The Old World Roccella species outside Europe and Macaronesia: Taxonomy, evolution and phylogeny

Abstract

With this paper the genus Roccella is complete and fully revised. The genus contains 24 species of which eight species from the Old World in Asia and Africa are here treated in detail, Roccella applanata, R. babingtonii, R. balfourii, R. boryi, R. minuta (newly described here), R. montagnei, R. phycopsioides (newly described here) and R. sinensis. Roccella tinctoria with a predominantly Macaronesian distribution has some outpost localities in south western Africa is also included in this treatise. A full species phylogeny is presented based on data from four molecular markers, RPB2, nuLSU, ITS 1 and 2 and an anonymous locus. The African–Asian species together with three Macaronesian species (R. allorgei, R. fuciformis, R. maderensis) form a poorly supported monophyletic clade. The sister group to that clade is the significantly supported group containing the American and European–Macaronesian species. Roccella montagnei is the most widespread of all Roccella species ranging from Australia, around the Indian Ocean and further on along the west coast of Africa up north to the Cape Verde Islands. It is a genetically variable species, but morphologically, anatomically and chemically distinct and it is here maintained as one species. The distribution of roccellic acid among the samples within the monophyletic species group Roccella applanata, R. babingtonii, R. boryi, R. montagnei was surprising: sorediate specimens contain roccellic acid whereas fertile specimens, except for four samples, lack roccellic acid.

Full text

Systematics and Biodiversity (2010), 8(2):223–246
Research Article
The Old World Roccella species outside Europe and Macaronesia:
taxonomy, evolution and phylogeny
ANDERS TEHLER1,MARTINIRESTEDT
2, MATS WEDIN1&DAMIENERTZ
3
1Naturhistoriska riksmuseet, Enheten f¨
or kryptogambotanik, Box 50007 S-104 05 Stockholm, Sweden
2Naturhistoriska riksmuseet, Molekyl¨
arsystematiska laboratoriet, Box 50007 S-104 05 Stockholm, Sweden
3Jardin botanique national de Belgique, D´
epartement Bryophytes et Thallophytes, Domaine de Bouchout, B-1860 Meise, Belgium
(Received 13 November 2009; revised 19 January 2010; accepted 19 February 2010)
With this paper the genus Roccella is complete and fully revised. The genus contains 24 species of which eight species from
the Old World in Asia and Africa are here treated in detail, Roccella applanata,R. babingtonii,R. balfourii,R. boryi,
R. minuta (newly described here), R. montagnei,R. phycopsioides (newly described here) and R. sinensis. Roccella tinctoria
with a predominantly Macaronesian distribution has some outpost localities in south western Africa is also included in this
treatise. A full species phylogeny is presented based on data from four molecular markers, RPB2, nuLSU, ITS 1 and 2 and
an anonymous locus. The African–Asian species together with three Macaronesian species (R. allorgei,R. fuciformis,R.
maderensis) form a poorly supported monophyletic clade. The sister group to that clade is the significantly supported group
containing the American and European–Macaronesian species. Roccella montagnei is the most widespread of all Roccella
species ranging from Australia, around the Indian Ocean and further on along the west coast of Africa up north to the Cape
Verde Islands. It is a genetically variable species, but morphologically, anatomically and chemically distinct and it is here
maintained as one species. The distribution of roccellic acid among the samples within the monophyletic species group
Roccella applanata,R. babingtonii,R. boryi,R. montagnei was surprising: sorediate specimens contain roccellic acid
whereas fertile specimens, except for four samples, lack roccellic acid.
Key words: Arthoniales, lichenized fungi, phylogenetic species, Roccellaceae, Roccella minuta,Roccella phycopsioides,
Roccella phylogeny, roccellic acid, species pairs
Introduction
The genus Roccella DC. contains 24 species; seven species
in Europe and Macaronesia, nine species in the Americas
and eight species in Africa and Asia. Here, we will treat
the taxonomy and nomenclature of the remaining Roccella
species from Asia and Africa not previously treated by
Teh l e r et al. (Tehler et al.,2004,2009a,2009b;Tehler&
Irestedt, 2007), by combining data from molecular phy-
logeny with morphological and chemical evidence. We will
also touch upon the biogeography of this exciting group in
aphylogeneticcontext.
Roccella is a subtropical genus of lichen fungi with a
preference for areas with Mediterranean climates extend-
ing into adjacent temperate and tropical regions, and it
is predominantly distributed over the northern hemisphere
(Tehler et al.,2004;Tehler&Irestedt,2007).Allspecies
are restricted to coastal habitats and may be very common
Correspondence to: Anders Tehler. E-mail: anders.tehler@nrm.se
locally. Roccella uses Trentepo hlia as photobiont and the
lichens are fruticose with a white-greyish to greyish-brown
usually pendent thallus of flattened or terete branches, nor-
mally between 5–15 cm long; the cortex is composed of
anticlinally arranged hyphae and the medulla is normally
white to pale brownish, several species have a character-
istically yellow medulla in the holdfast, e.g. R. phycopsis
and R. gracilis. The fruiting bodies are apothecioid, circu-
lar in outline, immersed or sessile, up to 2.5 mm in diam.,
with a pruinose disc. The thalline margin is more or less
developed; all species have a characteristically carbona-
ceous hypothecium, distinct from the neighbouring white
medulla. The asci are thick-walled, fissitunicate and con-
tain eight fusiform, 3-septate, hyaline spores, within the
size range of 15–35 ×4–8 µm. The conidia are filiform
and curved; most species produce soredia for vegetative
dispersal. Secondary metabolites are typically orcinol and
β-orcinol depsides, and aliphatic acids.
In Europe and Macaronesia Roccella is found through-
out the Mediterranean Ocean extending into the Black Sea
ISSN 1477-2000 print / 1478-0933 online
C
"2010 The Natural History Museum
DOI: 10.1080/14772001003789554
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224 A. Tehler et al.
(R. phycopsis), but the highest species diversity is found
in the islands of the Macaronesian Archipelago (Azores,
Canary Islands, Madeira and Cape Verde). The main Amer-
ican distribution is along the Pacific coast of Upper and
Lower California, Peru, the Galapagos Islands and various
localities around the Caribbean Sea. On the North Amer-
ican Pacific coast it extends to just north of Los Angeles
in California (c.34
◦north). On the Pacific coast of South
America it extends just south of the equator (south of Lima
in Peru, c.14
◦south). There are no Roccella species present
on the Atlantic coasts of North and South America or on
the Pacific coast south of the Atacama Desert. The species
previously known as Roccella portentosa from Chile was
shown unrelated to Roccella and recombined to Roccellina
portentosa (Tehler & Irestedt, 2007).
Materials and methods
Terminal taxa and taxon sampling
The study is based primarily on collections made by Anders
Teh l e r fr o m S ou t h A fr i c a (1 9 9 6) , M a ur i t iu s ( 2 00 3 ) , Ho n g
Kong (2004), India (2005), Kenya (2007), Socotra (2008),
Angola (2009); on collections made by Damien Ertz from
Madagascar (2008); on collections made by Andr´
eAptroot
from St Helena (2006); and on specimens from the herbaria:
B, BM, BR, Bradford, E, ESS, G, GLAM, H, HBG, KR, L,
LINN, M, OXF, PC, S, TUR, UPS.
Species sampled for DNA-sequencing were: Roccella
applanata,R. babingtonii,R. balfourii,R. boryi,R. min-
uta (newly described here), R. montagnei,R. phycopsioides
(newly described here), R. sinensis and R. tinctoria.We
tried to achieve a sample series with large morphological
variation and with a large and representative geographi-
cal distribution. A total of 163 samples including outgroup
were used for the molecular investigations representing all
hereby recognized Roccella species worldwide; 123 of the
Old World species’ DNA voucher collections were also used
for the TLC investigations. Designated voucher specimens
are deposited in (S) and (BR).
Molecular techniques and alignment
Four lo ci were sequenced and used in the phyl ogenetic
analyses, the nuclear ribosomal internal transcribed spac-
ers ITS 1 and ITS 2 (and the intermediate 5.8s region),
the nuclear large subunit ribosomal RNA gene (LSU), the
second largest RNA polymerase subunit (RPB2) and an
anonymous locus. The anonymous locus was amplified us-
ing the primers Cal-int2F (TTA CCT CTT TTC ACC TCC
CAC TTT GCA) or Cal-int3F (ACT TTG CAC GAG AGG
CCA AGC ATT) and Cal-int1R (AGT TGT CGC AGC
GGA ACA GTT GGA), Tehler et al. (2009a,2009b). For
further information about thermo cycling conditions, how
this locus was obtained and its potential affinity see Tehler
et al. (2009a). Extractions, amplifications and sequencing
procedures follow Tehler & Irestedt (2007), Tehler et al.
(2009a,2009b). The sequence fragments were assembled
to complete sequences with SeqMan II(tm) (DNASTAR
Inc.). Ambiguous nucleotide positions were coded with
the appropriate IUPAC codes. GenBank accession num-
bers are given in Table 1. Alignments were carried out
using the computer program ClustalX 2.0.9 (Larkin et al.,
2007) under Multiple Alignment Mode with all parame-
ters set to default. Before the final phylogenetic analyses
some minor manual adjustments were made in the ITS and
anonymous locus alignments at positions where ClustalX
obviously had failed to create the most parsimonious so-
lution. The aligned data sets are given in Appendix 1 (see
supplementary material which is available on the Supple-
mentary content tab of the article’s Informaworld page at
http://dx.doi.org/10.1080/14772001003789554).
Choice of outgroup
The outgroup taxa were chosen on the basis of earlier phylo-
genetic studies of the genus Roccella and the order Artho-
niales. They include at least two sample sequences from
all other recognized Roccella species (Tehler et al.,2004,
2009a,2009b; Tehler & Irestedt, 2007). The rooting taxon
was taken from the crustose genus Dirina which is the sister
group to Roccella.
Phylogenetic analysis
For the p hy logenetic analyses we u sed t he prog rams T.N.T.
Tree Analysis Using New Technology (Goloboff et al.,
2008) and; MrBayes (Ronquist & Huelsenbeck, 2003). In
all analyses gaps were treated as missing data.
The parsimony analyses used the New Technology search
with sectorial search, ratcheting, drifting, tree fusing and
driven search options in effect, all using default settings.
Resampling tree searches were done with parsimony jack-
knifing (Farris et al.,1996)undertheNewTechsearchas
implemented in T.N.T. (Goloboff et al., 2008); 1000 repli-
cates submitted to TBR branch swapping were conducted.
In parsimony jackknifing the data are internally resam-
pled with a jackknifing technique to find well-supported
groups. Resampling works by calculating a tree for each of
alargenumberofsubsamples(pseudoreplicates)ofchar-
acters from the data, then finding a summary tree, which
comprises the groups occurring in the majority of the trees
for subsamples. The tree for each pseudoreplicate is found
by parsimony analysis, and each pseudoreplicate is formed
by randomly selecting characters from the data without re-
placement, each character having a fixed chance 1/e (about
36%) of being excluded. With this resampling technique
the actual number of characters used may vary from repli-
cate to replicate. Groups found in less than 50% (search 1)
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The Old World Roccella species 225
Table 1. Specimens used in phylogenetic analyses. GenBank accession numbers in bold are new sequences for this study.
Herbarium Acc. no. Acc. no. Acc. no. Acc. no.
Species Locality Year Collector Coll.no. & id.no. RPB2 LSU ITS anonymous
Dirina catalinariae Mexico. Baja California, Ensenada, Pta Banda 2007 Tehler 9087 S-F148386 GU137543 GU137909 GU137787 GU137663
Dirina ceratoniae Spain. Mallorca, s’Amarodor 2007 Tehler 9021 S-F66044 GU137544 GU137910 GU137788 GU137664
Dirina ceratoniae Spain. Mallorca, Sa R´
apita 2007 Tehler 9047 S-F66048 FJ639025 FJ638966 FJ639084 FJ638893
Dirina paradoxa ssp.
africana
Mauritius. Black River, Mt St. Pierre 2003 Tehler 8524 S-L55029 DQ987629 EF081386 EF081375 GU137665
Roccella albida Ecuador. Galapagos Islands, Isla R´
abida 2005 Tehler 8780 S-F63119 FJ639026 FJ638967 EU449864 EU449775
Roccella albida Ecuador. Galapagos Islands, Santa F´
e2005Tehler8653-14S-L68229DQ987651EF081408EU449860EU449771
Roccella allorgei Portugal: Azores. S˜
ao Miguel 2000 Tehler 8163 S-L12507 DQ987640 EF081397 FJ639085 FJ638895
Roccella allorgei Portugal: Azores. S˜
ao Miguel, Ponta Delgada 2000 Tehler 8151 S-L12529 DQ987639 EF081396 AJ634005 FJ638894
Roccella applanata Madagascar. Toliara (Tul´
ear), Ifaty 2008 Ertz 13058 BR-8695-62 GU137545 GU137911 GU137789 GU137666
Roccella applanata Madagascar. Toliara (Tul´
ear), Ifaty 2008 Ertz 13060 BR-8697-64 GU137546 GU137912 GU137790 GU137667
Roccella applanata Madagascar. Toliara (Tul´
ear), Ifaty 2008 Ertz 13075 BR-8713-80 GU137547 GU137913 GU137791 GU137668
Roccella applanata Madagascar. Toliara (Tul´
ear), La Table 2008 Ertz 13104 BR-8743-13 GU137548 GU137914 GU137792 GU137669
Roccella applanata Madagascar. Toliara (Tul´
ear), Ifaty 2006 Emanuelsson 101 S-F58632 GU137549 GU137915 GU137793 GU137670
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9231 S-F146175 GU137550 GU137916 GU137794 GU137671
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9233 S-F146176 GU137551 GU137917 GU137795 GU137672
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9258 S-F146177 GU137557 GU137923 GU137801 GU137678
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9234 S-F146178 GU137552 GU137918 GU137796 GU137673
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Watamu 2007 Tehler 9242-18 S-F146179 GU137553 GU137919 GU137797 GU137674
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Watamu 2007 Tehler 9242-19 S-F146179 GU137554 GU137920 GU137798 GU137675
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Ngomeni 2007 Tehler 9251-27 S-F146180 GU137555 GU137921 GU137799 GU137676
Roccella babingtonii Kenya. Coast prov., Kilifi distr., Ngomeni 2007 Tehler 9251-28 S-F146180 GU137556 GU137922 GU137800 GU137677
Roccella bajasurensis Mexico. Baja California Sur, Playa San Pedro 2007 Tehler 9188 S-F113351 FJ639029 FJ638970 FJ639088 FJ638898
Roccella bajasurensis Mexico. Baja California Sur, La Palma 2007 Tehler 9192 S-F113352 FJ639030 FJ638971 FJ639089 FJ638899
Roccella balfourii Yemen. Socotra Island, Qalansiyah 2008 Tehler 9350 S-F146181 GU137564 GU137930 GU137808 GU137685
Roccella balfourii Yemen. Socotra Island, Homill 2008 Tehler 9302 S-F146182 GU137558 GU137924 GU137802 GU137679
Roccella balfourii Yemen. Socotra Island, Homill 2008 Tehler 9306-6 S-F146183 GU137559 GU137925 GU137803 GU137680
Roccella balfourii Yemen. Socotra Island, Homill 2008 Tehler 9306-7 S-F146183 GU137560 GU137926 GU137804 GU137681
Roccella balfourii Yemen. Socotra Island, in the south 2008 Tehler 9326 S-F146184 GU137561 GU137927 GU137805 GU137682
Roccella balfourii Yemen. Socotra Island, Sefflah 2008 Tehler 9335 S-F146185 GU137562 GU137928 GU137806 GU137683
Roccella balfourii Yemen. Socotra Island, Ras Ersel 2008 Tehler 9349 S-F146186 GU137563 GU137929 GU137807 GU137684
Roccella boryi Mauritius. Port Louis, Port Louis, Mt. Signal 2003 Tehler 8500-1 S-L55010 GU137565 GU137931 GU137809 GU137686
Roccella boryi Mauritius. Port Louis, Port Louis, Mt. Signal 2003 Tehler 8500-2 S-L55010 GU137566 GU137932 GU137810 GU137687
Roccella boryi Mauritius. Port Louis, Port Louis, Mt. Signal 2003 Tehler 8501 S-L55011 GU137567 GU137933 GU137811 GU137688
Roccella boryi Mauritius. Black River, Pointe Corail de la Prairie 2003 Tehler 8506 S-L55017 DQ987641 EF081398 EF081376 FJ638902
Roccella boryi Mauritius. Black River, Pointe Corail de la Prairie 2003 Tehler 8507 S-L55018 GU137568 GU137934 GU137812 GU137689
Roccella boryi Mauritius. Port Louis, Port Louis, Mt. Signal 2003 Tehler 8516 S-L55024 DQ987642 EF081399 FJ639092 FJ638903
Roccella boryi Mauritius. Port Louis, Port Louis, Mt. Signal 2003 Tehler 8518 S-L55026 GU137569 GU137935 GU137813 GU137690
Roccella boryi Mauritius. Black River, Mt St. Pierre 2003 Tehler 8526 S-L55032 GU137570 GU137936 GU137814 GU137691
Roccella boryi Mauritius. Black River, Mt St. Pierre 2003 Tehler 8527 S-L55033 GU137571 GU137937 GU137815 GU137692
Roccella boryi Mauritius. Plain Wilhelms, Corps de Garde 2003 Tehler 8532-28 S-L55036 GU137572 GU137938 GU137816 GU137693
Roccella boryi Mauritius. Plain Wilhelms, Corps de Garde 2003 Tehler 8532-29 S-L55036 GU137573 GU137939 GU137817 GU137694
Roccella boryi Mauritius. Plain Wilhelms, Corps de Garde 2003 Tehler 8533-30 S-L55037 GU137574 GU137940 GU137818 GU137695
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226 A. Tehler et al.
Table 1. Specimens used in phylogenetic analyses. GenBank accession numbers in bold are new sequences for this study. (Continued)
Herbarium Acc. no. Acc. no. Acc. no. Acc. no.
Species Locality Year Collector Coll.no. & id.no. RPB2 LSU ITS anonymous
Roccella boryi Mauritius. Plain Wilhelms, Corps de Garde 2003 Tehler 8533-31 S-L55037 GU137575 GU137941 GU137819 GU137696
Roccella boryi Mauritius. Savanne, Maconde 2003 Tehler 8538 S-L55038 GU137577 GU137943 GU137821 GU137698
Roccella boryi Mauritius. Savanne, Maconde 2003 Tehler 8537 S-L55039 GU137576 GU137942 GU137820 GU137697
Roccella decipiens Mexico. Baja California, Ensenada, Pta Banda 2007 Tehler 9080 S-F114301 FJ639034 FJ638975 FJ639095 FJ638907
Roccella decipiens Mexico. Baja California Sur, Punta Eugenia 2007 Tehler 9148 S-F114308 FJ639037 FJ638978 FJ639098 FJ638910
Roccella elisabethae Canary Islands, Tenerife, Anaga, Roque Enmedio 2000 Tehler 8272 S-L14811 DQ987646 EF081403 AJ634013 FJ638911
Roccella elisabethae Canary Islands, Tenerife, Buenavista del Norte 2001 Tehler 8306-45 S-L14846 DQ987647 EF081404 AJ634014 FJ638912
Roccella fuciformis Portugal: Azores. S˜
ao Miguel, Ponta Delgada 2000 Tehler 8171 S-L12540 FJ639038 FJ638979 AJ634015 Failed sequence
Roccella fuciformis Canary Islands, Tenerife, G¨
uimar 2000 Tehler 8255 S-L14794 DQ987649 EF081406 AJ634016 FJ638913
Roccella galapagoensis Ecuador. Galapagos Islands, Santa F´
e 2005 Tehler 8651-12 S-L68226 DQ987650 EF081407 EU449894 EU449805
Roccella galapagoensis Ecuador. Galapagos Islands, Isla Santa Cruz 2005 Tehler 8667 S-L72900 DQ987653 EF081410 EU449882 EU449793
Roccella gracilis Mexico. Baja California Sur, Cabo San Lucas 2007 Tehler 9190 S-F114058 FJ639072 FJ639013 FJ639131 FJ638946
Roccella gracilis Ecuador. Galapagos Islands, Santa F´
e 2005 Tehler 8654 S-L68231 DQ987655 EF081412 EU449895 EU449806
Roccella lirellina Peru. City of Lima, Morro Solar 2006 Tehler 8880 S-F79521 FJ639074 FJ639015 EU449898 EU449809
Roccella lirellina Peru. Chiclayo , Cerro de Reque 200 6 Tehler 8904 S-F 79525 FJ63 9075 FJ639016 EU449903 EU449814
Roccella maderensis Portugal: Azores. S˜
ao Miguel, Mosteiros 2000 Tehler 8157 S-L12561 DQ987660 EF081417 AJ634021 FJ638949
Roccella maderensis Portugal: Estremadura. Cabo da Roca 2000 Tehler 8140 S-L12563 DQ987658 EF081415 AJ634018 FJ638948
Roccella margaritifera Ecuador. Galapagos Islands, Isla R´
abida 2005 Tehler 8705 S-F63259 FJ639077 FJ639018 EU449924 EU449835
Roccella margaritifera Ecuador. Galapagos Islands, Santa F´
e 2005 Tehler 8656 S-L68234 DQ987657 EF081414 EU449907 EU449818
Roccella minuta Yemen. Socotra Island, Homill 2008 Tehler 9303 S-F146187 GU137578 GU137944 GU137822 GU137699
Roccella minuta Yemen. Socotra Island, Homill 2008 Tehler 9307-8 S-F146188 GU137579 GU137945 GU137823 GU137700
Roccella minuta Yemen. Socotra Island, Homill 2008 Tehler 9307-9 S-F146188 GU137580 GU137946 GU137824 GU137701
Roccella minuta Yemen. Socotra Island, Sefflah 2008 Tehler 9334 S-F146189 GU137581 GU137947 GU137825 GU137702
Roccella minuta Yemen. Socotra Island, Sefflah 2008 Tehler 9336 S-F146190 GU137582 GU137948 GU137826 GU137703
Roccella minuta Yemen. Socotra Island, Qalansiyah 2008 Tehler 9351 S-F146191 GU137583 GU137949 GU137827 GU137704
Roccella montagnei Madagascar. Isalo Massif near ANGAP house 2008 Ertz 13020 BR-7379-07 GU137660 GU138024 GU137906 GU137784
Roccella montagnei Madagascar. Isalo Massif near ANGAP house 2008 Ertz 13036 BR-7387-15 GU137661 GU138025 GU137907 GU137785
Roccella montagnei Madagascar. Toalanaro (Fort Dauphin) 2008 Ertz 13213 BR-7485-16 GU137588 GU137954 GU137832 GU137709
Roccella montagnei Madagascar. Toalanaro (Fort Dauphin) 2008 Ertz 13214 BR-7486-17 GU137589 GU137955 GU137833 GU137710
Roccella montagnei Madagascar. Toalanaro (Fort Dauphin) 2008 Ertz 13219 BR-7489-20 GU137653 GU138017 GU137899 GU137777
Roccella montagnei Madagascar. Toliara (Tul´
ear)f Ifaty 2008 Ertz 13054 BR-8691-58 GU137584 GU137950 GU137828 GU137705
Roccella montagnei Madagascar. Toliara (Tul´
ear), Ifaty 2008 Ertz 13069 BR-8707-74 GU137662 GU138026 GU137908 GU137786
Roccella montagnei Madagascar. Toliara (Tul´
ear)f Ifaty 2008 Ertz 13074 BR-8712-79 GU137585 GU137951 GU137829 GU137706
Roccella montagnei Madagascar. Toliara (Tul´
ear), La Table 2008 Ertz 13111 BR-8750-20 GU137586 GU137952 GU137830 GU137707
Roccella montagnei Madagascar. Toliara (Tul´
ear), La Table 2008 Ertz 13112 BR-8751-21 GU137587 GU137953 GU137831 GU137708
Roccella montagnei Namibia. Omaruru distr., Laguneberg 2001 Wirth 1 KR-s.n. GU137651 GU138015 GU137897 GU137775
Roccella montagnei Namibia. Omaruru distr., Laguneberg 2001 Wirth 2 KR-s.n. GU137652 GU138016 GU137898 GU137776
Roccella montagnei Kenya. Coast prov., Kwale distr., Shimoni 2007 Tehler 9221 S-F146192 GU137595 GU137961 GU137840 GU137718
Roccella montagnei Kenya. Coast prov., Kwale distr., Shimoni 2007 Tehler 9225 S-F146193 GU137597 GU137963 GU137842 GU137720
Roccella montagnei Kenya. Coast prov., Kilifi distr., Gedi, Gedi Ruins 2007 Tehler 9239-15 S-F146194 GU137605 GU137971 GU137850 GU137728
Roccella montagnei Kenya. Coast prov., Kilifi distr., Gedi, Gedi Ruins 2007 Tehler 9239-16 S-F146194 GU137606 GU137972 GU137851 GU137729
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The Old World Roccella species 227
Roccella montagnei Kenya. Coast prov., Kilifi distr., Watamu, Temple 2007 Tehler 9241 S-F146195 GU137607 GU137973 GU137852 GU137730
Roccella montagnei Kenya. Coast prov., Kilifi distr., Kilifi 2007 Tehler 9227 S-F146197 GU137598 GU137964 GU137843 GU137721
Roccella montagnei Kenya. Coast prov., Kwale distr., Kwale 2007 Tehler 9244 S-F146198 GU137608 GU137974 GU137853 GU137731
Roccella montagnei Kenya. Coast prov., Kwale distr., Moonlight Bay 2007 Tehler 9245 S-F146199 GU137609 GU137975 GU137854 GU137732
Roccella montagnei Kenya. Coast prov., Kwale distr., Diani Beach 2007 Tehler 9246 S-F146200 GU137610 GU137976 GU137855 GU137733
Roccella montagnei Kenya. Coast prov., Kwale distr., Shimba Hills 2007 Tehler 9247 S-F146201 GU137611 GU137977 GU137856 GU137734
Roccella montagnei Kenya. Coast prov., Kilifi distr., just N Vipingo 2007 Tehler 9248 S-F146202 GU137612 GU137978 GU137857 GU137735
Roccella montagnei Kenya. Coast prov., Kilifi distr., Ngomeni 2007 Tehler 9250 S-F146203 GU137614 GU137980 GU137859 GU137737
Roccella montagnei Kenya. Coast prov., Kilifi distr., Sabaki 2007 Tehler 9252 S-F146204 GU137615 GU137981 GU137860 GU137738
Roccella montagnei Kenya. Coast prov., Taita distr., Tsavo 2007 Tehler 9254 S-F146205 GU137617 GU137983 GU137862 GU137740
Roccella montagnei Kenya. Coast prov., Kilifi distr., Kenyatta Beach 2007 Tehler 9257 S-F146206 GU137619 GU137985 GU137864 GU137742
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9259 S-F146207 GU137620 GU137986 GU137865 GU137743
Roccella montagnei Kenya. Coast prov., Kilifi distr., Kilfi 2007 Tehler 9262 S-F146208 GU137622 GU137988 GU137867 GU137745
Roccella montagnei Kenya. Coast prov., Kilifi distr., Kilfi 2007 Tehler 9263 S-F146209 GU137623 GU137989 GU137868 GU137746
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9232-7 S-F146210 GU137599 GU137965 GU137844 GU137722
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9232-8 S-F146210 GU137600 GU137966 GU137845 GU137723
Roccella montagnei Kenya. Coast prov., Kilifi distr., Sabaki 2007 Tehler 9253 S-F146211 GU137616 GU137982 GU137861 GU137739
Roccella montagnei Kenya. Coast prov., Kwale distr., Shimoni 2007 Tehler 9222 S-F146212 GU137596 GU137962 GU137841 GU137719
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9235-11 S-F146213 GU137601 GU137967 GU137846 GU137724
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9235-14 S-F146213 GU137602 GU137968 GU137847 GU137725
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9236 S-F146214 GU137603 GU137969 GU137848 GU137726
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9237 S-F146215 GU137604 GU137970 GU137849 GU137727
Roccella montagnei Kenya. Coast prov., Kilifi distr., just N Vipingo 2007 Tehler 9249 S-F146216 GU137613 GU137979 GU137858 GU137736
Roccella montagnei Kenya. Coast prov., Taita distr., Tsavo 2007 Tehler 9255 S-F146217 GU137618 GU137984 GU137863 GU137741
Roccella montagnei Kenya. Coast prov., Kilifi distr., Mida Creek 2007 Tehler 9260 S-F146218 GU137621 GU137987 GU137866 GU137744
Roccella montagnei Yemen. Socotra Island, in the south 2008 Tehler 9325 S-F146219 GU137625 GU137991 GU137870 GU137748
Roccella montagnei Yemen. Socotra Island, in the south 2008 Tehler 9332 S-F146220 GU137626 GU137992 GU137871 GU137749
Roccella montagnei Yemen. Socotra Island, Sefflah 2008 Tehler 9337 S-F146221 GU137627 GU137993 GU137872 GU137750
Roccella montagnei Yemen. Socotra Island, Qalansiyah 2008 Tehler 9352a S-F146222 GU137628 GU137994 GU137873 GU137751
Roccella montagnei Angola. Luanda prov., Cacuaco 2009 Tehler 9702 S-F146224 GU137631 GU137997 GU137876 GU137754
Roccella montagnei Angola. Bengo prov., Barra do Dande 2009 Tehler 9708 S-F146225 GU137632 GU137998 GU137877 GU137755
Roccella montagnei Angola. Bengo prov., Barra do Dande 2009 Tehler 9714 S-F146227 GU137633 GU137999 GU137878 GU137756
Roccella montagnei Angola. Bengo prov., Lifune River 2009 Tehler 9718 S-F146229 GU137634 GU138000 GU137879 GU137757
Roccella montagnei Angola. Bengo prov., Tabi 2009 Tehler 9719 S-F146230 GU137635 GU138001 GU137880 GU137758
Roccella montagnei Angola. Bengo prov., Barra do Dande 2009 Tehler 9723-11 S-F146232 GU137636 GU138002 GU137881 GU137759
Roccella montagnei Angola. Bengo prov., Barra do Dande 2009 Tehler 9723-12 S-F146232 GU137637 GU138003 GU137882 GU137760
Roccella montagnei Angola. Bengo prov., Ambriz 2009 Tehler 9724 S-F146233 GU137638 GU138004 GU137883 GU137761
Roccella montagnei Angola. Luanda prov., Palmeirinhas 2009 Tehler 9728 S-F146234 GU137639 GU138005 GU137884 GU137762
Roccella montagnei Angola. Luanda prov., Palmeirinhas 2009 Tehler 9729 S-F146235 GU137640 GU138006 GU137885 GU137763
Roccella montagnei Angola. Luanda prov., Palmeirinhas 2009 Tehler 9735 S-F146237 GU137641 GU138007 GU137886 GU137764
Roccella montagnei Angola. Bengo prov., Caba Ledo 2009 Tehler 9740 S-F146239 GU137642 GU138008 GU137887 GU137765
Roccella montagnei Angola. Kwanza-Sul prov., Porto Amboim 2009 Tehler 9743 S-F146241 GU137643 GU138009 GU137888 GU137766
Roccella montagnei Angola. Kwanza-Sul prov., Sumbe 2009 Tehler 9747 S-F146243 GU137644 GU138010 GU137889 GU137767
(Continued on next page)
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228 A. Tehler et al.
Table 1. Specimens used in phylogenetic analyses. GenBank accession numbers in bold are new sequences for this study. (Continued)
Herbarium Acc. no. Acc. no. Acc. no. Acc. no.
Species Locality Year Collector Coll.no. & id.no. RPB2 LSU ITS anonymous
Roccella montagnei Angola. Luanda prov., Cacuaco 2009 Tehler 9701 S-F146244 GU137630 GU137996 GU137875 GU137753
Roccella montagnei Angola. Luanda prov., Cacuaco 2009 Tehler 9700 S-F146245 GU137629 GU137995 GU137874 GU137752
Roccella montagnei Yemen. Socotra Island, Homill 2008 Tehler 9309a S-F146247 GU137624 GU137990 GU137869 GU137747
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66650-109 S-F86656 GU137645 Fail ed se que n ce GU137890 GU137768
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66650-110 S-F86656 GU137646 Fail ed se que n ce GU137891 GU137769
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66616 S-F86659 GU137650 GU138014 GU137895 GU137773
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66253 S-F86661 GU137648 GU138012 GU137893 GU137771
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66251 S-F86668 GU137647 GU138011 GU137892 GU137770
Roccella montagnei South Atlantic Islands. St Helena 2006 Aptroot 66613 S-F86669 GU137649 GU138013 GU137894 GU137772
Roccella montagnei South Africa. Cape Peninsula, Cape Point 1996 Tehler 7709 S-L2872 GU137590 GU137956 AF110355 GU137711
Roccella montagnei India. Kancheepuram, Mamallapuram 2005 Tehler 8863 S-L67940 GU137591 GU137957 GU137834 GU137712
Roccella montagnei India. Kancheepuram, Mamallapuram 2005 Tehler 8864 S-L67946 GU137592 GU137958 GU137835 GU137713
Roccella montagnei India. Kancheepuram, Koovathur 2005 Tehler 8865 S-L67951 DQ987661 EF081418 FJ639133 FJ638950
Roccella montagnei India. Kancheepuram, Vadanemeli 2005 Tehler 8868 S-L67961 GU137593 GU137959 GU137836 GU137714
Roccella montagnei India. Kancheepuram, Vadanemeli 2005 Tehler 8869 S-L67962 GU137594 GU137960 GU137837 GU137715
Roccella montagnei India. Kancheepuram, Vadanemeli 2005 Tehler 8870 S-L67977 DQ987662 EF081419 GU137838 GU137716
Roccella montagnei India. Cuddalore, Pithcavaram 2005 Tehler 8878 S-L68093 DQ987663 EF081420 FJ639134 FJ638951
Roccella montagnei India. Cuddalore, Pithcavaram 2005 Tehler 8879 S-L68094 DQ987664 EF081421 GU137839 GU137717
Roccella montagnei South Africa. Cape District, Cape of Good Hope 2003 Feuerer s.n. S-L68743 DQ987665 EF081422 GU137896 GU137774
Roccella nigerrima Ecuador. Galapagos Islands, Santa F´
e 2005 Tehler 8655 S-L68233 DQ987656 EF081413 EU449926 EU449837
Roccella nigerrima Ecuador. Galapagos Islands, Isla Santa Cruz 2005 Tehler 8666 S-L72899 DQ987652 EF081409 EU449946 EU449857
Roccella phycopsioides Yemen. Socotra Island, Homill, in the east 2008 Tehler 9309b S-F146223 GU137655 GU138019 GU137901 GU137779
Roccella phycopsioides Yemen. Socotra Island, Homill, in the east 2008 Tehler 9308 S-F146246 GU137654 GU138018 GU137900 GU137778
Roccella phycopsioides Yemen. Socotra Island, in the south 2008 Tehler 9331 S-F146248 GU137656 GU138020 GU137902 GU137780
Roccella phycopsioides Yemen. Socotra Island, Qalansiyah 2008 Tehler 9352b S-F146249 GU137657 GU138021 GU137903 GU137781
Roccella phycopsis Portugal: Azores. Terceira, Ponta das Cavalas 2000 Tehler 8221 S-L12558 DQ987666 EF081423 AJ634031 FJ638952
Roccella phycopsis Canary Islands, Tenerife, El Tanque 2000 Tehler 8295 S-L14834 DQ987667 EF081424 AJ634033 FJ638953
Roccella sinensis China, Hong Kong Isl., Shek O, Shek O 2004 Tehler 8562-1 S-L60290 DQ987668 EF081425 FJ639135 FJ638954
Roccella sinensis China, Hong Kong Isl, Shek O, Shek O 2004 Tehler 8562-2 S-L60290 DQ987669 EF081426 FJ639136 FJ638955
Roccella sinensis China, Hong Kong Isl, Big Wave Bay 2004 Tehler 8564 S-L60292 GU137658 GU138022 GU137904 GU137782
Roccella sinensis China, Hong Kong Isl, South Bay Beach 2004 Tehler 8571 S-L60299 GU137659 GU138023 GU137905 GU137783
Roccella tinctoria Canary Islands, Tenerife, G¨
uimar 2000 Tehler 8257 S-L14796 DQ987671 EF081428 AJ634040 FJ638957
Roccella tinctoria Cape Verde. Boa Vista, Sal-Rei 2004 V¨
are 1784 S-L59866 DQ987644 EF081401 FJ639093 FJ638905
Roccella tuberculata Portugal: Estremadura. Cabo da Roca 2000 Tehler 8144 S-L12564 DQ987673 EF081430 AJ634043 FJ638959
Roccella tuberculata Canary Islands, Tenerife, Buenavista del Norte 2001 Tehler 8310-50 S-L14850 FJ639078 FJ639019 AJ634047 FJ638960
Roccella verruculosa Netherland Antilles, Bonaire, Washington-Slagbaai 2006 Tehler 8941-32 S-F69355 FJ639080 FJ639021 FJ639139 FJ638962
Roccella verruculosa Netherland Antilles, Bonaire, Washington-Slagbaai 2006 Tehler 8950-41 S-F69357 FJ639083 FJ639024 FJ639142 FJ638965
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The Old World Roccella species 229
and 95% (search 2) of the trees for pseudoreplicates were
discarded, thus eliminating unjustified (poorly supported)
resolution caused by ambiguous data sets.
Bayesian inference (Huelsenbeck et al.,2001;Holder
&Lewis,2003)wasalsousedtoestimatethephylogenetic
relationships. The models for nucleotide substitutions used
in the analyses were selected for each gene individually by
applying the Akaike Information Criterion (AIC) (Akaike,
1973) and the program MrModeltest 2.2 (Nylander, 2005)
in conjunction with PAUP∗(Swofford, 1998). Posterior
probabilities of trees and parameters in the substitution
models were approximated with MCMC and Metropolis
coupling using the program MrBayes 3.2 (Ronquist &
Huelsenbeck, 2003). Analyses were performed for both
the individual gene partitions and the combined data
set. In the analysis of the combined data set the models
selected for the individual gene partition were used, but
the topology was constrained to be the same. All chains
were r un for 100 million generations, with trees sampled
every 1000th generations. The trees sampled during
the burn-in phase (i.e. before the chain had reached its
apparent target distribution) were discarded, and after
checking for convergence, final inference was made from
the concatenated output from the two runs (180 000 trees).
The a priori selection of nucleotide substitution models
suggested that the GTR +Gmodelhadthebestfitforthe
RPB2 data set while the GTR +I+Gmodelhadthebest
fit for ITS, LSU and the anonymous locus.
Chemistry
For rou tin e scr eening of secondary chemist r y, ‘spot-test s’
on thalli and apothecia were performed with a water solu-
tion of potassium hydroxide (‘K’) and with a commercial
solution of potassium hypochlorite (‘C’). Thin-layer chro-
matography (TLC) of acetone extracts was performed on
20 ×20 cm silica gel 60 F254 layer aluminium or glass
plates using solvent systems A, B, C, EA and G. For the
visualization of the spots, 10% sulphuric acid was used as
a reagent (Orange et al.,2001).Lepraria membranacea,
Punctelia subrudecta and Trap elia involu ta were used as
control species for the identification of respectively roccel-
lic, lecanoric and gyrophoric acids
Taxonomy
Key to Old World Roccella species outside Europe
(for generic description and keys to other Roccella species the
reader is referred to Tehler et al.,2004,2009a,2009b)
1. Medulla at least in some parts
yellow
2
1. Medulla never yellow 4
2. Soralia present; apothecia absent Roccella phycopsioides
2. Apothecia present; soralia absent 3
3. Medulla yellow right below
hypothecium, usually spreading
into branches; thallus long,
pendulous, corticolous
Roccella babingtonii
3. Medulla only yellow in lowermost
holdfast; thallus short, erect,
saxicolous (endemic to Socotra
and Africa’s Horn)
Roccella minuta
4. Apothecia present; soralia absent 5
4. Soralia present; apothecia absent 10
5. Ascospores (20) 21 (22) µm long 6
5. Ascospores 24 µmorlonger 7
6. Branch tips K+reddish (endemic
to Mascarene Islands)
Roccella boryi
6. Branch tips K−(southern China) Roccella sinensis
7. Ascospore width 4–4,5 (5) µm;
branches strap-like or flattened
8
7. Ascospore width 6–7 µm;
branches approximately terete
9
8. Branches rarely more than 5 mm
wide
Roccella montagnei
8. Branches more than 5 mm up to
20(–30) mm wide
Roccella applanata
9. Branches, whitish with chalky
surface (endemic to Socotra and
Africa’s Horn)
Roccella balfourii
9. Branches, brownish, non-chalky
(a Macaronesian species with
outposts in Angola, Senegal,
Sierra Leone)
Roccella tinctoria
10. Soralia C+red (a Macaronesian
species with outposts in Senegal,
Sierra Leone, Angola)
Roccella tinctoria
10. Soralia C−11
11. Thallus strap-like or more or less
flattened, greenish-grey
Roccella montagnei
11. Thallus approximately terete,
whitish, greyish, beige or
brownish (endemic to Mascarene
Islands)
Roccella boryi
The species
For rep res ent ative spe cim ens examin ed re fer t o Ap-
pendix 2, List of specimens, see supplementary
material which is available on the Supplementary
content tab of the article’s Informaworld page at
http://dx.doi.org/10.1080/14772001003789554.
Roccella applanata Choisy (Fig. 1)
Ann. Cryptog. Exot.2:67(1929).Type:Madagascar.s.ann.,
s.coll.s.n.(LYJB-2323,holotype)
Thallus pendent to erect, branches broadly flattened up
to 30 mm wide, smooth to slightly foveate, 5–15 cm long,
greyish to green-greyish; cortex palisade plectenchyma;
epicortex absent; soralia absent; medulla white; in hold-
fast usually brown. Apothecia present, sessile; ascospores
24 ×4(L.max=25.6 µm; L.min =23.2 µm).
Chemistry: spot tests, thallus surface K−,C+red; medulla
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230 A. Tehler et al.
Fig. 1. Roccella applanata.Madagascar,Distr.Toliara.Toliara,
2006, E. Emanuelsson 101 (S-F58632). Scale mm.
K−,C−; secondary metabolites erythrin, lecanoric acid
and the unidentified substances 1 and 5.
Distribution and habitat (Fig. 2). Roccella applanata is
endemic to south and south-western parts of Madagascar.
It grows on shrubs and trees near the sea.
Remarks. Roccella applanata is similar to R. montagnei
but is easily distinguished by its broad flat branches often
as wide as 10–20 mm.
Roccella babingtonii Mont. (Figs 3, 4)
Ann. Sci. Nat. Bot. 18: 305 (1852). Type: according to
protolog ‘Hab. in Peruvia ad frutices’, label type collec-
tion says’Ex America meridionalis ut videtur: sed locus
specialis haud innotuit crescit in ligno’. s.ann., Churchill
Babington s.n. (PC, lectotype sel. here).
Roccella endocrocea Choisy, Ann. Cryptog. Exot.2:68
(1929). Type: Tanzania ‘Zanzibar, Afrique. sur les arbes’.
s.ann, Dumeny s.n. (LYJB-2322, holotype)
Fig. 2. Roccella applanata, white circles; R. babingtonii,black
circles; R. boryi squares; R. balfourii stars.
Fig. 3. Roccella babingtonii.Kenya,Kilifidistr.,MidaCreek,
2007, A. Tehler 9231 (S-F146175), DNA voucher specimen. Scale
mm.
Roccella intermedia Choisy, Ann. Cryptog. Exot.2:68
(1929). Type: Tanzania ‘Zanzibar, Afrique’. s.ann., s.coll.
s.n. (LYJB-2321, holotype)
Thallus pendent, branches flattened, foveate to smooth,
5–30 cm long, greyish to brown-greyish; cortex palisade
plectenchyma; epicortex absent; soralia absent; medulla
white or partly rusty yellow usually yellowish, also in hold-
fast zone. Apothecia present, sessile, deep yellow immedi-
ately under the hypothecium; ascospores 30 ×4µm(L.max
=33 µm; L.min =27 µm). Chemistry: spot tests, thallus
surface K−or if medulla is rusty yellow K+wine-red, C+
red; white parts of medulla K−,yellowpartsK+purple,
C−; secondary metabolites skyrin, erythrin, lecanoric acid,
and the unidentified substances 1 and 5.
Fig. 4. Roccella babingtonii. Cross-section of apothecium show-
ing the yellow medulla just below the black hypothecium (right),
also showing yellow medulla in branch (left). Kenya, Kilifi distr.,
Mida Creek, 2007, Tehler 9231 (S-F146177), DNA voucher spec-
imen. The sectioned apothecium, margin to margin, is 1mm.
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The Old World Roccella species 231
Distribution and habitat (Fig. 2). Roccella babingtonii
is known from the African coasts of Kenya, Tanzania,
Mozambique and from the islands Comores and Seychelles
in the Indian Ocean. It grows on shrubs and trees near the
sea often on mangrove.
Remarks. Roccella babingtonii is similar to fertile R. mon-
tagnei but is readily distinguished from that latter species
by its conspicuously deep yellow medulla. The deep yel-
low colour often breaks through the surface around the
apothecia and colours the apothecial margin orange yellow.
However, the medulla is not always yellow throughout the
thallus and often the medulla is simply white. Immediately
under the hypothecium the medulla is always deep yellow
(Fig. 4).
Roccella babingtonii is a unique species in many re-
spects. It is one of only two obligately corticolous Roc-
cella species and it is also obligately sexual; vegetative
reproduction by means of soredia or other propagules is
not known. The only other Roccella species possessing
those characteristics is the closely related species Roc-
cella applanata.Furthermore,Roccella babingtonii is the
only Roccella species and one of very few Roccellaceae
species and one of very few Arthoniales species to pro-
duce an anthraquinone, skyrin, as a secondary product,
a feature which has evolved as an autapomorphy for R.
babingtonii only. The crustose species Roccellina cinerea,
endemic to Socotra, is the only other species in Roccel-
laceae which has been found to produce skyrin. Skyrin is
responsible for the K+wine-red reaction of the yellow
medulla. The name Roccella babingtonii has been fre-
quently but incorrectly used for the sorediate taxon, R.
gracilis (Tehler, 2006), even though Montagne (1852) de-
scribed it as a fertile taxon with apothecia only, soredia
were not mentioned by him. A specimen found in PC
obviously belongs to the type material although the lo-
cality specifications slightly differ from that of the pro-
tolog. It has obviously been studied by Montagne since it
bears Montagne’s handwriting. A major problem with the
collection is the specimen and label data do not match.
The locality information ‘America meridionalis’ is with-
out any doubt incorrect since the specimen belongs to an
East African Roccella species usually called R. endocro-
cea and not to any New World species of Roccella.Ex-
cept for the type specimen there are no other records
in South America of Roccella babingtonii.Wesuggest
that label data has been confused and that the origin of
the specimen is from somewhere in East Africa. We re-
gard it as highly implausible that a single specimen of
this unique skyrin-producing species would have dispersed
all the way from the African east coast to the South
American west coast. According to the rules of priority
(Art. 11) the name Roccella babingtonii should be ap-
plied to East African species up until now referred to as R.
endocrocea.
Fig. 5. Roccella balfourii. Yemen. Socotra Island, Qalansiyah,
2008, A. Tehler 9350 (S-F146181). Scale mm.
Roccella balfourii M¨
ull. Arg. (Fig. 5)
Proc. Roy. Soc. Edinburgh 11: 458 (1882). Type: Yemen,
Socotra. 1881, Balfour 1338 (G, lectotype sel. here).
Roccella hertelii Mies & Schultz, Biblioth. Lichenol.88:
446. Type: Yemen, Socotra Island, Haggier Mountains,
Aduno ridge, cliffs in the east, on granite blocks. 1200
m; expos SW. 1996, Mies 499,1 (M-90344, holotype)
Thallus erect or pendent, branches terete rarely somewhat
flattened with a chalky surface, 5–10 cm long, white grey-
ish, smooth or foveate; cortex palisade plectenchyma; epi-
cortex absent; soralia absent; medulla compact or byssoid,
white or with central parts dirty white; in holdfast usually
brown. Apothecia present, sessile; ascospores 26 ×7µm
(L.max =33 µm, L.min =22 µm). Chemistry: spot tests,
thallus surface K+reddish in branch tips, C+red; medulla
K−,C−;secondarymetabolitesroccellicacid,erythrin,
lecanoric acid and the unidentified substances 1 and 5.
Distribution and habitat (Fig. 2). Roccella balfourii is
endemic to Socotra and Africa’s Horn in Somalia. It grows
on cliffs and rocks near the sea.
Remarks. Small specimens of Roccella balfourii are con-
fusingly similar to R. minuta but can be readily distin-
guished by the completely brownish medulla in the hold-
fast. Roccella minuta has pale yellowish medullary hyphae
in the immediate area next to the substrate. After examin-
ing the holotype of Roccella hertelii we conclude that this
collection is most probably specimens from poorly grown,
off habitat, R. balfourii.
Roccella boryi Delise ex F´
ee (Fig. 6)
Essai Crypt. ´
Ecorc. CI. (1824). Type: Ile de Bourbon
[R´
eunion, France]. s.ann., s.coll.s.n.(PC,lectotypesel.
Teh l e r & Ir e s te d t ( 20 0 7 )) .
Roccella flaccida Bory, Dict. Class. Hist. Nat. 14: 631
(1828).—Roccella flaccida Delise ex Darb., Biblioth.
Bot. 9: 44. (1898) nom.hom. Type: Del’ile de France
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232 A. Tehler et al.
Fig. 6. Roccella boryi. Mauritius. Savanne Distr., Maconde, 2003,
A. Tehler 8509 (S-L55020). Scale mm.
[Mauritius]. 1826, Jussieu 2444 (PC, lectotype sel. Tehler
& Irestedt (2007); isolectotypes PC).
Roccella montagnei f. teretior Cromb., J. Linn. Soc.,Bot.
15: 433 (1877). Type: Mauritius. Island of Rodriguez. On
rocks; infertile. 1874, Balfour s.n. (BM-674747, lectotype
sel. Tehler & Irestedt (2007); isolectotypes BM-674746,
H-NYL36728, H-NYL36735).
Thallus pendent or erect, branches approximately terete
but often also flattened, smooth or foveate, 5–15 cm long,
white greyish; cortex palisade plectenchyma; epicortex ab-
sent; soralia present but rarely seen together with apothe-
cia; medulla byssoid; in holdfast usually brown. Apothe-
cia present, sessile, rarely seen together with soralia;
ascospores 21 ×5.5 µm(L.max=24 µm; L.min =16 µm).
Chemistry: spot tests, thallus surface C+red, K+pinkish
red in branch tips; soralia C–; secondary metabolites roc-
cellic acid (present in sorediate specimens only), erythrin,
lecanoric acid and the unidentified substances 1 and 5.
Distribution and habitat (Fig. 2). Roccella boryi is a saxi-
colous species endemic to the Mascarene Islands (Mauri-
tius, R´
eunion and Rodriguez). The fertile morph is much
more common than its sorediate morph.
Remarks. Roccella boryi is most similar to R. montagnei,
however, the two species are not sympatric. Roccella boryi
is best distinguished from R. montagnei by its more terete
thallus branches and its shorter ascospores.
Roccella minuta Teh le r, s p . nov. ( Fi g. 7 )
Ty p e : Ye m e n . S o c o t r a I s l a n d , H o m i l l n e a r t h e v i l l a g e , i n
the eastern part of the island, on the ridge facing the sea
towards the east, alt. 350 m., 22 March 2008, Anders Tehler
9307 (S-F146188, holotype)
Thallus fruticosus, erectus, albido-cinereus; rami sparsi,
teretes, 5–10 cm longi. Cortex hyphis verticalibus. Asco-
carpia numerosa, lateralia vel apicalia. Soralia non visa.
Medulla albida autem basaliter pallidus-lutea prope sub-
stratum. Thallus et ascocarpia C+rubra et K non reagens.
Thallus erect, branches terete, smooth, 1–5 cm long,
white to white greyish; cortex palisade plectenchyma; epi-
cortex absent; soralia absent; medulla byssoid, white, in
the holdfast zone in the contact area to the substrate pale
light yellow. Apothecia present, sessile usually lateral but
also apical; ascospores 25 ×7(L.max=30 µmL.min=
22 µm). Chemistry: spot tests, thallus surface K+slightly
Fig. 7. Roccella minuta.Holotype,Yemen.SocotraIsland,Homill,2008,A.Tehler9307(S-F146188).Scalemm.
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The Old World Roccella species 233
Fig. 8. Distribution of Roccella minuta.
yellowish, C+red; medulla K−(incl. yellow hyphae in
holdfast), C−;secondarymetaboliteserythrin,lecanoric
acid and the unidentified substances 1, 2 and 5.
Etymology.Thespecificnameminuta refers to the small
growth of the thallus.
Distribution and habitat (Fig. 8). Roccella minuta is only
known from Socotra and from Cape Guardafui in Somalia,
Bari Region. It grows on cliffs and rocks near the sea.
Remarks. Roccella minuta is distinguished from R. bal-
fourii by its smaller often minute thallus and its char-
acteristic light yellow medullary hyphae in the lower-
most part of the holdfast in immediate contact with the
substrate.
Roccella montagnei B´
el. (Figs 9, 10)
Vo y. I n d e s O r. 2 C r y p t o g .: 117 (1838). India. Pondicherry,
ad Mangiferas indicas.1838,Belanger s.n.(PC, lectotype
sel. Awasthi (1981); isolectotypes B, BM-660964, PC).
Fig. 9. Roccella montagnei sorediate specimen. Sri Lanka, Ham-
bantota distr., Mawella, 1975, A. Tehler 43a (S-L9731) Scale mm.
Fig. 10. Roccella montagnei fertile specimen. Kenya. Coast prov.,
Kilifi distr., Gedi, 2007, A. Tehler 9239 (S-F146194), DNA
voucher specimen. Scale mm.
Roccella fuciformis ßlinearis Ach., Lichenogr. Univer-
salis 440 (1810).—Roccella mauritiana Darb., Ber.
Deutsch. Bot. Ges. 16: 11 (1898).—Roccella linearis
(Ach.) Vainio, in Hiern et al.,Cat. Afr. Pl. 2(2): 434
(1901).—Nemaria fuciformis var. linearis (Ach.) Nav´
as,
Broteria, Ser. Bot. 11: 32 (1913). Type: Sumatra,
s.ann., Gr¨
ondahl s.n. (S-L52524, lectotype sel. here;
isolectotypes H-ACH, H-NYL1329, UPS).
Roccella montagnei f. angustior Nyl., Syn. Meth. Lich.1:
261 (1860). Type: Indonesia, Java. 1856, Zollinger 2716
(PC, lectotype sel here; isolectotypes PC, UPS).
Roccella tinctoria f. minor Krempelh., Ve r h . Z o o l . - B o t .
Ges. Wien 18: 311 (1868). Type: Mexico, Los Ba ˜
nos,
an B¨
aumen. s.ann., Heller 155 (PC, lectotype sel. Tehler
(2006); isolectotype G).
Roccella montagnei v. rigidula M¨
ull. Arg., Flora 62: 294
(1880). Type: Angola, Chintoxo, Afr. occid., in trunco
Adansoniae. s.ann., Pech ue l - L oe s c h e s.n. (BM-660965,
lectotype sel here; isolectotype M-24562).
Roccella africana Vain., in Hiern et al.,Cat. Afr. Pl.
2(2): 432 (1901). Type: Angola. Benguella. Habit. inter
Roccellam ad portum: Novo Rodondo. 1857, Lopes de
Silva 405 (BM-661017, lectotype sel. here; isolectotypes
TUR-V-26953, B-103554, B-20983, B).
Roccella podocarpa Vain., in Hiern et al.,Cat. Afr. Pl.
2(2): 435 (1901). Type: Angola. Ambriz ‘Ad Adanso-
niam digit. retro Mubango in interioribus de Ambriz.
1854, We l w i t s c h 407 (BM-660962, lectotype sel. here;
isolectotypes B, BM-660961, TUR-V-26951; syntypes
B-57086, BM-660960, TUR-V-26952).
Roccella linearis var. guineensis Vain., in Hiern et al.,
Cat. Afr. Pl. 2(2): 434 (1901).—Roccella montagnei var.
guineensis (Vain.) v. d. Byl, Ann. Univ. Stellenbosch,
Reeks A,Wis.- Na turk 11(6): 3 (1933). Type: Guinea.
Insula S. Thom´
einsinuGuinea.Adrupesmaritimasin-
sulae S. Thomae sin. Guineensis. 1859, We l w i t s c h 421
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234 A. Tehler et al.
Fig. 11. Roccella montagnei, generalized distribution, black; R. sinensis stars. Localities in Indonesia are unspecified (Sumatra, Java and
Bali).
(BM-660971, lectotype sel. here; isolectotypes B, TUR-
V- 2 6 9 4 6 ) .
Roccella linearis var. hyperchromatica Vain., in Hiern et
al.,Cat. Afr. Pl. 2(2): 434 (1901). Type: Angola. Luanda.
Ad rupes maritimas, non procul a Morro de Sta Catha-
rina ast non frequens. 1854, We l w i t s c h 422 (BM-660970,
lectotype sel. here; syntypes B, TUR-V 26938).
Roccella linearis var. primaria Vain., in Hiern et al.,Cat.
Afr. Pl. 2(2): 434 (1901). Type: Angola. Luanda Distr.
Frequentiss. ad ramos Adansoniae,Euphorbiae Candel.
aliarumque arborum inter Loanda & Quicuje. 1854, We l -
witsch 413 (BM-660972, lectotype sel. here; syntypes
B, BM-660948, BM-660975, BM-660976, BM-660977,
BM-660978, BM-660979, BM-660980, TUR-V-26944,
TUR-V-26945).
Roccella montagnei f. obtusa Vain., in Hiern et al.,Cat.
Afr. Pl. 2(2): 435 (1901). Type: Angola. Ambriz distr. Ad
Acacias et Anacardias territ. Ambrizensis versus =terras
de Mossul. 1853, We l w i t s c h 408 (BM-661010, lectotype
sel. here; isolectotype B).
Roccella tinctoria f. complanata Vain., in Hiern et al.,Cat.
Afr. Pl. 2(2): 432 (1901).—Roccella subpodicellata f.
complanata (Vain.) Follmann & Huneck, J. Hattori Bot.
Lab. 32: 39 (1969). Type: Angola: Namibe ‘Mossamedes.
Ad rupis Oceani Atlantici prope Cazimba (via versus
Cabo Negro). 1859, We l w i t s c h 406 (BM 660956, lecto-
type sel here; isolectotypes TUR-V 26948, B).
Roccella pattensis Dodge, Beih. Nova Hedwigia 12: 88
(1964). Type: ‘Kenya. Lamu District. Patta Island, Tan-
gawanda cortiole, Greenway & Rawlings 8926 (not
found, not seen). Paratype: Kenya–Somalia border, Li-
boi 600’ epiphyte on Commiphora in tree-savanna. 1954,
Bally B9491 (B-128663).
Roccella pembensis Dodge, Beih. Nova Hedwigia 12: 89
(1964). Type: Tanzania. Pemba Island, Jundoni coast.
1901, Lyn e 116 (BM 660963, lectotype sel. here; isolec-
totypes B, BM).
Roccella belangeriana Awasthi, Bryologist 84: 219 (1981).
Ty p e : I n d i a , P o n d i c h e r r y , P i c h a v a r a m , p r o p r o o t s o f Rhi-
zophora,20June1979,Thanikaimoni 1524 p.p. (sepa-
rated as 1524B) (hb Awasthi holotype, unable to receive
on loan); isotype Inst. Franc¸ais, Pondicherry, unable to
receive on loan).
Thallus pendent or erect, branches flattened, strap like,
smooth, foveate or wrinkled, 5–15 cm long, greyish; cor-
tex palisade plectenchyma; epicortex absent; soralia present
but rarely seen together with apothecia; medulla byssoid,
white; in holdfast usually brown. Apothecia present, ses-
sile, rarely seen together with soralia; ascospores 28 ×4
µm(L.max=33 µm, L.min =24 µm). Chemistry: spot
tests, thallus surface K−or faintly pinkish red in branch
tips, C+red; medulla K−,C−;secondarymetabolitesroc-
cellic acid (in sorediate specimens), erythrin, lecanoric acid
and the unidentified substances 1 and 5.
Distribution and habitat (Fig. 11). Roccella montagnei
is one of the most common and widespread of all Roc-
cella species. We have seen collections from New Caledo-
nia, Northern Australia, Philippines over Indonesia (Java,
Sumatra) all around the coasts of the Indian Ocean includ-
ing many of its islands, continuing around the Cape of
Good Hope in South Africa and north along the Atlantic
coast up to the Democratic Republic of the Congo and S˜
ao
To m ´
ewithoutpostlocalitiesasfarnorthasTogo,Cape
Ver d e .
Remarks. Roccella montagnei has been considered the
sorediate counterpart in a so-called species pair relation-
ship with the fertile counterpart Roccella belangeriana
(Awasthi, 1981). The present results do not support that
hypothesis (see Discussion below) and the two taxa are
here considered conspecific.
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The Old World Roccella species 235
Roccella africana which is represented only by its type
specimen differs from the common habitus of R. montag-
nei by their approximately terete thallus branches. However,
their thallus surface is conspicuously C+red which is char-
acteristic for R. montagnei.
We have s e e n five ve r y ol d s pec i men s o f Roccella mon-
tagnei said to be collected from Chile (G), Peru (G, PC),
Mexico (G, PC), and California, USA (G) but these are ob-
viously mislabelled. Label data are also questioned for an-
other two very old collections recorded from New Zealand
and the south coast of Tasmania (BM). A specimen col-
lected by Gaudichaud in Montevideo (PC) is also dubi-
ous since it is the single record from anywhere along the
South American Atlantic coast and it will be considered
mislabelled until Roccella montagnei can be confirmed by
modern collections from that region.
As found from our TLC-investigation roccellic acid is
always present in sterile, sorediate specimens. Fertile spec-
imens, with few exceptions, lack roccellic acid.
Roccella phycopsioides Tehl e r , s p. nov. (Fi g . 12)
Ty p e : Ye m e n . S o c o t r a I s l a n d , H o m i l l n e a r t h e v i l l a g e , i n
the eastern part of the island, on the ridge facing the sea
towards the east, alt. 400–450 m, 22 March 2008, Anders
Tehl e r 9308 (S-F146246, holotype).
Thallus fruticosus, erectus vel pendulus, albido-cinereus;
rami teretes, 5–10 cm longi. Cortex hyphis verticalibus. As-
cocarpia non visa. Soralia numerosa, lateralia. Medulla al-
bida autem basaliter pallidus-lutea prope substratum. Thal-
lus C+rubra et K non reagens; soralia C- reagens. Thallus
acidum roccellicum continens.
Fig. 12. Roccella phycopsioides, note yellowish medulla in hold-
fast. Holotype, Yemen, Socotra Island, Homill, 2008, A. Tehler
9308 (S-F146246), DNA voucher specimen. Scale mm.
Fig. 13. Distribution of Roccella phycopsioides.
Thallus erect or pendent, branches terete rarely some-
what flattened, 5–10 cm long, greyish-greenish, smooth or
foveate; cortex palisade plectenchyma; epicortex absent;
soralia present; medulla compact or byssoid, white or with
central parts dirty white; in holdfast with yellowish medulla.
Apothecia absent. Chemistry: spot tests, thallus surface K+
reddish in branch tips, C+red; medulla K−,C−;secondary
metabolites roccellic acid, erythrin, lecanoric acid and the
unidentified substances 1 and 5.
Etymology.Thespecificnamephycopsioides refers to its
similarity with Roccella phycopsis sharing the features with
ayellowmedullaintheholdfastandC−soredia.
Distribution and habitat (Fig. 13). Roccella phycopsioides
is endemic to Socotra. It grows on cliffs and rocks near the
sea. It may also be corticolous.
Remarks. Roccella phycopsioides is most easily confused
with sorediate specimens of R. montagnei but can be read-
ily distinguished by the yellowish medulla in the hold-
fast. Roccella minuta, which also has a yellowish medulla
in the holdfast, is best distinguished by the absence of
soredia.
Roccella sinensis Nyl. (Fig. 14)
Syn. Meth. Lich.1:261(1860).Type:Chine.Cantonet
Macao. 1837, Gaudichaud 33 (PC, lectotype sel. in Tehler
& Irestedt (2007)).
Thallus erect or pendent, branches terete, smooth to
foveate, 5–10 cm long, white greyish; cortex palisade
plectenchyma; epicortex absent; soralia absent; medulla
compact to byssoid, white; in holdfast white or some-
times brownish. Apothecia present, sessile; ascospores
21 ×5.5 µm(L.max=23 µm; L.min =18 µm). Chem-
istry: spot tests, thallus surface K−,C+red sometimes
reaction seen only in upper branches; medulla K−,C−;sec-
ondary metabolites erythrin, lecanoric acid and the uniden-
tified substances 1 and 5.
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236 A. Tehler et al.
Fig. 14. Roccella sinensis. Hong Kong, Kowloon District, Clear
Water Bay, 2004, A. Tehler 8565 (S-L60293). Scale mm.
Distribution and habitat (Fig. 11). Roccella sinensis is
only known from the southern coast of China, Hong Kong,
Macau, Hainan and northern Vietnam in the bay of Tongk-
ing.
Remarks. Roccella sinensis is one of few Roccella species
that is only found fertile; soredia are not present.
Roccella tinctoria DC.
Fl. Franc¸. ed.3,2:334.(1805).–Lichen roccella L., Sp. Pl.:
1154. (1753). – Parmelia roccella (L.) Ach., Methodus:
274 (1803). – Thamnium roccella (L.) St. Hilaire, Expos.
Fam . N a t . 1: 21 (1805). – Roccella purpura-antiquorum
Bory, Dict. Class. Hist. Nat. 14: 630 (1828). – Roccella
tinctoria f. filiformis C. Bab., Primit. Fl. Sarnic.:123
(1839). – Roccella arnoldii Vain., in Hiern et al.,Cat. Afr.
Pl. (2): 433 (1901). – Nemaria roccella (L.) Nav´
as, Bro-
teria 8: 4 (1909). – Roccella fucoides var. arnoldii (Vain.)
Zahlbr., Catal. Lich. Univ. 2: 519 (1923). – Type: Roc-
cella ex insula Fyal, s.ann., s.coll. s.n. (LINN 1273.263,
lower specimen lectotype; upper specimen isolecto-
type), [lectotype designated by Howe (1912); lectotype
designation narrowed (Art. 9.14) by Jørgensen et al.
(1994)].
Physcia dichotoma Pers., Ann. Wetterauischen Ges.
Gesammte Naturk., Frankfurt. 1(2): 18 (1810). – Roc-
cella tinctoria var. dichotoma (Pers.) Ach., Syn. Meth.
Lich.:337(1814).–Roccella dichotoma (Pers.) Ach.,
Syn. Meth. Lich.:337(1814),prosyn.–Type:(searched
for but not found in L). – Neotype: ‘Roccella dichotoma’
in herb Persoon, s.loc., s.ann., s.coll. s.n. (L-0246980
neotype designated by Tehler (2002); isoneotype L-
0246981).
Roccella fastigiata Bory, Dict. Class. Hist. Nat.14:631
(1828). – Roccella tinctoria var. fastigiata (Bory) Delise
ex Darbishire, Biblioth. Bot.45:32(1898a),prosyn.et
comb. nov. inval. – Coll. orig.: [France] Ouessant, Pylaie;
cˆ
otes de Saint-Malo, s.ann. Br´
ebisson s.n. (searched for
but not found in PC). – Neotype: Canary Islands, Tener-
ife, 1822, Delise s.n. (PC designated by Tehler (2002)).
Roccella patellata Stirton, J. Linn. Soc. Bot.14:366
(1875). – Type: [Cape Verde Is.], ‘ad saxa, Bird Is-
land’, 1843, Stirton s.n. (BM-660991, lectotype selected
by Tehler (2002); isolectotypes B, BM-660954, BM-
660955, GLAM-STIRTON).
Roccella canariensis Darb., Ber. Deutsch. Bot. Ges.15:5
(1897) nom.nud.; Ber. Deutsch. Bot. Ges.16:11(1898b)
nom. nud.; Biblioth. Bot. 45: 50 (1898a) nom. valid.
–Roccella tinctoria f. valida Hampe ex Darbishire, Ber.
Deutsch. Bot. Ges.15:5(1897),pro.syn.;Ber. Deutsch.
Bot. Ges. 16: 11 (1898b) pro. syn.; Biblioth. Bot.45:
50 (1898a) pro. syn. – Type: The Canary Islands ‘Roc-
cella tinctoria forma valida,Ins.Canariens,exhbHampe
1877’. (G lectotype selected here; isolectotype G).
Roccella caribeae Darb., Ber. Deutsch. Bot. Ges.16:11
(1898) nom. nud; Biblioth. Bot. 9: 53, nom. valid. – Roc-
cella bellicosa Darb., Biblioth. Bot.9:53,prosyn.et
nom. nud. (1898). Type Venezuela. 1898, Schuchardt
s.n. (HBG, lectotype sel Tehler & Irestedt (2007); isolec-
totypes B-103557, B-128642, B, G).
Roccella tinctoria var. subpodicellata Vain., in Hiern et al.,
Cat. Afr. Pl.2(2):431(1901).–Roccella subpodicellata
(Vain.) Follm. & Huneck, J. Hattori Bot. Lab .32:39
(1969). Type: Angola: Namibe ‘Mossamedes. Ad rupis
Oceani Atlarici prope Cazimba (via versus Cabo Negro).
1859, We l w i t s c h ,406(BM-660956lectotypeselhere;
isolectotypes TUR-V 26947, B-57101, B).
Roccella tuberculata var. vincentina Vain., in Hiern et
al.,Cat. Afr. Pl. (2): 433 (1901). – Roccella canarien-
sis var. vincentina (Vain.) Zahlbr., Deutsch. S¨
udpolar-
Exp.1901-1903.8:21(1906).–Roccella vincentina
(Vain.) Follmann in Follmann & S´
anchez-Pinto, Courier
For s ch un g si n st . S e nc k e nb e rg 95: 184 (1987). [ut ‘vi-
centina’] – Type: Cape Verde Is., ‘ad rupes maritimas
insulae S. Vicentii, Cap. Virid.’, January 1861, We l w i t s c h
402 (BM-660953 lower left specimen, lectotype selected
by Tehler (2002); isolectotypes B, BM-660953, TUR-V-
26958).
Roccella hypomecha var. isabellina Vain., in Hiern et
al.,Cat. Afr. Pl.(2):432(1901).–Type:Angola.
Mossamedes. Ad rupes maritimas inter Mossamedes et
Porto Pinda loco dicto Cazimba, parce; Ad rupes mar-
itimas. 1859-09, We l w i t s c h 404. (BM-660982, lectotype
selected by Tehler (2004); isolectotypes B-20901, TUR-
V- 2 6 9 5 5 ) .
Roccella boergesenii Vain i o , Kongel. Danske Vidensk.
Selsk. Skr.,Naturvidensk. Math. Afd.8:397(1924).–
Ty p e : C a n a r y I s l a n d s , Te n e r i f e , O r o t a v a ( Ta o r o ) , 1 9 2 1 ,
Børgesen s.n. (TUR-V-26919, lectotype selected by
Teh l e r (2 0 0 2) ) .
Roccella guanchica G. B. Feige & Viethen, Herzogia 7:
611 (1987). – Type: Canary Islands. Tenerife, ad saxa
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The Old World Roccella species 237
vulcanica in territorio ‘Malpais de G¨
uimar’, ad mare,
circ. 20 m.s.m., expositione septentriale usque ad occi-
dentale, 1985, Feig e & Viethen s.n. (ESS-3447, holotype;
isotype B).
Thallus erect to pendent, stiff, branches terete, foveate
or nearly smooth, brown-greyish-brown; cortex hyphae
slightly interlaced, anticlinally arranged but not forming
aconspicuouspalisadeplectenchyma;epicortexpresentas
asurfacegel;soraliapresentbutrarelyseentogetherwith
apothecia; medulla compact or byssoid, white; in holdfast
white but usually streaked with brown or black. Apothecia
present and usually numerous but rarely seen together with
soredia; ascospores 24 ×6µm(L.max=27 µm; L.min =
21 µm). Chemistry: spot tests, thallus surface C−rarely C+
red, K−; soralia C+red and cortex under epicortical layer
C+red; secondary metabolites erythrin (sometimes traces
or absent), lecanoric acid (Viethen & Feige 1986 by HPLC
and TLC, annotated voucher specimens [BM-660953, ESS-
3447, M-24629, M-24628, M-24638, M-24695]).
Distribution and habitat.Roccella tinctoria is one of the
most common and widely distributed species in Roccella.
It grows on the vertical side of cliffs and rocks near the sea.
Locally it may be very abundant not only in its central distri-
bution area of Macaronesia, the Canary Islands, the Azores,
Madeira and in Estremadura, Portugal but also in Cape
Ver d e . I t al s o ra n ge s i n t o t he we s te r n M e di t e r r a ne a n Re g io n
(Sardinia) and into the South Atlantic Region with Cape
Ver d e , S en e g a l a nd a s f a r s ou t h as B e n g ue l a in A n g o l a .
Remarks. Roccella tinctoria was emended by Tehler et al.
(2009a)toalsoincludeR. canariensis.Thebestknown
so-called ‘species pair’ in Roccella is the Macaronesian R.
canariensis/R. tinctoria which belongs to the significantly
supported R. phycopsis group (Tehler et al.,2004;Tehler&
Irestedt, 2007). The R. canariensis/R. tinctoria-pair is the
typical case in which a monophyletic, evolutionary lineage
contains both sexual and vegetative specimens intermin-
gled in the phylogenetic tree (Tehler et al.,2004).There
is neither support for a phylogenetic sexual species nor
aphylogeneticvegetative,sorediatespeciessincenoneof
the two reproductive modes form monophyletic groups.
Instead R. canariensis/R. tinctoria represents one polymor-
phic lineage regarding its reproductive strategy. On basis of
molecular evidence Roccella tinctoria (asexual) was united
with R. canariensis (sexual) into one single species with
the name R. tinctoria.
Occasionally, the thallus surface of Roccella tinctoria
yields a more or less C+red reaction. Specimens reacting
C+red are rarely seen within its core distribution area in
Macaronesia comprising Estremadura (Portugal), the Ca-
nary Islands, Madeira and the Azores, even though oc-
casionally such specimens have been collected there (see
below). In marginal areas and in outpost localities spec-
imens with C+red thallus surface are more common as
have for example been observed in a specimen originating
from Sardinia in the Mediterranean. More often specimens
growing south or far south of main Macaronesia are more
susceptible to a positive thallus surface chlorine reaction
i.e. in Cape Verde, Senegal and Angola. The populations in
Cape Verde and Senegal may react both C−and C+red in
different parts of the thallus. The upper part of the branches
more often react C+red while the lower parts react C−.
The variation in C reaction is seen particularly well in Roc-
cella tinctoria specimens from the southernmost locality in
Angola and from which two new subspecific taxa were de-
scribed by Vainio (1901). His type specimens, R. tinctoria
var. subpodicellata and R. hypomecha var. isabellina show
agradientCthallussurfacereactionfromcompletelyneg-
ative (best seen in R. hypomecha var. isabellina)topartly
negative (best seen in R. tinctoria var. subpodicellata)to
reacting weakly C+red after a few seconds (best seen in
R. tinctoria var. subpodicellata). These taxa are only repre-
sented by their type specimens collected on the Welwitsch
expedition to Angola 1859.
Why some populations, usually those from marginal dis-
tribution areas, react C+red is that the epicortex surface
gel layer is poorly developed or even missing and the chlo-
rine solution may then break through the epicortex and the
surface becomes C+red. A well-developed epicortex is
otherwise completely impenetrable to chorine solution.
Tw o s p e c i m e n s o f Roccella tinctoria that reacted C+
red from the Azores and Cape Verde were sequenced and
analysed in Tehler et al. (2004) and Tehler et al. (2009b)
respectively. Both specimens clustered with high support to-
gether with all other Roccella tinctoria specimens. On basis
of that and on the general morphological evidence we find
that all the four taxa described by Vainio from Angola above
should be synonymized with Roccella tinctoria.Unfortu-
nately, we had no fresh material of any Angolan Roccella
tinctoria and thus we have no sequence evidence to corrob-
orate our morphological hypothesis that this population is
conspecific with the Macaronesian main population.
Results
The studied loci and alignments
One hundred and sixty-three ingroup and outgroup se-
quences from four loci, RPB2, nuLSU rDNA, ITS rDNA
and an anonymous locus were used for this study. Newly
produced were: 122 sequences from the ITS 1 and 2 regions
including 5.8S rDNA; 118 sequences from the nuLSU,
120 sequences from the RPB2 and 124 sequences from
the anonymous locus; altogether 484 new sequences (Ta-
ble 1). We were unable to sequence the sample Roccella
fuciformis 8171 for the anonymous locus and we were un-
able to sequence two samples of R. montagnei 66650-109
and 66650-110 for LSU; they were included in the data
set as missing data. The aligned data set contained 163
terminals and 3216 aligned sites. The sequences obtained
from the anonymous locus were compared with calmod-
ulin sequences of Sclerotinia sclerotiorum and Ve r t i c i l l i u m
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238 A. Tehler et al.
dahliae published in GenBank by Carbone & Kohn (1999).
ABLASTsearchwasalsoexecuted,butnoobviousmatches
could be found. We thus consider this unidentified locus as
anonymous, although a potential relationship to the calmod-
ulin gene cannot be excluded (Tehler et al., 2009a). (See
also Materials and methods above.).
Take n in t o a cc o u n t th a t s ho r t fragments in the 5 &or 3&
ends of the studied loci are missing for some taxa (pri-
marily in the anonymous locus and ITS), the length of the
sequences and sequence divergences varied as follows. The
LSU loci varied in length between 886–898 bp and the
sequence divergence ranged between 0.0–6.0%; the ITS1
and ITS2 region varied in length between 576–636 bp and
the sequence divergence ranged between 0.0–25.4%; the
anonymous locus varied in length between 409–466 bp and
the sequence divergence ranged between 0.0–38.7%; and
the sequence divergence for the 923 bp long region from
the coding RPB2 gene ranged between 0.0–14.5%. The con-
catenated alignment consists of 3163 bp. In the final align-
ments of the ITS and the anonymous locus some regions are
still ambiguous. However, if only the Old World Roccella
species (the primary study group in this paper) are consid-
ered in these alignments very few positions are ambiguous
in the anonymous locus, and also in the ITS region the
number of ambiguous positions is much reduced. Analyses
of the ITS region before and after the manual adjustments
also produced trees that were topologically fully congru-
ent (only nodal support values were marginally changed),
suggesting that modest adjustments in ambiguous regions
in ITS only have minor effects on the topological results.
Similarly, Tehler et al. (2009a)foundthatanalysesofalter-
native ITS alignments of New World Roccella species had
minor effects on the topological results. In addition, the
independent gene trees were found to be very similar with
no significant topological conflicts (see Results). Based on
these observations we found it warranted to not exclude any
positions for the final analyses of the concatenated data set,
as the most variable regions may be potentially important
for resolving particularly terminal branches in the tree.
Phylogenetic analyses
There were no significant conflicts between the differ-
ent gene tree topologies. The eight Old World Roccella
outside Europe and Macaronesia were non-monophyletic
(Figs 15–18). The largest clade contain four recognized
species in a significantly supported group, here called
‘montagnei-group’ containing the species Roccella ap-
planata,R. babingtonii,R. boryi and R. montagnei (these
names includes synonyms such as R. africana,R. belan-
geriana,R. endocrocea,R. flaccida,R. intermedia,R. lin-
earis,R. mauritiana,R. pattensis,R. pembensis and R.
podocarpa).
The PJ tree and the Bayes tree were mutually without con-
flict. In the PJ tree the ‘montagnei-group’ formed part of an
unresolved trichotomy but in the Bayes tree the ‘montagnei-
group’ was resolved with full PP support: (a) Roccella
balfourii/R. phycopsioides;(b)R. minuta/R.sinensis and R.
fuciformis/R. maderensis (the two latter treated in Tehler
et al.,2004);(c)themajorgroupincludingallAmerican
species and the rest of the Macaronesian species (Tehler
et al., 2004; Tehler et al.,2009a,2009b). The group b)
above were included together with the ‘montagnei-group’
(a) above into a major but insignificantly supported clade
(Figs 15, 16).
The PJ tree retaining only significantly supported groups
begins with a polytomy consisting of seven clades (Fig.
18): two major groups of which one includes four of the
presently studied Old World species, Roccella applanata/R.
babingtonii/R. boryi/R. montagnei; two sister pairs of which
one was the Roccella balfourii/R. phycopsioides pair from
the present study and; three single species, two of which
are the Old World species, R. minuta and R. sinensis.
The only noteworthy topological conflict between the
parsimony trees (Fig. 17) and the Bayes tree is the position
of Roccella allorgei which is placed as the sister taxon to
the rest of the Roccella species in the parsimony trees, but in
the Bayes tree it is placed as sister taxon to the sub group R.
minuta/R. sinensis/R. fuciformis/R. maderensis four nodes
higher up the tree. Neither of the two hypotheses is confi-
dently supported.
The widespread and genetically diverse species Roccella
montagnei is retained as one species on basis of its mor-
phological, anatomical and chemical homogeneity.
Seven of the species, R. balfourii,R. phycopsioides,R.
minuta (Socotra), R. sinensis (Hong Kong and Tonkin), R.
babingtonii (East Africa), R. applanata (Madagascar) and
R. boryi (Mascarene Islands) have relatively restricted dis-
tribution areas and can be considered endemic to their par-
ticular areas. One species, Roccella montagnei,hasavery
large distribution area ranging from Australia, Indonesia
and all around the Indian Ocean continuing into the At-
lantic Ocean north to the Cape Verde.
Morphology
Morphological characteristics of the Old World species are
mainly seen in the shape and size of the thallus branches,
ascospore size, medulla colour, and hypomedulla colour in
the holdfast.
Roccella montagnei is the most common of the Old World
species and it has the widest geographical distribution range
of all species in Roccella.Morphologically,anatomically
and chemically it is very homogeneous featuring usually
strap-shaped thallus branches and numerous apothecia or
soralia. Ascospore size varies only a few µminlength.
Chemically it contains erythrin, lecanoric acid and roccel-
lic acid and a few unidentified substances. Specimens and
populations of Roccella montagnei are morphologically and
chemically indistinguishable.
Downloaded By: [Tehler, Anders] At: 13:31 16 August 2010

The Old World Roccella species 239
1
2
3
0.1
R. montagnei 8865 sor In
R. montagnei 8870 ap In
R. montagnei 8864 sor In
R. montagnei 8869 sor In
R. montagnei 8863 ap In
R. montagnei 8879 sor In
R. montagnei 8878 ap In
R. montagnei 9248 sor Ke
R. montagnei 9352a sor So
R. montagnei 9337 sor So
R. montagnei 9332 sor So
R. montagnei 9309b-13 sor So
R. montagnei 9227 sor Ke
R. montagnei 9257 sor Ke
R. montagnei 9263 sor Ke
R. montagnei 9225 sor Ke
R. montagnei 9244 sor Ke
R. montagnei 8868 sor In
R. montagnei 9246 sor Ke
R. montagnei 9253 sor Ke
R. montagnei 9254 sor Ke
R. montagnei 9255 apsor Ke
R. montagnei 9241 sor Ke
R. montagnei 9250 sor Ke
R. montagnei 9221 sor Ke
R. montagnei 9325 ap So
R. montagnei 9260 sor Ke
R. montagnei 9247 sor Ke
R. montagnei 9245 sor Ke
R. montagnei 9235-14 ster Ke
R. montagnei 9222 ster Ke
R. montagnei 9232-7 sor Ke
R. montagnei 9232-8 sor Ke
R. montagnei 13214 sor Mdg
.57
R. montagnei 13069 ap Mdg
.57
R. montagnei 13074 sor Mdg
.55
R. montagnei 7709 sor SAfr
R. montagnei Feur sor SAfr
R. montagneiagnei 13219 sor Mdg
R. montagnei 9236 sor Ke
R. montagnei Apt 66650-109 sor He
R. montagnei Apt 66650-110 sor He
R. montagnei 9235-11 ster Ke
R. montagnei 13213 sor Mdg
R. montagnei 9237 apsor Ke
R. montagnei 13112 ap Mdg
R. montagnei 13054 sor Mdg
1
R. montagnei 13111 ap Mdg
.90
R. montagnei 9701 sor Ang
.92
1
R. montagnei 13020 ap Mdg
R. montagnei 13036 ap Mdg
.98
R. montagnei 9708 sor Ang
R. montagnei 9729 sor Ang
R. montagnei 9718 sor Ang
R. montagnei 9702 sor Ang
R. montagnei 9743 sor Ang
R. montagnei 9724 sor Ang
R. montagnei 9723 12 sor Ang
R. montagnei 9747 ster Ang
R. montagnei Wirth KR1 sor Nami
R. montagnei Wirth KR2 sor Nami
R. montagnei 9723-11 sor Ang
R. montagnei 9740 sor Ang
R. montagnei 9735 sor Ang
R. montagnei Aptr 66616 sor He
R. montagnei Aptr 66253 sor He
R. montagnei Aptr 66251 ap He
R. montagnei Aptr 66613 ap He
R. montagnei 9714 ap Ang
R. montagnei 9719 ap Ang
R. montagnei 9700 ap Ang
R. montagnei 9728 ap Ang
1
R. montagnei 9239-16 ap Ke
R. montagnei 9259 ap Ke
1
1
R. montagnei 9252 ap Ke
R. montagnei 9262 ap Ke
R. montagnei 9239-15 ap Ke
R. montagnei 9249 ap Ke
1
R. boryi 8500-1 ap
R. boryi 8500-2 ap
R. boryi 8506 sor
R. boryi 8527 sor
R. boryi 8507 ap
R. boryi 8538 ap
R. boryi 8526 ap
R. boryi 8537 sor
R. boryi 8532-29 sor
R. boryi 8533-31 ap
R. boryi 8533-30 ap
R. boryi 8518-18 ap
R. boryi 8532-28 sor
R. boryi 8501 ap
R. boryi 8516 ster
1
R. applanata Erik101 Mdg
R. applanata 13104 Mdg
R. applanata 13075 Mdg
R. applanata 13060 Mdg
R. applanata 13058 Mdg
1
1
R. babingtonii 9231
R. babingtonii 9258
R. babingtonii 9233
R. babingtonii 9251-28
R. babingtonii 9242-18
R. babingtonii 9234
R. babingtonii 9242-19
R. babingtonii 9251-27
1
1
1
R. minuta 9307 9
R. minuta 9336
R. minuta 9351
R. minuta 9307-8
R. minuta 9334
R. minuta 9303
1
R. sinensis 8562-1
R. sinensis 8564
R. sinensis 8562-2
R. sinensis 8571
1
.85
R. fuciformis 8255
R. fuciformis 8171
.99
R. maderensis 8140
R. maderensis 8157
1
1
1
R. allorgei 8151
R. allorgei 8163
1
.95
R. balfourii 9306-6
R. balfourii 9326
R. balfourii 9302
R. balfourii 9306-7
R. balfourii 9349
R. balfourii 9350
R. balfourii 9335
1
R. phycopsioides 9308
R. phycopsioides 9331
R. phycopsioides 9309a-12
R. phycopsioides 9352b
1
1
.74
.66
R. galapagoensis 8651-12
R. galapagoensis 8667
1
R. albida 8653-14
R. albida 8780
1
.85
R. nigerrima 8655
R. nigerrima 8666
1
R. margaritifera 8656
R. margaritifera 8705
1
1
1
R. lirellina 8880
R. lirellina 8904
1
1
R. gracilis 8654
R. gracilis 9190
1
R. decipiens sax 9080
R. decipiens sax 9148
1
1
R. bajasurensis 9188
R. bajasurensis 9192
1
1
R. verruculosa 8941-32
R. verruculosa 8950-41
1
.92
1
R. tinctoria 8257
R. tinctoria L1784a
R. tuberculata 8144
R. tuberculata 8310-50
1
.97
R. elisabethae 8272
R. elisabethae 8306-45
1
1
R. phycopsis 8295
R. phycopsis 8221
1
1
1
1
D. ceratoniae 9021
D. ceratoniae 9047
1
1
D. paradoxa ssp. africana 8524
D. catalinariae 9087
1
1
1
1
1
1
1
1
.91
Fig. 15. Tree from Bayesian analysis showing species relationships in the genus Roccella. Species in coloured squares are the Old World
species outside Macaronesia recognized in the present study. PP-values on major nodes only. sor =sorediate, ap =apothecia, apsor =
both apothecia and soredia, ster =neither soredia nor apothecia. Ang =Angola, HK =Hong Kong, In =India, Ke =Kenya, Mdg =
Madagascar, Mau =Mauritius, Na =Namibia, He =Saint Helena, Soc =Socotra, SAfr =South Africa.
Downloaded By: [Tehler, Anders] At: 13:31 16 August 2010

240 A. Tehler et al.
D. ceratoniae 9047
D. ceratoniae 9021
D. paradoxa ssp. africana 8524
D. c catalinariae 9087
R. allorgei 8163
R. allorgei 8151
R. phycopsis 8295
R. phycopsis 8221
R. elisabethae 8306-45
R. elisabethae 8272
R. tuberculata 8310-50
R. tuberculata 8144
R. tinctoria L1784a
R. tinctoria 8257
R. verruculosa 8950-41
R. verruculosa 8941-32
R. bajasurensis 9192
R. bajasurensis 9188
R. gracilis 9190
R. gracilis 8654
R. decipiens 9148
R. decipiens 9080
R. lirellina 8904
R. lirellina 8880
R. nigerrima 8666
R. nigerrima 8655
R. margaritifera 8705
R. margaritifera 8656
R. galapagoensis 8667
R. galapagoensis 8651-12
R. albida 8780
R. albida 8653-14
R. maderensis 8157
R. maderensis 8140
R. fuciformis 8255
R. fuciformis 8171
R. sinensis 8571 HK
R. sinensis 8564 HK
R. sinensis 8562-2 HK
R. sinensis 8562-1 HK
R. minuta 9303 Soc
R. minuta 9334 Soc
R. minuta 9307-8 Soc
R. minuta 9351 Soc
R. minuta 9336 Soc
R. minuta 9307-9 Soc
R. phycopsioides 9352b Soc
R. phycopsioides 9331 Soc
R. phycopsioides 9309a-12 Soc
R. phycopsioides 9308 Soc
R. balfourii 9350 Soc
R. balfourii 9335 Soc
R. balfourii 9349 Soc
R. balfourii 9326 Soc
R. balfourii 9306-7 Soc
R. balfourii 9306-6 Soc
R. balfourii 9302 Soc
R. babingtonii 9242-19 Ke
R. babingtonii 9251-27 Ke
R. babingtonii 9251-28 Ke
R. babingtonii 9242-18 Ke
R. babingtonii 9234 Ke
R. babingtonii 9233 Ke
R. babingtonii 9258 Ke
R. babingtonii 9231 Ke
R. applanata 13058 Mdg
R. applanata Erik101 Mdg
R. applanata 13104 Mdg
R. applanata 13075 Mdg
R. applanata 13060 Mdg
R. boryi 8532-28 sor Mau
R. boryi 8518-18 ap Mau
R. boryi 8516 ster Mau
R. boryi 8501 ap Mau
R. boryi 8533-31 ap Mau
R. boryi 8533-30 ap Mau
R. boryi 8532-29 sor Mau
R. boryi 8537 sor Mau
R. boryi 8526 ap Mau
R. boryi 8538 ap Mau
R. boryi 8507 ap Mau
R. boryi 8527 sor Mau
R. boryi 8506 sor Mau
R. boryi 8500-2 ap Mau
R. boryi 8500-1 ap Mau
R. montagnei 9249 ap Ke
R. montagnei 9239-15 ap Ke
R. montagnei 9262 ap Ke
R montagnei 9252 ap Ke.
R. montagnei 9259 ap Ke
R. montagnei 9239-16 ap Ke
R. montagnei 9728 ap Ang
R. montagnei9700 ap Ang
R. montagnei 9719 ap Ang
R. montagnei 9714 ap Ang
R. montagnei Aptr 66616 sor He
R. montagnei Aptr 66253 sor He
R. montagnei Aptr 66613 ap He
R. montagnei Aptr 66251 ap He
R. montagnei 9740 sor Ang
R. montagnei 9735 sor Ang
R. montagnei 9723-11 sor Ang
R. montagnei Wirth KR2 sor Na
R. montagnei Wirth KR1 sor Na
R. montagnei 9747 ster Ang
R. montagnei 9743 sor Ang
R. montagnei 9724 sor Ang
R. montagnei 9723-12 sor Ang
R. montagnei 9702 sor Ang
R. montagnei 9729 sor Ang
R. montagnei 9718 sor Ang
R. montagnei 9708 sor Ang
R. montagnei 13069 ap Mdg
R. montagnei 9701 sor Ang
R. montagnei 13214 sor Mdg
R. montagnei 13111 ap Mdg
R. montagnei 13074 sor Mdg
R. montagnei 13112 ap Mdg
R. montagnei 13054 sor Mdg
R. montagnei Apt 66650-110 sor He
R. mont _Apt 66650-109 sor He
R. montagnei 9237 apsor Ke
R. montagnei 9235-11 ster Ke
R. montagnei 13213 sor Mdg
R. montagnei 9236 sor Ke
R. montagnei 13219 sor Mdg
R. montagnei Feur sor SAfr
R. montagnei 7709 sor SAfr
R. montagnei 9352a sor So
R. montagnei 9337 sor So
R. montagnei 9332 sor So
R. montagnei 9325 ap So
R. montagnei 9309b-13 sor So
R. montagnei 9263 sor Ke
R. montagnei 9260 sor Ke
R. montagnei 9255 apsor Ke
R. montagnei 9254 sor Ke
R. montagnei 9253 sor Ke
R. montagnei 9248 sor Ke
R. montagnei 9247 sor Ke
R. montagnei 9245 sor Ke
R. montagnei 9235-14 ster Ke
R. montagnei 9225 sor Ke
R. montagnei 9222 ster Ke
R. montagnei 9221 sor Ke
R. montagnei 8879 sor In
R. montagnei 8878 ap In
R. montagnei 8869 sor In
R. montagnei 8864 sor In
R. montagnei 8863 ap In
R. montagnei 9257 sor Ke
R. montagnei 9227 sor Ke
R. montagnei 9250 sor Ke
R. montagnei 9241 sor Ke
R. montagnei 9246 sor Ke
R. montagnei 8868 sor In
R. montagnei 9232-8 sor Ke
R. montagnei 9232-7 sor Ke
R. montagnei 8870 ap In
R. montagnei 8865 sor In
R. montagnei 9244 sor Ke
R. montagnei 13036 ap Mdg
R . montagnei 13020 ap Mdg
3
1
2
Fig. 16. Consensus tree showing species relationships in the genus Roccella.SpeciesincolouredsquaresaretheOldWorldspeciesoutside
Macaronesia recognized in the present study. sor =sorediate, ap =apothecia, apsor =both apothecia and soredia, ster =neither soredia
nor apothecia. Ang =Angola, HK =Hong Kong, In =India, Ke =Kenya, Mdg =Madagascar, Mau =Mauritius, Na =Namibia, He =
Saint Helena, Soc =Socotra, SAfr =South Africa.
Downloaded By: [Tehler, Anders] At: 13:31 16 August 2010

The Old World Roccella species 241
Fig. 17. Parsimony jackknife tree showing relationships (cut frequency 50%) in the genus Roccella.Speciesincolouredsquaresarethe
Old World species outside Macaronesia recognized in the present study. sor =sorediate, ap =apothecia, apsor =both apothecia and
soredia, ster =neither soredia nor apothecia. Ang =Angola, HK =Hong Kong, In =India, Ke =Kenya, Mdg =Madagascar, Mau =
Mauritius, Na =Namibia, He =Saint Helena, Soc =Socotra, SAfr =South Africa.
Chemistry
We fo u n d er y th r i n, l e c an o r ic a c i d, r o cce l lic a c id, s ky ri n
and six unidentified substances in the Old World Roccella
samples. Erythrin, lecanoric acid and the unidentified sub-
stances 1 and 5 were present in all species with the exception
for a few samples in some species. Skyrin was present only
in Roccella babingtonii.Roccellicacidwaspresentinthe
sister pair Roccella balfourii (fertile) and R. phycopsioides
(sorediate) and in R. montagnei and R. boryi.Roccellic
acid was absent from Roccella applanata,R. babingtonii,
R.minuta and R. sinensis. In the two closely related species
Roccella montagnei and R. boryi only sorediate specimens
were shown to contain roccellic acid, except for four fer-
tile specimens of R. montagnei.Outof73TLCinvestigated
Roccella montagnei specimens roccellic acid was present in
49 strictly sorediate specimens; it was absent in 15 strictly
fertile specimens. Two specimens with both soredia and
apothecia and two specimens with neither apothecia nor
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242 A. Tehler et al.
R. montagnei 13069 ap Mdg
R. montagnei 13219 sor Mdg
R. montagnei Feur sor SAfr
R. montagnei Apt 66650-110 sor He
100 R. montagnei Apt 66650-109 sor He
R. montagnei 9701 sor Ang
R. montagnei 9237 apsor Ke
R. montagnei 9236 sor Ke
R. montagnei 9235-11 ster Ke
R. montagnei 7709 sor SAfr
R. montagnei 13214 sor Mdg
R. montagnei 13213 sor Mdg
R. montagnei 13112 ap Mdg
R. montagnei 13111 ap Mdg
R. montagnei 13074 sor Mdg
R. montagnei 13054 sor Mdg
R. montagnei 13036 ap Mdg
R. montagnei 13020 ap Mdg
R. montagnei 9262 ap Ke
R. montagnei 9259 ap Ke
R. montagnei 9252 ap Ke
R. montagnei 9249 ap Ke
R. montagnei 9239-16 ap Ke
R. montagnei 9239-15 ap Ke
R. montagnei 9728 ap Ang
R. montagnei 9719 ap Ang
R. montagnei 9714 ap Ang
R. montagnei 9700 ap Ang
R. montagnei Aptr 66616 sor He
R. montagnei Aptr 66613 ap He
R. montagnei Aptr 66253 sor He
R. montagnei Aptr 66251 ap He
R. montagnei Wirth KR2 sor Na
R. montagnei Wirth KR1 sor Na
R. montagnei 9747 ster Ang
R. montagnei 9743 sor Ang
R. montagnei 9740 sor Ang
R. montagnei 9735 sor Ang
R. montagnei 9729 sor Ang
R. montagnei 9724 sor Ang
R. montagnei 9723-12 sor Ang
R. montagnei 9723-11 sor Ang
R. montagnei 9718 sor Ang
R. montagnei 9708 sor Ang
R. montagnei 9702 sor Ang
R. montagnei 9352a sor Soc
R. montagnei 9337 sor Soc
R. montagnei 9332 sor Soc
R. montagnei 9325 ap Soc
R. montagnei 9309b-13 sor Soc
R. montagnei 9263 sor Ke
R. montagnei 9260 sor Ke
R. montagnei 9257 sor Ke
R. montagnei 9255 apsor Ke
R. montagnei 9254 sor Ke
R. montagnei 9253 sor Ke
R. montagnei 9250 sor Ke
R. montagnei 9248 sor Ke
R. montagnei 9247 sor Ke
R. montagnei 9246 sor Ke
R. montagnei 9245 sor Ke
R. montagnei 9244 sor Ke
R. montagnei 9241 sor Ke
R. montagnei 9235-14 ster Ke
R. montagnei 9227 sor Ke
R. montagnei 9225 sor Ke
R. montagnei 9222 ster Ke
R. montagnei 9221 sor Ke
R. montagnei 8879 sor In
R. montagnei 8878 ap In
R. montagnei 8870 ap In
R. montagnei 8869 sor In
R. montagnei 8868 sor In
R. montagnei 8865 sor In
R. montagnei 8864 sor In
R. montagnei 8863 ap In
R. montagnei 9232-8 sor Ke
R. montagnei 9232-7 sor Ke
R. boryi 8516 ster Mau
D. ceratoniae 9047
D. ceratoniae 9021
D. paradoxa ssp. africana 8524
D. catalinariae 9087
R. allorgei 8163
R. allorgei 8151
R. sinensis 8571 HK
R. sinensis 8564 HK
R. sinensis 8562-2 HK
R. sinensis 8562-1 HK
R. maderensis 8157
R. maderensis 8140
R. fuciformis 8255
R. fuciformis 8171
R. minuta 9334 Soc
R. minuta 9303 Soc
1 R. minuta 9351 Soc
R. minuta 9336 Soc
R. minuta 9307-9 Soc
R. minuta 9307-8 Soc
R. phycopsioides 9352b Soc
R. phycopsioides 9331 Soc
R. phycopsioides 9309a-12 Soc
R. phycopsioides 9308 Soc
R. balfourii 9350 Soc
R. balfourii 9349 Soc
R. balfourii 9335 Soc
R. balfourii 9326 Soc
R. balfourii 9306-7 Soc
R. balfourii 9306-6 Soc
R. balfourii 9302 Soc
R. phycopsis 8295
R. phycopsis 8221
R. tuberculata 8310-50
R. tuberculata 8144
R. tinctoria L1784a
R. tinctoria 8257
R. elisabethae 8306-45
100 R. elisabethae 8272
R. verruculosa 8950-41
R. verruculosa 8941-32
R. bajasurensis 9192
R. bajasurensis 9188
R. gracilis 9190
R. gracilis 8654
R. decipiens 9148
R. decipiens 9080
R. nigerrima 8666
R. nigerrima 8655
R. margaritifera 8705
R. margaritifera 8656
R. lirellina 8904
R. lirellina 8880
R. galapagoensis 8667
R. galapagoensis 8651-12
R. albida 8780
R. albida 8653-14
R. babingtonii 9258 Ke
R. babingtonii 9251-28 Ke
R. babingtonii 9251-27 Ke
R. babingtonii 9242-19 Ke
R. babingtonii 9242-18 Ke
R. babingtonii 9234 Ke
R. babingtonii 9233 Ke
R. babingtonii 9231 Ke
R. applanata Erik101 Mdg
R. applanata 13104 Mdg
R. applanata 13075 Mdg
R. applanata 13060 Mdg
R. applanata 13058 Mdg
R. boryi 8538 ap Mau
R. boryi 8537 sor Mau
R. boryi 8533-31 ap Mau
R. boryi 8533-30 ap Mau
R. boryi 8532-29 sor Mau
R. boryi 8532-28 sor Mau
R. boryi 8527 sor Mau
98
100
100
98
100
100
100
100
98
98
99
100
100
100
100
100 98
00
100
100
100 100
100
100
100 100
100 99
99 100
100
100 100
99
95 100
100
99
100
100 99
100
99
100
99
100
100 R. boryi 8526 ap Mau
R. boryi 8518-18 ap Mau
R. boryi 8507 ap Mau
R. boryi 8506 sor Mau
R. boryi 8500-2 ap Mau
R. boryi 8500-1 ap Mau
R. boryi 8501 ap Mau
3
2
Fig. 18. Parsimony jackknife tree showing significantly supported (<95%) relationships in the genus Roccella all species inclusive.
Species in coloured squares are the Old World species outside Macaronesia recognized in the present study. sor =sorediate, ap =
apothecia, apsor =both apothecia and soredia, ster =neither soredia nor apothecia. Ang =Angola, HK =Hong Kong, In =India, Ke =
Kenya, Mdg =Madagascar, Mau =Mauritius, Na =Namibia, He =Saint Helena, Soc =Socotra, SAfr =South Africa.
Downloaded By: [Tehler, Anders] At: 13:31 16 August 2010

The Old World Roccella species 243
soredia contained roccellic acid. Roccellic acid was miss-
ing in one specimen that lacked apothecia and soredia. The
same pattern as in R. montagnei was observed in Roccella
boryi in which roccellic acid was similarly absent in fertile
specimens (7) and present in sorediate specimens (3).
Discussion
With the present study all 24 recognized species of the genus
Roccella have been revised taxonomically, nomenclaturally
and phylogenetically. The European-Macaronesian and the
New World species were treated in Tehler et al. (2004,
2009a,2009b).
Phylogeny
The resulting tree topologies from Parsimony analyses (par-
simony jackknifing and strict consensus) and Bayesian
inference analysis are in most respects nearly identical.
In general Bayesian inference gave higher resolution and
partly better support values than Parsimony Jackknifing.
Most important major groups were common to all the vari-
ous trees and there were no statistically remarkable conflict
between the Parsimony and Bayesian hypotheses. The most
noteworthy difference was the position of the species Roc-
cella allorgei.InParsimony,althoughinsignificantlysup-
ported, R. allorgei was placed as sister taxon to the rest of
the species in Roccella;intheBayesanalysisitwasplaced
as sister species to Roccella fuciformis/R. maderensis/R.
sinensis/R. minuta four nodes up the tree although most
nodes were poorly supported and none received highly sig-
nificant PP-support. Morphologically the Bayes hypotheses
make sense since R. allorgei is similar to R. fuciformis and
R. maderensis and ascospores size is similar for all species
of that group except R. sinensis which has shorter spores.
Concerning the fully supported American and Macarone-
sian clade (Tehler et al.,2004,2009a,2009b)thepresent
Bayes tree is topologically identical with earlier published
Parsimony trees and nearly all nodes have full PP-support.
The suggested basal relationships are indicated by both the
Parsimony hypotheses (Figs 16, 17) and the Bayes hypoth-
esis (Fig. 15) where all the Old World species of the present
study together with the two Macaronesian species, Roccella
fuciformis and R. maderensis,joinsintoacladewiththerest
of the Old World species to which the Macaronesian species
and the American species forms the sister clade. Both the
Jackknife hypothesis and the Bayes hypothesis are insignif-
icantly supported at this level and they merely confirm the
uncertainty at this part of the tree and that more data are
needed to corroborate sister group relationships.
Distribution and biogeography
Both the species and the species relationships in the
‘montagnei-group’ [ =R. applanata/R. babingtonii/R. bo-
ryi/R. montagnei]werefullyresolvedandsignificantlysup-
ported at all nodes in the Bayesian tree (Fig. 15) whereas
in the PJ tree (Fig. 17) the base of the group formed an un-
resolved trichotomy. The Strict consensus tree (Fig. 16)
is similar to the Bayes tree in that Roccella montagnei
is the sister taxon to R. babingtonii/R. applanata/R. bo-
ryi. Thus, the widespread Roccella montagnei is the most
basal species of that clade and it seems plausible that a
R. montagnei progenitor some time during the evolution-
ary history dispersed to one of the island groups today
known as the Comoros or Seychelles and then further on
to the Mascarenes and southern Madagascar radiating into
the endemic species Roccella babingtonii,R. boryi and
R. applanata respectively. Later Roccella babingtonii dis-
persed back to the mainland Africa. Roccella montagnei is
recorded from many of the Indian Ocean islands such as
Seychelles, Madagascar, Socotra but with the noteworthy
exception of the Mascarene Islands, Mauritius, Rodrigues
and R´
eunion where it strangely enough has not become
established. It cannot be argued that Roccella montagnei
is replaced in the Mascarenes by the closely related en-
demic species R. boryi since R. boryi is a strictly saxicolous
species and thus is not competing for substrate with the
predominantly corticolous R. montagnei. Considering sub-
strate specificity as a feature, we can conclude that the clear
majority of all Roccella species are obligately saxicolous.
Out of 24 recognized species only two are obligately cor-
ticolous, Roccella applanata and R. babingtonii,andtwo
are facultatively but predominantly corticolous, R. gracilis
and R. montagnei.Oneofthecorticolousspecies,Roccella
gracilis,isdistributedintheNewWorldandthethreeoth-
ers are found in Africa–Asia (Figs 2, 9). The ‘montagnei-
group’ is interesting considering substrate specificity. All
the remaining corticolous species belong here and the cor-
ticolous state is synapomorphic for that group, Roccella
applanata,R. babingtonii and R. montagnei. One species
in the ‘montagnei-group’, R. boryi, has reverted to being
saxicolous. Thus, obviously after reverting to the saxicolous
state there has been no successful dispersal event of Roc-
cella montagnei to the Mascarenes which furthermore in-
dicates that this species, although widely distributed, is not
as dispersive as may be anticipated.
Regarding both the Bayes tree and the Consensus-tree a
possible centre of origin and line of descent concerning the
Roccella species would be one traced back to a facultatively
sexual/sorediate progenitor with a distribution in an ancient
limited area which today ranges from the Macaronesian re-
gion to the North East African region presently known as
Africa’s Horn and Socotra. Some time in the evolution-
ary history progenitors of the former group dispersed to
the Americas and progenitors of the latter group dispersed
south along the coasts of the early Indian Ocean.
Since the species of Roccella are mainly restricted to
maritime coastal habitats they consequently often prefer
island substrates such as those found in, for example, the
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244 A. Tehler et al.
Galapagos Islands, the Canary Islands, the Azores, Socotra
and Mauritius, all islands from which the age can be accu-
rately determined and thus could serve as calibration points
for the nodes in a phylogenetic tree. This situation will be
investigated and analysed in more detail in a forthcoming
publication now that all species in the genus Roccella have
been thoroughly studied and circumscribed.
Roccella montagnei species delimitation
Sample groups worthy of taxonomic recognition should not
only form distinct monophyletic groups representing phy-
logenetically distinct units in different single-gene phyloge-
nies (phylogenetic species recognition), it is also preferred
that such groups correlate with external diagnosable fea-
tures (morphological species) and not only internal features
inferred from DNA-data.
Within the ‘montagnei-group’ three taxa, Roccella ap-
planata,R. babingtonii and R. boryi are clear-cut species
that can be distinguished and diagnosed by morphologi-
cal, chemical and molecular data which correlate with ge-
ographical distribution patterns. The fourth species, Roc-
cella montagnei,isamuchmoredisparateandvariable
species. Phylogenetically, as evident from both Parsimony
and Bayesian inference the Roccella montagnei sample
group, which received full support was found to inter-
nally contain an aggregate of several evolutionary lineages.
Some of these were significantly supported whereas others
were poorly supported (Fig. 19). Numerous single samples
were retained in a polytomy at the base of the R. montagnei
node (Figs 17, 18) or insignificantly supported at internal
nodes (Fig. 15).
As evident from both the Bayes tree and the Strict con-
sensus tree molecular data divide the Roccella montagnei
specimens into two major groups, Group 2 and Group 1+3
indicating that at least two of the numerous evolutionary
lineages within the clade should be interpreted as species
and receive species rank.
The fully supported clade in the eastern lineage (Group
3) contains both fertile and sterile specimens. The group
contains sterile specimens which are more closely related to
fertile specimens from the same group than they are to other
sterile specimens within that group or vice versa. Also there
is no phylogenetic distinction between the morphs since
they are mixed in the same clade. Specimens in the eastern
lineage are frequently fertile in India but more rarely fertile
in Africa. The areas of the African south and east coasts,
Madagascar and Socotra Island are dominated by sterile
sorediate Roccella montagnei specimens although fertile
specimens may also be found. The distribution area of the
fully supported eastern lineage stretches around the Indian
Ocean and judging from examined herbarium specimens
the same lineage continues along the coasts of southern In-
donesia and northern Australia. However, no samples from
the latter regions were available to molecularly corroborate
the hypothesis that specimens from those far-away regions
are included in the same clade.
Some of the populations growing in the western part of
Africa, Namibia, Angola and the Atlantic Island St He-
lena form a significantly supported group (Group 2, Fig.
19) which is furthermore internally resolved into smaller
and highly supported subgroups. One of those subgroups
contains strictly fertile specimens with a noteworthy dis-
junction on the opposite coasts of the African continent,
Angola–Kenya. The sister group to the fertile clade contains
both sorediate and fertile specimens but fertile specimens
are confined to the Mid-Atlantic island St. Helena forming
its own significantly supported group. It should be noted
though that some of the sorediate specimens on St. Helena
belong to another, internal sister group within R. montag-
nei (Group 1, Figs 15–19) and are not related to Group
2. A large, significantly supported, strictly sterile sorediate
clade within Group 2 and sympatric with the disjunct fer-
tile clade (Group 2) inhabits the West African coast south to
Namibia. Perhaps the herbarium specimens examined from
Cape Verde should belong to this clade. Even though this
clade is significantly distinct from the fertile and soredi-
ate samples growing on the African east and south coasts
there are no significant morphological, anatomical and
chemical discontinuities to distinguish specimens from one
another.
Although we have not had the opportunity to sample
across the enormous geographical range of Roccella mon-
tagnei we have had a large enough sampling size to be able
to draw the conclusion that there are two major and sig-
nificantly supported intra-specific evolutionary lineages:
Group 3 with an easterly distribution around the Indian
Ocean and Group 2 with a westerly distribution on the
African west coast apart from the disjunction of the fertile
lineage Angola–Kenya within that group.
The situation is complex and far from conclusive since
there is no significantly supported resolution at the base
of the clade. A number of morphologically very similar
but geographically widespread specimens ranging from St.
Helena, South Africa, Kenya and Madagascar group with-
out support or with poor support to the easterly distributed
population. Any attempt to distinguish two or more species
within the Roccella montagnei sample clade would be cer-
tain to fail since we have no means to differentiate between
specimens other than by molecular data. Further, molecular
data is unsupportive for important parts of the present sam-
ple group. We have found neither morphological, anatomi-
cal or chemical diagnostic character discontinuities or geo-
graphical patterns to distinguish or corroborate the genetic
groups obtained from analyses of the molecular data. The
more or less strap-shaped thallus morphology with either
apothecia or soralia in Roccella montagnei is very similar;
the cortex anatomy consist of typical palisade plectenchy-
matic cortex, and; the ascospores are of the same length
and width.
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The Old World Roccella species 245
Fig. 19. Roccella montagnei, distribution and relationships obtained from parsimony jackknife. Numbers on branches show significant
jackknife frequencies. Number within squares =number of DNA samples; s =sorediate; a =with apothecia; a/s =both apothecia and
soredia; st =sterile, neither soredia nor apothecia. Groups 2 and 3, see text and Figs 15–18.
Chemically all Roccella montagnei specimens contain
erythrin and lecanoric acid and often roccellic acid. How-
ever, the distribution pattern of roccellic acid among the
samples within the species turned out to be very surpris-
ing. All sorediate specimens contain roccellic acid whereas
fertile specimens nearly always lack roccellic acid. Only
four fertile specimens deviated from this pattern. However,
presence and absence of roccellic acid as well as presence
or absence of soralia/apothecia cuts right through the very
well-supported molecular groups within both the species
Roccella montagnei as here defined as well as the clade
including all species of the ‘montagnei-group’ and can ob-
viously not be used for distinguishing species. Sorediosity
has evolved separately in closely related species, e.g. in Roc-
cella montagnei and R. boryi and the genetic predisposition
for the production of roccellic acid and development of
soredia obviously co-occur in these two species. However,
in other closely related species such as Roccella balfourii
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246 A. Tehler et al.
(fertile) and R. phycopsioides (sorediate) this is not the case
since both of these species produce roccellic acid.
The geographical distribution pattern of Roccella mon-
tagnei was inconclusive. The polytomy at the base of the
node from Parsimony jackknifing (Fig. 16), consists of 15
single samples and one sister pair from Madagascar, South
Africa, Kenya and Angola (Fig. 19). That polytomy fal-
sifies all attempts to propose any convincing hypotheses
of internal species delimitation even though the Bayesian
analysis gave better support values. It seems reasonable
to suggest that various lineages within Roccella montag-
nei have both radiated and merged during the evolution-
ary history. Speciation is obviously an ongoing process
within this widespread critical group, and without any phe-
netic diagnostic features to distinguish between groups
of specimens, we will for the time being take a prag-
matic approach to the complex and consider it as a single
species.
Acknowledgements
We ar e g ra t e fu l t o F r ed r i k Ro n q ui s t f or g e n er o u sly h e l p-
ing with the Bayesian analyses and we want to thank
Jens Klackenberg for revising the Latin diagnoses. Keyvan
Mirbakhsh, Wim Baert, Francine Demuylder and Patrick
Vivignis are thanked for technical support in the labora-
tory. We thank the curators of the herbaria mentioned in the
Materials and methods for the loan of specimens. D.E. is
indebted to the staff of the Parc Botanique et Zoologique
de Tsimbazaza in Antananarivo and to Prof. E. Fischer, E.
S´
erusiaux and Dr D. Killmann for their guidance and logis-
tical support during the field trip in Madagascar. We would
like to thank the Minist`
ere des Eaux et Forˆ
ets for collection
and export permits of scientific material in Madagascar.
The study was supported financially by Vetenskapsr ˚
adet
(Swedish Science Research Council) grant 621-2005-5275.
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