On the status of Umbilicaria aprina var. halei and U. canescens (Umbilicariaceae, lichenized Ascomycota)

Abstract

To clarify the nomenclature of Umbilicaria aprina var. halei and U. canescens, type material from LE and MIN was investigated. Description, measurements and photos of the thalloconidia of the type specimens are provided. According to our investigation Umbilicaria aprina var. halei is a synonym of U. rhizinata, and U. canescens is a synonym of U. aprina. Both species belong to the U. aprina group and are distinguished by the septation of their thalloconidia. Additional specimens identified by previous authors as Umbilicaria aprina var. halei and U. canescens were re-identified. Umbilicaria rhizinata is reported as new to Xinjiang, China.

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Phytotaxa 533 (1): 091–097
https://www.mapress.com/pt/
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Frank Bungartz: 22 Jan. 2022; published: 4 Feb. 2022
https://doi.org/10.11646/phytotaxa.533.1.6
91
On the status of Umbilicaria aprina var. halei and U. canescens (Umbilicariaceae,
lichenized Ascomycota)
EVGENY A. DAVYDOV
Altai State University, Lenin Ave. 61, 656049 Barnaul, Russia; Central Siberian Botanical Garden, Siberian Branch of the Russian
Academy of Sciences, Zolotodolinskaya, 101, Novosibirsk, 630090, Russia
�
eadavydov@yandex.ru; https://orcid.org/0000-0002-2316-8506
Abstract
To clarify the nomenclature of Umbilicaria aprina var. halei and U. canescens, type material from LE and MIN was inves-
tigated. Description, measurements and photos of the thalloconidia of the type specimens are provided. According to our
investigation Umbilicaria aprina var. halei is a synonym of U. rhizinata, and U. canescens is a synonym of U. aprina. Both
species belong to the U. aprina group and are distinguished by the septation of their thalloconidia. Additional specimens
identified by previous authors as Umbilicaria aprina var. halei and U. canescens were re-identified. Umbilicaria rhizinata
is reported as new to Xinjiang, China.
Keywords: Altai, China, Mongolia, thalloconidia, Umbilicaria rhizinata
Introduction
Species of Umbilicariaceae Chevall. are predominantly saxicolous lichens mostly found in regions of higher latitudes
or at higher elevations worldwide. Multilocus phylogenies resulted in a new generic concept of the family, which
currently includes three genera comprising together about one hundred species of mostly umbilicate growth habit
(Bendiksby & Timdal 2013, Davydov et al. 2017). The Umbilicaria aprina group is a monophyletic group of
morphologically similar species within Umbilicaria Hoffm. subg. Umbilicaria (Davydov et al. 2017). Species of
the Umbilicaria aprina group have a slightly areolate or reticulate centre of the grey upper thallus surface, scarce to
dense, simple or branched rhizinomorphs, non-septate to 10-septate thalloconidia, and omphalodisc apothecia with
asci containing hyaline, unicellular ascospores.
Thalloconidia have been shown to be highly species-specific and are the most important diagnostic trait in the
Umbilicaria aprina group (Hasenhüttl & Poelt 1978, Krog & Swinscow1986, Hestmark 1990, 1991). Hestmark (1991:
356), discussing the phylogenetic relationship between U. aprina Nyl. and U. africana (Jatta) Krog & Swinscow,
mentioned that “data on conidium septation perhaps…suggest the existence of intermediate forms (taxa)” between U.
aprina with unicellular thalloconidia and U. africana possessing 4–10 celled ones. Regarding thalloconidia septation,
U. rhizinata (Frey & Poelt) Krzewicka (Krzewicka 2010) represents such an intermediate species first described as
U. decussata (Vill.) Zahlbr. var. rhizinata Frey & Poelt (Poelt 1977). According to Krzewicka (2010), U. rhizinata
is mainly distinguished from the morphologically similar U. decussata, U. krascheninnikovii (Savicz) Zahlbr. and
U. formosana Frey by its 3- or 4-celled thalloconidia produced on the lower surface in patches, whereas the three
latter species lack thalloconidia or possess non-septate ones. This character was also constant for specimens with
similar morphology from Asia, Africa and South America (Krzewicka 2010). The species status of U. rhizinata and its
inclusion in U. aprina group was supported by molecular phylogenetic analyses (Davydov et al. 2017).
Thalloconidia of Umbilicaria aprina Nyl. var. halei Llano and Gyrophora canescens Dombr. both related to the
U. aprina group have not been investigated so far and the status of such taxa therefore remains unclear.
Umbilicaria aprina var. halei was described by Llano (1956) from Baffin Island (North American Arctic). The
main characters distinguishing U. aprina var. halei from the type variety were its diminutive size, slate to grey-black
colour of the upper surface, and scarce, small, dichotomous rhizinomorphs. Ryvarden (1968) noted that specimens

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from Baffin Island are more or less similar with rather variable Scandinavian material of U. aprina. Since that time, U.
aprina var. halei has been rarely mentioned in the literature (e.g., Goward et al. 1994). However, Wei & Jiang (1993)
accepted this variety in their monograph of Asian Umbilicariaceae, and suggested morphological and geographical
differences from U. aprina var. aprina. They were not able to examine the type material, and recognized U. aprina var.
halei by its blackish-grey upper surface, sometimes bearing apothecia, with a distribution in North-East Asia, North
Europe and North America. Moreover, they reduced U. decussata var. rhizinata to synonymy with U. aprina var.
halei.
Gyrophora canescens was described from the Kola Peninsula (Dombrovskaya 1970a) and transferred to
Umbilicaria by Golubkova (Golubkova & Savicz 1978). All the above authors treated the species as related to
Umbilicaria cylindrica (L.) Delise, but differing by its dark lower surface. Byazrov (1986) accepted U. canescens
(Dombr.) Golubk. as a species, while Santesson (1993) synonymized U. canescens with U. cylindrica var. delisei Nyl.
Wei & Jiang (1993) placed U. canescens (Dombr.) Golubk. to the “excluded species” section and did not propose
synonymy. Latter J. C. Wei referred the type material of Gyrophora canescens to U. aprina var. halei (LE–L266, L267,
L268, tested in 1995). Davydov & Zhurbenko (2008) supported the opinion that Gyrophora canescens is conspecific
with Umbilicaria aprina, considering superfluous the segregation of var. halei. Clerc (2004), Nimis (2016), Nimis et
al. (2018) referred U. canescens to U. cylindrica s. lat., and Urbanavichus et al. (2008) and Roux et coll. (2020) – to
U. cylindrica var. delisei.
Thus, the specimens recognised by different authors as Umbilicaria aprina var. halei seem to represent at least
two morphologically similar species, namely U. aprina and U. rhizinata, and Umbilicaria canescens treated by authors
as U. aprina var. aprina, U. aprina var. halei, U. cylindrica var. delisei or U. cylindrica s.lat. The goal of the present
paper is to clarify the taxonomic status of U. aprina var halei and U. canescens.
Material and methods
The holotype of Gyrophora canescens was studied in the herbarium of the Komarov Botanical Institute, Saint Petersburg
(LE), the lectotype of Umbilicaria decussata var. rhizinata – in Staatliche Naturwissenschaftliche Sammlungen
Bayerns (M), the holotype of U. aprina and the holotype of U. aprina var. halei were received on loan from the
University of Helsinki (H) and University of Minnesota (MIN), respectively. Additional specimens were collected
in the Altai Mountains as well as studied in herbaria ALTB, KPABG, LE, and M. Specimens were examined using a
stereomicroscope (Zeiss Stemi 2000-C) and a compound microscope (Zeiss Axio Lab.A1). Anatomical examination
was undertaken using hand-cut sections mounted in water. Thalloconidia were brushed off from the lower surface of
thalli, mounted in water, then examined and photographed in the light microscope Zeiss Axio Imager A1; measurements
were performed on the photographs using Zeiss software. Due to significant differences in size, thalloconidia with
different septation were measured separately. Measurements of thalloconidia are presented as follows: (smallest value
recorded) (X-SE)–X–(X+SE) (largest value recorded), where X is the (arithmetic) sample mean, and SE is the sample
error of mean.
Taxonomy
Umbilicaria rhizinata (Frey & Poelt) Krzewicka (2010: 491) Mycobank no. 548351
Umbilicaria decussata (Vill.) Zahlbr. var. rhizinata Frey & Poelt (Poelt 1977: 419).
Type:—NEPAL. Mahalangur: Himal Khumbu, Moränen des Lobuche-Gletschers bei Lobuche, elev. 4950–5000 m, September 1962, J.
Poelt L247a (lectotype M–0035585!; isolectotype M–0035586!).
Umbilicaria aprina Nyl. var. halei Llano (1956: 183), syn. nov. Type:—CANADA. N.W.T.: Baffin Island, head of Clyde Fiord, on exposed
gneiss boulders, 26 August 1950, M. E. Hale 450 (holotype MIN–664956!, Fig. 1).
The holotype specimen of U. aprina var. halei has a similar morphology to the Himalayan U. rhizinata: small size,
dark grey granulate upper surface, lower surface covered with black patches of thalloconidia in the central part only,
and scarce dichotomous pale rhizinomorphs (Fig. 1). Thalloconidia are sphaerical to ellipsoid, brown to dark brown
3- or 4-celled when mature, but often also 2- or rarely 1-celled (Fig. 2). The sizes were (12.6–)15.0–16.2–17.5(–18.7) ×

UMBILICARIACEAE, LICHENIZED ASCOMYCOTA Phytotaxa 533 (1) © 2022 Magnolia Press • 93
(10.7–)12.6–13.7–14.9(–16.5) μm for (3-)4-celled (n=25), (11.6–)12.8–13.7–14.9(–16.5) × (8.0–)8.7–9.7–10.7(–11.7)
μm for 2-celled (n=25), and (8.1–)9.3–10.3–11.4(–12.0) × (7.6–)8.5–9.5–10.6(–11.4) μm for 1-celled thalloconidia
(n=25). The wall was 1.3–1.8(2.8) μm. The average size for all types of thalloconidia is 10.9–13.6–16.4 × 8.8–11.0–
13.2 μm which agrees well with the data presented by Krzewicka (2010) for U. rhizinata. All the remaining characters
of U. aprina var. halei also perfectly fit the circumscription of U. rhizinata, except, none of the three specimens of the
U. aprina var. halei holotype has a distinct reticulate pattern at the centre. They are coarsely pruinose with crystals
positioned centrally near the umbo, but lack a reticulate pattern. Additionally, examined specimens from the Altai
Mts. are only occasionally reticulate at the centre. This can be explained by the smaller size of both the holotype and
the Altaian specimens (0.8–1.3 cm v. 1–2(–3) cm of U. rhizinata). As has been shown in phylogenies (Davydov et
al. 2017), U. rhizinata is closely related to U. aprina, and only distantly related to U. decussata, which consistently
possesses this reticulate pattern. Therefore, this character should be used with care as a diagnosis for U. rhizinata.
Thus, it is proposed that U. aprina var. halei be reduced to synonymy with U. rhizinata.
Umbilicaria rhizinata resembles immature U. aprina, but differs in its 3–4 celled thalloconidia. Due to the
different conception of U. aprina var. halei by Wei & Jiang (1993) who did not pay attention to thalloconidium
septation, material cited by them should be re-identified. Based on the information on the apothecia of U. aprina var.
halei given in Wei & Jiang (1993) their material probably belongs to U. aprina. Umbilicaria rhizinata was previously
known from Nepal and Stanovoye Nagor’e Highlands (South Siberia, Russia) in Asia, Tanzania in Africa, and Bolivia
in South America (Poelt 1977, Krzewicka 2010, Davydov et al. 2019a). Further populations in the Altai Mts. both in
Russia and China are recorded here. So, we have at least one verified locality for China – in Xinjiang.
Selected specimens examined. RUSSIA. Altai Territory: Altai Mts., Korgonsky range, headwaters of Sentelek
River, 51°03’ N, 83°43’ E, elev. 1900–2200 m, mountain tundra, 19 August 1996. E. A. Davydov 5350 (ALTB);
Republic of Altai: Katunsky range, Ak-Kem River, stone fields and rocks, 49°59’15.6’’ N, 86°35’09.3’’ E, elev. 2686
m, 9 August 2009. E. A. Davydov 6879 (ALTB).—CHINA. Xinjiang: South Altai range, 49°01’41’’ N, 86°50’38’’ E,
elev. 2300–2600 m, 17 July 2007. Davydov 6542 (ALTB).
FIGURE 1. Umbilicaria aprina var. halei (MIN-664956—lectotype). Scale=1 cm

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FIGURE 2. Umbilicaria aprina var. halei (MIN-664956—lectotype). Individual 3-, 4-celullar thallocondia (light microscopy).
Scale=10μm
Umbilicaria aprina Nyl. (Nylander 1869: 12) Mycobank no. 408077
Type:—ETHIOPIA (=Abyssinia). Dedschen Mts, elev. 14200 ft. W. Schimper (holotype H–NYL 31742!).
Gyrophora canescens Dombr. (Dombrovskaya: 1970a: 131), syn. nov. Type:—RUSSIA. [Murmansk Region]: Peninsula Kolaёnsis,
montes Chibinensis, ad declivitatem australem montis Vudjavrczorr, elev. 380 m, in schistosis ad lapides nudis, No 6, 5 June 1965,
A. V. Dombrovskaya (holotype LE-L266!, Fig. 3)
Umbilicaria canescens (Dombr.) Dombr. (Dombrovskaya: 1970b: 131), comb. inval., ICN Art. 41.1 (Turland et al. 2018).
Umbilicaria canescens (Dombr.) N. S. Golubk. (Golubkova & Savicz 1978: 115).
Thalloconidia produced on most of lower surface of the holotype specimen of Gyrophora canescens, spherical to
ellipsoid, brown to dark brown, mainly unicellular, rarely 2-cellular (Fig. 4). The sizes were (7.3–)7.9–8.7–9.6(–9.9) ×
(6.2–)7.0–7.6–8.2(–8.7) μm for 1-celled (n=25), and (10.6–)10.6–11.2–11.8(–12.3) × (7.2–)7.8–8.4–9.0(–9.0) μm for
2-celled (n=10). The wall was 1.2–1.8 μm. The size and septation of thalloconidia correspond to those known for U.
aprina (Hestmark 1990). The morphology of the holotype (Fig. 3) and paratype specimens of Gyrophora canescens
also agree with the circumscription of U. aprina except for the dark grey colour of the upper surface. However, this is
not a reliable distinguishing character because it may reflect ecological variances and varies even among specimens of
the same population. Thus, it is proposed to reduce U. canescens (Dombr.) Golubk. into synonymy with U. aprina.
Aside from the locus classicus in the Kola Peninsula Umbilicaria canescens was reported for the Kamchatka
Peninsula (Dombrovskaya 1970a), and Mongolia (Byazrov 1986). All the specimens cited by Dombrovskaya (1970a)
and Byazrov (1986) were examined and refer to U. aprina. Umbilicaria aprina Nyl. is distributed in the high Arctic, the
Antarctic and high mountain habitats worldwide (Davydov et al. 2011, 2019b, Hestmark 2015, 2016). References to U.
aprina in the Russian Altai (Davydov & Zhurbenko 2008) actually refer to U. rhizinata. According to its distribution
pattern I suspect U. aprina may be present in Altai but I haven’t seen any verified specimens yet.
Selected specimens examined. Exiccata: Feige & Lumbsch, Umbilicariaceae Exs. no. 27, 67; Lichenotheca
Afganica no. 16, 25, 58.—RUSSIA. Murmansk Region: Monchegorsk District, Nyavka-Tundra Range, at the middle
section of Liva River, 9 km upstream from Livozero Lake, elev. 140 m, spruce forest, on stone, 21 July 1973. A.
V. Dombrovskaya (KPABG–2707); Kirovsk District, Khibiny Mts, the slope of Vuonnemyok River valley near the

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Yuksporlak pass, mountain tundra, on rock. 21 July 1974. A. V. Dombrovskaya (KPABG–4434); Novosibirskie Is.,
Bennett Island, c. 76°40’N 149°00’E, 130 m, 23 July 1989. M. P. Zhurbenko 8990 (LE–L6191); Republic of Buryatia:
Khamar-Daban range, Baikalsky reserve, elev. 1600–1700 m, 26 July 1993. G. P. Urbanavichus (ALTB–L178);
Kamchatka Region: Elizovsky District, Khalaktyrka River, vicinity of Petropavlovsk-Kamchatsky, 12 July 1980, I.
M. Antonova (KPABG–4410).—MONGOLIA. Kosogol Lake, Mundu-Sardyka Mt. 23 July 1902. A. A. Elenkin (LE–
L5286); Uvs aimag, Han-Hu-Hei Range, Dulga-Ula Mt., elev. 2928 m, 9 July 1976. L. G. Byazrov 4584 (LE–L5287);
Zavkhan aimag: Somon Tudevtei, elev. 2000 m., 11 July 1976. L. G. Byazrov 4585 (LE–L5288).—TAJIKISTAN.
Turkistansky range, elev. 3000–3400 m, 15 July 1986. I. Kudratov (LE–L6239).
FIGURE 3. Umbilicaria canescens (LE-L266—holotype). Upper and lower surfaces. Scale=1 cm

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FIGURE 4. Umbilicaria canescens (LE-L266—holotype). Non-septate and one-septate thallocondia (light microscopy). Scale=10μm
Acknowledgements
We gratefully acknowledge help from the curators and staff of cited herbaria and to Professor Mark Seaward and two
anonymous reviewers for improving the text. The reported study was funded by Russian Science Foundation according
to the research project № 22-24-00283.
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