattered 
remnants  of  a  formerly  larger,  continuous  range,  that  have  been 
unable  to  adapt  to  the  changing  climate  and  still  exhibit  the  vernal 
phenology  type.  Relict  status  based  on  Tertiary  processes,  however, 
is  normally  attributed  to  groups  of  far  wider  disjunct  ranges  than  is 
the  case  here.  Hawkes  ( 1972a)  based  his  definition  of  M.  officinarum 
L.  on  these  northern  Italian  and  Dalmatian  populations  only,  using 
corolla  colour  and  phenology.  On  the  evidence  of  our  multivariate 
analyses,  Hawkes's  position  is  untenable,  since  the  plants  fall  within 
the  range  of  variation  exhibited  by  the  rest  of  the  Mediterranean 
plants.  It  could  be  argued  that  the  different  corolla  colour  (white)  of 
the  northern  Italian  and  Dalmatian  plants,  coupled  with  their  dis- 
crete distribution  is  sufficient  to  recognize  them  at  least  at  subspecific 
rank.  We  have  already  pointed  out  that  the  corolla  colour  is  only  an 


MANDRAGORA 


29 


apparent  distinction,  with  similar  colours  occurring  in  populations 
outside  northern  Italy  and  Dalmatia.  The  somewhat  isolated  distri- 
bution alone  is  insufficient  reason  to  recognize  the  Italian  and 
Dalmatian  plants  as  a  separate  taxon. 

The  close  relationship  between  the  Mediterranean  and  the 
Turkmenian  mandrake  as  suggested  by  the  cladistic  analysis  is 
reflected  in  a  linked  history  of  the  regions  particularly  during  the 
Tertiary.  It  can  be  proposed  that  in  this  period,  after  the  Tethys  Sea 
receded  and  the  middle  Asian  mountains  became  exposed  during 
their  simultaneous  uplift,  a  vicariance  event  of  the  Mediterranean 
and  Turkmenian  ancestors  took  place.  Both  extant  species  show  a 
comparable  adaptation  to  the  aridization  and  change  from  a  summer 
to  a  winter  rainfall  regime.  In  other  words,  Mandragora  turcomanica 
can  be  seen  as  an  example  of  a  plant  that  has  been  evolving  in 
isolation  in  Turkmenistan  since  the  Tertiary  from  ancient  Tethyan 
stock  (Kurbanov,  1994).  It  is  of  some  interest  to  note  that  there  is  a 
highly  congruent  pattern  within  Atropa  where  the  disjunct  distribu- 
tion of  the  Turkmenian  A tropa  komarovii  Blin.  &  Schal.,  consists  of 
a  very  narrow  range  south  of  the  river  Sumbar  only  a  few  miles  away 
from  the  Mandragora  turcomanica  population  north  of  the  river 
(Kurbanov,  1994). 

The  Sino-Himalayan  area  is  of  particular  interest  because  of  its 
position  at  the  transition  between  the  Palearctic  and  the  Oriental 
floristic  realms.  The  period  of  the  collision  of  the  Indian  sub- 
continent with  Asia  is  contentious  but  probably  occurred  in  the 
Paleogene  approximately  50  million  years  ago  (Scotese  et  al.,  1988) 
with  the  penetration  of  the  Eurasian  border  continuing  during  the 
early  Neogene  (c.  20  millions  years  ago)  (Dercourt  et  al.,  1986).  The 
connection  between  Tethys  and  the  Mediterranean,  on  the  other 
hand,  probably  did  not  close  until  the  mid-Miocene  (c.  15  million 
years  ago)  (Hallam,  1994).  Climate  changes  during  the  Paleogene 
were  severe.  Temperatures  increased  up  to  the  early  Eocene  (c.  55 
million  years  ago)  followed  by  a  global  cooling  which  marks  the 
passage  from  a  warm  domain  of  the  Mesozoic  and  early  Palaeogene 
to  the  cool  domain  thereafter  (Hallam,  1994).  According  to  Rud- 
diman  et  al.  (1989)  much  of  the  uplift  in  the  Xizang  plateau  region 
occurred  only  in  the  last  5  to  10  million  years  (Plio-Pleistocene 
uplift).  It  must  be  assumed  that  the  glaciation  of  the  high  altitude 
regions  of  the  Sino-Himalayan  area  during  the  Pleistocene  (Ice- 
Ages)  was  more  or  less  complete,  posing  the  question  whether 
refugia  further  south  and  at  lower  altitudes  might  have  served  as 
speciation  centres. 

Schuster  (1976)  argued  that  the  Indian  plate  served  as  a  vehicle  for 
the  dissemination  of  Gonwanan  taxa  into  Laurasia.  But  at  the  same 
time  he  expected  that  due  to  the  migration  through  several  climatic 
zones  the  Indian  plate  probably  arrived  with  a  rather  depleted  and 
altered  flora  and  therefore  only  remnants  of  the  original  Gondwanan 
flora  are  recognizable  in  India  today.  If  angiosperms  were  indeed  on 
the  Indian  plate  in  early  Cretaceous  time,  they  could  probably  not  be 
assigned  to  presently  recognizable  families.  On  the  basis  of  a 
proposed  late  Triassic  (c.  210  million  years  ago)  start  of  the  Indian 
plate  migration,  serious  doubt  is  cast  on  the  possibility  that  early 
Solanaceae  could  have  been  'on  board'.  The  main  objection  is  that 
the  plate  drift  simply  occurred  too  early,  before  modern  groups  were 
in  existence.  However,  other  data  (Smith  et  al.,  1994)  suggest  that 
India  did  not  break  free  from  Gondwana  until  the  late  Jurassic  to 
early  Cretaceous  (130-140  million  years)  or  even  near  the  Creta- 
ceous-Tertiary boundary  only  60-65  million  years  ago  (Hallam, 
1994).  Lu  &  Zhang  (1986)  proposed  southwestern  China  as  a  later, 
secondary  speciation  centre  of  the  whole  tribe  Hyoscyameae  on  the 
grounds  of  its  high  species-richness. 

A  different  biogeographical  scenario  proposes  a  migration  from 
Gondwana  to  the  Mediterranean  basin.  Migration  routes  that  are 


closed  today  were  wide  open  in  the  Mesozoic.  Indeed,  massive  and 
relatively  unimpeded  migrations  from  Gondwana  to  Laurasia  might 
have  been  feasible  until  the  late  Cretaceous  (Schuster,  1976).  Con- 
trary to  the  first  scenario,  the  mandrake  ancestors  would  therefore 
have  spread  eastwards  from  the  Protomediterranean  to  the  develop- 
ing eastern  Asian  mountain  ranges  where  the  massive  uplift  possibly 
resulted  in  a  vicariance  event.  The  Mediterranean  populations  on  the 
other  hand  would  have  had  to  adapt  to  the  increasing  aridization  in 
the  western  part  of  the  continent  during  the  late  Tertiary  (Weinert, 
1972)  and  thus  slowly  changed  their  mode  of  development  from  a 
summer  flowering  to  a  autumn- winter-spring  flowering  rhythm. 
Proskuryakova  &  Belyanina  (1985)  claim  that  the  slow  succession 
of  the  development  phases,  particularly  of  the  germination,  the  long 
growth  and  flowering  period  and  the  large  size  of  the  plant  speak  for 
the  great  age  of  the  Mediterranean  group.  They  draw  the  conclusion 
that  the  Mediterranean  section  must  be  older,  representing  a  Terti- 
ary relic,  than  the  apparently  highly  adapted  Himalayan  mandrake. 
The  Himalayan  mandrake,  in  fact,  might  have  arisen  from  now- 
extinct  ancestors  as  late  as  the  Quaternary  glaciations. 

To  conclude,  the  massive  timespan  covered  in  the  Tertiary,  the 
many  fluctuations  in  climate  and  transgressions  of  the  seas  and  the 
current  very  inaccurate  timing  of  even  the  major  events  render 
definitive  statements  or  falsifications  concerning  the  above  scenar- 
ios and  the  history  of  Mandragora  hardly  possible.  In  this  context 
the  distinction  between  a  cladogram  (what  is  more  closely  related?) 
and  the  actual  phylogenetic  tree  (what  is  the  ancestor?)  must  be 
stressed.  The  cladogram  (Fig.  3)  suggesting  a  more  recent  common 
ancestry  of  Mandragora  officinarum  and  M.  turcomanica  is  con- 
gruent with  only  one  of  the  many  possible  trees  equally  resulting  in 
this  particular  cladogram,  which  therefore  represents  a  far  less 
informative  (more  general)  mode  of  expression.  Thus,  it  does  not 
allow  for  statements  concerning  the  ancestor  and  does  not  imply  that 
Mandragora  caulescens  is  the  older  taxon.  Nevertheless,  it  can  be 
proposed  that  the  major  events  affecting  speciation  and  present 
distribution  of  Mandragora  in  the  Mediterranean-Turkmenian  area 
were  probably  the  receding  of  the  Tethys  and  the  aridization  of  the 
climate  in  the  Tertiary.  In  the  Sino-Himalayan  area,  on  the  other 
hand,  the  collision  of  the  Indian  plate  and  the  subsequent  uplift  of  the 
Himalayan  range,  as  well  as  the  later  glaciation  periods  during  the 
Quaternary,  are  probably  most  important. 


TAXONOMIC  TREATMENT 

Mandragora  L.,  Sp.  pi.   1:   181  (1753).    Type  species:  M. 
officinarum  L. 

Perennial  herbs.  Stout  tap-root,  often  branched,  sometimes  vaguely 
anthropomorphic.  Stem  very  short  or  with  secondary  growth  during 
flowering  and  fruiting,  sometimes  branched.  Leaves  simple,  nor- 
mally entire,  occasionally  toothed.  Inflorescence  of  typically  solitary 
axillary,  pedicellate  flowers.  Floral  envelope  actinomorphic.  Calyx 
5(6)-lobed,  persistent,  enlarged  in  fruit.  Corolla  5(6)-lobed.  Sta- 
mens 5(6),  included,  inserted  in  lower  half  of  corolla-tube;  filaments 
filiform,  pubescent  at  base;  anthers  dorsifixed,  dehiscing  longitudi- 
nally. Ovary  2-locular;  ovules  numerous;  style  elongate;  stigma 
capitate  or  somewhat  bilobed.  Fruit  a  fleshy,  many  seeded  berry, 
globose  or  ovoid,  borne  beneath  the  leaves  on  nodding  pedicel  or 
lying  on  the  ground.  Seeds  compressed;  embryo  curved. 

FLORAL  FORMULA.    K(5)  C(5)  A5  G(2) 

DIVERSITY  AND  DISTRIBUTION.  Three  disjunct  species:  Mediterra- 
nean region,  central  Asia,  Sino-Himalayan  region. 


30 

PHENOLOGY  AND  HABITAT.  All  three  Mandragora  species  are 
geophytes  with  a  conspicuous  tap-root  which  is  the  perennating 
organ  of  these  herbs,  linking  the  annual  vegetative  periods.  The 
Turkmenian  mandrake  with  its  prolonged  dry  season  dormancy  has 
been  referred  to  as  an  ephemeroid  (Proskuryakova  &  Belyanina, 
1985),  its  rhythm  of  development  more  closely  related  to  the  Medi- 
terranean than  to  the  Himalayan  climatic  cycle.  Propagation  is 
usually  by  seeds  or  vegetatively  by  underground  buds  on  the  tap- 
root. The  species  grow  in  open  woodland,  olive  groves,  grassland, 
fallow  land,  waysides,  railway  embankments,  ruins,  in  rock  crev- 
ices, grassy  and  stony  mountain  slopes  and  screes.  The  altitudinal 
range  is  0-1200  and  3000-4900  m. 

ETHNOBOTANY.  Once  important  medicinal  plants  with  a  long  tra- 
dition in  mythology  and  superstition.  All  parts  of  the  plant  contain 
potent  narcotic  alkaloids. 

ETYMOLOGY.  Origin  uncertain,  possibly  from  the  Greek  |0,otv- 
Spcryopaa,  original  meaning  controversial;  alternatively  from  the 
Sanskrit  mandros,  meaning  sleep  and  agora,  meaning  object  or 
substance  (Thompson,  1934). 

Key  to  the  species  of  Mandragora 

1  Young  leaves  arranged  in  a  cluster  on  top  of  a  stem  with  scales,  later 
along  the  secondarily  elongated,  sometimes  branched  stem;  corolla 
yellow  to  dark  purple;  style  not  exceeding  stamens.  Sino-Himalayan. 
Alt.  3000-4900  m 3.  M.  caulescens 

Leaves  permanently  arranged  in  a  rosette,  the  stem  not  elongating; 
corolla  white  to  pale  blue  or  violet;  style  exceeding  stamens 2 

2  Mature  lower  leaves  in  a  large  rosette  of  1 50- 1 80  cm;  berries  40-60  mm 
in  diameter.  Turkmenian.  Alt.  500-700  m 2.  M.  turcomanica 

Mature  lower  leaves  in  a  rosette  of  less  than  1 00  cm;  berries  less  than  40 
mm  in  diameter.  Mediterranean.  Alt.  0-1200  m  ...  1.  M.  officinarum 

1.  Mandragora  officinarum  L.,  Sp.  pi.  1:  181  (1753).  Type:  Herb. 

Burser  IX:  26  (UPS-lectotype,  designated  by  Knapp  in  Jarvis  et 

al.  (1993),  microfiche-BM!). 
Figs  7,  8. 

=   Atropa  mandragora  L.,  Syst.  nal   10th  ed.:  933  (1759),  nom. 

illeg.  (Art.  52.1). 
=  Mandragora  foemina  Garsault,  Fig.  pi.  med.  3:  221,  t.  363  p.p. 

(1764). 
=   Mandragora  mas  Garsault,  Fig.  pi.  med.  3:  221,  t.  363  p.p. 

(1764). 
=   Mandragora  acaulis  Gaertn.,  Fruct.  sem.  pi.  2:  236,  t.   131 

(1791). 

=  Atropa  humilis  Salisb.,  Prodr.  stirp.  Chap.  Allerton:  132  (1796). 
=   Atropa  acaulis  Stokes,  Bot.  mat.  med.  1:  396  (1812). 
=   Mandragora  autumnalis  Bertol.,  Elench.  plant,  viv.:  6  (1820). 
=  Mandragora  vernalis  Bertol.,  Virid.  Bonon.  veg.:  6  (1824). 
=  Mandragora  praecox  Sweet,  Brit.fl.  gard.  2:  t.  198  (1827). 
=  Mandragora  neglecta  G.  Don  ex  Loudon,  Hon.  brit.:  71  (1830). 
=  Mandragora  microcarpa  Bertol.,  Comm.  Mandrag.:   12,  t.  3 

(1835). 
=  Mandragora  haussknechtii  Heldr.  in  Mitt.  Geogr.  Ges.  Jena.  4: 

77,  app.  (1886). 
=   Mandragora  x  hybrida  Hausskn.  &  Heldr.  in  Mitt.  Geogr.  Ges. 

Jena.  4:77(1886). 
=   Mandragora  hispanica  Vierh.  in  Osterr.  Bot.  Z.  65:  132-133 

(1915). 

ILLUSTRATIONS.     Wettstein  (1895:  fig.  12M-O);  Feinbrun-Dothan 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 

(1978:  fig.  278);  Pignatti  (1982:  figs  on  p.  518);  Hoare  &  Knapp 
( 1997:  fig.  5c). 

Plant  virtually  stemless.  Root  stout,  often  branched,  very  long. 
Leaves  arranged  in  a  rosette,  leaf  shape  and  size  very  variable,  but 
usually  elliptic  to  obovate,  max.  45  cm  long,  length/width  ratio  1 .5- 
10,  variously  pubescent.  Flowers  borne  in  leaf  axils.  Pedicels  very 
variable  in  length,  max.  15  cm.  Calyx  6-28  mm  long,  lobed  '/2-2/3  of 
the  way  to  the  base,  lobes  3-15  mm,  acuminate,  only  slightly 
accrescent  in  fruit.  Corolla  12-65  mm  long,  campanulate,  greenish 
white  to  pale  blue  or  violet,  lobed  l/2  to  nearly  to  the  base,  the  lobes 
6-60  mm.  Stamens  adnate  to  lower  half  of  corolla;  filaments  7-15 
mm  long;  anthers  2.5^.0  mm  long,  yellow  to  brown,  sometimes 
pale  blue.  Style  8-20  mm,  exceeding  stamens;  stigma  capitate.  Fruit 
5^0  mm  in  diameter,  globose  to  ellipsoid,  glossy  yellow  to  orange 
when  ripe,  juicy,  edible.  Seeds  2.5-6.0  mm  long,  reniform,  the 
surface  reticulate,  yellow  to  light  brown. 

HABITAT.  Open  woodland,  olive  groves,  fallow  land,  waysides, 
railway  embankments,  ruins,  crevices.  Altitudinal  range  0-1200  m. 

VERNACULAR  NAME.     Mediterranean  mandrake. 

PHENOLOGY.  Flowering  September  to  April.  Fruiting  November 
to  June. 

CONSERVATION  STATUS.  Populations  are  scattered  throughout  the 
species  range.  Northern  Italian  populations  are  vulnerable  (Cellinese 
et  al.,  1994)  as  they  are  apparently  known  only  from  old  herbarium 
records  and  the  present  occurrence  and  abundance  of  these 
populations  are  unclear. 

DISTRIBUTION.  Circum-Mediterranean.  (Tunisia,  Algeria,  Mo- 
rocco, Portugal,  Spain,  Italy,  former  Yugoslavia,  Greece,  Cyprus, 
Turkey,  Syria,  Lebanon,  Israel,  Jordan). 

SELECTED  SPECIMENS  EXAMINED.    TUISIA.  Cap  Bon,  roadsides  and 

orchards,  0-150  m,  6  October  1969,  Tomkinson,  M.J.  72,  72A  (BM).  Nabel, 
60  m,  14  October  1907,  Gandoger,  M.  84  (K). 

MOROCCO.  Guercif,  144  km  E.  of  Fes  near  'La  Kazbah',  38  km 
before  the  town  of  Guercif,  34°16'N,  3°45'W,  510  m,  disturbed  gravelly 
area  by  road,  3  November  1994,  Jury,  S.L.  et  al.  15434  (RNG);  Gharb,  c.  4 
km  NNE  of  Souk-El-Arbaa-du-Rharb  on  road  to  Ksar-El-Kebir,  by  bridge 
and  salt  works,  34°43'N,  6°00'W,  30  m,  pasture  between  fields  at  side  of 
salt  pans,  7  November  1994,  Jury,  S.L.  et  al.  15701  (RNG);  Zerhoun,  c.  3 
km  NW  of  Moulay  Idriss  and  33  km  from  Meknes  along  road  to  Sidi 
Kacem  and  Tanger,  34°04'N,  5°33'W,  390  m,  ruins  of  Roman  city  of 
Volubilis,  1  November  1994,  Jury,  S.L  et  al.  15398  (RNG);  Prerif  Central, 
N.  of  Fes,  on  the  road  to  Ouezzane,  34°22'N,  5°09'W,  210  m,  roadside 
ruderal  community,  24  October  1993,  Jury,  S.L  et  al.  12634  (RNG);  Haul 
Ouerrha,  c.  50  km  N.  of  Fes  on  the  road  between  Fes-el-Bali  and  Ourtzarh, 
34°33'N,  5°02'W,  120  m,  in  olive  plantation  on  steep  N-facing  slope  with 
Zizyphus  lotus,  24  October  1993,  Jury,  S.L  et  al.  12653  (RNG);  Temara,  10 
km  S.  of  Rabat,  0-30  m,  sandy  roadside,  14-20  October  1981,  Lewalle,  J. 
10054  (BM,  RNG);  5  km  W.  of  Oued,  Beht  on  Rabat-Meknes  road, 
33°52'N,  5°58'W,  400  m,  27  September  1991,  Gardner,  M.F.  et  al.  4885  (E, 
RNG);  Loukkos,  SW  of  Chefchaouen,  on  road  to  Ksar-el-Kebir,  35°1'N, 
5°45'W,  120  m,  in  cultivated  fields  of  wheat,  21  October  1993,  Jury,  S.L.  et 
al.  12542  (RNG);  Loukkos,  c.  1  km  E.  of  Larache  on  road  to  Asilah,  c.  200 
m  from  Oued  Loukkos,  35°12'N,  6°08'W,  5  m,  at  side  of  road  by  saltpans, 
28  October  1994,  Jury,  S.L  et  al.  15237  (RNG),  Tanga,  just  N.  of  airport, 
road  to  Cap  Spartel,  35°45'N,  5°55'W,  35  m,  fallow  area  at  side  of  ploughed 
field  with  Urginea  maritima,  16  October  1993,  Jury,  S.L.  et  al.  12217 
(RNG),  Tanga,  E.  on  road  to  Sebta  (Ceuta),  Oued  Dliane,  35°50'N,  5°39'W, 
5  m,  nitrophilous  areas  on  fallow  cultivated  land  on  heavy  clay  alluvial 
plain,  17  October  1993,  Jury,  S.L.  et  al.  12238  (RNG);  Allal-Tazi,  close  to 
the  river  Sedou,  1  November  1965,  Talon,  A.  1630  (BM). 

SPAIN.  Baleares,  Palma,  between  S.  Sardina  and  S.  Bernardo,  22  Septem- 
ber 1913,  Bianor  &  Sabasien  1877  (BM,  E,  P,  RNG);  Province  Cadiz, 


MANDRAGORA 


Fig.  7     Mandragora  officinarum.  a)  Flowering,  cultivated  at  RBG  Kew,  b)  Fruiting,  cultivated  at  RBG  Kew. 


32 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


MANDRAGORA 


33 


Chiclana,  on  clayey  soil  by  the  river,  10  January  1 855,  Bourgeau,  E.  2304  (E, 
K,  P);Torredonjimeno,  600 m,  10 December  1986,  Espinosa,  A.  870252  (K); 
alrededores  de  Cordoba,  Cuesta  del  Espino,  2  December  1967,  Galiano,  E.F. 
&  Valdes,  B.  820  (E,  RNG);  Province  Malaga,  C  341  from  Campillos  to 
Ronda,  36°52'N,  5°00'W,  525  m,  disturbed  ground  near  new  road  cutting,  27 
October  1990,  Gardner,  M.F.  &  Knees,  S.G.  4769  (RNG);  Province  Cadiz,  23 
km  from  Jerez  de  la  Frontera,  36°32'N,  5°59'W,  1 30  m,  ruderal  area  by  road 
with  abundant  Silybum  marianum  below  ruined  castle  of  El  Pedroso,  22 
December  1993,  Jury,  S.L.  &  Jury,  L.C.  13111  (BM,  RNG);  Province  Ja6n, 
Torredonjimeno,  between  Porcuna  andTorredonjimeno,  near  Pilar  de  Moya, 
500  m,  roadside  on  chalky  soil,  3  November  1979,  Ladero,  M.  et  al.  10605 
(RNG);  Malaga,  near  Churiana,  3  December  1957,  Partridge,  F.  2  (BM); 
Campamento,  9  November  1911,  Wolley-Dod,  A.M.  22  (BM). 

ITALY.  Sicily,  Province  Palermo,  Madonie,  near  Polizzi,  950  m,  rocky 
limestone  slopes,  1 8  August  1964,  Davis,  P.M.  40134  (E);  Sicily,  Trapani,  200 
m,  steep  slopes,  25  August  1964,  Davis,  P.M.  40182  (RNG);  Sicily,  Province 
Siracusa,  Augusta,  October-November  1911,  Vaccari,  A.  2326  (BM,  K). 

FORMER  YUGOSLAVIA.  Hercegovinae,  Doljnja  Glavska,  6  May 
1908,  Reiser,  C.  s.n.  (K);  Silieski,  near  Ragusa,  500  m,  29  April  1909, 
Maly,  K.  s.n.  (K). 

GREECE.  Near  Corinth,  30  m,  waste  grounds  and  waysides,  April  1931, 
Atchley,  S.C.  819  (K);  Crete,  Frangokastello,  29  March  1973,  Barclay,  C. 
3004  (K);  Crete,  Canea,  1  December  1943,  Bickerich,  G.  15324  (BM,  K); 
Crete,  Kolokithia,  December  1977,  Bowen,  H.J.M.  694  (RNG);  Crete,  Agios 
Nikolaos,  December  1983,  Bowen,  H.J.M.  3088  (RNG);  Peloponnese,  5  km 
from  Pilos  on  the  road  from  Kalamata,  250  m,  in  cultivated  fields  near  stream, 
12  November  1973,  Brickell,  C.  &  Mathew,  B.  8088  (K);  Crete,  Canea, 
frequent  in  hills  and  ditches  throughout  Crete  up  to  300  m,  1  January  1940, 
Davis,  P.H.  1100(E,K);Cyclades,Amorgos,  lOApril  1940,  Davis,  P.M.  1438 
(K);  Dodecanese,  Kos,  Asfendion,  300-600  m,  rocky  slopes,  27  March  1965, 
Davis,  P.H.  40478  (K);  Crete,  near  Candia,  Hagios-Deka,  December  1913, 
Duffour,  C.  1501  (P);  Cyclades,  Kea,  30  April  1961,  Gathorne-Hardy,  E.  14 
(E);  Dodecanese,  Tilos,  near  chapel  of  Ayios  Nicolaos  c.  180  m  above 
Livadia,  26  April  1962,  Gathorne-Hardy,  E.  139  (E);  Crete,  near  Sitia,  150- 
200  m,  on  clayey  soil,  30  October  1966,  Greuter,  W.  7821  (E);  Near  Corinth, 
Examillia,  8  February  1931,  Guiol,  F.  1731  (BM);  Crete,  Canea,  on  waste- 
land, 10  March  1883,  Reverchon,  E.  117  (BM,  E,  K,  P,  PAL);  Dodecanese, 
Kos,  by  track  near  Asclepeion,  23  March  1985,  Townsend,  C.C.  85/11  (K); 
Dodecanese,  Rhodes,  9  February  1914,  Vaccari,  A.  1141  (K);  Crete,  Nomos 
Lasithiou,  Monastiraki,  roadside,  5  April  1980,  Walree,  A.M.  22226  (BM). 

CYPRUS.  Kyrenia,  fallow  olive  grove,  13  February  1949,  Casey,  B.C. 
221  (K). 

TURKEY.  Province  Mugla,  Kemer-Kestep,  50  m,  edge  of  fields,  29 
March  1956,  Davis,  P.H.  &  Polunin,  O.  25469  (BM,  E,  K);  Side,  22  February 
1966,  Baytop,  A.  et  al.  8504  (E). 

SYRIA.  Baniyas,  300  m,  foot  of  hill,  beneath  tree,  11  March  1945, 
Norris,  F.H.  s.n.  (BM). 

ISRAEL.  Jerusalem,  fields,  800  m,  20  February  1908,  Meyers,  F.  4377 
(E);  Mount  Carmel,  upper  Nahal  Nesher,  2  km  S.  of  Nesher,  32°45'N, 
35°31'E,  280  m,  terra  rossa  on  hard  limestone,  6  April  1989,  Danin,  A.  & 
Knees,  S.G.  1709  (RNG);  Mount  Gerizim,  near  Nablus,  fields,  700  m,  22 
December  1910,  Meyers,  F.S.  &  Dinsmore,  J.E.  6377  (E);  District  Haifa,  near 
Waldheim  (Alloney  Aba),  open  places  in  Quercus  ithaburensis  forest,  28 
March  1942,  Davis,  P.H.  4166  (E,  K). 

The  confusion  surrounding  specific  epithets  for  the  taxon  we  are 
here  calling  Mandragora  officinarum  is  incredible.  This  morass  has 
arisen  for  two  basic  reasons,  firstly  orthographic  errors,  and  sec- 
ondly, the  confusion  over  the  phenological  status  of  type  specimens. 
Orthographic  errors  made  by  early  authors  persist  through  floristic 
works,  adding  and  compounding  any  ambiguities  that  may  have 
arisen.  A  name  often  used  in  synonymy  (or  as  a  valid  name)  of  M. 
officinarum  is  Mandragora  officinalis  Mill.,  said  to  be  validly 
published  in  1768  (see  Jackson  &  Berry,  1979:  508,  footnote). 
However,  in  the  eighth  edition  of  The  gardener's  dictionary  (Miller, 
1768),  where  Miller  used  Linnaean  epithets  for  the  first  time,  Miller 
actually  used  the  epithet  officinarum  ('MANDRAGORA 
(officinarum)'),  notofficinalis.  In  the  ninth  edition  (Miller  &  Martyn, 


1807),  the  taxon  was  classified  as  Atropa  mandragora,  following 
Systema  naturae  (Linneaus,  1759)  and  the  second  edition  of  Species 
plantarum  (Linnaeus,  1762).  Mandragora  officinalis  was  cited  in 
synonymy,  with  reference  made  to  the  illustrations  produced  to 
compliment  the  earlier  editions  (Miller,  1760).  These  illustrations 
bear  no  specific  epithets,  thus  the  1807  reference  to  the  epithet 
officinalis  is  almost  certainly  an  orthographic  error  for  the  1768  use 
of  Linneaus's  epithet  officinarum,  which  was  not  cited  in  synonymy 
in  the  ninth  edition  (Miller  &  Martyn,  1807).  Thus  the  name 
Mandragora  officinalis  Mill,  has  persisted  in  the  literature,  but  has 
no  real  nomenclatural  standing. 

Similarly,  Bertoloni  (1820)  attributed  the  epithet  officinalis  (see 
below)  to  Willdenow,  explicitly  citing  'W.  En.\  Willdenow  (1809), 
in  his  Enumeratio  plantarum,  attributed  the  epithet  officinalis  to 
DeCandolle  (in  Lamarck  &  DeCandolle,  1805),  where  it  is  again 
attributed,  in  error,  to  Miller's  Gardener's  dictionary.  Thus  it  is 
apparent  that  the  numerous  occurences  of  the  epithet  officinalis, 
attributed  to  many  authors,  are  traceable  to  a  single  orthographic 
error  from  Miller's  1767  edition  of  the  Gardener's  dictionary.  Even 
if  the  epithet  officinalis  had  nomenclatural  standing,  it  would  be 
invalid  under  Article  52.1  (see  above),  as  it  is  homotypic  with 
Mandragora  officinarum  L.:  in  DeCandolle  (in  Lamarck  &  DeCan- 
dolle, 1805)  and  Willdenow  (1809),  Atropa  mandragora  L.  is  cited 
in  synonymy. 

Assumptions  as  to  the  phenological  status  of  type  specimens  has 
also  led  to  confusion  about  the  correct  application  of  epithets  (e.g. 
Jackson  &  Berry,  1979).  Bertoloni  (1820,  1824,  1835)  contributed 
greatly  to  the  confusion  surrounding  names  for  the  Mediterranean 
mandrake.  In  a  list  of  plants  cultivated  in  the  Botanic  Gardens  at 
Bologna  (Bertoloni,  1820),  he  included  two  species  of  mandrakes: 
Mandragora  autumnalis,  which  he  coined  as  a  new  name,  and  M. 
officinalis,  which  he  attributed  to  Willdenow.  In  a  later  work  (Berto- 
loni, 1824)  he  designated  two  different  taxa:  M.  vernalis  and  M. 
officinarum,  explicitly  citing  Linnaeus  as  the  source  of  the  latter 
epithet.  He  put  M.  officinalis  in  synonymy  with  M.  vernalis  and  put 
M.  autumnalis  in  synonymy  with  M.  officinarum.  In  Commentarius 
de  Mandragoris  (Bertoloni,  1835),  he  revised  the  genus  and  recog- 
nized three  species:  M.  vernalis,  M.  officinarum  andM.  microcarpa. 
In  synonymy  he  left  out  both  of  the  previously  used  epithets, 
autumnalis  and  officinalis,  which  he  may  have  considered  to  be 
superfluous. 

The  lectotype  of  Mandragora  officinarum,  selected  from  amongst 
the  elements  used  by  Linneaus  in  his  description,  bears  no  pheno- 
logical information  at  all.  The  specimen  in  Burser's  herbarium  is 
labelled  'Misnia,  Lusatia',  indicating  it  came  from  Germany.  It  is 
probable  therefore  that  it  is  from  a  botanical  garden,  and  since  plants 
of  Mandragora  in  northern  European  gardens  usually  bloom  in 
spring,  it  has  been  assumed  that  this  one  did  as  well.  This  is  not 
founded  in  fact,  thus  rendering  much  of  the  debate  as  to  which  name 
is  the  vernal  or  autumnal  taxon  irrelevant. 

2.  Mandragora  turcomanica  Mizg.  in  Trudy  Turkmen.  Fil.  Akad. 
Nauk  SSSR.Ashkabad2:  165  (1942).  Type:  Turkmenistan,  west- 
ern Kopet  Dag,  Kara  Kala  region,  southern  foot  of  Mt  Syunt, 
Schevlan  valley,  on  scree  slopes,  26  November  1938,  O.G. 
Mizgireva,  M.  Nastacalicz  &  G.  Nastacalicz  s.n.  (ASH-holotype; 
LE-isotype(?)). 

ILLUSTRATIONS.    Mizgireva  (1942:  figs  1, 2;  1955:  figs  2-7, 9,  14, 
15,  18,  19,  24);  Belyanina  (1982:  figs  1,  2). 

Plant  virtually  stemless.  Leaves  arranged  in  a  rosette:  lower  leaves  c. 
90  x  60  cm,  length/width  ratio  c.  1 .3-1 .5,  broadly  elliptical  or  ovate, 


34 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


usually  with  large,  irregular  teeth  in  upper  half,  upper  leaves  smaller, 
oblong-ovate  or  broadly  lanceolate,  usually  without  large  teeth,  on 
both  sides  sparsely  hairy,  the  hairs  mostly  arranged  along  veins. 
Flowers  borne  l(-3)  in  leaf  axils,  occasionally  the  pedicels  up  to  2- 
6-flowered.  Pedicels  2-3  cm  long,  7-18  cm  in  fruit.  Calyx  15-20 
mm  long,  lobes  10-15  mm,  acuminate,  accrescent  in  fruit.  Corolla 
20-25  mm  long,  campanulate,  violet  or  purple,  with  three  narrow 
white  stripes  at  base,  the  lobes  c.  15  mm.  Stamens  c.  10  mm  long, 
attached  to  lower  half  of  corolla;  filaments  6-7  mm,  densely  pubes- 
cent at  the  base  and  in  the  lower  Vr,  anthers  c.  4  mm  long,  pale  blue. 
Style  exceeding  stamens;  stigma  capitate,  slightly  bilobed,  green. 
Fruit  40-60  mm  in  diameter,  glossy  yellow  to  orange  when  ripe, 
pulpy,  juicy,  smelling  of  melons,  edible.  Seeds  4-5  x  6-7  mm, 
reniform,  yellow  to  light  brown. 

HABITAT.  Shrubland  in  ravines  and  on  stony  mountain  slopes, 
especially  among  Paliurus  spina-christi  Mill.  (Rhamnaceae). 
Altitudinal  range  500-700  m. 

VERNACULAR  NAME.    Turkmenian  mandrake. 

PHENOLOGY.  Flowering  October  to  March.  Fruiting  until  June. 
Summer  dormancy  until  autumn  rains  begin.  Xerothermic 
ephemeroid. 

CONSERVATION  STATUS.  Only  small  populations  in  a  restricted 
area  within  the  Syunt-Khassardagh  Reserve,  and  in  the  valleys  of 
Chokhadgh,  Shevlan,  Altybai,  Ekechinar,  Sarymsakli  and  Dagdanli 
are  known  (Kurbanov,  1994).  According  to  Mizgireva  (1978)  the 
population  in  the  Syunt-Khassardagh  reserve  consists  of  fewer  than 
a  thousand  plants. 

DISTRIBUTION.     Southwestern  Kopet  Dag.  (Turkmenistan,  ?Iran). 

SELECTED  SPECIMENS  EXAMINED.  No  specimens  were  available 
for  examination:  the  data  here  are  drawn  exclusively  from  the 
literature. 

In  the  original  article  in  which  this  species  was  described  (Mizgireva, 
1942)  the  type  is  cited  as  being  in  Ashkabad  (ASH).  Subsequently, 
in  the  Flora  of  the  USSR  (Lincevskij,  1955),  the  type  was  said  to  be 
in  Leningrad  (now  St.  Petersburg,  LE)  and  this  citation  has  been 
followed  in  other  floristic  works  (Schonbeck-Temesy,  1972).  Since 
we  have  been  unable  to  obtain  specimens  from  either  herbarium,  it 
is  uncertain  whether  or  not  the  holotype  remains  in  ASH  or  has  been 
transfered  to  LE,  thus  we  have  cited  its  location  as  in  the  original 
publication. 

Mizgireva's  (1955)  detailed  study  of  this  species  documented  the 
flowering  times  (Mizgireva,  1955:  fig.  16)  of  individual  plants  and 
clearly  shows  that  some  individuals  flower  in  autumn  (October  and 
November)  while  others  flower  in  spring  (February  and  early  March). 
This  is  the  same  situation  as  that  in  Mandragora  officinarum,  and  is 
markedly  different  from  that  found  in  much  of  the  rest  of  the  flora  of 
Kopet  Dag  (Mizgireva,  1978).  Mizgireva  (1955)  stated  that  the 
differences  between  her  species,  M.  turcomanica,  and  the  Mediter- 
ranean mandrakes  (separated  by  her  into  M.  officinarum  and  M. 
autumnalis)  were  largely  size  of  plant,  colour  of  anthers  and  relative 
sizes  of  calyx  and  corolla.  It  is  clear  from  her  work  that  these 
characteristics  are  consistent  throughout  the  range  of  wild  and 
greenhouse  grown  material  she  examined,  but  since  she  cites  no 
specimens,  nor  have  we  been  able  to  examine  any  of  the  known 
specimens  of  M.  turcomanica,  an  in-depth  analysis  of  these  charac- 
ters relative  to  the  range  of  variation  found  in  M.  officinarum  awaits 
future  study. 


3.  Mandragora  caulescens  C.B.  Clarke,  Fl.  Brit.  Ind.  4:  242 
(1883).  Type:  India,  Sikkim,  Lachen,  Hooker  s.n.  (Kl-lectotype, 
designated  here  by  R.  Mill). 

Fig.  9. 

=  Anisodus  mariae  Pascher  in  Feddes  Repert.  Spec.  Nov.  7:  227 

(1909). 

=  Mairella  yunnanensis  H.  Lev.,  Cat.  pi.  Yun-Nan:  199  (1916). 
=  Anisodus  caulescens  (C.B.  Clarke)  Diels  in  Feddes  Repert.  Spec. 

Nov.Beih.  12:480(1922). 
=  Mandragora  tibetica  Grubov  in  Rast.  Tsentral.  Azii.  5:  108-109 

(1970). 
=  Mandragora  caulescens  subsp.  brevicalyx  Grierson  &  D.G.  Long 

in  Notes  Roy.  Bot.  Card.  Edinburgh  36:  144  (1978). 
=  Mandragora  caulescens  subsp.  flavida  Grierson  &  D.G.  Long  in 

Notes  Roy.  Bot.  Card.  Edinburgh  36:  143  (1978). 
=  Mandragora  caulescent  subsp.  purpurascens  Grierson  &  D.G. 

Long  in  Notes  Roy.  Bot.  Card.  Edinburgh  36:  143  (1978). 
=  Mandragora  chinghaiensis  Kuang  &  A.M.  Lu,  Fl.  reip.  pop.  Sin. 

67:  159  (1978). 

ILLUSTRATIONS.  Grubov  (1970:  pi.  IV,  fig.  2;  pi.  V,  fig.  2);  Kuang 
&  Lu  (1978:  fig.  36,  1-6).  Hoare  &  Knapp  (1997:  fig.  5d). 

Root  stout,  often  branched.  Stem  present,  max.  60  cm  long.  Leaves 
arranged  in  a  cluster  on  top  of  the  young  stem  above  scale  leaves, 
later  along  the  secondary  growth  of  the  stem,  max.  30  cm  long, 
length/width  ratio  1.9-5.5,  oblanceolate  or  spathulate,  glabrous  to 
densely  pubescent  beneath,  especially  along  the  veins.  Flowers 
borne  in  leaf  axils,  often  nodding,  pleasantly  scented.  Pedicels  5-10 
cm  long.  Calyx  5-30  mm  long,  often  nearly  as  long  as  corolla,  lobed 
c.  l/4  of  the  way  to  the  base,  accrescent  in  fruit.  Corolla  5-30  mm 
long,  campanulate  to  cup-shaped,  yellow  to  purple,  the  venation 
usually  darker,  lobed  about  1A  of  the  way  to  the  base,  the  lobes  2-10 
mm.  Stamens  adnate  in  lower  half  of  corolla;  filaments  3-10  mm; 
anthers  1-5  mm  long,  white  to  yellow.  Style  3-6  mm  long,  not 
exceeding  the  stamens;  stigma  capitate,  somewhat  bilobed.  Ovary 
2.5-5  mm  in  diameter,  globose.  Fruit  10-25  mm  in  diameter, 
globose,  pale  greenish  white  or  greenish  yellow.  Seeds  1.5-3.0  mm 
long,  numerous,  reniform,  the  surface  pitted,  yellow  to  light  brown. 

HABITAT.  Grassy  and  stony  slopes,  open  moorland,  pastures  and 
screes.  Subalpine  and  alpine  zone,  especially  among  Rhododendron 
shrubs.  Altitudinal  range  3000-4900  m. 

VERNACULAR  NAMES.     Himalayan  mandrake.  China:  qie  shen. 

PHENOLOGY.  Flowering  April  to  September.  Fruiting  August  to 
October. 

CONSERVATION  STATUS.    Locally  common.  Not  threatened. 

DISTRIBUTION.  Sino-Himalayan.  (Nepal,  India,  Bhutan,  Myanmar, 
China). 

SELECTED  SPECIMENS  EXAMINED.    NEPAL.  Topke  Goia,  27°39'N, 

87°35'E,  3810  m,  in  short  grassland,  3  July  1971,  Beer,  LW.  8264  (BM); 
Tarakot,  28°53'N,  82°59'E,  3900  m,  25  May  1969,  Dobremez,  J.F.  175  (BM); 
Jangla  Banyang,  wet  place,  3800  m,  5  June  1973,  Einarsson,  L.  et  al.  430 
(BM);  Ganesh  Himalaya,  Shior  Khola,  4270  m,  bouldery  alpine  meadow, 
Gardner,  P.C.  479  (BM);  Topke  Gola-Tasagon,  3600-4350  m,  17  August 
1977,  Ohashi,  H.  et  al.  775165  (BM);  Langtang  Valley,  28°14'N,  85°32'E, 
4420  m,  among  rocks  in  turf,  June  1949,  Polunin,  O.  63 1  (BM);  5  miles  E.  of 
Timure,  oak  wood,  41 10  m,  3  July  1949,  Polunin,  O.  780  (BM);  pass  NW  of 
Jumla,  6  miles  Pansae  Dara,  3050  m,  growing  in  loose  soil  recently  free  of 
snow,  30  April  1952,  Polunin,  O.  et  al.  878  (BM);  Dozam  Schola,  near 
Simikot,  3510  m,  grass  slopes,  2  June  1952,  Polunin,  O.  et  al.  4250  (BM); 
near  Jumla,  3810  m,  open  grassy  slopes,  14  July  1952,  Polunin,  O.  et  al. 


MANDRAGORA 


35 


X    S 

\U*iin    I 


I 


36 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


4668  (BM);  Arun-Tamur  watershed,  Topke  Gola,  3960  m,  11  May  1956, 
Stainton,  J.D.A.  254  (BM);  Balangra  Pass,  W.  of  Tibrikot,  29°06'N,  82°41'E, 
3810  m,  28  May  1963,  Stainton,  J.D.A.  4263  (BM);  Rolwaling,  27°55'N, 
86°23'E,  3810  m,  beneath  small  shrubs,  25  June  1964,  Stainton,  J.D.A.  4676 
(BM);Khunde-Khumjung,3820m,6Junel952,Z//n/w^r/nann,A.  688  (BM). 

INDIA.  Sikkim:  Changu,  3960  m,  in  crevices  of  rocks,  28  June  1913, 
Cooper,  R.E.  80  (E);  Gnatong,  3960  m,  peaty  marsh,  31  August  1913, 
Cooper,  R.E.  759  (E);  Jongri,  4000-4200  m,  22  May  1960,  Hara,  H.  et  al. 
443  (BM,  K);  Tungu,  3660-3960  m,  23  July  1849,  Hooker,  J.D.  s.n.  (K); 
Jongri,  4420  m,  campsite  towards  Yakhut  amongst  Juniperus  scrubs,  29 
May  1990,  Kirkpatrick,  G.  53  (E);  District  Darjeeling,  Phalut,  3350-3660 
m,  30  May  1902,  Lace,  J.H.  2266  (E);  West  District,  Bikbari,  Choktsering 
Chhu  valley;  27°30'53"N,  88°08'28"E,  3950  m,  amongst  scrub,  12  July 
1992,  Long,  D.G.  et  al.  266  (E);  Chulong,  4570  m,  12  September  1912, 
Rhomoo,  L  210  (E);  Gamothang,  3960  m,  12  August  1913,  Rhomoo,  L. 
1059  (E);  Giagong,  3960  m,  6  September  191 1,  Ribu  &  Rhomoo  5492  (BM, 
E,  K);  Phalut,  3350-3660  m,  27  May  1911,  Smith,  W.W.  4593  (E);  Jongri, 
4000  m,  in  shelter  of  dwarf  Rhododendron  and  Juniperus,  23  June  1983, 
Starling,  B.N.  et  al.  92  (K);  Tangshing  campsite,  4000  m,  open  grazed  areas 
among  Rhododendron  and  around  campsite,  24  June  1983,  Starling,  B.N.  et 
al.  106  (K);  summit  of  Sandakphu,  3660  m,  small  dense  tufts  amongst 
grass,  18  May  1881,  Watt,  G.  5373  (E,  K).  Arunachal  Pradesh  (Assam): 
Orka  La,  Bhutan  frontier,  3960  m,  on  stony  slopes,  scattered,  7  June  1938, 
Kingdon  Ward,  E  13840  (BM). 

BHUTAN.  Paco  Chu,  4110m,  loose  sandy  crevice,  15  May  1966,  Bowes 
Lyon,  S.  3 156  (BM);  Phajudin  Timpu,  3960  m,  5  August  1914,  Cooper,  R.E. 
3232  (BM,  E);  Tare-La  area,  3660  m,  19  May  1938,  Gould,  B.J.  109  (K); 
Laum  Thang,  3900  m,  in  alpine  herbage  with  moderate  humidity,  18  May 
1967,  Hara,  H.  et  al.  1 1973  (BM);  Western  Bhutan,  3510  m,  among  boulders 
above  tree  line,  17  June  1933,  Ludlow,  F.  &  Sheriff,  G.  107  (BM);  Byasu  La, 
4110m,  growing  among  stones  on  open  hillside,  18  May  1937,  Ludlow,  F.  & 
Sherriff,  G.  3073  (BM);  Cheli  La,  3810  m  in  open  moorland,  4  May  1949, 
Ludlow,  F.  et  al.  16139  (BM);  Pangotang,  3960  m,  open  hillside  above  fir 
zone,  27  May  1949,  Ludlow,  Eel  al.  18986(BM);  Kantanang,Tsampa,4110 
m,  among  shrubs  on  steep  hillside,  3  June  1949,  Ludlow,  Eetal.  19040(BM); 
Shingbe,  Me  La,  3350  m,  growing  amongst  other  herbs  and  small  shrubs,  16 
May  1949,  Ludlow,  F.  et  al.  20645  (BM,  E). 

MYANMAR  (Burma).  Chenochi  Pass,  3600  m,  3  July  1920,  Farrer,  J.R. 
1694  (E). 

CHINA.  Yunnan:  Diqing  Prefecture,  E.  slope  of  Bai  Ma  Shan,  28°19'N, 
99°05'E,  3895  m,  among  stones  in  Abies  forest,  9  June  1993,  Alden,  B.  et  al. 
1021  (E);  Diqing  Prefecture,  Zhongdian  County,  above  Napa  Hai,  N.  of 
Zhongdian,  27°55'N,  99°34'E,  4000  m,  forested  ridge,  snow-bed  vegetation 
and  dwarf  Rhododendron  scrub,  12  June  1993,  Alden,  B.  et  al.  1185  (E); 
western  flank  of  the  Lichiang  range,  27°20'N,  3050-3350  m,  open  alpine 
pasture,  June  1910,  Forrest,  G.  5999  (E,  K);  Lichiang  range,  27°25'N, 
3660-3960  m,  July  1913,  Forrest,  G.  10474  (BM,  E,  K);  western  slopes  of 
Likiang  snow  range,  Yangtze  watershed,  30  May-6  June  1922,  Rock,  J.F. 
4211  (BM);  Mo-ting  shan,  ME  of  Atuntze,  eastern  slopes  of  Yangtze- 
Mekong  divide,  3660  m,  along  trail,  June  1932,  Rock,  J.F.  22771  (E);  slopes 
of  Mt  Gyi-na  lo-ko,  the  second  peak  of  the  Yu-lung  shan,  3660  m,  in  alpine 
meadows,  April-May  1932,  Rock,  J.F.  25005  (BM,  E);  Chungtien,  Chiren, 
2200  m,  grass  slope,  17  May  1937,  Yu,  T.T.  11327  (KUN);  Mekong- 
Salween  divide,  Chingpoh,  3800  m,  alpine  pasture  lands,  14  June  1938,  Yu, 
T.T.  19078  (E);  Upper  Kiukiang  valley,  Clulung  Lungtsahmuru,  3700  m, 
mountain  slope,  open  and  grassy  place,  9  August  1938,  Yu,  T.T.  19843  (E, 
KUN).  Sichuan:  Tien-chuan-hsien,  3500  m,  14  June  1936,  Chu,  K.L.  2791 
(BM);  mountains  E.  of  Yungning,  27°48'N,  101°  E,  3660  m,  open  stony 
pasture,  June  1922,  Forrest,  G.  21407  (BM,  E,  K);  mountains  around  Muli, 
28°12'N,  101°  E,  3660  m,  July  1930,  Forrest,  G.  28416  (BM,  E);  Liuku- 
liangdse,  27°48'N,  3700-4200  m,  18  May  1914,  Handel-Mazetti,  H.  von 
2349  (K);  Minya  Konka  snow  range,  S.  of  Tatsienlu,  4450  m,  in  meadows, 
July  1929,  Rock,  J.F.  17530  (E);  Mu-li-ka-bu,  3600  m,  mountain  slope, 
grassy  place,  18  May  1937,  Yu,  T.T.  5496  (KUN).  Qinghai:  Dari  (Darlag) 
Xian,  just  N.  of  Manzhang,  along  the  Manzhang  He,  between  Dari  and 
Banma,  33°17'51"N,  100°25'55"E,  4000  m,  on  moist  rodent  disturbed 
slope,  12  August  1993,  Ho,  T.N.  et  al.  1 172  (E);  Chindu  Xian,  Xiwu  Xiang, 
E.  of  pass  on  road  between  Xiwu  and  the  Szechwan  border,  33°09'14"N, 
97°32'35"E,  4250  m.  damp  slope  with  extensive  pika  warrens,  1 1  Septem- 
ber 1996,  Ho,  T.N.  et  al.  3063  (BM).  Xizang:  Dotha,  3960  m,  20  June  1945, 


Bar  &  Kirat  Ram  20511  (K);  Mt.  Demula,  Yiwei  County,  4100  m,  23 
August  1973,  Chinese  Tibet  Expedition  1973  1214  (KUN);  District 
Chengtang,  Dingjie  County,  Ladang  to  Xingeng,  3800-4000  m,  9  June 
1975,  Chinese  Tibet  Expedition  1975  5679  (KUN);  Londre  Pass,  Tsarong, 
Mekong-Salween  divide,  28°14'N,  98°40'E,  3960  m,  July  1921,  Forrest,  G. 
19630  (UC/JEPS,  P,  K);  Tha  Chu  valley,  3660  m,  in  clumps,  alpine  region, 
in  stony  pastures  and  on  grassy  slopes,  11  July  1950,  Kingdon  Ward,  F. 
19613  (BM);  N.  of  Sanga  Choling,  28°41'N,  93°02'E,  4110  m,  on  open 
grassy  bank,  14  May  1936,  Ludlow,  E  &  Sherriff,  G.  1590  (BM);  hills  S.  of 
Lhasa,  4420  m,  ground  left  bare  by  melting  snow,  6  June  1942,  Ludlow,  F. 
&  Sherriff,  G.  8676  (BM);  hills  S.  of  Lhasa,  4720  m,  herb  growing  under 
boulders,  21  August  1942,  Ludlow,  F.  &  Sherriff,  G.  9014  (BM);  Pome, 
above  Showa  Dzong,  29°55'N,  95°25'E,  3350  m,  open  hillsides  midst  grass, 
1 1  June  1947,  Ludlow,  F.  et  al.  13149  (BM,  E);  Sobhe  La,  near  Tongyuk 
Dzong,  Pome,  30°07'N,  94°54'E,  3510  m,  21  May  1947,  Ludlow,  F.  et  al. 
13750  (BM,  E);  Province  Kongbo,  Nyima  La,  29°38'N,  94°52'E,  3350  m, 
on  dry  ground  under  trees,  21  May  1947,  Ludlow,  F.  et  al.  15051  (BM,  E); 
Lo  La,  Pachakshiri,  28°49'N,  93°59'E,  3810  m,  in  openings  of  Rhododen- 
dron forest,  mostly  above  Abies  zone,  15  May  1938,  Ludlow,  F.  et  al.  3776 
(BM);  Province  Kongbo,  south  side  of  Lusha  Chu,  29°20'N,  94°35'E,  3810 
m,  in  Lonicera  and  Berberis  scrub,  13  June  1938,  Ludlow,  F.  et  al.  4591  A 
(BM,  E);  valley  above  Tse,  Tsangpo  valley,  29°23'N,  94°22'E,  3350  m,  in 
grassy  banks  under  Rhododendron,  Ludlow,  F.  et  al.  4591  (BM). 

The  status  and  identity  of  the  previously  proposed  subspecies  of 
Mandragora  caulescens  (Grierson  &  Long,  1978)  are  discussed  in 
the  section  on  the  species  delimitation  (p.  25).  It  is  clear  from  our 
studies  that  M.  caulescens  is  extremely  variable  and  that  a  detailed 
field-based  study  over  the  entire  species  range  is  necessary. 


ACKNOWLEDGEMENTS.  This  study  was  undertaken  as  partial  fulfilment  of 
the  requirements  for  an  M.Sc.  in  Advanced  Methods  in  Taxonomy  and 
Biodiversity  at  Imperial  College,  London  and  The  Natural  History  Museum, 
London  undertaken  by  S.  Ungricht  and  supervised  by  S.  Knapp  and  J.R. 
Press.  The  work  has  profited  from  discussions  with  many  members  of  the 
staff  at  The  Natural  History  Museum,  particularly  S.  Cafferty,  N.C.  Garwood, 
CJ.  Humphries,  C.E.  Jarvis,  C.  Moncrieff,  N.K.B.  Robson,  K.  Sidwell,  A.R. 
Vickery,  D.M.Williams,  and  C.Whitefoord.  We  are  grateful  to  Z.  Frankiel,  K. 
Rottcher  and  Zhan  Ren-bin  for  their  translation  of  the  crucial  Russian  and 
Chinese  papers  respectively.  Furthermore,  we  are  indebted  to  M.  Gilbert  and 
R.R.  Mill  for  their  sharing  of  personal  observations  and  to  N.  Turland  for  a 
thoughtful  and  detailed  review.  Thanks  are  also  due  to  R.  Baldini  in  Florence 
and  the  directors  and  staff  of  the  following  herbaria  who  made  their  collec- 
tions available:  E,  JEPS,  K,  KUN,  P,  PAL,  RNG  and  UC.  This  study  has  been 
supported  by  a  grant  from  the  Park  Fund  of  the  Botany  Department  of  The 
Natural  History  Museum,  London. 


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EXSICCATAE 

Arranged  by  collector,  with  collection  numbers  (s.n.:  collections 
without  collection  number),  species  and  herbaria. 

Al-Eisawi,  D.  &  Al-Jallad,  W.  2221,  M.  officinarum  (BM) 

Alden,  B.,  Alexander,  J.C.M.,  Long,  D.G.,  McBeath,  R.J.D.,  Noltie,  H.J.  & 

Watson,  M.E  1021,  M.  caulescens  (E);  1185,  M.  caulescens  (E) 
Atchley,  S.C.  s.n.,  M.  officinarum  (BM);  315,  M.  officinarum  (K);  819,  M. 

officinarum  (K) 


Atherton,  G.E.  793,  M.  officinarum  (K);  812,  M.  officinarum  (K);  998  M. 

officinarum  (K) 

Baker,  E.W.  149,  M.  officinarum  (BM) 
Ball,  J.  s.n.,  M.  officinarum  (E) 
Barclay,  C.  3004,  M.  officinarum  (K) 
Bauer,  K.  &  Spitzenberger,  F.  53,  M.  officinarum  (BM) 
Baytop,  A.,  Tanker,  N.  &  Sezik,  E.  8504,  M.  officinarum  (E) 
Beer,  L.W.  8264,  M.  caulescens  (BM) 
Bianor  &  Sabasien  1877  ,  M.  officinarum  (BM,  E,  P,  RNG) 
Bickerich,  G.  15324,  M.  officinarum  (BM,  K) 
Biro,  L.  s.n.,  M.  officinarum  (K) 

Bar  &  Kirat  Ram  20481,  M.  caulescens  (K);  2051 1,  M.  caulescens  (K) 
Bornmiiller,  J.  s.n.,  M.  officinarum  (K) 

Bourgeau,  E.  s.n.,  M.  officinarum  (K,  P),  2304,  M.  officinarum  (E,  K,  P) 
Bowen,  H.J.M.  694,  M.  officinarum  (RNG);  3088,  M.  officinarum  (RNG) 
Bowes  Lyon,  S.  3156,  M.  caulescens  (BM);  3239,  M.  caulescens  (BM) 
Brickell,  C.  &  Mathew,  B.  8088,  M.  officinarum  (K) 
Carr,  J.W.  s.n.,  M.  officinarum  (RNG) 
Casey,  E.  C.  22 1 ,  M.  officinarum  (K) 
Chaboisseau,  C.  575,  M.  officinarum  (BM) 
Chatterjee,  D.  75,  M.  caulescens  (BM) 
Chinese  Tibet  Expedition  1973  1214,  M.  caulescens  (KUN) 
Chinese  Tibet  Expedition  1975  5679,  M.  caulescens  (KUN) 
Chu,  K.L.  2791,  M.  caulescens  (BM) 
Clarke,  C.B.  34977,  M.  caulescens  (K);  34983  A,  M.  caulescens  (BM); 

35704,  M.  caulescens  (K) 
Clarke,  E.D.  s.n.,  M.  officinarum  (BM) 
Coincy,  M.A.  de  s.n.,  M.  officinarum  (P) 
Cooper,  R.E.  80,  M.  caulescens  (E);  759,  M.  caulescens  (E);  3232,  M. 

caulescens  (BM,  E) 

Danin,  A.  &  Knees,  S.G.  1709,  M.  officinarum  (RNG) 
Davis,  P.H.  1 100,  M.  officinarum  (E,  K);  1438,  M.  officinarum  (K);  2108, 

M.  officinarum  (E,  K);  4166,  M.  officinarum  (E,  K);  40134,  M. 

officinarum  (E);  40182,  M.  officinarum  (RNG);  40478,  M.  officinarum 

(K) 

Davis,  P.H.  &  Polunin,  O.  25469,  M.  officinarum  (BM,  E,  K) 
Delavay,  J.M.  s.n.,  M.  caulescens  (P);  263,  M.  caulescens  (P);  1026,  M. 

caulescens  (P) 

Dhwoj,  L  513,  M.  caulescens  (BM,  E) 
Dinsmore,  J.E.  B  377,  M.  officinarum  (E) 
Dobremez,  J.F.  175,  M.  caulescens  (BM) 
Duff  our,  C.  1501,  M.  officinarum  (P) 
Economides,  S.  1055,  M.  officinarum  (K) 

Einarsson,  L,  Skarby,  L.  &  Wetterhall,  B.  430,  M.  caulescens  (BM) 
Espinosa,  A.  870252,  M.  officinarum  (K) 

Ex  Herb.  Postian.  apud  Colleg.  Syriens.  Protest.  952,  M.  officinarum  (BM) 
Farrer,  J.R.  1694,  M.  caulescens  (E) 
Fay,  J.M.  815,  M.  officinarum  (K);  864,  M.  officinarum  (K) 
Forbes  s.n.,  M.  officinarum  (UC/JEPS) 
Forrest,  G.  569,  M.  caulescens  (E);  5999,  M.  caulescens  (E,  K);  10474,  M. 

caulescens  (BM,  E,  K);  19630,  M.  caulescens  (UC/JEPS,  P,  K);  21407, 

M.  caulescens  (BM,  E,  K);  26951,  M.  caulescens  (P);  28416,  M. 

caulescens  (BM,  E) 

Galiano,  E.F.  &  Valdes,  B.  820,  M.  officinarum  (E,  RNG) 
Gammie,  G.A.  s.n.,  M.  caulescens  (BM,  E) 
Gandoger,  M.  s.n.,  M.  officinarum  (K);  84,  M.  officinarum  (K);  8400,  M. 

officinarum  (K) 

Gardner,  M.E,  Jury,  S.L.  &  Rejdali,  M.  4885,  M.  officinarum  (E,  RNG) 
Gardner,  M.F.  &  Knees,  S.G.  4769,  M.  officinarum  (RNG) 
Gardner,  PC.  479,  M.  caulescens  (BM) 

Garnett,  C.S.  33/5,  M.  officinarum  (BM);  158/7,  M.  officinarum  (BM) 
Gathorne-Hardy,  E.  14,  M.  officinarum  (E);  139,  M.  officinarum  (E);  194, 

M.  officinarum  (E) 
Girgenti  s.n.,  M.  officinarum  (BM) 
Gould,  B.J.  109,  M.  caulescens  (K) 
Greuter,  W.  7821,  M.  officinarum  (E) 
Guiol,  F.  1731,  M.  officinarum  (BM) 
Haig,  E.  s.n.,  M.  officinarum  (BM) 
Handel-Mazetti,  H.  von  2349,  M.  caulescens  (K) 


MANDRAGORA 


39 


Hara,  H.,  Kanai,  H.,  Murata,  G.  Ohashi,  H.,  Tanaka,  O.  &  Yamazaki,  T. 

1 1973,  M.  caulescent  (BM) 
Hara,  H.,  Kanai,  H.,  Murata,  G.,  Togashi,  M.  &  Tuyama,  T.  443,  M. 

caulescens  (BM,  K) 
Hardy,  E.  s.n.,  M.  officinarum  (BM) 
Hart,  H.C.  s.n.,  M.  officinarum  (BM) 

Hay,  J.H.  1707,  M.  officinarum  (K);  H.  2484,  M.  officinarum  (K) 
Helbaek,  H.  74,  M.  officinarum  (K) 
Heldreich,  T.  von  s.n.,  M.  officinarum  (BM,  E,  K,  P);  257  [1860],  M. 

officinarum  (BM);  257  [1857],  M.  officinarum  (P);  257  [1859],  M. 

officinarum  (K);  2817  [1853],  M.  officinarum  (P);  2817  [1860],  M. 

officinarum  (K,  P) 

Higgins,  E.K.  s.n.,  M.  officinarum  (BM) 

Ho,  T.N.,  Bartholomew,  B.  &  Gilbert,  M.  1 172,  M.  caulescens  (E) 
Ho,  T.N.,  Bartholomew,  B.,  Watson,  M.  &  Gilbert,  M.  3063,  M.  caulescens 

(BM) 

Hooker,  J.D.  s.n.,  M.  caulescens  (E,  K-lectotype,  P) 
Huguenin  s.n.,  M.  officinarum  (BM) 
/wry,  S.L.,  Achhal,  A.  &  Mouks,  H.  &  Upson,  T.M.  12634,  M.  officinarum 

(RNG);  12653,  M.  officinarum  (RNG) 
Jury,  S.L.,  Aitlafkih,  M.,  Hedderson,  T.  &  Kahouadji,  A.  15237,  M. 

officinarum  (RNG) 
Jury,  S.L.,  Aitlafkih,  M.,  Hedderson,  T.  &  Rutherford,  R.  W.  15398,  M. 

officinarum  (RNG);  15434,  M.  officinarum  (RNG);  15701,  M. 

officinarum  (RNG) 

Jury,  S.L.  &  Jury,  L.C.  13111,  M.  officinarum  (BM,  RNG) 
Jury,  S.L,  Rejdali,  M.,  Taleb,  A.  &  Upson,  T.M.  12542,  M.  officinarum 

(RNG) 
Jury,  S.L,  Taleb,  A.  &  Upson,  T.M.  12217,  M.  officinarum  (RNG);  12238, 

M.  officinarum  (RNG) 
Jussieu,  A.  de  s.n.,  M.  officinarum  (P) 
King,  G.  s.n.,  M.  caulescens  (E);  4368,  M.  caulescens  (P) 
Kingdon  Ward,  F.  13840,  M.  caulescens  (BM);  19613,  M.  caulescens  (BM) 
Kirkpatrick,  G.  53,  M.  caulescens  (E) 

Lacaita,  C.C.  s.n.,  M.  officinarum  (BM);  s.n.,  M.  caulescens  (BM) 
Lace,  J.H.  2266,  M.  caulescens  (E) 
Ladero,  M.,  Lopez  Guadalupe,  M.,  Molero,  J.  &  Perez  Raya,  F.  10605,  M. 

officinarum  (RNG) 
Lange,  J.  s.n.,  M.  officinarum  (K,  P) 
Laukkonen,  P.  55,  M.  officinarum  (K) 
Le  Testu,  G.  s.n.,  M.  officinarum  (BM) 
Lewalle,  J.  10054,  M.  officinarum  (BM,  RNG) 
Lojacono-Pojero,  M.  s.n.,  M.  officinarum  (BM) 
Long,  D.G.,  McBeath,  R.J.D.,  Noltie,  H.J.  &  Watson,  M.F.  266,  M. 

caulescens  (E) 

Lowne,  B.T.  s.n.,  M.  officinarum  (BM) 
Ludlow,  F.&  Sheriff,  G.  107,  M.  caulescens  (BM);  1590,  M.  caulescens 

(BM);  3073,  M.  caulescens  (BM);  8676,  M.  caulescens  (BM);  9014,  M. 

caulescens  (BM) 
LHd/ovv,  E,  S/zernjft  G.  cfe  E///O?,  //.//.  13149,  M.  caulescens  (BM,  E); 

13750,  M.  caulescens  (BM,  E);  15051,  M.  caulescens  (BM,  E) 
LwoVow,  E,  Sherriff,  G.  &  Hicks,  J.H.  16139,  M.  caulescens  (BM);  18986, 

M.  caulescens  (BM);  19040,  M.  caulescens,  (BM);  20645,  M. 

caulescens  (BM,  E) 
Ludlow,  F.,  Sherriff,  G.  &  Taylor,  G.  3776,  M.  caulescens  (BM);  4591  A, 

M.  caulescens  (BM,  E);  4591,  M.  caulescens  (BM) 
Maty,  K  s.n.,  M.  officinarum  (K) 
Manberg,  T.  s.n.,  A/,  caulescens  (K) 
Mavzomoustakis,  G.A.  8,  M.  officinarum  (BM) 
Me  Laren  '5  collectors  AC  67,  M.  caulescens  (E,  P) 
Meyers,  E  4377,  M.  officinarum  (E) 
Meyers,  F.S.  &  Dinsmore,  J.E.  6377,  M.  officinarum  (E);  B  6377,  M. 

officinarum  (K) 

M;7/,  y.S.  s.n.,  M.  officinarum  (K) 
Moreau,  W.M.  62,  M.  officinarum  (K) 
Norris,  F.H.  s.n.,  M.  officinarum  (BM) 

,  H.,  ATanai,  H.,  0/z6a,  //.  c£  rafm/z/,  K  775165,  M.  caulescens 


(BM) 

Olivier  &  Bruguiere  s.n.,  M.  officinarum  (P) 
Orphanides,  T.G.  75,  M.  officinarum  (BM,  E,  K,  P) 
Pa/ne,  /A.  s.n.,  M.  officinarum  (K) 

Pantling,  R.  s.n.,  M.  caulescens  (K);  46387,  M.  caulescens  (K) 
Partridge,  F.  2,  M.  officinarum  (BM) 
Pasta/;  7.  <6  Va/des,  B.  s.n.,  M.  officinarum  (RNG) 
Perraudiere,  H.  de  la  s.n.,  M.  officinarum  (P) 
Pette;;  E  s.n.,  M.  officinarum  (BM) 
Pichler  s.n.,  M.  officinarum  (BM,  E,  K) 
Pignant  s.n.,  M.  officinarum  (K) 
P/fard,  C.-y.  1757,  M.  officinarum  (K) 
Polunin,  0.  631,  M.  caulescens  (BM);  780,  M.  caulescens  (BM);  6616,  M. 

officinarum  (K) 
Po/Mrtw,  O.,  Sytes,  W/?.  <fe  Williams,  L.H.J.  878,  M.  caulescens  (BM,  P); 

4250,  M.  caulescens  (BM);  4668,  M.  caulescens  (BM);  4696,  M. 

caulescens  (BM) 

Pratt,  A.E.  755,  M.  caulescens  (BM,  K,  P) 
Probyn,  P.M.  49,  M.  officinarum  (K) 
Raulin,  V.  s.n.,  M.  officinarum  (P);  302,  M.  officinarum  (P) 
Rechinger,  K.H.  90,  M.  officinarum  (BM);  3630,  M.  officinarum  (BM,  K) 
Reiser,  C.  s.n.,  M.  officinarum  (K) 
Reverchon,  E.  s.n.,  M.  officinarum  (BM,  E,  P,  K);  1 17,  M.  officinarum 

(BM,  E,  K,  P,  PAL);  469  [1888],  M.  officinarum  (BM,  P);  469  [1890], 

M.  officinarum  (P) 

Rhomoo,  L  210,  M.  caulescens  (E);  1059,  M.  caulescens  (E) 
/?/£«  <fe  Rhomoo  5492,  M.  caulescens  (BM,  E,  K) 
flocfc,  y.E  3597,  M.  caulescens  (E);  3985,  M.  caulescens  (P);  421 1,  M. 

caulescens  (BM);  17530,  M.  caulescens  (E);  22771,  M.  caulescens  (E); 

24750,  M.  caulescens  (BM,  E);  25005,  M.  caulescens  (BM,  E) 
s,  //.  s.n.,  M.  officinarum  (BM) 
s,  H.  266,  M.  officinarum  (E,  K,  P) 
Schneider,  C.  3501,  M.  caulescens  (E,  K) 

Sintenis  &  Rigo  8,  M.  officinarum  (K,  PAL);  8a,  M.  officinarum  (K) 
Smtt/z,  WW  4593,  M.  caulescens  (E) 
S0H/j<?',  y.-A.  849,  M.  caulescens  (P);  1252,  M.  caulescens  (P);  2102,  M. 

caulescens  (P) 
Stainton,  J.D.A.  254,  M.  caulescens  (BM);  4263,  M.  caulescens  (BM); 

4676,  M.  caulescens  (BM) 
Starling,  B.N.,  Upward,  EM.,  Brickell,  C.D.  &  Mathew,  B.  92,  M. 

caulescens  (K);  106,  M.  caulescens  (K) 
Syngrassides,  A.  1464,  M.  officinarum  (K) 
,  A.  38,  M.  officinarum  (RNG) 
,  A.  1630,  M.  officinarum  (BM) 
Thomas,  P.  s.n.,  M.  officinarum  (K) 
TOman,  //.  W  23,  M.  caulescens  (K) 
Todaro,  A.  s.n.,  M.  officinarum  (P) 
Tbdd,  E.  s.n.,  M.  officinarum  (BM) 
Tomkinson,  M.J.  72,  72A,  M.  officinarum  (BM) 
Townsend,  C.C.  85/1 1,  M.  officinarum  (K) 
Tbze/;  //.E  s.n.,  M.  officinarum  (K) 

Treffewy,  A.  W  1 1,  M.  officinarum  (K);  20,  M.  officinarum  (K) 
Vaccarz,  A.  1 141,  M.  officinarum  (K);  2326,  M.  officinarum  (BM,  K) 
Virantonio  s.n.,  M.  officinarum  (BM) 
Wa/ree,  A.M.  22226,  M.  officinarum  (BM) 
WaM,  £.113,  M.  caulescens  (K) 

Watt,  G.  5373,  M.  caulescens  (E,  K);  5436,  M.  caulescens  (E,  K) 
Wfe/wttscfc,  EM.y.  s.n.,  M.  officinarum  (P);  159,  M.  officinarum  (BM);  337, 

M.  officinarum  (P) 

White,  M.F.  s.n.,  M.  officinarum  (BM) 
Williams,  LH.J.  700,  M.  caulescens  (BM) 
W/son,  E.//.  4192,  M.  caulescens  (BM,  K,  P);  4194,  M.  caulescens  (BM, 

K,  P);  4195,  M.  caulescens  (K) 
WW/ey-ZW,  A.//.  22,  M.  officinarum  (BM) 
Ku,  rr  5496,  M.  caulescens  (KUN);  1 1327,  M.  caulescens  (KUN);  19078, 

M.  caulescens  (E);  19843,  M.  caulescens  (E,  KUN) 
Zimmermann,  A.  688,  M.  caulescens  (BM) 


40 


S.  UNGRICHT,  S.  KNAPP  AND  J.R.  PRESS 


TAXONOMIC  INDEX 


Accepted  names  are  given  in  roman,  synonyms  in  italics.  An  asterisk  denotes 
a  figure. 

Anisodus  caulescens  (C.B.  Clarke)  Diels  34 

A.  luridus  Link  22 

A.  mariae  Pascher  34 

Atropa  acaulis  Stokes  30 

A.  belladonna  L.  30 

A.  humilis  Salisb.  30 

A.  mandragora  L.  30 

Lycium  chinense  Mill.  22 

Mairella  yunnanensis  H.  LeY  34 

Mandragora  acaulis  Gaertn.  30 

M.  autumnalis  Bertol.  30 

M.  caulescens  C.B.  Clarke  34,  35* 

M.  caulescens  subsp.  brevicalyx  Grierson  &  D.G.  Long  34 


M.  caulescens  C.B.  Clarke  subsp.  caulescens  34 

M.  caulescens  subsp.  flavida  Grierson  &  D.G.  Long  34 

M.  caulescens  subsp.  purpurascens  Grierson  &  D.G.  Long  34 

M.  chinghaiensis  Kuang  &  A.M.  Lu  34 

M.  foemina  Garsault  30 

M.  haussknechtii  Heldr.  30 

M.  hispanica  F.  Vierh.  30 

M.  x  hybrida  Hausskn.  &  Heldr.  30 

M.  mas  Garsault  30 

M.  microcarpa  Bertol.  30 

M.  neglecta  G.  Don  ex  Loudon  30 

M.  officinarum  L.  30,  31*.  32* 

M.  praecox  Sweet  30 

M.  shebbearei  C.E.C.  Fisch.    19 

M.  tibetica  Grubov  34 

M.  turcomanica  Mizg.  33 

M.  vemalis  Bertol.  30 

Nicotiana  glauca  Graham  22 

Przewalskia  Grubov   19 


Bull.  nat.  Hist.  Mus.  Land.  (Bot.)  28(1):  41-66  Issued  25  June  1998 

The  pteridophytes  of  Sao  Tome  and  Principe 
(Gulf  of  Guinea) 

ESTRELA  FIGUEIREDO^  ,_ 

Centra  de  Botdnica,  Institute  de  Investigacao  Cientifica  Tropical,  R.  da  Junqueira  86,  1300  Lisboa,  Portugal 

CONTENTS 

Introduction 41 

Diversity  42 

Conservation 42 

Collections 42 

Historical  collections 42 

Recent  collections 42 

Catalogue 43 

Acrostichaceae 43 

Adiantaceae 43 

Aspidiaceae 45 

Aspleniaceae 48 

Athyriaceae 50 

Blechnaceae 51 

Cyatheaceae 51 

Davalliaceae 51 

Dennstaedtiaceae 51 

Elaphoglossaceae 52 

Gleicheniaceae 53 

Grammitaceae 53 

Hemionitidaceae 54 

Hymenophyllaceae 54 

Lomariopsidaceae 56 

Loxogrammaceae  56 

Lycopodiaceae 5" 

Marattiaceae  57 

Nephrolepidaceae 5£ 

Oleandraceae 58 

Ophioglossaceae 55 

Polypodiaceae 5> 

Psilotaceae "" 

Pteridaceae 60 

Selaginellaceae 61 

sr\ 

Sinopteridaceae DZ 

f\) 
Thelypteridaceae 

Vittariaceae "- 

Doubtful  records 


References 


64 


Systematic  index 

SYNOPSIS.  A  catalogue  of  the  pteridophytes  of  Sao  Tome  and  Principe  is  presented,  with  156  taxa,  12  of  which  are  endemic  to 
the  islands.  Recent  synonymy  for  the  area  is  included.  All  collections  examined  are  cited. 


-  '  ---  The  pteridophytes  of  the  neighbouring  island  of  Bioko  (Fernando 

INTRODUCTION  _  _  _        Po)  have  been  recently  revised  by  Benl  (1978,  1980,  1982,  1988, 

1  99  1  )  who  provided  descriptions  and  keys  to  the  taxa.  The  lack  of  an 

The  most  recent  account  of  the  pteridophytes  of  Sao  Tome  and  updated  checklist  for  Sao  Tome  and  Principe  and  the  difficulty  in 
Principe  (Gulf  of  Guinea)  was  produced  by  Alston  in  1944.  Since  compiling  the  dispersed  information  on  this  group  of  plants  have 
then  several  nomenclatural  changes  have  taken  place  and  new  precluded  its  inclusion  in  recent  floristic  and  biodiversity  studies 
records  have  been  added  to  the  flora  of  the  islands  (e.g.  Alston  1958,  (Figueiredo,  1994). 

The  incorporation  of  several  recent  collections  from  Sao  Tome 


©  The  Natural  History  Museum,  1998 


42 


E.  FIGUEIREDO 


and  Principe  at  LISC,  has  revealed  further  new  records  and  new 
areas  of  distribution  for  some  taxa,  and  provided  the  basis  for  a 
revision  of  Alston's  (1944)  account. 


DIVERSITY 

There  are  153  indigenous  taxa  of  pteridophytes  in  Sao  Tome  and 
Principe  occurring  from  sea  level  (e.g.  Acrostichum  aureum  L., 
found  in  swamps  and  mangroves)  to  the  highest  peaks  of  the  islands 
(e.g.  Grammitis  spp.),  and  belonging  to  28  families  and  60  genera. 
Three  taxa  were  probably  introduced,  and  became  naturalized, 
Pityrogramma  calomelanos  (L.)  Link  (Alston,  1944),  Adiantum 
raddianum  C.  Presl  from  the  Neotropics  (Alston,  1956a),  and  Pteris 
vittata  L.  (Alston  1958,  1959). 

One  hundred  and  thirty-seven  taxa  occur  in  Sao  Tome  and  75 
occur  in  Principe;  12  taxa  are  endemic  to  the  islands.  In  Bioko,  the 
pteridophyte  richness  is  higher  than  in  either  Sao  Tome  or  Principe, 
with  208  taxa  in  31  families  (Benl,  1978).  However,  only  four  taxa 
(1.9%)  are  endemic  to  Bioko,  while  eight  taxa  are  endemic  to  Sao 
Tome  (5.8%)  and  three  are  endemic  to  Principe  (4%). 

This  difference  in  the  percentage  of  endemics  is  less  striking  than 
the  difference  found  for  the  angiosperms.  In  that  group,  Bioko  has 
3.6%  endemic  taxa,  Sao  Tome  has  13.4%,  and  Principe  has  8.2% 
(Figueiredo,  1994). 

Asplenium  is  the  genus  of  pteridophytes  with  the  largest  number 
of  taxa  in  Sao  Tome  and  Principe.  Twenty-four  taxa  are  represented, 
of  which  three  are  endemic. 


CONSERVATION 

Twenty-five  taxa  collected  in  Sao  Tome  and  Principe  in  the  nine- 
teenth century  have  not  been  collected  since,  even  though  extensive 
collecting  took  place  in  the  1950s  and  also  in  the  1990s.  Although 
some  of  these  taxa,  represented  in  the  herbarium  by  a  single  collec- 
tion, may  be  doubtful  records,  others  were  collected  more  than  once 
by  different  collectors.  The  absence  of  these  taxa  amongst  the 
collections  made  this  century  may  indicate  their  rarity  or  extinction. 
For  example,  Microlepia  speluncae  (L.)  T.  Moore  was  collected  in 
Sao  Tome  by  Don  (s.l.),  Quintas  (Ilheu  das  Rolas),  and  Welwitsch 
(Monte  Cafe).  Although  Monte  Cafe  is  a  well  collected  area,  visited 
by  most  botanists,  this  species  has  not  been  found  during  the  last  100 
years.  Elsewhere  in  Africa,  M.  speluncae  is  rare  and  sporadic, 
occurring  at  low  to  medium  altitudes. 

Sixteen  taxa  are  represented  in  herbaria  by  a  single  collection, 
either  from  Sao  Tome  or  from  Principe  (10  from  Sao  Tome  and  6 
from  Principe),  which  may  indicate  their  rarity  or  even  extinction. 
When  Principe  alone  is  considered,  27  of  the  total  of  75  taxa  have 
been  collected  only  once.  In  this  case,  mistaken  recordings  or 
undercollecting  are  also  a  possibility.  Several  species  are  repre- 
sented in  herbaria  by  a  single  collection  made  by  Barter,  who  visited 
Principe  when  stationed  at  Bioko,  and  therefore  it  is  possible  that 
there  were  some  errors  when  the  specimens  were  labelled.  Principe 
remains  poorly  collected  and  recent  collections  of  pteridophytes 
from  that  island  were  not  available  to  confirm  the  presence  of  these 
taxa. 

Seven  taxa  are  apparently  restricted  to  the  Pico  of  Sao  Tome 
[Asplenium  friesiorum  C.  Chr.,  Ctenitis  cirrhosa  (Schum.)  Ching, 
Grammitis  molleri  (Baker)  Schelpe,  G.  tomensis  Schelpe,  Huperzia 
ophioglossoides  (Lam.)  Rothm.,  Lycopodium  clavatum  L.,  and 


Xiphopteris  oosora  (Baker)  Alston]  and  five  taxa  are  confined  to  the 
Pico  of  Principe  [Grammitis  nigrocincta  Alston,  Hymenophyllum 
polyanthos  var.  mildbraedii  (Brause  ex  Brause  &  Hieron.)  Schelpe, 
Lomariopsis  warneckei  (Hieron.)  Alston,  Selaginella  squarrosa 
Baker,  and  Triplophyllum  jenseniae  (C.  Chr.)  Holttum].  Most  of 
these  are  high  altitude  pteridophytes  which  are  also  distributed  on 
other  African  mountains.  Two  of  these  taxa,  Grammitis  molleri  and 
G.  tomensis,  are  very  rare  and  endemic  to  the  Pico  of  Sao  Tome. 
Lycopodium  clavatum  is  cosmopolitan  but,  in  the  tropics,  it  is 
confined  to  mountains. 

Of  the  twelve  taxa  endemic  to  Sao  Tome  and  Principe,  two  are 
apparently  very  rare,  having  been  collected  only  once:  Selaginella 
monodii  Alston  and  Sphaerostephanos  elatus  subsp.  thomensis 
Holttum. 


COLLECTIONS 

Historical  collections 

Exell  (1944)  gives  a  lively  account  of  the  early  collectors  in  Sao 
Tome  and  Principe.  The  majority  of  the  earliest  collections  of 
pteridophytes  from  Sao  Tome  and  Principe  were  made  by  Don  (in 
1822),  Curror  (in  1839),  Welwitsch  (in  1853),  Barter  (in  1858), 
and  Mann  (in  1861).  These  collections  are  deposited  at  K  (Curror, 
Barter,  and  Mann),  BM  (Don  and  Welwitsch),  and  LISU 
(Welwitsch). 

In  the  late  nineteenth  century  extensive  collecting  was  made  by 
the  Portuguese  collectors  Moller,  Quintas,  and  Newton.  The  origi- 
nal set  of  these  collections  is  kept  at  COI,  but  duplicates  were  sent 
to  several  European  herbaria.  The  duplicates  were  distributed  as 
Flora  Africanae  Exsiccata  (F.A.E.},  a  series  in  which  different 
collections  of  the  same  taxon  were  grouped  and  distributed  with 
the  same  number.  For  this  reason,  most  F.A.E.  labels  refer  to 
several  localities  and  different  collectors  After  examining  the 
original  sets  of  the  Moller,  Quintas,  and  Newton  collections,  Exell 
(1944:  383-385)  tried  to  link  originals  with  duplicates.  However, 
it  seems  that,  for  the  pteridophytes,  in  most  cases  the  original 
collections  were  scarsely  labelled,  lacking  the  information  needed 
to  link  them  to  the  duplicates  distributed.  Many  F.A.E.  duplicates 
sent  from  COI  to  K  had  a  second  label  referring  to  a  single 
collector.  The  numbering  on  these  collector  labels  is  unique  to  K 
specimens,  as  the  original  sets  at  COI  are  mostly  unnumbered. 
Although  the  second  label  usually  refers  to  a  single  collector  and 
locality  there  is  no  certainty  as  to  the  correspondence  between  the 
specimen  and  the  data.  The  information  consisting  of  more  than 
one  locality,  found  on  some  F.A.E.  and  original  set  labels,  is 
worthless  and  misleading  because  it  is  not  possible  to  relate  it  to 
the  specimens  it  is  attached  to.  This  information  should  be  omitted 
when  those  collections  are  cited. 

Together  with  the  F.A.E.  collections,  the  collections  made  by 
Chevalier  in  1905  and  Exell  in  1932-33,  the  first  sets  of  which  are 
deposited,  respectively,  at  P  and  BM,  comprise  the  bulk  of  the 
material  examined  to  produce  Alston's  account  of  the  pteridophytes 
in  Exell's  Catalogue  (1944). 

Recent  collections 

Only  a  few  collections  were  made  during  the  twenty  years  that 
followed  Exell's  expedition  in  1932-33.  The  majority  of  recent 
collections  were  made  during  the  years  1954-1973,  and  in  the 
1990s. 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


43 


The  first  set  of  the  collections  made  by  Rozeira  in  Sao  Tome  and 
Principe  in  1954  is  deposited  at  PO.  However,  several  collections  of 
pteridophytes  cited  in  the  literature  (Alston,  1958)  could  not  be 
located  in  that  herbarium.  A  set  of  duplicates  of  Rozeira's  collection 
is  apparently  deposited  at  BM  (Exell  &  Rozeira,  1958)  but  could  not 
be  found  in  1997.  Some  duplicates  (with  the  name  of  the  collector 
missing)  exist  at  K.  Based  on  Rozeira's  collections,  new  records  for 
the  islands  were  published  by  Alston  (1958)  in  the  proceedings  of 
the  'Conferencia  Internacional  dos  Africanistas  Ocidentais',  which 
took  place  in  Sao  Tome  in  1956. 

During  1956  several  botanists  collected  in  Sao  Tome  and 
Principe.  As  part  of  a  Calypso  expedition  which  took  place  in 
1956-57,  Rose  made  several  collections  in  the  two  islands,  in- 
cluding living  material,  which  were  deposited  at  P.  In  the  same 
year,  Monod  and  Thorold  made  important  collections  in  Sao  Tome 
and  Principe,  which  are  deposited  at  BM.  These  collections  were 
studied  by  Alston  (1958,  1959)  and  resulted  in  several  new  records 
being  added  to  the  flora  of  the  islands.  Lains  e  Silva  also  collected 
in  Sao  Tome  in  1956.  Although  his  collections  have  been  cited  in 
the  literature,  several  numbers  could  not  be  found  either  at  LISJC 
or  LISU.  In  1968-1973,  Espirito  Santo  collected  in  both  islands. 
The  original  sets  of  his  collections  are  kept  at  LISJC  and  LISC, 
with  some  duplicates  at  BM  and  COI. 

Recent  collections  of  pteridophytes  from  Sao  Tome  examined 
during  this  study  were  mainly  made  by  Pinto  Basto  in  1990,  Matos 
and  Matos  &  Van  Essche  in  1994,  and  Lejoly  in  1994-1995,  and  are 
deposited  at  LISC.  Recent  collections  were  also  made  by  Paiva 
(COI),  but  these  were  not  available  for  examination  at  the  time  of 
this  study. 


CATALOGUE 

The  catalogue  includes  all  the  taxa  of  pteridophytes  known  to  occur 
in  Sao  Tome  and  Principe,  arranged  in  alphabetical  order,  with 
synonyms  and  references  to  the  literature  for  the  area  published 
since  1944.  Synonyms  already  cited  in  Alston's  (1944)  work  are 
omitted. 

All  the  collections  examined  are  cited  in  this  catalogue.  Collec- 
tions cited  in  the  literature  are  also  included.  Some  of  these  specimens 
were  not  located  (indicated  in  the  text  with  'not  found'),  and  others 
were  not  available  for  study.  Some  of  the  collections  cited  but  not 
seen  by  Alston  ( 1 944),  probably  at  B,  were  not  examined  during  this 
study;  these  are  marked  with  a  question  mark.  It  is  possible  that  a 
future  study  of  the  unseen  material  may  reveal  misidentifications  by 
previous  authors. 

For  each  taxon,  the  collections  are  cited  by  alphabetical  order  of 
collectors,  but  collections  from  the  same  locality  are  grouped. 
Collections  without  locality  are  placed  at  the  end  of  the  specimens 
cited.  These  include  F.A.E.  and  Moller  collections  with  more  than 
one  locality  on  the  labels.  F.A.E.  duplicates  at  K  are  cited  under  their 
F.A.E.  number  and  reference  is  made  to  other  labels  and  numbers  if 
they  exist. 

Most  of  the  localities  provided  by  the  collectors  consist  of  the 
name  of  a  plantation  ('ro9a')  or  a  river.  For  this  reason,  distribution 
maps  based  on  these  collections  would  be  very  inaccurate.  Maps 
with  'ro9as'  and  other  collecting  localities  mentioned  in  the  text  are 
provided,  to  allow  for  an  estimate  of  the  distribution  of  the  taxa  (Figs 
1  and  2). 

Data  on  habit,  habitat,  and  ecology  are  lacking  for  the  vast 
majority  of  the  collections  examined. 


Acrostichaceae 
Acrostichum  L. 

Acrostichum  aureum  L.,  Sp.  pi.  2:  1069  (1753);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  85  (1944);  Benl  inActa  Bot.  Barcinon.  38: 
22(1988). 

SAO  TOME.  Porto  Alegre,  Chevalier  14194bis  (P!),  14200  (P!),  salt 
marshes,  Espirito  Santo  5131  (LISC!,  LISJC!);  Ilheu  das  Rolas,  F.A.E. 
(Quintas)69  (BM!,  LISU!),  Quintass.n.  (COI!);  PortoAlegre,  Praia  Jal<£,  Rio 
Malanza,  swamp,  Matos  7663  (LISC!);  Perseveran?a,  Rose  538  (P!). 
PRINCIPE.  Santo  Antonio,  salt  marshes,  Espirito  Santo  5170  (LISC!, 
LISJC!),  mangrove,  Monod  12191  (BM!),  mangrove,  Rose  425  (P!);  Rio 
Papagaio,  swampy  border,  Quintas  1  (BM!,  COI!);  Praia  General  Fonseca, 
Rozeira  430  (PO!);  s.l.,  Welwitsch  15  (BM!,  LISU!). 

DISTRIBUTION.    Pantropical. 

Adiantaceae 
Adiantum  L. 

Adiantum  mettenii  Kuhn,  Filic.  afr.:  65  (1868)  nomen;  in  Jahrb. 
Konigl.  Bot.  Gart.  Berl  1:  338  (1881). 

Adiantum  soboliferum  sensu  auct.  non  Wall,  ex  Hook.,  Sp.fil.  2:  13, 
t.  74A  (1858);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  82  (1944). 

SAO  TOME.  Monte  Forte,  50  m,  Quintas  s.n.  (COI!,  K!);  s.l.,  Don  s.n. 

(BM!). 

DISTRIBUTION.  Tropical  Africa  and  Madagascar,  extending  to  In- 
dia and  Philippines. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Adiantum  philippense  L.,  Sp.  pi.  2:  1094  (1753);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  82  (1944);  Alston  in  Exell,  Suppl.  Cat. 
Vase.  PL  S.  Tome:  1  (1956);  Benl  in  Acta  Bot.  Barcinon.  32:  26 
(1980). 

SAO  TOME.  Casal  Pires,  waterfall,  200  m,  Espirito  Santo  1 10  (BM!,  COI!, 
LISJC!);  Boa  Entrada,  Chevalier  14518  (P);  Rio  do  Ouro,  1 10  m,  Quintas 
1249  (BM!,  COI!,  LISJC!);  Agua  Coimbra,  100  m,  Quintas  s.n.  (COI!), 
Moller  &  Quintas  s.n.  (LISU!);  Diogo  Vaz,  Rozeira  518  (PO!);  Santa 
Catarina,  between  main  house  and  Lemba,  Rozeira  840  (PO!). 

DISTRIBUTION.    Palaeotropical. 

Adiantum  raddianum  C.  Presl,  Tent,  pterid.:  158  (1836);  Alston  in 
Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  1  (1956);  Alston  in  Exell  & 
Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  79  (1958). 

SAO  TOME.  Quinta  da  Graca,  600  m,  Espirito  Santo  124  (BM!,  COI!, 
LISJC!);  Pouso  Alto,  490  m,  Espirito  Santo  4278  (LISC!,  LISJC!);  Monte 
Cafe,  Sao  Carlos,  secondary  forest,  Espirito  Santo  5048  (LISC!,  LISJC!); 
Sao  Nicolau,  waterfall,  on  wet  rocks,  Espirito  Santo  5178  (LISC!,  LISJC!); 
Saudade,  900  m,  Exell  399  (BM!,  COI!);  Ponta  Figo,  on  the  way  to  Morro 
Vilela,  Lains  e  Silva  357  (LISU!);  between  Pico  and  Monte  Castro,  1200- 
1800  m,  mountain  rainforest,  Lejoly  95/13  (LISC!);  Monte  Cafe,  1010  m, 
Monod  1 1694  (BM!,  COI!),  Oliveira  &  Noronha  s.n.  (LISU!);  NW  of  Pico, 
1000-1200  m,  Monod  12263  (BM!,  COI!);  Agua  Ize\  common,  Pinto  Basto 
199  (LISC!);  Tras-os-Montes,  Calvario,  Rozeira  582  (PO!). 
PRINCIPE.  Esperansa,  near  Ribeira  Camarao,  Rozeira  354  (PO!). 

DISTRIBUTION.  Cultivated  American  species,  escaped  and  natural- 
ized. 


44 


E.  FIGUEIREDO 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 

Adiantum  vogelii  Mett.  ex  Keyserl.  in  Mem.  Acad.  Imp.  Sci.  Saint 
Petersbourg  ser.  7,  22(2):  8,  3 1  ( 1 875);Alston  in  Exell,  Cat.  Vase. 
PI.  S.  Tome:  83  (1944);Tardieu-Blot  inAubrev.,  Fl.  Cameroun  3: 
147  (1964);  Benl  in  Acta  Bot.  Barcinon.  32:  26  (1980). 

PRINCIPE.  Santo  Antonio,  Monod  12072  (BM!,  COI!);  Berimbau,  Newton 
18  (BM!,  COI!,  K!),  Newton  s.n.  (K!);  Cimalo,  Quintas  36  (BM!,  COI!); 
Esperan?a,  between  Montalegre  and  Maquina,  Rozeira  378  (PO!);  s.l.,  Souza 
s.n.  (COI!). 

DISTRIBUTION.     W.  tropical  Africa  to  Angola,  Zanzibar. 


Aspidiaceae  (=Dryopteridaceae) 
Ctenitis  (C.  Chr.)  C.  Chr. 

Ctenitis  cirrhosa  (Schum.)  Ching  in  Sunyatsenia  5:  250  (1940); 
Benl  in  Acta  Bot.  Barcinon.  40:  41  (1991). 

Dryopteris  crinobulbon  (Hook.)  C.  Chr.,  Index  filic.,  Suppl.  3:  84 
(1934);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  61  (1944). 

SAO   TOME.  Pico,   1850  m,  Mann  s.n.  (B,  K!-type  of  Dryopteris 
crinobulbon). 

DISTRIBUTION.     Tropical  and  SE  Africa,  Madagascar,  Comoros 
and  Mascarene  Is. 

NOTE.     This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Ctenitis  lanuginosa  (Willd.  ex  Kaulf.)  Copel.,  Gen.  fit.:  124  (1947); 
Benl  in  Acta  Bot.  Barcinon.  40:  41  (1991). 

Dryopteris  lanuginosa  (Willd.  ex  Kaulf.)  C.  Chr.,  Index  filic.:  273 
(1905);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  62  (1944). 

SAO  TOME.  S.I.,  Mann  s.n.  (K-not  found). 

DISTRIBUTION.     Tropical  and  subtropical   Africa,  Madagascar, 
Comoros,  Seychelles  and  Mascarene  Is. 


45 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Didymochlaena  Desv. 

Didymochlaena  truncatula  (Sw.)  J.  Sm.  in  J.  Bot.  (Hooker)  4:  196 
(1841);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  66  (1944); 
Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  254  (1964);  Benl  in 
Acta  Bot.  Barcinon.  40:  35  (1991). 

SAO  TOME.  Rio  Contador,  1270-1350  m,  F.A.E.  (Moller)  36  (BM!, 
LISU!);Trds-os-Montes,  Gama  s.n.  (CO\!);Mollers.n.  (COI!);  Macambrani, 
1270  m,  Moller  s.n.  (COI!);  Sela  do  Camelo,  near  Lemba,  200  m,  Rozeira 
3450  (PO!);  Monte  Cafe,  760-850  m,  Welwitsch  61  (BM!,  K!,  LISU!). 

DISTRIBUTION.    Pantropical. 
Dryopteris  Adans. 

Dryopteris  pentheri  (Krasser)  C.  Chr.,  Index  filic.:  284  (1905); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  447 
(1959);  Benl  in  Acta  Bot.  Barcinon.  40:  36  (1991). 

Dryopteris  oligantha  sensu  auct.  non  (Desv.)  C.  Chr.,  Index  filic. 
Suppl.  3:  93  (1934);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  62 

(1944). 

SAO  TOME.  Pico,  Chevalier  13651  (P),  14290  (P),  Moller  s.n.  (COI-not 
found,  P!);  between  Monte  Cafe  and  Pico,  Chevalier  14580  (P);  Macambrara, 
Ere// 131  (BM),  1 35  (BM);Tras-os-Montes,  Gama  s.n.  (COI-not  found),  800 
m,  Quintas  1286  (BM!,  COI-not  found),  Rozeira  554  (K!,  PO!);  between 
Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain  rainforest,  1200- 
1800  m,  Lejoly  95/9  (LISC!);  between  Monte  Quinas  and  Quinas  Alias, 
Monod  12007  (BM!);  Monte  Cafe,  1010  m,  Monod  1 1696  (BM!);  Lagoa 
Amelia,  1300  m,  Quintas  1291  (BM!,  COI-not  found),  Rose  199  (P!);  Sao 
Nicolau,  near  waterfall,  Rozeira  182  (K!,  PO!);  Santa  Catarina,  Lemba, 
Rozeira  856  (K!);  s.l.,  1850  m,  Mann  (K!),  F.A.E.  (Moller)  39  (BM!,  LISU!), 
Moller  s.n.  (COI-not  found),  Quintas  10  (K!). 

DISTRIBUTION.    Tropical  and  southern  Africa. 


Fig.  1     Sao  Tome 

1 .  Agua  Cardoso-2B 

2.  Agua  Coimbra-3A 

3.  Agua  Grande-3B-4B-4A 

4.  Agua  Ize  (Ro?a)-4C 

5.  Agua  Pinhao-3B 

6.  Almas-4B 

7.  Antonio  Soares-4B 

8.  Bacelar-2D 

9.  Batepa-4B 

10.  Bemposta  (Ro9a)-3B 
ll.Benfica(Roca)-3B 

12.  Binda(Ro?a)-lC 

13.  Blu-blu-4B 

14.  Boa  Entrada  (Roca)-4A 

15.  Bom  Sucesso  (Ro?a)-3B 

16.  Bombaim  (Roca)-3C 

17.  Cabumbe-2C 

18.  Caixao  Grande-4B 

19.  Calvario-2B 

20.  Cantagalo  (Ro?a)-4C 
21.CaoPequeno-2D 
22.  Cascata-2B 


23.  Chami90-3B 

24.  Colonia  Acoreana 
(Roca)-4C 

25.  Cruzeiro  (Roca)-3C 

26.  Diana  (Ro?a)-4B 

27.  Diogo  Nunes-4A 

28.  Diogo  Vaz  (Roca)-2B 

29.  Dona  Amelia  (Roca)-2B 

30.  Dona  Augusta  (Roca)-3D 

3 1 .  Dona  Eugenia  (Roca)-3D 

32.  Ermelinda  (Roca)-2D 

33.  Estacao  Sousa-2B 

34.  Famosa-4B 

35.  Formoso-3C 

36.  Guaiaquil  (Ro?a)-3C 

37.  Guarda  (Ro?a)-3A 

38.  Ilheu  das  Rolas-2E-2F 

39.  Java  (Roca)-3B 

40.  Jou  (Roca)-2D 

41 .  Lagoa  Amelia-SB 

42.  Lemba  (Roca)- IB 

43.  Macambrara  (Roca)-3B 

44.  Maria  Fernandes-3C 


45.  Milagrosa  (Roca)-4B 

46.  Monte  Cafe  (Roca)-3B 

47.  Monte  Castro-2B 

48.  Monte  Forte  (Ro9a)-2A 

49.  Monte  Mario  (Roc.a)-2E 

50.  Monte  Quinas-2B 

5 1 .  Monte  Rosa  (Roca)-2D 

52.  Monte  Verde-2D 

53.  Morro  Cantagalo-3C 

54.  Morro  Vigia  -3B 

55.  Morro  Vilela-2B 

56.  Nova  Ceilao  (Ro9a)-3B 

57.  Nova  Moca  (Ro9a)-3B 

58.  Novo  Brasil  (Ro9a)-2E 

59.  Novo  Destine  (Ro9a)-3B 

60.  Paga  Fogo  (Ro9a)-2B 

61.  Pedroma  (Ro9a)-4B 

62.  Perseveran9a  (Ro9a)-3D 

63.  Pico-2B 

64.  Pinheira  (Ro9a)-4B 

65.  Ponta  Figo  (Ro9a)-2A 

66.  Porto  Alegre  (Ro9a)-2E 


68.  Praia  Jale-2E 

69.  Praia  Melao  (Ro9a)-5B 

70.  Praia  Pantufo-5B 
71.Quija-2D 

72.  Quinas  Altas  -2B 

73.  Quinta  da  Gra9a  (Roca)-3B 

74.  RibeiraMo9a-lB-2B 

75.  Ribeira  Peixe-3D 

76.  Rio  Abade-3C-4C 

77.  Rio  Abade  (bridge)  -4C 

78.  Rio  Campos-2C 

79.  Rio  Caue-2D 

80.  Rio  Contador-2A-2B 

8 1 .  Rio  do  Ouro  (Ro9a)-3  A 

82.  Rio  Malanza-2E 


89.  Rodia  (Ro9a)-3B 

90.  Santa  Catarina  (Ro9a)-lB 
9  I.Santa  Irene  (Ro9a)-2C 

92.  Santa  Margarida  (Ro9a)-3B 

93.  Santelmo  (Ro9a)-3C 

94.  Sao  Carlos  (Ro9a)-3B 

95.  Sao  Joao  dos  Angolares-3D 

96.  Sao  Miguel  (Ro9a)-lD 

97.  Sao  Nicolau  (Ro9a)-3B 

98.  Sao  Pedro-3B 

99.  Sao  Tome  -4A 

100.  Sao  Vicente-4B 
101.Saudade(Ro9a)-3B 

102.  Sela  do  Camelo-2C 

103.  Tras-os-Montes  (Ro9a)-3B 


83.  Rio  Manuel  Jorge-3B-4B-5B   104.  Trindade-4B 


84.  Rio  Manuel  Jorge 
(waterfall  )-3B 

85.  Rio  Miranda  Guedes-3C- 
3D 

86.  Rio  Paga  Fogo-lB-2B 

87.  Rio  Xufexufe-2D 

88.  Ro9a  Nova  (Ro9a)-3B 


105.  Vale  Carmo  (Ro9a)-3D 

106.  Vieira  Machado  (Roca)-3C 

107.  Vila  Aida  (Ro9a)-3D 

108.  Vila  Conceicao  (Roca)-2E 

109.  Vila  Jose  (Ro9a)-3D 

110.  Zagaia-2C 

111.  Zampalma  (Ro9a)-3B. 


67.  Pouso  Alto  (Ro9a)-3B 

The  following  collecting  localities  could  not  be  located:  Agua  Telha  (Diogo  Vaz),  Agua  Telha  (near  Ribeira  Mo9a),  Casa  do  Pico  (1935  m  alt.),  Casal  Pires 
Correia,  Monte  Figo,  Magodinho,  Morro  Pempem  (Tras-os-Montes),  Quingombo,  Santa  Maria,  Rio  Santo  Antonio. 


46 


E.  FIGUEIREDO 


Fig.  2     Principe 

1 .  Barriga  Branca- 1 C 

2.  Bela  Vista  (Ro?a)- IB 

3.  Berimbau-2B 

4.  Cajamanga- 1 B 

5.  Cimalo-2B 

6.  Dois  Irmaos-2C 

7.  Esperanca  (Ro9a)-lB 


8.  Fundao-lB 

9.  General  Fonseca-2C 
10.  Infante  D.  Henrique 

(Rosa)- 1C 
H.Lapa(Ro9a)-lB 

12.  Mamas- 1C 

13.  Maquina-lB 


14.  Mencorne-lC 

15.  Montalegre  (Roca)-lB 

16.  Morro  do  Leste-lB 

17.  Neves  Ferreira  (Rosa)- 1C 

18.  Oque  Caspar  (Rosa)- IB 

19.  6queNazare-lB 

20.  Oque  Pipi-lB 


21.Pico-lC 

22.  Pico  Papagaio-lB 

23.  Praia  da  Lapa-lB 

24.  Praia  das  Burras-2A 

25.  Precipfcio-2A 

26.  RibeiraBibi-lC 

27.  Ribeira  Cambungo-lC-2C 


28.  Ribeira  Chimboto-lC 

29.  Ribeira  Fria  (Ro?a)-2B 

30.  Ribeira  Jambere-lB-2B 
31.RioPapagaio-lB-2B 

32.  Santo  Antonio-2B 

33.  Sao  Joao  (Rosa)-2B 

34.  Sundi(Ro?a)-lA 
35.TerreiroVelho-2B. 


The  following  collecting  localities  could  not  be  located:  Porto  Real,  Ribeira  Camarao  and  Rio  Bambu-Porco. 


Lastreopsis  Ching 

Lastreopsis  aff .  currori  (Mett.  ex  Kuhn)  Tindale  in  Victoria  Natu- 
ralist 73:  184  (1957);  Tindale  in  Contr.  New  South  Wales  Natl. 
Herb.  3:  338(1965). 

Aspidium  nigritianum  Mett.  ex  Kuhn,  Filic.  afr.:  138  (1868). 
Dryopteris  nigritiana  (Mett.  ex  Kuhn)  Kuntze,  Revis.  gen.  pi.  2:  813 

(1891);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  62  (1944). 
Ctenitis  nigritiana  (Mett.  ex  Kuhn)  Alston  in  Bol.  Soc.  Brot.,  ser.  2, 

30:  12(1956). 

PRINCIPE.  Rio  Bambu-Porco,  60  m,  Navel  138  (LISU!),  141 B  (not  found); 
Pico,  Rozeira  2607  (PO!);  s.l.,  Barter  1906  (K-type  of  Aspidium  nigritianum, 
not  found). 

Tectaria  Cav. 

Tectaria  angelicifolia  (Schum.)  Copel.  in  Philipp.  J.  Sci.,  ser.  C, 
Bot.  2:  410  (1907);  Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  66 


(1944);  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  1  (1956); 
Benl  in  Acta  Bot.  Barcinon.  40:  55  (1991). 

SAO  TOME.  Sao  Vicente,  200  m,  Espirito  Santo  42  (BM!,  COI!,  LISJC!); 
Bombaim,  Monod    11886  (BM!);  between  Sao   Tome  and  Monte  Cafe, 
Welwitsch  62  (BM!,  LISU!);  s.l.,  F.A.E.  (Moller  &  Quintas)  43  p.p.  (BM!, 
COI!,  K  also  Moller  32!,  LISU!),  550  m,  Welwitsch  s.n.  (K!). 
PRINCIPE.  S.l.,  Welwitsch  12b  (BM!,  LISU!). 

DISTRIBUTION.     W.  and  Central  tropical  Africa 

Tectaria  camerooniana  (Hook.)  Alston  in  /  Bot.  77:  288  (1939); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  67  (1944);Tardieu-Blot  in 
Aubrev.,  Fl.  Cameroun  3:  29 1  ( 1 964);  Benl  in  Acta  Bot.  Barcinon. 
40:56(1991). 

SAO  TOME.  Macambrara  to  Zampalma,  900  m,  ground  fern,  Exell  293 
(BM!),  294  (BM!,  COI!);  between  Bom  Sucesso  and  LagoaAmelia,  primary 
forest,  1350-1420  m,Matos  &  Van  Esschel296  (LISC!);  above  Monte  Cafe, 
1010  m,  Monod  1 1693  (BM!);  between  Sao  Pedro  and  LagoaAmelia,  1050 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 

m,  Monod  11711  (BM!);  Calvario,  1200  m,  Monod  11815  (BM!),  11817 
(BM!);  Lagoa  Amelia,  Rose  232  (P!);  s.l.,  F.A.E.  (Moller)  44  (BM!,  COI!, 
LISJC!,  LISU!,  P!). 

DISTRIBUTION.     W.  and  Central  tropical  Africa. 

Tectaria  fernandensis  (Baker)  C.  Chr.,  Index  filic.,  Suppl.  3:  179 
(1934);Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  67  (1944);  Benl  in 
Acta  Bot.  Bardnon.  40:  57  (1991). 

SAO  TOME.  Boa  Entrada,  Chevalier  14360  (P!),  14517  (P!);  SW  region, 
Chevalier  14591  (P!),  14617  (P!);  Sao  Miguel,  50  m,  Espirito  Santo  4742 
(LISJC!);  between  Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain 
rainforest,  1200-1800  m,  Lejoly  95/10  (LISC!);  Sao  Miguel,  Monte  Verde, 
secondary  forest,  10-1 50  m,Lejoly95/%4  (LISC!);  Pedroma,Mocquerys  1 257 
126(P!);SaoJoaodosAngolares,80m,A'eH'fons.n.(COI!),50m,2M/n/a5S.n. 
(COI!);  Agualze,  common,  Pinto  Basto2\  1  (LISC!);  Perseveranca,/?o.se537 
(P!);  s.l.,  Don  s.n.  (BM!),  F.A.E.  (Moller  &  Quintas)  43  p.p.  (LISJC!). 
PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  564  (BM!,  COI!),  565  (BM!,  COI!);  Pico  Papagaio,  600 
m,  ground  fern,  Exell  684  (BM!);  s.l.,  Barter  1907  (K!),  Welwitsch  12  (BM!, 
Ki.LISU!). 

DISTRIBUTION.     W.  and  Central  tropical  Africa. 
Triplophyllum  Holttum 

Triplophyllum  buchholzii  (Kuhn)  Holttum  in  Kew  Bull.  41:  251 
(1986). 

Tectaria  buchholzii  (Kuhn)  Copel.  in  Philipp.  J.  Sci.  38:  138  (1929) 
'buckholzii';  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  67  (1944). 

Ctenitis  buchholzii  (Kuhn)  Alston  in  Bol.  Soc.  Brot.,  ser.  2,  30:  1 1 
(1956);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A, 
21:  448  (1959). 

PRINCIPE.  Pico,  500-600  m,  Monod  12119  (BM!),  800  m,  Rozeira  886 
(PO!),  91 1  (PO!),  2617  (PO!);  Rio  Bambu-Porco,  60  m,  Navel  139B  (LISU!); 
Infante  D.  Henrique,  250  m,  Rozeira  2300  (PO !);  Infante  D.  Henrique,  old  path 
to  Ribeira  Bibi,  Rozeira  2391  (PO!);  Mencorne,  200  m,  Rozeira  241 1  (PO!). 

DISTRIBUTION.     Ivory  Coast  to  Zaire. 

Triplophyllum  fraternum  (Mett.  ex  Kuhn)  Holttum  in  Kew  Bull. 
41:253(1986). 

Aspidium  fraternum  Mett.  ex  Kuhn,  Filic.  afr.:  132  (1868). 
Ctenitisfraterna(Meti.)Tardieu  inNotul.  Syst.  (Paris)  14: 342(1953). 

var.  elongatum  (Hook.)  Holttum  in  Kew  Bull.  41:  254  (1986). 

Nephrodium  subquinquefidum  (P.  Beauv.)  Hook.,  Sp.  fil.  4:  130 
(1862)  var.  elongatum  Hook.,  Sp.fil.  4:  130  (1862). 

PRINCIPE.  S.l.,  Mam  1861  (KMectotype  of  Nephrodium  subquinque- 
fidum var.  elongatum). 

DISTRIBUTION.    Endemic. 

NOTES.  The  following  collections  are  unlocalized  but  may  have 
been  collected  in  Principe,  since  there  are  no  further  collections  from 
the  continent:  Niger  Exped.,  Barter  s.n.  (K!),  s.l.,  Curror  s.n.  (K!). 

Based  on  the  same  type  specimen,  Pichi-Sermolli  (1985)  consid- 
ered this  fern  to  be  a  new  species  (Ctenitis  attenuata  Pic.  Serm.)  and 
later  (Pichi  Sermolli,  1991)  transferred  it  to  Triplophyllum  with  the 
name  T.  attenuatum  (Pic.  Serm.)  Pic.  Serm. 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Triplophyllum  jenseniae  (C.  Chr.)  Holttum  in  Kew.  Bull.  41:  253 
(1986). 


47 

PRINCIPE.  Pico,  850  m,  Rozeira  1021  (PO!). 
DISTRIBUTION.     W.  and  Central  tropical  Africa. 

Triplophyllum  principis  Holttum  in  Kew  Bull.  41:  246  (1986). 

Dryopteris  variabilis  (Hook.)Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome: 
64  (1944)  var.  barteri  (Hook.)  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  64(1944). 

PRINCIPE.  Oque  Pipi,  secondary  forest,  300  m,  ground  fem,  Exell  546 
(BM!,  COI!);  between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  562  (BM!,  COI!);  above  Infante  D.  Henrique,  second- 
ary forest,  300  m,  ground  fern,  Exell  618  (BM!,  COI!),  624  (BM!);  Rio 
Bambu-Porco,  60  m,  Navel  14 la  (LISU!);  Oque  Nazare,  500  m,  Newton  s.n. 
(COI!);  Infante  D.  Henrique,  Rose  402  (P!),  Rozeira  2370  (PO!);  Neves 
Ferreira,  Rozeira  2204A  (PO!),  2205  (PO!);  between  Barriga  Branca  and 
Mamas,  Rozeira  2552  (PO!);  Mencorne,  Rozeira  4299  (PO!);  s.l.,  Barter 
1927  (BM!,  K!-holotype),  Wilde  et  al.  371  (WAG,  cited  by  Holttum,  I.e.). 

DISTRIBUTION.     Endemic. 

Triplophyllum  protensum  (Afzel.  ex  Sw.)  Holttum  in  Kew  Bull. 
41:  247  (1986);  Benl  in  Acta  Bot.  Bardnon.  40:  45  (1991). 

Aspidium  subquinquefidum  P.  Beauv.,  Fl.  Oware  1:  34,  t.  19  (1805). 
Dryopteris protensa  (Afzel.  ex  Sw.)  C.  Chr.,  Indexfilic.:  286  (1905); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  63  (1944). 

SAO  TOME.  Bemposta,  600  m,  Moller  s.n.  (BM!,  COI!). 
PRINCIPE.  SW  Esperan9a,  200  m,  Exell  67 1  a  (BM ! );  Sao  Joao,  Newton  s.n. 
(COI!);  Infante  D.  Henrique,  old  path  to  Ribeira  Bibi,  Rozeira  2390A  (PO!); 
Pico  Papagaio,  Welwitsch  16  (BM!,  LISU!);  s.l.,  Souza  s.n.  (COI!). 

DISTRIBUTION.     Widespread  in  W.  and  Central  tropical  Africa. 

NOTE.  Pichi  Sermolli  (1991)  considered  that  the  type  of 
Triplophyllum  protensum  was  a  hybrid  and  described  this  taxon  as 
a  new  species,  T.  heudelotii  Pic.  Serm.  He  considered 
Triplophyllum  subquinquefidum  (P.  Beauv.)  Pic.  Serm.  to  be  a 
distinct  species.  Of  these  two  taxa  only  the  latter  was  recorded  for 
S.  Tome  and  Principe. 

Triplophyllum  securidiforme  (Hook.)  Holttum  in  Kew  Bull.  41: 
242  (1986);  Benl  in  Acta  Bot.  Bardnon.  40:  43  (1991). 

Dryopteris  securidiformis  (Hook.)  C.  Chr.,  Indexfilic.:  291  (1905). 
D.  securidiformis  var.  nana  Bonap.,  Not.  pterid.  14:  211  (1923); 

Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  64  (1944). 
Triplophyllum  securidiforme  var.  nanum  (Bonap.)  Holttum  in  Kew 

Bull.  41:243(1986). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  Exell  103  (BM!), 
151  (BM!);  Monte  Cafe,  800  m,  Moller  s.n.  (COI!);  between  Monte  Quinas 
and  Quinas  Altas,  Monod  12013  (BM!);  Sao  Joao  dos  Angolares,  200  m, 
Quintas  s.n.  (COI!);  Ribeira  Moca,  Rozeira  21 19  (PO!);  between  Ermelinda 
and  Cabumbe,  Rozeira  2819  (PO!),  2823  (PO!);  Cantagalo.  400  m,  Rozeira 
3152  (PO!);  Sela  do  Camelo,  200  m,  Rozeira  3451  (PO!);  Formoso,  Rozeira 
3537  (PO!);  near  Rio  Miranda  Guedes,  Rozeira  3639  (PO!);  Vieira  Machado, 
400  m,  Rozeira  3666  (PO!);  s.l.,  F.A.E  (Moller  &  Quintas)  40  (BM!,  COI!,  K 
also  Quintas  44!,  LISU!,  P!). 

PRINCIPE.  Rio  Bambu-Porco,  60  m,  Navel  1 39 A  (LISU !),  1 4 1 C  (not  found); 
Infante  D.  Henrique,  Rose  409  (P!),  414  (P!);  Neves  Ferreira,  Rozeira  2201 
(PO!);  Ribeira  Camarao,  Rozeira  22 19(PO!),  2220(PO!),  Rozeira  2268  (PO!); 
between  Ribeira  Jambere  and  Ribeira  Fria,  Rozeira  2237  (PO!);  near  Ribeira 
Cambungo,  Rozeira  2239  (PO !);  Infante  D.  Henrique,  old  path  to  Ribeira  Bibi, 
Rozeira  2305  (PO!),  2326  (PO!);  Infante  D.  Henrique,  Rozeira  2374  (PO!); 
Mencorne,  Rozeira  2408  (PO !);  between  Barriga  Branca  and  Mamas,  Rozeira 
2562  (PO!);  s.l.,  Barter  1929  (P!),  Mann  s.n.  (K-not  found). 

DISTRIBUTION.     W.  and  Central  tropical  Africa. 


48 

Aspleniaceae 
Asplenium  L. 

Asplenium  aethiopicum  (Burm.f.)  Bech.  in  Candollea  6:22(1 935); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  74  (1944);  Benl  in  Acta 
Bot.  Barcinon.  40:  24  (1991). 

SAO  TOME.  Pico,  primary  forest,  1987  m,  epiphyte,  Matos  &  Van  Essche 
7381  (LISC!);  Lagoa  Amelia,  1430  m,  epiphyte,  Matos  7495  (LISC!); 
between  Estasao  Sousa  and  Pico,  primary  forest,  1700  m,  epiphyte,  Matos 
7554  (LISC!);  Macambrara,  1270  m,  Moller  s.n.  (COI!);  Lagoa  Amelia  to 
Sao  Pedro,  1200  m,  Quintas  1392  (BM!,  COI!);  Monte  Cafe,  Quintas  1392 
(LISJC!);  s.l.,  F.A.E.  {Moller)  31  (BM!,  K!,  LISU!),  Moller  s.n.  (COI!). 

DISTRIBUTION.     Pantropical  and  subtropical. 

Asplenium  africanum  Desv.  in  Ges.  Naturf.  Freunde  Berlin  Mag. 
Neuesten  Entdeck.  Gesammten  Naturk.  5:  322  (1811);  Alston  in 
Exell,  Cat  Vase.  PI.  S.  Tome:  74  (1944);  Benl  in  Acta  Bot. 
Barcmon.  40:6(1991). 

SAO  TOME.  Ilheu  das  Rolas,  Greeffs.n.  (B);  near  Sao  Tome,  8  m,  Moller 
s.n.  (COI!);  Agua  Ize,  frequent  epiphyte,  Pinto  Basto2\S  (LISC!);  s.l.,  Don 
s.n.  (BM!),  F.A.E.  (Moller)  21  (BM!,  LISU!,  P!). 

PRINCIPE.  Above  Neves  Ferreira,  200-350  m,  epiphyte,  Exell  650  (BM!, 
COI!);  s.l.,  Newton  (not  found  at  COI). 

DISTRIBUTION.    W.,  Central,  and  E.  Africa. 

Asplenium  anisophyllum  Kunze  in  Linnaea  10:  511  (1836). 

Asplenium  geppii  Carruth.,  Cat.  Afr.  PI.  2:  269  (1901);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  78  (1944). 

SAO  TOME.  Near  Sao  Pedro,  1280  m,  F.A.E.  (Moller)  26  (BM!,  K  also 
Moller  23 !,  LISU!);  between  Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo, 
mountain  forest,  1200-1800  m,  Lejoly  95/18  (LISC!);  NW  of  Pico,  1350- 
1600  m,  Monod  12226  (BM!);  Lagoa  Amelia,  Quintas  1385  (BM!,  COI!), 
Rose  206  (P!). 

DISTRIBUTION.     W.  tropical  Africa. 

Asplenium  barteri  Hook.,  Sec.  cent,  ferns:  t.  75  (1860);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  75  (1944);Tardieu-Blot  inAubrev., 
Fl.  Cameroun  3:  192  (1964);  Benl  in  Acta  Bot.  Barcmon.  40:  13 
(1991). 

SAO  TOME.  Sao  Miguel,  Chevalier  1449 Ibis  (P-not  found);  SW  region, 
Chevalier  14618  (P!);  Porto  Alegre,  Chevalier  14619bis  (P!);  Saudade  and 
Pico,  F.A.E.  (Moller)  25  p.p.  (BM!);  Sao  Joao  dos  Angolares,  80  m,  Newton 
s.n.  (BM!,  COI!);  Cruzeiro,  300  m,  Thorold  2065  (BM!);  s.l.,  Matos  s.n. 
(LISC!). 

PRINCIPE.  Esperanca,  plantation,  100  m,  on  rocks, Exell  677  (BM!,  COI!); 
W.  of  Pico,  above  Lapa,  below  250  m,  Monod  12100  (BM!);  Pico,  near  Lapa, 
below  250  m,  Monod  12103  (BM!);  Sao  Joao,  450  m,  Newton  28  (BM!, 
COI!);  Lapa  (Oeste),  Quintas  25  (BM!,  COI!);  s.l.,  Barter  1909  (K!),  1910 
(K!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Asplenium  biafranum  Alston  &  F.  Ballard  in  Hooker's  Icon.  pi.  34: 
t.  3367  (1938);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  75  (1944); 
Tardieu-Blot  in  Aubrev.,  FL  Cameroun  3:  200  (1964);  Benl  in 
Acta  Bot.  Barcinon.  40:  23  (1991). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14543  (P!); 
Macambrara,  primary  forest,  1050-1200  m,  Exell  154  (BM!);  Pico,  primary 
forest,  1900-2020  m,  Lejoly  94/581  (LISC!);  between  Lagoa  Amelia  and 
Chamico,  Matos  7528  (LISC!);  Santa  Maria,  1350  m,  Moller  s.n.  (COI!); 


E.  FIGUEIREDO 

Lagoa  Amelia,  1400  m,  Monod  11747  (BM!);  Calvario,  1560  m,  Monod 
1 1 834  (B  M ! ) ;  between  Casa  do  Pico  and  Pico,  1 950  m,  Monod  1 1 958  (BM ! ). 

DISTRIBUTION.     Benin,  Bioko,  Nigeria,  Cameroun. 

Asplenium  cuneatum  Lam.,  Encycl.  2:  309  (1786);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  75  (1944);  Tardieu-Blot  in  Aubrev.,  FL 
Cameroun  3:  219(1964). 

SAO  TOME.  Tras-os-Montes,  Morro  Pempem,  1200  m,  Quintas  1387 
(BM!,  COI!). 

DISTRIBUTION.     W.  tropical  Africa,  tropical  America,  Polynesia. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Asplenium  currorii  Hook.,  Sp.  fil.  3:  82  (1860);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  75  (1944);  Alston  in  Exell  in  Bull.  Inst. 
Franc.  Afrique  Noire  ser.  A,  21:  445  (1959);  Tardieu-Blot  in 
Aubrev.,  FL  Cameroun  3:  180  (1964). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  epiphyte,  Exell 
172  (BM!);  between  Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain 
forest,  1200-1800  m,  Lejoly  95/29  (LISC!);  between  Bom  Sucesso  and 
Lagoa  Amelia,  primary  forest,  1350-1420  m,  Matos  &  Van  Essche  7291 
(LISC!);  Moller  s.n.  (COI!);  Lagoa  Amelia,  1500  m,  Monod  1 1722  (BM!); 
s.l.,  550  m,  Welwitsch  64  (BM!,  LISU!). 

PRINCIPE.  Pico  Papagaio,  Welwitsch  24a  (BM!);  s.l.,  Barter  1900  (K-not 
found),  Mann  1861  (K!),  Welwitsch  24  (BM!,  K-not  found,  LISU!). 

DISTRIBUTION.     W.  tropical  Africa. 

Asplenium  dregeanum  Kunze  in  Linnaea  10:517(1 836);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  76  (1944);  Alston  in  Exell,  Suppl 
Cat.  Vase.  PL  S.  Tome:  1  (1956). 

subsp.  brachypterum  (Kunze  ex  Houlston  &  T.  Moore)  Pic.  Serm. 
in  Bull.  Jard.  Bot.  Belg.  55:  130  (1985);  Benl  in  Acta  Bot.  Barci- 
non. 40:  28  (1991). 

SAO  TOME.  Monte  Cafe,  1200  m,  epiphyte,  Espirito  Santo  164  (BM!, 
COI!,  LISJC!);  Macambrara,  primary  forest,  1050-1200  m,  Exell  96  (BM!), 
126  (BM!,  COI!);  Lagoa  Amelia,  1200-1400  m,  Lains  e  Silva  83  (LISU!), 
Matos  7493  (LISC!),  1400  m,  Monod  1 1741  (BM!),  1400  m,  Quintas  1349 
(BM!,  COI!),  Rose  224  (P!),  240  (P!),  Rozeira  154  (PO!),  1 161  (PO!);  Pico, 
primary  forest,  1900-2020  m,  epiphyte,  Lejoly  94/580  (LISC!),  1950  m, 
Moller  s.n.  (COI!);  between  Bom  Sucesso  and  LagoaAmelia,  primary  forest, 
1350-1400  m,  epiphyte,  Matos  &  Van  Essche  7287  (LISC!);  between  Sao 
Pedro  and  Lagoa  Amelia,  1050  m,  Monod  11705  (BM!,  COI!);  Calvario, 
1400  m,  Monod  11852  (BM!);  s.l.,  F.A.E.  (Moller)  33  (BM!,  COI!,  K!, 
LISJC!,  LISU!),  Mann  s.n.  (K!),  Moller  s.n.  (COI!). 

DISTRIBUTION.     Zaire,  Rwanda,  Burundi. 

Asplenium  emarginatum  P.  Beauv.,  Fl.  Oware  2:  6,  t.  61  (1808); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  76  ( 1944);Alston  in  Exell, 
Suppl.  Cat.  Vase.  PL  S.  Tome:  1  (1956);  Tardieu-Blot  inAubrev., 
Fl.  Cameroun  3:  192  (1964);  Benl  in  Acta  Bot.  Barcinon.  40:  10 
(1991). 

SAO  TOME.  Sao  Vicente,  200  m,  common,  Esptrito  Santo  35  (BM!,  COI!, 
LISC!,  LISJC!);  Monte  Cafe,  800  m,    Moller  349  (BM!,  COI!);  Santa 
Margarida,  Antonio  Soares, /toze/ra  1225  (PO!);  s.l.,  Don.  s.n.  (BM!),800m, 
Quintas  8  (K!). 
PRINCIPE?  (Beauvois,  I.e.). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Asplenium  erectum  Bory  ex  Willd.,  Sp.  pi.  5:  328  (1810). 

van  usambarense  (Hieron.)  Schelpe  in  Bol.  Soc.  Brot.,  ser.  2,  41: 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


49 


207  (1967);  Schelpe  in  Exell  &  Launert,  Fl.  Zamb.,  Pteridophyta: 
176  (1970);  Benl  in  Acta  Bot.  Barcinon.  40:  16  (1991). 

Asplenium  quintasii  Gand.  in  Bull.  Soc.  Bot.  France  66:  305  (1919); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  79  (1944);Tardieu-Blot  in 
Aubrev.,  Fl.  Cameroun  3:  198  (1964). 

SAO  TOME.  Monte  Caf6,  Chevalier  13673  (BM!,  P!);  Pico,  1850  m,  Exell 
345a  (BM!),  Newton  s.n.  (BM!,  COI!),  1650  m,  Quintas  1342  (BM!,  COI!; 
type  of  Asplenium  quintasii);  between  Pico  and  Monte  Castro,  6  km  S.  of 
Ponta  Figo,  mountain  forest,  1200-1800  m,  Lejoly  95/19  (LISC!);  between 
Pico  and  Ponta  Figo,  primary  forest,  1600-1700  m,  epiphyte,  Matos  &  Van 
Essche  7389  (LISC!);  Calvario,  1450  m,  Monod  1 1826  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Asplenium  eurysorum  Hieron.  in  Engl.  in  Bot.  Jahrb.  Syst.  46:  364 
(191 1);  Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  76  (1944). 

SAO  TOME.  Pico,  Chevalier  13659  (P!),  primary  forest,  1987  m,  epiphyte, 
Matos  &  Van  Essche  7382  (LISC!);  Calvario,  920  m,  Espirito  Santo  5070 
(LISC!,  LISJC!);  between  Bacelar  and  Quija,  secondary  forest,  160  m, 
Espirito  Santo  5139  (LISC!,  LISJC!);  Macambrara,  primary  forest,  1050- 
1200  m,  ground  fern,  Exell  132  (BM!),  1385  m,  Moller  s.n.  (COI!);  Ribeira 
Peixe,  Lains  e  Silva  301  (LISU!);  between  Pico  and  Monte  Castro,  6  km  S.  of 
Ponta  Figo,  mountain  forest,  1200-1800  m,  Lejoly  95/1  (LISC!);  Santa 
Maria,  1260  m,  Moller  s.n.  (COI!);  between  Sao  Pedro  and  Lagoa  Amelia, 
1050  m,  Monod  11714  (BM!);  between  Monte  Quinas  and  Quinas  Alias, 
Monod  12008  (BM!);  s.l.,  F.A.E.  (Moller)  28  (BM!,  LISJC!,  LISU!,  P!). 
PRINCIPE.  Oque  Nazare,  Newton  27  (BM!,  COI!,  P!);  Pico  Papagaio, 
Welwitsch  14  (BM!,  K!,  LISU!);  s.l.,  Barter  1896b  (K!). 

DISTRIBUTION.     Endemic. 

Asplenium  exhaustum  (H.  Christ)  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  76,  f.  2  (1944);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique 
Noire  ser.  A,  21:  445  (1959). 

SAO  TOME.  Pico,  Chevalier  14587  (P!-type);  Cruzeiro,  300  m,  Thorold 
2072  (BM!);  s.l.,  Seabra  s.n.  (BM!,  COI!,  LISU!). 

DISTRIBUTION.     Endemic. 

Asplenium  formosum  Willd.,  Sp.  pi.  5:  329  (1810);  Alston  in  Exell 
&  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  79  (1958). 

SAO  TOME.  Diogo  Vaz,  near  Agua  Telha,  Rozeira  5 1 8a  (PO). 
DISTRIBUTION.    Pantropical. 

Asplenium  friesiorumC.  Chr.  inNotizbl.  Bot.  Gart.  Berlin-Dahlem 
9:  181  (1924);  Benl  in  Acta  Bot.  Barcinon.  40:  22  (1991). 

SAO  TOME.  Pico,  1950  m,  F.A.E.  (Moller)  27  (BM!,  K!,  LISU!,  P!); 
between  Casa  do  Pico  and  Pico,  1950  m,  Monod  1 1957  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Asplenium  hypomelas  Kuhn,  Filic.  afr.:  104  (1868);  Alston  in 
Exell,  Cat  Vase.  PL  S.  Tome:  78  (1944);  Benl  in  Acta  Bot. 
Barcinon.  40:  31  (1991). 

SAO  TOME.  Porto  Alegre,  Chevalier  13658  (P!);  between  Monte  Cafe  and 
Pico,  Chevalier  14579  (P!);  Lagoa  Amelia,  1400  m,  Espirito  Santo  4069 
(COI!,  LISJC!),  1400  m,  Monod  1 1735  (BM!,  COI!),  Rose 233  (P!),  Rozeira 
1 129  (PO!);  Calvario,  secondary  forest,  920  m,  Espirito  Santo  5074  (LISC!, 
LISJC!),  1200  m,Monod  11813  (BM\),Rose  150 (P!);  Macambrara,  primary 
forest,  1050-1200  m,  ground  fern,  Exell  120  (BM!,  COI!),  1200-1350  m, 
ground  fern,  235a  (BM!,  COI!);Trds-os-Montes,  Gamas.n.  (COI!);  between 
Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain  forest,  1200-1800 
m,  Lejoly  95/16  (LISC!);  between  Bom  Sucesso  and  Lagoa  Amelia,  primary 
forest,  1350-1420  m,Matos&  Van  £wc/je  7295  (LISC!);  Rio  Contador,  1400 


m,  Moller  s.n.  (COI!);  W.  of  Pico,  1225  m,  Monod  11996  (BM!);  Pico, 
Newton  s.n.  (BM!,  COI!);  Sao  Pedro,  Quintas  s.n.  (COI!);  s.l.,  F.A.E.  (Moller 
&  Quintas)  \\(BM\,  COI!,  K  also  Moller  10!,  LISJC!,  LISU!,  P\),Mann  s.n. 
(K!),  Quintas  s.n.  (LISU!),  Souza  s.n.  (COI!). 

DISTRIBUTION.  Widespread  in  tropical  Africa,  extending  to  S.  and 
SE  Africa. 

Asplenium  inaequilaterale  Bory  ex  Willd.,  Sp.  pi.  5:  322  (1810); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  78  (1944);  Benl  in  Acta 
Bot.  Barcinon.  40:  17  (1991). 

SAO  TOME.  Ribeira  Peixe,  Lains  e  Silva  253  (LISU!),  254  (LISU!);  Lagoa 
Amelia,  1430  m,  Matos  7492  (LISC!);  between  Bom  Sucesso  and 
Macambrara,  Matos  7605  (LISC!);  Saudade,  750  m,  Moller  s.n.  (COI!); 
Calvdrio,  Monod  1 1856  (BM!);  s.l.,  F.A.E.  (Moller)  25  p.p.  (BM!,  LISJC!, 
LISU!),  Moller  25  (BM!,  COI!). 

DISTRIBUTION.  Widespread  in  tropical  Africa,  extending  to  S.  and 
SE  Africa  and  Madagascar. 

Asplenium  lividum  Mett.  ex  Kuhn  in  Linnaea  36:  100  (1869). 

Asplenium  protensum  sensu  auct.  non  Schrad.  in  Gott.  Gel.  Anz. 
[1818]:  916  (1818);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  79 
(1944). 

SAO  TOME.  Macambrara  to  Zampalma,  900  m,  ground  fem,  Exell  296 
(BM!,  COI!);  Agua  Pinhao,  850  m,  Quintas  1283  (BM!,  COI!). 

DISTRIBUTION.     E.  and  S.  Africa,  and  S.  America. 

Asplenium  longicauda  Hook.,  Sec.  cent,  ferns:  t.  69  (1860);  Alston 
in  Exell,  Cat  Vase.  PL  S.  Tome:  78  (1944);  Tardieu-Blot  in 
Aubrev.,  FL  Cameroun  3:  1 86  ( 1 964);  Benl  in  Acta  Bot.  Barcinon. 
40:9(1991). 

PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  571  (BM!,  COI!);  Rio  Bambu-Porco,  60  m,  Navel  144 
(LISU!,  P!);  s.l.,  Barter  1900  (K!-syntype). 

DISTRIBUTION.     W.  tropical  Africa,  Madagascar. 

Asplenium  megalura  Hieron.  in  Brause  &  Hieron.  in  Mildbr.,  Wiss. 
Erg.  Deut.  Zentr.-Afr.  Exped.,  Bot.  2:  17(1910);  Benl  inActaBot. 
Barcinon.  40:  23  (1991). 

var.  molleri  (Hieron. )Tardieu  inMem.  Inst.  Franc.  Afrique  Noire  28: 
190(1953). 

Asplenium  molleri  Hieron.  in  Engl.  in  Bot.  Jahrb.  Syst.  46:  371 
(191 1);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  79  (1944). 

SAO  TOME.  Lagoa  Amelia,  1400  m,  epiphyte,  Espirito  Santo  4068  (COI!, 
LISJC!),  crater  rim,  1350  m,  epiphyte,  Exell  202  (BM!),  1430  m,  rare 
epiphyte,  Matos  7496  (LISC!),  1400  m,  Monod  1 1779  (BM!);  Macambrara, 
primary  forest,  1200  m,  epiphyte,  Exell  228  (BM!);  Bom  Sucesso,  1075  m, 
F.A.E.  (Moller)  29  (BM!,  COI!,  K!-type,  LISU!);  between  Bom  Sucesso  and 
Lagoa  Amelia,  secondary  forest,  1 150-1300  m,  Matos  &  Van  Essche  7310 
(LISC!);  Calvario,  1400  m,  Monod  1 1854  (BM!);  W.  of  Pico,  870-1225  m, 
Monod  1 1905  (BM!);  s.l.,  Matos  s.n.  (LISC!). 

DISTRIBUTION.  The  species  is  widespread  in  tropical  Africa.  The 
variety  is  endemic  to  Sao  Tome. 

Asplenium  nigritianum  Hook.,  Sp.fil.  3:  223  (1860)  et  Sec.  cent, 
ferns:  t.  44  (1860);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  79 
(1944);  Alston  in  Bol.  Soc.  Brot.,  ser.  2,  30:  10  (1956). 

SAO  TOME.  S.l.,  Mann  1 107  (K-not  found). 

PRINCIPE.  Lapa  (Leste),  Quintas  s.n.  (BM!,  COI!);  s.l., Barter  1895  (BM!, 

K-not  found),  (probably  Lapa),  Quintas  B  (K!). 


50 

DISTRIBUTION.    Tropical  Africa. 

NOTES.  As  shown  by  Alston  ( 1956£),  it  was  wrongly  assumed  that 
the  type  collection  came  from  Bioko.  In  fact,  Asplenium  nigritianum 
does  not  seem  to  occur  in  Bioko. 

Alston  (19566)  referred  to  the  type  collection  as  Barter  225, 
explaining  that  Hooker  mistook  the  date,  1 859,  for  the  number  and 
then  miscopied  it  as  1 895  (in  K  specimen)  or  1 898  (in  BM  speci- 
men). Since  the  specimen  at  K  was  not  found,  it  was  not  possible  to 
check  its  number.  The  collections  of  pteridophytes  made  by  Barter 
which  I  examined  range  from  no.  1886  to  1928,  so  it  is  likely  that 
these  figures  refer  to  the  number  and  not  the  date  of  collection.  The 
type  collection  should  be  referred  to  as  Barter  1895. 

The  species  is  poorly  represented  in  herbaria.  Neither  the  speci- 
men Barter  1895  (K)  nor  Mann  1107  (K)  could  be  located.  In  a 
handwritten  document  kept  at  BM,  reference  is  made  to  a  duplicate 
of  the  Mann  collection  kept  at  B. 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Asplenium  paucijugum  F.  Ballard  in  Hooker's  Icon.  pi.  33:  t.  3287 
(1935);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  79  (1944);  Benl  in 
Acta  Bot.  Barcinon.  40:  8  (1991). 

Asplenium  variabile  Hook.,  Sp.  fil.  3:  93,  t.  185  (1860)  var. 
paucijugum  (F.  Ballard)  Alston  in  Bol.  Soc.  Brot.,  ser.  2,  30:  7 
(1956);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  184  (1964). 

SAO  TOME.  Monte  Cafe,  800  m,  F.A.E.  (Moller)  24  (BM!,  COI!,  LISJC!, 
LISU!);  LagoaAmelia,  primary  forest,  1400-1420  m,  epiphyte,  Matos  &  Van 
Essche  7410  (LISC!);  NW  of  Pico,  1000-1350  m,  Monod  12216  (BM!);  s.l., 
800  m,  Moller  19  (K!),  550  m,  Welwitsch  63  (BM!,  LISU!). 
PRINCIPE.  Sundi,  Newton  29  (BM!,  COI!). 

DISTRIBUTION.     W.,  Central,  and  E.  tropical  Africa,  Madagascar. 

Asplenium  sandersonii  Hook.,  Sp.fil.  3:  147,  t.179  (1860);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  80  (1944);  Benl  in  Acta  Bot. 
Barcinon.  40:  27  (1991). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  epiphyte,  Exell 
144  (BM!,  COI!);  Sao  Miguel,  between  Xufexufe  and  Sao  Miguel,  400  m, 
epiphyte,  Matos  7658  (LISC!);  Pico,  1950  m,  Moller  s.n.  (COI!);  Bombaim, 
Rose  156  (P!);  DonaAugusta,  Dona  Eugenia,  epiphyte,  Rozeira  1096  (PO!); 
LagoaAmelia,  1400  m,  epiphyte,  Sergio  s.n.  (LISU!);  s.l.,  F.A.E.  (Moller)  23 
(BM!,  K  also  Moller  21 !,  LISU!),  Mann  (not  found),  Moller  s.n.  (COI!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical  Africa. 

Asplenium  unilaterale  Lam.,  Encycl.  2:  305  (1786);  Alston  in 
Exell,  Cat  Vase.  PL  S.  Tome:  80  (1944);  Benl  in  Acta  Bot. 
Barcinon.  40:  15  (1991). 

SAO  TOME.  Pico,  1300-1900  m,  EA.E.  (Moller)  30  (K-not  found,  LISU!); 
Pico,  1900  m,  Moller  s.n.  (COI!);  Rio  Contador,  1300  m,  Moller  s.n.  (COI!). 
PRINCIPE.  S.L,  Barter  1917  (K!). 

DISTRIBUTION.    Palaeotropical. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Asplenium  variabile  Hook.,  Sp.fil.  3:  93,  t.  185  (1860);  Alston  in 
Exell,  Cat  Vase.  PI.  S.  Tome:  80  (1944);  Benl  in  Acta  Bot. 
Barcinon.  40:  7  (1991). 

SAO  TOME.  16  Grande,  Sao  Joao  dos  Angolares,  50  m,  EA.E.  (Quintas)  22 
(K  also  Quintas  20!,  LISU!);  between  Xufexufe  and  Sao  Miguel,  400  m,  epi- 
phyte, Matos  7657  (LISC!);  Sao  Joao  dos  Angolares,  50  m,  Quintas  s.n. 
(COI!). 


E.  FIGUEIREDO 

PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  on  rocks,  Exell  567  (BM!,  COI!);  W.  of  Pico,  above  Lapa,  below  250  m, 
Monod  12099  (BM!);  Rio  Bambu-Porco,  60  m,  Navel  140  (LISU!,  P!);  Pico, 
Thorold  2085  (BM!);  s.l.,  Mann  s.n.  (K-not  found). 

DISTRIBUTION.    W.  tropical  Africa. 

NOTE.  Alston  (1944)  cited  the  collection  Barter  1910  (BM,  K).  At 
BM,  only  the  specimen  Barter  s.n.  (Niger  Expedition)  was  found. 
Barter  1910  (K)  is  Asplenium  barteri. 

Athyriaceae 
Athyrium  Roth 

Athyrium  newtonii  (Baker)  Diels  in  Engl.  &  Prantl,  Nat. 
Pflanzenfam.  1(4):  224  (1899);  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  73  (1944). 

SAO  TOME.  Pico,  1650  m,  Quintas  1393  (BM!,  COI!,  LISJC!),  1650  m, 
Quintas  s.n.  (COI!);  s.l.,  Newton  2  (K!-type),  Newton  s.n.  (COI!),  (probably 
Pico),  1650  m,  Quintas  9  (K!). 

DISTRIBUTION.     Also  in  Uganda  and  Tanzania,  above  1000  m. 

NOTES.  Newton  88  (K)  cited  by  Alston  (I.e.)  is  most  probably  a 
mistake  for  Quintas  9  (the  number  88  after  the  collector's  name  is 
the  date). 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Diplazium  Sw. 

Diplazium  arborescens  (Bory)  Sw.,  Syn.fiL:  92  (1806);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  73  (1944). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14290  (P!);  Pico, 
Chevalier  s.n.  (P!);  Macambrara,  primary  forest,  1050-1200  m,  Exell  128 
(BM!,  COI!);Tras-os-Montes,  Gama  s.n.  (COI!);  between  LagoaAmelia  and 
Chamico,  primary  forest,  1400  m,  Matos  7526  (LISC!);  Monte  Cafe,  Moller 
s.n.  (COI!);  LagoaAmelia,  1350-1380  m,  Monod  1 1772  (BM!,  COI!),  Rose 
234  (P!),  Rozeira  1139  (K!,  PO!);  Cantagalo,  400  m,  Rozeira  3157  (PO!); 
Formoso,  Rozeira  3236  (PO!);  Nova  Ceilao,  Rozeira  4905  (PO!);  s.l.,  EA.E. 
(Moller)  34  (BM!,  COI!,  K  also  Moller  28!,  LISJC!,  LISU!,  P!),  Mann  621 
(P!),  1200  m,  Mann  s.n.  (K!),  Moller  s.n.  (COI!),  Rozeira  2846  (PO!). 

DISTRIBUTION.     Also  in  Madagascar,  Comoros,  and  Mascarene  Is. 

Diplazium  proliferum  (Lam.)  Thouars,  Esquisse  fl.  Tristan 
d'Acugna:  35  (1804);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  73 
(1944);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  237  (1964). 

Callipteris  prolifera  (Lam.)  Bory,  Voy.  ties  Afrique  1:  283  (1804); 
Benl  in  Acta  Bot.  Barcinon.  38:  37  (1988). 

SAO  TOME.  Between  Bom  Sucesso  and  Lagoa  Amelia,  Almeida  s.n. 
(LISC!);  Boa  Entrada,  Chevalier  14333  (P!),  14361  (P!)  14361bis  (not 
found);  Ribeira  Peixe,  S.  of  Cabumbe,  secondary  forest,  350  m,  Espirito 
Santo  3916  (COI!,  LISJC!);  Morro  Vigia,  800  m,  Espirito  Santo  4253 
(LISC!,  LISJC!);  Monte  Rosa,  secondary  forest,  50  m,  Espirito  Santo  4748 
(LISC!,  LISJC  !);Tras-os-Montes,  Lains  e  Silva  229  (LISU!);  between  Monte 
Castro  and  Ponta  Figo,  secondary  forest,  500-1070  m,Lejoly95/45  (LISC!); 
Dona  Amelia,  Santa  Catarina,  near  Agua  Cardoso,  Matos  7593  (LISC!); 
Pedroma,  Mocquerys  89/90  (P!),  131/132  (P!);  Nova  Moca,  near  Monte 
Cafe,  90  m,  Moller  s.n.  (COI!);  Pinheira,  Moller  s.n.  (COI!);  Rodia,  540  m, 
Moller  s.n.  (COI!);  Calvario,  1200  m,  Monod  11816  (BM!);  waterfall  near 
Sao  Nicolau,  Monod  12037  (BM!,  COI!),  Rozeira  571  (PO!);  Cruzeiro, 
Rozeira  765  (PO!);  between  Milagrosa  and  Rio  Abade,  near  Formoso, 
Rozeira  3188  (PO!);  on  the  way  from  Santa  Irene  to  the  river,  Rozeira  3319 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 

(PO!);  Sela do  Camelo,  near  Lembd,  300-700  m,  Rozeira  3459  (PO!);  Monte 
Cafe,  760  m,  Welwitsch  60  (BM!,  K!,  LISU!);  s.l.,  Don  49  (BM!),  F.A.E. 
(Moller)  35  (BM!,  COI!,  LISJC!,  LISU!,  P!). 

PRINCIPE.  Between  Oque  Pipi  and  Morro  de  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  563  (BM!,  COI!);  Rio  Bambu-Porco,  60  m,  Navel  137 
(LISU!,  P!);  Infante  D.  Henrique,  Rose  413  (P!);  Infante  D.  Henrique,  near 
Ribeiro  Chimboto,  Rozeira  3 1 2  (PO ! );  Lapa,  way  to  Pico,  300  m,  Rozeira  9 1 5 
(PO!);  near  Ribeira  Camarao,  Rozeira  2213  (PO!);  Bela  Vista,  near  main 
house,  Rozeira  3916  (PO!). 

DISTRIBUTION.     Palaeotropical. 


Blechnaceae 
Blechnum  L. 

Blechnum attenuatum  (Sw.)  Mett.,  Fit.  hort.  hot.  Lips.:  (A,  t.  3,  figs 
1-6  (1856);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  80  (1944); 
Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  295  (1964);  Benl  in 
Acta  Bot.  Barcinon.  38:  63  (1988). 

SAO  TOME.  Macambrara  to  Zampalma,  primary  forest,  800-1050  m,  Exell 
437  (BM!,  COI!),  438  (BM!,  COI!);  between  Casa  do  Pico  and  Pico,  1 250  m, 
Mono</11960(BM!). 

DISTRIBUTION.     Tropical  and  S.  Africa,  Madagascar. 

Cyatheaceae 

Cyathea  Sm. 

Cyathea  camerooniana  Hook,  in  Hook.  &  Baker,  Syn.  fil:  21 
(1865);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  60  (1944). 

var.  currorii  Holttum  in  Kew  Bull.  36:  480  (1981). 

Alsophila  camerooniana  var.  camerooniana  sensu  auct.  non  (Hook.) 
R.M.  Tryon  in  Contr.  Gray  Herb.  200:  30  (1970);  Benl  in  Acta 
Bot.  Barcinon.  31:  26  (1978). 

PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  Exell  556  (BM!,  COI!);  Rio  Bambu-Porco,  60  m,  Navel  136  (COI!,  K!, 
LISU!,  P!,  WAG);  Infante  D.  Henrique,  Rose  400  (P!);  Infante  D.  Henrique, 
near  Ribeiro  Chimboto,  Rozeira  325  (PO!);  s.l.,  Strickland  s.n.  (BM!,  COI- 
not  found). 

DISTRIBUTION.  Apparently  this  variety  occurs  only  in  Principe, 
where  the  type  (W.  Africa,  S.  of  Tropics,  Curror  s.n.  (K!))  was 
probably  collected. 

Cyathea  manniana  Hook,  in  Hook.  &  Baker,  Syn.  fil.:  21  (1865); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  60  (1944);  Holttum  in 
Kew  Bull.  36:472(1981). 

Alsophila  manniana  (Hook.)  R.M.  Tryon  in  Contr.  Gray  Herb.  200: 
30  (1970);  Benl  in  Acta  Bot.  Barcinon.  31:  27  (1978). 

SAO  TOME.  Lagoa Amelia,  1400  m,  Espirito  Santo  5035  (LISC!,  LISJC!), 
1380  m,  Monod  11763  (BM!),  Quintas  s.n.  (COI!),  Rozeira  1150  (PO!); 
Macambrara,  primary  forest,  1050-1200  m,  Exell  244  (BM!);  Pico,  primary 
forest,  1900-2020  m,  Lejoly  94/577  (LISC!),  1850  m,  Moller  s.n.  (COI!); 
between  Quinas  Alias  and  Monte  Quinas,  Monod  12021  (BM!);  between 
Casa  do  Pico  and  Pico,  1950  m,  Monod  12242ter  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Cyathea  welwitschii  Hook,  in  Hook.  &  Baker,  Syn.  fil.:  21  (1865); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  60  (1944);  Holttum  in 
Kew  Bull.  36:475(1981). 


51 

Alsophila  welwitschii  (Hook.)  R.M. Tryon  in  Contr.  Gray  Herb.  200: 
31  (1970). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  Exell  \  1 1  (BM!, 
COI!),  233  (BM!,  COI!);  Bom  Sucesso,  1200  m,  F.A.E.  (Moller)  2  (K!, 
LISU!,  P!),  1200  m,  Moller  s.n.  (BM!,  COI!);  Pico,  1500-1850  m,  F.A.E. 
(Moller)  3  (K!,  LISU!),  1850  m,  Moller  s.n.  (COI!);  Tris-os-Montes,  Gama 
s.n.  (COI!),  900-1200  m,  Henriques  s.n.  (P!);  between  Quinas  Alias  and 
Monte  Quinas,  Monod  12010  (BM!);  Monte  Cafe\  600  m,  Welwitsch  66 
(BM!,  K!,  LISU!);  s.l.,  Mannl  41/1  (P!),  Mann  1 104  (KMectotype). 

DISTRIBUTION.     Endemic. 
Davalliaceae 
Davallia  Sm. 

Davallia  chaerophylloides  (Poir.)  Steud.,  Nomencl.  hot.  2:  146 
(1824);  Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  72  (1944);  Alston 
in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  1  ( 1 956);  Benl  in  Acta  Bot. 
Barcinon.  33:  27  (1982). 

SAOTOME.  Sao  Vicente,  200  m,  epiphyte,  Espirito  Santo  107  (BM!,  COI!, 
LISJC!);  Benfica,  350  m,  epiphyte,  F.A.E.(Moller)  9  (BM!,  LISU!);  Benfica, 
near  Trindade,  350  m,  Moller  s.n.  (COI!);  Rio  do  Ouro,  350  m,  Moller  s.n. 
(COI !);  near  Diana,  epiphyte,  Rozeira  1 2 1 0  (PO !);  s.l.,  460-600  m,  Welwitsch 
56  (BM!,  K!,  LISU!). 


Widespread  in  tropical  and  S.  Africa  and  Mascarene 


DISTRIBUTION. 
Is. 

NOTE.  Nooteboom  ( 1 994)  synonymizedDava///a  chaerophylloides 
with  D.  denticulata  (Burm.f.)  Mett.  ex  Kuhn  var.  denticulata. 

Humata  Cav. 

Humata  repens  (L.f.)  Diels  in  Engl.  &  Prantl,  Nat.  Pflanzenfam. 
1(4):  209  (1899);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  71 

(1944). 

SAO  TOME.  Sao  Miguel,  Newton  s.n.  (P!). 
DISTRIBUTION.     Palaeotropical. 

NOTES.  Nooteboom  (1994)  included  Humata  in  Davallia  [D. 
repens  (L.f.)  Kuhn]. 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Dennstaedtiaceae  (incl.  Hypolepidaceae) 

Blotiella  R.M.  Tryon 

Blotiella  currorii  (Hook.)  R.M.  Tryon  in  Contr.  Gray  Herb.  191: 99 
(1962);  Benl  in  Acta  Bot.  Barcinon.  38:  29  (1988). 

Lonchitis  currorii  (Hook.)  Mett.  ex  Kuhn  in  Von  der  Decken's 
Reisen  Ost-Afr.,  Bot.  3:  10  (1879);  Alston  in  Exell,  Cat.  Vase.  PI. 
S.  Tome:  85  ( 1944); Alston  in  Exell  &  Rozeira  in  Conf.  Int.Afr.  Oc. 
3:  79  (1958);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire 
ser.  A,  21:  441  (1959);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3: 
100(1964). 

SAO  TOME.  Nova  Moca,  1 100  m,  Quintas  6  (K!,  LISU!);  Lagoa  Amelia, 
1400  m,  Quintas  1262  p.p.  (P!),  1400  m, Quintas  1455  (BM!,  COI!,  K!);  s.l., 
Quintas  1287  p.p.  (P!). 

PRINCIPE.  Infante  D.  Henrique,  Lains  e  Silva  458  (LISU!),  459  (LISU!); 
Pico,  500-600  m,  Monod  12109  (BM!),  Rozeira  913  (PO). 


DISTRIBUTION. 
bia. 


W.  tropical  Africa,  extending  to  Sudan  and  Zam- 


52 


E.  FIGUEIREDO 


NOTE.  Newton  s.n.  (P!),  from  Principe,  cited  but  not  seen  by 
Alston  (1944),  consists  of  sterile  young  fronds. 

Blotiella  glabra  (Bory)  R.M.  Tryon  in  Contr.  Gray  Herb.  191:  99 
(1962);  Benl  in  Acta  Bot.  Barcinon.  38:  31  (1988). 

Lonchitis  glabra  Bory,  Voy.  ties  Afrique  1:  321  (1804);  Alston  in 

Exell,  Cat.  Vase.  PL  S.  Tome:  85  (1944). 
Lonchitis  gracilis  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  1 

(1956);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:102  (1964). 

SAO  TOME.  Porto  Alegre,  Chevalier  14311  (P!);  Pico,  1700-1900  m, 
F.A.E.  (Moller)  13  (BM!,  COI!,  LISJC!,  LISU!,  P!),  Mann  s.n.  (K!),  1750- 
1900  m,  Moller  s.n.  (COI!);  Lagoa  Amelia,  1400  m,  Quintas  1262  (BM!). 

DISTRIBUTION.     Tropical  and  S.  Africa,  Madagascar  and  Reunion. 

Blotiella  mannii  (Baker)  Pic.  Serm.  in  Webbia  31:  250  (1977);  Benl 
in  Acta  Bot.  Barcinon.  38:  30  (1988). 

Lonchitis  mannii  (Baker)  Alston  in  Bol.  Soc.  Brot.,  ser.  2,  30:  18 
(1956);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  80 
(1958). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  ground  fern,  Exell 
239  (BM!,  COI!),  Quintas  1262A  (BM!);  Tras-os-Montes,  Quintas  1287 
(BM!,  COI!):  s.l.,  Quintas  1262  p.p.  (COI!,  P!). 

PRINCIPE.  Pico,  Rozeira  903  (BM-not  found,  PO!),  800  m,  Rozeira  912 
(K!),  918  (K!,  PO!),  600  m,  Rozeira  1075  (K!). 

DISTRIBUTION.     W.  tropical  Africa. 

Histiopteris  (J.G.  Agardh)  J.  Sm. 

Histiopteris  incisa  (Thunb.)  J.  Sm.,  Hist.fil:  295  (1875);  Alston  in 
Exell,  Cat  Vase.  PL  S.  Tome:  86  (1944);  Benl  in  Acta  Bot. 
Barcinon.  38:  25  (1988). 

SAO  TOME.  Near  Sao  Pedro,  F.A.E.  (Moller)  19  (BM!,  LISU!);  Macambrara, 

1280m,  Moller  s.n.  (COI!). 

PRINCIPE.  Oque  Nazare,  Newton  s.n.  (COI!). 

DISTRIBUTION.     Pantropical. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Hypolepis  Bernh. 

Hypolepis  sparsisora  (Schrad.)  Kuhn,  Filic.  afr.:  120  (1868); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  82  (1944);  Benl  in  Acta 
Bot.  Barcinon.  38:  27  (1988). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  Exell  1 12  (BM!, 
COI!),  1200  m,  Quintas  4  (COI!-also  with  no.  1354,  K!),  1200  m,  Quintas 
1354  (BM!,  COI!,  LISJC!);  Tras-os-Montes,  Gama  s.n.  (COI!). 

DISTRIBUTION.  Tropical  and  S.  Africa,  Madagascar,  Comoros  and 
Mascarene  Is. 

Lonchitis  L. 

Lonchitis  occidentalis  Baker  in  Hook.  &  Baker,  Syn.  fil:  128 
(1867);  Benl  in  Acta  Bot.  Barcinon.  38:  34  (1988). 

Anisosorus  occidentalis  (Baker)  C.  Chr.,  Cat.  pi.  mad.,  pter.:  54 
(1932);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  85  (1944). 

SAO  TOME.  Macambrara,  1290  m,  Moller  s.n.  (COI!);  Sao  Joao  dos 
Angolares,  200  m  alt,  Quintas  s.n.  (COI!);  s.l.,  F.A.E  (Moller  &  Quintas)  14 
(BM!,  K  also  Moller  1 1 !,  LISU!). 

DISTRIBUTION.     Widespread  in  tropical  and  SE  Africa,  Madagascar. 


NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Microlepia  C.  Presl 

Microlepia  speluncae  (L.)T.  Moore,  Index  fil.:  93  (1857);  Alston  in 
Exell,  Cat  Vase.  PL  S.  Tome:  72  (1944);  Benl  in  Acta  Bot. 
Barcinon.  38:  23(1988). 

SAO  TOME.  Ilheu  das  Rolas,  Quintas  s.n.  (BM !,  COI!);  Monte  Cafe,  760  m, 
Welwitsch  56(2)  (BM!,  LISU!);  s.l.,  Don  s.n.  (BM!),  F.A.E.  (Moller  & 
Quintas)  10  (K-not  found,  LISU!),  760  m,  Welwitsch  56  (K!,  LISU!). 

DISTRIBUTION.     Palaeotropical,  rare  and  sporadic. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Odontosoria  Fee 

Odontosoria  chinensis  (L.)  J.  Sm.  in  Seem.,  Bot.  voy.  Herald:  430 

(1857). 

van  divaricata  H.  Christ  in  J.  Bot.  (Morot),  ser.  2,  2:  23  (1909). 

Stenoloma  chinense  (L.)  Bedd.,  Handb.  ferns  Brit.  Ind.:  70  (1883) 

var.  divarication  (H.  Christ)  Alston  in  Exell,  Cat  Vase.  PL  S. 

Tome:  73  (1944). 
Sphenomeris  chusana  (L.)  Copel.  in  Bernice  P.  Bishop  Mus.  Bull. 

59:  69  (1929)  var.  divaricata  (H.  Christ)  Tardieu  in  Humbert,  Fl. 

Madag.  et  Comoros  1,  fam.  5:  29  (1958). 
Sphenomeris  chinensis  (L.)Maxonin/  Wash.  Acad.  Sci.  3: 144(1913) 

var.  divaricata  (H.  Christ)  Kramer  in  Blumea  15:  572  (1968). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14309  (P!-type  of 
the  var.);  Calvario,  1500  m,  ground  fern,  Exell  321  (BM!,  COI!);  Pico,  1500- 
\900m, F.A.E.  (Moller)  12 (BM!, COI!, LISJC!, LISU!),  1900m,Mo//ers.n. 
(COI!),  Newton  s.n.  (BM!,  COI!);  Tras-os-Montes,  Gama  s.n.  (COI!);  be- 
tween Calvario  and  Esta?ao  Sousa,  primary  forest,  1500  m,  Matos  &  Van 
Essche  7357  (LISC!). 

DISTRIBUTION.  Occurs  also  sporadically  in  tropical  Asia  and 
Oceania,  and  in  Madagascar,  Reunion,  Mauritius  and  Seychelles. 

NOTE.  Kramer  (1972)  suggested  that  the  collectionsMo//e/;  Gama, 
Exell,  Newton  and  F.A.E.  (Moller),  cited  above,  might  represent  an 
undescribed  species  of  Sphenomeris  (=  Odontosoria),  noting  that 
more  material  and  field  studies  are  required  to  resolve  the  problem. 

Pteridium  Gled.  ex  Scop. 

Pteridium  aquilinum  (L.)  Kuhn  in  Von  der  Decken's  Reisen  Ost- 
Afr.,  Bot.  3:  1 1  (1879);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  86 
(1944);  Benl  in  Acta  Bot.  Barcinon.  38:  35  (1988). 

SAO  TOME.  Boa  Entrada,  Chevalier  13791  (P);  between  Macambrara  and 
Sao  Nicolau,  secondary  woodland,  1050  m,  common,  Exell  272  (BM!, 
COI!);  Nova  Moca,  800-900  m,  F.A.E.  (Moller)  17  (LISU!);  between  Morro 
Vigia  and  Guarda,  Rozeira  688  (PO!);  Pico,  1900  m,  Monod  1 1920  (BM!); 
Monte  Cafe,  760  m,  Welwitsch  59  (BM!,  LISU!);  s.l.,  Moller  s.n.  (COI!), 
Welwitsch  57  (LISU!). 

PRINCIPE.  Cajamanga,  Rozeira  633  (PO!);  Pico  Papagaio,  Welwitsch  18 
(BM!,  LISU!). 

DISTRIBUTION.     Cosmopolitan. 

Elaphoglossaceae 

Elaphoglossum  Schott  ex  J.  Sm. 

Elaphoglossum  acrostichoides  (Hook.  &  Grev.)  Schelpe  in  J.  S. 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


53 


African  Bot.  30:  196  (1964);  Schelpe  in  Contr.  Bolus  Herb.  1:  29 
(1969);  Benl  in  Acta  Bot.  Barcinon.  40:  71  (1991). 

SAO  TOME.   Macambrara,  Exell  153  (BM!);  Calvario,  Monod    11846 
(BM !);  Pico,  Rose  311  (P!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical  Africa, 
Madagascar,  Comoros  and  Reunion. 

Elaphoglossum  aubertii  (Desv.)  T.  Moore,  Index  fit.  2:  5  (1857); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  91  (1944);  Schelpe  in 
Contr.  Bolus  Herb.  1:  32  (1969);  Benl  in  Acta  Bot.  Barcinon.  40: 
75(1991). 

SAO  TOME.  Pico,  1900  m,  F.A.E.  (Moller)  63  (K  also  Moller  561,  LISU!), 
1900  m,  Moller  s.n.  (BM!,  COI!);  Calvario,  1550  m,  Monod  1 1841  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical   Africa, 
Madagascar,  Comoros  and  Mascarene  Is. 

Elaphoglossum  chevalieri  H.  Christ  in  7.  Bot.  (Morot),  ser.  2,  2:  23 
(1909);  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  8  (1956); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  447 
(1959);  Schelpe  in  Contr.  Bolus  Herb.  1:  33  (1969). 

Elaphoglossum  clarenceanum  sensu  auct.  non  (Baker)  C.  Chr., 
Index filic.:  304  (1905);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  92 

(1944). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14292bis  (P!-type); 

Pico,  1950  m,  Moller  s.n.  (COI!),  Rose  298  (P!);  Calvario,  1550  m,  Monod 

1 1837  (BM!);  near  Casa  do  Pico,  1930  m,  Monod  1 1953  (BM!);  E.  of  Pico, 

1700 m,  Quintas  1379 (BM !, COI!);  s.l., F.A.E.  (Moller)64 (LISU!), Henriques 

5/84  p.p.  (K  also  labelled  F.A.E.  (Moller)  64!). 

PRINCIPE.  Pico,  700-800  m,  Monod  12134  (BM!,  COI!),  above  300  m, 

77zoro«2087(BM!). 

DISTRIBUTION.     W.  tropical  Africa,  SE  Africa. 

Elaphoglossum  isabelense  Brause  in  Brause  &  Hieron.  in  Bot. 
Jahrb.  Syst.  53:  432  (1915);  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  92  (1944);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  297 
(1964);  Benl  in  Acta  Bot.  Barcinon.  40:  73  (1991). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14540  (not  found); 

Lagoa  Amelia,  Exell  221  (BM!),  Rose  195  (P!);  Macambrara,  Exell  227 

(BM!);  Calvario, Monod  1 1823  (BM!);  Rio Contador,  1450 m,  Quintas  1356 

(BM!,  P!);  s.l.,  F.A.E.  (Moller)  62  (BM!,  LISU!,  P!),  Henriques  5/84  p.p. 

(K!). 

PRINCIPE.  S.l.,  Barter  1903  (K!). 

DISTRIBUTION.    W.  tropical  Africa. 

Elaphoglossum  salicifolium  (Willd.  ex  Kaulf.)  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  92  (1944);  Tardieu-Blot  in  Aubrev.,  FL 
Cameroun  3:  302  (1964);  Benl  in  Acta  Bot.  Barcinon.  40:  73 
(1991). 

Elaphoglossum  petiolatum  (Sw.)  Urb.,  Symb.  antill.  4:  61  (1903) 
subsp.  salicifolium  (Willd.  ex  Kaulf.)  Schelpe  in  Contr.  Bolus 
Herb.  1:  34(1969). 

SAO  TOME.  Sao  Joao  dosAngolares,  F.A.E.  (Quintas)  61  p.p.  (BM!);  Pico, 
Rose  315  (P!). 

DISTRIBUTION.     Tropical   Africa,  Madagascar,  Comoros,  Sey- 
chelles and  Mascarene  Is. 


Gleicheniaceae 

Dicranopteris  Bernh. 

Dicranopteris  linearis  (Burm.f.)  Underw.  in  Bull.  Torrey  Bot.  Club 
34:  250  (1907);  Benl  in  Acta  Bot.  Barcinon.  31:  23  (1978). 

Gleichenia  linearis  (Burm.f.)  C.B.  Clarke  in  Trans.  Linn.  Soc. 
London,  Bot.  1:  428  (1880);  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  93  (1944);  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  8 
(1956). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14310  (P);  Famosa, 
400  m,  Espirito  Santoll  (BM!,  COI!,  LISJC!);  between  Java  and  Bombaim, 
secondary  forest,  520  m,  Espirito  Santo  501 3  (LISC!,  LISJC!);  Macambrara 
to  Zampalma,  1050  m,  ground  fern, Exell  298  (BM!,  COI!);  Sao  Nicolau,  900 
m,  Figueiredo  &  Arriegas  52  (LISC!);  Jou,  Lains  e  Silva  331  (LISU!); 
between  Monte  Castro  and  Ponta  Figo,  secondary  forest,  500-1070  m,Lejoly 
95/35  (LISC!);  Pico,  summit,  Mann  (K!),  1800  m,  Moller  s.n.  (COI!); 
between  Calvario  and  Esta^ao  Sousa,  primary  forest,  1500  m,  Matos  &  Van 
Essche  7358  (LISC!);  Rio  Contador,  1325  m,  Moller  s.n.  (COI!);  Calvano, 
1 100  m,  Monod  11810  (BM!,  COI!);  Agua  Ize,  common,  Pinto  Basto  198 
(LISC!);  Nova  Moca,  750  m,  Quintas  1465  (BM!,  COI!,  K!);  between  Vale 
Carmo  and  Guaiaquil,  Rozeira  18  (PO!);  Tras-os-Montes,  s.c.  s.n.  (COI!); 
s.l.,  F.A.E.  (Moller)  1  (BM!,  COI!,  LISJC!,  LISU!). 

PRINCIPE.  Infante  D.  Henrique,  200  m,  common  ground  fern,  Exell  654 
(BM!,  COI!),  240  m,  Lains  e  Silva  467  (LISU!);  Oque  Nazare,  Newton  s.n. 
(BM!,  COI!);  Pico  Papagaio,  760  m,  Welwitsch  19  (BM!,  LISU!). 

DISTRIBUTION.     Pantropical. 
Grammitaceae 

Grammitis  Sw. 

Grammitis  ebenina  (Maxon)  Tardieu  in  Mem.  Inst.  Franc.  Afrique 
Noire  28:  21 1,  t.  42,  figs  1 1-12  (1953);  Schelpe  in  Contr.  Bolus 
Herb.  1:  3  (1969). 

Polypodium  ebeninum  Maxon  in  Bull.  Torrey  Bot.  Club  42:  224 
(1915);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  88  (1944). 

SAO  TOME.  Cabumbe,  800  m,  epiphyte,  Newton  s.n.  (P!). 
DISTRIBUTION.    Also  in  St.  Helena. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Grammitis  molleri  (Baker)  Schelpe  in  Bol.  Soc.  Brot.,  ser.  2,  40: 
162  (1966);  Schelpe  in  Contr.  Bolus  Herb.  1:  5  (1969). 

Polypodium  molleri  Baker  in  Henriq.  in  Bol.  Soc.  Brot.  4:  154,  t.  2, 
fig.  B  (1887);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  88  (1944). 

SAO  TOME.  Pico,  1900  m,  F.A.E.  (Moller)  5\  (BM!,  COI!,  LISJC!,  LISU!, 

P!);  s.l.  (probably  Pico),  Moller  64  (K-holotype). 

DISTRIBUTION.     Endemic. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Grammitis  nigrocincta  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique 
Noire  ser.  A,  21:  444  (1959);  Schelpe  in  Contr.  Bolus  Herb.  1:  3 
(1969). 

Grammitis  sp.;  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  80 
(1958). 

PRINCIPE.  Pico,  600  m,  Thorold  2074  (BM !),  Monod  1 2 1 62  (BM  !-holotype, 
COl\),  Rozeira  921  (PO). 


54 


E.  FIGUEIREDO 


DISTRIBUTION.     Also  in  Madagascar. 

Grammitis  tomensis  Schelpe  in  Contr.  Bolus  Herb.  1:  6  (1969). 

SAO  TOME.  Pico,  2000  m,  Quintas  1 1  (BM!-holotype,  COI!);  s.l.  (prob- 
ably Pico),  Newton  1 1  p.p.  (K). 

DISTRIBUTION.    Endemic. 

NOTES.  At  BM  there  is  another  collection  labelled  Quintas  1 1 
which  consists  of  Pleris  tripartita. 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Xiphopteris  Kaulf. 

Xiphopteris  cultrata  (Willd.)  Schelpe  in  Bol.  Soc.  Brot,  ser.  2,  41: 
217  (1967);  Schelpe  in  Contr.  Bolus  Herb.  1:  9  (1969);  Benl  in 
Acta  Bot.  Barcinon.  33:  22  (1982). 

Polypodium  cultratum  Willd.,  Sp.  pi.  5:  187  (1810)  van  elasticum 
(Bory  ex  Willd.)  Baker  in  Hook.  &  Baker,  Syn.fil.:  327  (1867); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  87  (1944). 

SAO  TOME.  Macambrara  to  Zampalma,  primary  forest,  800-1 050  m,  Exell 
441  (BM!);  between  Bom  Sucesso  and  Lagoa Amelia,  epiphyte,  Matos  7585 
(LISC!);  Lagoa  Amelia,  primary  forest,  1400-1420  m,  epiphyte,  Matos  & 
Van  Essche  7412  (LISC!). 

DISTRIBUTION.     W.  and  Central  Africa  and  tropical  America. 

Xiphopteris  oosora  (Baker)  Alston  in  Bol.  Soc.  Brot.,  ser.  2,  30:  26 
(1956);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  325  (1964); 
Schelpe  in  Contr.  Bolus  Herb.  1:  12  (1969);  Benl  in  Acta  Bot. 
Barcinon.  33:  21  (1982). 

Polypodium  oosorum  Baker  in  Henriq.  in  Bol.  Soc.  Brot.  4:  154,  t.  2, 
fig.  A  (1887);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  89  (1944). 

SAO  TOME.  Pico,  1950  m,  Moller  s.n.  (BM!,  COI!);  Pico,  1950  m,  F.A.E. 
(Moller)  52  (BM!,  K?,  LISU!,  P!);  s.l.  (probably  Pico),  Moller  1337  (K- 
holotype). 

DISTRIBUTION.     W.  and  E.  tropical  Africa,  Madagascar. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Xiphopteris  villosissima  (Hook.)  Alston  in  Bol.  Soc.  Brot.,  ser.  2, 
30:  27  (1956);  Schelpe  in  Contr.  Bolus  Herb.  1:  8  (1969);  Benl  in 
Acta  Bot.  Barcinon.  33:  23  (1982). 

Polypodium  villosissimum  Hook.,  Sp.  fil.  4:  197  (1862);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  90  (1944). 

SAO  TOME.  S.l.,  Mann  s.n.  (K),  Newton  (not  found  at  COI). 
DISTRIBUTION.    W.,  E.,  and  SE  tropical  Africa. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Hemionitidaceae 

Coniogramme  Fee 

Coniogramme  africana  Hieron.  in  Hedwigia  57:  293  (1916);  Als- 
ton in  Exell,  Cat.  Vase.  PL  S.  Tome:  81  (1944);  Tardieu-Blot  in 
Aubrev.,  FL  Cameroun  3:  132  (1964);  Benl  in  Acta  Bot.  Barc- 
inon. 32:  26  (1980). 

SAO  TOME.  Macambrara  to  Zampalma,  900  m,  ground  fern,  Exell  292 


(BM!);  Macambrara,  primary  forest,  1050-1200  m,  ground  fern,  Exell  421 
(BM!,  COI!);  Pico,  1800  m,  Moller  s.n.  (COI!);  W.  of  Pico,  1225  m,  Monod 
1 1994  (BM!);  NW  of  Pico,  1225  m,  Monod  1 1995  (BM!);  Lagoa  Amelia, 
Rozeira  1 182  (PO!);  s.l.,  F.A.E.  (Moller)  58  (BM!,  LISU!),  Mann  s.n.  (K!), 
Moller  s.n.  (COI!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Pityrogramma  Link 

Pityrogramma  calomelanos  (L.)  Link,  Handbuch  3:  20  (1833) 
'calomelas';  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  81  (1944); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  442 
(1959). 

van  calomelanos;  Benl  in  Acta  Bot.  Barcinon.  32:  30  (1980). 

SAO  TOME.  Between  Bom  Sucesso  and  Macambrara,  Matos 761 1  (LISC!); 
Agua  Ize,  Pinto  Basto  215  (LISC!);  near  Bombaim,  Rozeira  499  (PO!). 
PRINCIPE.  Santo  Antonio,  Monod  12064  (BM!,  COI!);  s.l.,  Barter  1912 
(K-not  found). 

DISTRIBUTION.     A  weed  introduced  from  tropical  America. 


Hymenophyllaceae 

Hymenophyllum  Sm. 

Hymenophyllum  capillare  Desv.  in  Mem.  Soc.  Linn.  Paris  6:  333 
(1827);  Alston  in  Exell,  Cat  Vase.  PL  S.  Tome:  59  (1944); 
Tardieu-Blot  inAubrev.,  Fl.  Cameroun  3:  75  (1964);  Benl  inActa 
Bot.  Barcinon.  32:  7  (1980). 

SAO  TOME.  Pico,  1 500-2000  m,  Moller  s.n.  (BM!);W.  of  Pico,  1225-1430 
m,  Monod  11913  (BM!,  COI!);  Calvario,  Rose  155  (P!);  s.l.,  F.A.E.  (Moller 
&  Quintas)  5  p.p.  (K  also  Moller  4!,  P!). 

DISTRIBUTION.  Widespread  in  tropical  Africa,  Mascarene  Is.  and 
Tristan  da  Cunha. 

Hymenophyllum  hirsutum  (L.)  Sw.  in  J.  Bot.  (Schrader)  1800(2): 
99  (1801),  emend.  C.V.  Morton  in  Contr.  U.S.  Nad.  Herb.  29: 
155,  173  (1947);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  76 
(1964);  Benl  in  Acta  Bot.  Barcinon.  32:  9  (1980). 

Hymenophyllum  ciliatum  Sw.  in  J.  Bot.  (Schrader)  1800(2):  100 
(1801)  var.  boryanum  (Willd.)  Mett.  ex  Kuhn,  Filic.  afr.:  39 
(1868);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  59  (1944). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14292  p.p.  (P!); 
Calvario,  1500  m,  ground  fern,  Exell  327  (BM!,  COI!),  1560  m,  Monod 
1 1835  (BM!,  COI!);  Lagoa  Amelia,  primary  forest,  1400-1420  m,  epiphyte, 
Matos  &  Van  Essche  7414  (LISC!);  Pico,  1500-2000  m,  Moller  s.n.  (BM!, 
COI!);  Bom  Sucesso,  1 150  m,  Moller  &  Quintas  s.n.  (COI!);  Binda,  Newton 
(P-specimen  is  missing  from  sheet);  Sao  Nicolau,  900  m,  Quintas  s.n. 
(LISU!);  s.l.,  F.A.E.  (Moller  &  Quintas)  5  p.p.  (BM!,  COI!,  K!,  LISJC!, 
LISU!,  P!),  Quintas  s.n.  (K!). 

DISTRIBUTION.     Pantropical  and  subtropical. 

Hymenophyllum  polyanthos  Sw.  in  J.  Bot.  (Schrader)  1800(2): 
102(1801). 

var.  kuhnii  (C.  Chr.)  Schelpe  in  Bol.  Soc.  Brot.,  ser.  2,  40:  156 
(1966);  Benl  in  Acta  Bot.  Barcinon.  32:  6  (1980). 

Hymenophyllum  kuhnii  C.  Chr.,  Index filic.:  363  (1905);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  59  (1944);  Alston  in  Exell  &  Rozeira 
in  Conf.  Int.  Afr.  Oc.  3:  78  (1958);  Alston  in  Exell  in  Bull.  Inst. 
Franc.  Afrique  Noire  ser.  A,  21:  441  (1959);  Tardieu-Blot  in 
Aubrev.,  FL  Cameroun  3:  72  (1964). 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


55 


SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14292  p.p.  (P!), 
14292ter(BM!,  P!),  14551  (P!);  SW  region,  Chevalier  14588  (P!);  Calvario, 
1 500  m,  ground  fern,  Exell  323  (BM !);  between  Lagoa  Amelia  and  Change, 
primary  forest,  1 250  m,  epiphyte,  Matosl534  (LISC!);  Pico,  1950  m,  Moller 
s.n.  (COI!),  Rose  316  (P!),  319  (P!),  Rozeira  967  (PO!);  Tras-os-Montes, 
800- 1 200  m,  Quintas  1315  (BM !,  COI-not  found);  s.l.,  F.A.E.  (Moller)  4  p.p. 
(BM!,  COI!,  K  also  Moller  3!,  LISJC!,  LISU!,  P!). 

PRINCIPE.  W.  of  Pico,  500-600  m,Monod\2\  13  (BM!);  Pico,  500-600  m, 
Monod  121 16  (BM!),  800-900  m,  Rozeira  973  (BM-not  found,  PO),  600  m, 
epiphyte,  Thorold  2073 A  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Hymenophyllum  polyanthos  Sw.  in  J.  Bot.  (Schroder)  1800(2): 
102(1801). 

var.  mildbraedii  (Brause  ex  Brause  &  Hieron.)  Schelpe  in  Garcia  de 
Orta,  ser.  Bot.  3:  54(1976). 

Hymenophyllum  mildbraedii  (Brause  ex  Brause  &  Hieron.)  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  60  (1944);  Alston  in  Exell  in  Bull. 
Inst.  Franc.  Afrique  Noire  ser.  A,  21:  441  (1959). 

PRINCIPE.  Pico,  Rozeira  965  (K!),  600  m,  epiphyte,  Thorold  2073  p.p. 
(BM!),  900  m,  Thorold  2084  p.p.  (BM!). 

DISTRIBUTION.     Angola  and  Annobon. 

Hymenophyllum  splendidum  Bosch  in  Ned.  Kruidk.  Arch.  5(3): 
192  (1863);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  60  (1944); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  441 
(1959);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  76  (1964); 
Benl  in  Acta  Bot.  Barcinon.  32:  8  (1980). 

SAO  TOME.  Macambrara,  primary  forest,  800-1050  m,  Exell  439  (BM!, 
COI!);  s.l.,  F.A.E.  (Moller  &  Quintas)  5  p.p.  (P!). 

PRINCIPE.  Pico,Rozeira  966  (K!),  600  m,  epiphyte,  Thorold  2013B  (BM!), 
900  m,  epiphyte,  Thorold  2084A  (BM!). 

DISTRIBUTION.     Tropical  Africa,  extending  to  Tanzania. 

Trichomanes  L. 

NOTE.  Recent  work  (Dubuisson,  1996)  supports  the  division  by 
earlier  workers  of  the  genus  Trichomanes  s.l.  into  several  genera. 
Although  the  genus  is  treated  here  in  its  sensu  lato,  the  names  of  the 
taxa  in  the  segregate  genera  are  also  provided,  when  available. 

Trichomanes  borbonicum  Bosch  in  Ned.  Kruidk.  Arch.  5(2):  158 
(as  '185')  (1861);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  57 
(1944);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  88  (1964); 
Benl  in  Acta  Bot.  Barcinon.  32:  22  (1980). 

[Vandenboschia  borbonica  (Bosch)  G.  Kunkel  in  Nova  Hedwigia 

6:213(1963)] 

SAO  TOME.  Pico,  1 500-2000 m,Moller s.n.  (BM!,COI!);Tras-os-Montes, 
1200  m,  Quintas  1269A  (BM!,  COI!). 

DISTRIBUTION.  Widespread  in  tropical  Africa,  SE  and  S.  Africa, 
Madagascar  and  Reunion. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Trichomanes  crispiforme  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome: 
57  (1944);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  78 
(1958);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  90  (1964). 

SAO  TOME.  Cabumbe,  Newton  (P?). 

PRINCIPE.  Oque  Pipi,  secondary  forest,  300  m,  epiphyte,  Exell  501  (BM!, 
COI!);  W.  of  Pico,  500-600  m,  Monod  12123  (BM!),  700-800  m,  Monod 
12143  (BM!);  Pico,  Rozeira  905  (BM-not  found,  PO);  s.l.,  Barter  1918  (K- 
not  found). 


DISTRIBUTION.    W.  and  Central  tropical  Africa. 

Trichomanes  digitatum  Sw.,  Syn.fil:  370  (1806);  Alston  in  Exell 
in  Bull  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  441  (1959). 

[Microtrichomanes  digitatum  (Sw.)  Copel.  in  Philipp.  J.  Sci.  67: 
36(1938)] 

SAO  TOME.  Calvario,  1400  m,  Monod  1 1855  (BM!,  COI!). 

DISTRIBUTION.  Also  in  Madagascar,  Mauritius,  Reunion,  Malaya 
and  Borneo. 

Trichomanes  erosum  Willd.,  Sp.  pi.  5:  501  (1810);  Alston  in  Exell, 
Cat.  Vase.  PI.  S.  Tome:  58  (1944);  Benl  in  Acta  Bot.  Barcinon.  32: 
16(1980). 

[Microgonium  erosum  (Willd.)  C.  Presl,  Gefassbiindel  Farm:  21 

(1847)] 

Trichomanes  chamaedrys  Taton  in  Bull.  Soc.  Roy.  Bot.  Belgique  78: 
29,  pi.  2,  figs  K-L  (1946);  Alston  in  Exell  &  Rozeira  in  Conf.  Int. 
Afr.  Oc.  3:78(1958). 

SAO  TOME.  Porto  Alegre,  Chevalier  14178  (P);  Pico,  1900  m,  Moller  s.n. 
(COI!);  Sao  Joao  dosAngolares,  &Om,Newton  s.n.  (BM!,  COI!);  Macambrara, 
1200  m,  epiphyte,  Quintas  1269B  (BM!,  COI!);  Bom  Sucesso,  Quintas  s.n. 
(BM!,  COI!);  Ribeira  Peixe,  Rozeira  205  (BM-not  found,  PO);  s.l.,  F.A.E. 
(Moller  &  Quintas)  6  (K!,  LISU!),  Quintas  3  (COI!). 
PRINCIPE.  S.l.,  Barter  1919  (K!). 

DISTRIBUTION.    Tropical  Africa. 

Trichomanes  mannii  Hook,  ex  Hook.  &  Baker,  Syn.fil.:  75  (1867); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  58  (1944);  Benl  in  Acta 
Bot.  Barcinon.  32:  17  (1980). 

[Gonocormus  mannii  (Hook,  ex  Hook.  &  Baker)  G.  Kunkel  in 
Nova  Hedwigia  6:  212  (1963)] 

SAO  TOME.  Lagoa  Amelia,  Exell  203  (BM!,  COI!);  between  Sao  Pedro  and 
Lagoa  Amelia,  1050  m,  Monod  11706  (BM!);  Tras-os-Montes,  850  m, 
Quintas  1269  (BM!,  COI!). 

DISTRIBUTION.  W.,  Central  and  E.  Africa,  Madagascar  and 
Comoros. 

Trichomanes  pyxidiferum  L.,  Sp.  pi.  2:  1098  (1753). 

[Vandenboschia  pyxidifera  (L.)  Copel.  in  Philipp.  J.  Sci.  67:  53 

(1938)] 

var.  melanotrichum  (Schltdl.)  Schelpe  in/  5.  African  Bot.  30:  181 
(1964);  Benl  in  Acta  Bot.  Barcinon.  32:  24  (1980). 

Trichomanes  melanotrichum  Sch\td\.,Adumbr.pL:  56  (1832); Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  58  (1944). 

SAO  TOME.  Sao  Nicolau,  900  m,  F.A.E.  (Moller)  1  (BM!,  K!,  LISU!),  950 
m,  Moller  s.n.  (COI!);  Lagoa  Amelia,  Rozeira  1084  (K!);  s.l.  (probably  Sao 
Nicolau),  F.A.E.  (Moller)  4  p.p.  (P!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical  Africa. 

Trichomanes  radicans  Sw.  in  J.  Bot.  (Schroder)  1800(2):  97 
(1801);  Benl  in  Acta  Bot.  Barcinon.  32:  14  (1980). 

[Vandenboschia  radicans  (Sw.)  Copel.  in  Philipp.  J.  Sci.  67:  54 

(1938)] 

Trichomanes  giganteum  sensu  auct.  non  Bory  ex  Willd.,  Sp.  pi.  5: 
514  (1810);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  58  (1944). 

SAO  TOME.  Macambrara,  primary  forest,  1050- 1200m,  ground  fern,  Exell 


56 


E.  FIGUEIREDO 


185  (BM!,  COI!),  Moller  s.n.  (COI!);  near  Sao  Tome,  20  m,  Mailer  s.n. 
(COI!);  Pico,  1800  m,  Moller  s.n.  (COI!),  Newton  s.n.  (BM!,  COI!);  Sao 
Pedro,  Quintas  s.n.  (COI!);  s.l.,  F.A.E.  (Moller  &  Quintas)  8  (BM-not  found, 
COI!,  LISU!,  P!),  Mann  s.n.  (K!). 

DISTRIBUTION.  W.,  Central  and  E.  Africa,  Madagascar,  Comoros 
and  Reunion. 

Trichomanes  rigidum  Sw.,  Prodr.:  137  (1788). 

[Selenodesmium  rigidum  (Sw.)  Copel.  in  Philipp.  J.  Sci.  67:  81 
(1938)] 

Trichomanes  cupressoides  Desv.  in  Mem.  Soc.  Linn.  Paris  6(2):  330 
(1827);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  57  (1944);  Alston 
in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  78  (1958). 

SAO  TOME.  Maria  Femandes,  secondary  forest,  275  m,  Carvdho  2 
(LISC!);  Sao  Miguel,  Monte  Verde,  secondary  forest,  10-150  m,  Lejoly  95/ 
96  (LISC!). 

PRINCIPE.  Oque  Nazare,  Newton  s.n.  (BM!,  COI!);  Pico,  above  Neves 
Ferreira,  Rozeira  650  (BM-not  found,  PO),  683  (BM-not  found,  PO);  Pico, 
Rozeira  907  (BM-not  found,  PO). 

DISTRIBUTION.  Widespread  in  tropical  and  subtropical  Africa, 
tropical  America. 

Lomariopsidaceae 

Bolbitis  Schott 

Bolbitis  acrostichoides  (Afzel.  ex  Sw.)  Ching  in  C.  Chr.,  Index filic. 
Suppl.  3:  47  (1934);  Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  68 
(1944);  Hennipman,  Monogr.  fern  genus  Bolbitis:  149  (1977); 
Benl  in  Acta  Bot.  Barcinon.  40:  61  (1991). 

SAO  TOME.  Near  Sao  Tome,  F.A.E.  (Moller)  67  (K  also  Moller  41!, 
LISU!);  Monte  Cafe,  800  m,  Moller  s.n.  (COI!);  Welwitsch  63b  (BM!, 
LISU!);  Sao  Pedro,  1 100  m,  Quintas  s.n.  (COI!). 

DISTRIBUTION.     Tropical  Africa,  Madagascar. 

NOTES.  Moller  63  (K),  cited  by  Hennipman  (1977),  is  probably  a 
mistake  for  F.A.E.  (Moller)  67. 

This  taxon  has  not  been  recorded  in  the  islands  since  the  nine- 
teenth century. 

Bolbitis  auriculata  (Lam.)  Ching  in  C.  Chr.,  Index  filic.  Suppl.  3:  47 
(1934)  et Alston  in/  Bot.  72,  Suppl.  2:  3  (1934);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  68  (1944);  Hennipman,  Monogr.  fern  genus 
Bolbitis:  136  (1977);  Benl  in  Acta  Bot.  Barcinon.  40:  59  (1991). 

SAO  TOME.  Boa  Entrada,  Chevalier  14335  (P!);  Monte  Cafe,  Moller  s.n. 
(COI!),  800  m,  F.A.E.  (Moller)  66  (BM!,  LISU!,  P!);  Diogo  Vaz,  150  m, 
Quintas  s.n.  (BM!,  COI!,  K!,  LISJC!,  P!);  above  Monte  Cafe,  Welwitsch  58 
(BM!,  LISU!). 

PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  572  (BM!);  s.l.,  Barter  1899  (K-not  found). 

DISTRIBUTION.     Tropical  Africa,  Madagascar,  Mascarenes. 

Bolbitis  fluviatilis  (Hook.)  Ching  in  C.  Chr.,  Index  filic.,  Suppl.  3: 
48  (1934);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  69  (1944); 
Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  78  (1958); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  447 
(1959);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  316  (1964); 
Hennipman,  Monogr.  fern  genus  Bolbitis:  142  (1977);  Benl  in 
Acta  Bot.  Barcinon.  40:  60  (1991). 

SAO  TOME.  Sao  Joao  dos  Angolares,  200  m,  F.A.E.  (Quintas)  68  (BM!, 
COI!,  LISU!);  Rio  Abade,  1300  m,  Quintas  s.n.  (COI!). 


PRINCIPE.  Pico,  700-800  m,  Monod  12137  (BM!);  between  Lapa  and 
Pico,  Rozeira  893  (PO);  s.l.,  Barter  1897  (K),  1897bis  (K!). 

DISTRIBUTION.     W.  tropical  Africa. 

Lomariopsis  Fee 

Lomariopsis  guineensis  (Underw.)  Alston  in/  Bot.  72,  Suppl.  2:  5 
(1934);  Alston  in  Exell,  Cat  Vase.  PL  S.  Tome:  81  (1944); 
Tardieu-Blot  in  Aubrev.,  FL  Cameroun  3:  309  (1964);  Benl  in 
Acta  Bot.  Barcinon.  40:  64  (1991). 

SAO  TOME.  Ilheu  das  Rolas,  1 20  m,  Quintas  1164  (BM!,  COI  also  labelled 
no.  35  from  Monte  Cafe!),  F.A.E.  (Quintas)  65  p.p.  (BM!,  COI!,  K!-also 
Quintas  35);  Monte  Cafe,  Welwitsch  68  (BM!,  LISU!). 
PRINCIPE.  Esperanc,a,  600  m,  Rozeira  892  (K!). 

DISTRIBUTION.     W.  tropical  Africa  to  Angola. 

Lomariopsis  hederacea  Alston  in  J.  Bot.  72,  Suppl.  2:  5  (1934); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  81  (1944);  Benl  in  Acta 
Bot.  Barcinon.  40:  70  (1991). 

SAO  TOME.  Macambrara,  1050-1200  m,  epiphyte,  Exell  189  (BM!), 
primary  forest,  1050-1200  m,  ground  fern,  Exell  315  (BM!,  COI!);  Rio 
Contador,  1 300  m,  Moller  s.n.  (COI!);  s.l.,  F.A.E.  (Moller  &  Quintas)  65  p.p. 
(BM!,  COI!),  Moller  s.n.  (COI!). 

DISTRIBUTION.     W.  tropical  Africa  to  Angola,  Central  Africa. 

NOTE.  Barter  s.n.  (K)  from  Principe  was  cited  by  Alston  in  the 
protologue  of  the  species  (paratype).  Barter  1 899  (K),  also  from 
Principe,  was  cited  by  Alston  in  1944.  The  same  collection  (Barter 
1 899)  was  cited  by  Alston  ( 1 944)  under  Bolbitis  auriculata.  Neither 
of  these  collections  was  found  under  Lomariopsis  hederacea  at  K. 
Barter  1899  (K)  is  Bolbitis  auriculata.  It  is  possible  that  both 
citations  refer  to  Barter  1902  (K),  a  specimen  of  L.  warneckei. 

Lomariopsis  warneckei  (Hieron.)  Alston  in  J.  Bot.  72,  Suppl.  2:  6 
(1934);  Benl  in  Acta  Bot.  Barcinon.  40:  70  (1991). 

Lomariopsis  nigrescens  Holttum  in  Kew  Bull.  1939:  627,  figs  13  & 

14  (1940);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser. 

A,  21:  447  (1959). 
Lomariopsis  hederacea  sensu  auct.  non  Alston  in  J.  Bot.  72,  Suppl. 

2:  5  (1934);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  79 

(1958). 

PRINCIPE.  Pico,  above  Neves  Ferreira,  epiphyte,  Rozeira  663  (BM-not 
found,  PO!);  Pico,  epiphyte,  77zoroW2086  (BM!);  s.l.,  Barter  1902  p.p.  (K!). 

DISTRIBUTION.    E.  and  SE  Africa  and  Bioko. 

NOTE.  Barter  1 902  (K)  consists  of  two  herbarium  sheets.  The  one 
cited  above  is  apparently  a  young  plant  (as  noted  by  Holttum  on  the 
herbarium  sheet),  with  pinnae  smaller  than  in  typical  Lomariopsis 
warneckei.  The  other  sheet  contains  a  larger  specimen,  referred  to  L. 
hederacea  by  Holttum,  but  with  much  larger  and  separated  pinnae. 
Although  the  distinction  between  this  species  and  the  previous 
seems  to  be  clear  in  material  from  the  continent,  some  specimens 
from  Sao  Tome  and  Principe  show  intermediate  features. 


Loxogrammaceae 

Loxogramme  (Blume)  C.  Presl 

Loxogramme  lanceolata  (Sw.)  C.  Presl,  Tent,  pterid.:  215,  t.  9,  fig. 
8  (1836);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  90  (1944);  Benl 
in  Acta  Bot.  Barcinon.  33:  24  (1982). 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 

SAO  TOME.  Macambrara,  primary  forest,  1200  m,  epiphyte,  Exell  230 
(BM!);  between  Estacao  Sousa  and  Pico,  1700  m,  Matos  7551  (LISC!); 
between  Bom  Sucesso  and  Lagoa  Amelia,  primary  forest,  1350-1420  m, 
Matos  &  Van  Essche  7290  (LISC!);  Saudade,780  m,  Moller  s.n.  (COI!);  NW 
of  Pico,  1600-1800  m,  Monod  12239  (BM!);  Tras-os-Montes,  Quintas  s.n. 
(BM!,  COI!);  Novo  Brasil,  Monte  Mario,  Rozeira  4  (PO!);  Cruzeiro,  300  m, 
77wrt»W2069(BM!). 
PRINCIPE.  S.I.,  Barter  1898  (K!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical  Africa. 
Lycopodiaceae 

Huperzia  Bernh. 

Huperzia  mildbraedii  (Herter)  Pic.  Serm.  in  Webbia  23:  163 
(1968);  Benl  in  Acta  Bot.  Barcinon.  31:  10  (1978). 

Lycopodium  mildbraedii  Herter  in  Hedwigia  49:  90  (1909);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  95  (1944). 

SAO  TOME.  Lagoa  Amelia,  epiphyte,  Espirito  Santo  5032  (LISC!, 
LISJC!),  primary  forest,  1400-1420  m,  epiphyte,  Matos  &  Van  Essche 
7409  (LISC!),  1350  m,  epiphyte,  Moller  s.n.  (COI!),  1400  m,  Monod  1 1734 
(BM!);  Calvario,  1500  m,  Exell  317  (BM!,  COI!),  1550  m,  Monod  11832 
(BM!);  Macambrara,  primary  forest,  1050-1200  m,  epiphyte,  Exell  427 
(BM!,  COI!);  Pico,  1700-1900  m,  F.A.E.  (Moller)  75  (BM!,  K!,  LISU!,  P!), 
1850  m,  epiphyte,  Moller  s.n.  (COI!),  Moller  s.n.  p.p.  (COI!),  Rose  323 
(P!);  Bom  Sucesso,  1250  m,  epiphyte,  Moller  s.n.  (COI!);  Tras-os-Montes, 
1100  m,  Quintas  1321  p.p.  (COI!);  s.l.,  F.A.E.  (Moller)  74  p.p.  (BM!,  COI!, 
LISU!),  1 100-1400  m,  Moller  76  (K!). 

PRINCIPE.  Pico  Papagaio,  350  m,  Exell  702  (BM!,  COI!);  W.  of  Pico, 
below  250  m,  Monod  12097  (BM!);  Pico,  common  up  to  900  m,  Thorold 
2083  (BM!). 

DISTRIBUTION.    W.  tropical  Africa. 

Huperzia  ophioglossoides  (Lam.)  Rothm.  in  Feddes  Repert.  Spec. 
Nov.  Regni  Veg.  54:  62  (1944). 

SAO  TOME.  Pico,  primary  forest,  1900-2020  m,  Lejoly  94/538  (LISC!), 
1850m,  epiphyte,  Moller  s.n.  p.p.  (COll),  Rose  328  (P!);  s.l.  (probably  Pico), 
F.A.E.  (Moller)  74  p.p.  (LISU!). 

DISTRIBUTION.  Tropical,  SE  and  S.  Africa,  Madagascar  and 
Mascarene  Is. 

Huperzia  phlegmaria  (L.)  Rothm.  in  Feddes  Repert.  Spec.  Nov. 
Regni  Veg.  54:  62  (1944);  Benl  in  Acta  Bot.  Barcinon.  31:  9 

(1978). 

Lycopodium  phlegmaria  L.,  Sp.  pi.  2:  1101  (1753);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  96  (1944). 

SAO  TOME.  Pico,  Chevalier  13660  (P);  Lagoa  Amelia,  Chevalier  14307 
(P!);  Ribeira  Peixe,  Vila  Irene,  epiphyte,  Espirito  Santo  4601  (LISC!);  Sao 
Joao  dos  Angolares,  F.A.E.  (Quintas)  76  (BM!,  K!,  LISU!,  P!),  Quintas  s.n. 
(COI!);  Perseveran?a,  Seabra  183b  (LISU!). 

PRINCIPE.  Between  Terreiro  Velho  and  Infante  D.  Henrique,  plantation, 
100  m,  Exell  587  (BM!,  COI!);  Porto  Real,  Rose  483  (P!). 

DISTRIBUTION.     Palaeotropical. 

Huperzia  verticillata  (L.f.)  Trevis.  in  Atti  Soc.  Ital.  Sci.  Nat.  17: 
248  (1874);  Benl  in  Acta  Bot.  Barcinon.  31:  8  (1978). 

Lycopodium  verticillatum  L.f.,  Suppl.  pi.:  448  (1782);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  96  (1944);  Tardieu-Blot  in  Aubrev., 
Fl.  Cameroun3:  12(1964). 

SAO  TOME.  Bom  Sucesso,  1200  m,  F.A.E.  (Moller)  73  (BM!,  COI!,  K!, 
LISU!,  P!);  Macambrara,  1 100  m,  Moller  s.n.  (COI!). 


57 

DISTRIBUTION.  Tropical,  SE  and  S.  Africa,  Comoros  and  Reunion, 
tropical  America,  and  Polynesia. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Huperzia  warneckei  (Herter  ex  Nessel)  Pic.  Serm.  in  Webbia  23: 
163(1968). 

Lycopodium  warneckei  (Herter  ex  Nessel)  Alston  in  Bol.  Soc.  Brot., 
ser.  2,  30:  20  (1956);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr. 
Oc.  3:81  (1958). 

SAO  TOME.  Between  Sao  Miguel  and  Zagaia,  primary  forest,  170-400  m, 
Matos  &  Van  Essche  7247  (LISC!);  Tras-os-Montes,  1 100  m,  Quintas  1321 
p.p.  (COI!);  Ribeira  Peixe,  between  Vila  Jose  and  Ermelinda,  Rozeira  288 
(PO!). 

DISTRIBUTION.    W.  tropical  Africa. 

Lycopodiella  Holub 

Lycopodiellacernua(L.) Pic.  Serm.  inWebbia23:  166 (1968); Benl 
in  Acta  Bot.  Barcinon.  31:  1 1  (1978). 

Lycopodium  cernuum  L.,  Sp.  pi.  2:  1103  (1753);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  95  (1944);  Sobrinho  in  Portugaliae  Acta 
BioL,  Ser.  B,  Sist.  3:  393  (1952). 

SAO  TOME.  Porto  Alegre,  Chevalier  14196  (P);  Sao  Nicolau,  1000  m, 
Espirito  Santo4448  (LISC!,  LISJC!),900m,Mo/ters.n.  (COI!);  Macambrara, 
primary  forest,  1200  m,  Exell  94  (BM!);  Santa  Maria,  1300  m,  Exell  196 
(BM!,COI!),  1380m,  Moller  s.n.  (COI!);  Lagoa  Amelia,  1 300  m,  Exell  206 
(BM!,  COI!),  crater  swamp,  1300m,  Exell  209  (BM!),  1412  m,  Matos  7504 
(LISC!);  Tras-os-Montes,  Gama  s.n.  (COI!);  road  to  Monte  Mario,  Lains  e 
Silva279  (LISU!);  Pico,  primary  forest,  1900-200  m,L<yo/y94/546  (LISC!); 
Bom  Sucesso,  secondary  forest,  1 150  m,  Matos  &  Van  Essche  7459  (LISC!); 
Saudade,  950  m,  Moller  s.n.  (COI!);  above  Monte  Cafe,  1050  m,  Monod 
11679  (BM!);  between  Cascata  and  Morro  Vilela,  700-1000  m,  Monod 
12202  (BM!);  Santelmo,  Oliveira  &  Noronha  6-B  (LISU!);  Monte  Cafe,  in 
forest  and  plantations,  Oliveira  &  Noronha  s.n.  (LISU!);  Tras-os-Montes, 
between  Calvario  and  main  house,  Rozeira  527  (PO!);  s.l.,  Campos  16 
(COI!),  F.A.E.  (Moller)  77  (BM!,  COI!,  K!,  LISU!). 
PRINCIPE.  Oque  Pipi,  secondary  forest,  300  m,  Exell  551  (BM!,  COI!); 
Oque  Caspar,  Newton  s.n.  (COI!);  Ribeira  Jambere,  Rozeira  593  (PO!);  Pico 
Papagaio,  900  m,  Welwitsch  10  (BM!,  LISU!). 

DISTRIBUTION.    Cosmopolitan. 

Lycopodium  L. 

Lycopodium  clavatum  L.,  Sp.  pi.  2:  1101  (1753);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  95  (1944);  Tardieu-Blpt  in  Aubrev.,  Fl. 
Cameroun  3:  16  (1964);  Benl  in  Acta  Bot.  Barcinon.  31:  10 

(1978). 

SAO  TOME.  Pico,  2000  m,  Chevalier  13676  (P!),  2000  m.  F.A.E.  (Moller) 
78  (BM!,  COI!,  K!,  LISU!,  P!),  1850  m,  Monod  1 1919  (BM!,  COI!),  Rose 

320  (P!). 

DISTRIBUTION.    Cosmopolitan. 
Marattiaceae 

Marattia  Sw. 

Marattia  fraxinea  Sm.,  PL  icon.  ined.  2:  t.  48  (1790);  Alston  in 
Exell,  Cat  Vase.  PL  S.  Tome:  94  (1944);  Benl  in  Acta  Bot. 
Barcinon.  32:  3  (1980);  Pichi  Sermolli  in  Webbia  23:  332  (1969). 

Marattia  robusta  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  8 
(1956). 


58 

SAO  TOME.  Monte  Cafe,  primary  forest,  1 200  m,  Espirito Santo  191  (BM!- 
holotype  of  Marattia  robusta,  COI!,  LISJC!),  760  m,  Welwitsch  70  (BM!, 
LISU!);  Ribeira  Peixe,  S.  of  Cabumbe,  secondary  forest,  Espirito  Sanfo3899 
(COI!,  LISJC!);  Macambrara,  primary  forest,  1050-1200  m,Exell  129  (BM!, 
COI!),  1250  m,  Moller  s.n.  (COI!);  Lagoa  Amelia,  1200-1400  m,  very 
common,  Lains  e  Silva  88  (LISU!),  88a  (LISU!),  1350-1380  m,  Monod 
\  1771  (BM!,  COI!),  Rozeira  1091  (PO!);  Sao  Miguel,  Monte  Verde,  second- 
ary forest,  10-150  m,  Lejoly  95/88  (LISC!);  between  Sao  Miguel  and  Zagaia, 
primary  forest,  170-400  m,  Matos  &  Van  Essche  7239  (LISC!);  Bom 
Sucesso,  1250  m,  Moller  s.n.  (COI!);  between  Sao  Pedro  and  Lagoa  Amelia, 
1050  m,  Monod  11717  (BM!);  Sao  Joao  dos  Angolares,  150  m,  Quintas  s.n. 
(COI!);  Porto  Alegre,  near  Cao  Pequeno,  Rozeira  5 1  (PO!);  between  Lagoa 
Amelia  and  Calvario,  Rozeira  1171  (PO!);  Dona  Augusta,  Rozeira  1179 
(PO);  s.l.,  EA.E.  (Moller  &  Quintas)  71  (BM!,  COI!,  LISU!). 
PRINCIPE.  S.I.,  Barter  1916  (K!). 

DISTRIBUTION.     Palaeotropical. 

NOTE.  Pichi  Sermolli  ( 1 969)  considered  that  Marattia  robusta  was 
a  strong  form  of  M.  fraxinea  and  that  the  plants  from  Sao  Tome 
might  be  recognized  at  infraspecific  level.  The  collection  from 
Principe  is  from  a  smaller  plant  (M.  fraxinea  s.s.). 


Nephrolepidaceae 

Arthropteris  J.  Sm. 

Arthropteris monocarpa  (Cordem.) C.  Chr.,  Cat.  pi.  mad.,  pter.:  32 
(1932);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  69  (1944);  Alston 
in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  445  (1959); 
Benl  in  Acta  Bot.  Barcinon.  33:  35  (1982). 

SAO  TOME:.  Pico,  Campos  77  (COI!),  Chevalier  s.n.  (P!),  primary  forest, 
1987  m,  Matos  &  Van  Essche  7385  (LISC!);  Macambrara,  virgin  forest, 
1050-1200  m,  Exell  121  (BM!),  1200-1350  m,  122a  (BM!);  Lagoa  Amelia, 
crater  swamp,  1300  m,  ground  fern,  Exell  210  (BM!),  1430  m,  epiphyte, 
Matos  7494  (LISC!);  between  Macambrara  and  Sao  Nicolau,  secondary 
woodland,  1050  m,  Exell  403  (BM!,  COI!);  Tras-os-Montes,  Gama  s.n. 
(COI!);  between  Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain 
rainforest,  1200-1800  m,  Lejoly  95/23  (LISC!);  Calvario  and  surroundings, 
primary  forest,  1600-1620  m,  epiphyte,  Matos  &  Van  Essche  7349  (LISC!); 
Sao  Joao  dos  Angolares,  80  m,  Newton  s.n.  (BM!,  COI!);  s.l.,  Moller  s.n. 
(COI!),  F.A.E.  (Moller)  38  p.p.  (BM!,  LISJC!,  LISU!,  P!). 
PRINCIPE.  Porto  Real,  Rose  450  (P!);  Pico,  950  m,  Thorold  2082  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa  and  Mascarene  Is. 

Arthropteris  orientalis  (J.F.  Gmel.)  Posth.  in  Recueil  Trav.  Bot. 
Neerl.  21:  218  (1924);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  70 
(1944);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  116  (1964); 
Benl  in  Acta  Bot.  Barcinon.  33:  34  (1982). 

SAO  TOME.  Between  Monte  Cafe  and  Pico,  Chevalier  14562  (P!);  Pico, 
Moller  s.n.  (COI-not  found);  Cruzeiro,  300  m,  Thorold  2068  (BM!);  s.l., 
EA.E.  (Moller)  38  p.p.  (BM!,  COI!,  LISJC!,  P!). 
PRINCIPE.  Rio  Bambu-Porco,  60  m,  epiphyte,  Navel  143  (P!). 


E.  FIGUEIREDO 
DISTRIBUTION.     Tropical  Africa,  tropical  Asia,  Philippines. 

Nephrolepis  Schott 

Nephrolepis  biserrata  (Sw.)  Schott,  Gen.fiL:  ad  t.  3  (1834);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  70  ( 1 944);  Sobrinho  inPortugaliae 
Acta  BioL,  ser.  B,  Sist.  3:  392  (1952);  Benl  in  Acta  Bot.  Barcinon. 
33:36(1982). 

SAO  TOME.  Boa  Entrada,  Chevalier  13789  (P);  between  Monte  Caf£  and 
Pico,  Chevalier  14291bis  (P);  Sao  Miguel,  50  m,  not  frequent,  Espirito  Santo 
4743  (LISC!,  LISJC!);  Agua  Ize,  littoral,  ground  fern,  Exell  482  (BM!, 
COI!),  common  epiphyte,  Pinto  Basto  214  (LISC!);  Rio  Contador,  1300  m, 
Moller  s.n.  (COI!);  Bombaim,  Monod  11887  (BM!);  W.  of  Pico,  Monod 
1 1895  (BM!);  between  Quinas  Alias  and  Monte  Quinas,  520-750  m,  Monod 
12021  (BM!),  12022  (BM!);  Santelmo,  secondary  forest,  very  abundant 
epiphyte,  Oliveira  &  Noronha  23-A  (LISU!);  Sao  Nicolau,  Dependencia 
Roca  Nova,  road  to  Tras-os-Montes,  Oliveira  &  Noronha  s.n  (LISU!);  Ilheu 
das  Rolas,  Quintas  s.n.  (COI!);  Cruzeiro,  300  m,  Thorold  2063  (BM!);  s.l., 
EA.E.  (Moller)  46  (BM!,  COI!,  LISU!),  Moller  s.n.  (COI!),  Rattray  s.n.  (E), 
Souza  s.n.  (COI!). 

PRINCIPE.  Terreiro  Velho,  plantation,  200  m,  ground  fern,  Exell  525  (BM !, 
COI!),  526  (BM!,  COI!);  s.l.,  Rattray  s.n.  (E),  Souza  s.n.  (COI!),  Welwitsch 
23  (BM!,  K!,  LISU!),  Welwitsch  23b  (BM!,  LISU!). 

DISTRIBUTION.    Pantropical. 

Nephrolepis pumicicola  F.  Ballard  inKewBulL  [1955]:  467  (1955); 
Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  445 
(1959);  Tardieu-Blot  in  Aubrev.,  FL  Cameroun  3:  108  (1964); 
Benl  in  Acta  Bot.  Barcinon.  33:  37  (1982). 

SAO  TOME.  Lagoa  Amelia,  Chevalier  13650  (P!),  1412  m,  Matos  7505 
(LISC!),  1380  m,  Monod  11759  (BM!);  Pico,  Chevalier  13659ter  (P!); 
between  Monte  Cafe  and  Pico,  Chevalier  14581  (P!);  s.l.,  1500  m,  Mann 
1 106  (K!),  Mann  s.n.  (P!). 

DISTRIBUTION.    Cameroun,  Bioko. 

Nephrolepis  undulata  (Afzel.  ex  Sw.)  J.  Sm.  in  Bot.  Mag.  72, 
Comp.:  37  [as  '35']  (1846);  Alston  in  Exell,  Cat.  Vase.  PL  S. 
Tome:  71  (1944);  Benl  in  Acta  Bot.  Barcinon.  33:  39  (1982). 

SAO  TOME.  Sao  Nicolau,  870  m,  Espirito  Santo  5053  (LISC!,  LISJC!); 
Macambrara,  coffee  plantation,  1200  m,  epiphyte,  Exell  116  (BM!,  COI!), 
primary  forest,  1050-1200  m,  epiphyte,  Exell  190a  (BM!);  Lagoa  Amelia, 
1300  m,  ground  fern,  Exell  212  (BM!);  Tras-os-Montes,  Gama  s.n.  (COI!); 
Santa  Catarina,  Lains  e  Silva  387  (LISU!);  between  Monte  Castro  and  Ponta 
Figo,  secondary  forest,  500-1070  m,  Lejoly  94/39  (LISC!);  Pico,  primary 
forest,  1900-2020  m,  Lejoly  94/540  (LISC !);  Nova  Moca,  850  m,  Moller  s.n. 
(COI!);  Saudade,  Moller  s.n.  (COI!);  Monte  Cafe,  690  m,  Monod  11669 
(BM!),  600  m,  Thorold  2061  (BM!);  Bombaim,  Monod  11885  (BM!); 
between  Monte  Quinas  and  Quinas  Alias,  Monod  12016  (BM!);  NW  of  Pico, 
above  Cascata,  1000  m,  Monod  12267  (BM!);  Agua  Ize,  Pinto  Basto  209 
(LISC!);  s.l.,  Brentel  s.n.  (COI!),  EA.E.  (Moller)  45  (BM!,  LISU!),  Mann 
(not  found). 
PRINCIPE.  Pico,  summit,  Rozeira  924  (PO!). 


DISTRIBUTION.     Widespread  in  tropical  Africa  and  Mascarene  Is.          DISTRIBUTION.     Widespread  in  tropical  and  S.  Africa. 


Arthropteris  palisotii  (Desv.)  Alston  inBol.  Soc.  Brot.,  ser.  2, 30:  6 
(1956);  Benl  in  Acta  Bot.  Barcinon.  33:  31  (1982). 

Arthropteris  obliterata  sensu  auct.  non  (R.  Br.)  J.  Sm.,  Ferns  Brit, 
for.:  163  (1866);  C.  Chr.,  Index filic.:  62  (1905);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  70  (1944). 

SAO  TOME.  Sao  Joao  dos  Angolares,  120  m,  EA.E.  (Quintas)  47  (BM!, 
LISU!),  80  m,  Newton  s.n.  (BM!,  COI!),  120  m,  Quintas  s.n.  (COI!);  Agua 
Pinhao,650m,  Quintas  1 454  (BM!,  COI!);  Monte  Cafe,  760m,  Welwitsch  55 
(BM!,  K!,LISU!). 


Oleandraceae 

Oleandra  Cav. 

Oleandra  distenta  Kunze  in  Bot.  Zeitung  (Berlin)  9:  347  (1851); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  69  (1944);  Benl  in  Acta 
Bot.  Barcinon.  33:  28  (1982). 

Oleandra  annetii  sensu  auct.  non  Tardieu  in  Notul.  Syst.  (Paris)  14: 
332  (1953);  Pichi  Sermolli  in  Webbia  20:  763  (1965)  p.p.  quoad 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 

distr.  Sao  Tome;  Benl  in  Acta  Bot.  Bardnon.  33:  30  (1982)  p.p. 
quoad  distr.  Sao  Tome. 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  epiphyte,  Exell 
237  (BM!);  Nova  Moca,  950  m,  Matter  s.n.  (COI!);  Bom  Sucesso,  1250  m, 
Matter  s.n.  (COI!);  Calvario,  Espirita  Santo  5058  p.p.  (LISC!);  Sao  Joao  dos 
Angolares,  Newton  s.n.  (BM!,  COI!);  Tra"s-os-Montes,  Quintas  s.n.  (COI!); 
Monte  Cafe,  Welwitsch  52  (BM!,  LISU!);  s.l.,  F.A.E.  (Matter  &  Quintas)  48 
(BM!,  G,  K  also  Moller  43\,  LISU!). 
PRINCIPE.  S.I.,  Barter  1928  (K!). 

DISTRIBUTION.  Widespread  in  tropical  and  subtropical  Africa  and 
Mascarene  Is. 

NOTE.  Some  collections  from  Sao  Tome  previously  identified  as 
Oleandra  annetii  Tardieu  have  squarrose  rhizomes  with  abruptly 
subulate  to  gradually  tapering  scales  and  for  this  reason,  they  are 
here  referred  to  O.  distenta  s.l. 

Ophioglossaceae 

Ophioglossum  L. 

Ophioglossum  costatum  R.  Br.,  Prodr.:  163  (1810);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  94  (1944). 

SAO  TOME.  Between  Macambrara  and  Sao  Nicolau,  Exell  307  (BM-not 
found). 

DISTRIBUTION.  Widespread  in  tropical  Africa,  Asia,  and  Australa- 
sia. 

Ophioglossum  reticulatum  L.,  Sp.  pi.  2:  1063  (1753);  Alston  in 
Exell,  Cat  Vase.  PI.  S.  Tome:  94  (1944);  Benl  in  Acta  Bot. 
Bardnon.  33:  3  (1982). 

SAO  TOME.  Near  Sao  Pedro,  900-1200  m,  Chevalier  14532  (P!); 
Magodinho,  50  m,  Espirito  Santo  4929  (LISC!,  LISJC!);  Macambrara, 
primary  forest,  1050-1200  m,  Exell  444  (BM!);  Nova  Moca,  800  m,  F.A.E. 
(Quintas)  72  (BM!,  COI!,  K!,  LISU!,  P!),  Quintas  s.n.  (COI!);  Diogo  Nunes, 
Newton  s.n.  (BM!,  COI!,  P!);  Diogo  Vaz,  between  Paga  Fogo  and  Rio  Paga 
Fogo,  Rozeira  698  (PO!);  s.l.  (probably  Nova  Moca),  Quintas  62  (K!), 
Welwitsch  30  (BM\,  LISU!). 

DISTRIBUTION.     Pantropical  and  subtropical. 

Polypodiaceae 

Anapeltis  J.  Sm. 

Anapeltis  lycopodioides  (L.)  J.  Sm.,  Cult,  ferns:  6  (1857). 

var.  owariensis  (Desv.)  Benl  in  Acta  Bot.  Bardnon.  33:  18  (1982). 

Polypodium  lycopodioides  sensu  auct.  non  L.,  Sp.  pi.  2:  1082  (1753); 

Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  88  (1944);  Sobrinho  in 

Portugaliae  Acta  BioL,  Ser.  B,  Sist.  3:  393  (1952). 
Microgramma  owariensis  (Desv.)  Alston  in  Bol.  Soc.  Brot.,  ser.  2, 

30:  20  (1956);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3: 

80(1958). 

SAO  TOME.  Pico,  Chevalier  13639  (P);  Porto  Alegre,  Chevalier  1421  Ibis 
(P);  Vila  Conceicao,  100  m,  Matos  7718  (LISC!);  Caixao  Grande,  200  m, 
Moller  s.n.  (COI!);  Trindade,  250  m,  Matter  s.n.  (COI!);  near  Sao  Tome, 
Mailer  s.n.  (COI!);  Ribeira  Peixe,  epiphyte,  Oliveira  &  Noronha  35-A 
(LISU!);  Agua Ize,  epiphyte  on  plantation,  Pinto Basto206  (LISC!);  Correia, 
50  m,  Quintas  1246  (BM!,  COI!);  Ilheu  das  Rolas,  Quintas  s.n.  (COI!);  Sao 
Nicolau,  900  m,  Quintas  s.n.  (LISU!);  Ribeira  Peixe,  between  Vila  Jose  and 
Ermelinda,  epiphyte,  Rozeira  236  (PO!);  Diogo  Vaz,  Rozeira  869  (PO);  s.l., 
F.A.E.  (Moller  &  Quintas)  54  (COI!,  BM!,  LISJC!,  LISU!),  Seabra  s.n. 
(COI!,  LISU!). 


59 

PRINCIPE.  Terreiro  Velho,  200  m,  epiphyte,  Exell  534  (BM!,  COI!);  Porto 
Real,  epiphyte,  Lains  e  Silva  420  (LISU!);  Sundi,  epiphyte,  Monod  12076 
(BM!),  Thorold  2080  (BM!);  Berimbau,  Newton  23  (BM!,  COI!);  Cimalo, 
Quintas  43  (BM!,  COI!);  Infante  D.  Henrique,  at  the  base  of  Dois  Irmaos, 
epiphyte,  Rozeira  435  (BM-not  found,  PO!);  Esperan?a,  1 20  m,  Rozeira  996 
(PO),  Thorold2016  (BM!);  Santo  Antdnio,  sea  level,  epiphyte,  Thorold2019 
(BM!);  s.l.,  Rattray  s.n.  (E,  K!). 

DISTRIBUTION.    W.  tropical  Africa. 

Belvisia  Mirb. 

Belvisia  spicata  (L.f.)  Mirb.  ex  Copel.,  Gen.fiL:  192  (1947);  Alston 
in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  444  (1959); 
Tardieu-Blot  in  Aubrev.,  FL  Cameroun  3:  342  (1964);  Benl  in 
Acta  Bot.  Bardnon.  33:  12  (1982). 

Hymenolepis  spicata  (L.f.)  C.  Presl,  Epimel.  hot.:  159  (1851); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  90  (1944);  Sobrinho  in 
Garcia  de  Orta  7:  89  (1959). 

SAO  TOME.  Calvario,  secondary  forest,  920  m,  epiphyte,  Espirito  Santo 
5075  (LISJC!);  Ribeira  Peixe,  140  m,  epiphyte,  Lains  e  Silva  248  (LISU!), 
257  (LISJC-not  found),  284  (LISJC-not  found);  Tras-os-Montes,  700  m, 
Quintas  1322  p.p.  (BM!,  COI!);  Nova  Moca,  1 100  m,  Quintas  s.n.  (LISU!); 
Ribeira  Peixe,  between  Vila  Jose  and  Ermelinda,  Rozeira  272  (PO!);  Cruzeiro, 
300  m,  Thorold2067  (BM!),  300 m,  2071  (BM!);  s.l.,  900  m,  Quintas?,  (K!). 

DISTRIBUTION.     Tropical  Africa  and  Mascarene  Is. 

Drynaria  (Bory)  J.  Sm. 

Drynaria  laurentii  (H.  Christ  ex  De  Wild.  &  T.  Durand)  Hieron. 
in  Engl.  &  Drude,  Veg.  Erde  9(2):  57,  fig.  54  (1908);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  90  (1944);  Benl  in  Acta  Bot. 
Bardnon.  33:  10(1982). 

SAO  TOME.  Monte  Cafe,  Chevalier  14291  (P!),  600  m,  Thorold  2062 
(BM!),  600-760  m,  Welwitsch  69  (BM!,  K!,  LISU!);  Blu-blu,  50  m,  Moller 
s.n.  (COI!);  Agua  Ize,  epiphyte,  Pinto  Basto2\l  (LISC!);  Diana,  Rozeira  957 
(PO!);  Perseveranca,  Seabra  173  (LISU!);  s.l.,  Don  s.n.  (BM!),  F.A.E. 
(Moller)  57  (BM!,  K  also  Moller  52!,  LISU!). 
PRINCIPE.  S.l.,  Welwitsch  17  (BM!,  K!,  LISU!). 

DISTRIBUTION.    Tropical  Africa. 

Drynaria  volkensii  Hieron.  in  Engl.  &  Drude,  Veg.  Erde  9(2):  57 
(1908)  nomen;  in  Bot.  Jahrb.  Syst.  46:  393  (1911);  Alston  in 
Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  80  (1958);  Alston  in 
Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  444  (1959); 
Benl  in  Acta  Bot.  Bardnon.  33:  1 1  (1982). 

SAO  TOME.  Nova  Moca,  900  m,  Moller  s.n.  (COI!);  Cruzeiro,  300  m, 
Thorold  2066  (BM!);  Agua  Ize,  180  m,  Thorold  2092  (BM!). 
PRINCIPE.  Praia  da  Lapa,  epiphyte,  Rozeira  1902  (BM-not  found,  PO!); 
s.l.,  Barter  1921  (K!). 

DISTRIBUTION.     Widespread  in  tropical  Africa. 

Microsorum  Link 

Microsorum  punctatum  (L.)  Copel.  in  Univ.  Calif.  Publ.  Bot.  16: 
1 1 1  (1929);  Benl  in  Acta  Bot.  Bardnon.  33:  14  (1982). 

Polypodium  irioides  Poir.,  Encycl.  5:  513  (1804);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  88  (1944). 

SAO  TOME.  Between  Bom  Sucesso  and  Lagoa  Amelia,  Almeida  s.n. 
(LISC!);  Boa  Entrada,  Chevalier  13788  (P!),  14375  (not  found);  Sao  Joao 
dos  Angolares,  F.A.E.  (Quintas)  55  (LISU!),  Quintas  s.n.  (COI!);  Agua  Ize", 
common  epiphyte,  Pinto  Basto  203  (LISC!);  s.l.,  Mendonca  s.n.  (P!),  Souza 
s.n.  (BM!,  COI!). 


60 


E.  FIGUEIREDO 


PRINCIPE.  SantoAntonio,  sea  level,  epiphyte,  Monod  1 2 1 89  (BM !);  Sundi, 
150  m,  Quintas  62  (COI!);  Esperan?a,  120  m,  Thorold  2077  (BM!);  Lapa, 
near  sea  level,  Thorold  2089  (BM!),  2090  (BM!);  s.l.,  Rattray  s.n.  (E), 
Welwitsch  25  (BM\,USU\). 

DISTRIBUTION.    Palaeotropical. 

Phymatosorus  Pic.  Serm. 

Phymatosorus  scolopendria  (Burm.f.)  Pic.  Serm.  in  Webbia  28: 
460  (1973);  Benl  in  Acta  Bot.  Barcinon.  33:  13  (1982). 

Polypodium phymatodes  L.,  Mant.  pi.  2:  306  ( 1 77 1 );  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  89  (1944);  Alston  in  Exell,  Suppl.  Cat. 
Vase.  Pl.S.  Tome:  8(1956). 

SAO  TOME.  Boa  Entrada,  Chevalier  13786  (P);  Porto  Alegre,  Chevalier 
14243  (K!);  Famosa,  400  m,  Espirito  Santo  69  (BM!,  COI!,  LISJC!);  Ilheu 
das  Rolas,  plantation,  70  m,  epiphyte,  Espirito  Santo  5 1 10  (LISC!,  LISJC!), 
Quintas  s.n.  (COI!);  Agua  Ize,  epiphyte,  Exell  480  (BM!,  COI!),  epiphyte, 
Pinto  Basto  201  (LISC!);  bridge  over  R.  Abade,  Figueiredo  &  Arriegas  34 
(LISC!);  Rio  SantoAntonio,  Matos  7646  (LISC!);  Praia  Pantufo,  Moller  s.n. 
(COI!);  nearBatepa,  500  m,  Moller  s.n.  (COI!);  Bombaim,  epiphyte,  Monod 
1 1880(BM!);NW  of  Pico,  1000  m,M0/i<«/12268(BM!,  COI!);  Quingombo, 
100  m,  Newton  s.n.  (COI!);  Cruzeiro,  300  m,  Thorold  2070  (BM!);  Monte 
Cafe,  850m,  Welwitsch  5\  (BM!,  LISU!);  s.l, Don  s.n.  (BM!),  F.A.E.  (Moller 
&  Quintas)  56  (BM!,  COI!,  LISJC!,  LISU!),  Mendonfa  (COI!,  K!),  Moller 
s.n.  (COI!),  Rozeira  1  (PO!),  Seabra  s.n.  (COI!). 

PRINCIPE.  Terreiro  Velho,  plantation,  200  m,  ground  fern,  Exell  523  (BM!, 
COI!);  Esperanca,  120  m,  Thorold2015  (BM!),  2078  (BM!);  Lapa,  near  sea 
level,  Thorold  2091  (BM!);  s.l.,  Rattray  s.n.  (E),  Welwitsch  9  (BM!,  K!, 
LISU!),  9bis  (K!). 

DISTRIBUTION.     Palaeotropical. 

Platycerium  Desv. 

Platycerium  stemaria  (P.  Beauv.)  Desv.  \nMem.  Soc.  Linn.  Paris  6: 
213  (1827);  Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  91  (1944); 
Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  8  (1956);  Benl  in 
Acta  Bot.  Barcinon.  33:  9  (1982);  Hennipman  &  Roos,  Monogr. 
fern  genus  Platycerium:  108  (1982). 

SAO  TOME.  Sao  Vicente,  200  m,  epiphyte,  Espirito  Santo  24  (BM!,  COI!, 
LISJC!);  near  Sao  Tome,  Chevalier  13726  (P!);Trindade,  250  m,  Moller  s.n. 
(COI!),  Quintas  s.n.  (COI!);  Rio  do  Ouro,  380  m,  Moller  s.n.  (COI!);  Diogo 
Vaz,  epiphyte,  Rozeira  829  (PO!);  s.l.,  250-350  m,  F.A.E.  (Moller)  70 
(LISU!),  Mendonfa  s.n.  (K!,  P!),  250  m,  Moller  60  (K!). 
PRINCIPE.  Precipfcio,  Monod  12178  (BM!);  Praia  das  Burras,  epiphyte, 
Rozeira  1029  (PO!);  s.l.,  Barter  1886  (K!),  Welwitsch  20  (K-not  found, 
LISU!),  20b  (BM-not  found,  LISU!). 

DISTRIBUTION.     W.  and  Central  tropical  Africa. 

Pleopeltis  Humb.  &  Bonpl.  ex  Willd. 

Pleopeltis  excavata  (Bory  ex  Willd.)  Sledge  in  Bull.  Br.  Mus.  (Nat. 
Hist.)  Bot.  2:  138  (1960);  Benl  in  Acta  Bot.  Barcinon.  33:  16 
(1982). 

Polypodium  preussii  Hieron.  in  Engl.  in  Bot.  Jahrb.  Syst.  46:  386 
(1911);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  89  (1944). 

SAO  TOME:  Calvario,  Espirito  Santo  5058  p.p.  (LISC!,  LISJC!); 
Macambrara,  primary  forest,  1050-1200  m,  epiphyte,  Exell  113  (BM!); 
Lagoa  Amelia,  Exell  216  (BM-not  found),  1412  m,  Matos  751 1  (LISC!), 
1380  m,  Monod  11758  (BM!),  1450  m,  Monod  11768  (BM!,  COI!); 
Macambrara  to  Zampalma,  900  m,  epiphyte,  Exell  295  (BM!);  Pico,  light 
woodland,  2000m,  epiphyte,  Exell  341  (BM!),  primary  forest,  1 900-2020  m, 
Lejoly  94/536  (LISC!),  Mann  (not  found),  primary  forest,  1987  m,  epiphyte, 
Matos  &  Van  Essche  7386  (LISC!);  Bom  Sucesso,  1080  m,  F.A.E.  (Moller  & 
Quintas)  53  (BM!,  LISU!),  Moller  s.n.  (COI!);  near  Sao  Nicolau  waterfall, 


800  m,  Lains  e  Silva  129  (LISU!);  Casa  do  Pico,  1930  m,  Monod  12250 
(BM!);s.l.,  Souzas.n.  (COI!). 

DISTRIBUTION.     Palaeotropical. 

Pleopeltis  macrocarpa  (Bory  ex  Willd.)  Kaulf.  in  Berlin.  Jahrb. 
Pharm.  Verbundenen  Wiss.  21:  41  (1820). 

SAO  TOME.  Lagoa  Amelia,  primary  forest,  1400-1420  m,  epiphyte,  Matos 
&  Van  Essche  741 1  (LISC!). 

DISTRIBUTION.     Pantropical  and  subtropical 

Pyrrosia  Mirb. 

Pyrrosia  lanceolate  (L.)  Farw.  in  Amer.  Midi.  Naturalist  12:  245 
(1931);  Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  80 
(1958);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  340  (1964); 
Hovenkamp,  Monogr.  fern  genus  Pyrrosia:  191  (1986). 

Cyclophorus  spissus  (Bory  ex  Willd.)  Desv.  in  Ges.  Naturf.  Freunde 
Berlin  Mag.  Neuesten  Entdeck.  Gesammten  Naturk.  5:  301  (1811) 
var.  continentalis  Hieron.  ex  Engl.  in  Engl.  &  Drude,  Veg.  Erde 9(2): 
55,  fig.  53  (1908);  Alston  in  Exell,  Cat .  Vase.  PI.  S.  Tome:9\  (1944). 

SAO  TOME.  Cruzeiro,  Rozeira  690  (BM-not  found,  PO). 
PRINCIPE.  Cimalo,  Quintas  30  (BM!,  COI!,  P!,  Z);  Esperan?a,  way  to 
Montalegre,  Rozeira  392  (PO!);  Praia  da  Lapa,  Rozeira  1944  (PO!);  s.l., 
Keulemans  s.n.  (L),  Quintas  A  (K!). 

DISTRIBUTION.     Palaeotropical. 

Psilotaceae 

Psilotum  Sw. 

Psilotum  nudum  (L.)  P.  Beauv.,  Prodr.  aetheogam.:  106,  112 
(1805);  Alston  in  Exell,  Suppl.  Cat.  Vase.  PL  S.  Tome:  9  (1956); 
Benl  in  Acta  Bot.  Barcinon.  31:  21  (1978). 

SAO  TOME.  Praia  Melao,  Rozeira  57  (PO!);  s.l.,  Don  s.n.  (BM!). 

DISTRIBUTION.  Widespread  in  tropical  and  subtropical  regions 
worldwide. 

Pteridaceae 

Pteris  L. 

Pteris  atrovirens  Willd.,  Sp.  pi.  5:  385  (1810);  Alston  in  Exell,  Cat. 
Vase.  PL  S.  Tome:  83  (1944);  Tardieu-Blot  in  Aubrev.,  FL  Cam- 
eroun 3:  166  (1964);  Benl  in  Acta  Bot.  Barcinon.  38:  16  (1988). 

SAO  TOME.  Boa  Entrada,  Chevalier  13792  p.p.  (P!);  Pico,  Chevalier  14617 
(P!);  Tras-os-Montes,  Gama  s.n.  (COI!);  between  Quinas  Altas  and  Monte 
Quinas,  Monod  12009  (BM!,  COI!);  Agua  Ize,  Pinto  Basto  210  (LISC!); 
Pedroma,  Rozeira  1370  (K!);  s.l.,  Don  s.n.  (BM!),  F.A.E.  (Moller)  18  p.p. 
(BM!,  COI!,  LISU!,  P!),  Moller  s.n.  p.p.  (COI!),  Moller  s.n.  (COI!),  Rattray 
s.n.  (E,  K!). 

PRINCIPE.  Esperanga,  Navel  146  (P!);  Sundi,  Newton  42  (COI!);  Pico 
Papagaio,  Welwitsch  22  (BM!,  LISU!);  s.l.,  Jardim  (cited  by  Kuhn,  1868), 
Rattray  s.n.  (E). 

DISTRIBUTION.    W.  tropical  Africa. 

Pteris  burtonii  Baker  in  Ann.  Bot.  (London)  5:  218  (1891);  Alston 
in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  442  (1959); 
Benl  in  Acta  Bot.  Barcinon.  38:  7  (1988). 

PRINCIPE.  S.l.,  Fox  s.n.  (BM!). 
DISTRIBUTION.     W.  and  E.  tropical  Africa. 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


61 


NOTES.     This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

The  collection  Walg  1 1  (WAG),  from  Sao  Tome,  was  interpreted 
as  a  possible  hybrid  between  Pteris  atrovirens  and  P.  burtonii  by 
Benl(1988:  17). 

Pteris  dentata  Forssk.,  Fl.  aegypt.-arab.:  186  (1775);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  84  (1944). 

subsp.  flabellata  (Thunb.)  Runemark  in  Bot.  Not.  115:  190  (1962); 
Benl  in  Acta  Bot.  Bardnon.  38:  1 1  (1988). 

Pteris  communata  sensu  auct.  non  Kuhn  in  Von  der  Decken's  Reisen 
Ost-Afr.,  Bot.  3:  20  (1879);  Alston  in  Exell  in  Bull.  Inst.  Franc. 
Afrique  Noire  ser.  A,  21:  442  (1959). 

SAO  TOME.  Lagoa  Amelia,  1400  m,  Monod  1 1749  (BM!). 
PRINCIPE.  Esperanca,  600  m,  Rozeira  890  (K!). 

DISTRIBUTION.     Subtropical  Africa,  Madagascar,  Mascarene  Is., 
Ascension,  St  Helena. 

Pteris  linearis  Poir.,  Encyd.  5:  723  (1804);  Alston  in  Exell  & 
Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  79  (1958);  Benl  in  Acta  Bot. 
Bardnon.  38:  18(1988). 

Pteris  biaurita  sensu  auct.  non  L.,  Sp.  pi.  2:  1076  (1753);  Alston  in 
Exell,  Cat.  Vase.  PI.  S.  Tome:  83  (1944). 

SAO  TOME.  Boa  Entrada,  Chevalier  13792  p.p.  (P);  Ilheu  das  Rolas,  palm 
plantation,  Espirito  Santo  5 145  (LISC!,  LISJC!),  Rozeira  74  (BM-not  found, 
PO!);  Bombaim,  Rozeira  495  (PO);  Santa  Catarina,  palm  plantation,  Rozeira 
790  (K!);  s.l.,  Don  s.n.  (BM!),  F.A.E.  (Moller)  16  (BM!,  COI!,  LISU!),  900 m, 
Moller  14  (K!),  Moller  s.n.  (COI!),  Rattray  s.n.  (E). 
PRINCIPE.  Oque  Nazare,  Carvalho  s.n.  (COI!);Terreiro  Velho,  plantation, 
200  m,  ground  fern,  Exell  529  (BM!,  COI!);  s.l.,  Welwitsch  22b  (LISU!). 

DISTRIBUTION.     Palaeotropical. 

Pteris  paucipinnata  Alston  in  Exell  in  Bull.  Inst.  Franf.  Afrique 
Noire  ser.  A,  21:  443  (1959);  Benl  in  Acta  Bot.  Bardnon.  38:  14 
(1988). 

Pteris  catoptera  sensu  auct.  non  Kunze  in  Linnaea  18:  119  (1844); 

Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  79  (1958). 
Pteris  quadriaurita  sensu  auct.  non  Retz,  Observ.  hot.  6:  38  (1791); 

Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  84  (1944)  p.p.  quoad  spec. 

Sao  Tome. 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  ground  fern,  Exell 
187  (BM !);  between  Pico  and  Monte  Castro,  6  km  S.  of  Ponta  Figo,  mountain 
rainforest,  1200-1 800  m,Lejoly 95/1 1  (LISC!);  Monte  Cafe,  lQlOm,Monod 
11690  (BM!-holotype);  Colonia  A?oreana,  Rozeira  267  (PO);  Diogo  Vaz, 
Rozeira  516  (PO);  s.l.,  350  m,  Welwitsch  54  (BM!,  LISU!). 
PRINCIPE.  Pico  Papagaio,  600  m,  ground  fern,  Exell  686  (BM!). 

DISTRIBUTION.    Bioko. 

Pteris  pteridioides  (Hook.)  F.  Ballard  in  Kew  Bull.  [1937]:  348 
(1937);  Alston  in  Exell,  Cat  Vase.  PL  S.  Tome:  84  (1944); 
Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  170  (1964);  Benl  in 
Acta  Bot.  Bardnon.  38:  9  (1988). 

SAO  TOME.  Tras-os-Montes,  Gama  s.n.  (COI!);  Macambrara,  1200  m, 
Quintas  5  (K!),  1200  m,  Quintas  1352  (BM!,  COI!,  K!,  P!). 

DISTRIBUTION.     Widespread  in  tropical  and  subtropical  Africa. 

NOTE.     This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 


Pteris  similis  Kuhn  in  Von  der  Decken's  Reisen  Ost-Afr.,  Bot.  3:  21 
(1879);  Benl  in  Acta  Bot.  Bardnon.  38:  18  (1988). 

Pteris  atrovirens  sensu  auct.  nonWilld.,5/?.  pi.  5:  385  (1810);Alston 
in  Exell,  Cat.  Vase.  PI.  S.  Tome:  83  (1944)  p.p. 

SAO  TOME.  Sao  Miguel,  Monte  Verde,  secondary  forest,  10-150  m,  Lejoly 
95/98  (LISC!);  Agua  Ize,  common,  Pinto  Basto  200  (LISC!);  Monte  Cafe, 
760  m,  Welwitsch  53  (BM!,  LISU!);  s.l.,  Don  s.n.  (BM!). 
PRINCIPE.  Between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350 
m,  ground  fern,  Exell  561  (BM!,  COI!);  Sundi,  150  m,  Quintas  45  (BM!, 
COI!,  K!,  P!);  Infante  D.  Henrique,  Rose  407  (P!). 

DISTRIBUTION.    W.  tropical  Africa  to  E.  Africa. 

Pteris  togoensis  Hieron.  in  Brause  &  Hieron.  in  Engl.  in  Bot.  Jahrb. 
Syst.  53:  402  (1915);  Benl  in  Acta  Bot.  Bardnon.  38:  13  (1988). 

PRINCIPE.  S.l.,  Barter  1926  (K!). 
DISTRIBUTION.    W.  tropical  Africa  to  E.  Africa. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Pteris  tripartita  Sw.  in  J.  Bot.  (Schroder)  1800(2):  67  (1801); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  84  (1944);  Benl  in  Acta 
Bot.  Bardnon.  38:  15  (1988). 

SAO  TOME.  Boa  Entrada,  Chevalier  13720  (P!);  Ilheu  das  Rolas,  EA.E. 
(Quintas)  20  (BM!,  LISU!),  Quintas  s.n.  (COI!);  s.l.,  Don  s.n.  (BM!);  s.l., 
Rattray  s.n.  (E). 
PRINCIPE.  Sundi,  littoral,  Quintas  1 1  (BM!,  COI!,  K!,  P!). 

DISTRIBUTION.    Pantropical. 

Pteris  vittata  L.,  Sp.  pi.  2:  1074  (1753);  Alston  in  Exell  &  Rozeira 
in  Conf.  Int.  Afr.  Oc.  3:  79  (1958);  Alston  in  Exell  in  Bull.  Inst. 
Franc.  Afrique  Noire  ser.  A,  21:  443  (1959). 

SAO  TOME.  Monte  Cafe,  690  m,  Monod  1 1671  (BM!),  Rozeira  s.n.  (PO); 
Agua  Ize,  Pinto  Basto  216  (LISC!). 

DISTRIBUTION.  Pantropical.  Alston  (1958,  1959)  suggested  that 
this  might  be  an  escape  from  cultivation. 

Selaginellaceae 
Selaginella  P.  Beauv. 

Selaginella  cathedrifolia  Spring  in  Mem.  Acad.  Roy.  Sd.  Belgique 
24:  112  (1849);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  96 
(1944);  Tardieu-Blot  in  Aubrev.,  FL  Cameroun  3:  30  (1964); 
Benl  in  Acta  Bot.  Bardnon.  31:  15  (1978). 

PRINCIPE.  Above  Infante  D.  Henrique,  secondary  forest,  300  m,  ground 
fern,  Exell  629  (BM!);  Pico  Papagaio,  600  m,  ground  fern,  Exell  687  (BM!, 
COI!);  near Lapa, Monod  12094 (BM!);  s.l., Barter  1931  (K\),Curror 3 (K!- 
type),  Mann  s.n.  (K!). 

DISTRIBUTION.    W.  and  Central  tropical  Africa. 

Selaginella  mannii  Baker  in  J.  Bot.  23:  1 80  ( 1 885);  Alston  in  Exell, 
Cat.  Vase.  PL  S.  Tome:  96  (1944). 

SAO  TOME.  Between  Monte  Caf<§  and  Pico,  Chevalier  14552  (P!); 
Macambrara,  primary  forest,  1050-1200  m,  Exell  95  (BM!,  COI!),  106 
(BM!),  168  (BM!),  1260  m,  Moller  s.n.  (COI!);  between  Bom  Sucesso  and 
Calvario,  primary  forest,  1350-1400  m,  Matos  &  Van  Essche  7347  (LISC!); 
Bom  Sucesso,  1 250  m,Moller  s.n.  (COI!);  Pico,  1900m,Afo//ers.n.  (COI!); 
Rio  Contador,  1350  m,  Moller  s.n.  (COI!);  Lagoa  Amelia,  1400  m,  Monod 


62 


E.  FIGUEIREDO 


1 1784  (BM!);  W.  of  Pico,  870-1225  m,  Monod  1 1904  (BM!);  Calvario,  Rose 
151  (P!);  s.l.,  F.A.E.(Moller)  80  (BM!,  COI!,  K  also  Matter  75!,  LISU!,  P!), 
Henriques  4  (K!),  1500  m,  Mann  1 108  (K!-type). 
PRINCIPE.  Pico  Papagaio,  600  m,  ground  fern,  Exell  683  (BM!). 

DISTRIBUTION.     Endemic. 

Selaginella  molleri  Hieron.  in  Engl.  &  Prantl,  Nat.  Pflanzenfam. 
1(4):  697  (1901);  Alston  in  Exell,  Cat  Vase.  PL  S.  Tome:  97 
(1944);  Benl  in  Acta  Bot.  Barcinon.  31:  20  (1978). 

SAO  TOME.  Pico,  Chevalier  13668  (P!);  Macambrara,  primary  forest, 
1050-1200  m,  Exell  180  (BM!,  COI!);  Bom  Sucesso,  1250  m,  Moller  s.n. 
(COI!);  Rio  Manuel  Jorge,  waterfall,  800  m,  Moller  s.n.  (COI!);  s.l.,  F.A.E. 
(Moller)  79  p.p.  (BM  type!,  COI!,  K!,  LISU!). 

DISTRIBUTION.     Liberia,  Nigeria,  Bioko. 

Selaginella  molliceps  Spring  in  Mem.  Acad.  Roy.  Sci.  Belgique  24: 
257  (1849);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  97  (1944); 
Alston  in  Exell  &  Rozeira  in  Conf.  Int.  Afr.  Oc.  3:  8 1  ( 1 958);  Benl 
in  Acta  Bot.  Barcinon.  31:  20  (1978). 

SA  TOME.  Between  Bacelar  and  Quija,  secondary  forest,  160  m,  Espirito 
Santo  5140  (LISC!,  LISJC!);  Ribeira  Peixe,  Cao  Grande,  primary  forest, 
Matos  7636  (LISC!);  between  Sao  Miguel  and  Zagaia,  primary  forest,  170- 
400  m,  Matos  &  Van  Essche  7238  (LISC!);  Diogo  Vaz,  Rozeira  879  (PO). 
PRINCIPE.  Oque  Pipi,  secondary  forest,  300  m,  ground  fern,  Exell  500 
(BM!,  COI!);  Sao  Joao,  450  m,  Newton  s.n.  (COI!),  Newton  4  (K!);  Pico 
Papagaio,  900  m,  Welwitsch  1 1  (BM!,  LISU!);  s.l.,  Curror  2  p.p.  (K!-type). 

DISTRIBUTION.     W.  and  Central  tropical  Africa. 

Selaginella  monodii  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique 
Noire  ser.  A,  21:  440  (1959). 

SAO  TOME.  Blu-blu,  Monod  1 1806  (BM!-holotype,  COI!). 
DISTRIBUTION.     Endemic. 

Selaginella  myosurus  (Sw.)  Alston  in  J.  Bot.  70:  64  (1932). 
PRINCIPE.  S.l.,  Curror  2  p.p.  (K!). 
DISTRIBUTION.    Tropical  Africa. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 

Selaginella  squarrosa  Baker  in  J.  Bot.  23:  180  (1885);  Alston  in 
Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser.  A,  21:  440  (1959). 

PRINCIPE.  Pico,  500-600  m,  Monod  12111  (BM!). 
DISTRIBUTION.     Cameroun,  Equatorial  Guinea. 

Selaginella  thomensis  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  97,  f. 
3  (1944);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  40  (1964). 

SAO  TOME.  Between  Santa  Maria  and  Zampalma,  1200  m,  Exell  261 
(BM!,  COI!);  Macambrara,  primary  forest,  1050-1200  m,  Exell  423  (BM!- 
holotype,  COI!);  road  Monte  Cafe  to  Sao  Nicolau,  Figueiredo  &  Arriegas 
121  (LISC!);  Nova  Moca,  890  m,  Moller  s.n.  (COI!,  K!);  Pico,  1950  m, 
Moller  s.n.  (COI!);  s.l.,  F.A.E.  (Moller)  79  p.p.  (BM!,  K!,  LISU!). 

DISTRIBUTION.     Sierra  Leone,  Nigeria,  Cameroun. 

Sinopteridaceae 

Pellaea  Link 

Pellaea  doniana  Hook.,  Sp.fil.  2:  137,  t.  125  fig.  A  (1858);  Alston 


in  Exell,  Cat  Vase.  PL  S.  Tome:  82  (1944);  Tardieu-Blot  in 
Aubrev.,  FL  Cameroun  3:  140  (1964);  Benl  in  Acta  Bot. 
Barcinon.  32:  29  (1980). 

SAO  TOME.  Pico,  Don  s.n.  (BM!-type);  Ilheu  das  Rolas,  plantation, 
Espirito  Santo  5106  (LISC!,  LISJC!),  F.A.E.  (Quintas)  15  (K  also  Quintas 
13!,  LISU!),  Quintas  s.n.  (COI!),  Rozeira  92  (PO!). 

PRINCIPE.  Between  Esperan9a  and  Fundao,  primary  forest,  300-350  m, 
Ere// 739  (BM!). 

DISTRIBUTION.     Widespread  in  tropical  Africa 
Thelypteridaceae 

Christella  H.  Lev. 

Christella  dentata  (Forssk.)  Brownsey  &  Jermy  in  Brit.  Fern  Gaz. 
10:  338  (1973);  Benl  in  Acta  Bot.  Barcinon.  38:  53  (1988). 

Dryopteris  dentata  (Forssk.)  C.  Chr.  in  Kongel.  Danske  Vtdensk. 
Selsk.  Naturvidensk.  Math.  Afh.  8(6):  24  (1920);  Alston  in  Exell, 
Cat.  Vase.  PI.  S.  Tome:  61  (1944);  Alston  in  Exell,  Suppl.  Cat. 
Vase.  PL  S.  Tome:  7(1956). 

SAO  TOME.  Boa  Entrada,  Chevalier  13787  (P!),  14336  (P!);  Sao  Vicente, 
200  m,  Espirito  Santo  41  p.p.  (BM!,  COI!,  LISJC!);  Novo  Destino,  760  m, 
Welwitsch  61  (BM!,  K!,  LISU!);  Monte  Cafe,  Welwitsch  65  (BM!,  LISU!); 
s.l.,  F.A.E.  (Moller)  42  p.p.  (COI!,  BM!),  Rattray  s.n.  (BM!,  E). 
PRINCIPE.  6queNazare,A^ewtons.n.(BM!,COI!);s.l.,^a«ra>'s.n.(BM!,E). 

DISTRIBUTION.     Palaeotropical. 

Christella  hispidula  (Decne.)  Holttum  in  Kew  Bull.  31:  312  (1976); 
Benl  in  Acta  Bot.  Barcinon.  38:  51  (1988). 

Dryopteris  quadrangularis  (Fee)  Alston  in  J.  Bot.  75:  253  (1937); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  63  (1944). 

SAO  TOME.  Agua  Grande,  100  m,  Moller  s.n.  (COI!);  Saudade,  720  m, 
Moller  s.n.  (COI!);  above  Monte  Cafe,  1010  m,  Monod  11691  (BM!); 
between  Monte  Quinas  and  Quinas  Alias,  Monod  12006  (BM!). 

DISTRIBUTION.    Pantropical. 

Cyclosorus  Link 

Cyclosorus  striatus  (Schum.)  Ching  in  Bull.  Fan  Mem.  Inst.  Biol. 
10:  249  (1941);  Benl  in  Acta  Bot.  Barcinon.  38:  55  (1988). 

Dryopteris  striata  (Schum.)  C.  Chr.,  Indexfilic.:  294  (1905);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  64  (1944). 

SAO  TOME.  Lagoa  Amelia,  Chevalier  13653  (P). 
DISTRIBUTION.    Tropical  Africa. 

Metathelypteris  (H.  Ito)  Ching 

Metathelypteris  fragilis  (Baker)  Holttum  in  /.  S.  African  Bot.  40: 
127  (1974). 

Dryopteris  fragilis  (Baker)  C.  Chr.,  Indexfilic.:  266  (1905);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  62  (1944). 

subsp.  guineensis  Benl  in  Nova  Hedwigia  27:  150  (1976);  Benl  in 
Acta  Bot.  Barcinon.  38:  43  (1988). 

SAO  TOME.  Macambrara,  primary  forest,  1050-1200  m,  Exell  133  (BM!); 
s.l.,  Newton  1  (K!). 

DISTRIBUTION.    Bioko. 

NOTE.  The  collection  cited  by  Holttum  (I.e.)  as  Newton  s.n.  (K  ex 
COI),  1885,  is  Newton  1  (K). 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 
Pneumatopteris  Nakai 

Pneumatopteris  afra  (C.  Chr.)  Holttum  in  Blumea  21:  306  (1973); 
Benl  in  Acta  Bot.  Barcinon.  38:  57  (1988). 

Dryopteris  oppositifolia  sensu  auct.  non  (Hook.)  C.  Chr.,  Index 

filic.:  281  (1905);  Alston  in  Exell,  Cat  Vase.  PL  S.  Tome:  63 

(1944)  p.p.  quoad  pi.  ex  Principe. 
Cyclosorus  afer  (H.  Christ)  Ching  in  Bull.  Fan  Mem.  Inst.  Biol.  10: 

242  ( 1 94 1 );  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser. 

A,  21:446(1959). 

SAO  TOME.  Sao  Vicente,  Espirito  Santo  41  p.p.  (LISC!);  Sao  Miguel,  50m, 
Espirito  Santo  4744  (LISC!);  between  Pico  and  Monte  Castro,  6  km  S.  of 
Ponta  Figo,  mountain  rainforest,  1200-1800  m,  Lejoly  95/30  (LISC!); 
Pedroma,  Mocquerys  39/40  (P!);  between  Quinas  Altas  and  Monte  Quinas, 
Monod  12023  (BM!);  Agua  Ize,  Pinto  Basto  213  (LISC!);  Lagoa  Amelia, 
Rose  204  (P\),  231  (P!). 

PRINCIPE.  Terreiro  Velho,  plantation,  200  m,  Exell  530a  (BM!,  COI!); 
between  Oque  Pipi  and  Morro  do  Leste,  secondary  forest,  350  m,  ground 
fern,  Exell  566  (BM!,  COI!);  Infante  D.  Henrique,  Rose  398  (P!);  s.l.,  Barter 
1924  (K!),  Rattray  s.n.  (E),  300  m,  Welwitsch  21  (BM!,  K!,  LISU!),  21bis 
(K!). 

DISTRIBUTION.     Widespread  in  W.  tropical  Africa. 

Pneumatopteris  oppositifolia  (Hook.)  Holttum  in  Blumea  21:  304 
(1973);  Holttum  in/  S.  African  Bot.  40:  157  (1974);  Benl  inActa 
Bot.  Barcinon.  38:  62  (1988). 

Dryopteris  oppositifolia  (Hook.)  C.  Chr.,  Index  filic.:  281  (1905); 

Alston  in  Exell,  Cat.  Vase.  PI.  S.  Tome:  63  (1944)  p.p.  quoad  pi.  ex 

Sao  Tome. 
Cyclosorus  oppositifolius  (Hook.)Tardieu  inNotul.  Syst.  (Paris)  14: 

346  ( 1 953);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire  ser. 

A,  21:445(1959). 

SAO  TOME.  Between  Lagoa  Amelia  and  Calvario,  primary  forest,  150  m, 
ground  fern,  Exell  259  (BM!,  COI!);  Macambrara  to  Zampalma,  900  m, 
ground  fern,  Exell  297  (BM!);  Tras-os-Montes,  Gama  s.n.  (COI!);  Ribeira 
Peixe,  140  m,  frequent,  Lains  e  Silva  268  (LISU!);  Pico,  1500  m,  Mann  s.n. 
(K!-type);  Lagoa  Amelia,  1400  m,  Monod  11748  (BM!);  Calvario,  1200  m, 
Monod  11814  (BM!);  between  Quinas  Altas  and  Monte  Quinas,  Monod 
12018  (BM!);  s.l.,  F.A.E.  (Moller)  41  p.p.  (BM!,  COI!,  LISU!,  P!). 

DISTRIBUTION.     Bioko,  Annobon. 

Pneumatopteris  venulosa  (Kuntze)  Holttum  in  Blumea  21:  315 
(1973);  Benl  in  Acta  Bot.  Barcinon.  38:  59  (1988). 

Dryopteris  venulosa  Kuntze,  Revis.  gen.  pi.  2:  814  (1891);  Alston  in 

Exell,  Cat.  Vase.  PL  S.  Tome:  66  (1944). 
Cyclosorus  elatus  (Mett.  ex  Kuhn)  Alston  in  Bol.  Soc.  Brot.,  ser.  2, 

30:  13  (1956);  Alston  in  Exell  in  Bull.  Inst.  Franc.  Afrique  Noire 

ser.  A,  21:  446  (1959). 

SAO  TOME.  Boa  Entrada,  Chevalier  13680  (P!);  Macambrara,  primary 
forest,  1050-1200  m,  ground  fern,  Exell  242  (BM!,  COI!);  Lagoa  Amelia, 
1400  m,  Monod  1 1750  (BM!);  s.l.,  Don  s.n.  (BM-not  found),  F.A.E.  (Moller) 
42  p.p.  (K  also  Moller  39!,  P!). 
PRINCIPE.  Esperan9a,  Navel  145  (LISU!,  P!). 

DISTRIBUTION.     Equatorial  Guinea  (continent,  Bioko  and  Annobon). 

Pseudophegopteris  Ching 

Pseudophegopteris  henriquesii  (Baker)  Holttum  in  Blumea  17:  15 
(1969);  Holttum  in  J.  S.  African  Bot.  40:  128  (1974). 

Polypodium  henriquesii  Baker  in  Henriq.  in  Bol.  Soc.  Brot.  4:  154, 
t.l  (1887). 


63 

Dryopteris  henriquesii  (Baker)  C.  Chr.,  Index  filic.:  270  (1905); 

Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  62  (1944). 
Thelypteris  henriquesii  (Baker)  Tardieu  in  Notul.  Syst.  (Paris)  14: 

344(1953). 
Macrothelypteris  henriquesii  (Baker)  Pic.  Serm.  in  Webbia  23:  179 

(1968). 

SAO  TOME.  Tras-os-Montes,  Gama  s.n.  (COI!);  Nova  Moca,  Moller  s.n. 
(COI!),  750  m,  Quintas  1464  (COI!,  LISJC!,  P!);  Macambrard,  1250  m, 
Moller  s.n.  (COI!);  Sao  Nicolau,  waterfall,  Rozeira  162  (PO!);  Trds-os- 
Montes,  Calvario,  Rozeira  562  (PO!);  s.l.,  F.A.E.  (Moller)  49  (BM!,  K!-also 
Moller  45,  type,  LISU!,  P!),  Moller  1  (BM!). 

DISTRIBUTION.     Endemic. 

Sphaerostephanos  J.  Sm. 

Sphaerostephanos  elatus  (Bojer)  Holttum  in  J.  5.  African  Bot.  40: 
167(1974). 

subsp.  thomensis  Holttum  in  J.  S.  African  Bot.  40:  167  (1974). 

Thelypteris  mauritiana  (Fee)  C.F.  Reed  in  Phytologia  17:  291 
(1968)  subsp.  thomensis  (Holttum)  Viane  in  Bull.  Soc.  Roy.  Bot. 
Belg.  118:54(1985). 

SAO  TOME.  S.l.,  F.A.E.  (Moller)  41  p.p.  (COI!,  P!-type). 
DISTRIBUTION.     Endemic. 

NOTE.  This  taxon  has  not  been  recorded  in  the  islands  since  the 
nineteenth  century. 


Vittariaceae 

Anthrophyum  Kaulf. 

Anthrophyum  immersum  (Bory  ex  Willd.)  Mett.  ex  Hook,  in 
Hook.  &  Baker,  Syn.fiL:  393  (1868);  Alston  in  Exell  in  Bull.  Inst. 
Franc.  Afrique  Noire  ser.  A,  21:  442  (1959). 

SAO  TOME.  Monte  Verde,  secondary  forest,  10-150  m,  Lejoly  95/86 
(LISC!);  between  Estacao  Sousa  and  Pico,  primary  forest,  1700  m,  rare 
epiphyte,  Matos  7553  (LISC!);  Cruzeiro,  300  m,  Thorold  2064  (BM!). 
PRINCIPE.  Pico,  950  m,  Thorold  2081  (BM!). 

DISTRIBUTION.     Also  in  Liberia  and  Mascarene  Is. 

Anthrophyum  mannianum  Hook.,  Sec.  cent,  ferns:  t.  73  (1860); 
Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  87  (1944);  Benl  in  Acta 
Bot.  Barcinon.  33:  6  (1982). 

SAO  TOME.  Pico,  light  woodland,  2000  m,  epiphyte,  Exell  340  (BM!);  near 
Rio  Campos,  primary  forest,  epiphyte,  Matos  &  Van  Essche  7441  (LISC!); 
between  Sao  Pedro  and  Lagoa  Amelia,  1250  m,  Moller  s.n.  (COI!); 
Macambrara,  1 200  m,  epiphyte,  Moller  s.n.  (COI!);  Sao  JoaodosAngolares, 
200  m,  Quintas  54A  (K!),  Quintas  s.n.  (COI!);  Lagoa  Amelia,  epiphyte, 
Rozeira  1097  (PO!);  s.l.,  F.A.E.  (Moller  &  Quintas)  60  (BM!,  COI!,  K!, 
LISJC!,  LISU!),  1250  m,  Moller  54  (K!). 

DISTRIBUTION.    Widespread  in  tropical  Africa. 

Vittaria  Sm. 

Vittaria  guineensis  Desv.  in  Ges.  Naturf.  Freunde  Berlin  Mag. 
Neuesten  Entdeck.  Gesammten  Naturk.  5:  325  (1811);  Alston  in 
Exell,  Cat.  Vase.  PL  S.  Tome:  86  (1944);Tardieu-Blot  inAubrev., 
Fl.  Cameroun  3:  124  (1964);  Benl  in  Acta  Bot.  Barcinon.  33:  7 
(1982). 

SAO  TOME.  SW  region,  Chevalier  14598  (P!);  Calvario,  970  m,  epiphyte, 


64 


E.  FIGUEIREDO 


Espi'rito  Santo5Q59  (LISC!,  LISJC!);  between  LagoaAmelia  and  Esperanca, 
1300  m,  epiphyte,  Espi'rito  Santo5\56  (LISC!,  LISJC!);  between  Sao  Carlos 
and  Bom  Sucesso,  secondary  forest,  Espi'rito  Santo  5 1 90  (LISJC!);  Sao  Joao 
dosAngolares,  littoral,  F.A.E.  (Newton)  61  p.p.  (BM!),  80  m,  F.A.E.  (Quintas) 
61  p.p.  (BM!,  COI!,  K!-mixed  with  Barter  1908,  LISU!),  Newton  s.n.  p.p. 
(COI!,  P!);  Ilheu  das  Rolas,  Greeffs.n.  (B?);  between  Pico  and  Monte  Figo, 
6  km  S.  of  Ponta  Figo,  mountain  rainforest,  1200-1800  m,  epiphyte,  Lejoly 
95/6  (LISC!);  W.  of  Pico,  870-1225  m,  Monod  1 1906  (BM!);  Macambrara, 
1200  m,  Quintas  s.n.  (COI!);  Ribeira  Peixe,  between  Vila  Aida  and  main 
house,  epiphyte,  Rozeira  291  (PO!). 

PRINCIPE.  Pico  Papagaio,  primary  forest,  450  m,  epiphyte,  Exell  719 
(BM!,  COI!);  Oque  Caspar,  Newton  25  p.p.  (BM!);  s.l.,  Barter  1908  (K!- 
mixed  with  F.A.E.  61). 

DISTRIBUTION.     W.  tropical  Africa  to  Uganda. 

Vittaria  owariensis  Fee,  Mem.foug.  3:  2 1 ,  t.  3,  fig.  2  ( 1 852);  Alston 
in  Exell,  Cat.  Vase.  PL  S.  Tome:  87  (1944). 

SAO  TOME.  Sao  Joao  dosAngolares,  F.A.E.  (Newton)6\  p.p.  (BM!),  F.A.E. 
(Quintas)  61  p.p.  (BM!,  P!),  Newton  s.n.  p.p.  (P!),  Quintas  s.n.  (COI!);  Vila 
Concei9§o,  100  m,  epiphyte,  Matos  7716  (LISC!);  Rio  Caue,  epiphyte, 
Rozeira  30  (PO!);  Rio  Manuel  Jorge,  between  Pinheira  and  Almas,  epiphyte, 
Rozeira  83  (PO!);  s.l.,  Seabra  s.n.  (COI!). 

PRINCIPE.  Oque  Caspar,  Newton  25  p.p.  (BM!,  COI!);  Porto  Real,  Rose 
467  (P!);  Infante  D.  Henrique,  at  the  base  of  Dois  Irmaos,  epiphyte,  Rozeira 
480  (PO!);  s.l.,  Welwitsch  13  (BM-not  found,  K!,  LISU!). 

DISTRIBUTION.    W.  tropical  Africa. 


DOUBTFUL  RECORDS 

Asplenium  hemitomum  Hieron.  in  Engl.  in  Bot.  Jahrb.  Syst.  46: 
365  (1911);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  78  (1944); 
Benl  in  Acta  Bot.  Barcinon.  40:  20  (1991). 

PRINCIPE  ? 

NOTE.  Alston  (I.e.)  recorded  this  species  based  on  the  collection 
Barter  1896a  (K!)  from  Principe.  Barter  1896  is  a  mixed  collec- 
tion, containing  A.  hemitomum  and  A.  eurysorum.  Since  Barter 
also  collected  A.  hemitomum  in  Bioko  it  is  possible  that  a  specimen 
of  this  species,  from  Bioko,  was  attached  to  the  herbarium  sheet 
with  A.  eurysorum  from  Principe. 

Ceratopteris  cornuta  (P.  Beauv.)  Lepr.  in  Ann.  Sci.  Nat.  (Paris)  19: 
103,  t.  4A  (1830);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  92 
(1944). 

PRINCIPE  ?  Barter  (Kuhn,  1868). 

NOTE.  No  specimens  were  seen  at  BM,  K  or  P.  However,  at  BM 
there  is  a  specimen  collected  by  Barter  during  the  Niger  Expedition, 
unlocalized,  which  could  have  been  collected  in  Principe. 

Dryopteris  manniana  (Hook.)  C.  Chr.,  Index  filic.:  276  (1905); 
Benl  in  Acta  Bot.  Barcinon.  40:  38  (1991). 

SAO  TOME  ? 

NOTE.     No  specimens  were  seen. 

Lastreopsis  nigritiana  (Baker)  Tindale  in  Contr.  New  South  Wales 
Natl.  Herb.  3:  245  (1963);Tardieu-Blot  inAubrev.,  Fl.  Cameroun 
3:  280  (1964);  Benl  in  Acta  Bot.  Barcinon.  40:  51  (1991). 

Polypodium  nigritianum  Baker  in  Hook.  &  Baker,  Syn.  fil.:  313 
(1867). 


PRINCIPE  ? 

NOTE.  No  specimens  were  seen.  Although  there  are  references  in 
the  literature  to  the  occurrence  of  this  species  in  Principe,  Tindale 
(1965)  did  not  record  it  for  Sao  Tome  and  Principe  in  his  monograph 
of  the  genus. 

Lygodium  microphyllum  (Cav.)  R.  Br.,  Prodr.:  162  (1810). 

Lygodium  scandens  sensu  auct.  non  (L.)  Sw.  in  /.  Bot.  (Schrader) 
1800(2):  106  (1801);  Alston  in  Exell,  Cat.  Vase.  PL  S.  Tome:  93 

(1944). 

PRINCIPE  ?  (Barter,  1860). 

NOTE.  No  specimens  were  seen  at  BM  or  K.  There  are  specimens 
of  Lygodium  microphyllum  collected  by  Barter  in  1859  during  the 
Niger  Expedition,  some  of  which  are  unlocalized  and  could  have 
been  collected  on  the  island.  Being  a  lowland  species,  L. 
microphyllum  might  be  extinct  due  to  the  extensive  cultivation  of 
these  areas. 

Ophioglossum  gramineum  Willd.  in  Schrift.  Akad.  Erfurt:  18,  t.l, 
f.l  (1802);  Tardieu-Blot  in  Aubrev.,  Fl.  Cameroun  3:  46  (1964). 

SAO  TOME  ? 

NOTE.     No  specimens  seen. 

Trichomanes  africanum  H.  Christ  in  J.  Bot.  (Morot),  ser.  2,  2:  21 
(1909);  Benl  in  Acta  Bot.  Barcinon.  32:  23  (1980). 

[Vandenboschia  africana  (H.  Christ)  G.  Kunkel  in  Nova  Hedwigia 
6:213(1963)] 

SAO  TOME  ? 

NOTE.     No  specimens  seen. 


ACKNOWLEDGEMENTS.  Thanks  are  due  to  Fundagao  Calouste  Gulbenkian 
for  a  research  grant.  I  would  also  like  to  thank  the  directors  of  BM,  K,  and  P 
for  providing  all  the  facilities  during  my  stay  in  their  herbaria,  and  also 
Frederic  Badre,  Gustav  Benl,  Josephine  Camus,  Peter  Edwards,  Elisa 
Folhadela,  Mary  Gibby,  Bob  Johns,  and  Alison  Paul. 


REFERENCES 

Alston,  A.H.G.  1944.  Pteridophyta.  In  A.  Exell,  Catalogue  of  the  vascular  plants  ofS. 
Tome  (with  Principe  and  Annobon):  57-99.  London. 

1956a.  Pteridophyta.  In  A.  Exell,  Supplement  to  the  catalogue  of  the  vascular 

plants  ofS.  Tome  (with  Principe  and  Annobon):  7-9.  London. 

1956/7.  New  African  ferns.  Boletim  da  Sociedade  Broteriana,  serie  2,  30:  5-27. 

1958.  Pteridophyta.  In  A.  Exell  &A.  Rozeira,  Aditamento  a  Flora  das  ilhas  de  Sao 

Tome  e  Principe.  Conferencia  Internacional  dos  Africanistas  Ocidentais  3:  78-81. 

1959.  Pteridophyta.  In  A.  Exell,  Additions  to  the  flora  of  S.  Tome  and  Principe. 

Bulletin  de  I'lnstitut  Franfaise  d'Afrique  Noire,  serie  A,  21:  440-448. 

Barter,  C.  1 860.  Letter  from  Mr  Charles  Barter,  natural  history  collector  to  the  Niger 
Expedition.  Journal  of  the  Proceedings  of  the  Linnean  Society,  Botany.  4:  17-23. 

Benl,  G.  1 978.  The  Pteridophyta  of  Fernando  Po  I.  Acta  Botanica  Barcinonensia  31:  1- 
31. 

1980.  The  Pteridophyta  of  Fernando  Po  II.  Acta  Botanica  Barcinonensia  32:  1- 

34. 

1982.  The  Pteridophyta  of  Fernando  Po  III.  Acta  Botanica  Barcinonensia  33:  1- 

46. 

1 988.  The  Pteridophyta  of  Bioko  (Fernando  Po)  IV.  Acta  Botanica  Barcinonensia 

38:  1-69. 

199 1 .  The  Pteridophyta  of  Bioko  (Fernando  Po)  V.  Acta  Botanica  Barcinonensia 

40:  1-106. 

Dubuisson,  J.-Y.  1996.  Evolutionary  relationships  within  the  genus  Trichomanes 
sensu  lato  (Hymenophyllaceae)  based  on  anatomical  and  morphological  characters 
and  a  comparison  with  rbcL  necleotide  sequences:  preliminary  results.  In  J.  Camus 
et  al.  (Eds)  Pteridology  in  perspective:  285-287.  Kew. 


PTERIDOPHYTES  OF  SAO  TOME  AND  PRINCIPE 


65 


Exell,  A.  1944.    Catalogue  of  the  vascular  plants  of  S.  Tome  (with  Principe  and 

Annobon).  London. 
Exell,  A.  &  Rozeira,  A.  1958.  Aditamento  a  flora  das  ilhas  de  Sao  Tome  e  Principe. 

Conferencia  International  dos  Africanistas  Ocidentais  3:  77-91. 
Figueiredo,  E.  1994.  Diversity  and  endemism  of  angiosperms  in  the  Gulf  of  Guinea 

islands.  Biodiversity  and  Conservation  3:  785-793. 
Kramer,  K.  1972.  The  Lindsaeoid  ferns  of  the  Old  World-IX  Africa  and  its  islands. 

Bulletin  du  Jardin  Botanique  National  de  Belgique  42:  305-345. 
Kuhn,  M.  1 868.  Filices  africanae.  Lipsiae. 
Nooteboom,  H.P.  1994.  Notes  on  Davalliaceae.  II.  A  revision  of  the  genus  Davallia. 


Blumea39:  151-214. 

Pichi  Sermolli,  R.E.G.  1969.   Adumbratio  Florae  Aethiopicae.  16.   Marattiaceae. 
Webbia  23:  329-351. 

1985.  On  the  taxonomy  and  nomenclature  of  some  species  of    Ctenitis 

(Aspidiaceae)  from  Tropical  Africa.  Webbia  39:  1-28. 

1991.  On  the  taxonomy  and  nomenclature  of  some  species  from  Tropical 

Africa  of  the  genus  Triplophyllum  Holttum  (Dryopteridaceae).  Webbia  45:  117- 
135. 

Tindale,  M.D.  1 965.  A  monograph  of  the  genus  Lastreopsis  Ching.  Contributions  from 
the  New  South  Wales  National  Herbarium  3:  249-339. 


SYSTEMATIC  INDEX 


Accepted  names  are  in  roman  and  synonyms  in  italic. 


Acrostichum  aureum  L.  42,  43 

Adiantum  mettenii  Kuhn  43 

Adiantum  philippense  L.  43 

Adiantum  raddianum  C.  Pres  42,  43 

Adiantum  soboliferum  auct.  43 

Adiantum  vogelii  Mett.  ex  Keyserl.  45 

Alsophila  camerooniana  van  camerooniana  auct.  51 

Alsophila  manniana  (Hook.)  R.M.  Tryon  5 1 

Alsophila  welwitschil  (Hook.)  R.M.  Tryon    51 

Anapeltis  lycopodioides  van  owariensis  (Desv.) 

Benl  59 

Anisosorus  occidentalis  (Baker)  C.  Chr.  52 
Anthrophyum  immersum  (Bory  ex  Willd.)  Mett.  ex 

Hook.  63 

Anthrophyum  mannianum  Hook.  63 
Arthropteris  monocarpa  (Cordem.)  C.  Chr.  58 
Arthropteris  obllterata  auct.  58 
Arthropteris  orientalis  (J.F.  Gmel.)  Posth.  58 
Arthropteris  palisotii  (Desv.)  Alston  58 
Aspidium  fraternum  Mett.  ex  Kuhn  47 
Aspidium  nigritlanum  Mett.  ex  Kuhn  46 
Aspidium  subquinquefidum  P.  Beauv.  47 
Asplenium  aethiopicum  (Burm.f.)  Bech.  48 
Asplenium  africanum  Desv.  48 
Asplenium  anisophyllum  Kunze  48 
Asplenium  barteri  Hook.  48,  50 
Asplenium  biafranum  Alston  &  F.  Ballard  48 
Asplenium  cuneatum  Lam.  48 
Asplenium  currorii  Hook.  48 
Asplenium  dregeanum  subsp.  brachypterum  (Kunze 

ex  Houlston  &  T.  Moore)  Pic.  Serm.  48 
Asplenium  emarginatum  P.  Beauv.  48 
Asplenium  erectum  var.  usambarense  (Hieron.) 

Schelpe  48 

Asplenium  eurysorum  Hieron.  49,  64 
Asplenium  exhaustum  (H.  Christ)  Alston  49 
Asplenium  formosum  Willd.  49 
Asplenium  friesiorum  C.  Chr.  42,  49 
Asplenium  geppii  Carruth.  48 
Asplenium  hemitomum  Hieron.  64 
Asplenium  hypomelas  Kuhn  49 
Asplenium  inaequilaterale  Bory  ex  Willd.  49 
Asplenium  lividum  Mett.  ex  Kuhn  49 
Asplenium  longicauda  Hook.  49 
Asplenium  megalura  var.  molleri  (Hieron.) 

Tardieu  49 

Asplenium  molleri  Hieron.  49 
Asplenium  nigritianum  Hook.  49,  50 
Asplenium  paucijugum  F.  Ballard  50 
Asplenium  protensum  auct.  49 
Asplenium  quintasil  Gand.  49 
Asplenium  sandersonii  Hook.  50 
Asplenium  unilaterale  Lam.  50 
Asplenium  variabile  Hook.  50 
Asplenium  variabile  var.  paucijugum  (F.  Ballard) 

Alston  50 

Athyrium  newtonii  (Baker)  Diels  50 
Belvisia  spicata  (L.f.)  Mirb.  ex  Copel.  59 
Blechnum  attenuatum  (Sw.)  Mett.  51 
Blotiella  currorii  (Hook.)  R.M.  Tryon  51 


Blotiella  glabra  (Bory)  R.M.  Tryon  52 
Blotiella  mannii  (Baker)  Pic.  Serm.  52 
Bolbitis  acrostichoides  (Afzel.  ex  Sw.)  Ching  56 
Bolbitis  auriculata  (Lam.)  Ching  56 
Bolbitis  fluviatilis  (Hook.)  Ching  56 
Callipteris  prolifera  (Lam.)  Bory  50 
Ceratopteris  cornuta  (P.  Beauv.)  Lepr.  64 
Christella  dentata  (Forssk.)  Brownsey  &  Jermy  62 
Christella  hispidula  (Decne.)  Holttum  62 
Coniogramme  africana  Hieron.  54 
Ctenitis  attenuata  Pic.  Serm.  47 
Ctenitis  buchholzii  (Kuhn)  Alston  47 
Ctenitis  cirrhosa  (Schum.)  Ching  42,  45 
Ctenitis  fraterna  (Mett.)  Tardieu  47 
Ctenitis  lanuginosa  (Willd.  ex  Kaulf.)  Copel.  45 
Ctenitis  nigritiana  (Mett.  ex  Kuhn)  Alston  46 
Cyathea  camerooniana  var.  currorii  Holttum  5 1 
Cyathea  manniana  Hook.  51 
Cyathea  welwitschii  Hook.  5 1 
Cyclophorus  spissus  var.  continentalis  Hieron.  ex 

Engl.  60 

Cyclosorus  afer  (H.  Christ)  Ching  63 
Cyclosorus  elatus  (Mett.  ex  Kuhn)  Alston  63 
Cyclosorus  oppositifolius  (Hook.)  Tardieu  63 
Cyclosorus  striatus  (Schum.)  Ching  62 
Davallia  chaerophylloides  (Poir.)  Steud.  51 
Davallia  denticulata  var.  denticulata  (Burm.f.)  Mett. 

ex  Kuhn  51 

Davallia  repens  (L.f.)  Kuhn  51 
Dicranopteris  linearis  (Burm.f.)  Underw.  53 
Didymochlaena  truncatula  (Sw.)  J.  Sm.  45 
Diplazium  arborescens  (Bory)  Sw.  50 
Diplazium  proliferum  (Lam.)  Thouars  50 
Drynaria  laurentii  (H.  Christ  ex  De  Wild.  &  T. 

Durand)  Hieron.  59 
Drynaria  volkensii  Hieron.  59 
Dryopteris  crinobulbon  (Hook.)  C.  Chr.  45 
Dryopteris  dentata  (Forssk.)  C.  Chr.  62 
Dryopteris  fragilis  (Baker)  C.  Chr.  62 
Dryopteris  henriquesii  (Baker)  C.  Chr.  63 
Dryopteris  lanuginosa  (Willd.  ex  Kaulf.)  C.  Chr.  45 
Dryopteris  manniana  (Hook.)  C.  Chr.  64 
Dryopteris  nigritiana  (Mett.  ex  Kuhn)  Kuntze  46 
Dryopteris  oligantha  auct.  45 
Dryopteris  oppositifolia  auct.  63 
Dryopteris  oppositifolia  (Hook.)  C.  Chr.  63 
Dryopteris  pentheri  (Krasser)  C.  Chr.  45 
Dryopteris  protensa  (Afzel.  ex  Sw.)  C.  Chr.  47 
Dryopteris  quadrangularis  (Fee)  Alston  62 
Dryopteris  securidiformis  (Hook.)  C.  Chr.  47 
Dryopteris  securidiformis  var.  nana  Bonap.  47 
Dryopteris  striata  (Schum.)  C.  Chr.  62 
Dryopteris  variabilis  var.  barteri  (Hook.)  Alston  47 
Dryopteris  venulosa  Kuntze  63 
Elaphoglossum  acrostichoides  (Hook.  &  Grev.) 

Schelpe  52 

Elaphoglossum  aubertii  (Desv.)  T.  Moore  53 
Elaphoglossum  chevalieri  H.  Christ  53 
Elaphoglossum  clarenceanum  auct.  53 
Elaphoglossum  isabelense  Brause  53 


Elaphoglossum  petiolatum  subsp.  salicifolium  (Willd. 

ex  Kaulf.)  Schelpe  53 
Elaphoglossum  salicifolium  (Willd.  ex  Kaulf.) 

Alston  53 

Gleichenia  linearis  (Burm.f.)  C.B.  Clarke  53 
Gonocormus  mannii  (Hook,  ex  Hook.  &  Baker)  G. 

Kunkel  55 

Grammitis  ebenina  (Maxon)  Tardieu  53 
Grammitis  molleri  (Baker)  Schelpe  42,  53 
Grammitis  nigrocincta  Alston  42,  53 
Grammitis  sp.  53 

Grammitis  tomensis  Schelpe  42,  54 
Histiopteris  incisa  (Thunb.)  J.  Sm.  52 
Humata  repens  (L.f.)  Diels  51 
Huperzia  mildbraedii  (Herter)  Pic.  Serm.  57 
Huperzia  ophioglossoides  (Lam.)  Rothm.  42,  57 
Huperzia  phlegmaria  (L.)  Rothm.    57 
Huperzia  verticillata  (L.f.)  Trevis.  57 
Huperzia  warneckei  (Herter  ex  Nessel)  Pic.  Serm.  57 
Hymenolepis  spicata  (L.f.)  C.  Presl  59 
Hymenophyllum  capillare  Desv.  54 
Hymenophyllum  ciliatum  var.  boryanum  (Willd.) 

Mett.  ex  Kuhn  54 

Hymenophyllum  hirsutum  (L.)  Sw.  54 
Hymenophyllum  kuhnii  C.  Chr.  54 
Hymenophyllum  mildbraedii  (Brause  ex  Brause  & 

Hieron.)  Alston  55 
Hymenophyllum  polyanthos  var.  kuhnii  (C.  Chr.) 

Schelpe  54 
Hymenophyllum  polyanthos  var.  mildbraedii  (Brause 

ex  Brause  &  Hieron.)  Schelpe  42,  55 
Hymenophyllum  splendidum  Bosch  55 
Hypolepis  sparsisora  (Schrad.)  Kuhn  52 
Lastreopsis  aff.  currori  (Mett.  ex  Kuhn)  Tindale  46 
Lastreopsis  nigritiana  (Baker)  Tindale  64 
Lomariopsis  guineensis  (Underw.)  Alston  56 
Lomariopsis  hederacea  Alston  56 
Lomariopsis  hederacea  auct.  56 
Lomariopsis  nigrescens  Holttum  56 
Lomariopsis  warneckei  (Hieron.)  Alston  42,  56 
Lonchitis  currorii  (Hook.)  Mett.  ex  Kuhn  51 
Lonchitis  glabra  Bory  52 
Lonchitis  gracilis  Alston  52 
Lonchitis  mannii  (Baker)  Alston  52 
Lonchitis  occidentalis  Baker  52 
Loxogramme  lanceolata  (Sw.)  C.  Presl  56 
Lycopodiella  cernua  (L.)  Pic.  Serm.  57 
Lycopodium  cemuum  L.  57 
Lycopodium  clavatum  L.  42,  57 
Lycopodium  mildbraedii  Herter  57 
Lycopodium  phlegmaria  L.  57 
Lycopodium  verticillatum  L.f.  57 
Lycopodium  warneckei  (Herter  ex  Nessel)  Alston  57 
Lygodium  microphyllum  (Cav.)  R.  Br.  64 
Lygodium  scandens  auct.  64 
Macrothelypteris  henriquesii  (Baker)  Pic.  Serm.  63 
Marattia  fraxinea  Sm.  57,  58 
Marattia  robusta  Alston  57,  58 
Metathelypteris  fragilis  subsp.  guineensis  Benl  62 
Microgonium  erosum  (Willd.)  C. Presl  55 


66 


E.  FIGUEIREDO 


Microgramma  owariensis  (Desv.)  Alston  59 
Microlepia  speluncae  (L.)  T.  Moore  42,  52 
Microsorum  punctatum  (L.)  Copel.  59 
Microtrichomanes  digitatum  (Sw.)  Copel.  55 
Nephmdium  subquinquefidum  var.  elongatum 

Hook.  47 

Nephrolepis  biserrata  (Sw.)  Schott  58 
Nephrolepis  pumicicola  F.  Ballard  58 
Nephrolepis  undulata  (Afzel.  ex  Sw.)  J.  Sm.  58 
Odontosoria  chinensis  var.  divaricata  H.  Christ  52 
Oleandra  annetii  auct.  58 
Oleandra  annetii  Tardieu  59 
Oleandra  distenta  Kunze  58 
Ophioglossum  costatum  R.  Br.  59 
Ophioglossum  gramineum  Willd.  64 
Ophioglossum  reticulatum  L.  59 
Pellaea  doniana  Hook.  62 

Phymatosorus  scolopendria  (Burm.f.)  Pic.  Serm.  60 
Pityrogramma  calomelanos  var.  calomelanos  (L.) 

Link  42,  54 

Platycerium  stemaria  (P.  Beauv.)  Desv.  60 
Pleopeltis  excavata  (Bory  ex  Willd.)  Sledge  60 
Pleopeltis  macrocarpa  (Bory  ex  Willd.)  Kaulf.  60 
Pneumatopteris  afra  (C.  Chr.)  Holttum  63 
Pneumatopteris  oppositifolia  (Hook.)  Holttum  63 
Pneumatopteris  venulosa  (Kuntze)  Holttum  63 
Polypodium  cultratum  var.  elasticum  (Bory  ex  Willd.) 

Baker  54 

Polypodium  ebeninum  Maxon  53 
Pol\podium  henriquesii  Baker  63 
Polypodium  irioides  Poir.  59 
Polypodium  lycopodioides  auct.  59 
Polypodium  molleri  Baker  53 
Polypodium  nigritianum  Baker  64 
Polypodium  oosorum  Baker  54 
Polypodium  phymatodes  L.  60 
Polypodium  preussii  Hieron.  60 
Polypodium  villosissimum  Hook.  54 
Pseudophegopteris  henriquesii  (Baker)  Holttum  63 


Psilotum  nudum  (L.)  P.  Beauv.  60 

Pteridium  aquilinum  (L.)  Kuhn  52 

Pteris  atrovirens  Willd.  60,  61 

Pteris  atmvirens  auct.  61 

Pteris  biaurita  auct.  61 

Pteris  burtonii  Baker  60,  61 

Pteris  catoptera  auct.  61 

Pteris  communata  auct.  6 1 

Pteris  dentata  subsp.  flabellata  (Thunb.)  Runemark  61 

Pteris  linearis  Poir.  61 

Pteris  paucipinnata  Alston  61 

Pteris  pteridioides  (Hook.)  F.  Ballard  61 

Pteris  quadriaurita  auct.  61 

Pteris  similis  Kuhn  61 

Pteris  togoensis  Hieron.  61 

Pteris  tripartita  Sw.  54,  61 

Pteris  vittata  L.  42,  61 

Pyrrosia  lanceolata  (L.)  Farw.  60 

Selaginella  cathedrifolia  Spring  61 

Selaginella  mannii  Baker  61 

Selaginella  molleri  Hieron.  62 

Selaginella  molliceps  Spring  62 

Selaginella  monodii  Alston  42,  62 

Selaginella  myosurus  (Sw.)  Alston  62 

Selaginella  squarrosa  Baker  42,  62 

Selaginella  thomensis  Alston  62 

Selenodesmium  rigidum  (Sw.)  Copel.  56 

Sphaerostephanos  elatus  subsp.  thomensis 

Holttum  42,  63 
Sphenomeris  chinensis  var.  divaricata  (H.  Christ) 

Kramer  52 
Sphenomeris  chusana  var.  divaricata  (H.  Christ) 

Tardieu  52 
Stenoloma  chinense  var.  divaricatum  (H.  Christ) 

Alston  52 

Tectaria  angelicifolia  (Schum.)  Copel.  46 
Tectaria  buchholzii  (Kuhn)  Copel.  47 
Tectaria  camerooniana  (Hook.)  Alston  46 
Tectaria  fernandensis  (Baker)  C.  Chr.  47 


Thelypteris  henriquesii  (Baker)  Tardieu  63 
Thelypteris  mauritiana  subsp.  thomensis  (Holttum) 

Viane  63 

Trichomanes  africanum  H.  Christ  64 
Trichomanes  borbonicum  Bosch  55 
Trichomanes  chamaedrys  Taton  55 
Trichomanes  crispiforme  Alston  55 
Trichomanes  cupressoides  Desv.  56 
Trichomanes  digitatum  Sw.  55 
Trichomanes  erosum  Willd.  55 
Trichomanes  giganteum  auct.  55 
Trichomanes  mannii  Hook,  ex  Hook.  &  Baker  55 
Trichomanes  melanotrichum  Schltdl.  55 
Trichomanes  pyxidiferum  var.  melanotrichum 

(Schltdl.)  Schelpe  55 
Trichomanes  radicans  Sw.  55 
Trichomanes  rigidum  Sw.  56 

Triplophyllum  attenuatum  (Pic.  Serm.)  Pic.  Serm.  47 
Triplophyllum  buchholzii  (Kuhn)  Holttum  47 
Triplophyllum  fraternum  var.  elongatum  (Hook.) 

Holttum  47 

Triplophyllum  heudelotii  Pic.  Serm.  47 
Triplophyllum  jenseniae  (C.  Chr.)  Holttum  42,  47 
Triplophyllum  principis  Holttum  47 
Triplophyllum  protensum  (Afzel.  ex  Sw.)  Holttum  47 
Triplophyllum  securidiforme  (Hook.)  Holttum  47 
Triplophyllum  securidiforme  var.  nanum  (Bonap.) 

Holttum  47 
Triplophyllum  subquinquefidum  (P.  Beauv.)  Pic. 

Serm.  47 

Vandenboschia  africana  (H.  Christ)  G.  Kunkel  64 
Vandenboschia  borbonica  (Bosch)  G.  Kunkel  55 
Vandenboschia  pyxidifera  (L.)  Copel.  55 
Vandenboschia  radicans  (Sw.)  Copel.  55 
Vittaria  guineensis  Desv.  63 
Vittaria  owariensis  Fee  64 
Xiphopteris  cultrata  (Willd.)  Schelpe  54 
Xiphopteris  oosora  (Baker)  Alston  42,  54 
Xiphopteris  villosissima  (Hook.)  Alston  54 


Bulletin  of  The  Natural  History  Museum 
Botany  Series 

Earlier  Botany  Bulletins  are  still  in  print.  The  following  can  be  ordered  from  Intercept  (address  on  inside  front  cover).  Where  the  complete  backlist  is  not  shown, 
this  may  also  be  obtained  from  the  same  address. 


Volume  2 

No.  1  New  Himalayan  species  of  Pedicularis  with  special  reference 

to  those  from  the  eastern  Himalaya.  P.C.  Tsoong.  1955.  Pp.  1- 
34.  Facsimile  edition.  £4.35 

No.  2  Mosses  of  Dominica,  British  West  Indies.  E.B.  Bartram.  1955. 
Pp.  37-49.  Mosses  of  the  Ecuadorian  Andes  collected  by  P.R. 
Bell.  E.B.  Bartram.  1955.  Pp.  51-64.  Facsimile  edition.  £4.00 

No.  3          Novitates  Himalaicae-1.  F.  Ludlow  &  W.T.  Stearn.  1956.  Pp. 

65-81,  8  plates,  1 1  figs.  Facsimile  edition.  £5.25 

No.  4          Saxifraga  of  the  Himalaya  1.  Section  Kabschia.  H.  Smith. 

1958.  Pp.  83-129,  14  figs.  £9.65 

No.  5          The  Polypodiaceae  and  Grammitidaceae  of  Ceylon.  W.A. 

Sledge.  1960.  Pp.  131-158,  4  figs.  Facsimile  edition.         £3.90 

No.  6          Allium  and  Milula  in  the  central  and  eastern  Himalaya.  W.T. 

Stearn.  1960.  Pp.  159-191,  4  plates,  10  figs.  Facsimile  edition. 

£4.35 

No.  7          The  identity  of  Isopyrum  aquilegioides.  L.G.  de  Beer  &  W.T. 

Stearn.  1960.  Pp.  193-202,  3  figs.  Facsimile  edition.         £3.15 

No.  8          On  the  geographical  relationships  of  the  angiosperm  flora  of 
New  Guinea.  R.  Good.  1960.  Pp.  203-226,  1  fig.  Facsimile 
edition.  £3.60 

No.  9          Saxifraga  of  the  Himalaya  II.  Some  new  species.  H.  Smith. 

1960.  Pp.  227-260.  9  plates,  17  figs.  Facsimile  edition.     £5.65 

No.  10        New  species  of  Taraxacum  from  the  Himalayan  region.  J.L. 
van  Soest.  1961.  Pp.  261-273,  8  plates.  Facsimile  edition. 

£6.15 

No.  1 1        The  athyriod  ferns  of  Ceylon.  W.A.  Sledge.  1962.  Pp.  275-323. 
3  plates.  Facsimile  edition.  £5.25 

No.  12        The  genus  Epilobium  in  the  Himalayan  region.  PH.  Raven. 

1962.  Pp.  325-382,  7  plates,  13  figs.  Facsimile  edition.     £5.90 

Volume  3 

No.  1  A  revision  of  the  genera  Buchenavia  and  Ramatuella.  A.W. 

Exell  &  C.A.  Stace.  1963.  Pp.  1-46,  5  figs.  Facsimile  edition. 

£5.15 
No.  2          The  diatom  genus  Capartogramma  and  the  identity  of 

Schizostauron.  R.  Ross.  1963.  Pp.  47-92,  2  plates,  38  figs. 

Facsimile  edition.  £5.15 

No.  3          Angiosperms  of  the  Cambridge  Annobon  Island  Expedition. 
A.W.  Exell.  1963.  Pp.  93-1 18,  10  plates.  Facsimile  edition. 

£3.75 

No.  4          A  revision  of  the  genus  Petrorhagia.  P.W.  Ball  &  V.H. 

Hey  wood.  1964.  Pp.  1 19-172,  3  plates,  22  figs.  Facsimile 
edition.  £4.90 

No.  5          Marine  algae  of  Gough  Island.  Y.M.  Chamberlain.  1965.  Pp. 

173-232,  4  plates,  80  figs.  £12.50 

No.  6          The  Ceylon  species  ofAsplenium.  W.A.  Sledge.  1965.  Pp.  233- 
277,  1  plate,  3  figs.  Facsimile  edition.  £5.15 

Volume  4 

No.  1          Cuticular  studies  as  an  aid  to  plant  taxonomy.  C.A.  Stace. 

1965.  Pp.  1-78,  5  plates,  10  figs.  Facsimile  edition.  £7.20 

No.  2  The  genus  Elaphoglossum  in  the  Indian  peninsula  and  Ceylon. 
W.A.  Sledge.  1967.  Pp.  79-96.  Facsimile  edition.  £3.25 

No.  3  Fungi  of  recent  Nepal  expeditions.  F.L.  Balfour-Browne.  1968. 
Pp.  97-141,  4  figs.  Facsimile  edition.  £3.75 


No.  4 

No.  5 

No.  6 
No.  7 


A  synopsis  of  Jamaican  Myrsinaceae.  W.T.  Stearn.  1969.  Pp. 
143-178,  8  plates,  25  figs.  £4.55 

The  Jamaican  species  of  Columnea  and  Alloplectus 
(Gesneriaceae).  W.T.Stearn.  1969.  Pp.  179-236,  8  plates,  29 
figs.  £6.40 

New  or  little  known  Himalayan  species  of  Swertia  and 
Veratrilla  (Gentianaceae).  H.  Smith.  1970.  Pp.  237-258,  16 
plates,  7  figs.  £8.25 

A  survey  of  the  tropical  genera  Oplonia  and  Psilanthele 
(Acanthaceae).  W.T.  Stearn.  1971.  Pp.  259-323,  10  plates,  18 
figs.  £12.40 

Angiosperms  of  the  islands  of  the  Gulf  of  Guinea  (Fernando 
Po,  Principe,  S.  Tome,  and  Annobon).  A.W.  Exell.  1973.  Pp. 
325-411.  £12.50 


Volume  5 

No.  1          The  dryopteroid  ferns  of  Ceylon.  W.A.  Sledge.  1973.  Pp.  1-43, 
4  figs.  £6.90 

No.  2          New  Himalayan  and  Tibetan  species  of  Corydalis 

(Papaveraceae).  F.  Ludlow  &  W.T.  Stearn.  1975.  Pp.  45-69,  15 
plates,  14  figs.  £8.40 

No.  3          The  marine  algae  of  Trinidad,  West  Indies.  W.D.  Richardson. 

1975.  Pp.  71-143,  12  plates,  2  figs.  £13.45 

No.  4  A  revision  of  the  Macaronesian  genus  Argyranthemum  Webb 
ex  Schultz  Bip.  (Compositae-Anthemideae).C.J.  Humphries. 

1976.  Pp.  145-240,  2  plates,  26  figs.  £14.20 

No.  5          Frank  Ludlow  (1885-1972)  and  the  Ludlow-Sherriff  expedi- 
tions to  Bhutan  and  south-eastern  Tibet  of  1933-1950.  W.T. 
Stearn.  1976.  Pp.  243-268,  1  fig.  Reliquiae  botanicae 
himalaicae.  F.  Ludlow.  1976.  Pp.  269-289,  7  plates,  8  figs. 
Facsimile  edition.  £11.10 

No.  6          Studies  in  the  genus  Hypericum  L.  (Guttiferae).  1 .  Infrageneric 
classification.  N.K.B.  Robson.  1977.  Pp.  291-355,  9  figs. 

£14.20 

No.  7          Sphagnales  of  tropical  Asia.  A.Eddy.  1977.  Pp.  357^45,  4 

plates,  17  maps,  25  figs.  £17.80 

Volume  6 

No.  1          The  handwriting  of  Joseph  Banks,  his  scientific  staff  and 

amanuenses.  J.B.  Marshall.  1978.  Pp.  1-85,  62  figs.        £18.30 

No.  2  Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  II.  Phaeophyta.  J.H.  Price,  D.M. 
John  &  G.W.  Lawson.  1978.  Pp.  87-182,  1  fig.  £24.40 

No.  3          The  lichenicolous  Hyphomycetes.  D.L.  Hawksworth.  1979.  Pp. 
183-300,  47  figs.  £24.40 

No.  4          The  species  of  Chisocheton  (Meliaceae).  D.J.  Mabberley.  1979. 
Pp.  301-386,  3  plates,  10  figs.  £24.40 

Volume  7 

No.  1          The  distribution  of  Padina  pavonica  (L.)  Lamour.  (Phaeophyta: 
Dictyotales)  on  British  and  adjacent  European  shores.  J.H. 
Price,  I.  Tittley  &  W.D.  Richardson.  1979.  Pp.  1-67,  3  plates,  2 
figs.  £17.40 

No.  2          Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  111.  Rhodophyta 
(Bangiophyceae).  D.M.  John,  J.H.  Price,  C.A.  Maggs,  G.W. 
Lawson.  1979.  Pp.  69-82,  1  fig.  £5.40 


No.  3          A  revision  of  the  genus  Anacyclus  L.  (Compositae:  Anthemi- 

deae).  C.J.  Humphries.  1979.  Pp.  83-142,  27  figs.  £15.60 

Volume  8 

No.  1          The  Thelypteridaceae  of  Ceylon.  W.A.  Sledge.  Pp.  1-54,  5  figs. 
1981.  £15.15 

No.  2          Studies  in  the  genus  Hypericum  L.  (Guttiferae)  2.  Characters  of 
the  genus.  N.K.B.  Robson.  1981.  Pp.  55-226,  73  figs.     £33.55 

No.  3  A  revision  of  the  lichen  family  Thelotremataceae  in  Sri  Lanka. 
M.E.  Hale,  Jr.  1981.  Pp.  227-332,  20  figs.  £24.80 

No.  4  Vascular  plant  collections  from  the  Tristan  da  Cunha  group  of 
islands.  E.W.  Groves.  Pp.  333^20,  33  figs.  £21.40 

Volume  9 

No.  1  The  lichenicolous  Coelomycetes.  D.L.  Hawksworth.  1981.  Pp. 
1-98,  36  figs.  £22.70 

No.  2          The  genus  Callithamnion  (Rhodophyta:  Ceramiaceae)  in  the 
British  Isles.  PS.  Dixon  &  J.H.  Price.  1981.  Pp.  99-141,5  figs. 

£12.50 

No.  3          Parmelia  subgenus  Amphigymnia  (lichens)  in  East  Africa.  H. 

Krog  &  T.F.V.  Swinscow.  1981.  Pp.  143-231,  31  figs.      £21.05 

No.  4          The  genus  Selaginella  in  tropical  South  America.  A.H.G. 

Alston,  A.C.  Jermy  &  J.M.  Rankin.  1981.  Pp.  233-330,  18  figs. 

£23.05 

Volume  10 

No.  1          Taxonomic  studies  in  the  Labiatae  tribe  Pogostemoneae.  J.R. 

Press.  1982.  Pp.  1-89,  33  figs.  £21.85 

No.  2          The  typification  of  Hudson's  algae:  a  taxonomic  and 

nomenclatural  reappraisal.  L.M.  Irvine  &  PS.  Dixon.  1982.  Pp. 
91-105.  £5.40 

No.  3          Seaweeds  of  the  Faroes.  Various  authors.  1982.  Pp.  107-225, 

13  figs.  £27.15 

No.  4  The  lichen  genus  Steinera.  A.M.  Henssen  &  P.W.  James.  1982. 
Pp.  227-256,  24  figs.  £9.70 

Volume  11 

No.  1          The  algae  of  Lightfoot's  Flora  scotica.  PS.  Dixon.  1983.  Pp. 

1-15,  2  figs.  £5.55 

No.  2  A  taxonomic  study  of  the  lichen  genus  Micarea  in  Europe.  B.J. 
Coppins.  1983.  Pp.  17-214,  57  figs,  28  maps.  £37.75 

No.  3          The  hepatics  of  Sierra  Leone  and  Ghana.  E.W.  Jones  &  A.J. 

Harrington.  1983.  Pp.  215-289,  8  figs.  £18.15 

No.  4  Studies  in  the  Corallinaceae  with  special  reference  to  Fosliella 
and  Pneophyllum  in  the  British  Isles.  Y.M.  Chamberlain.  1983. 
Pp.  291^63,  89  figs.  £33.45 

Volume  12 

No.  1  A  revision  of  the  Morinaceae  (Magnoliophyta-Dipsacales). 

M.J.  Cannon  &  J.F.M.  Cannon.  1984.  Pp.  1-35,  9  figs.    £11.40 

No.  2  An  introduction  to  fern  genera  of  the  Indian  subcontinent.  C.R. 
Fraser- Jenkins.  1984.  Pp.  37-76,  1  fig.  £12.50 

No.  3  A  revision  of  African  Sphagnales.  A.  Eddy.  1985.  Pp.  77-162, 
47  figs.  £23.05 

No.  4  Studies  in  the  genus  Hypericum  L.  (Guttiferae)  3.  Sections  1. 
Campylosporus  to  6a.  Umbraculoides.  N.K.B.  Robson.  1985. 
Pp.  163-325,  24  figs,  34  maps.  £39.20 

Volume  13 

No.  1          The  lichen  genus  Usnea  subgenus  Neuropogon.  F.J.  Walker. 

1985.  Pp.  1-130,  39  figs.  £31.85 

No.  2  Cytotaxonomic  studies  of  the  ferns  of  Trinidad.  A.C.  Jermy  & 
T.G.  Walker.  1985.  Pp.  131-276,  69  figs.  £31.50 


No.  3          Some  genera  of  the  Biddulphiaceae  (diatoms)  with  interlocking 
linking  spines.  R.  Ross  &  PA.  Sims.  1985.  Pp.  277-381,  33 
plates.  £28.75 

Volume  14 

No.  1  Cytological  observations  on  Indian  subcontinent  and  Chinese 
Dryopteris  and  Polystichum  (Pteridophyta:  Dryopteridaceae). 
M.  Gibby.  1985.  Pp.  1^2,  78  figs.  £12.50 

No.  2          A  redisposition  of  the  species  referred  to  the  ascomycete  genus 
Microthelia.  D.L.  Hawksworth.  1985.  Pp.  43-181,  73  figs. 

£34.25 

No.  3          A  classification  of  the  genus  Dryopteris  (Pteridophyta: 

Dryopteridaceae).  C.R.  Fraser- Jenkins.  1986.  Pp.  183-218,  4 
figs.  £11.75 

No.  4          Evolutionary  cladistics  of  marattialean  ferns.  C.R.  Hill  &  J.M. 
Camus.  1986.  Pp.  219-300,  27  figs.  £24.65 

Volume  15 

No.  1  Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 

islands:  a  critical  assessment.  IV.  Rhodophyta  (Florideae)  1. 
Genera  A-F.  J.H.  Price,  D.M.  John  &  G.W.  Lawson.  1986.  Pp. 
1-122,  1  fig.  £33.60 

No.  2          Cytology  of  the  fern  flora  of  Madeira.  I.  Manton,  J.D.  Lovis, 

G.  Vida  &  M.  Gibby.  1986.  Pp.  123-161,  12  plates.         £14.70 

No.  3  A  revision  of  the  lichen  genus  Xanthoparmelia  in  Australasia. 
J.A.  Elix,  J.  Johnston  &  P.M.  Armstrong.  1986.  Pp.  163-362, 
42  figs,  117  maps.  £42.90 

Volume  16 

No.  1  Studies  in  the  genus  Hypericum  L.  (Guttiferae)  7.  Section  29. 

Brathys  (part  1).  N.K.B.  Robson.  1987.  Pp.  1-106,  14  figs,  25 
maps.  £28.75 

No.  2          The  lichen  genus  Ramalina  in  Australia.  G.N.  Stevens.  1987. 

Pp.  107-233,  15  plates,  31  figs.  £32.20 

No.  3          An  annotated  list  of  vascular  plants  collected  in  the  valleys 
south  of  Mt  Everest.  G.  Miehe.  1987.  Pp.  225-268,  4  figs. 

£16.50 

No.  4          Further  genera  of  the  Biddulphiaceae  (diatoms)  with  interlock- 
ing linking  spines.  R.  Ross  &  A.Sims.  1987.  Pp.  269-31 1,  13 
plates.  £15.60 

Volume  17 

No.  1  Studies  in  Pseudocyphellaria  (lichens)  1 .  The  New  Zealand 

species.  D.J.  Galloway.  1988.  Pp.  1-267,  124  figs.  £55.65 

Volume  18 

No.  1  An  illustrated  catalogue  of  the  type  specimens  in  the  Greville 
diatom  herbarium.  D.M.  Williams.  1988.  Pp.  1-148,  74  plates. 

£38.00 

No.  2          Erik  Acharius  and  his  influence  on  English  lichenology.  D.J. 

Galloway.  1988.  Pp.  149-194,  18  figs.  £17.05 

No.  3          Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  IV.  Rhodophyta  (Florideae)  2. 
Genera  G.  J.H.  Price,  D.M.  John  &  G.W.  Lawson.  1988.  Pp. 
195-273,  1  fig.  £25.10 

No.  4          Some  Cretaceous  and  Palaeogene  Trinacria  (diatom)  species. 

PA.  Sims  &  R.  Ross.  1988.  Pp.  275-322,  13  plates.         £17.55 

No.  5          A  monograph  of  Dryopteris  (Pteridophyta:  Dryopteridaceae)  in 
the  Indian  subcontinent.  C.R.  Fraser- Jenkins.  1989.  Pp.  323- 
477,  79  figs.  £34.10 

No.  6          Corydalis  (Papaveraceae:  Fumarioideae)  in  Nepal.  M.  Liden. 

1 989.  Pp.  479-538,  26  figs.  £19.35 


Volume  19 


A  new  species  of  Maytenus  (Celastraceae)  in  Ethiopia.  Sebsebe 
Demissew.  1989.  Pp.  1-3,  1  fig. 


Central  American  Araliaceae  -  a  precursory  study  for  the  Flora 

Mesoamericana.  M.J.  Cannon  &  J.F.M.Cannon.  1989.  Pp.  5- 

6 1,36  figs. 

A  revision  of  the  Solarium  nitidum  group  (section  Holophylla 

pro  parte):  Solanaceae.  S.  Knapp.  1989.  Pp.  63-102,  21  figs. 

Six  new  species  of  Solatium  sect.  Geminata  from  South 

America.  S.  Knapp.  1989.  Pp.  103-1 12,  8  figs. 

The  application  of  names  of  some  Indian  species  of  Ocimum 

and  Geniosporum  (Labiatae).  J.R.  Press  &  V.V.  Sivarajan. 

1989.  Pp.  11 3- 11 6,  4  figs. 

Revision  of  Piper  (Piperaceae)  in  the  New  World  1 .  Review  of 

characters  and  taxonomy  of  Piper  section  Macrostachys.  M.C. 

Tebbs.  1989.  Pp.  1 17-158,  41  figs.  Facsimile  edition.      £52.45 

Volume  20 

No.  1  Studies  in  the  genus  Hypericum  L.  (Guttiferae)  8.  Sections  29. 

Brathys  (part  2)  and  30.  Trigynobrathys.  N.K.B.  Robson.  1990. 
Pp.  1-151,  22  figs,  46  maps.  Facsimile  edition.  £49.25 

No.  2          The  marine  algal  flora  of  Namibia:  its  distributions  and 

affinities.  G.W.  Lawson,  R.H.  Simons  and  W.E.  Isaac.  1990. 

Pp.  153-168,  1  fig,  7  plates. 

The  infrageneric  classification  of  Gentiana  (Gentianaceae).  T- 

N.  Ho  and  S.-W.  Liu.  1990.  Pp.  169-192,  13  figs. 

Revision  of  Piper  (Piperaceae)  in  the  New  World.  2.  The 

taxonomy  of  Piper  section  Churumayu.  M.C.  Tebbs.  1990.  Pp. 

193-236,  49  figs.  £34.10 

Volume  21 

No.  1  Historical  and  taxonomic  studies  in  the  genus  Titanoderma 

(Rhodophyta,  Corallinales)  in  the  British  Isles.  Y.M.  Chamber- 
lain. 1991.  Pp.  1-80,  247  figs.  £42.35 

No.  2          Early  collections  of  the  Holy  Thorn  (Crataegus  monogyna  cv. 
Biflora).  A.R.  Vickery.  1991.  Pp.  81-83,  1  fig. 
A  taxonomic  study  of  the  species  referred  to  the  ascomycete  genus 
Leptorhaphis.  B.  Aguirre-Hudson.  1991 .  Pp.  85-192, 76  figs. 
The  typification  and  identification  of  Calymperes 
crassilimbatum  Renauld  &  Cardot  (Musci:  Calymperaceae). 
L.T.  Ellis.  1991.  Pp.  193-194,  1  fig.  £42.35 

Volume  22 

No.  1  An  account  of  southern  Australian  species  of  Lithophyllum 

(Corallinaceae,  Rhodophyta).  Wm.  J.  Woelkerling  and  S.J. 
Campbell.  1992.  Pp.  1-107,  63  figs.  £41.25 

No.  2          Palynological  evidence  for  the  generic  delimitation  of  Sechium 
(Cucurbitaceae)  and  its  allies.  J.L.  Alvarado,  R.  Lira-Saade  & 
J.  Caballero.  1992.  Pp.  109-121. 

Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  IV.  Rhodophyta  (Florideae)  3. 
Genera  H-K.  J.H.  Price,  D.M.  John  &  G.W.  Lawson.  1992.  pp. 
123-146. 

Two  new  species  of  Solarium  section  Geminata  (Solanaceae) 
from  Cerro  del  Torr  in  western  Colombia.  S.  Knapp.  1992.  Pp. 
147-152. 

Fissidens  ceylonensis  Dozy  &  Molkenb.  (Musci: 
Fissidentaceae)  and  some  allied  taxa  from  southern  India.  L.T. 
Ellis.  1992.  Pp.  153-156,  2  figs. 

New  species  of  Piper  (Piperaceae)  from  Central  America.  M. 
Tebbs.  1992.  Pp.  157-158. 

Studies  on  the  Cretan  flora  1 .  Floristic  notes.  N.J.  Turland. 
1992.  Pp.  159-164. 

Studies  on  the  Cretan  flora  2.  The  Dianthus  juniperinus  complex 
(Caryophyllaceae).  N.J.  Turland.  1992.  Pp.  165-169.       £41.25 

Volume  23 

No.  1  Revision  of  Piper  (Piperaceae)  in  the  New  World  3.  The 

taxonomy  of  Piper  sections  Lepianthes  and  Radula.  M.C. 
Tebbs.  1993.  Pp.  1-50,  18  figs. 

Mounting  techniques  for  the  preservation  and  analysis  of 
diatoms.  S.J.  Russell.  1993.  Pp.  51-54.  1  fig.  £43.25 

No.  2          New  taxa  of  Gentiana  (Gentianaceae)  from  Western  China  and 


the  Himalayan  region.  T.-N.  Ho  and  S.-W.  Liu.  1993.  Pp.  55- 

60.  2  figs. 

New  combinations,  names  and  taxonomic  notes  on  Gentianella 

(Gentianaceae)  from  South  America  and  New  Zealand.  T.-N. 

Ho  and  S.-W.  Liu.  1993.  Pp.  61-66. 

Studies  in  Hypericum:  validation  of  new  names.  N.K.B. 

Robson.  1993.  Pp.  67-70. 

Generic  monograph  of  the  Asteraceae-Anthemideae.  K. 

Bremer  and  C.J.  Humphries.  1993.  Pp.  71-177.  12  figs.  £43.25 

Volume  24 

No.  1  Pre-Linnaean  references  for  the  Macaronesian  flora  found  in 

Leonard  Plukenet's  works  and  collections.  J.  Francisco-Ortega, 
A.  Santos-Guerra  and  C.E.  Jarvis.  Pp.  1-34. 
Studies  on  the  lichen  genus  Sticta  (Schreber)  Ach.:  II. 
Typification  of  taxa  from  Swartz's  Prodromus  of  1788.  D.J. 
Galloway.  Pp.  35-48. 

Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  IV.  Rhodophyta  (Florideae)  4. 
Genera  L-O.  D.M.  John,  G.W.  Lawson,  J.H.  Price,  W.F. 
Prud'homme  van  Reine  and  W.J.  Woelkerling.  Pp.  49-90. 
Studies  on  the  Cretan  flora  3.  Additions  to  the  flora  of  • 

Karpathos.  N.J.  Turland  and  L.  Chilton.  Pp.  9 1-100.        £43.25 

No.  2          Observations  on  the  benthic  marine  algal  flora  of  South 

Georgia:  a  floristic  and  ecological  analysis.  D.M.  John,  P.J.A. 

Pugh  and  I.  Tittley.  Pp.  101-1 14. 

Studies  in  Pseudocyphellaria  (Lichens)  IV.  Palaeotropical 

species  (excluding  Australia).  D.J.  Galloway.  Pp.  1 15-160. 

Morphology  and  ecology  of  seedlings,  fruits  and  seeds  of 

Panama:  Bixaceae  and  Cochlospermaceae.  N.C.  Garwood.  Pp. 

161-172. 

A  study  of  Bixa  (Bixaceae),  with  particular  reference  to  the  leaf 

undersurface  indumentum  as  a  diagnostic  character.  R.E. 

Dempsey  and  N.C.  Garwood.  Pp.  173-180.  £43.40 

Volume  25 

No.  1  A  revision  of  Rutilaria  Greville  (Bacillariophyta).  R.  Ross.  Pp. 

1-94. 

William  Roxburgh's  St  Helena  plants.  Q.C.B.  Cronk.  Pp.  95-98. 

£43.40 

No.  2          Seaweeds  of  the  western  coast  of  tropical  Africa  and  adjacent 
islands:  a  critical  assessment.  IV.  Rhodophyta  (Florideae)  5. 
Genera  P.  G.W.  Lawson,  W.J.  Woelkerling,  J.H.  Price,  W.F. 
Prud'homme  Van  Reine  and  D.M.  John.  Pp.  99-122. 
A  new  species  of  Odontorrhynchos  (Orchidaceae, 
Spiranthinae)  from  Boliva.  D.L.  Szlachetko.  Pp.  123-125. 
Linnaeus's  interpretation  of  Prospero  Alpino's  De  plantis 
exoticis,  with  special  emphasis  on  the  flora  of  Crete.  N.J. 
Turland.  Pp.  127-159. 
Book  review.  M.G.  Gilbert.  P.  1 6 1 .  £43.40 

Volume  26 

No.  1  A  morphological  study  of  Chaetoceros  species 

(Bacillariophyta)  from  the  plankton  of  the  Pacific  ocean  of 
Mexico.  D.U.  Hernandez-Becerril.  1996.  Pp.  1-73.          £43.40 

No.  2          Studies  in  the  genus  Hypericum  L.  (Guttiferae)  6.  Sections  20. 
Myriandra  to  28.  Elodes.  N.K.B.  Robson.  1996.  Pp.  75-217. 

£43.40 


Volume  27 

No.  1  Notes  on  the  diatom  species  Tetracyclus  castellum  (Ehrenb.) 

Grunow  with  a  description  of  Tetracyclus  pseudocastellum  nov. 

sp.  D.M.  Williams.  Pp.  1-5. 

A  new  species  of  Calymperes  (Musci:  Calymperaceae)  from 

Peninsular  Malaysia.  L.T.  Ellis.  Pp.  7-9. 

A  phylogenetic  conspectus  of  the  tribe  Hyoscyameae 

(Solanaceae).  A.L.  Hoare  and  S.  Knapp.  Pp.  1 1-29. 

A  revision  of  Solanum  section  Ptemidea:  Solanaceae.  S. 

Knapp  and  T.  Helgason.  Pp.  3 1-73.  1997.  £43.40 

No.  2          Systematics  of  Pogostemon  (Labiatae)  G.R.  Bhatti  and  M. 

Ingrouille.  Pp.  77-147.  1997  £43.40 


CONTENTS 

1        Morphology  and  ecology  of  seedlings,  fruits  and  seeds  of  Panama:  Vochysiaceae 

Nancy  C.  Garwood 
17      A  revision  of  the  genus  Mandragora  (Solanaceae) 

S.  Ungricht,  S.  Knapp  and  J.R.  Press 

41       The  pteridophytes  of  Sao  Tome  and  Principe  (Gulf  of  Guinea) 

Estrela  Figueiredo 


tural  History  Musei 


BOTANY  SERIES 

Vol.28,  No.  1,  June  1998