Orobanche Clausonis Pomel (Orobanchaceae) in the Iberian Península

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. hesperina (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 parasitic upon Asperula hirsuta (Rubiaceae). Since then, germine records have been scarce and although 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). 320 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996 identification and under-recording rather than O. clausonis, although it was included in the a reflection of extreme rarity, although it is subsequent supplement (WILKOMM, 1893: undoubtedly uncommon. 188) based on BECK'S (1890:235) verification In his original description of O. clausonis, of the Granada specimen. Later, when Gui- POMEL (1874: 107) remarked on several im- maraes published his monograph on Portu- portant characters: the rather dense inflores- guese Orobanche he included under O. luco- cence, the tubular to campanulate corollas, rum A. Braun (GUIMARAES, 1904: 85) a new the calyx divided into four shortly lanceolate, variety hesperina based on plants from four subequal divisions, and the insertion of the fil- collections by Welwitsch and by Daveau. aments in the lowest quarter of the corolla These were parasitic upon Rubia spp. and in tube, the filaments themselves being slightly one case upon Quercus pseudococcifera and pubescent at their base. Of these characters, it were collected from a rather limited área of is the shape of the broad, relatively short, div- central Portugal. From elsewhere in his mono- ided calyx as evidenced in the type specimen graph it is clear that Guimaraes was aware of (Mouzaía, 1858; Clauson, MPU), which es- the principal characters of Pomel's O. clauso- pecially separates O. clausonis from similar nis since he occasionally quotes such charact- taxa. ers in comparison to other taxa and he placed it Following Pomel's diagnosis, there appe- cióse to O. mauretanica Beck (grex Minores). ars to be no mention of O. clausonis until a However he did not consider O. clausonis to be a Portuguese plant ñor did he refer to it in dismissive comment by BATTANDDER & TRA- BUT (1890: 661) that it was merely a glabres- relation to his O. lucorum var. hesperina. His cent form of O. galii Vaucher var. atlantica attribution of the Welwitsch and Daveau spe- having more hispid filaments and hairy ant- cimens to O. lucorum is intriguing and a pos- hers. This latter taxon, apparently parasitic sible explanation may lie in a combination of upon Galium tunetanum, is a small-flowered factors. True O. lucorum, parasitic on Berberis variant of the widespread European O. caryo- spp., has a narrow distribution in central Eur- phyllacea Sm., a very different and distinctive ope, occurring essentially over a limited área plant to O. clausonis, again apparently restric- of the eastern Alps. It is readily separated from ted in its host to members of the Rubiaceae. It Guimaraes' Portuguese var. hesperina on is not clear whether Battandier & Trabut's calyx shape alone - the lobes of O. lucorum comments were based upon actual experience being much longer and narrower that in var. of O. clausonis, but any affinities between hesperina. On the relatively limited informa- it and variants of O. caryophyllacea can be tion available at the time, it is possible that dismissed upon morphological grounds. In Guimaraes associated his var. hesperina with O. lucorum based on similar colouration (yel- the same year, BECK (1890: 234) included O. clausonis in his monograph of the genus low). Also, a variant or segregate of O. luco- Orobanche, listing Clauson's Algerian loca- rum from south-eastern France parasitic upon lity, although not having seen the original spe- Rubus spp. had been described as O. rubi cimen. He also added a record of Boissier's (DUBY, 1828:350). This, together with the fact and Reuter's from Granada, Spain, in 1849 that Welwitsch had annotated one of his sheets (specimen now in LE - corolla dissections in "O. rubii", may have led to a possible misin- PRC!) which at that time appeared to be the terpretation of their respective hosts and there- first and only European record for the plant. fore to the connection with O. lucorum. Ne- He surprisingly placed O. clausonis cióse to vertheless, Guimaraes did indícate that the taxa of the O. minor group (i.e. in his grex Mi- specimens of Portuguese "O. rubi Duby" nores) despite its very different calyx shape, which he had seen differed from typical O. lu- which contrasts with the elongated, tapering corum in several respects and for that reason calyces of members of his grex Minores. he gave them the varietal rank hesperina. In Prodromus Florae Hispanicae, WlL- In Flore de Portugal, COUTINHO (1913: KOMM & LANGE (1870) made no reference to 569) took a different view and accepted M.J.Y. FOLEY: OROBANCHE CLAUSONIS 321

O. clausonis as a Portuguese plant. This was been contacted for the loan of material. From presumably based upon the same specimens the response it is evident that collections are considered by Guimaraes under his O. luco- very scarce and this possibly reflects the plant' s rum var. hesperina, since he usted the locali- overall rarity in nature. Loan specimens and ties and hosts of Welwitsch's collections, an- specimen photographs have been obtained notated some of the sheets and added a taxo- from the following herbaria: E, COI, FI (pho- nomic description. He did not include O. lu- to), G, LISE, LISU, LTR, MPU, PRC, RNG. corum for Portugal. This stance has been A range of quantitative morphological maintained subsequently for Portugal (COU- characters which are considered to be of taxo- TINHO, 1939:673; SAMPAIO, 1946:498; FRAN- nomic significance within the genus were CO, 1984:280) but O. clausonis is excluded in examined on these specimens and where ap- works covering Algeria and Morocco (SAU- propriate single detached corollas were sof- VAGE, 1961; NÉGRE, 1962; QUEZEL & SANTA, tened prior to dissection by soaking for five 1963) and most regions of Spain (COSTA, minutes in boiling water containing a small 1864; FERNÁNDEZ-GALIANO & HEYWOOD, amount of surfactant. Measurements in- 1960; VALDÉS & al., 1987; ROMO, 1989; cluded: plant height, inflorescence length, in- BOLOS & al., 1990; SANZ, 1990, 1992). Ho- florescence height, corolla length, corolla wever, in eastern Andalucía there is a single width, overall calyx length, calyx length to record by SAGREDO (1987: 421) of plants the división of the teeth, and typical height of parasitic upon Galium verrucosum at Barran- filament insertion above the corolla base. co del Caballar, Almería and another by MU- Estimates have also been made of inflores- ÑOZ-MEDINA (1944:121) from near Lanjarón, cence density, corolla shape, calyx shape and Granada province. Two collections from Gal- of filament pubescence. The results are shown do, north-west Spain, and mentioned by ME- in Table 1. It should be borne in mind that di- RINO (1906: 45) under O. lucorum, may also mensions of width are likely exceed to those be O. clausonis. FRANCO (1984: 280) recently found in living material. extended the Portuguese range of O. clausonis to include the Atlantic off-shore Ilhas Ber- lengas. RESULTS AND DISCUSSION

In Flora Europaea, CHATER & WEBB From the results shown in Table 1 it is clear (1972: 292) listed it as present in Spain, Por- that the specimens possess many of the main tugal and Malta, based upon information gi- characters indicated by POMEL (1874: 107- ven in BECK'S monographs (1890: 235; 1930: 108) in his original diagnosis of O. clausonis, 218). However, the solé Maltese record ap- viz: corollas tubular to campanulate, calyces pears to be that of Sommier's collection (FI) divided into four, short, subequal divisions, which Beck named f. parviflora (BECK, 1922: and filaments pubescent at the base and inser- 39). This attribution appears to be dubious ted in the lowest quarter of the corolla tube. In and is discussed further below. This Maltese all specimens the short, broadly bifid calyx lo- record is perpetuated by HASLAM & al. (1970: bes, divided to approximately half their length, 299) in their Flora of the island. are particularly characteristic and the height of In addition to these herbarium and litera- filament insertion and degree and distribution ture records, other herbarium specimens have of filament pubescence is fairly constant been traced. All of these will now be exami- throughout. Nevertheless, despite the very limi- ned in more detail below. ted amount of material available, it is particularly noticeable that compared to the Alge- MATERIALS AND METHODS rian specimens, those from the Iberian penin- sula have a much laxer inflorescence, in which Most major European herbaria considered the corolla is more patent (very erect in the likely to hold specimens of this taxon have type) and the corollas themselves appear 322 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996 M.J.Y. FOLEY: OROBANCHE CLAUSONIS 323 slightly larger and perhaps more tubular; these CLASSMCATION, DESCROTTON AND SUMMARY plants are also more robust. It therefore seems appropriate to give the Iberian plants subspe- Orobanche clausonis Pomel, Nouv. Mat. Fl. cific recognition under O. clausonis. Atiant.: 107-108 (1874) As well as those Usted in Table 1, other Typus: Parasitic upon Asperula hirsuta at specimens previously referred to O. clauso- Maziz, Mouzaía [Algeria], 1858, leg. Clau- nis have been examined. Material collected son (MPU!). by Sommier from Malta was described as O. clausonis f. parvifolia BECK (1922: 39). Plant 14-50 cm high, stem pale yellow, Although not available for loan, detailed pho- slightly glandular-pubescent; leaves lanceo- tographs of this collection [Sao Paolo, Malta, late, sparse; inflorescence 3-16 cm long, fairly 1907, Sommier (FI)] have been examined. It lax to dense; corollas pale yellow, tubular to would appear not to be O. clausonis but is slightly campanulate, 13-22 mm long, usually possibly referable to a form of O. caryophyl- 15-20 mm, erecto-patent to suberect, dorsal lacea, resembling it particularly in corolla line curved, lower corolla lip prominent, shape. Another collection of very similar sometimes recurved; bracts lanceolate, 11- 20 mm long, glandular-pubescent; calyx lob- plants (also referred to O. clausonis by the col- es broad, short, 7-9 mm long, each divided to lector) is that from the Sierra del Pinar, Cádiz approximately half their length into two short, Province, Spain [1973, Smythies 829 (LTR)]. more or less equal teeth; filaments hairy be- These specimens, although larger than those low, glabrous above, inserted 2-4.5 mm above from Malta, have very noticeable elongated base of corolla; stigma lobes pink or red. calyx teeth and are clearly not O. clausonis. Flowering: April - May (-July). Chromosome They again possess some morphological sim- number: - not known. ilarities to O. caryophyllacea. A further specimen referred to O. clausonis is unlocalised, Geographical distribution: Apparently undated and on an unrecorded host [Morís in very local in Portugal, Spain and Algeria. herb. Pomel (MPU)]. Again, it would appear Possibly also in the Balearics. not to be that taxon, but rather a variant of Habitat: Mainly parasitic upon members of O. caryophyllacea. A single, rather inade- the Rubiaceae, but in Portugal recorded also quate specimen from the Balearics [Majorca, 1985, Beckett (RNG)], initially referred to O. loricata Rchb., could in fact be O. clausonis, but no host is indicated and other impor- tant details are lacking. The scarcity of records and herbarium material and comments in the literature (BATTAN- DIER & TRABUT, 1890: 661) and on several herbarium labels suggest that the plant is of rare occurrence. It does not appear to have been found in large populations, usually only as scattered or isolated individuáis. At the present time, political considerations in Algeria prevent field work from being carried out but when the opportunity arises, a comparison of living Iberian and Algerian plants should be made. The currently known geographical Fig. 1.-Geographical distribution of Orobanche clausonis Pomel, based upon U.T.M. 50 km squares (O subsp. distribution of O. clausonis is mapped in Fi- clausonis; • subsp. hesperina; ® unconfirmed literature gure 1. records). 324 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

upon Quercus coccifera (= Q. pseudococci- che Rubiae (Welw.N.Sp.) (ad radices rubii)". fera) which may be a secondary host. This as- Serra da Arrabida, Valle entre El Carmen & sociation led Welwitsch to annotate his speci- Portinho, Mai 1882, leg. /. Daveau (LISU). men from Serra de Luiz (LISU) as "O. dryo- (Lectotype here designated: left hand spec- phila n.sp." [nomen nudum]. Amongst the Ru- imen). biaceae, Asperula hirsuta, Galium tunetanum, G. verrucosum and Rubia peregrina have lllustrations: Figure 2; Beck (1890) Taf. been recorded. The main habitat is that of its IV, no. 72; Sagredo (1987:421). host - scrub, mattoral, open grassland. Description: Differs from the type in its Morphometric affinities and discontinui- laxer inflorescence, slightly longer, erecto- ties: The plant has been variously placed in patent (not erect/suberect) and perhaps more the O. minor group (BECK, 1890: 234, 1930: tubular corollas and in its generally more ro- 218) as well as in association with the dis- bust habit. similar Orobanche lucorum (GUIMARAES, Geographical distribution: Occurs locally 1904: 85; BECK, 1930: 266). The similarity in in central Portugal, and southern Spain. Pos- colouration and other misconceptions may be sibly also in north-west Spain. a reason for its inclusión in the latter but there seems to be no justification for any morpho- Specimens seen logical association with the central European SPAIN: Granada, 1849, Boissier & Reuter (PRC) (dis- O. lucorum. There are undoubted similarities section), entire specimen in LE (not seen), parasitising Rubia sp. Near Ronda, 1975, herb. Carr (plus illustra- between O. clausonis and taxa within the tion) (RNG), host not recorded. O. minor group, but the very distinctive shape PORTUGAL: Near Vendas, Serra de S. Luiz, 1845, Wel- of the short ovate calyx teeth distinguish witsch (LISU), parasitising Quercus pseudococcifera. O. clausonis from all members of this group Serra de Montejunto, 1848, Welwitsch (LISU), parasi- as well as from many other Orobanche taxa. tising "Rubiae". Serra da Arrabida, 1849, Welwitsch (LISU), parasitising "Rubiae". Between El Carmen and Since certain other morphological differences Portinho, Serra da Arrabida, 1882, Daveau (LISU), para- are detectable in Orobanche clausonis when sitising'TÍMMí", annotated "Orobanche Rubiae n.spec.!" examined across its geographical range, (lectotype). Mata do Vidal, Serra da Arrabida, 1952, Sil- O. clausonis has been separated into two va, Fontes & Silva 5080a (LISE), on Rubia peregrina. Bombarral, Serra Mamede, 1948, Rainka 1525 (G, subspecies. LISE), parasitising Rubia peregrina. Bombarral prox. de Requengo Grande, 1950, Rainha 1955 (LISE), parasitising Rubia peregrina. Nazaré, 1994, Sales & Neves 138 a. subsp. clausonis (E), parasitising Rubia peregrina. Geographical distribution: Occurs locally Note: The occurrence of two host families in northern Algeria, where it is probably rare. for O. clausonis subsp. hesperina is perhaps a little unexpected, but simultaneous attach- Specimens seen ment to the roots of up to three different host ALGERIA: Mouzaüa, 1858, Clauson (MPU), parasitising Asperula hirsuta (type). Colonne-Voirol, s.d., herb. families is not unknown in Orobanche (HIP- Battandier (MPU), parasitising Asperula hirsuta. Medea KIN, 1992:117) and other parasitised taxa may Nador, 1887, herb. Battandier (MPU), parasin'c on Ga- have been overlooked during collection. At the lium tunetanum [poor specimen], one recently recorded locality [Nazaré, Portu- gal, 1994, Sales & Neves 138 (E)], where Rubia b. subsp. hesperina (J.A. Guim.) M.J.Y. Fo- peregrina was abundant, the observed host ley, comb. & stat. nov. was initíally thought to be Quercus coccifera = O. lucorum A. Braun var. hesperina but on re-examination was later confirmed to J.A. Guim in Brotéria 3: 85 (1904); O. hes- be Rubia peregrina (pers. comm. F. Sales). perina (J.A. Guim.) Beck in Engl., Pflan- The original Granada specimen collected by zenr.96:266(1930) Boissier & Reuter in 1849, from which Beck dissected corollas (now in PRC), is in LE but Typus: The specimen annotated "Oroban- has unfortunately not been available for exa- M.J.Y. FOLEY: OROBANCHE CLAUSONIS 325

Fig. 2. — Orobanche clausonis subsp. hesperina, Ronda, Andalucia, Spain (RNG): A, whole plant; B, corolla, calyx and bract (side view); C, corolla (front view); D, lowerlip of corolla (expanded); E, section of corolla (side view); F, filament attachment to corolla base (expanded). 326 ANALES JARDÍN BOTÁNICO DE MADRID, 54.1996

mination. However Beck's dissections have BOLOS, O. DE, R.M. MASALLES, J.M. NINOT & J. VIGO been examined and are clearly referrable to (1990). Flora Manual deis Paisos Catalans. Barcelona. CHATER, A.O. & D.A. WEBB (1972). Orobanche L. In: T.G. O. clausonis. The subsp. hesperina is finely Tu™, V.H. HEYWOOD, N.A. BURGES, D.M. MOORE, depicted in S.R. Edwards' painting (Figure 2) D.H. VALENTINE, S.M. WALTERS & D.A. WEBB (eds.). made from the living plant of Carr's collec- Flora Europaea. Vol. 3. Cambridge. tion from near Ronda, Spain in 1975 (RNG) COSTA, A.C. (1877). Introducción á la Flora de Catalu- (illustration also in RNG). A colour photo- ña. Barcelona. COUTINHO, A.X.P. (1913). A Flora de Portugal. Lisboa. graph of the living plants of Sales & Neves COUTINHO, AX.P. (1939). Flora de Portugal, ed. 2. Lisboa. 138 (E) from Nazaré, Portugal has been at- DUBY, J.E. (1828). Aug. Pyrami de Candolle Botanicon tached to the specimen sheet and other photo- gallicum seu Synopsis Plantarum in Flora gallica des- graphs (!) of this collection are held by the criptarum. Vol. 1. Paris. collectors and others. Other records for FERNANDEZ-GALIANO, E.F. & V.H. HEYWOOD (1960). O. clausonis from Galdo, Spain (MERINO, Catálogo de plantas de la provincia de Jaén. Jaén. FRANCO, J.A. (1984). Nova flora de Portugal. Vol. 2. Lis- 1906:45), from near Lanjarón, Granada (MU- boa. ÑOZ MEDINA, 1944: 121) and from Barranco GUIMARAES, J.D'A. (1904). Monographia das Oroban- del Caballar, Almería (SAGREDO, 1987: 421) chaceas Portuguezas. Brotéria 3. appear to be unsupported by voucher spe- HASLAM, S., P.D. SELL & P.A. WOLESEY (1970). A Flora cimens. ofthe Maltese Islands. Msida, Malta. HIPKIN, C.R. (1992). Host range and specificity of Oro- banche minor Sm. on Crymlyn Burrows. Watsonia 19: 117. ACKNOWLEDGMENTS MERINO, B. (1906). Flora Descriptiva é Ilustrada de Ga- licia. Vol. 3. Santiago de Compostela. I am very grateful to the curators of the herbaria MUÑOZ-MEDINA, J.M. (1944). Lanjarón como localidad at E, COI, FT, G, LISU, LTR, MPU, PRC and RNG botánica. Farmacognosia 3(5): 121. for their help with loan specimens, to Dr F. Sales NÉGRE, R. (1962). Petite Flore des Régions Arides du (Coimbra) for the loan of colour photographs of the Maroc Occidental. Vol. 2. Paris. plant in si tu, to Dr S.R. Edwards (Manchester) for POMEL, A.N. (1874). Nouveaux Matériaux pour la Flore permission to publish his drawing of O.clausonis Atlantique. Paris & Argel. QUEZEL, P. & S. SANTA (1963). Nouvelle Flore de VAlgé- subsp. hesperina. I would also like to thank rie et des Régions Désertiques Meridionales. Vol. 2. Dr N.K.B. Robson (London) for his advice on no- Paris. menclature, Dr G. Halliday (Lancaster) for com- ROMO, A.M. (1989). Flora i Vegetado delMontsec (Pre- menting on the draft of this paper, and the Trustees Pirineus Catalans). Barcelona. of the Botanical Research Fund and of the Welch SAGREDO, R. (1987). Flora de Almería. Almería. Bequest for financial assistance towards this SAMPAIO, G. (1947). Flora Portuguesa. Porto. project. SANZ, G.M. (1990). Catálogo florístico de la provincia de Teruel. Teruel. SANZ, G.M. (1992). Claves para la flora de la provincia REFERENCES de Teruel. Teruel. SAUVAGE, CH. (1961). Flore des.Subéraines Marocaines. BATTANDER, J-A. & L. TRABUT (1890). Flore de l'Algé- Rabat. rie. Vol. 4. Alger & Paris. VALDÉS, B., S. TALAVERA & E. FERNANDEZ-GALIANO BECK, G. (1890). Monographie der Gattung Orobanche. (eds.). (1987). Flora Vascular de Andalucía Occiden- Biblioth. Bot. 19. tal. Vol. 2. Barcelona. BECK, G. (1922). Orobancheae novae. Repert. Spec. Nov. WILLKOMM, M. (1893). Supplementum Prodromi Florae RegniVeg.U-.39. Hispanicae. Stuttgart. BECK, G. (1930). Orobanchaceae. In: A. Engler (ed.), WILLKOMM, M. & J. LANGE (1870). Prodromus Florae Das Pflanzenreich. Vol. 96. Leipzig. Hispanicae. Vol. 2. Stuttgart.