Lagascalia 26: 71-81 (2006)
THE STATUS OF PORTULACA OLERACEA L. IN
TENERIFE, THE CANARY ISLANDS
A. DANIN* & J. A. REYES-BETANCORT**
* Depa men o E olu ion, Sys ema ics, and Ecology. The Alexande Silbe man
Ins i u e. The Heb ew Uni e si y o Je usalem, Is ael 91904. ** Ja dín de Aclima a-
ción de La O o a a (ICIA). Pue o de La C uz 38400. Islas Cana ias, Spain
(Recibido el 10 de Ene o de 2006)
Resumen. La ecolección de ma e ial pe enecien e a Po ulaca ole acea y especies a ines en
Tene i e y algunas o as islas del a chipiélago ea i ó nues o in e és en es e complejo poliploide.
Con la in ención de homogeneiza el a amien o de los axones elacionados a Po ulaca ole acea
L., se p opone ele a el ango subespecí ico de alguno de ellos a ango especí ico. Se desc ibe po
p ime a ez pa a la ciencia Po ulaca cana iensis. El es udio de Po ulaca ole acea y especies a ines
en Tene i e y o as islas del a chipiélago e elan los siguien es hechos: a. T es ni eles de ploidía
(diploide, e aploide y hexaploide) es án p esen es en las islas, b. Se obse a on poblaciones
simpá icas de especies diploides, de e aploides, de e aploides más hexaploides, de hexaploides
y de diploides más e aploides y hexaploides y c. Las especies e aploides ue on las más ecuen es,
seguidas de las diploides, siendo las hexaploides las menos ecuen es. Los esul ados de los ecuen-
os c omosómicos concue dan con los es udios p e ios ealizados
Summa y. Following a comp ehensi e collec ion o Po ulaca ole acea- ela ed species in
Tene i e and a ew o he Cana y Island enewed in e es his polyploidy complex led o ano he
p og ess in i s in es iga ions. In o de o equalize he ea men o he axa ela ed o Po ulaca
ole acea L., i s subspecies a e aised o he speci ic ank. A new species, P. cana iensis is desc ibed.
Ou s udy o he Po ulaca ole acea- ela ed species ound in Tene i e and a ew o he Cana y Island
e eal he ollowing ac s: a. The h ee ploidy le els (diploids, e aploids, and hexaploids) a e well
ep esen ed he e, b. Sympa ic popula ions o diploid plus diploid, e aploids plus e aploids,
e aploids plus hexaploids, hexaploids plus hexaploids, and diploid plus e aploid plus hexaploid
we e disco e ed he e and c. The mos equen species ound we e e aploid, hen he diploid and
he leas we e hexaploids. Ch omosome coun ollowed ou indings in p e ious s udies.
INTRODUCTION
Po ulaca ole acea is well known o be a cosmopoli an species. In ea lie s udies
(DANIN & al., 1978; DANIN & ANDERSON, 1986; DANIN, 1990) we ha e shown ha
i is an une enly dis ibu ed polyploid complex. The e is a high ege a i e
esemblance among he a ious axa al eady desc ibed. Thei seed-coa cha ac e s,
seed size, and ch omosomes numbe can be used o dis inguishing he axa. When
ou i s s udy ook place, a ew colleagues (e.g. D . Pe e Ra en, 1978, and 2003 pe s.
comm.) s a ed:” I hese axa a e dis inc , ecognized, and ep esen di e en ploidy
le els, hey should be ecognized a he speci ic le el, e en i he di e ences a e
limi ed”. This is because hey a e ecognizable by hei seed-coa mo phology and
72 The s a us o Po ulaca ole acea L. in Tene i e, he Cana y Islands
biologically isola ed by hei ploidy le el ba ie s. Ou indings we e dis ega ded by
MATTHEWS & al. (1993) who concluded ha “P. ole acea exis s as a polymo phic
species ha is no di isible in o subspecies based on seed su ace as p ima y
mo phological ai ”. MATTHEWS & al. (l.c.) o go conside ing he seeds-size and
ch omosome numbe cha ac e s which a e linked o he seed su ace mo phology in
ou s udies. They did no check he he ba ium specimens we quo ed and made
hemsel es easy li e by concluding wi hou in es ing ime in inding new ac s. They
claim ha ou indings a e no eliable and ha Po ulaca ole acea is in ac a
polymo phic plan wi h popula ions ha con ain se e al se s o ch omosome
numbe s and mixed seed-su ace ypes. GEREAU (2001), who ollowed MATTHEWS
& al. (1993) and in a simila way o he la e did no was e ime on inding new ac s,
sunk he wo subspecies eco ded om Nica agua in o s ic synonyms o he species,
wi hou gi ing any explana ion. In pa icula , Po ulaca ole acea L. subsp.
nica aguensis Danin & H. G. Bake , a common diploid o he Gul o Mexico
coun ies, was no ecognized by GEREAU (2001), and no was subsp. g anula o-
s ellula a collec ed in ha coun y. He p obably ailed o use a high magni ica ion
dissec ing mic oscope wi h di used ligh , as ecommended by DANIN & al. (1978,
p. 178), and he e o e could no see he unique seed coa p ope ies o hese axa.
Consequen ly GEREAU (2001) desc ibed o he Nica aguan Po ulaca seed
p ope ies o Po ulaca ole acea L. subsp. ole acea which do no exis , o ou bes
knowledge, in Nica agua. On he o he hand, RICCERI & ARRIGONI (2000) ook he
ad ice gi en o us by D . Ra en and aised wo o ou subspecies o he species le el.
The new names o hese axa a e lis ed below.
The p esen s udy epo s he axonomic analysis o samples o Po ulaca
ole acea g oup om 56 popula ions in Tene i e, The Cana y Islands. A new species
o Po ulaca is desc ibed, and he axa which we e no aised o he species le el a e
aised he e o equalize he ea men .
MATERIAL AND METHODS
New collec ion o seeds om Tene i e was made in Oc obe 2003. The loca ions
a e lis ed in appendix 1 wi h hei unning numbe s. Thei dis ibu ion is shown in
Fig. 1. We also analyzed he specimens deposi ed in he he ba ium o he Depa men
o Biologia Vege al, o he Uni e si y o La Laguna, Tene i e (TFC). Popula ions
we e de e mined using seeds size and seed-coa mo phology. They we e compa ed
o he SEM igu es in DANIN & al. (1978). All he specimens quo ed he e we e seen
(!). Addi ional SEM images we e aken a e being coa ed wi h ca bon-coa ing. A ew
o hem a e p esen ed he e.
Chomosome coun s we e made o oo ips om seeds o sample T30 (see
appendix) in o de o e i y he conclusions om he 1978 s udy wi h he Cana y
Islands ma e ial. Seeds om all popula ions eco ded in appendix a e deposi ed a
HUJ and TFC. Seeds om popula ions T04, T05, T10, T14, T25, T32, T34, T38, T40,
Lagascalia 26, 2006 73
LZ2 we e plan ed and indi iduals we e aised om mos o hem in a g eenhouse a
La Laguna. Seed mo phology o F1 was de e mined and compa ed o he pa en seed
ype. Addi ional ch omosome coun s we e done in D . B. Gemeinholze labo a o y
a he Be lin-Dahlem Bo anical Ga den.
Fig. 1. Loca ions o he collec ion si es o Po ulaca species in Tene i e.
RESULTS
Accep ed axa
Following RICCERI & ARRIGONI (2000) hese a e he Po ulaca axa ecognized:
Po ulaca ole acea L. (named Po ulaca ole acea L. subsp. ole acea by DANIN &
al., 1978). Fig. 2.
Po ulaca nica aguensis (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. nica aguensis Danin &
H. G. Bake in: A. DANIN, I. BAKER, & H. G. BAKER, Is ael J. Bo . 27: 186-187.
1978. Fig. 3.
74 The s a us o Po ulaca ole acea L. in Tene i e, he Cana y Islands
Fig. 2. SEM o Po ulaca ole acea seeds (sou ce – T38, T40).
Fig. 3. SEM o Po ulaca nica aguensis seeds (sou ce – T32, T34).
Lagascalia 26, 2006 75
Fig. 4. SEM o Po ulaca g anula o-s ellula a seeds (sou ce – T04).
Fig. 5. SEM o Po ulaca ni ida seeds (sou ce – T14).

76 The s a us o Po ulaca ole acea L. in Tene i e, he Cana y Islands
Po ulaca g anula o-s ellula a (Poelln.) C. Ricce i & P.V. A igoni (named
Po ulaca ole acea L. subsp. g anula o-s ellula a (Poelln.) Danin & H. G.
Bake by DANIN & al. 1978). Fig. 4.
Po ulaca ni ida (Danin & H. G. Bake ) C. Ricce i & P.V. A igoni (named Po ulaca
ole acea L. subsp. ni ida Danin & H. G. Bake by DANIN & al., 1978). Fig. 5.
Po ulaca a icana (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. a icana Danin & H. G.
Bake , in: A. DANIN, I. BAKER, & H. G. BAKER, Is ael J. Bo 27: 187-189. 1978.
Po ulaca ube cula a (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. ube culal a Danin &
H. G. Bake , in: A. DANIN, I. BAKER & H. G. BAKER, Is ael J. Bo . 27: 194, 1978.
Po ulaca impoli a (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. impoli a Danin & H. G.
Bake , in: A. Danin, I. Bake & H. G. Bake , Is ael J. Bo . 27: 195-196. 1978.
Po ulaca s ella a (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. s ella a Danin and H.
G. Bake , in: A. DANIN, I. BAKER & H. G. BAKER Is ael J. Bo . 27: 198-
200.1978. Fig. 6
Po ulaca papilla o-s ellula a (Danin & H. G. Bake ) Danin, s a . no .
Basionym and synonym: Po ulaca ole acea L. subsp. papilla o-s ellula a
Danin & H. G. Bake , in: A. DANIN, I. BAKER & H. G. BAKER, Is ael J. Bo . 27:
200-201. 1978.
Fig. 6. SEM o Po ulaca s ella a seeds (sou ce – LZ02).
Lagascalia 26, 2006 77
Po ulaca cana iensis Danin & Reyes-Be anco species no .
Species seminibus supe icie impoli a g isea ple umque ni o e me allico p edi a.
Ab Po ulaca impoli a (Danin and H. G. Bake ) Danin cellulis la i udine 2-3 plo
longio e (non isodiame icis nec as e i o mibus) adii la i udine b e is (nec
equalibus ad la i udine 2-3 plo longio e). Fig. 7.
Type: Tene i e, Fasnia, La Hondu a, 30.3.1996, C uz T ujillo 39.452 (Holo -
TFC; Iso - HUJ). Fig. 6.
Addi ional specimens seen: Cana y Islands, Tene i e, El Escobonal, 24.3.1981,
Rod iguez 12847 (TFC); Fasnia, La Hondu a, 30.3.1996, C uz T ujillo 39.452 (TFC);
Ba anco de Ijuana (Lade a S.) 250m, 12.5.1996, Racca 39.164 (TFC). Lanza o e,
Tinajo, Casas del Isole e, 24.4.1996, Reyes-Be anco 40.328 (TFC).
Con a y o he si ua ion o all he o he axa o Po ulaca in he Cana y Islands,
P. cana iensis g ows in habi a s ha a e no much dis u bed. Mos o ou specimens
we e collec ed in he he In amedi e anean dese ic-xe ic a id-semia id bioclima ic
bel s, sensu RIVAS-MARTINEZ & al. (1993).
Ch omosome coun s
The only sample which was success ully ge mina ed and p o ided oo ips
wo h o ch omosomes coun in La Laguna was T30. I is a sample o Po ulaca
nica aguensis and he numbe o ch omosomes ound was 18, as in he p e ious
coun s (DANIN & al., 1978). The coun in Be lin o sample T32 om Los Ch is ianos
Fig. 7. SEM o Po ulaca cana iensis seeds (sou ce – LZ02).
78 The s a us o Po ulaca ole acea L. in Tene i e, he Cana y Islands
(P. nica aguensis) ga e 18 ch omosomes, T41 om A a o (P. g anula o-s ellula a)
ga e 36 ch omosomes, T52 om O o a a (P. ni ida) ga e 36 ch omosomes, T44 om
Fasnia (P. ole acea) ga e 54 ch omosomes. All hese coun s a e in allacco d wi h
ou p e ious coun s (DANIN & al. 1978).
Seed su ace in o sp ing
The esul s o aising F1 seeds om pa en ma e ial collec ed in Tene i e and
aised he e in a g eenhouse a e lis ed in Table 1. Un o una ely, samples No. T04,
T25, T32, T34, and LZ02 did no ge mina e o did no de elop o p oduce seeds.
Checking again he esul s o simila F1 aising o ou 1978 ma e ial e ealed 100%
inhe i ence o seed su ace cha ac e s in one popula ion o P. nica aguensis ( om
Nica agua), one popula ion o P. ni ida ( om Cali o nia), wo popula ions o P.
g anula o-s ellula a ( om Nica agua and om Is ael), and wo popula ions o P.
papilla o-s ellula a ( om Palm Sp ings and om Be keley in Cali o nia).
Table 1. Summa y o o sp ing seed coun s in popula ions o P. ole acea complex om he Cana y
Islands.
DISCUSSION
The i s expe imen o aising he F1 seeds om seeds collec ed in he ield was
al eady done o he ma e ial epo ed in DANIN & al. (1978) and epo ed again he e.
Howe e , in o de o double check he s a us o he axa in his complex, we epea ed
aising plan s om seeds collec ed in he Cana y Islands. The o sp ings did no p o-
Species Locali y Seeds
coun ed
Seeds o he
same species
P. g anula o-s ellula a San a C uz 0 0
P. ni ida San And es 33 33
P. g anula o-s ellula a Bajama 25 25
P. ni ida Buena Vis a del No e 37 37
P. nica aguensis Guaza 0 0
P. nica aguensis Los C is ianos 0 0
P. nica aguensis Playa de la A ena 0 0
P. ole acea Las E as 6 6
P. ole acea Escabonal 8 8
P. cana iensis Timan aya, Lanza o e 0 0
Lagascalia 26, 2006 79
duce seeds o mixed mo pho ypes; his p ope y seems o be inhe i ed. Each o he
h ee species which became ma u e p oduced seeds which we e o he same species.
Seed a iabili y in he axa o he P. ole acea g oup and he ew hyb ids be ween axa
ecognized we e al eady discussed by DANIN & al. (1978).
When conside ing which le el o gi e he axa we disco e ed (DANIN & al.
1978) we had wo possibili ies – ha sugges ed by D . P. Ra en (pe s. comm. 1978
and 2003) and ou own conside a ion. In 1978 we we e hinking o he common
bo anis , inding a plan om his g oup. We wan ed he /him o be able o
di e en ia e i a a ce ain le el. The isibili y o “li ing wi h Po ulaca ole acea”
was he ligh mo i e. In la e s udies I conduc ed (DANIN 1981, 1990; DANIN &
ANDERSON 1986) I had he eeling ha ou way was he igh way. Howe e , he e
we e people who ailed o unde s and i and sunk he subspeci ic axa o he
“synonymy le el” (MATTHEWS & al., 1993; GEREAU, 2001). The speci ic le el,
sugges ed by D . Ra en, is adop ed by RICCERI & ARRIGONI (2000). This e mina es
he imes when a collec o could know wha he holds in his hand wi hou using a
dissec ing mic oscope o de e mine a species om he Po ulaca ole acea g oup o
axa. Raising he es o he axa in o a species le el became now an ine i able s ep.
The esul s o he s udy o Po ulaca species ound in he Cana y Islands, and
in pa icula he occu ence o P. nica aguensis, indica e he possibili y o seed
anspo a ion by cu en s in he A lan ic Ocean. The p e iously known dis ibu ion
a ea o P. nica aguensis is om he coun ies nea he Gul o Mexico, om Nica a-
gua h ough sou he n Flo ida (DANIN & al., 1978; DANIN & ANDERSON, 1986).
Howe e , he inc easing isi s o ou is s and he possible an h opogenic dispe sal
o seeds can no be excluded.
The disco e y o he new species P. cana iensis calls o u he esea ch. The
seeds we had a ailable while p epa ing he p esen pape , we e de i ed om old
collec ions and passed umiga ion be o e hei deposi ion in he he ba ium. This
migh ha e in luenced hei ge minabili y and we could no s udy hei ch omosome
numbe . Howe e , seed size o 1020-1114 x 880-1000 ¼m may indica e a high
possibili y, ollowing DANIN & al. (1978), ha we deal he e wi h a hexaploid. DANIN
& al. (1978) assume ha phylogene ic ela ionships among he axa ela ed o P.
ole acea may be d awn when seed su ace mo phology is compa ed. The closes
axon o P. cana iensis in i s seed su ace mo phology is P. nica aguensis. The
epide mal cells o bo h ha e sho and wide a ms which in e inge wi h hose o he
neighbou ing cells in a simial way. Howe e , P. cana iensis su ace e lec s me alic
sheen and has minu e bumps (1-3 ¼m) seen only a high magni ica ion (Fig. 6d). P.
nica aguensis is unique in he wax co e o mos o i s seeds (Fig. 3d) o hei o ally
smoo h su ace (Fig 3c). Fu u e esea ch wi h mo e sophis ica ed me hods may assis
in ull unde s anding o he ac ual ela ionships o axa in his di e s and in e es ing
g oup o axa.
Acknowledgemen s. We hank he keepe o he he ba ium a La Laguna (TFC) o enabling
us o use he ma e ial. We hank M . M. DVORACHEK, The Geological Ins i u e, Je usalem o he