Two new common, previously unrecognized species in the Sticta weigelii morphodeme (Ascomycota: Peltigeraceae)

Abstract

Sticta is a subcosmopolitan genus most diverse in the tropics. Traditionally, many taxa were considered to be widespread and morphologically variable, following broadly circumscribed morphodemes. Among these is the S. weigelii morphodeme, characterized by a cyanobacterial photobiont and rather narrow, flabellate to truncate or tapering lobes producing predominantly marginal isidia. Molecular phylogenetic analyses focusing on the ITS fungal barcoding marker revealed that this morphodeme represents several species, some of which are only distantly related to each other. Here we describe two species and one subspecies of this morphodeme as new to science, based on analysis of 400 specimens, for 344 of which we generated ITS barcoding data. The two new species, S. andina and S. scabrosa, are broadly distributed in the Neotropics and also found in Hawaii, where the latter is represented by the new subspecies, S. scabrosa subsp. hawaiiensis; in the case of S. andina, the species is also found in the Azores. Sticta andina exhibits high phenotypic variation and reticulate genetic diversification, whereas the phenotypically rather uniform S. scabrosa contains two main haplotypes, one restricted to Hawaii. Sticta andina occurs in well-preserved montane to andine forests and paramos, whereas the two subspecies of S. scabrosa are found in tropical lowland to lower montane forests, tolerating disturbance and extending into anthropogenic habitats.Citation: Moncada B., Mercado-Díaz J. A., Smith C. W., Bungartz F., Sérusiaux E., Lumbsch H. T. & Lücking R. 2021: Two new common, previously unrecognized species in the Sticta weigelii morphodeme (Ascomycota: Peltigeraceae). – Willdenowia 51: 35–45.Version of record first published online on 24 February 2021 ahead of inclusion in April 2021 issue.

Full text

Two new common, previously unrecognized species in
the Sticta weigelii morphodeme (Ascomycota:
Peltigeraceae)
Authors: Moncada, Bibiana, Mercado-Díaz, Joel A., Smith, Clifford W.,
Bungartz, Frank, Sérusiaux, Emmanuël, et al.
Source: Willdenowia, 51(1) : 35-45
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
URL: https://doi.org/10.3372/wi.51.51103
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Willdenowia
Annals of the Botanic Garden and Botanical Museum Berlin
BIBIANA MONCADA1, JOEL A. MERCADO-DÍAZ2, CLIFFORD W. SMITH3, FRANK BUNGARTZ4,
EMMANUËL SÉRUSIAUX5, H. THORSTEN LUMBSCH6 & ROBERT LÜCKING7*
Two new common, previously unrecognized species in the Sticta weigelii morphodeme
(Ascomycota: Peltigeraceae)
Version of record first published online on 24 February 2021 ahead of inclusion in April 2021 issue.
Abstract: Sticta is a subcosmopolitan genus most diverse in the tropics. Traditionally, many taxa were considered
to be widespread and morphologically variable, following broadly circumscribed morphodemes. Among these is
the S. weigelii morphodeme, characterized by a cyanobacterial photobiont and rather narrow, flabellate to truncate
or tapering lobes producing predominantly marginal isidia. Molecular phylogenetic analyses focusing on the ITS
fungal barcoding marker revealed that this morphodeme represents several species, some of which are only distantly
related to each other. Here we describe two species and one subspecies of this morphodeme as new to science, based
on analysis of 400 specimens, for 344 of which we generated ITS barcoding data. The two new species, S. andina
and S. scabrosa, are broadly distributed in the Neotropics and also found in Hawaii, where the latter is represented by
the new subspecies, S. scabrosa subsp. hawaiiensis; in the case of S. andina, the species is also found in the Azores.
Sticta andina exhibits high phenotypic variation and reticulate genetic diversification, whereas the phenotypically
rather uniform S. scabrosa contains two main haplotypes, one restricted to Hawaii. Sticta andina occurs in well-
preserved montane to andine forests and paramos, whereas the two subspecies of S. scabrosa are found in tropical
lowland to lower montane forests, tolerating disturbance and extending into anthropogenic habitats.
Key words: Ascomycota, Azores, Hawaii, Neotropics, Peltigeraceae, Sticta, Sticta weigelii
Article history: Received 27 May 2020; peer-review completed 14 August 2020; received in revised form 2 Septem-
ber 2020; accepted for publication 3 September 2020.
Citation: Moncada B., Mercado-Díaz J. A., Smith C. W., Bungartz F., Sérusiaux E., Lumbsch H. T. & Lücking R.
2021: Two new common, previously unrecognized species in the Sticta weigelii morphodeme (Ascomycota: Pelti-
geraceae). – Willdenowia 51: 35 – 45. doi: https://doi.org/10.3372/wi.51.51103
Introduction
The genus Sticta is among the most conspicuous mac-
rolichens, recognized by its usually large, leathery thalli
with well-defined pores (cyphellae) on the lower side
(Galloway 1994, 2001, 2007; Brodo & al. 2001; Mon-
cada & al. 2014). Prior to molecular phylogenetic stud-
ies, approximately 115 species had been distinguished in
this genus (Kirk & al. 2008), many of them presumed
to be widespread, and common taxa had been circum-
1 Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26D-54, Torre de Laboratorios, Herbario,
Bogotá D.C., Colombia; Research Associate, Negaunee Integrative Research Center, The Field Museum, 1400 South Lake Shore,
Chicago, IL 60605, U.S.A.
2 Committee on Evolutionary Biology, University of Chicago, 1025 E. 57th Street, Chicago, Illinois 60637, U.S.A.; Negaunee Inte-
grative Research Center, The Field Museum, 1400 South Lake Shore, Chicago, IL 60605, U.S.A.; Oahu Army Natural Resources
Program, 3190 Maile Way, St. John Hall Room #409, Honolulu, HI 96822, U.S.A.
3 Oahu Army Natural Resources Program, 3190 Maile Way, St. John Hall Room #409, Honolulu, HI 96822, U.S.A.
4 Biodiversity Knowledge Integration Center & School of Life Sciences, Arizona State University, P.O. Box 874108, Tempe, AZ
85287-4108, U.S.A.; Charles Darwin Foundation for the Galapagos Islands, Puerto Ayora, Ecuador; Instituto Nacional de Biodi-
versidad (INABIO), Quito, Ecuador.
5 University of Liège, Sart Tilman B22, InBioS Research Center, Evolution and Conservation Biology, B-4000 Liège, Belgium.
6 Negaunee Integrative Research Center, The Field Museum, 1400 South Lake Shore, Chicago, IL 60605, U.S.A.
7 Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Königin-Luise-Straße 6 – 8, 14195 Berlin, Germany;
Research Associate, Negaunee Integrative Research Center, The Field Museum, 1400 South Lake Shore, Chicago, IL 60605,
U.S.A.; *e-mail: r.luecking@bgbm.org (author for correspondence).
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36 Moncada & al.: Two new species in the Sticta weigelii morphodeme
scribed through broadly defined morphologies. For in-
stance, cyanobacterial forms with broadly rounded lobes
producing laminal isidia have been identified with the
name S. fuliginosa (Dicks.) Ach. and those with nar-
row lobes producing predominantly marginal isidia as S.
weigelii (Ach.) Vain. Other commonly employed names
are S. limbata (Sm.) Ach. for cyanobacterial forms with
marginal soredia, and S. canariensis (Bory) Delise, S.
damicornis (Sw.) Ach. or S. dichotoma (Bory) Delise
for species associating with green algae and forming
dichotomously branched thalli (Joshi & Awasthi 1982;
Swinscow & Krog 1988; Galloway & al. 1995; Büdel
& al. 2000; Oliveira & al. 2002; Farkas 2003; Galloway
& Thomas 2004; Jørgensen & Tønsberg 2007; Makryi
2008; Smith & al. 2009).
Molecular phylogenetic and revisionary studies showed
that such broadly circumscribed taxa represent several to
sometimes numerous, often only distantly related species
(McDonald & al. 2013; Moncada & al. 2013, 2014, 2018;
Magain & Sérusiaux 2015; Dal Forno & al. 2018; Simon
& al. 2018; Ekman & al. 2019; Mercado-Díaz & al. 2020).
The Sticta weigelii morphodeme evolved multiple times
independently in various clades, resulting in the descrip-
tion of several new species, e.g. S. borinquensis Merc.-
Díaz & Lücking and S. rhizinata B. Moncada & Lücking,
and the reinstatement of some old names, such as S. beau-
voisii Delise (Moncada & Lücking 2012; McDonald & al.
2003; Moncada & al. 2014; Mercado-Díaz & al. 2020).
These more narrowly delimited species are not only phy-
logenetically supported but also exhibit diagnostic pheno-
typic characters that had previously been considered envi-
ronmental or ontogenetical variation (Swinscow & Krog
1988; Galloway 1994, 1997).
As result of a broad sampling of mostly neotropical
representatives of Sticta, we accumulated a large amount
of data for two undescribed lineages corresponding to the
S. weigelii morphodeme. Among all globally recognized
clades corresponding to this morphodeme, the two newly
recognized lineages were the most abundant. Yet, no names
were found in the literature that could be applied to them
and they are here formally described as new to science,
under the names S. andina B. Moncada, Lücking & Sérus.
and S. scabrosa B. Moncada, Merc.-Díaz & Bungartz, the
latter with two subspecies, subsp. scabrosa from the Neo-
tropics and subsp. hawaiiensis B. Moncada, Lücking & C.
W. Sm. from Hawaii. Judging from pre-molecular treat-
ments including specimens now assigned to these taxa,
both had previously identified with the broadly delimited
name S. weigelii and more recently also with the name S.
beauvoisii (Benner & Vitousek 2012). However, the two
species are not closely related to either S. weigelii s.str.
or S. beauvoisii and both are also only distantly related to
each other. The two lineages were first informally recog-
nized using a broad ITS-based phylogeny (Moncada & al.
2014). In that study, what is now recognized as S. andina
was believed to represent seven dierent species, labelled
“andina”, “colombiana”, “dioica”, “paramuna”, “phyl-
lidiata”, “a. phyllidiata” and “squamifera”. However,
the data now available are more consistent with merging
three of these (“andina”, “colombiana”, “paramuna”)
into a single lineage, with apothecia and/or cylindric to
mostly flattened isidia or phyllidia (Moncada & al. 2014:
220, fig. 4, 223, fig. 8; Moncada & al. 2020). In contrast,
S. scabrosa, previously recognized as a single species
(Moncada & al. 2014: 218, fig. 3), even with the now
much expanded data set was found to be morphologically
and genetically rather uniform, except for a unique sur-
face morphodeme occurring solely in Hawaii.
Material and methods
ITS barcoding sequences of the genus Sticta were assem-
bled for a much expanded data set of ingroup 677 OTUs
(Suppl. File S1; Moncada & al. 2020), as compared to 370
OTUs published previously (Moncada & al. 2014). The
S. andina complex initially comprised 19 OTUs, all sam-
pled in Colombia, as mentioned above corresponding to
three OTUs informally labelled “andina”, “colombiana”
and “paramuna” (Moncada & al. 2014). For an updated
ITS-based phylogeny, we assembled a total of 164 OTUs
from Central America, Colombia, Ecuador, Brazil, the
Azores and Hawaii (Moncada & al. 2020). Sticta scabrosa
was initially based on nine OTUs from Colombia and the
Dominican Republic (Moncada & al. 2014), while the up-
dated set included 180 OTUs from Mexico, Costa Rica,
the Dominican Republic, Puerto Rico, Colombia, Brazil,
Argentina, Galapagos and Hawaii (Moncada & al. 2020).
The updated alignment was assembled in BIOEDIT
7 (Hall 1999) and sequences were aligned with MAFFT
7 (Katoh & Standley 2013) using the [ –auto] option. The
alignment included 677 ingroup OTUs and was 626 bas-
es long. We did not detect critical alignment ambiguity
and so included all sites, in order to maximize terminal
resolution. Phylogenetic analysis was performed using
maximum likelihood in RAxML 8.2.12 on the CIPRES
Science Gateway (Stamatakis 2015; Miller & al. 2010),
applying the GTR-Gamma model and 98 bootstrap pseu-
doreplicates according to an automated saturation crite-
rion. The resulting tree were visualized in FigTree 1.4.2
(Rambaut 2016).
Morphological characters of specimens of Sticta
andina and S. scabrosa were assessed at the Universidad
Distrital Francisco José de Caldas (Bogotá), the Field
Museum (Chicago), the Université de Liège and the Bo-
tanischer Garten und Botanisches Museum, Freie Uni-
versität Berlin using standard microscopical techniques
described in Moncada (2012) and Ranft & al. (2018).
Results and Discussion
Sticta andina and S. scabrosa form two large clades in
the terminal portion of a global ITS-based Sticta phylog-
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37Willdenowia 51 – 2021
eny (Fig. 1; Suppl. File S2). Both species are geographi-
cally largely overlapping, exhibiting similar distribution
ranges, predominantly across the Neotropics and Hawaii,
with S. scabrosa also including the Caribbean (Domini-
can Republic and Puerto Rico) and the Galapagos Is-
lands, whereas S. andina is further present in the Azores
(Moncada & al. 2020). Sticta andina includes apotheci-
ate and isidiate to phyllidiate specimens, as well as sun
and shade forms (Fig. 2). Sticta scabrosa is uniformly
phyllidiate but, besides most specimens having an un-
even lobe surface, those from Hawaii may also present
an unique surface with foveolate-pitted lobe tips (Fig. 3).
Until recently, representatives of Sticta with a cyano-
bacterial photobiont and predominantly marginal isidia
were considered a single species, S. weigelii, presumably
with a subcosmopolitan distribution (Joshi & Awasthi
1982; Swinscow & Krog 1988; Galloway 1994, 2001,
2007; Galloway & al. 1995; Büdel & al. 2000; Brodo &
al. 2001; Farkas 2003; Galloway & Thomas 2004). How-
ever, like other morphodemes in the genus (Moncada &
al. 2014; Magain & Sérusiaux 2015; Simon & al. 2018),
the S. weigelii morphodeme appears to have evolved mul-
tiple times within the genus, and most of the sequences
previously deposited under this name do not represent
this taxon (Moncada & al. 2014). Sticta weigelii s.str. is
widely distributed in the Neotropics including the Car-
ibbean (Fig. 1; Suppl. File S2). Material identified with
this name from other regions does not represent this
Fig. 1. Global phylogeny (circle cladogram) of the largest clade of the genus Sticta containing the target taxa, based on the ITS
barcoding marker, showing the position of the newly recognized taxa with the S. weigelii morphodeme in relation to S. weigelii s.str.
For full tree with bootstrap support values, see Suppl. File S2 (see also Moncada & al. 2020).
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38 Moncada & al.: Two new species in the Sticta weigelii morphodeme
taxon. For instance, the ITS accession identified as S.
weigelii from Taiwan (AB245124) is an undescribed rel-
ative of the latter, whereas the sample from South Korea
(KF730791) represents the cosmopolitan S. fuliginosa
s.str. Another specimen from Guyana (AF524905) is
an undescribed relative of S. scabrosa (see Suppl. File
S2). Sticta weigelii s.str. is a species of tropical climates,
similar in altitudinal zonation to S. scabrosa, but more
ane to well-preserved forest, characterized by dark to
blackened, marginal isidia, a rather thin and rather dark
lower tomentum and partly yellow cyphellae (Galloway
2006; Moncada 2012; Mercado-Díaz & al. 2020). The
two species newly recognized here, S. andina and S.
scabrosa, represent the S. weigelii morphodeme in hav-
ing a cyanobacterial photobiont and elongate lobes with
marginal isidia and/or phyllidia, but are phylogenetically
unrelated to each other and to S. weigelii (Fig. 1; Suppl.
File S2). Sticta andina agrees with S. weigelii s.str. in
the slightly shiny thallus surface and rather dark vegeta-
tive propagules, but it diers in the frequent formation
of phyllidia; it also has a much thicker lower tomentum,
persistently white cyphellae and frequently produces
apothecia, and further diers in its preference for upper
montane to andine habitats. Sticta scabrosa has a simi-
lar ecology as S. weigelii, although it is more commonly
found in disturbed and anthropogenic habitats, but its
thallus surface is opaque and thinly scabrose, especially
toward the margins. The lower tomentum is grey-brown
and much thicker, and its cyphellae are consistently
white to at most cream-coloured but never yellow. The
three species are therefore not only phylogenetically
distinct but also morphologically and ecologically well
distinguished.
Although Sticta andina and S. scabrosa have broadly
overlapping geographic ranges, they exhibit dierent
evolutionary histories, which may be explained by their
autecology. As a species largely confined to undisturbed
andine forests and paramos, S. andina underwent frag-
mentation and partial isolation in the recent past, due to
the insular nature of these habitats both in space and time
(Moncada & al. 2020). Similar eects on genetic diver-
sification have been shown for andine orchids, bromeli-
ads and paramo plants (Küper & al. 2004; Givnish 2010;
Madriñan & al. 2013; Givnish & al. 2014, 2015). The
level of genetic and morphological diversification in S.
andina is indeed much higher than in other widespread
species of the genus, particularly S. fuliginosa s.str. and
S. limbata s.str. (Moncada 2012; Moncada & al. 2014;
Magain & Sérusiaux 2015). Based on an earlier ITS-
based phylogeny of Sticta (Moncada & al. 2014), the
clade now recognized as S. andina, at that time with 19
accessions from Colombia, had been tentatively divided
into three taxa (“andina”, “colombiana”, “paramuna”).
In our much-expanded dataset, with more than eight
times as many accessions, the original “andina” corre-
sponds to a small group in one of the subclades, whereas
“colombiana” and “paramuna” largely represent two
other subclades. Despite the observed phylogenetic
structure and morphological variation, separating these
subclades at species level is not warranted, because the
internal topology is not supported and there is no clear-
cut correlation between the subclades and the observed
morphological variation.
In contrast, Sticta scabrosa was found to be phy-
logenetically less complex, with a distinctive haplotype
present only in Hawaii, and morphologically more uni-
form, except for a deviating morphodeme with foveo-
late surface in part of the Hawaiian material. Because
this species is frequent in tropical settings and tolerates
disturbances, it may retain more eective genetic inter-
change between populations (Moncada & al. 2020). Stic-
ta scabrosa is thus rather well-defined as a taxon and the
Hawaiian metapopulation is formally best classified as a
subspecies.
Taxonomic treatment
Sticta andina B. Moncada, Lücking & Sérus., sp. nov. –
MycoBank 836871. – Fig. 2.
Holotype: Colombia, Cundinamarca, Mun. Chipaque,
Vereda Marilandia, vía Santuario, 04°26'N, 74°01'W,
2400m, 8 Sep 2011, B. Moncada 4802 (UDBC).
– “Sticta andina” B. Moncada & Lücking in Mon-
cada, El Género Sticta (Schreb.) Ach. en Colombia:
Taxonomía, Ecogeografía e Importancia: 44. 2012,
nom. inval. (Turland & al. 2018: Art. 29.1, F.5).
– “Sticta colombiana” B. Moncada & Lücking in Mon-
cada, El Género Sticta (Schreb.) Ach. en Colombia:
Taxonomía, Ecogeografía e Importancia: 61. 2012,
nom. inval. (Art. 29.1, F.5).
– “Sticta paramuna” B. Moncada & Lücking in Mon-
cada, El Género Sticta (Schreb.) Ach. en Colombia:
Taxonomía, Ecogeografía e Importancia: 147. 2012,
nom. inval. (Art. 29.1, F.5).
Diagnosis — Diering from Sticta weigelii in the forma-
tion of mostly flattened or dorsiventral isidia and phyl-
lidia instead of cylindric isidia, a thicker lower tomen-
tum, consistently white cyphellae, and the upper montane
versus lower montane-tropical habitat preferences.
ITS barcoding marker accession — KC732688 (holo-
type).
Description — Thallus forming suborbicular rosettes or
becoming irregular, up to 15 cm in diam., moderately
branched, with 3 – 5 branches per 5 cm radius; ramifica-
tion anisotomous to pleurotomous or rarely appearing
polytomous. Lobes leathery, flabellate to ligulate with
rounded tips, adnate to horizontal or slightly ascend-
ing, adjacent to imbricate, undulate to slightly canal-
iculate; margins entire to sinuose or shallowly crenate,
not thickened; lobe internodes (5 –)7 – 12(– 15)mm long,
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39Willdenowia 51 – 2021
Fig. 2. Sticta andina, morphology and anatomy. – A – C: specimens in situ (A: Colombia, Moncada 7452; B: Colombia, Moncada
7948; C: Colombia, Moncada 4802, holotype); D, E: lobe tip and phyllidia enlarged (Colombia, Moncada 4802, holotype); F: lower
surface tomentum and cyphellae (Colombia, Moncada 4936); G: section showing lower primary tomentum (Colombia, Moncada
4936); H: section showing lower secondary tomentum (Colombia, Moncada 4592); I: thallus section showing upper and lower cor-
tex, photobiont layer and medulla (Colombia, Moncada 4936); J: close-up of section showing upper cortex (Colombia, Moncada
4592); K: close-up of medulla showing crystals (Colombia, Moncada 4592); L: mature apothecia (Colombia, Moncada 4592); M:
immature apothecium (Colombia, Moncada 4936); N: ascus with ascospores (Colombia, Moncada 4592); O: ascospores (Colom-
bia, Moncada 4936). – Scale bars: E, F = 1mm; L, M = 0.1mm; G, I = 100µm; H, J, K, N, O = 10µm.
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40 Moncada & al.: Two new species in the Sticta weigelii morphodeme
(3 –)7 – 15(– 20)mm broad. Upper surface smooth to ru-
gose or shallowly scrobiculate, glabrous, without or with
scattered papillae, olive when fresh, with various shades
of brown when dry, shiny, with lobe margins of same
colour and with abundant but usually indistinct, irregu-
lar, cream-coloured maculae. Medulla compact, white to
cream-coloured, sometimes with yellowish patches, K+
yellow, C−, KC−, P−. Marginal cilia rare, up to 0.5mm
long, agglutinate to fasciculate, dark brown to blackish;
tomentum sometimes projecting laterally beyond lobe
margins to resemble cilia. Lower surface usually uneven
to undulate, dark brown to blackish. Lower tomentum
composed of two types; primary tomentum dense up to
lobe margins, rather thick and spongy, becoming thin-
ner toward lobe margins, dark brown to blackish brown,
composed of fascicles of 12 – 20 agglutinate, branched
and apically intertwined hyphae, 0.2 – 1 mm long; sec-
ondary tomentum appressed to surface, arachnoid, pale,
composed of individual, branched, moniliform hyphae
10 – 35µm long. Rhizines abundant, developed centrally
to submarginally along lobe undersides, fasciculate to
squarrose or anziform to hapteriform, dark brown, up
to 4mm long. Cyphellae abundant, about (40 –)60 – 100
percm2 toward periphery and 1 – 20 per cm2 toward cen-
tre of thallus, erumpent to prominent or becoming sessile,
below level of tomentum, rounded, broadly urceolate,
(0.1 – )0.3 – 1(– 1.8)mm in diam., with inner, basal mem-
brane somewhat wider and cavity 60 – 150(– 300)µm high;
margin light to dark brown or blackish, glabrous; basal
membrane white, pubescent, K+ yellow, C−, KC−, P−;
cells of basal membrane lacking papillae on outer side.
Upper cortex paraplectenchymatous, 25 – 45 µm
thick, composed of two dierentiated strata; upper stra-
tum brownish, composed of a single cell layer, with
small, pachydermatous cells 3 – 5µm in diam., with their
walls 1.3 – 2.5µm thick; lower stratum composed of 3 – 5
cell layers, with larger, leptodermatous cells 6 – 12µm in
diam., with their walls 0.7 – 1.3 µm thick. Photobiont a
species of Nostoc Vaucher ex Bornet & Flahault; pho-
tobiont layer 35 – 75µm thick, with individual photobi-
ont cells 10 – 17µm in diam. Medulla 50 – 100(– 170)µm
thick, with individual hyphae 2 – 2.5 broad, inspersed
with yellow-orange crystals. Lower cortex paraplecten-
chymatous, 17 – 35µm thick, composed of 2 or 3 cell lay-
ers; cells 5 – 14µm in diam., with their walls 0.7 – 2.5µm
thick.
Vegetative propagules (flattened) isidia to phyllidia,
formed densely along lobe margins and extending onto
lamina, richly branched and becoming palmate to coral-
loid, up to 1mm long and 0.5mm broad, with terminal
parts dorsiventrally flattened to squamiform and basal
part forming a short, cylindric stipe, often with cyphellae
primordia; phyllidia of same colour as thallus or often
somewhat darker on upper side, somewhat paler on un-
derside, nitidous.
Apothecia common, often on thalli lacking vegeta-
tive propagules, rarely both developed on same thallus;
apothecia biatorine, mostly submarginal, sometimes lam-
inal or marginal, usually dispersed, sessile to substipitate
with pronounced invagination on underside, 2 – 4mm in
diam. and 0.5 – 0.6mm high; disc orange-brown, opaque
to slightly shiny; proper margin verrucose to crenulate,
sometimes thinly pilose when young, dark brown. Ex-
cipulum 100 – 150 µm broad. Hymenium 115 – 155 µm
high; epithecium 2.5 – 5 µm thick, orange-brown. As-
cospores broadly fusiform, 1(– 3)-septate, 27 – 38 ×
9 – 13µm. Pycnidia immersed.
Etymology — The epithet was selected among the names
originally considered for this complex (Moncada & al.
2014), because it best fits the centre of distribution and
the ecology of this species.
Distribution and ecology — Naturally distributed across
the Neotropics, with its centre in the northern Andes.
Given that the records from Hawaii and the Azores rep-
resent the same haplotype as one of the common haplo-
types present in the Andes, these disjunct records may be
the result of recent, perhaps anthropogenic long-distance
dispersal. Biogeographic connections between the Neo-
tropics, Hawaii and the Azores have been reported for
other lichens and bryophytes, e.g. the Sticta ciliata Taylor
complex (Magain & Sérusiaux 2015; Mercado-Díaz &
al. 2020; Moncada & al. 2021) and the liverworts Lepto-
scyphus azoricus Grolle (Vanderpoorten & Long 2006;
Devos & Vanderpoorten 2009) and Syzygiella rubricau-
lis (Nees) Steph. (Maciel-Silva & al. 2016). However, in
these cases, the material found in dierent regions repre-
sents distinct lineages and not identical haplotypes as in
S. andina.
In South America, Sticta andina is an upper montane
to andine species mostly found above 2500m elevation
and usually confined to well-preserved forest and para-
mo habitats, typically growing epiphytically on trunks
and branches of trees and shrubs. In Hawaii and in the
Azores, the species is also found in humid montane for-
ests, although generally at lower altitudes due to a mass
elevation eect.
Remarks — Sticta andina is one of a number of partially
unrelated species that share the S. weigelii morphodeme,
i.e. associated with a cyanobacterial photobiont and form-
ing marginally isidiate-phyllidiate lobes. From S. weigelii
s.str. (Galloway 2006; Moncada 2012; Mercado-Díaz &
al. 2020), S. andina can be dierentiated by the often flat-
tened to dorsiventral isidia and phyllidia (vs. consistently
cylindric isidia in S. weigelii), the much thicker lower to-
mentum, the uniformly white cyphellae (vs. partially yel-
low in S. weigelii) and the upper montane versus lower
montane-tropical habitat. Sticta rhizinata is also similar
but can be distinguished by the narrower lobes, the more
or less cylindric, rather dark isidia, the formation of con-
spicuous, large rhizines on the underside, and the predomi-
nant growth on the ground between bryophytes (Moncada
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41Willdenowia 51 – 2021
& Lücking 2012). Sticta scabrosa (see below) diers by
the marginally thinly scabrose lobe surface, the lighter
brown phyllidia that are concolorous with the thallus, and
the light grey-brown lower tomentum, together with a pre-
ferred growth in tropical, often disturbed or anthropogenic
habitats.
Additional specimens examined — See Suppl. File S1.
Sticta scabrosa B. Moncada, Merc.-Díaz & Bungartz,
sp. nov. – MycoBank 836872. – Fig. 3A – O.
Holotype: Colombia, Cesar, Mun. Río de Oro,
08°16'51"N, 73°25'01"W, 1714m, 15 Oct 2010, B. Mon-
cada 4403 (UDBC; isotype: COL).
– “Sticta scabrosa” B. Moncada & Lücking in Mon-
cada, El Género Sticta (Schreb.) Ach. en Colombia:
Taxonomía, Ecogeografía e Importancia: 172. 2012,
nom. inval. (Turland & al. 2018: Art. 29.1, F.5).
Diagnosis — Diering from Sticta beauvoisii in the thin-
ly scabrose versus glabrous lobe surface, the formation
of dorsiventrally flattened phyllidia instead of cylindric
isidia, and the more greyish lower tomentum.
Description — Thallus forming suborbicular rosettes or
becoming irregular, up to 20 cm in diam., frequently
branched, with 6 – 10 branches per 5cm radius; ramifi-
cation anisotomous to polytomous. Lobes leathery, fla-
bellate to ligulate with rounded tips, adnate to horizon-
tal, imbricate, undulate to slightly canaliculate; margins
entire to shallowly crenate, not thickened; lobe inter-
nodes 5 – 7(– 10)mm long, 3 – 10(– 15)mm broad. Upper
surface uneven to weakly scrobiculate toward older por-
tions of thallus (subsp. scabrosa and subsp. hawaiiensis
p.p.) to rarely foveolate-pitted toward lobe tips (subsp.
hawaiiensis p.p.), often sparsely scabrous toward mar-
gins, without or rarely with scattered papillae featuring
trichomes (subsp. hawaiiensis p.p.), olive when fresh,
with various shades of brown when dry, usually opaque,
with lobe margins of same colour and with usually indis-
tinct, irregular, cream-coloured maculae. Medulla com-
pact, light cream-coloured, K− to K+ pale ochraceous-
yellow, C−, KC−, P−. Marginal cilia rare, when present
up to 0.5mm long, fasciculate, pale to golden brown;
lower tomentum often protruding beyond margins and
then resembling short cilia. Lower surface undulate,
pale brown to brownish yellow. Lower tomentum com-
posed of two types; primary tomentum dense but spars-
er toward lobe margins, rather thick and spongy, becom-
ing thin toward lobe margins, pale to dark grey-brown,
composed of fascicles of 6 – 12 agglutinate, branched
and apically intertwined hyphae, 0.2 – 1mm long; sec-
ondary tomentum appressed to surface, arachnoid, pale,
composed of individual, branched, moniliform hyphae
15 – 25 µm long. Rhizines sparse, dispersed, fascicu-
late to fibrillose, grey-brown with pale tips, up to 2mm
long. Cyphellae abundant, about 40 – 60 per cm2 toward
periphery and 20 – 40 percm2 toward centre of thallus,
erumpent to prominent, below level of tomentum, round-
ed, broadly urceolate, (0.3 –)0.5 – 1.2(– 1.8)mm in diam.,
with inner, basal membrane somewhat wider and cavity
90 – 150(– 250)µm high; margin cream-coloured to light
brown sometimes with a yellowish tinge, usually tomen-
tose; basal membrane cream-coloured to pale yellowish,
pubescent, K+ ochraceous, C−, KC−, P−; cells of basal
membrane lacking papillae on outer side.
Upper cortex paraplectenchymatous, 25 – 30 µm
thick, homogeneous, composed of 3 or 4 cell layers,
cells 4 – 12 µm in diam., with their walls 0.7 – 2.5 µm
thick. Photobiont a species of Nostoc; photobiont lay-
er 50 – 70 µm thick, with individual photobiont cells
10 – 15µm in diam. Medulla 80 – 170µm thick, with in-
dividual hyphae 2 – 2.5 broad, lacking crystals. Lower
cortex paraplectenchymatous, 20 – 30 µm thick, com-
posed of 2 or 3 cell layers; cells 4 – 12µm in diam., with
their walls 0.7 – 2.5µm thick.
Vegetative propagules phyllidia, formed densely
along lobe margins but sometimes extending onto lam-
ina, richly branched and becoming palmate to coralloid,
up to 0.5 mm long and 0.3 mm broad, with terminal
parts dorsiventrally flattened to squamiform and basal
part forming a short, flattened stipe, without cyphellae
primordia; phyllidia of same colour as thallus or some-
what darker on upper side, somewhat paler on underside,
slightly nitidous.
Apothecia not observed. Pycnidia immersed.
Etymology — The epithet refers to the often thinly scab-
rose lobe surface toward the tips.
Distribution and ecology — Sticta scabrosa is widely
distributed in the Neotropics, including the Caribbean
and the Galapagos Islands (subsp. scabrosa) and also
found in Hawaii (subsp. hawaiiensis, see below). In con-
trast to S. andina, it is a lowland to lower montane, tropi-
cal species found both in forest habitats and in disturbed
or anthropogenic situations, e.g. on planted trees along
roads, sometimes with a weedy character. Both subspe-
cies have a similar ecology (Moncada & al. 2021).
Remarks — Like Sticta andina, S. scabrosa also corre-
sponds to the S. weigelii morphodeme, but is more sim-
ilar to S. beauvoisii than to S. weigelii s.str., a species
only recently removed from synonymy under S. weigelii
(McDonald & al. 2003). Sticta scabrosa is not closely re-
lated to either S. weigelii s.str. or S. beauvoisii (Moncada
& al. 2014; this paper) and diers from the latter in the
thinly scabrose versus glabrous lobe surface, the forma-
tion of dorsiventrally flattened phyllidia instead of mostly
cylindric isidia, and the more greyish lower tomentum.
The two subspecies, subsp. scabrosa and subsp. hawai-
iensis, are morphologically mostly identical, but the lat-
ter includes a distinctive lobe surface morphodeme (see
below).
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42 Moncada & al.: Two new species in the Sticta weigelii morphodeme
Fig. 3. Sticta scabrosa, morphology and anatomy. – A – L: S. scabrosa subsp. scabrosa; M – O: S. scabrosa subsp. hawaiiensis;
A: thallus lobes (Colombia, Moncada 4310); B: lobes with phyllidia (Colombia, Moncada 4403, holotype); C: phyllidia enlarged
(Colombia, Moncada 4403, holotype); D: lower tomentum with cyphellae (Colombia, Moncada 4403, holotype); E: section show-
ing lower tomentum (Colombia, Moncada 4403, holotype); F – O: specimens in situ, in part showing lower tomentum with cyphel-
lae (F: Colombia, Cauca, Moncada 7357; G: Costa Rica, Lücking 34607; H: Colombia, Cauca. Moncada 7366; I – K: Brazil, Lü-
cking 40042; L: Brazil, Lücking 40087; M – O: Hawaii, Moncada & al. 6937, 7011, 7015). – Scale bars: C, D = 1mm; E = 10µm.
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43Willdenowia 51 – 2021
Sticta scabrosa subsp. scabrosa – Fig. 3A – L.
Diagnosis — Diering from subsp. hawaiiensis in two
positions in the ITS (see Suppl. File S3), namely position
143 (T > C) and position 401 (T > C). Upper lobe surface
smooth to uneven, never pitted.
ITS barcoding marker accession — MT936608,
MT936611 (both holotype).
Description — See above.
Distribution and ecology — Widely distributed in the
Neotropics, lowlands to lower montane zones, often in
exposed situations (Moncada & al. 2021).
Remarks — See above.
Additional specimens examined — See Suppl. File S1.
Sticta scabrosa subsp. hawaiiensis B. Moncada, Lücking
& C. W. Sm., subsp. nov. – MycoBank 836873. – Fig.
3M – O.
Holotype: U.S.A., Hawaii, Oahu, Koolau Range, Manoa
Valley, 6km ENE of Honolulu and 8km WSW of Kane-
ohe, Manoa Clis Trail, Moleka trailhead to forestry ex-
closure, 21°19'55"N, 157°48'43"W, 410 – 575m, partially
disturbed secondary forest with some exposed vegetation
and some planted trees, 9 Jun 2013, B. Moncada & al.
6915 (BISH; isotypes: B, F).
Diagnosis — Diering from subsp. scabrosa in two po-
sitions in the ITS (see Suppl. File S3), namely position
143 (C > T) and position 401 (C > T). Upper lobe surface
variable, smooth to uneven but in some forms distinctly
foveolate-pitted.
ITS barcoding marker accession — MT132639 (holo-
type).
Description — See above.
Etymology — The subspecific epithet refers to the geo-
graphic distribution of this subspecies.
Distribution and ecology — Restricted to the Hawaiian
archipelago, where it has been found on all five major
islands (Hawaii or Big Island, Oahu, Molokai, Maui and
Kauai). In lowlands to lower montane zones, often in ex-
posed microhabitats (Moncada & al. 2021). Its ecology
is the same as in the species as a whole. It is the only
taxon of the genus present in Hawaii consistently found
in disturbed habitats.
Remarks — Sticta scabrosa subsp. hawaiiensis is here sep-
arated as a formal taxon due to its consistent phylo genetic
dierences with subsp. scabrosa, with a clear geographic
correlation. Because both subspecies cannot be separated
morphologically (with exception of the additional lobe
surface morphodeme present in Hawaii) and the phyloge-
netic dierences are small (two substitutions out of 550 in
a pairwise ITS alignment, i.e. 99.6 % similarity; see Suppl.
File S3), we consider the rank of subspecies appropriate,
to reflect the phylogenetic distinctiveness of this lineage
and its geographic distribution in an isolated archipelago
distant from the geographic range of subsp. scabrosa.
Additional specimens examined — See Suppl. File S1.
Acknowledgements
Funding for field and laboratory work for this study
was provided by two grants from the National Science
Foundation (NSF) to The Field Museum: DEB-1025861
“ATM– Assembling a Taxonomic Monograph: the lichen
family Graphidaceae” and DEB-1354884 “Collabora-
tive research: evolution, diversification, and conserva-
tion of a megadiverse flagship lichen genus”. For recent
collections gathered in Colombia (Bogotá), BM and
RL are also grateful for the financial support from the
Bundesministerium für Bildung und Forschung (BMBF;
Kooperation mit dem Botanischen Garten Bogotá und
der Universidad del Norte Barranquilla, ColBioDiv, FK
01DN17006; Pilotprojekt Kooperation mit dem Bota-
nischen Garten Bogotá, FK 01DN13030) and the Center
for International Cooperation of the Freie Universität
Berlin (CIC; FM Ex3-2017-013). The following col-
leagues and students are thanked for providing addition-
al material or aiding in the logistics of field work: Wil-
son Álvaro, C. J. Arango, M. Cáceres, L. F. Coca, M. Dal
Forno, L. Ferraro, M. A. Herrera-Campos, L. Katib, Y.
Lozano, N. Marín, R. Peláez, S. Silano, D. F. Simijaca,
E. Soto-Medina, A. Suárez, L. Tisnes, J. M. Torres and
L. Vargas. Galapagos material of Sticta was examined as
part of the Galapagos Lichen Inventory by the Charles
Darwin Foundation (CDF) and the Galapagos National
Park (DPNG) as part of a national biodiversity assess-
ment program “Biodiversidad Genética del Ecuador” led
by the Instituto Nacional de Biodiversidad del Ecuador
(INABIO). The authors are indebted to our colleagues
and collaborators at these institutions, especially Daniel
Lara Solís, Galo Quedaza and Victor Carrión (DPNG),
Arturo Izurieta and Maria-José Barragan Paladines
(CDF), and Diego Inclán and Rosa Batallas (INABIO)
for research permits and logistical support. This publi-
cation is contribution number 2363 of the Charles Dar-
win Foundation for the Galapagos Islands. All material
used in this study was collected under the corresponding
permits in collaboration with JAMD (Puerto Rico), BM
(Colombia), FB (Galapagos), Emerson Gumboski (Bra-
zil) and CWS (Hawaii). Specimens from Kauai (Hawaii)
were collected under Special Use Permit K2013-090cc
of the Division of State Parks. Finally we thank Manuela
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44 Moncada & al.: Two new species in the Sticta weigelii morphodeme
Dal Forno (BRIT) and an anonymous reviewer for their
comments on an earlier version of this paper.
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Supplemental content online
See https://doi.org/10.3372/wi.51.51103
Suppl. File S1. Voucher table for the specimens of the
newly described taxa used in this study. For GenBank ac-
cessions of additional species used in the global phylog-
eny, see Suppl. File S2.
Suppl. File S2. Best-scoring maximum likelihood tree of
the largest clade of the genus Sticta containing the target
taxa, based on the ITS barcoding marker, showing the
position of the newly recognized taxa with the S. weigelii
morphodeme in relation to S. weigelii s.str. Branches are
thickened relative to support and bootstrap support val-
ues are indicated.
Suppl. File S3. Alignment of ITS sequences of Sticta
scabrosa subsp. scabrosa and subsp. hawaiiensis (in
FASTA format), showing diagnostic dierences in posi-
tions 143 and 401.
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