![World distribution of Cetreliopsis species. A. C. asahinae. B. C. endoxanthoides. C. C. hypotrachyna. D. C. laeteflava. E. C. papuae (•) and C. thailandica (■). F. C. rhytidocarpa [C. r. ssp. rhytidocarpa (o) and C. r. ssp. langtangi (•)].](https://www.researchgate.net/profile/Tiina-Randlane/publication/267978660/figure/fig2/AS:669404002021391@1536609759458/World-distribution-of-Cetreliopsis-species-A-C-asahinae-B-C-endoxanthoides-C-C_Q320.jpg)
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Distribution patterns of some cetrarioid species 359
Symb. Bot. Ups. 34:1
Distribution patterns of some primary and secondary
cetrarioid species
Tiina Randlane and Andres Saag
Randlane, T. & Saag, A. 2004. Distribution patterns of some primary and secondary cetrari-
oid species. – Acta Univ. Ups. Symb. Bot. Ups. 34:1, 359–376. Uppsala. ISBN 91-554-6025-9.
World-wide distribution maps for 39 cetrarioid lichen species from four genera, Allocetraria,
Cetrelia, Cetreliopsis and Tuckneraria, are presented. The taxa chosen represent both primary
and secondary species in the same genus, which makes it possible to compare the global
distribution patterns of closely allied taxa with different strategies of reproduction. The centers
of diversity are suggested for each genus on the basis of distribution maps. Presumed pairs of
primary and secondary species are presented for Allocetraria, Cetrelia and Cetreliopsis.The
majority of primary species (22 taxa) have restricted distribution areas in the region where the
diversity and speciation centers of these genera are most probably situated (eastern and south-
eastern Asia). Still, this is not the absolute rule as two primary Allocetraria species (A. ma-
dreporiformis, A. stracheyi) have spread from their distribution center (Asia) to North Amer-
ica. Of the 15 secondary species treated here, six sorediate lichens (Allocetraria oakesiana,
Cetrelia cetrarioides, C. chicitae, C. monachorum, C. olivetorum and Tuckneraria laureri)
occur extensively on two, three or even four continents, although the diversity centers of all
these genera lie in rather restricted areas in Asia. Only one sorediate taxon (Cetreliopsis laete-
flava) has a limited distribution area close to the diversity center of the genus Cetreliopsis.
Isidiate species are less successful in their distribution. All four discussed species (Alloce-
traria isidiigera, Cetrelia braunsiana, C. isidiata and Tuckneraria togashii) have "stayed" in
Asia. Distribution of lobulate species demonstrates a pattern similar to that of isidiate taxa.
Key words:Allocetraria, Cetrelia, Cetreliopsis, Tuckneraria, distribution maps.
Tiina Randlane and Andres Saag, Institute of Botany and Ecology, University of Tartu, Lai
Street 38, 51005, Tartu, Estonia. E-mail: tiina.randlane@ut.ee
Introduction
General knowledge about the global distribution
of lichens is still poor, although an enormous
amount of data concerning the localities of differ-
ent species is available. Any generalisation of geo-
graphical distribution of taxa can be based only on
their detailed mapping. Very precise systems for
the mapping of lichens have been initiated in some
West-European countries (Austria, Great Britain,
Italy), and now also in northern Europe (Finland,
Sweden, Norway, Denmark, Iceland) since the
project Nordic Lichen Flora was started (Nordic
Lichen Flora vol. 1 1999). The distribution of only
a few lichen species has been mapped on a trans-
European scale recently (Wirth & Oberhollenzer
1990, Trass 1998, Litterski & Otte 2002) while
world-wide distribution maps of species are some-
times presented in systematic treatments of limited
groups.
In this paper we introduce world-wide distribu-
tion maps for 39 cetrarioid lichen species. The taxa
chosen represent both primary and secondary spe-
cies of the same genus, which makes it possible to
compare the global distribution patterns of closely
allied taxa with different strategies of reproduc-
tion. The cetrarioid lichens (Parmeliaceae, Asco-
mycota) include 143 species distributed among 24
genera (Randlane & Saag 2000). In eight of these
both primary and secondary species are repre-
360 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
Table 1. Cetrarioid genera including both “primary” and “secondary” species (species are listed alpha-
betically, phylogenetic affinities between primary and secondary species are not stated here).
Genus Primary species Apothecia
present Secondary species Vegetative
propagules
Allocetraria Kurok. &
M.J.Lai A. ambiqua (Bab.) Kurok. &
M.J.Lai yes A. isidiigera Kurok. & M.J Lai
A. oakesiana (Tuck.) Randlane & isidia
soredia
10 species A. endochrysea (Lynge) Kärnefelt
& A.Thell no A.Thell
A. denticulata (Hue) A.Thell &
Randlane no
A. flavonigrescens A.Thell &
Randlane yes
A. globulans (Nyl.) A.Thell &
Randlane yes
A. madreporiformis (Ach.)
Kärnefelt & A.Thell yes
A. sinensis X.Q.Gao no
A. stracheyi (Bab.) Kurok. &
M.J.Lai yes
Cetrelia W.L.Culb. &
C.F.Culb. C. alaskana W.L.Culb. &
C.F.Culb. no C. braunsiana (Müll.Arg.)
W.L.Culb. & C.F.Culb. isidia
17 species C. collata (Nyl.) W.L.Culb. &
C.F.Culb. yes C. cetrarioides (Duby) W.L.Culb.
& C.F.Culb. soredia
C. davidiana W.L.Culb. &
C.F.Culb. yes C. chicitae (W.L.Culb.)
W.L.Culb. & C.F.Culb. soredia
C. delavayana W.L.Culb. &
C.F.Culb. yes C. isidiata (Asahina) W.L.Culb.
& C.F.Culb. isidia
C. nuda (Hue) W.L.Culb. &
C.F.Culb. yes C. japonica (Zahlbr.) W.L.Culb.
& C.F.Culb. lobules
C. pseudocollata Randlane &
Saag no C. monachorum (Zahlbr.)
W.L.Culb. & C.F.Culb. soredia
C. sanguinea (Schaer.) W.L.Culb.
& C.F.Culb. yes C. olivetorum (Nyl.) W.L.Culb.
& C.F.Culb. soredia
C. orientalis Randlane & Saag lobules
C. pseudolivetorum (Asahina)
W.L.Culb. & C.F.Culb. lobules
C. sinensis W.L.Culb. &
C.F.Culb. lobules
Cetreliopsis M.J. Lai C. asahinae (M.Satô) Randlane &
A.Thell yes C. laeteflava (Zahlbr.) Randlane
& Saag soredia
7 species C. endoxanthoides D.D.Awasthi yes
C. hypotrachyna (Müll.Arg.)
Randlane & Saag yes
C. papuae Randlane & Saag no
C. rhytidocarpa (Mont. & Bosch)
M.J.Lai yes
C. thailandica Elix & M.J.Lai yes
Platismatia W.L.Culb.
& C.F.Culb. P. formosana (Zahlbr.) W.L.Culb.
& C.F.Culb. yes P. erosa W.L.Culb. & C.F.Culb.
P. glauca (L.) W.L.Culb. & isidia
soredia &
10 species P. lacunosa (Ach.) W.L.Culb. &
C.F.Culb. yes C.F.Culb.
P. herrei (Imshaug) W.L.Culb. & isidia
P. stenophylla (Tuck.) W.L.Culb.
& C.F.Culb. yes C.F.Culb.
P. interrupta W.L.Culb. & isidia
soredia &
P. tuckermanii (Oakes) W.L.Culb.
& C.F.Culb. yes C.F.Culb.
P. norvegica (Lynge) W.L.Culb. isidia
& C.F.Culb. isidia
P. regenerans W.L.Culb. &
C.F.Culb. lobules
Distribution patterns of some cetrarioid species 361
Symb. Bot. Ups. 34:1
Tuckermanella Essl. T. arizonica Essl. yes T. coralligera (W.A.Weber) Essl. isidia
T. fendleri (Nyl.) Essl. yes
6 species T. pseudoweberi Essl. yes
T. subfendleri (Essl.) Essl. yes
T. weberi (Essl.) Essl. yes
Tuckermanopsis
Gyeln. T. americana (Spreng.) Hale yes T. chlorophylla (Willd.) Hale
T. gilva (Asahina) M.J.Lai soredia
soredia
T. ciliaris (Ach.) Gyeln. yes T. ulophylloides (Asahina) soredia &
12 species T. inermis (Nyl.) Kärnefelt yes M.J.Lai isidia
T. microphyllica (W.L.Culb. &
C.F.Culb.) M.J.Lai yes
T. orbata (Nyl.) M.J.Lai yes
T. platyphylla (Tuck.) Hale yes
T. platyphylloides (Asahina)
M.J.Lai yes
T. subalpina (Imshaug) Kärnefelt yes
T. weii (X.Q.Gao & L.H.Chen)
Randlane & Saag yes
TucknerariaRandlane
& A.Thell T. ahtii Randlane & Saag
T. laxa (Zahlbr.) Randlane & yes T. laureri (Kremp.) Randlane &
A.Thell soredia
5 species A.Thell
T. pseudocomplicata (Asahina) yes T. togashii (Asahina) Randlane &
A.Thell isidia
Randlane & Saag yes
Vulpicida J.-E.Matts-
son & M.J.Lai V. canadensis (Räsänen) J.-
E.Mattsson & M.J.Lai yes V. pinastri (Scop.) J.-E.Mattsson
& M.J.Lai soredia
6 species V. juniperinus (L.) J.-E.Mattsson
& M.J.Lai yes
V. tilesii (Ach.) J.-E.Mattsson &
M.J.Lai yes
V. tubulosus (Schaer.) J.-E.Matts-
son & M.J.Lai yes
V. viridis (Schwein.) J.-E.Mattsson
& M.J.Lai yes
Table 1. Cetrarioid genera including both “primary” and “secondary” species (species are listed alpha-
betically, phylogenetic affinities between primary and secondary species are not stated here).
Genus Primary species Apothecia
present Secondary species Vegetative
propagules
sented (Table 1). Taxa with uncertain generic posi-
tion are not taken into consideration. Distribution
maps for four mainly Asian genera (Allocetraria,
Cetrelia, Cetreliopsis and Tuckneraria) are pre-
sented here, while the distribution of four further,
mainly North American genera, Platismatia, Tuck-
ermanella, Tuckermanopsis and Vulpicida, will be
analysed in another paper.
The cetrarioid lichens are phylogenetically het-
erogenous (Mattsson & Wedin 1998, Saag et al.
2002, Thell et al. 2002). This also applies to the the
four genera studied here. Allocetraria, Cetrelia,
Cetreliopsis and Tuckneraria do not form a mono-
phyletic group and no phylogenetic affinities be-
tween these genera are suggested. The attention is
mainly paid to the presumed connections within
the genera.
The criterion for a species pair concept was first
and clearly formulated by Du Rietz in 1924; he
used the term "Artenpaaren" to indicate the phe-
nomenon that pairs of lichen species exist, which
differ from each other in just one character – hav-
ing or lacking soredia (or isidia). He also drew
attention to the fact that sorediate and usually ster-
Table 1 (cont.)
362 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
ile taxa have a different ecology and distribution
from their nonsorediate fertile allies (Mattsson &
Lumbsch 1989). Hale (1965) pointed out that sore-
diate species often have a wider distribution than
their fertile counterparts. Poelt (1970, 1972) ex-
tended the species pair concept and declared that
to each sorediate taxon there is a corresponding
nonsorediate counterpart, and he considered sore-
diate taxa evolutionary dead ends. The genus Cet-
relia was shown to fit rather well into such treat-
ment since species in this genus have been inter-
preted as combinations of different morpho- and
chemotypes (Randlane & Saag 1991) and for
every secondary species an existing primary taxon
can be found. However, not only pairs of species
but also triplets or even tetrades of species are ob-
served in Cetrelia. It was also pointed out that all
taxa with sexual reproduction appear to be quite
rare with very restricted distribution.
The taxonomic significance of soredia and
isidia is a problem closely connected with the
ranking of species pairs (or triplets etc.). Different
solutions have been offered (Tehler 1982, Matts-
son & Lumbsch 1989), but the usual practice is
still to treat them as separate species. Today the
situation has become much more complex due to
the accumulation of molecular data. According to
Grube & Kroken (2000) some studies carried out
on a single locus indicate that species pairs are not
real, while other studies based on multilocus se-
quence analyses support the species pair concept,
at least in some genera (e.g. Letharia).
The objective of this paper is not to propose or
evaluate the taxonomic ranking of the species pair
concept. We just present detailed world-wide
maps of some taxa belonging to the same genus
(i.e. should be phylogenetically closely allied) but
varying in their reproductive means, and point out
the possible differences in these distribution pat-
terns.
However, to distinguish between primary and
secondary species using the presence of apothecia
versus soredia, isidia or lobules is not as simple as
it seems to be. Some species in Allocetraria,Cet-
relia and Cetreliopsis lack both vegetative repro-
ductive propagules and apothecia (Table 1). Some
of these can still be considered secondary species,
e.g. Cetrelia alaskana and Cetreliopsis papuae.
Contrary to the other species in these genera they
grow on the ground and presumably reproduce by
thallus fragmentation. The same does not with cer-
tainty apply to some other taxa, e.g. Allocetraria
endochrysea,A. denticulata and A. sinensis, al-
though they also grow on soil. Most Allocetraria
species usually grow on the ground, including spe-
cies with apothecia. The species mentioned above
are rare, A. denticulata even exceedingly rare, and
it is quite possible that fertile specimens might be
found in the future. In this paper, the status of "sec-
ondary species" is only assigned to taxa which
clearly produce either soredia, isidia or secondary
lobules. Taxa for which neither vegetative nor sex-
ual reprocuctive structures have been observed are
not included in that group.
Material and methods
Distribution maps were compiled using the com-
puter program DMAP. Both herbarium and litera-
ture data were used as sources for the localities.
Data of the checklists published on the Internet
(Feuerer 2004) were taken into consideration only
in exceptional cases, when no other data were
available. The sources for locality data of each li-
chen species are listed below (the corresponding
database is preserved in the Institute of Botany and
Ecology, University of Tartu).
Allocetraria ambigua: B, BM, COLO, GZU, H,
LD, S, TU; Randlane et al. (2001), Thell et al.
(1995b).
A. denticulata: Hue (1899).
A. endochrysea: GZU, H; Follmann et al.
(1968), Kärnefelt & Thell (1996), Randlane et al.
(2001).
A. flavonigrescens: GZU, UPS; Randlane et al.
(2001), Thell et al. (1995b).
A. globulans: GZU, H, UPS; Randlane et al.
(2001), Thell et al. (1995b).
A. isidiigera: TNS; Kurokawa & Lai (1991).
A. madreporiformis: GZU, TU; Golubkova et
al. (1996), Kärnefelt & Thell (1996), Randlane et
al. (2001), Thell et al. (2004).
A. oakesiana: B, GZU, FH, LD, S, TU, UPS;
Randlane et al. (2001), Thell et al. (1995b).
Distribution patterns of some cetrarioid species 363
Symb. Bot. Ups. 34:1
A. sinensis: COLO, GZU, LD, UPS; Randlane
et al. (2001), Thell et al. (1995b).
A. stracheyi: B, COLO, GZU, H, KW, LD, LE,
S, TU, UPS; Randlane et al. (2001), Thell et al.
(1995b).
Cetrelia alaskana: TU, UPS; Culberson & Cul-
berson (1968), Kurokawa (2003), Randlane &
Saag (1991), Urbanavichene & Urbanavichus
(2001).
C. braunsiana: B, C, GZU, LD, LE, TU; Cul-
berson & Culberson (1968), Feuerer (2004), Kash-
iwadani et al. (2002), Park (1990), Randlane &
Saag (1991), Sipman (1993), Wei (1991).
C. cetrarioides: B, GZU, H, KW, LD, LE, TU;
Barbero et al. (1995); Culberson & Culberson
(1968), Feuerer (2004), Fadeyeva et al. (1997),
Hafellner & Türk (2001), Kurokawa (2003), Rand-
lane & Saag (1991), Wei (1991), Wirth (1995).
C. chicitae: B, C, GZU, LD, TU; Barbero et al.
(1995), Culberson & Culberson (1968), Hafellner
& Türk (2001), Kurokawa (2003), Randlane &
Saag (1991), Wei (1991), Wirth (1995).
C. collata: Culberson & Culberson (1968),
Feuerer (2004), Wei (1991).
C. davidiana: Culberson & Culberson (1968),
Wei (1991).
C. delavayana: DUKE; Culberson & Culberson
(1968), Wei (1991).
C. isidiata: Culberson & Culberson (1968), Wei
(1991).
C. japonica: CANB, TU, UPS; Culberson &
Culberson (1968), Kashiwadani et al. (2002), Park
(1990), Randlane & Saag (1991), Sipman (1993).
C. monachorum: GZU, KW, TU; Barbero et al.
(1995), Culberson & Culberson (1968), Park
(1990), Randlane & Saag (1991).
C. nuda: H, TU, UPS; Culberson & Culberson
(1968).
C. olivetorum: C, GZU, H, KW, LD, TU; Bar-
bero et al. (1995), Culberson & Culberson (1968),
Fadeyeva et al. (1997), Elix (1994), Golubkova
(1981), Hafellner & Türk (2001), Park (1990),
Randlane & Saag (1991), Santesson et al. (2004),
Thell et al. (2004), Vitikainen et al. (1997), Wei
(1991).
C. orientalis: GZU, TU; Randlane & Saag
(1991).
C. pseudocollata: LD; Randlane & Saag
(1991).
C. pseudolivetorum: GZU, TU; Culberson &
Culberson (1968), Park (1990), Randlane & Saag
(1991), Wei (1991).
C. sanguinea: CANB, TU, UPS; Culberson &
Culberson (1968), Wei (1991).
C. sinensis: Culberson & Culberson (1968),
Wei (1991).
Cetreliopsis asahinae: H, GZU, KW, LD, LE,
PC, TU, WU; Randlane et al. (1995), Randlane et
al. (2001).
C. endoxanthoides: Herb. Awasthi; Randlane et
al. (1995).
C. hypotrachyna: BM; Randlane & Saag
(2003).
C. laeteflava: H, B, GZU, TAIM, TU, US, W;
Randlane et al. (1995), Räsänen (1949).
C. papuae: Herb. Aptroot, CANB, GZU, TU,
US; Randlane et al. (1995).
C. rhytidocarpa ssp. rhytidocarpa: FH, H, PC,
TNS, US; Lai & Elix (2002), Randlane et al.
(1995).
C. rhytidocarpa ssp. langtangi: GZU; Randlane
et al. (1995).
C. thailandica: TU; Lai & Elix (2002).
Tuckneraria ahtii: B, FH, GZU, H, S, TAIM,
TU, UPS, US, WU; Randlane et al. (1994), Rand-
lane et al. (2001).
T. laureri: B, GZU, LD, KW, M, MB, S, TAIM,
TNS, TU, UPS, US, W; Randlane et al. (1994),
Randlane et al. (2001).
T. laxa: H, TAIM; Lai (1980), Randlane et al.
(1994).
T. pseudocomplicata: DUKE, H, LD, M, TAIM,
TNS, TU, UPS, US, W; Randlane et al. (1994).
T. togashii: H, M, TAIM, TNS, TU; Thell et al.
(1995a), Wei (1991).
Results
Distribution of the genus Allocetraria
Of the ten Allocetraria species known at present
(Thell et al. 1995b, Randlane et al. 2001), eight are
here regarded as primary species. Six of these, A.
ambigua, A. denticulata, A. endochrysea, A. flavo-
nigrescens, A. globulans and A. sinensis, have lim-
364 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
ited distribution areas in the high mountains of
south-eastern Asia, historically called Tibet (at
present administratively divided between China,
India, Nepal, Bhutan and Myanmar) (Fig. 1). Two
primary species, A. madreporiformis and A. stra-
cheyi, are more widely dispersed and occur on two
continents, Eurasia and North America (Fig. 2).
Their distribution patterns are still not the same. A.
madreporiformis is sparsely but widely distributed
in Eurasia and somewhat more extensively in
western North America, whereas A. stracheyi is
concentrated to south-eastern Asia only and has a
few scattered localities in western North America
(Colorado, Yukon, Alberta and New Mexico). All
primary Allocetraria species except A. globulans
are mainly terricolous but rarely found also on
bases and twigs of small shrubs. A. globulans is
mainly corticolous and only occasionally terri-
colous.
Two species from this genus are regarded as
secondary, A. isidiigera and A. oakesiana. Still, the
status is doubtful for A. isidiigera. According to
the original description (Kurokawa & Lai 1991)
the lichen has scattered cylindrical isidia together
with apothecia. The importance or frequency of
these structures can not be evaluated as only the
type specimen is known for the species. The distri-
bution of A. isidiigera is of the same type as that of
most of the primary Allocetraria species. So far
the species is known from one single locality in
Xizang province, China, and it grows on soil like
most of the primary species.
A. oakesiana is the only sorediate taxon in the
genus. It also occurs on two continents (Eurasia
and North America) but has a more restricted dis-
tribution than the primary species A. madrepori-
formis. It occurs in the montane forests of Central
Europe, in the central part of North America (but
mainly in its eastern areas) and rarely in Asia
(Hubei and Xizang provinces in China, Omsk in
Russia, Japan). The species is corticolous on co-
niferous and deciduous trees and has never been
found growing on soil.
Thus, among the ten Allocetraria species, seven
are restricted to a rather limited area in south-east-
ern Asia, while three species are widely distributed
in Eurasia and North America. Both distribution
patterns are represented by primary as well as by
secondary species. The most extensively distrib-
uted taxon is primary A. madreporiformis. The
center of diversity of this genus evidently lies in
Tibet.
Distribution of the genus Cetrelia
At present 17 species of Cetrelia are recognised on
the basis of combinations of five morphotypes and
six chemotypes (Culberson & Culberson 1968,
Randlane & Saag 1991). The genus is a classic
example for the species pair concept (Poelt 1972,
Culberson & Culberson 1976). Two primary mor-
photypes can be identified, both having thalli with-
out any vegetative propagules: the collata-mor-
photype with large pseudocyphellae (C. nuda, C.
pseudocollata, C. collata) and the davidiana-mor-
photype with small pseudocyphellae (C. alaskana,
C. davidiana, C. delavayana, C. sanguinea). Of
these seven primary species six have more or less
limited distribution areas in Asia (Figs 3, 5 & 6). C.
collata is known from Bhutan, China and Nepal;
C. davidiana,C. delavayana and C. pseudocollata
from China; C. nuda from China, Japan and Tai-
wan; and C. sanguinea from China, Indonesia and
Japan. The seventh taxon, C. alaskana, is known
both from Asia and North America (Japan,
Khamar-Daban Mts and peninsula Tschukotka in
Russian Far East, and the west coast of Alaska;
Fig. 3). This species differs essentially from the
other members of the genus in its ecology: it grows
on the ground in tundra vegetation. All the other
Cetrelia species are epiphytic in various types of
forests. The distribution area of C. alaskana can be
considered close to Amphi-Beringian.
The secondary species in Cetrelia are repre-
sented by three morphotypes which are character-
ised by the presence of different vegetative prop-
agules: the sorediate cetrarioides-morphotype (C.
cetrarioides, C. chicitae, C. monachorum, C. olive-
torum); the isidiate isidiata-morphotype (C. braun-
siana, C. isidiata); and the sinensis-morphotype
with small flat lobules (C. japonica, C. orientalis,
C. pseudolivetorum, C. sinensis). The sorediate
taxa are most widespread and occur on two conti-
nents, Eurasia and North America (Fig. 4). C. ol-
Distribution patterns of some cetrarioid species 365
Symb. Bot. Ups. 34:1
Figure 1. World distribution of Allocetraria species I (after Randlane et al. 2001, complemented).
A. A. ambiqua. B. A. denticulata. C. A. endochrysea. D. A. flavonigrescens. E. A. globulans. F. A. isidiigera.
AB
CD
EF
366 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
F
igure 2. World distribution of Allocetraria species II (after Randlane et al. 2001, complemented). A. A. madrepori-
f
ormis. B. A. oakesiana. C. A. stracheyi. D. A. sinensis.
A
B
C
D
Distribution patterns of some cetrarioid species 367
Symb. Bot. Ups. 34:1
Figure 3. World distribution of Cetrelia species I (after Culberson & Culberson 1968, complemented).
A. C. alaskana. B. C. davidiana. C. C. delavayana. D. C. braunsiana. E. C. collata.
A
BC
D
E
368 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
Figure 4. World distribution of Cetrelia species II (after Culberson & Culberson 1968, complemented).
A. C. cetrarioides. B. C. chicitae. C. C. monachorum. D. C. olivetorum [C. o. s. str. (●) and C. o. s. l. (o)].
A
B
C
D
Distribution patterns of some cetrarioid species 369
Symb. Bot. Ups. 34:1
Figure 5. World distribution of Cetrelia species III (after Culberson & Culberson 1968, complemented).
A. C. isidiata. B. C. nuda. C. C. orientalis. D. C. pseudocollata. E. C. japonica. F. C. pseudolivetorum.
AB
CD
EF
370 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
Figure 6. World distribution of Cetrelia species IV (after Culberson & Culberson 1968, complemented).
A. C. sanguinea. B. C. sinensis.
AB
ivetorum has even been reported from South Amer-
ica (Argentina) and Australia. Species of the isidate
morphotype are distributed in Asia, C. braunsiana
in Bhutan, China, India, Malaysia, Nepal, Japan,
Philippines, Russia, South Korea and Taiwan, and
C. isidiata in China, Japan and Taiwan (Figs 3 &
5). The distribution areas of species with lobules
are also situated in Asia but their size varies. C.
japonica is known from Indonesia, Japan, Korea,
Malaysia and Taiwan; C. orientalis from the type
locality in the Far East of Russia and from Nepal;
C. pseudolivetorum from China, India, Japan, Ne-
pal, Russian Far East, South Korea and Taiwan; and
C. sinensis from China and Taiwan (Figs 5 & 6).
The center of diversity of the genus Cetrelia lies
in eastern Asia, probably in the southern parts of
China.
Distribution of the genus Cetreliopsis
Cetreliopsis includes seven species according to
our present knowledge (Randlane et al. 1995, Lai
& Elix 2002, Randlane & Saag 2003) and six of
them are treated as primary here. C. asahinae,
which has the widest distribution area in the genus,
occurs in the eastern part of Asia (Bhutan, China,
India, Japan, Nepal, Russian Far East, South Korea
and Vietnam) (Fig. 7). C. endoxanthoides and C.
hypotrachyna, the latter recently transferred from
Cetraria (Randlane & Saag 2003), are both known
from their type localities only, C. endoxanthoides
from Nepal and C. hypotrachyna from north-east-
ern India (Fig. 7). Cetreliopsis papuae is the only
member of the genus, which is not corticolous but
grows on the ground. It has been found in two sep-
arate localities in Papua New Guinea. C. rhytido-
carpa has two subspecies; ssp. rhytidocarpa is
known from Indonesia, Malaysia, Papua New
Guinea and Philippines, and ssp. langtangi from
Bhutan, Nepal and India. The recently described C.
thailandica (Lai & Elix 2002) is reported from
Thailand (type collection only).
The only secondary species in this genus, the
sorediate C. laeteflava, has a similar, rather re-
stricted distribution area (Taiwan and the Philip-
pines).
The center of diversity of the genus Cetreliopsis
seems to be double – one in Tibet and another
clearly more southern (Philippines, Indonesia).
Distribution of the genus Tuckneraria
The genus Tuckneraria comprises five species
(Randlane et al. 1994, Thell et al. 1995a), three of
which are considered primary. T. ahtii is distrib-
uted in Bhutan, southern provinces of China (Xi-
Distribution patterns of some cetrarioid species 371
Symb. Bot. Ups. 34:1
Figure 7. World distribution of Cetreliopsis species. A. C. asahinae. B. C. endoxanthoides. C. C. hypotrachyna.
D. C. laeteflava. E. C. papuae (●) and C. thailandica (■). F. C. rhytidocarpa [C. r. ssp. rhytidocarpa (o) and
C. r. ssp. langtangi (●)].
AB
CD
E F
372 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
zang, Sichuan, Yunnan), Taiwan and Nepal. T. laxa
is regarded as a Taiwanese endemic, known from
several localities (Lai 1980). T. pseudocomplicata
has the widest distribution area among the primary
Tuckneraria species, ranging from the Russian Far
East (Kuril islands, Sakhalin island, Primorye re-
gion) and Japan to Taiwan (Fig. 8).
Secondary species are represented by the isidi-
ate T. togashii and the sorediate T. laureri. The
former is known from several localities in Japan,
and a few localities have also been reported from
China. The latter has a very extensive distribution
area, covering Central Europe (Austria, Germany,
Italy, Poland, Romania, Slovakia, Switzerland,
Ukraine), Asia (Bhutan, China, India, Japan, Mon-
golia, Nepal, Russia) and South America (Colom-
bia, Venezuela).
All taxa of this genus are corticolous and grow
on coniferous or deciduous trees. The center of
diversity of the genus Tuckneraria is in Eastern
Asia (Japan, Taiwan).
Discussion
Primary species dominate in Allocetraria,Cetreli-
opsis,Tuckermanella,Tuckermanopsis,Tuckner-
aria and Vulpicida. The only cetrarioid genera
where secondary species dominate are Cetrelia
and Platismatia (Table 1). We have presented
world-wide distribution maps for 24 primary spe-
cies from four genera, Allocetraria,Cetrelia,Cet-
reliopsis and Tuckneraria. The majority of them
(22 taxa) have restricted distribution areas in the
region where the diversity and speciation centers
of these genera are most probably situated. How-
ever, two primary Allocetraria species, A. ma-
dreporiformis and A. stracheyi, are more wide-
spread and have spread from Asia to North Amer-
ica (Fig. 9a).
Of the 15 secondary species treated here, six
sorediate lichens, Allocetraria oakesiana, Cetrelia
cetrarioides, C. chicitae, C. monachorum, C. ol-
ivetorum and Tuckneraria laureri, occur exten-
sively on two, three or even four continents, al-
though the diversity and most probably also the
speciation centers of all these genera lie in rather
rectricted areas in Asia (Fig. 9c). Only one soredi-
ate taxon, Cetreliopsis laeteflava, has a limited dis-
tribution area close to the diversity center of the
genus Cetreliopsis.
Isidiate species are much less successful in ex-
tending their distribution (Fig. 9b). The four spe-
cies discussed, Allocetraria isidiigera,Cetrelia
braunsiana, C. isidiata and Tuckneraria togashii,
all occur exclusively in Asia. A. isidiigera and T.
togashii are endemic in China and Japan while C.
braunsiana and C. isidiata occur more widely.
The morphotypes with lobules in the genus Ce-
trelia are also grouped under secondary taxa.
Probably there is not much difference in the repro-
ductive efficiency of isidia and lobules. The distri-
bution of lobulate species demonstrates a pattern
similar to that of isidiate Cetrelia species. Some
occur in very limited or rather limited areas and
some more widely within Asia.
The determination of species pairs (noted as
primary species-secondary species below) which
phylogenetically would be real sister taxa is not
possible without phylogenetic analyses. Still,
some suggestions can be made based on morpho-
logical, chemical, ecological and distributional
data. In the genus Allocetraria, two presumed
pairs would be A. globulans-A. oakesiana and A.
ambigua-A. isidiigera. The primary species of
both pairs have very limited distribution while the
secondary species are distributed either narrowly
(A. isidiigera) or widely (A. oakesiana).
In the genus Cetrelia possible species pairs, tri-
plets or even tetrads of species have been identified
according to the chemosyndromes and suggested
direction of chemical evolution (Randlane & Saag
1991). These are C. nuda-C. chicitae, C. braun-
siana, C. orientalis;C. pseudocollata-C. japonica;
C. davidiana-C. olivetorum, C. pseudolivetorum;
C. sanguinea-C. isidiata;C. delavayana-C. ce-
trarioides; and C. collata-C. monachorum, C. sin-
ensis. All primary species are rare lichens with a
very restricted distribution. The distribution of the
secondary species varies from very wide (the sore-
diate C. olivetorum is known from four continents)
to rather limited (the lobulate C. sinensis is known
from China and Taiwan only).
Of the seven Cetreliopsis species only the sore-
diate C. laeteflava can be considered secondary. Its
Distribution patterns of some cetrarioid species 373
Symb. Bot. Ups. 34:1
Figure 8. World distribution of Tuckneraria species. A. T. laureri. B. T. ahtii. C. T. laxa. D. T. pseudocomplicata.
E. T. togashii.
A
B
C
DE
374 Tiina Randlane and Andres Saag
Symb. Bot. Ups. 34:1
Figure 9. Distribution patterns of primary and secondary species of the genera Allocetraria,Cetrelia,Cetreliopsis and
Tuckneraria. A. “primary” species (24 taxa). B. isidiate and lobulate “secondary” species (8 taxa). C. sorediate
“secondary” species (7 taxa).
A
B
C
Distribution patterns of some cetrarioid species 375
Symb. Bot. Ups. 34:1
primary counterpart could be either C. endoxan-
thoides or C. thailandica, representing the same
chemotype, with quaesitic acid as major medullary
substance (Randlane & Saag 2003). Both of them
are only known from type localities, while the sec-
ondary taxon C. laeteflava has a rather restricted
distribution area.
In the genus Tuckneraria species pairs cannot
with certainty be identified at the present level of
knowledge. Furthermore, the monophyly of the
genus is considered doubtful by some authors
(Thell et al. 2002) while in another study the ana-
lysed Tuckneraria species form a monophyletic
clade (Saag et al. 2002).
Acknowledgements
The authors are grateful to the keepers of the her-
baria mentioned in the text for kindly sending the
lichen specimens. Sincere thanks to the anony-
mous reviewers of the manuscript. Triin Randlane
is acknowledged for revising the English. This
study was financially supported by the Estonian
Science Foundation (grants 5505 and 5823).
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