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Investigations on thegenus Cerastium L.(Caryophyllaceae): typus, locusclassicus and distribution of C.lineare AllNeda Bechi a & Giuliana Forneris ba Dipartimento di Biologia Vegetale dell'Università,Via La Pira 4, I-50121, Firenzeb Erbario del Dipartimento di Biologia Vegetaledell'Università, Viale P.A. Mattioli 25, I-10125, Torino

To cite this article: Neda Bechi & Giuliana Forneris (1998): Investigations on thegenus Cerastium L. (Caryophyllaceae): typus, locus classicus and distribution of C.lineare All, Webbia: Journal of Plant Taxonomy and Geography, 53:1, 13-25

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Webbia 53(1): 13-25. 1998

Investigations on the genus Cerastium L. (Caryophyllaceae): typus, locus classicus and distribution of C. lineare All.*

NEDA BEcm 1, GruUANA FoRNERis 2

'Dipartimento eli Biologia Vegetale dell'Universita Via La Pira 4, 1-50121 Firenze 2 Erbario del Dipartimento eli Biologia Vegetale dell'Universita Viale P.A. Mattioli 25,1-10125 Torino

Ricevuto il13 Giugno 1997 Accettato il23 Gennaio 1998

Investigations on Cerastium L. (Caryophyllaceae): typus, locus classicus and distribution of C. lineare All. - The name Cerastium lineare All. was published by Allioni in 1785. A specimen belonging to Bellardi's collection, preserved in TO, is here proposed as lectotype. Remarks on the identification of the locus classicus are also given. Some aspects of hairs, capsules and seeds seem to show that this diploid, endemic to the Western Alps, is related to the Cerastium alpinum L. group. A detailed description, a new iconography, and information on its ecology and distribution are also provided.

Key words: Caryophyllaceae, Cerastium lineare, chorology, systematics, typifica­tion.

During a biosystematic, taxonomic and phytogeographic study on some critical groups of the genus Cerastium - the C. banaticum (Rochel) Heuffel group [ex C. scaraniiTen. sensu BuscHMANN (1938) group], C. arvense L. group and the C. tomentosum L. group (BECHI et al., 1992; BARBERIS et al., 1994, 1995; BEcHI & MICELI, 1995; BECHI, 1998a, 1998b; MicELI et al., 1997; BECHI 1998c; BECHI et al., 1998)- C. lineare, endemic to the Western Alps, was examined. Several authors have expressed controversial opinions on its systematic position (NEGRI, 1904; BuscHMANN, 1938; HEss et al., 1967; FAVARGER, 1973; MoscHL,

1982). In order to discuss the matter satisfactorily, it was necessary necessary to designate the type on which the Allionian name is based. This procedure, however, required in-depth nomenclatural, morphological and chorological investigations, particularly with regard to its locus classicus.

MATERIALS AND METHODS

In order to proceed with the typification of C. lineare, the references in the works of Allioni as well as the iconographic tables that the Author commissioned were examined for necessary

*Research funded with contribution from the Ministry for University and Scientific and Technological Research (40% M.U.R.S.T. funds).

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14 NEDA BECHI, GIULIANA FORNERIS

comparisons. Obviously, we examined all the material in the collections at TO (Allioni, Bellardi, HP e HG), FI, FI-W, BOLO, G and G-DC, all possible centres where some of Allioni's specimens could have turned up following exchanges which the Botanist had with other contem­porary scholars (SINISCALCO & FoRNERIS, 1985-86) or after subsequent reorganization of his herbarium (FORNERIS, 1993, pp. 276-277).

The most important Italian and European Floras were consulted regarding the nomenclature and respective taxonomical classifications. Moreover, a detailed description of the morphology of C. lineare is given from observations on herbarium specimens and fresh material, seen under the newly proposed figure, together with references to the most important iconographic tables available in the literature. On the basis of the herbarium material inspected (BM, Fl, FI-W, PAD, TO) the distribution of C. lineare has been defined for Piedmont, Valle d'Aosta and France using the Map of the Floristic, Ecological and Vegetation Sectors proposed by MoNTACCHINI & FoRNERis (1980).

TYPIFICATION

The publication of the name Cerastium lineare by ALLIONI (1785), in Flora Pedemontana, must be considered as two distinct parts. The first part is included in the text at n° 1729 (2:117) where the Author cites Cerastium strictum L. with the Linnaean diagnostic phrase and the previous polynomial synonyms. In the detailed description which follows, Allioni underlines certain morphological characters which differentiate his specimens both from the analysis given by Linnaeus (" .. .fructus cylindricus ... " " ... sed fructus globosi non sunt ... ") and the illustration published by HALLER (1768, tab. XIV) for C. strictum. Accordingly, he proposed a new iconographical reference, citing fig. 1 from table 18 in vol. XII in the Iconographia Taurinensis 1•

ALuoNI (1785) mentions the Valli Valdesi Alps (Val Chisone and Val Germanasca) and Mount Cenisio (Vaile di Sus a) as localities where the occur­rence of this species was observed.

The second part - the protologue - can be found in "Addenda et Emendanda" (2:365) where Cerastium lineare N[obis] is cited at n° 1729*2 and is completed with a new diagnostic phrase: "Cerastium foliis linearibus flac­cidis aristatis, pedunculis subunifloris tomentosis, capsulis oblongis". Allioni states clearly that the description given under C. strictum is valid for C. lineare, and the localities where the species occurs are also reconfirmed, to which,

1 For an exhaustive reference on the collection of water colour tables in the Iconographia Taurinensis, the reader is referred to CHIAPusso Vou ( 1904). The importance given by Allioni to the illustrations in the Iconographia Taurinensis, as scientific documents, can be seen in FORNERIS (1996). The title page of vol. XII of the Iconographia Taurinensis mentioned by Allioni, bears the date 1770.

2 In this part of the Allionian text, the Author gives corrections and additional data to the citations contained in Flora Pedemontana before the work was published. The asterisk after number "1729" indicates that a previously non-cited species should be inserted following the systematic order proposed by Allioni.

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INVESTIGATIONS ON THE GENUS CERASTIUM L. (CARYOPHYLLACEAE) 15

however, he adds the Viu Alps (Valli di Lanzo) following information from Bellardi3 .

As further support to his own description, Allioni.publishes fig. 4, table 88 in Flora Pedemontana, but also refers to table 46 in val. XXVII (sphalm. for :xx\TIII) of the Iconographia Taurinensis. This latter illustration can be consid­ered to be the original drawing commissioned by Allioni, which was used as the matrix for the print; in fact the two coincide and are mirror images of each other (FORNERIS, 1985-86) (fig. la, lb).

In Auctarium ad Floram Pedemontanam (31: n° 1729*), ALLIONI (1789) reconfirms the occurrence of C. lineare in Valli di Lanzo and adds the Alpi di Giaveno (Val Sangone). Our research in G-DC and BOLO did not highlight any autoptic material, although during the 19th century, many specimens of Allioni arrived at these Herbaria 4 •

For type identification, our research therefore concentrated on specimens listed sub C. lineare All. and sub C. strictum L. in TO:

1- "Cerastium lineare" (TO- Herb. Allioni sub C. lineare)

The binomial at the foot of the sheet was written by I. Molineri. There are two flowering stems complete with basal leaves, flowers and capsules.

2- "Cerastium strictum" (TO-HG sub C. strictum)

The specimen, found in the Herbarium Generate under C. strictum, is part of the Allioni collection that can be seen from the sheet typology, the prepara­tion of the specimen and from the handwriting of Allioni who only wrote the binomial. The exsiccatum corresponds to C. lineare and consists of five flower­ing stems two of which are arranged under the same paper strip.

3 - "Cerastium strictum L. et Allion. Stirp. Pedem." (TO - Herb. Bellar eli)

The sheet reports, manu Bellardi, a short diagnosis: "Cerastium foliis linearibus acuminatis, glabris, pedunculis unifloris subtomentosis, capsulis

3 For the Bellardi's contribution to the scientific work of Allioni, see FoRNERIS eta!. (1989). 4 A.P. De Candolle often exchanged herbarium specimens with G.B. Balbis, who owned the Allioni

collection untill831. Material from the I. Molineri collection (who was the closest collaborator of Allioni) are kept in the A. Bertoloni collection (FORNERIS et al., 1991 pp.108-109). In BOLO a specimen of C.lineare All. exists with a label manu Molineri.

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Fig. 1 - 1a: Cerastium lineare All. in Iconographia Taurinensis, vol. XXVIII, tab. 46. Watercolour painting attributed to A. Bottione. 1b: Cerastium lineare All. in Flora Pedemontana by C. Allioni (1785): tab. LXXXVIII, fig. 4. Engraving.

globosis binfr. M[. .. ] p. 3 [. .. ]"; some observations: "Habitus totius planta laxus, foliorum glabritias absque acumine, capsularum figura non globosa, sed oblonga repugnant Cerastio stricto."; a very significant phrase: "Meo suasu hanc plantam distinxit Cl. Allionius, dixitque Cerastium lineare Nob. Fl. ped. "; and refers to the collection site of the specimen: "lnveni in alpibus Viucensibus. D. CatochiP." (fig. 2).

5 The note "D. Catochii" refers to a scholar originally from Valle di Vii\ with whom Bellardi exchanged material and informations. The collaboration between Catochi and Bellardi is often mentioned in the herbarium.

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INVESTIGATIONS ON THE GENUS CERASTIUM L. (CARYOPHYLLACEAE) 17

Fig. 2 - Handwritten notes by L. Bellardi on the folder containing the specimen of Cerastium lineare All. kept in his collection.

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There are two flowering stems pinned together: they are complete with leaves, flowers and fruits and are clearly identifiable as C. lineare.

4- "Cerastium lineare Fl. ped. tab. 88 fig. 4" (TO- Herb. Bellardi)

The notes, written in hand by Bellardi, refer to the publication of C. lineare in Flora Pedemontana and the relative printed illustration. The collecting site is mentioned at the foot of the sheet: "Reperi abunde in alpibus di Viti alibique similibus locis". The specimen consists of three flowering stems pinned to­gether and of three free fragments, corresponding to C. lineare.

CONCLUSIONS

Observations on the examined material are as follows: 1 - although exhibiting close resemblance to the drawing which Allioni originally chose as the iconographic reference (Icon. Taur. vol. XII, tab. 18 fig. 1), the first specimen described as n. 1 does not bear traces of intervention by the Author. 2- the second (n. 2) was definitely analysed by Allioni, who designated it as C. strictum. However, the lack of any correlation with the doubts already ex­pressed when publishing C. strictum - or subsequent corrections - plus the fact that the collecting sites mentioned in the first description by Allioni in Flora Pedemontana (2: 117) are not reported, justify the preference for other material. 3 - the specimen described as n. 3, although not included in the Allioni collection, bears obvious evidence of study by the Author, referred to by Bellardi and cited in the protologue (fig. 3 ). 4 -the exsiccatum n. 4 does not exhibit any proof of study by Allioni.

We therefore retain that the specimen here referred to as n. 3 should be designated as lectotype of the name Cerastium lineare. This choise is based on the personal observations of Bellardi about the poor correlation with the morphological characters referred to by Linnaeus for C. strictum - also published by Allioni- and the note "Meo suasu hanc plantam distinxit Cl. Allionius, dixitque Cerastium lineare Nob. Fl. ped. ".

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The formal citation is:

Cerastium lineare All., Fl. Pedem. 2:365, no 1729*. 1785.

LT. (here designated): "Cerastium strictum L. et Allion. Stirp. Pedem." (TO, Herb. Bellardi!) (fig. 3 ). The locality referred to on the specimen "alpibus Viucensibus", reported in the protologue, is therefore the locus classicus of Cerastium lineare All.

(=) Cerastium strictum L. var. lineare (All.) DC., Fl. franc. 5: 610. 1815 (=) Cerastium repens L. ssp. lineare (All.) Nyman, Consp. fl. europ.: 108. 1878 (=) Cerastium arvense L. ssp.lineare (All.) Arcang.,Comp. fl. Ital.: 99. 1882 (=) Cerastium repens L. var. lineare (All.) Cesati, Passerini & Gibelli, Comp. Fl. Ital.: 784. 1883 (=) Cerastium arvense L. var. lineare (All.) Fiori, Fl. Anal. Ital. 1: 354. 1898 (=) Cerastium tomentosum L. var. lineare (All.) Giirke in Richter, Pl. Europaeae 2: 216. 1899 (=) Cerastium alpinum L. ~ /oliis lanceolato-linearibus, linearibusque Bertol., Fl. Ital. 4: 762. 1839. (-) Cerastium laricifolium ViTI. sensu Schur, Enum. pl. trans.: 124. 1866, pro parte.

IcoNOGRAPHY- Ar.uoNI (1785) t. 88, fig. 4; NEGRI (1904) t. 8; HEss et al. (1967) fig. 818; FAVARGER (1973) pl. 35, fig. 478; MoscHL (1982) n. 482; our illustration (fig. 4).

DESCRIPTION- Perennial, 5-40 em, stoloniferous; stem sparsely tomentose to glabrescent, with hairs arranged in two rows. Leaves of flowering stems (15-) 20-35 (-50) x (1-) 1.9-2.5 (-4) mm, linear, acute, glabrous or glabrescent, ciliate at the base. Leaves of procumbent non-flowering stems (1.5-) 2.5-4 (-9) x (1-) 1.5-2.5 (-4) mm, ovate-lanceolate, densely tomentose on upper surface and margins, glabrescent on lower surface. Inflorescence generally pauciflorous, (1) 2-3 (9) flowers; peduncles sparsely tomentose, sepals and bracts glabres­cent. Capsules (6.5-) 7.7-11.4 (-12.5) mm, fine, straight, cylindrical, erect on the peduncle. Seeds (0.9-) 1-1.2 (-1.4) mm, testa close, acutely tuberculate.

PHENOLOGY- Flowers in July and August. CHROMOSOME NUMBER- 2n = 36 (FAVARGER, 1969). EcoLOGY - Growing in arid and stony grassland between 1,300 and

2,500m. DISTRIBUTION- Endemic to the Western Alps, Cozie and Graie in Italian

and French territories (fig. 5).

SPECIMINA SELECTA.

ITALY: Piedmont - In Alpibus Viucensibus, D. Catochii, s. die, Bellardi (TO-Herb. Bellardi- Typus); In Alpibus di Ceresole, s. die, Re (TO-HP). -Pascoli dumetosi, Alpi di Salsa eire. Pinerolo, s. die, Rostan (TO-HP). - In alpibus Bussolini, la Gardinera, 1810, Balbis (TO-HP). -In valle di Ala, 1824, Bertero (TO-HP). - Nelle rocce erbose delle Alpi di Maniglia, 9.1861, Rostan (TO-HP).- Broussailles des alpes de Massel, 7.1863, Rostan (PI, TO-HP).­Alpi di Viii, M. Solera, 7.8.1863, Beccari(FI).- Valle dell'Orco allo Scalone di Ceresol, 9.8.1863, s. col!. (FI).- Ex Alpibus Pedemontanis prope Ceresol, 9.8.1863,]. Ball (FI).- Vallone di Lea sopra Forno Alpi Graie (Valle Stura di Lanzo), 8.1870, Vallino (TO-HP).- Alpi Graie, sopra Forno, 8.1870, s. col!. (TO-HP).- Alpi Graie, Valli di Lanzo, Valle d'Ala, Pian della Mussa, 7.8.1881, Ungern-Sternberg (TO-HP).- In dumetis herbidis vallis Germanasca supra Perrero,

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Fig. 3 - Cerastium lineare All., lectotypus designated here ITO, herb. Bellardi).

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INVESTIGATIONS ON THE GENUS CERASTIUM L. (CARYOPHYLLACEAE) 21

Fig. 4- Cerastiumlineare All., iwn. nova from: "pascoli pietrosi Val Germanasca sopra Masse!, S.1880, Rostan" (Fl).

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8.1884, Rostan (FI).- Alpes Cottiennes, Vallees Vaudoises, pascoli boschivi, alpi di Salsa, 7 e 8.1887, Rostan (TO-HP, FI). -Alpes du Val di St. Martin, 8.1888, Rostan (TO-HP). -Paturage boissoneux des Alpes Val Germanasca, 8.1890, s. coll. (TO-HP).- Bussoleno tra la Balmetta e il Balmarot in siti erbosi e boschivi (Val di Susa), 8.8.1898, Ferrari & Vallino (TO-HP). - Colle di Crest tra Ronco Canavese (Val Soana) e Riboldone (Val Locana), 7.1904, [?] (TO-HP).- Salendo al colle del Sabbione dall'Alpe del Balmarotto (Valle di Susa), 27.8.1904, Andreis (TO-HP). -Villar Focchiardo al Piano delle Cavalle a circa 1800 m (Val di Susa), 16.8.1905, Fontana (TO-HP).- Valle di Pianprato, frail colle Bocchetta (M. Marzo e Grange Santanel) 2000-2600 m, 30.8.1906, Vaccari (FI).- Villar Focchiardo rupi presso al Mustione, 25.8.1907, Fontana (PAD).- Campiglia Canavese fra santuario diS. Demo e Col della Dalma, 2200-2900 m, 3.9.1908, Vaccari (FI).- Campiglia Soana salita al Colle di Rancio, 1350-2100 m, 1.9.1908, Vaccari (FI).- Dal colle della Portia a ovest scendendo a Usseglio, 22.6.1909, Fontana (PAD).- Ceresole Reale, Alpe di Broglio, 2200-2400 m, 20.7.1910, Vaccari (FI).- Bussoleno rupi trail piano Tuglie e l'Alpe Orsiera (Val di Susa), 11.9.1911, Fontana (TO-HP, PAD).- Val Soana sopra Pianprato sulle rupi nel vallone Giasset, 20.8.1913, Santi (TO-HP).- Balme: piano della Mussa sopra la palazzina Sigismondi, 21.8.1915, Fe"ari & Santi (TO-HP). - Giaveno, pascoli sopra l'Alpe Rocciavre, 17.8.1923, Fontana (TO-HP).- Fenestrelle, rupi cespugliose fra 1' Alpe Balma e del Fondo, 20.8.1937, Negri & Fontana (TO-HP).- Chargeoir (Pinerolo) vallone Borsetto rupi a destra del torrente salendo all'Alpe Comba del Torno, 1550 m, 10.8.1939, Fontana & Oreglia (TO-HP).- Val Susa, vallone del Gravio, 15.9.1974, Merlo (TO-HP).- Lac Falin (V. di Viti) 1691 m, pendii sassosi (serpentiniti), 6.8.1992, Miserere (TO-HP).

Val d' Aosta- Val de Champorcher, Comba della Legna, tra Trome e Mt. Digny, 1300-2000 m, 16.8.1902, Vaccari (TO-HP).- Val de Champorcher, in umbrosis inter Trome e Mont Digny, 1600-2100 m, 24.8.1904, Vaccari (TO-HP, FI, BM).- Champorcher, Bois de Panola, 2000 m, calcescisto, 19.8.1904, Vaccari (TO-HP).- Champorcher fra Dondena e Bois de Boile, 1900-2100 m, 19.8.1904, Vaccari (FI).- Secus viam in silva Panosa inf. 1700 m, 28.7.1905, Levier (FI). -Col di Lavizza, versante di Dondena, 2400 m, 31.8.1906, Vaccari (FI).

FRANCE: Bonneval, s. die, s. colt. (TO-HG).- In alpibus Bonneval in Sabaudia, s. die, Balbis (FI-W).

OBSERVATIONS

Cerastium lineare, now considered an autonomous species (MoscHL, 1982, GREUTER et al., 1984,}ALAS, 1993 ), was in the past referred to as C. strictum L., C. arvense L., C. boissieri Grenier, C. repens L., C. tomentosum L. at infraspe­cific level (cfr. NEGRI, 1904). Most of authors have included C. lineare in the C. tomentosum L. group (NEGRI, 1904, BusCHMANN, 1938, HEss et al., 1967, FAVARGER, 1969). According to MoscHL (1982), C. lineare is similar to C. alpinum L. subsp. lanatum (Lam.) Graebn. but with dimorphous leaves. The presence in C. lineare of long, soft and twisted hairs, tending towars a feldike mat, is not sufficient in itself to affirm systematic affinity between this species and C. tomentosum L., from which it differs on account of the close testa seeds and the lack of lateral shoots. On the other hand, the straight, and erect capsules, the larger seeds with higher and almost spinescent tubercules and the absence of lateral shoots, also set C. lineare apart from C. arvense s. 1.

C. lineare shares systematically important aspects e.g. the densely villous peduncles and rosettes, the lack of axillary shoots, the cylindrical capsules, the

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Fig. 5- Distribution of Cerastium lineare All. in Piedmont, Valle d'Aosta and France, determined from the examined exsiccata and drawn on the Map of the Floristic, Ecological and Vegetation Sectors proposed by MoNTACCHINI & FoRNERIS (1980). The marked sectors correspond to: Valli Chisone and Germanasca (13); Valle di Susa (14); Valli di Lanzo (15); Valli dell'Orco and Val Soana (16); Valle d'Aosta (17); Val Sangone (23 ). The symbol(*) refers to the locus classicus.

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close testa seeds, with the taxa belonging to C. alpinum L. group. Even though the shape of the flowering stem leaves which are linear and glabrescent rather than ovate and generally pubescent, are distinguishing features, we retain that the most pertinent collocation is under the C. alpinum group.

FAVARGER (1969) considers C. lineare as primitive on account of its low ploidy level (diploid, 2n = 36) and the numerous archesporia! cells in the ovules (FAVARGER, 1956). The diploid entities so far ascertained as belonging to the C. alpinum group are C. theophrasti Merxm. & Strid (MERXMULLER & STRID, 1977; STRID, 1997), endemic to the summit of Mount Olympus in Greece and C. villosum Baumg. (BoscAIU & EHRENDORFER, 1996), recently found in the Carpathian Mountains and Eastern Alps. The seeds of C. lineare are very similar to those of C. theophrasti both in size and in the aspect of the acutely tuberculate testa, but the shape of the flowering stem leaves is different.

AcKNOWLEDGEMENTs

The authors wish to thank Prof. F. Garbari (Pisa) and Prof. G. Moggi (Florence) for revising the text and Dr. U. Mossetti (Bologna) for his kind assistance.

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Riassunto

n nome Cerastium lineare All. fu pubblicato da Allioni nel1785. Un Campione appartenente all'erbario di Bellardi conservato a TOe stato designata come suo lectotipo. ll locus classicus di questa specie e individuato nelle Alpi di Viu (Valli di Lanzo). C. lineare, entita diploide di antica differenziazione ed endemica del settore occidentale delle Alpi, presenta aspetti morfologici comuni al gruppo artico-alpino di C. alpinum L. nell' ambito del quale, oltre ai citotipi poliploidi piu ampiamente diffusi, sono presenti anche entita diploidi dalla distribuzione localizzata. Vengono fornite una descrizione dettagliata, una nuova iconografia e informazioni sull'ecologia e la distribuzione di questa specie.

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