An updated checklist of the lichens of St. Eustatius, Netherlands Antilles

Abstract

In the course of a multi-taxon biodiversity inventory for the island of St. Eustatius, lichens were collected from 11 plots representing different vegetation types. From these collections, 126 lichen species are reported, 54 of which are new reports for St. Eustatius. Most species could be identified to species level based on morphological and chemical characters. In a few cases, mtSSU DNA sequences were generated for a preliminary molecular identification and future phylogenetic studies. In total, 263 identified lichen species are currently known from St. Eustatius, as well as some additional genera with yet unidentified species and lichenicolous fungi.

Full text

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 69
An updated checklist of the lichens of St. Eustatius,
Netherlands Antilles
André Aptroot1, Michael Stech2,3
1 ABL Herbarium, Gerrit van der Veenstraat 107, NL-3762 XK Soest, e Netherlands 2 Naturalis Bio-
diversity Center, P.O. Box 9517, NL-2300 RA Leiden, e Netherlands 3 Leiden University, Leiden, e
Netherlands
Corresponding author: André Aptroot (andreaptroot@gmail.com)
Academic editor: G. Rambold|Received 26 January 2018|Accepted 17 March 2018|Published 3 April 2018
Citation: Aptroot A, Stech M (2018) An updated checklist of the lichens of St. Eustatius, Netherlands Antilles.
MycoKeys 33: 69–84. https://doi.org/10.3897/mycokeys.33.23911
Abstract
In the course of a multi-taxon biodiversity inventory for the island of St. Eustatius, lichens were col-
lected from 11 plots representing dierent vegetation types. From these collections, 126 lichen species are
reported, 54 of which are new reports for St. Eustatius. Most species could be identied to species level
based on morphological and chemical characters. In a few cases, mtSSU DNA sequences were generated
for a preliminary molecular identication and future phylogenetic studies. In total, 263 identied lichen
species are currently known from St. Eustatius, as well as some additional genera with yet unidentied
species and lichenicolous fungi.
Keywords
Biodiversity inventory, lichens, mtSSU, St. Eustatius
Introduction
Sint Eustatius is a small island (21 km2) in the northern Leeward Islands part of the
West Indies. It is one of the six islands of the Netherlands Antilles and, since 2010, a spe-
cial municipality of the Netherlands. Sint Eustatius is roughly divided into three parts,
the Northern Hills, the urbanised central area (‘Cultuurvlakte’) and the southern part
dominated by the steep dormant volcano e Quill (600 m elev.). Although the whole
island of St. Eustatius has been heavily impacted by human activities, the northern and
Copyright A. Aptroot and M. Stech. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
MycoKeys 33: 69–84 (2018)
doi: 10.3897/mycokeys.33.23911
http://mycokeys.pensoft.net
A peer-reviewed open-access journal
MycoKeys
Launched to accelerate biodiversity research
CHECKLIST

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
70
southern parts are nowadays designated as National Parks with varied vegetation types
especially on the slopes of e Quill. e latter comprise, for example, thorny wood-
lands, deciduous to evergreen seasonal forests, dry evergreen forest, montane thickets
and eln woodland (Stoers 1956). A re-classication of the vegetation of St. Eustatius,
based on cluster analysis of sample plots, resulted in 13 vegetation types characterised by
dierent combinations of individual vascular plant species (Freitas et al. 2014).
Just as in almost all other groups of organisms, lichens are most diverse in the tropics
(Sipman and Aptroot 2001). On St. Eustatius, they are commonly present on various
substrates, including tree bark and twigs, siliceous rock, limestone, dead wood and liv-
ing leaves. Despite their abundance, the lichen ora of St. Eustatius is still incompletely
known. e authors are aware of only nine publications citing in total 14 lichen spe-
cies from St. Eustatius, viz. Phyllopsora corallina (Eschw.) Müll. Arg. and P. parvifoliella
(Nyl.) Müll. Arg. (Brako 1991), Anisomeridium excellens (Müll. Arg.) R.C. Harris (Har-
ris 1995), Syncesia glyphysoides (Fée) Tehler (Tehler 1997), Pseudopyrenula subnudata
Müll. Arg. (Harris 1998, as P. diluta var. degenerans Vain.), Stirtonia neotropica Aptroot,
described based on material from St. Eustatius and Costa Rica (Aptroot 2009), Syncesia
subintegra Sipman, described based on material from St. Eustatius, as well as S. farinacea
(Fée) Tehler, S. glyphysoides and S. graphica (Fr.) Tehler (Sipman 2009), Roccella gracilis
Bory (Aptroot and Schumm 2011), Dirina paradoxa (Fée) Tehler (Tehler et al. 2013),
as well as Astrothelium bicolor (Taylor) Aptroot & Lücking, A. phlyctaena (Fée) Aptroot
& Lücking and Pseudopyrenula subnudata Müll. Arg. (Aptroot and Lücking 2016).
e main source of information about the lichen ora of St. Eustatius is the on-
line portal ‘Plants and Lichens of St. Eustatius’ (Boom et al. 2009). It contains a list
with 209 lichen species, based on specimens collected by H. Sipman and W.R. Buck
in 2008, identied by H. Sipman and mostly hosted in B (some in NY). In addition,
some identied specimens are present in various other herbaria, some of which can be
searched online. For instance, the database of BR cites the following identied speci-
mens from St. Eustatius, collected by R. Hensen in 1991 and identied by the rst
author: Megalaria bengalensis Jagadeesh Ram & Aptroot, Porina mastoidea (Ach.) Müll.
Arg. and Sticta xanthotropa (Kremp.) D.J. Galloway.
In 2015, a plot-based, multi-taxon biodiversity inventory of St. Eustatius was car-
ried out by Naturalis Biodiversity Center, the European Invertebrate Survey (EIS) and
dierent Dutch non-governmental organisations, together with St. Eustatius National
Parks Foundation (STENAPA) and students from dierent Dutch universities. Here,
the lichen records of that inventory are reported and an updated checklist of the li-
chens known from St. Eustatius is presented.
Materials and methods
As part of a multi-taxon inventory, lichens were collected on St. Eustatius from 11
plots (25 m × 25 m) in dierent main vegetation types according to Freitas et al.
(2014). Two plots (H1, H2) were situated in the Northern Hills area, eight (M1−M5,

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 71
M7−M9) on and around e Quill in the southern part of the island and one (U1) in
the central urban area. Details concerning the location and vegetation characteristics of
the plot locations and the multi-taxon sampling approach are described in van Andel et
al. (2016). Lichens were collected using a knife or hammer and chisel and subsequently
air-dried and stored in paper bags.
Specimens were observed and identied by the rst author using an Olympus SZX7
stereomicroscope and an Olympus BX50 compound mic roscope with interference con-
trast, connected to a Nikon Coolpix digital ca mera. Sections were mounted in tap
water, in which all measurements were also taken. e chemistry of selected specimens
was investigated by thin-layer chromatography (Orange et al. 2001), using solvent A.
DNA analysis based on mitochondrial ribosomal small subunit (mtSSU) sequenc-
es was carried out for ten unidentiable or provisionally identied specimens of good
quality (indicated in Table 1). Although the nuclear ribosomal ITS region is the gener-
ally accepted fungal DNA barcode locus (Schoch et al. 2012), mtSSU was chosen since
more mtSSU than ITS sequences have yet been published for several of the genera or
families to which the respective specimens putatively belong.
Genomic DNA was extracted using the NucleoMag 96 Plant kit (Macherey-Nagel)
on the KingFisher Flex Purication System (ermoFisher Scientic). e mtSSU re-
gion was PCR-amplied following Zoller et al. (1999) in terms of primers (mrSSU1/
mrSSU3R) and the PCR protocol. PCR products were puried and sequenced at Base-
Clear B.V. (www.baseclear.com) using the amplication primers. Sequences were as-
sembled and edited using Geneious v8.1.8 (Biomatters Ltd.) and subjected to a BLAST
search (Altschul et al. 1990) against the GenBank database (megablast; considering,
where possible, BLAST results with E value of 0.0 and query cover >90 %). Sequences
are available in GenBank under accession numbers MH028639−MH028646.
To compile an updated list of the lichens of St. Eustatius, literature and internet
sources were exhaustively consulted for previous reports and previous collectors were
contacted for additional information.
Results and discussion
In total, 126 lichen species (and one identiable lichenicolous fungus) were found in
243 collections (Table 1). e vast majority (113 species) could be identied to species
level based on morphological and chemical characters, even though no identication
book exists for any region nearby. However, many species have been described from
other islands in the Caribbean, which can be expected to have many species in com-
mon. ese were often already described in the 19th century and partly never studied
again, but illustrations of their types are increasingly available. e authors also had
access to various unpublished sources, such as the unpublished keys, descriptions and
specimen citation (by H. Sipman) that was the basis of the internet checklist of St.
Eustatius lichens and keys to the lichens from Puerto Rico (Harris 1989) and Guade-
loupe (Øvstedal 2010), the latter with many illustrations of type and other specimens.

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
72
Table 1. Lichenised and lichenicolous fungi recorded in 11 plots on St. Eustatius. Species names in earlier
publications are indicated in brackets. Author names are given in Table 2. Plots H1 and H2 are situated
in the Northern Hills, M2−M9 on and around the volcano e Quill in the southern part of the island
and U1 in the central urban area. Substrates occupied by each taxon are indicated per plot; b: bark, k:
limestone, l: leaves, r: siliceous rock, s: soil, w: wood. Asterisks indicate rst records for St. Eustatius (aster-
isks in brackets indicate additional taxa that are not yet identied to species level). Black dots () indicate
specimens from which DNA was extracted.
Taxon H1 H2 M1 M2 M3 M4 M5 M7 M8 M9 U1
(*)Acanthothecis sp.  b
Alyxoria culmigena (Opegrapha
herbarum)r w b
*Alyxoria varia b b b b
*Amandinea multispora b
*Anisomeridium subprostans b
Anisomeridium tamarindi b
(*)Anisomeridium sp. corticate c.
pycnidia • b
*Anisomeridium terminatum b
Arthonia antillarum b b
Arthonia caribaea b
Arthonia conferta b b b b b b b
Arthonia cyrtodes b
Arthonia minuta b b
*Arthonia parantillarum b b
Arthothelium macrothecum b b
*Bacidia medialis b, r r
(*)Bacidia sp. apotheciate • b, w
(*)Bacidia sp. sorediate • b
Bactrospora denticulata b b b b b
*Bactrospora jenikii b
*Bogoriella annonacea b b
*Brigantiaea leucoxantha b
Buellia dejungens r r r r r
*Buellia griseovirens w w
Buellia mamillana (Buellia
glaziouana)r
Caloplaca leptozona r r
*Caloplaca obscurella w
Coenogonium linkii b b
*Coenogonium saepincola w
Coenogonium strigosum b
Coniocarpon cinnabarinum
(Arthonia cinnabarina)b
*Crespoa carneopruinata b, r
*Cresponea ava r
*Cryptothecia punctosorediata b
Cryptothecia striata bb, r
(*)Cryptothecia sp. isidiate • b
(*)Cryptothecia sp. sterile • b
*Dactylospora saxatilis
(lichenicolous on Pertusaria
praetervisa)
r
Dichosporidium nigrocinctum b, r b, r

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 73
Taxon H1 H2 M1 M2 M3 M4 M5 M7 M8 M9 U1
*Dictyomeridium amylosporum b b
Diorygma hieroglyphicum b
Diorygma poitaei b
*Diorygma pruinosum b
*Endocarpon pallidulum r r r, k r, s r
Enterographa pallidella b b
Enterographa subserialis b
Flakea papillata bb, r, w b, r r
Glyphis scyphulifera b
Graphis caesiella b
*Graphis cincta b
Graphis dendrogramma b b
*Graphis librata b
Gyalolechia bassiae (Caloplaca
bassiae)bb, r
*Hafellia curatellae b
Hyperphyscia adglutinata bb, r b
Lathagrium neglectum (Collema
neglectum)b
Lecanora legalloana r r r r r
*Lecanora leproplaca bbb
Lecanora leprosa r
Lecanora prosecha r r r r r r
Lecanora sulfurescens r
*Lepraria nkii b
*Leprocollema nova-caledonianum w
Letrouitia domingensis b
Malmidea piperis (Malcolmiella
piperis)b
*Malmidea psychotrioides b
Malmidea vinosa (Malcolmiella
vinosa)b
Mazosia carnea (Mazosia ocellata) b b
(*)Melaspilea sp. (lichenicolous on
Pyrenula dissimulans)b
*Microtheliopsis uleana l
*Mycoporum eschweileri b b b b b
Nyungwea anguinella (Enterographa
anguinella)b
*Opegrapha astraea b b
*Opegrapha lithyrgiza r
*Opegrapha quintana b
(*)Opegrapha sp. k
Peltula bolanderi r
Peltula obscurans r r r
Pertusaria coccopoda r r
Pertusaria praetervisa r r
*Pertusaria texana b
Pertusaria xanthodes b
*Phaeographis crispata b
Phaeographis dendritica b
*Phyllopeltula corticola b
Phyllopsora corallina bb, r r
Physcia atrostriata b

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
74
Taxon H1 H2 M1 M2 M3 M4 M5 M7 M8 M9 U1
*Physcia erumpens b
*Physcia integrata b b
Physcia sorediosa b
*Porina conspersa b, r r
Porina epiphylla l
Porina internigrans b b
*Porina leptalea r
Porina nucula b, r
*Porina rubentior l
Porina tetracerae b b r
*Porina thaxteri l
(*)Psorotichia cf. americana r
*Pyrenopsis antillarum r
*Pyrenula adacta b b b b
Pyrenula breutelii (Pyrenula
macularis)b b
Pyrenula cocoes b b b b b
*Pyrenula cruenta b
*Pyrenula dissimulans b b b b b
Pyrenula nitidula b
Pyxine cocoes b, r r r r b
*Ramalina stoersii r
*Rinodina antillarum r
*Rinodina colobinoides b b
Rinodina pyxinoides r r r r r r
Sarcographa cf. tricosa • b
Squamulea subsoluta (Caloplaca
subsoluta)r r r r
*Staurolemma dussii b
*Sticta xanthotropa r r
*Stigmatochroma gerontoides w
(*)Stigmidium cf. schaereri k
*Strigula decipiens r
*Strigula phaea b, r
Strigula smaragdula l
*Syncesia decussans b
*elenella luridella r r r
(*)elidium cf. decipiens • k
(*)Verrucaria cf. dolosa • r
*Verrucaria nigrescens r
(*)Wetmoreana cf. appressa • r r
Somewhat to the authors’ surprise, as many as 54 (almost 50 %) of the identied
species are new records for St. Eustatius. is includes mostly relatively common and
widespread tropical or Neotropical species, but also some rare species, notably Stauro-
lemma dussii (Vain.) P.M. Jørg. & Henssen, which was so far only known from its type
from Guadeloupe. Furthermore, it is remarkable that Cresponea ava (Vain.) Egea &
Torrente was found on siliceous rock. e presence of so many additional species with-
in the limited surface area of the plots, totalling 6875 m2 (0.03% of the total island sur-
face), suggests that the exploration of the lichen ora of St. Eustatius has not yet been

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 75
exhaustive. However, no clearly undescribed species were found in the material and the
number of species described based on material from St. Eustatius remains low with two,
viz. Stirtonia neotropica (Aptroot 2009) and Syncesia subintegra (Sipman 2009).
Several specimens could not be identied with certainty in the present material
but represent additional species (and in several cases additional genera). ese are, for
instance, Lichinaceae and Verrucariaceae, of which the taxonomy of the tropical taxa
is incompletely known. Rather than describing them as new, they were listed with the
name of the species that is morphologically most similar, preceded by “cf”. e BLAST
results from the mtSSU sequences obtained from eight of these specimens in most
cases allowed preliminary insights into their phylogenetic position.
e sequence of the Anisomeridium specimen with only conidia from St. Eustatius
receives the highest BLAST hits with other representatives of the Monoblastiaceae in
Nelsen et al. (2009, 2011), viz. Anisomeridium ubianum (Vain.) R.C. Harris, A. cf. wil-
leyanum (R.C. Harris) R.C. Harris, Megalotremis verrucosa (Makhija & Patw.) Aptroot
and Trypetheliopsis kalbii (Lücking & Sérus.) Aptroot. e low sequence identities of
86−93% clearly indicate that the St. Eustatius specimen belongs to another species in
that family, but too few mtSSU sequences are yet available for a more precise molecular
identication.
In the Graphidaceae, the top ve BLAST hits for the specimen of Acanthothecis
sp. were all with Acanthothecis peplophora (M. Wirth & Hale) E. Tripp & Lendemer
specimens (97% identity), whereas the identity with the sequence of the type species
of Acanthothecis, A. hololeucoides (Nyl.) Staiger & Kalb, was only 89%. e speci-
men from St. Eustatius thus most probably does not belong to Acanthothecis s.str., but
may represent a species of ‘Acanthothecis 2’ in the Carbacanthographis clade (cf. Rivas
Plata et al. 2013, Medeiros et al. 2017). e Sarcographa cf. tricosa specimen received
BLAST hits of 97% identity with Sarcographina glyphiza (Nyl.) Kr.P. Singh & D.D.
Awasthi and Pallidogramme chlorocarpoides (Nyl.) Staiger, Kalb & Lücking, both situ-
ated in the Graphioideae tribe Graphidae p.p. clade of Rivas Plata et al. (2013). How-
ever, another GenBank sequence of P. chlorocarpoides, as well as several species of other
genera of same clade, were 96 % identical, including the single other specimen of S.
tricosa in GenBank (but not the species of the Sarcographa s.str. clade sensu Rivas Plata
et al. 2013). e identity of the St. Eustatius specimen thus remains ambiguous based
on the presently available mtSSU sequence data.
Both the apotheciate and sorediate Bacidia specimens are closest to sequences of
species of the Toninia-Bacidia p.p. clade in Miadlikowska et al. (2014), the former to
Toninia sedifolia (Scop.) Timdal (94 % identity) and the latter to Bacidia californica S.
Ekman and B. phacodes Körb. (88−89 % identity), respectively. Consequently, they do
not belong to Bacidia s.str., which forms a separate clade (including the type species,
B. rosella (Pers.) De Not.) in Miadlikowska et al. (2014).
In the Verrucariaceae, Verrucaria was resolved as polyphyletic and elidium mixed
with Polyblastia, Staurothele p.p. and Verrucaria p.p. (Polyblastia clade) in molecular
phylogenetic reconstructions (Gueidan et al. 2007, Muggia et al. 2010, üs et al.
2011). e sequence of the elidium cf. decipiens specimen from St. Eustatius, how-

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
76
Table 2. Updated checklist of the lichens of St. Eustatius.
Species References
Lichens
Acarospora chrysops (Tuck.) H.Magn. Boom et al. (2009) as Acarospora dissipata H.Magn.
Alyxoria culmigena (Lib.) Ertz Boom et al. (2009) as Opegrapha herbarum Mont., present
study
Alyxoria ochrocheila (Nyl.) Ertz & Tehler Boom et al. (2009) as Opegrapha ochrocheila Nyl.
Alyxoria varia (Pers.) Ertz & Tehler present study
Amandinea eorescens (Müll. Arg.) Marbach Boom et al. (2009)
Amandinea multispora (Kalb & Vězda) Marbach present study
Amandinea prospersa (Nyl.) Elix & H. Mayhofer Boom et al. (2009) as Buellia prospersa (Nyl.) Riddle
Anisomeridium americanum (A.Massal.) R.C. Harris Boom et al. (2009)
Anisomeridium excellens (Müll. Arg.) R.C. Harris Boom et al. (2009), Harris (1995)
Anisomeridium subprostans (Nyl.) R.C. Harris present study
Anisomeridium tamarindi (Fée) R.C. Harris Boom et al. (2009), present study
Anisomeridium terminatum (Nyl.) R.C. Harris present study
Anisomeridium tuckerae (R.C. Harris) R.C. Harris Boom et al. (2009)
Arthonia antillarum (Fée) Nyl. Boom et al. (2009), present study
Arthonia caribaea (Ach.) A. Massal. Boom et al. (2009), present study
Arthonia conferta (Fée) Nyl. Boom et al. (2009), present study
Arthonia cyanea Müll. Arg. Boom et al. (2009)
Arthonia cyrtodes Nyl. Boom et al. (2009), present study
Arthonia minuta Vain. Boom et al. (2009), present study
Arthonia parantillarum Aptroot present study
Arthothelium macrothecum (Fée) A. Massal. Boom et al. (2009), present study
Astrothelium bicolor (Taylor) Aptroot & Lücking Boom et al. (2009) as Trypethelium nitidiusculum (Nyl.)
R.C. Harris, Aptroot and Lücking (2016)
Astrothelium phlyctaena (Fée) Aptroot & Lücking Boom et al. (2009) as Tr ypethelium ochroleucum (Eschw.)
Nyl., Aptroot and Lücking (2016)
Bacidia medialis (Tuck.) Zahlbr. present study
Bactrospora denticulata (Vain.) Egea & Torrente Boom et al. (2009), present study
Bactrospora jenikii (Vězda) Egea & Torrente present study
Bactrospora myriadea (Fée) Egea & Torrente Boom et al. (2009)
Baculifera intermedioides Marbach Boom et al. (2009)
Blastenia brittonii Zahlbr. Boom et al. (2009) as Caloplaca brittonii (Zahlbr.) ined.
Bogoriella annonacea (Müll. Arg.) Aptroot & Lücking present study
Brigantiaea leucoxantha (Spreng.) R. Sant. & Hafellner present study
Brownliella cinnabarina (Ach.) S.Y. Kondr., Kärnefelt, A.
ell, Elix, J.Kim, A.S.Kondr. & J.-S.Hur
Boom et al. (2009) as Caloplaca cinnabarina (Ach.) Zahlbr.
Buellia boergesenii Imshaug Boom et al. (2009)
Buellia dejungens (Nyl.) Vain. Boom et al. (2009), present study
Buellia griseovirens (Turner & Borrer ex Sm.) Almb. present study
Buellia mamillana (Tuck.) W.A. Weber Boom et al. (2009) as Buellia glaziouana (Kremp.) Müll.
Arg., present study
Buellia posthabita (Nyl.) Zahlbr. Boom et al. (2009)
Bulbothrix scortella (Nyl.) Hale Boom et al. (2009)
Bulbothrix suxa (Stirt.) Hale Boom et al. (2009)
Byssoloma leucoblepharum (Nyl.) Vain. Boom et al. (2009)
Caloplaca diplacia (Ach.) Riddle Boom et al. (2009)
Caloplaca leptozona (Nyl.) Zahlbr. Boom et al. (2009), present study
Caloplaca obscurella (J. Lahm) . Fr. present study
Canoparmelia martinicana (Nyl.) Elix & Hale Boom et al. (2009)
Carbacanthographis triphoroides (M. Wirth & Hale)
Lücking
Boom et al. (2009)

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 77
Species References
Chapsa cinchonarum (Fée) Frisch Boom et al. (2009)
Chrysothrix xanthina (Vain.) Kalb Boom et al. (2009)
Cladonia corymbites Nyl. Boom et al. (2009)
Cladonia didyma (Fée) Vain. Boom et al. (2009)
Cladonia subradiata (Vain.) Sandst. Boom et al. (2009)
Coccocarpia palmicola (Spreng.) Arv. & D.J. Galloway Boom et al. (2009)
Coccocarpia pellita (Ach.) Müll. Arg. Boom et al. (2009)
Coenogonium interpositum Nyl. Boom et al. (2009)
Coenogonium leprieurii (Mont.) Nyl. Boom et al. (2009)
Coenogonium linkii Ehrenb. Boom et al. (2009), present study
Coenogonium saepincola Aptroot, Sipman & Lücking present study
Coenogonium strigosum Rivas Plata, Lücking & Chaves Boom et al. (2009), present study
Coenogonium subdilutum (Malme) Lücking, Aptroot &
Sipman
Boom et al. (2009)
Coniocarpon cinnabarinum DC. Boom et al. (2009) as Arthonia cinnabarina (DC.) Wallr.,
present study
Cratiria lauricassiae (Fée) Marbach Boom et al. (2009)
Crespoa carneopruinata (Zahlbr.) Lendemer & B.P. Hodk. present study
Cresponea ava (Vain.) Egea & Torrente present study
Cresponea leprieurii (Mont.) Egea & Torrente Boom et al. (2009)
Cresponea proximata (Nyl.) Egea & Torrente Boom et al. (2009)
Cryptolechia carneolutea (Tuck.) A. Massal. Boom et al. (2009)
Cryptothecia megalocarpa (Müll. Arg.) R. Sant. Boom et al. (2009)
Cryptothecia punctosorediata Sparrius present study
Cryptothecia striata G. or Boom et al. (2009), present study
Dichosporidium nigrocinctum (Ehrenb.) G. or Boom et al. (2009), present study
Dictyomeridium amylosporum (Vain.) Aptroot, M.P. Nelsen
& Lücking
present study
Diorygma hieroglyphicum (Pers.) Staiger & Kalb Boom et al. (2009), present study
Diorygma poitiaei (Fée) Kalb, Staiger & Elix Boom et al. (2009), present study
Diorygma pruinosum (Eschw.) Kalb, Staiger & Elix present study
Diploschistes actinostomus (Ach.) Zahlbr. Boom et al. (2009)
Diploschistes aeneus (Müll. Arg.) Lumbsch Boom et al. (2009)
Diploschistes prominens (Vain.) Lumbsch Boom et al. (2009)
Dirina paradoxa (Fée) Tehler Boom et al. (2009) as Dirina approximata subsp. hioramii
(B. de Lesd.) Tehler, Tehler et al. (2013)
Dirinaria aegialita (Ach.) B.J. Moore Boom et al. (2009)
Endocarpon pallidulum (Nyl.) Nyl. present study
Enterographa compunctula (Nyl.) Redinger Boom et al. (2009)
Enterographa multilocularis (Müll. Arg.) Sparrius Boom et al. (2009)
Enterographa pallidella (Nyl.) Redinger Boom et al. (2009), present study
Enterographa perez-higaredae Herrera-Camp. & Lücking Boom et al. (2009)
Enterographa sipmanii Sparrius Boom et al. (2009)
Enterographa subserialis (Nyl.) Redinger Boom et al. (2009), present study
Eremothecella microcephalica Sipman Boom et al. (2009)
Fellhanera santessonii Barillas & Lücking Boom et al. (2009)
Fissurina dumastii Fée Boom et al. (2009)
Flakea papillata O.E. Erikss. Boom et al. (2009), present study
Glyphis cicatricosa Ach. Boom et al. (2009)
Glyphis scyphulifera (Ach.) Staiger Boom et al. (2009), present study
Graphis caesiella Vain. Boom et al. (2009), present study
Graphis chondroplaca (Redinger) Lücking Boom et al. (2009)
Graphis cincta (Pers.) Aptroot present study

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
78
Species References
Graphis dendrogramma Nyl. Boom et al. (2009), present study
Graphis furcata Fée Boom et al. (2009)
Graphis glaucescens Fée Boom et al. (2009)
Graphis librata C. Knight present study
Graphis lineola Ach. Boom et al. (2009)
Graphis tenella Ach. Boom et al. (2009)
Graphis tenellula Vain. Boom et al. (2009)
Gyalectidium licinum Müll. Arg. Boom et al. (2009)
Gyalolechia bassiae (Ach.) Søchting, Frödén & Arup ex Ahti Boom et al. (2009) as Caloplaca bassiae (Ach.) Zahlbr.,
present study
Hafellia bahiana (Malme) Sheard Boom et al. (2009)
Hafellia curatellae (Malme) Marbach present study
Herpothallon aurantiacoavum (B. de Lesd.) Aptroot,
Lücking & G.or
Boom et al. (2009)
Heterodermia albicans (Pers.) Swinscow & Krog Boom et al. (2009)
Heterodermia galactophylla (Tuck.) W.L. Culb. Boom et al. (2009)
Heterodermia lutescens (Kurok.) Follmann Boom et al. (2009)
Heterodermia obscurata (Nyl.) Trevis. Boom et al. (2009)
Heterodermia squamulosa (Degel.) W.L. Culb. Boom et al. (2009)
Heterodermia verrucifera (Kurok.) W.A. Weber Boom et al. (2009)
Hyperphyscia adglutinata (Flörke) H. Mayrhofer & Poelt Boom et al. (2009), present study
Hyperphyscia minor (Fée) D.D. Awasthi Boom et al. (2009)
Lathagrium neglectum (Degel.) Otálora, P.M. Jørg. &
Wedin
Boom et al. (2009) as Collema neglectum Degel., present
study
Lecanactis epileuca (Nyl.) Tehler Boom et al. (2009)
Lecanora galactiniza Nyl. Boom et al. (2009)
Lecanora legalloana Elix & Øvstedal Boom et al. (2009), present study
Lecanora leproplaca Zahlbr. present study
Lecanora leprosa Fée Boom et al. (2009), present study
Lecanora prosecha Ach. Boom et al. (2009), present study
Lecanora sulfurescens Fée Boom et al. (2009), present study
Leiorreuma exaltatum (Mont. & Bosch) Staiger Boom et al. (2009)
Lepraria nkii (B. de Lesd.) R.C. Harris present study
Leprocollema novacaledonianum A.L. Sm. present study
Leptogium austroamericanum (Malme) C.W. Dodge Boom et al. (2009)
Leptogium azureum (Sw.) Mont. Boom et al. (2009)
Leptogium cyanescens (Rabenh.) Körb. Boom et al. (2009)
Leptogium marginellum (Sw.) Gray Boom et al. (2009)
Letrouitia domingensis (Pers.) Hafellner & Bellem. Boom et al. (2009), present study
Leucodecton bisporum (Nyl.) Sipman & Lücking Boom et al. (2009)
Leucodecton compunctum (Ach.) A. Massal. Boom et al. (2009)
Loammia gabrielis (Müll. Arg.) Vezda Boom et al. (2009)
Malmidea piperis (Spreng.) Kalb, Rivas Plata & Lumbsch Boom et al. (2009) as Malcolmiella piperis (Spreng.) Kalb
& Lücking, present study
Malmidea psychotrioides (Kalb & Lücking) Kalb, Rivas
Plata & Lumbsch
present study
Malmidea vinosa (Eschw.) Kalb, Rivas Plata & Lumbsch Boom et al. (2009) as Malcolmiella vinosa (Eschw.) Kalb &
Lücking, present study
Mazosia carnea (Eckfelt) Aptroot & M. Cáceres Boom et al. (2009) as Mazosia ocellata (Nyl.) R.C. Harris,
present study
Mazosia phyllosema (Nyl.) Zahlbr. Boom et al. (2009)
Megalaria bengalensis Jagadeesh Ram & Aptroot Hensen (BR)
Melanotrema meiospermum (Nyl.) Frisch Boom et al. (2009)
Microtheliopsis uleana Müll. Arg. present study

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 79
Species References
Mycoporum eschweileri (Müll. Arg.) R.C. Harris present study
Myriostigma candidum Kremp. Boom et al. (2009) as Cryptothecia candida (Kremp.) R.
Sant.: incorrect report
Myriotrema myriotremoides (Nyl.) Hale Boom et al. (2009)
Myriotrema olivaceum Fée Boom et al. (2009)
Nyungwea anguinella (Nyl.) Aptroot Boom et al. (2009) as Enterographa anguinella (Nyl.)
Redinger, present study
Ocellularia depressa (Mont.) Hale Boom et al. (2009)
Ocellularia interposita (Nyl.) Hale Boom et al. (2009)
Ocellularia terebrata (Ach.) Müll. Arg. Boom et al. (2009)
Opegrapha astraea Tuck. present study
Opegrapha lithyrgiza Vain. present study
Opegrapha quintana Redinger present study
Pannaria prolicans Vain. Boom et al. (2009)
Parmotrema crinitum (Ach.) M. Choisy Boom et al. (2009)
Parmotrema endosulphureum (Hillmann) Hale Boom et al. (2009)
Parmotrema praesorediosum (Nyl.) Hale Boom et al. (2009)
Parmotrema tinctorum (Nyl.) Hale Boom et al. (2009)
Parmotrema ultralucens (Krog) Hale Boom et al. (2009)
Peltula bolanderi (Tuck.) Wetmore Boom et al. (2009), present study
Peltula obscurans (Nyl.) Gyeln. Boom et al. (2009), present study
Pertusaria coccopoda Vain. Boom et al. (2009), present study
Pertusaria leioplacella Nyl. Boom et al. (2009)
Pertusaria praetervisa Vain. Boom et al. (2009), present study
Pertusaria texana Müll. Arg. present study
Pertusaria xanthodes Müll. Arg. Boom et al. (2009), present study
Phaeographis crispata Kalb & Matthes-Leicht present study
Phaeographis dendritica (Ach.) Müll. Arg. Boom et al. (2009), present study
Phaeographis intricans (Nyl.) Staiger Boom et al. (2009)
Phaeographis scalpturata (Ach.) Staiger Boom et al. (2009)
Phyllopeltula corticola (Büdel & R. Sant.) Kalb present study
Phyllopsora chlorophaea (Müll. Arg.) Zahlbr. Boom et al. (2009)
Phyllopsora corallina (Eschw.) Müll. Arg. Brako (1991), Boom et al. (2009), present study
Phyllopsora glaucescens Timdal Boom et al. (2009)
Phyllopsora parvifoliella (Nyl.) Müll. Arg. Brako (1991), Boom et al. (2009)
Physcia atrostriata Moberg Boom et al. (2009), present study
Physcia crispa Nyl. Boom et al. (2009)
Physcia erumpens Moberg present study
Physcia integrata Moberg present study
Physcia sinuosa Moberg Boom et al. (2009)
Physcia sorediosa (Vain.) Lynge Boom et al. (2009), present study
Physcia tenuis Moberg Boom et al. (2009)
Platythecium colliculosum (Mont.) Staiger Boom et al. (2009)
Platythecium leiogramma (Nyl.) Staiger Boom et al. (2009)
Polymeridium quinqueseptatum (Nyl.) R.C. Harris Boom et al. (2009)
Porina conspersa Malme present study
Porina epiphylla (Fée) Fée Boom et al. (2009), present study
Porina internigrans (Nyl.) Müll. Arg. Boom et al. (2009), present study
Porina leptalea (Durieu & Mont.) A.L. Sm. present study
Porina mastoidea (Ach.) Müll. Arg. Hensen (BR), Boom et al. (2009)
Porina nitidula Müll. Arg. Boom et al. (2009)
Porina nucula Ach. Boom et al. (2009), present study
Porina octomera (Müll. Arg.) F.Schill. Boom et al. (2009)

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
80
Species References
Porina rubentior (Stirt.) Müll. Arg. present study
Porina tetracerae (Ach.) Müll. Arg. Boom et al. (2009), present study
Porina thaxteri R. Sant. present study
Pseudochapsa dilatata (Müll. Arg.) Parnmen, Lücking &
Lumbsch
Boom et al. (2009) as Chapsa dilatata (Müll. Arg.) Kalb
Pseudopyrenula subgregaria Müll. Arg. Boom et al. (2009)
Pseudopyrenula subnudata Müll. Arg. Harris (1998) as Pseudopyrenula diluta (Fée) Müll. Arg.
var. degenerans Vain, Boom et al. (2009) as Pseudopyrenula
diluta, Aptroot and Lücking (2016)
Pyrenopsis antillarum Vain. present study
Pyrenula adacta Fée present study
Pyrenula astroidea (Fée) R.C. Harris Boom et al. (2009)
Pyrenula bahiana Malme Boom et al. (2009) as Pyrenula concatervans (Nyl.) R.C.
Harris
Pyrenula breutelii (Müll. Arg.) Aptroot Boom et al. (2009) as Pyrenula macularis (Zahlbr.) R.C.
Harris, present study
Pyrenula cinerea Zahlbr. Boom et al. (2009)
Pyrenula cocoes Müll. Arg. Boom et al. (2009), present study
Pyrenula connis (Nyl.) R.C. Harris Boom et al. (2009)
Pyrenula cruenta (Mont.) Vain. present study
Pyrenula dissimulans (Müll. Arg.) R.C. Harris present study
Pyrenula duplicans (Nyl.) Aptroot Boom et al. (2009)
Pyrenula leucostoma Ach. Boom et al. (2009)
Pyrenula mamillana (Ach.) Trevis. Boom et al. (2009) as Pyrenula xyloides (Eschw.) Müll. Arg.
Pyrenula massariospora (Starbäck) R.C. Harris Boom et al. (2009)
Pyrenula microtheca R.C. Harris Boom et al. (2009)
Pyrenula nitidula (Bres.) R.C. Harris Boom et al. (2009), present study
Pyrenula septicollaris (Eschw.) R.C. Harris Boom et al. (2009)
Pyxine cocoes (Sw.) Nyl. Boom et al. (2009), present study
Ramalina anceps Nyl. Boom et al. (2009)
Ramalina complanata (Sw.) Ach. Boom et al. (2009)
Ramalina dendroides (Nyl.) Nyl. Boom et al. (2009)
Ramalina furcellata (Ach.) Zahlbr. Boom et al. (2009)
Ramalina stoersii Sipman present study
Rinodina antillarum Vain. present study
Rinodina colobinoides (Nyl.) Müll. Arg. present study
Rinodina pyxinoides Vain. Boom et al. (2009), present study
Roccella gracilis Bory Boom et al. (2009), Aptroot and Schumm (2011)
Roccellographa circumscripta (Leight.) Ertz & Tehler Boom et al. (2009) as Peterjamesia circumscripta (Taylor)
D. Hawksw.
Sarcographa heteroclita (Mont.) Zahlbr. Boom et al. (2009)
Sarcographa labyrinthica (Ach.) Müll. Arg. Boom et al. (2009)
Sarcographa ramicans (Kremp.) Staiger Boom et al. (2009)
Sarcographa tricosa (Ach.) Müll. Arg. Boom et al. (2009), present study
Sclerophyton elegans Eschw. Boom et al. (2009)
Sclerophyton trinidadense Sparrius Boom et al. (2009)
Sporopodium phyllocharis (Mont.) A. Massal. Boom et al. (2009)
Squamulea subsoluta (Nyl.) Arup, Søchting & Frödén Boom et al. (2009) as Caloplaca subsoluta (Nyl.) Zahlbr.,
present study
Staurolemma dussii (Vain.) P.M. Jørg. & Henssen present study
Stegobolus auberianus (Mont.) Frisch & Kalb Boom et al. (2009)
Stegobolus granulosus (Tuck.) Frisch Boom et al. (2009)
Stegobolus subcavatus (Nyl.) Frisch Boom et al. (2009)
Sticta xanthotropa (Kremp.) D.J. Galloway Hensen (BR), present study

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 81
Species References
Stigmatochroma gerontoides (Stirt.) Marbach present study
Stirtonia neotropica Aptroot Aptroot (2009)
Strigula decipiens (Malme) P.M. McCarthy present study
Strigula macrospora Vain. Boom et al. (2009)
Strigula nemathora Mont. Boom et al. (2009)
Strigula obducta (Müll. Arg.) R.C. Harris Boom et al. (2009)
Strigula phaea (Ach.) R.C. Harris present study
Strigula smaragdula Fr. Boom et al. (2009), present study
Synalissa lichinella Vain. Boom et al. (2009)
Syncesia decussans (Nyl.) Tehler present study
Syncesia farinacea (Fée) Tehler Boom et al. (2009), Sipman (2009)
Syncesia glyphysoides (Fée) Tehler Tehler (1997), Boom et al. (2009), Sipman (2009)
Syncesia graphica (Fr.) Tehler Boom et al. (2009), Sipman (2009)
Syncesia subintegra Sipman Boom et al. (2009), Sipman (2009)
Teloschistes avicans (Sw.) Norman Boom et al. (2009) as Teloschistes avicans var. crocea (Ach.)
Müll. Arg.
alloloma hypoleptum (Nyl.) Staiger Boom et al. (2009)
elenella luridella (Nyl.) H. Mayrhofer present study
elotrema porinoides Mont. & Bosch Boom et al. (2009)
Toninia submexicana B. de Lesd. Boom et al. (2009)
Trapelia coarctata (Sm.) M. Choisy Boom et al. (2009)
Usnea baileyi (Stirt.) Zahlbr. Boom et al. (2009)
Varicellaria velata (Turner) I. Schmitt & Lumbsch Boom et al. (2009)
Verrucaria nigrescens Pers. present study
Xanthoparmelia succedans Elix & J. Johnst. Boom et al. (2009)
Lichenicolous fungus
Dactylospora saxatilis (Schaer.) Hafellner (lichenicolous on
Pertusaria praetervisa)
present study
Additional genera (species uncertain)
Acanthothecis sp. present study
Bacidina sp. present study
Melaspilea sp. (lichenicolous fungus) present study
Psorotichia sp. present study
Stigmidium sp. (lichenicolous fungus) present study
elidium sp. present study
Wetmoreana sp. present study
ever, is closest to the Catapyrenium-Placidiopsis-Verrucaria p.p. (V. caerulea DC., V.
praetermissa (Trevis.) Anzi) clade (Muggia et al. 2010) with sequence identities of
96−97 %. e placement of the Verrucaria cf. dolosa specimen is more dicult to
assess, since its sequence shows lower similarities of 92−94 % to representatives of dif-
ferent Verrucariceae genera, such as Agonimia, Bagliettoa and Verrucaria spp.
Finally, the mtSSU sequence of the Wetmoreana cf. appressa specimen from St.
Eustatius is dicult to interpret, since it matches more closely with sequences of the
Xanthorioideae (sequence identity 97−99 %) than with Teloschistoideae, in which
Wetmoreana is placed (e.g. Arup et al. 2013).
e lichen ora of St. Eustatius can be characterised as lowland, relatively dry
Caribbean. As can be seen from Table 1, most species were found on one substratum

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
82
type, but some are less specialised. Also, there is a marked dierence between the li-
chens of the dierent plots and the three main areas on St. Eustatius (Northern Hills,
central urban area, e Quill). However, the authors refrain from performing statisti-
cal comparisons of the lichen diversity between plots, since the number of plots per
main area diers and is still low and the sampling strategy was devised by specialists of
other organism groups. Nevertheless, the lichen data will be useful for an island-wide,
plot-based comparison of diversity amongst all organism groups sampled during the
2015 inventory.
In Table 2, an updated checklist is presented of the lichens of St. Eustatius, citing
only identied species, but based on all available sources and with their taxonomy
(nomenclature and sometimes species concept) updated where necessary. According
to this list, a total of 263 species are currently known from St. Eustatius. As a side ef-
fect of revising the existing records, one record becomes questionable, viz. Myriostigma
candidum Kremp., which is not known from the Neotropics. It is intended to continue
the exploration of the lichens of this island in the near future.
Acknowledgements
is research was funded by Naturalis Biodiversity Center. Fieldwork was co-organized
by T. van Andel, B. van der Hoorn and J. Miller (Naturalis) and facilitated by the Car-
ibbean Netherlands Science Institute (CNSI), St. Eustatius National Parks (STENA-
PA), DCNA Island Conservationists H. Madden and A. van Zanten and eld guide
C. Gibbs. Students E. Haber, T. Huijts, C. Posthouwer, T. Verheijden, R. Vogel and S.
Zwartsenberg of the Leiden University course Tropical Biodiversity and Field Methods
as well as R. Butôt (Naturalis) contributed to collecting. Student participation was fa-
cilitated by the Alberta Mennega Stichting and the Van Eeden Fonds. Koos Biesmeijer
facilitated the Naturalis Caribbean programme and participated in the eldwork. Mar-
lon Murray and the St. Eustatius Historical Foundation allowed us to sample the urban
plot. Molecular lab work was carried out by R. Butôt. We are indebted to H. Sipman
(Berlin) for providing a copy of the unpublished keys, descriptions and specimen cita-
tion that was the basis for the internet checklist by Boom et al. (2009).
References
Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ (1990) Basic local alignment search
tool. Journal of Molecular Biology 215: 403‒410. https://doi.org/10.1016/S0022-
2836(05)80360-2
Andel T van, Hoorn B van der, Stech M, Bantjes Arostegui S, Miller J (2016) A quantitative
assessment of the vegetation types on the island of St. Eustatius, Dutch Caribbean. Global
Ecology and Conservation 7: 59–69. https://doi.org/10.1016/j.gecco.2016.05.003

An updated checklist of the lichens of St. Eustatius, Netherlands Antilles 83
Aptroot A (2009) A revision of the lichen genus Stirtonia. e Lichenologist 41: 615‒625.
https://doi.org/10.1017/S0024282909990107
Aptroot A, Lücking R (2016) A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypethe-
liales). e Lichenologist 48: 763‒982. https://doi.org/10.1017/S0024282916000487
Aptroot A, Schumm F (2011) Fruticose Roccellaceae, an anatomical-microscopical atlas and
guide with a worldwide key and further notes on some crustose Roccellaceae or similar
lichens. Books on demand, Norderstedt.
Arup U, Søchting U, Frödén P (2013) A new taxonomy of the family Teloschistaceae. Nordic
Journal of Botany 31: 16–83. https://doi.org/10.1111/j.1756-1051.2013.00062.x
Boom BM, Buck WR, Gracie CA, Tulig M (2009) Plants and Lichens of St. Eustatius. http://
sweetgum.nybg.org/st_eustatius/index.php
Brako L (1991) Phyllopsora (Bacidiaceae). Flora Neotropica Monograph 55: 1‒66.
Freitas JA de, Rojer AC, Nijhof BSJ, Debrot AO (2014) A landscape ecological vegetation map
of Sint Eustatius (Lesser Antilles). Den Helder.
Gueidan C, Roux C, Lutzoni F (2007) Using a multigene phylogenetic analysis to assess ge-
neric delineation and character evolution in Verrucariaceae (Verrucariales, Ascomycota).
Mycological Research 111: 1145‒1168. https://doi.org/10.1016/j.mycres.2007.08.010
Harris RC (1989) Working keys to the lichen forming fungi of Puerto Rico. Tropical lichen
workshop, Presented at Catholic University of Puerto Rico. [Published and distributed by
the author]
Harris RC (1995) More Florida lichens. [Published and distributed by the author]
Harris RC (1998) A preliminary revision of Pseudopyrenula Müll. Arg. (lichenized Ascomycetes,
Trypetheliaceae) with a redisposition of the names previously assigned to the genus. In: Glenn
MG, Harris RC, Dirig R, Cole MS (Eds) Lichenographia omsoniana: North American
Lichenology in Honor of John W. omson. Mycotaxon Ltd., Ithaca, New York, 133‒148.
Medeiros ID, Kraichak E, Lücking R, Mangold A, Lumbsch HT (2017) Assembling a Tax-
onomic Monograph of Tribe Wirthiotremateae (Lichenized Ascomycota: Ostropales:
Graphidaceae). Fieldiana Life and Earth Sciences 9: 1‒31. https://doi.org/10.3158/2158-
5520-16.9.1
Miadlikowska J, Kau F, Högnabba F, Oliver JC, Molnár K, Fraker E, Gaya E, Hafellner J,
Hofstetter V, Gueidan C, Otálora MAG, Hodkinson B, Kukwa M, Lücking R, Björk C,
Sipman HJM, Burgaz AR, ell A, Passo A, Myllys L, Goward T, Fernández-Brime S,
Hestmark G, Lendemer J, Lumbsch HT, Schmull M, Schoch CL, Sérusiaux E, Maddison
DR, Arnold AE, Lutzoni F, Stenroos S (2014) A multigene phylogenetic synthesis for the
class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317
genera and 66 families. Molecular Phylogenetics and Evolution 79: 132–168. https://doi.
org/10.1016/j.ympev.2014.04.003
Muggia L, Gueidan C, Grube M (2010) Phylogenetic placement of some morphologically unu-
sual members of Verrucariales. Mycologia 102: 835–846. https://doi.org/10.3852/09-153
Nelsen MP, Lücking R, Grube M, Mbatchou JS, Muggia L, Rivas Plata E, Lumbsch HT (2009)
Unravelling the phylogenetic relationships of lichenised fungi in Dothideomyceta. Studies
in Mycology 64: 135–144. https://doi.org/10.3114/sim.2009.64.07

André Aptroot & Michael Stech / MycoKeys 33: 69–84 (2018)
84
Nelsen MP, Lücking R, Mbatchou JS, Andrew CJ, Spielmann AA, Lumbsch HT (2011) New
insights into relationships of lichen-forming Dothideomycetes. Fungal Diversity 51: 155–
162. https://doi.org/10.1007/s13225-011-0144-7
Orange A, James PW, White FJ (2001) Microchemical Methods for the Identication of Li-
chens. British Lichen Society, London.
Øvstedal DO (2010) e lichens of Guadeloupe, French Antilles. Bergen Museum, Bergen.
Rivas Plata E, Parnmen S, Staiger B, Mangold A, Frisch A, Weerakoon G, Hernández MJE,
Cáceres MES, Kalb K, Sipman HJM, Common RS, Nelsen MP, Lücking R, Lumbsch
HT (2013) A molecular phylogeny of Graphidaceae (Ascomycota, Lecanoromycetes, Os-
tropales) including 428 species. In: Boonpragob K, Crittenden P, Lumbsch HT (Eds) Li-
chens: from genome to ecosystems in a changing world. MycoKeys 6: 55–94. https://doi.
org/10.3897/mycokeys.6.3482
Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA, Chen W, Fungal
Barcoding Consortium (2012) Nuclear ribosomal internal transcribed spacer (ITS) region
as a universal DNA barcode marker for Fungi. Proceedings of the National Academy of
Sciences of the USA 109: 6241–6246. https://doi.org/10.1073/pnas.1117018109
Sipman HJM (2009) e lichen genus Syncesia (Arthoniales) on Saba and St. Eustatius (West
Indies). Opuscula Philolichenum 7: 55‒60.https://doi.org/10.1017/S0953756201004932
Sipman HJM, Aptroot A (2001) Where are the missing lichens? Mycological Research 105:
1433‒1439.
Stoers AL (1956) e vegetation of the Netherlands Antilles. Mededelingen van het Botanisch
Museum van het Herbarium van de Rijksuniversiteit Utrecht 135: 1‒142. [+ 28 pl]
Tehler A (1997) Syncesia (Arthoniales, Euascomycetidae). Flora Neotropica Monograph 74:
1‒48.
Tehler A, Ertz D, Irestadt M (2013) e genus Dirina (Roccellaceae, Arthoniales) revisited. e
Lichenologist 45: 427‒476. https://doi.org/10.1017/S0024282913000121
üs H, Muggia L, Pérez-Ortega S, Favero-Longo SE, Joneson S, O’Brien H, Nelsen MP,
Duque-üs R, Grube M, Friedl T, Brodie J, Andrew CJ, Lücking R, Lutzoni F, Gueidan
C (2011) Revisiting photobiont diversity in the lichen family Verrucariaceae (Ascomy-
cota). European Journal of Phycology 46: 399–415. https://doi.org/10.1080/09670262.
2011.629788
Zoller S, Scheidegger C, Sperisen C (1999) PCR primers for the amplication of mitrochon-
drial small subunit ribosomal DNA of lichen-forming Ascomycetes. e Lichenologist 31:
511‒516. https://doi.org/10.1006/lich.1999.0220