orobanche clausonis pomel …...orobanche clausonis pomel (orobanchaceae) in the iberian penÍnsula...

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OROBANCHE CLAUSONIS POMEL (OROBANCHACEAE) IN THE IBERIAN PENÍNSULA by MICHAEL JAMES YATES FOLEY* Resumen FOLEY, MJ.Y. (1996). Orobanche clausonis Pomel (Orobanchaceae) en la Península Ibérica. Anales Jard. Bot. Madrid 54:319-326 (en inglés). Orobanche clausonis Pomel fue descrita sobre plantas recolectadas en Argelia, donde parasi- taba a Asperula hirsuta (Rubiaceae). Desde entonces, ha sido colectada ocasionalmente en va- rias localidades del sudoeste de Europa, especialmente en la Península Ibérica. Sin embargo, es aún mal conocida. En este trabajo se estudian la morfología y la taxonomía de la especie y se propone que las plantas europeas queden cobijadas bajo el trinomen O. clausonis subsp. hes- perina (J.A. Guim.) MJ.Y. Foley, comb. & stat. nov. Palabras clave: Spermatophyta, Orobanchaceae, Orobanche, taxonomía, Península Iberica, Argelia. Abstract FOLEY, MJ.Y. (1996). Orobanche clausonis Pomel (Orobanchaceae) in the Iberian Península. Anales Jard. Bot. Madrid 54:319-326. Orobanche clausonis Pomel was first described frorn Algeria where it was thought to be para- sitic upon Asperula hirsuta (Rubiaceae). Since then, germine records have been scarce and al- though occasionally collected from various localities in south-westera Europe (especially the Iberian península), where it is mainly parasitic upon members of the Rubiaceae, its identity and taxonomy have been poorly understood. Based principally on the limited number of preserved specimens available, the general morphology and taxonomy of O. clausonis has been investi- gated. As a result, it is proposed that the European plants be separated as Orobanche clausonis subsp. hesperina (J.A. Guim.) MJ.Y. Foley, comb. & stat. nov. Key words: Spermatophyta, Orobanchaceae, Orobanche, taxonomy, Iberian Península, Algeria. INTRODUCTION AND HISTORY of this results from the scarcity of subsequent records and the dearth of specimens retained Since first described by POMEL (1874:107) in herbaría. In the present work an attempt has based upon material collected in 1858 by been made to reassess the taxonomic status of Clauson and growing on Asperula hirsuta at the plant based mainly upon the relatively Mouzaia, Algeria, there has been much con- limited quantity of preserved material avail- fusion regarding the identity and taxonomic able. Only one present-day locality has so far status of Orobanche clausonis, a holoparasite been definitely identified for the plant, but mainly on members of the Rubiaceae. Much such apparent scarcity may be a result of mis- * Institute of Environmental and Biological Sciences, División of Biological Sciences, University of Lancaster. Lancaster, LA14YQ (United Kingdom).

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Page 1: OROBANCHE CLAUSONIS POMEL …...OROBANCHE CLAUSONIS POMEL (OROBANCHACEAE) IN THE IBERIAN PENÍNSULA by MICHAEL JAMES YATES FOLEY* Resumen FOLEY, MJ.Y. (1996). Orobanche clausonis Pomel

OROBANCHE CLAUSONIS POMEL (OROBANCHACEAE)IN THE IBERIAN PENÍNSULA

by

MICHAEL JAMES YATES FOLEY*

Resumen

FOLEY, MJ.Y. (1996). Orobanche clausonis Pomel (Orobanchaceae) en la Península Ibérica.Anales Jard. Bot. Madrid 54:319-326 (en inglés).

Orobanche clausonis Pomel fue descrita sobre plantas recolectadas en Argelia, donde parasi-taba a Asperula hirsuta (Rubiaceae). Desde entonces, ha sido colectada ocasionalmente en va-rias localidades del sudoeste de Europa, especialmente en la Península Ibérica. Sin embargo,es aún mal conocida. En este trabajo se estudian la morfología y la taxonomía de la especie yse propone que las plantas europeas queden cobijadas bajo el trinomen O. clausonis subsp. hes-perina (J.A. Guim.) MJ.Y. Foley, comb. & stat. nov.

Palabras clave: Spermatophyta, Orobanchaceae, Orobanche, taxonomía, Península Iberica,Argelia.

Abstract

FOLEY, MJ.Y. (1996). Orobanche clausonis Pomel (Orobanchaceae) in the Iberian Península.Anales Jard. Bot. Madrid 54:319-326.

Orobanche clausonis Pomel was first described frorn Algeria where it was thought to be para-sitic upon Asperula hirsuta (Rubiaceae). Since then, germine records have been scarce and al-though occasionally collected from various localities in south-westera Europe (especially theIberian península), where it is mainly parasitic upon members of the Rubiaceae, its identity andtaxonomy have been poorly understood. Based principally on the limited number of preservedspecimens available, the general morphology and taxonomy of O. clausonis has been investi-gated. As a result, it is proposed that the European plants be separated as Orobanche clausonissubsp. hesperina (J.A. Guim.) MJ.Y. Foley, comb. & stat. nov.

Key words: Spermatophyta, Orobanchaceae, Orobanche, taxonomy, Iberian Península,Algeria.

INTRODUCTION AND HISTORY of this results from the scarcity of subsequentrecords and the dearth of specimens retained

Since first described by POMEL (1874:107) in herbaría. In the present work an attempt hasbased upon material collected in 1858 by been made to reassess the taxonomic status ofClauson and growing on Asperula hirsuta at the plant based mainly upon the relativelyMouzaia, Algeria, there has been much con- limited quantity of preserved material avail-fusion regarding the identity and taxonomic able. Only one present-day locality has so farstatus of Orobanche clausonis, a holoparasite been definitely identified for the plant, butmainly on members of the Rubiaceae. Much such apparent scarcity may be a result of mis-

* Institute of Environmental and Biological Sciences, División of Biological Sciences, University of Lancaster.Lancaster, LA14YQ (United Kingdom).

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320 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

identification and under-recording rather thana reflection of extreme rarity, although it isundoubtedly uncommon.

In his original description of O. clausonis,POMEL (1874: 107) remarked on several im-portant characters: the rather dense inflores-cence, the tubular to campanulate corollas,the calyx divided into four shortly lanceolate,subequal divisions, and the insertion of the fil-aments in the lowest quarter of the corollatube, the filaments themselves being slightlypubescent at their base. Of these characters, itis the shape of the broad, relatively short, div-ided calyx as evidenced in the type specimen(Mouzaía, 1858; Clauson, MPU), which es-pecially separates O. clausonis from similartaxa.

Following Pomel's diagnosis, there appe-ars to be no mention of O. clausonis until adismissive comment by BATTANDDER & TRA-BUT (1890: 661) that it was merely a glabres-cent form of O. galii Vaucher var. atlanticahaving more hispid filaments and hairy ant-hers. This latter taxon, apparently parasiticupon Galium tunetanum, is a small-floweredvariant of the widespread European O. caryo-phyllacea Sm., a very different and distinctiveplant to O. clausonis, again apparently restric-ted in its host to members of the Rubiaceae. Itis not clear whether Battandier & Trabut'scomments were based upon actual experienceof O. clausonis, but any affinities betweenit and variants of O. caryophyllacea can bedismissed upon morphological grounds. Inthe same year, BECK (1890: 234) includedO. clausonis in his monograph of the genusOrobanche, listing Clauson's Algerian loca-lity, although not having seen the original spe-cimen. He also added a record of Boissier'sand Reuter's from Granada, Spain, in 1849(specimen now in LE - corolla dissections inPRC!) which at that time appeared to be thefirst and only European record for the plant.He surprisingly placed O. clausonis cióse totaxa of the O. minor group (i.e. in his grex Mi-nores) despite its very different calyx shape,which contrasts with the elongated, taperingcalyces of members of his grex Minores.

In Prodromus Florae Hispanicae, WlL-KOMM & LANGE (1870) made no reference to

O. clausonis, although it was included in thesubsequent supplement (WILKOMM, 1893:188) based on BECK'S (1890:235) verificationof the Granada specimen. Later, when Gui-maraes published his monograph on Portu-guese Orobanche he included under O. luco-rum A. Braun (GUIMARAES, 1904: 85) a newvariety hesperina based on plants from fourcollections by Welwitsch and by Daveau.These were parasitic upon Rubia spp. and inone case upon Quercus pseudococcifera andwere collected from a rather limited área ofcentral Portugal. From elsewhere in his mono-graph it is clear that Guimaraes was aware ofthe principal characters of Pomel's O. clauso-nis since he occasionally quotes such charact-ers in comparison to other taxa and he placed itcióse to O. mauretanica Beck (grex Minores).However he did not consider O. clausonis tobe a Portuguese plant ñor did he refer to it inrelation to his O. lucorum var. hesperina. Hisattribution of the Welwitsch and Daveau spe-cimens to O. lucorum is intriguing and a pos-sible explanation may lie in a combination offactors. True O. lucorum, parasitic on Berberisspp., has a narrow distribution in central Eur-ope, occurring essentially over a limited áreaof the eastern Alps. It is readily separated fromGuimaraes' Portuguese var. hesperina oncalyx shape alone - the lobes of O. lucorumbeing much longer and narrower that in var.hesperina. On the relatively limited informa-tion available at the time, it is possible thatGuimaraes associated his var. hesperina withO. lucorum based on similar colouration (yel-low). Also, a variant or segregate of O. luco-rum from south-eastern France parasitic uponRubus spp. had been described as O. rubi(DUBY, 1828:350). This, together with the factthat Welwitsch had annotated one of his sheets"O. rubii", may have led to a possible misin-terpretation of their respective hosts and there-fore to the connection with O. lucorum. Ne-vertheless, Guimaraes did indícate that thespecimens of Portuguese "O. rubi Duby"which he had seen differed from typical O. lu-corum in several respects and for that reasonhe gave them the varietal rank hesperina.

In Flore de Portugal, COUTINHO (1913:569) took a different view and accepted

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M.J.Y. FOLEY: OROBANCHE CLAUSONIS 321

O. clausonis as a Portuguese plant. This waspresumably based upon the same specimensconsidered by Guimaraes under his O. luco-rum var. hesperina, since he usted the locali-ties and hosts of Welwitsch's collections, an-notated some of the sheets and added a taxo-nomic description. He did not include O. lu-corum for Portugal. This stance has beenmaintained subsequently for Portugal (COU-TINHO, 1939:673; SAMPAIO, 1946:498; FRAN-CO, 1984:280) but O. clausonis is excluded inworks covering Algeria and Morocco (SAU-VAGE, 1961; NÉGRE, 1962; QUEZEL & SANTA,1963) and most regions of Spain (COSTA,1864; FERNÁNDEZ-GALIANO & HEYWOOD,1960; VALDÉS & al., 1987; ROMO, 1989;BOLOS & al., 1990; SANZ, 1990, 1992). Ho-wever, in eastern Andalucía there is a singlerecord by SAGREDO (1987: 421) of plantsparasitic upon Galium verrucosum at Barran-co del Caballar, Almería and another by MU-ÑOZ-MEDINA (1944:121) from near Lanjarón,Granada province. Two collections from Gal-do, north-west Spain, and mentioned by ME-RINO (1906: 45) under O. lucorum, may alsobe O. clausonis. FRANCO (1984: 280) recentlyextended the Portuguese range of O. clauso-nis to include the Atlantic off-shore Ilhas Ber-lengas.

In Flora Europaea, CHATER & WEBB(1972: 292) listed it as present in Spain, Por-tugal and Malta, based upon information gi-ven in BECK'S monographs (1890: 235; 1930:218). However, the solé Maltese record ap-pears to be that of Sommier's collection (FI)which Beck named f. parviflora (BECK, 1922:39). This attribution appears to be dubiousand is discussed further below. This Malteserecord is perpetuated by HASLAM & al. (1970:299) in their Flora of the island.

In addition to these herbarium and litera-ture records, other herbarium specimens havebeen traced. All of these will now be exami-ned in more detail below.

MATERIALS AND METHODS

Most major European herbaria consideredlikely to hold specimens of this taxon have

been contacted for the loan of material. Fromthe response it is evident that collections arevery scarce and this possibly reflects the plant' soverall rarity in nature. Loan specimens andspecimen photographs have been obtainedfrom the following herbaria: E, COI, FI (pho-to), G, LISE, LISU, LTR, MPU, PRC, RNG.

A range of quantitative morphologicalcharacters which are considered to be of taxo-nomic significance within the genus wereexamined on these specimens and where ap-propriate single detached corollas were sof-tened prior to dissection by soaking for fiveminutes in boiling water containing a smallamount of surfactant. Measurements in-cluded: plant height, inflorescence length, in-florescence height, corolla length, corollawidth, overall calyx length, calyx length tothe división of the teeth, and typical height offilament insertion above the corolla base.Estimates have also been made of inflores-cence density, corolla shape, calyx shape andof filament pubescence. The results are shownin Table 1. It should be borne in mind that di-mensions of width are likely exceed to thosefound in living material.

RESULTS AND DISCUSSION

From the results shown in Table 1 it is clearthat the specimens possess many of the maincharacters indicated by POMEL (1874: 107-108) in his original diagnosis of O. clausonis,viz: corollas tubular to campanulate, calycesdivided into four, short, subequal divisions,and filaments pubescent at the base and inser-ted in the lowest quarter of the corolla tube. Inall specimens the short, broadly bifid calyx lo-bes, divided to approximately half their length,are particularly characteristic and the height offilament insertion and degree and distributionof filament pubescence is fairly constantthroughout. Nevertheless, despite the very limi-ted amount of material available, it is partic-ularly noticeable that compared to the Alge-rian specimens, those from the Iberian penin-sula have a much laxer inflorescence, in whichthe corolla is more patent (very erect in thetype) and the corollas themselves appear

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M.J.Y. FOLEY: OROBANCHE CLAUSONIS 323

slightly larger and perhaps more tubular; theseplants are also more robust. It therefore seemsappropriate to give the Iberian plants subspe-cific recognition under O. clausonis.

As well as those Usted in Table 1, otherspecimens previously referred to O. clauso-nis have been examined. Material collectedby Sommier from Malta was described asO. clausonis f. parvifolia BECK (1922: 39).Although not available for loan, detailed pho-tographs of this collection [Sao Paolo, Malta,1907, Sommier (FI)] have been examined. Itwould appear not to be O. clausonis but ispossibly referable to a form of O. caryophyl-lacea, resembling it particularly in corollashape. Another collection of very similarplants (also referred to O. clausonis by the col-lector) is that from the Sierra del Pinar, CádizProvince, Spain [1973, Smythies 829 (LTR)].These specimens, although larger than thosefrom Malta, have very noticeable elongatedcalyx teeth and are clearly not O. clausonis.They again possess some morphological sim-ilarities to O. caryophyllacea. A further spe-cimen referred to O. clausonis is unlocalised,undated and on an unrecorded host [Morís inherb. Pomel (MPU)]. Again, it would appearnot to be that taxon, but rather a variant ofO. caryophyllacea. A single, rather inade-quate specimen from the Balearics [Majorca,1985, Beckett (RNG)], initially referred toO. loricata Rchb., could in fact be O. clauso-nis, but no host is indicated and other impor-tant details are lacking.

The scarcity of records and herbarium mat-erial and comments in the literature (BATTAN-

DIER & TRABUT, 1890: 661) and on severalherbarium labels suggest that the plant is ofrare occurrence. It does not appear to havebeen found in large populations, usually onlyas scattered or isolated individuáis. At the pre-sent time, political considerations in Algeriaprevent field work from being carried outbut when the opportunity arises, a comparisonof living Iberian and Algerian plants shouldbe made. The currently known geographicaldistribution of O. clausonis is mapped in Fi-gure 1.

CLASSMCATION, DESCROTTON AND SUMMARY

Orobanche clausonis Pomel, Nouv. Mat. Fl.Atiant.: 107-108 (1874)

Typus: Parasitic upon Asperula hirsuta atMaziz, Mouzaía [Algeria], 1858, leg. Clau-son (MPU!).

Plant 14-50 cm high, stem pale yellow,slightly glandular-pubescent; leaves lanceo-late, sparse; inflorescence 3-16 cm long, fairlylax to dense; corollas pale yellow, tubular toslightly campanulate, 13-22 mm long, usually15-20 mm, erecto-patent to suberect, dorsalline curved, lower corolla lip prominent,sometimes recurved; bracts lanceolate, 11-20 mm long, glandular-pubescent; calyx lob-es broad, short, 7-9 mm long, each divided toapproximately half their length into two short,more or less equal teeth; filaments hairy be-low, glabrous above, inserted 2-4.5 mm abovebase of corolla; stigma lobes pink or red.Flowering: April - May (-July). Chromosomenumber: - not known.

Geographical distribution: Apparentlyvery local in Portugal, Spain and Algeria.Possibly also in the Balearics.

Habitat: Mainly parasitic upon members ofthe Rubiaceae, but in Portugal recorded also

Fig. 1.-Geographical distribution of Orobanche clauso-nis Pomel, based upon U.T.M. 50 km squares (O subsp.clausonis; • subsp. hesperina; ® unconfirmed literaturerecords).

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324 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

upon Quercus coccifera (= Q. pseudococci-fera) which may be a secondary host. This as-sociation led Welwitsch to annotate his speci-men from Serra de Luiz (LISU) as "O. dryo-phila n.sp." [nomen nudum]. Amongst the Ru-biaceae, Asperula hirsuta, Galium tunetanum,G. verrucosum and Rubia peregrina havebeen recorded. The main habitat is that of itshost - scrub, mattoral, open grassland.

Morphometric affinities and discontinui-ties: The plant has been variously placed inthe O. minor group (BECK, 1890: 234, 1930:218) as well as in association with the dis-similar Orobanche lucorum (GUIMARAES,1904: 85; BECK, 1930: 266). The similarity incolouration and other misconceptions may bea reason for its inclusión in the latter but thereseems to be no justification for any morpho-logical association with the central EuropeanO. lucorum. There are undoubted similaritiesbetween O. clausonis and taxa within theO. minor group, but the very distinctive shapeof the short ovate calyx teeth distinguishO. clausonis from all members of this groupas well as from many other Orobanche taxa.Since certain other morphological differencesare detectable in Orobanche clausonis whenexamined across its geographical range,O. clausonis has been separated into twosubspecies.

a. subsp. clausonis

Geographical distribution: Occurs locallyin northern Algeria, where it is probably rare.

Specimens seenALGERIA: Mouzaüa, 1858, Clauson (MPU), parasiti-

sing Asperula hirsuta (type). Colonne-Voirol, s.d., herb.Battandier (MPU), parasitising Asperula hirsuta. MedeaNador, 1887, herb. Battandier (MPU), parasin'c on Ga-lium tunetanum [poor specimen],

b. subsp. hesperina (J.A. Guim.) M.J.Y. Fo-ley, comb. & stat. nov.

= O. lucorum A. Braun var. hesperinaJ.A. Guim in Brotéria 3: 85 (1904); O. hes-perina (J.A. Guim.) Beck in Engl., Pflan-zenr.96:266(1930)

Typus: The specimen annotated "Oroban-

che Rubiae (Welw.N.Sp.) (ad radices rubii)".Serra da Arrabida, Valle entre El Carmen &Portinho, Mai 1882, leg. / . Daveau (LISU).(Lectotype here designated: left hand spec-imen).

lllustrations: Figure 2; Beck (1890) Taf.IV, no. 72; Sagredo (1987:421).

Description: Differs from the type in itslaxer inflorescence, slightly longer, erecto-patent (not erect/suberect) and perhaps moretubular corollas and in its generally more ro-bust habit.

Geographical distribution: Occurs locallyin central Portugal, and southern Spain. Pos-sibly also in north-west Spain.

Specimens seenSPAIN: Granada, 1849, Boissier & Reuter (PRC) (dis-

section), entire specimen in LE (not seen), parasitisingRubia sp. Near Ronda, 1975, herb. Carr (plus illustra-tion) (RNG), host not recorded.

PORTUGAL: Near Vendas, Serra de S. Luiz, 1845, Wel-witsch (LISU), parasitising Quercus pseudococcifera.Serra de Montejunto, 1848, Welwitsch (LISU), parasi-tising "Rubiae". Serra da Arrabida, 1849, Welwitsch(LISU), parasitising "Rubiae". Between El Carmen andPortinho, Serra da Arrabida, 1882, Daveau (LISU), para-sitising'TÍMMí", annotated "Orobanche Rubiae n.spec.!"(lectotype). Mata do Vidal, Serra da Arrabida, 1952, Sil-va, Fontes & Silva 5080a (LISE), on Rubia peregrina.Bombarral, Serra Mamede, 1948, Rainka 1525 (G,LISE), parasitising Rubia peregrina. Bombarral prox. deRequengo Grande, 1950, Rainha 1955 (LISE), parasiti-sing Rubia peregrina. Nazaré, 1994, Sales & Neves 138(E), parasitising Rubia peregrina.

Note: The occurrence of two host familiesfor O. clausonis subsp. hesperina is perhaps alittle unexpected, but simultaneous attach-ment to the roots of up to three different hostfamilies is not unknown in Orobanche (HIP-KIN, 1992:117) and other parasitised taxa mayhave been overlooked during collection. At theone recently recorded locality [Nazaré, Portu-gal, 1994, Sales & Neves 138 (E)], where Rubiaperegrina was abundant, the observed hostwas initíally thought to be Quercus cocciferabut on re-examination was later confirmed tobe Rubia peregrina (pers. comm. F. Sales).The original Granada specimen collected byBoissier & Reuter in 1849, from which Beckdissected corollas (now in PRC), is in LE buthas unfortunately not been available for exa-

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M.J.Y. FOLEY: OROBANCHE CLAUSONIS 325

Fig. 2. — Orobanche clausonis subsp. hesperina, Ronda, Andalucia, Spain (RNG): A, whole plant; B, corolla, calyx andbract (side view); C, corolla (front view); D, lowerlip of corolla (expanded); E, section of corolla (side view); F, fila-ment attachment to corolla base (expanded).

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326 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

mination. However Beck's dissections havebeen examined and are clearly referrable toO. clausonis. The subsp. hesperina is finelydepicted in S.R. Edwards' painting (Figure 2)made from the living plant of Carr's collec-tion from near Ronda, Spain in 1975 (RNG)(illustration also in RNG). A colour photo-graph of the living plants of Sales & Neves

138 (E) from Nazaré, Portugal has been at-tached to the specimen sheet and other photo-graphs (!) of this collection are held by thecollectors and others. Other records forO. clausonis from Galdo, Spain (MERINO,1906:45), from near Lanjarón, Granada (MU-ÑOZ MEDINA, 1944: 121) and from Barrancodel Caballar, Almería (SAGREDO, 1987: 421)appear to be unsupported by voucher spe-cimens.

ACKNOWLEDGMENTS

I am very grateful to the curators of the herbariaat E, COI, FT, G, LISU, LTR, MPU, PRC and RNGfor their help with loan specimens, to Dr F. Sales(Coimbra) for the loan of colour photographs of theplant in si tu, to Dr S.R. Edwards (Manchester) forpermission to publish his drawing of O.clausonissubsp. hesperina. I would also like to thankDr N.K.B. Robson (London) for his advice on no-menclature, Dr G. Halliday (Lancaster) for com-menting on the draft of this paper, and the Trusteesof the Botanical Research Fund and of the WelchBequest for financial assistance towards thisproject.

REFERENCES

BATTANDER, J-A. & L. TRABUT (1890). Flore de l'Algé-rie. Vol. 4. Alger & Paris.

BECK, G. (1890). Monographie der Gattung Orobanche.Biblioth. Bot. 19.

BECK, G. (1922). Orobancheae novae. Repert. Spec. Nov.RegniVeg.U-.39.

BECK, G. (1930). Orobanchaceae. In: A. Engler (ed.),Das Pflanzenreich. Vol. 96. Leipzig.

BOLOS, O. DE, R.M. MASALLES, J.M. NINOT & J. VIGO

(1990). Flora Manual deis Paisos Catalans. Barcelona.CHATER, A.O. & D.A. WEBB (1972). Orobanche L. In: T.G.

Tu™, V.H. HEYWOOD, N.A. BURGES, D.M. MOORE,

D.H. VALENTINE, S.M. WALTERS & D.A. WEBB (eds.).

Flora Europaea. Vol. 3. Cambridge.COSTA, A.C. (1877). Introducción á la Flora de Catalu-

ña. Barcelona.COUTINHO, A.X.P. (1913). A Flora de Portugal. Lisboa.COUTINHO, AX.P. (1939). Flora de Portugal, ed. 2. Lisboa.DUBY, J.E. (1828). Aug. Pyrami de Candolle Botanicon

gallicum seu Synopsis Plantarum in Flora gallica des-criptarum. Vol. 1. Paris.

FERNANDEZ-GALIANO, E.F. & V.H. HEYWOOD (1960).

Catálogo de plantas de la provincia de Jaén. Jaén.FRANCO, J.A. (1984). Nova flora de Portugal. Vol. 2. Lis-

boa.GUIMARAES, J.D'A. (1904). Monographia das Oroban-

chaceas Portuguezas. Brotéria 3.HASLAM, S., P.D. SELL & P.A. WOLESEY (1970). A Flora

ofthe Maltese Islands. Msida, Malta.HIPKIN, C.R. (1992). Host range and specificity of Oro-

banche minor Sm. on Crymlyn Burrows. Watsonia 19:117.

MERINO, B. (1906). Flora Descriptiva é Ilustrada de Ga-licia. Vol. 3. Santiago de Compostela.

MUÑOZ-MEDINA, J.M. (1944). Lanjarón como localidadbotánica. Farmacognosia 3(5): 121.

NÉGRE, R. (1962). Petite Flore des Régions Arides duMaroc Occidental. Vol. 2. Paris.

POMEL, A.N. (1874). Nouveaux Matériaux pour la FloreAtlantique. Paris & Argel.

QUEZEL, P. & S. SANTA (1963). Nouvelle Flore de VAlgé-rie et des Régions Désertiques Meridionales. Vol. 2.Paris.

ROMO, A.M. (1989). Flora i Vegetado delMontsec (Pre-Pirineus Catalans). Barcelona.

SAGREDO, R. (1987). Flora de Almería. Almería.SAMPAIO, G. (1947). Flora Portuguesa. Porto.SANZ, G.M. (1990). Catálogo florístico de la provincia de

Teruel. Teruel.SANZ, G.M. (1992). Claves para la flora de la provincia

de Teruel. Teruel.SAUVAGE, CH. (1961). Flore des.Subéraines Marocaines.

Rabat.VALDÉS, B., S. TALAVERA & E. FERNANDEZ-GALIANO

(eds.). (1987). Flora Vascular de Andalucía Occiden-tal. Vol. 2. Barcelona.

WILLKOMM, M. (1893). Supplementum Prodromi FloraeHispanicae. Stuttgart.

WILLKOMM, M. & J. LANGE (1870). Prodromus FloraeHispanicae. Vol. 2. Stuttgart.