The status of Portulaca oleraceal in Tenerife, the Canary Islands

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