Constancea 83, 2002
University and Jepson Herbaria
P.C. Silva Festschrift

Notes on Lectotypification of Usnea fulvoreagens, U. gigantea, U. sulcata, and U. cavernosa

Isabelle I. Tavares
University Herbarium,
University of California, Berkeley,
California 94720-2465

From time to time, for the past thirty years, I have ventured to discuss nomenclatural problems with my next-door neighbor in the University Herbarium, Paul Silva. This could be done in the brief moments when one could find him relatively unoccupied and not discussing nomenclatural problems with botanists in various fields in person, over the phone, or by e-mail.

Among these questions have been the following examples concerning holotypes, lectotypification, and homonyms (see International Code of Botanical Nomenclature, St. Louis Code [Greuter et al., 2001], Art. 9, and the recommendation 9A.2 that all aspects of the protologue should be considered when a lectotype is designated; Art. 53.1 concerning the illegitimacy of later homonyms). The St. Louis Code, Art. 9.2, states that a lectotype is to be selected if the holotype is not indicated in the protologue or if it is missing. Only specimens that are part of the original material used for the description of a new taxon are candidates for lectotypification. The lectotype must be chosen from among isotypes if they exist, or from syntypes or paratypes (Art. 9.10, which states that if no cited specimens exist, the lectotype must be chosen from among illustrations and uncited specimens which comprise the remaining original material). Art. 9.3 defines isotype as any duplicate of the holotype. A specimen having significant disagreement with the protologue ought not to be considered for lectotypification even though it was part of the same mixed collection (see Art. 9.17, which indicates that the choice may be superseded if it is in serious conflict with the protologue and another element is available that is not in conflict).

Usnea gigantea Vain. ex Räsänen (Räsänen, 1932, p. 7).

Usnea gigantea is a name that has not been encountered in recent taxonomic literature, except for its inclusion in the “Checklist of Chilean lichen-forming and lichenicolous fungi” (Galloway and Quilhot, 1998). It was not listed by Motyka (1936, 1937, 1938). Application of the name is uncertain, because of differing opinions as to lectotypification.

Protologue: “Usnea gigantea Vain., Herb. Vainio in Turku, No. 84, 467. Thallus sublaxus, inflatus, laevis, parce fibrillosus et ramosus; soredia et verruculae destituta. Rami crassiores articulati. Medulla laxa, K -. Ster. Prov. de Chiloë: Peninsula Tres Montes, Puerto Barroso, ab cortice N. nitidae suspendens, fq., ut videtur.” [Thallus somewhat flaccid, inflated, smooth, sparingly fibrillose and branched; lacking soredia and verruculae. Thicker branches jointed. Medulla loose, K-. Thallus sterile; on Nothofagus nitida.]

In “Index Nominum Lichenum” (Lamb, 1963), Lamb indicated that the locality of the type was Chile (repeated in Galloway and Quilhot, 1998). My first questioning of Paul resulted in the comment that Lamb's statement seemed to be true at first glance. However, upon investigation, the situation becomes less clear-cut. Räsänen listed two specimens from Vainio's Herbarium at Turku: no. 84 and no. 467. In addition to these, the specimen seen by Räsänen from Chiloé (possibly at Helsinki) would be regarded as a syntype. This specimen was collected by H. Roivainen, April 4, 1929 from bark, Puerto Barroso, Peninsula Tres Montes, Chiloé, Chile (Räsänen, 1932, p. IV).

Of the three syntypes, no. 84 is an Umbilicaria, unrelated to Usnea (Dr. Soili Stenroos, personal correspondence); no. 467 (near Bogota, Colombia, leg. Fr. Emilion, 1898) is an inflated Usnea, broken off at the base, which differs from the diagnosis by the presence of indistinct verruculae and sparse, small fibrils bearing inconspicuous soralia; the Chiloé specimen I have not seen.

If a strict view were to be taken of Räsänen's statement that no soralia or verruculae are present, the explicitly cited specimen of Usnea (no. 467) would seem to be ruled out as a possible lectotype. If specimens can be found that were available to Räsänen from the cited locality in Chiloé, then one of these would be a logical lectotype, if it is concordant with the protologue. If such specimens cannot be found, then a neotype must be designated. My assumption is that an appropriate lectotype or neotype should have a strong resemblance to the Vainio herbarium specimen 467 (perhaps resembling U. fuegiana Motyka, which also is slightly verruculose).

Although Räsänen's diagnoses ordinarily provide little detail, when verruculae or papillae were present, he usually mentioned them (for example in U. comosa f. fuegica Räsänen [Räsänen, 1932, p. 9]). Generally speaking, his descriptions were brief and imprecise, and measurements of anatomical structures were lacking. Consequently, I do not regard no. 467 as seriously in conflict with the protologue; its verruculae and soralia are inconspicuous and the statement “soredia et verruculae destituta” could be interpreted as “not visible under the magnification used when description was written.”

The list of taxa from southern Chile (Räsänen, 1932) in which Räsänen described Usnea gigantea included a large number of species, many previously described by various authors. Because Räsänen provided Latin descriptions for many of the latter, it is not possible to use the presence of Latin to infer that he was proposing a new species. Furthermore, he listed Chilean localities for the specimens he examined. Räsänen's attribution of Usnea gigantea to Vainio suggests that his intent may have been to validly publish this herbarium name, basing it on the Colombian specimen in Vainio's Herbarium (no. 467), and adding the Chilean locality record, representing material not studied by Vainio (date of death—May 14, 1929; see Ann. Crypt. Exot. 3: 5. 1930). Nevertheless, the Chiloé specimen is also a possible lectotype.

Usnea glabrescens (Nyl. ex Vain.) Vain. var. fulvoreagens Räsänen
Die Flechten Estlands, Suomalaisen Tiedeakatemian Toimituksia ser. A, 34: 20. 1931.

Räsänen's diagnosis: “U. glabrescens var. fulvoreagens Räs. (nova).— medulla crebra, K + lutescens, deinde fulvescens.–Vi. Vägeva (Kar.). Sa. Kuresaare an Lärche und Kihelkonna an Picea excelsa (Räs.). Steril. Sehr selten.” (i.e., medulla dense, becoming yellowish with KOH, then becoming yellowish-brown; thallus sterile, uncommon.)

Three localities in Estonia were given: 1) Vi. Vägeva (Kar.); 2) Sa. Kuresaare an Lärche (Larix); 3) Sa. Kihelkonna an Picea excelsa (Räs.). (Sa. referred to Saaremaa, the island; Vi referred to Virumaa near the coast of the Gulf of Finland; see Halonen and Ahti, 2002). An apparent syntype has been found at TUR (Vägeva, 3 July 1924, Kari s.n., TUR) and this specimen, identified as U. subfloridana Stirt. (a taxon having isidiate soralia–cf. Räsänen, 1939), has been designated as the lectotype by Halonen and Ahti (2002).

In 1935, Räsänen used the new combination Usnea fulvoreagens when distributing thalli collected at Otsanlahti, Kurkijoki, Finland (now Kurkiyoki, Russia, west of Lake Ladoga [Laatokka]) as Lichenes Fenniae Exsiccati a Museo botanico Universitatis Helsinkiensis editi. Fasc. 1 (nr. 1–50) curavit V. Räsänen a. 1935. 13. Usnea fulvoreagens Räs. syn. U. glabrescens v. fulvoreagens Räs., Die Flechten Estlands I (1931), p. 20. Karelia ladogensis: Kurkijoki, Otsanlahti. Ad corticem Alni incanae. 7.XII.1932. leg. Veli Räsänen. This label was printed in the Schaedae ad Fasciculos I–III (Nr. 1–150), quos curavit Veli Räsänen, Helsinki, 1935, 50 p. Räsänen cited the variety fulvoreagens as the basionym.

Räsänen (1939) augmented the descriptive information of the protologue (Räsänen, 1931) under the name Usnea fulvoreagens Räsänen in a survey of lichens in the area bordering Lake Laatokka. Among the localities given were Lapinlahti and Otsanlahti. Räsänen described the species as scarcely branched and fibrillose, with the lateral branchlets bearing non-isidiate soralia. The medulla was dense to sublax, at the base ca. KOH-, toward the apex KOH+ fulvescent. It is not clear whether this description applied more closely to those specimens near Lake Laatokka than to those from Estonia, but he reported a similar chemical reaction for both.

Motyka (1936) also proposed the new combination Usnea fulvoreagens, but one year after Räsänen. He cited Räsänen as author of the basionym, but he incorrectly cited as type a specimen not mentioned by Räsänen, namely a specimen from Kurkijoki, Lapinlahti, Finland.

Clerc (1987), cited as “type (?)” [query indicating uncertainty on the part of Clerc] of U. fulvoreagens a specimen at Helsinki from Kl. Kurkijoki Lapinlahti and indicated that it has norstictic, stictic, and other acids. This specimen is not an acceptable lectotype, since it was not part of the original material. Ohmura and Kashiwadani (2000 p. 170) followed Clerc in listing (but without questioning its status) the specimen from Lapinlahti as the type of U. fulvoreagens (Räsänen) Räsänen. Halonen and Ahti (2002) have now proposed to conserve the name U. fulvoreagens with this specimen as the conserved type.

My photograph of what I assumed to be the type specimen of U. fulvoreagens in 1975 at Helsinki (i.e., thalli from Lapinlahti) shows 3 specimens mounted together—two with elongate branch apices (one having norstictic acid was marked “1.” possibly by Räsänen) and one having short apices, with crowded fibrils, the chemistry of which is not known to me. If these specimens represent the proposed conserved type, then the authors of the proposal should have indicated which of the specimens best represents their concept of U. fulvoreagens.

Motyka's description was of a taxon having a narrow, loose medulla with a KOH yellow, then subrubescent reaction, minute, briefly cylindrical papillae, dense fibrils, and indistinct branch apices that are densely soraliate. This description fits more closely the specimen having short apices and crowded fibrils mentioned above.

Usnea cavernosa Tuckerman
in Agassiz, Enumeratio Lichenum a D. Prof. Agassiz ad Lacum Superiorem, anno 1848.

Tuckerman's original description (in Agassiz, 1850) gave two Lake Superior syntypes—one collected by C.T. Jackson, 1845, and one by Agassiz, 1848. In addition he cited one from the White Mts. [New Hampshire], 1843 (collected by Tuckerman, according to Eileen Wozek, Farlow Herbarium—personal communication), and also mentioned a syntype from Madras, India (obviously another taxon).

Herrera-Campos et al. (Bryologist 101: 315. 1998) cited as type “America Septentr. ad oras Lacus superioris, Tuckerman (FH!, holotype)”.

In order for lectotypification to be effected properly, it will be necessary to specify which of the original specimens is referred to, because neither the 1845 nor the 1848 specimen was collected by Tuckerman, as implied by the citation.

Usnea sulcata Motyka, 1937, P. 478.

“Typus in Museo. Botan. Univ. Fennicae in Turku.—Locus classicus: Brasilia, Minas Geraës, Sitio, 1885 Vainio. Exsiccata: Vainio, Brasil. 388, 389, 922, 953, 1387 (Tu.)—Wright, L. Cubae 48 (U.. Y.).—Zahlbr., Lich. rar. 36 (V.)....Apothecia vulgo praesentia...sat parva, 0.5–0.8 cm. lata,...” [apothecia commonly present...moderately small, 0.5–0.8 cm. wide,...]. Motyka cited collections from Cuba, Jamaica, and Mexico to Brazil and Argentina, as well as South Africa. In his discussion, Motyka (p. 479) stated: “Magis affinis est U. paradoxa colore fusco, medulla K rubescente; est tamen multo crassior, sorediosa” [more completely allied is U. paradoxa, dark brown in color, medulla KOH reddish; it is nevertheless much thicker and sorediose].

Awasthi (1986) lectotypified U. sulcata on 00450 TUR-VAIN. She cited U. sulcata as a synonym of U. angulata, describing U. sulcata as similar to U. angulata, but more thickened, and stating “but other characters are identical”. She also examined specimen no. 453; i.e., Exs. no. 922. Herrera-Campos et al. (1998) followed Awasthi's lectotypification and they also considered U. sulcata conspecific with U. angulata.

It is pertinent to point out, however, that Usnea angulata Ach. is a soraliate species, whereas U. sulcata was described by Motyka (1937) as apotheciate, with no soralia or isidia mentioned in the protologue. Although he did not specifically state that U. sulcata is esorediate, he contrasted it with the soraliate species U. paradoxa (see above). Herrera-Campos et al. did not refer to apothecia in their description of U. angulata, and they later mentioned the apotheciate species, U. alata Motyka, as the primary species of U. angulata (p. 327), i.e., the corresponding apotheciate member of a species pair. No. 00450 (Vainio exs. 388) appears to have some small, poorly developed apothecia, and is also sorediate-isidiate, whereas no apothecia are visible on Vainio exs. nos. 389 (451), 922 (453), 953 (452) (Dr. Soili Stenroos, personal communication). Motyka listed the type as from Sitio, 1885; he gave no other information (see also Alava, 1986, p. 156). It should be noted that Vainio (1890, pt. 1, p. 11), under U. angulata, stated that Sitio specimens were sterile. This statement might have been used to refer either to pycnidial specimens on which apothecia had not yet developed, or to specimens producing isidia or soralia.


I wish to thank the directors and curatorial staffs of the following herbaria: H, TUR, and FH for their assistance during visits in the 1970's and for information sent by correspondence. Especially I want to express my appreciation for the invaluable advice and counsel given me by Paul Silva, and for the use of his library. I very much appreciate the editorial advice and assistance of Richard L. Moe.


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