United States Department of Agriculture
Lichens in Puerto Rico: An
Ecosystem Approach
Joel A. Mercado-Díaz, William A. Gould, Grizelle González, and Robert Lücking
Forest
Service
International Institute
of Tropical Forestry
General Technical Report
IITF-GTR-46
April
2015
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Authors
Joel A. Mercado-Díaz is an associate researcher, William A. Gould is a research
ecologist, and Grizelle González is a project leader, U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry, Jardín Botanico
Sur, 1201 Calle Ceiba, San Juan, PR 00926-1119; Robert Lücking is a collections
manager (fungi) and adjunct curator (lichens), Science & Education, The Field
Museum, 1400 South Lake Shore Drive, Chicago, IL 60605-2496.
Cover: Herpothallon rubrocinctum growing on a bamboo shoot (Bambusa vulgaris)
in the Central Mountain Range, Puerto Rico. Photo by J.A. Mercado-Díaz.
Lichens in Puerto Rico: An Ecosystem Approach
Joel A. Mercado-Díaz, William A. Gould, Grizelle González, and Robert Lücking
U.S. Department of Agriculture
Forest Service
International Institute of Tropical Forestry
San Juan, Puerto Rico
General Technical Report IITF-GTR-46
April 2015
Abstract
Mercado-Díaz, Joel A.; Gould, William A.; González, Grizelle; Lücking,
Robert. 2015. Lichens in Puerto Rico: an ecosystem approach. Gen. Tech. Rep.
IITF-GTR-46. San Juan, PR: U.S. Department of Agriculture, Forest Service,
International Institute of Tropical Forestry. 76 p.
This work presents basic information on tropical lichenology. It also describes
general aspects about the ecology and biodiversity of these organisms in eight
forest ecosystems present along an elevational gradient in northeastern Puerto Rico.
These ecosystems consist of elin woodlands, palo colorado, sierra palm, tabonuco,
lowland moist, dry, mangrove, and Pterocarpus forests. Lichen communities are
mainly described in terms of general ecological attributes (e.g., species richness,
common species, etc.). Basic information about the environment and vegetation
found in these forests is also provided. The information presented is supplemented
with ield and microscopic photographs of species and their habitats.
Keywords: Tropical lichens, Puerto Rico, forests, conservation.
ii
Contents
1
1
7
8
11
11
13
14
17
Introduction
What Are Lichens?
Lichenology in Puerto Rico
Lichens in Puerto Rican Ecosystems
Methods
Study Areas
Sampling Scheme
Forest Health and Thelotremoid Lichens
Organization and Other Considerations
18
18
22
26
29
33
36
62
65
68
69
69
70
75
Forests
Elin Woodland Forest
Palo Colorado Forest
Sierra Palm Forest
Tabonuco Forest
Lowland Moist Forest
Dry Forest
Mangrove Forest
Pterocarpus Forest
Concluding Remarks
Acknowledgments
English Equivalents
Literature Cited
Glossary
iii
Preface
This report aims to introduce the reader to general concepts in lichenology with a
particular emphasis in tropical lichens and the study of lichens in Puerto Rico. It
also intends to describe in a broad manner the main characteristics of lichen communities present in different forest ecosystems of the island, and the environmental
conditions in which they thrive. Although care was taken in following basic rules
for responsible scientiic investigation and writing, the information presented here
is aimed to appeal to a more general audience. Moreover, this work presents ideas
that may stimulate the formation of new research questions concerning the lichen
lora of Puerto Rico, and consequently increase interest in and appreciation for the
study of this amazing group of symbiotic organisms.
iv
Lichens in Puerto Rico: An Ecosystem Approach
Introduction
What Are Lichens?
For centuries, lichens have been an intriguing group of organisms for scientists
mostly because of the difficulties that botanists and taxonomists have faced in their
classification (fig. 1). Today, the symbiotic nature of lichens is well understood. In
1982, the International Association of Lichenology described lichens as “an association between a fungus and a “photosynthetic partner,” which results in a stable
thallus with a specific structure.” Most of the fungi that become lichenized belong
to the division Ascomycota. Only 1 percent of lichens have a fungal component
from the division Basidiomycota. The fungal partner in a lichenological association
is better known as the mycobiont, which is also the source for the species name in
lichens. The photosynthetic partner, or photobiont, is usually an alga within the
classes Trebouxiophyceae and Ulvophyceae (Leliaert et al. 2012). Cyanobacteria,
formerly called “bluegreen algae,” also participate in lichenization. In some lichens,
such as species in the genus Stereocaulon, both algae and cyanobacteria may be
found within the same thallus.
Prompted by the symbiotic nature of lichens, scientists have been interested
in studying how the mycobiont and the photobiont behave outside the lichen association. In fact, individual components of lichen species have been successfully
separated in vitro; not surprisingly, their appearance is usually in sharp contrast
to the lichenized form (fig. 2). The mycobiont is usually amorphous and has slow
growth rates; therefore, it is believed that, for most of the mycobionts, lichenization
is obligate (Nash 2008).
Depending on the species, the photobiont may grow successfully in a freeliving state. For example, tropical filamentous algae Trentepohlia is commonly
found free-living (fig. 3), but for the green algae Trebouxia, survival outside
lichenization is rare.
Lichens come in a variety of colors and growth forms and are able to colonize
almost every type of substrate that an environment could provide. Although the
most common colors in lichens are grey, pale green, and yellowish-green, some
species are capable of producing secondary compounds that make them orange,
red, or yellow (Brodo et al. 2001). There are six main types of growth forms in
lichens: crustose, foliose, squamulose, fruticose, dimorphic, and pendulous (fig.
4). The size of these growth forms is highly variable, from less than a few millimeters in some crustose species to more than 2 m in some pendulous Usnea species.
Although the majority of species fit into one of these types, it is not uncommon
to find intermediate forms.
1
Lichens are a symbiotic
association between
a fungus and a photosynthethic partner.
Note: terms in boldfaced type are deined in the glossary (page 75).
1
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 1—Lichens growing on a tree stump in Aibonito, Puerto Rico.
2
Vernon Ahmadjian
A
B
Figure 2— (A) Acarospora fuscata in lichenized
state; (B) (lask on left) the photobiont of A. fuscata;
(lask on right) the mycobiont of A. fuscata.
J.A. Mercado-Díaz
Stephen Sharnoff
Lichens in Puerto Rico: An Ecosystem Approach
Figure 3—Colonies of Trentepohlia algae colonizing the branches of a tree in an elin woodland forest, El Yunque National Forest,
Puerto Rico.
3
D
F
Figure 4—Common growth forms found in lichens. (A) Crustose [Graphis spp.], El Yunque National Forest, Puerto Rico; (B)
Foliose [Sticta spp.], El Yunque National Forest, Puerto Rico; (C) Squamulose [Acarospora socialis], Sierra Bermeja, Puerto Rico;
(D) Fruticose [Cladina sandstede], Maricao, Puerto Rico; (E) Dimorphic [Cladonia subradiata], El Yunque National Forest, Puerto
Rico; (F) Pendulous [Usnea spp.] Maricao, Puerto Rico. Photos by: J.A. Mercado-Díaz, R. Lücking, and A. Cuevas-Padró.
4
R. Lücking
A. Cuevas-Padró
B
A. Cuevas-Padró
E
J.A. Mercado-Díaz
C
R. Lücking
A
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Lichens in Puerto Rico: An Ecosystem Approach
J.A. Mercado-Díaz
Rocks and the bark of trees are usually the preferred colonization substrates for
lichens; however, they have been found growing on all types of natural and artificial substrates including leaves, wood, concrete, glass, plastic, and metal (fig. 5) or
even the carapace of some insects (Lücking et al. 2010). Lichens produce a number
of morphological and anatomical characteristics that their component organisms
would not develop in their free-living forms (Ahmadjian 1993, Brodo et al. 2001,
Nash 2008). Examples of these features include cilia, rhizines, and cyphellae.
These attributes are also used for taxonomic identification.
Figure 5—Several foliose and crustose lichens growing on human-made structures, Jardín Botánico Norte, Río Piedras, Puerto Rico.
In terms of nutrition, the photobiont provides most of the organic nutrients to
the mycobiont. For green algal lichens, carbohydrates are transferred to the mycobiont in the form of sugar alcohols, and for cyanobacterial lichens in the form of
glucose (Nash 2008, Purvis 2000). Benefits gained for the photobiont from lichenization are not so apparently obvious, especially because it has been observed that,
in a lichenized state, some photobionts grow more slowly than in their free-living
form (Nash 2008). In spite of this, studies have found that the mycobiont facilitates
the absorption of mineral nutrients, enhances water uptake to the photobiont, and
protects it from light damage in sun-exposed environments, which suggests that
the photobiont does actually gain some benefits from lichenization.
5
GENERAL TECHNICAL REPORT IITF-GTR-46
Because of the diversity
of microorganisms
that have been found
to inhabit the lichen
thallus, lichens
are often regarded
as “miniature
ecosystems.”
6
It remains undisputed that in the lichenized state, both the mycobiont and the
photobiont are capable of thriving in environments in which they would not survive
individually. This would probably explain why lichens colonize almost every
ecosystem in the world, from tropical rain forests to polar environments (Brodo et
al. 2001, Purvis 2000). In this respect, lichens were initially believed to be more
diverse in temperate ecosystems, a notion supported by several misleading lichen
surveys (Lücking et al. 2011). However, at smaller scales, (e.g., looking at uniform
area sizes of 100 km2, 10 km2, 1 km2, or even 1 ha), lichen species richness clearly
increases toward lower latitudes (Lücking et al. 2011). Estimates of world species
richness of lichens range from 13,500 (Hawksworth and Hill 1984) to 18,000
species (Sipman and Aptroot 2001). Yet, a more recent study has suggested that
the extent of world species richness may be about 28,000, with at least half of the
species (14,000) occurring in the tropics (Lücking et al. 2009a). Compared to their
fungal counterpart, photobiont diversity is less conspicuous, with nearly 54 genera
and about 100 species of algae and cyanobacteria reported (Ahmadjian 1967, Büdel
1992, Frey 2012, Tschermak-Woess 1988). For years lichenologists believed that
lichen photobionts had little influence on thallus morphology. Nevertheless, recent
discoveries, like the continually increasing description of new species forming
photosymbiodemes (Moncada et al. 2013), shows that at least for some groups,
photobionts have a stronger influence on thallus morphology than was previously
supposed.
The symbiotic association of lichens is still a controversial topic among
lichenologists and taxonomists, mostly because the nature of this association does
not fit precisely in any of the known examples of ecological symbiosis. Perhaps
the most appropriate categorization of the lichen symbiosis would be “controlled
parasitism,” because the mycobiont negatively affects the physiognomy and metabolism of the photobiont without killing it (Ahmadjian 1993, Schwendener 1869).
Goward (1994) referred to this relationship as “fungi that have discovered agriculture.” Evidence from a recently described lineage of cyanobacteria (Rhizonema)
strongly supports this view of the lichen symbiosis (Lücking et al. 2009b). However,
recent discoveries have shown that lichens are made of much more than fungal
and photobiont components. A number of endolichenic bacteria and fungi with
distinctive chemical properties have been found in different lichen thalli (Arnold
2007, Cardinale et al. 2006); hence lichens may be more appropriately regarded as
miniature ecosystems (Nash 2008).
Lichens in Puerto Rico: An Ecosystem Approach
Lichenology in Puerto Rico
The field of lichenology in Puerto Rico began in 1820
with the identification of two species from Puerto
Rico in a report by Sprengel (1820), and a subsequent
report titled “Lichenes Portoricenses,” published by
Jean Müller Argoviensis in 1888 (fig. 6). However, not
until the beginning of the 20th century were significant
contributions to Puerto Rican lichenology made by
scientists from the United States and Europe. Some
of these included world-renowned lichenologists of
that era, such as Bruce Fink, Edvard A. Vainio, and
Alexander Zahlbruckner (Mercado-Díaz and SantiagoValentín 2010). Lichenology on the island was nearly
abandoned until 1972 when a monograph of the lichen
genus Ramalina in Puerto Rico and the Caribbean
was presented by Landrón-Concepción (1972), then in
the late 1980s with the publication of Working Keys of
Lichenized Fungi from Puerto Rico (Harris 1989).
Recent efforts promoting lichenology in Puerto
Rico include the research project under which this
report is supported: “Characterizing lichen communities along an elevational gradient in Puerto
Rico: assessing their role as indicators of forest
health, biodiversity and microclimate,” led by Joel A.
Mercado-Díaz and William Gould. Also, a workshop
Figure 6—Cover of the report Lichenes Portorricenses,
published by Jean Müller Argoviensis in 1888.
titled “Taller de Liquenología Tropical: los micro y
macro-líquenes de Puerto Rico,” held in 2011 at the
University of Puerto Rico and organized by Joel A. Mercado-Díaz and Robert
Lücking, resulted in the identification of at least 5 new lichen species and documentation of 25 species previously unknown to Puerto Rico, as well as the training of
14 participants in tropical lichenology (Mercado-Díaz 2011).
Finally, Lichens of Puerto Rico, a website at http://lichensofpuertorico.herbario.
upr.edu/, presents a list of the currently known lichens and lichenicolous fungi
species that occur in Puerto Rico (Mercado-Díaz 2009). For more information
about the history of lichenology in Puerto Rico, refer to Mercado-Díaz and
Santiago-Valentín (2010).
7
GENERAL TECHNICAL REPORT IITF-GTR-46
Lichens in Puerto Rican Ecosystems
By colonizing and
accelerating the
erosion of newly
exposed surfaces,
lichens facilitate the
J.A. Mercado-Díaz
R. Seavey and J. Seavey
formation of soils.
Only occasionally do lichens comprise most of the biomass in forests; however,
they play many ecological roles in these ecosystems (Will-Wolf et al. 2002). Lichens
occupy nearly 8 percent of the Earth’s terrestrial surface (Ahmadjian 1993), and it
is highly likely that their most prominent role in terrestrial ecosystems is related to
their ability to facilitate primary succession. In this respect, lichens are among the
first organisms to colonize newly exposed surfaces, such as rocks formed after the
cooling of recently erupted lava or soil exposed by a landslide. Lichen growth and
expansion result in mechanical and chemical abrasion that speeds up the natural
weathering process of these surfaces and therefore accelerates soil formation.
Colonization over smooth, inhospitable rock surfaces is therefore the most important contribution lichens make to soil formation and consequently to primary
colonization (Brodo et al. 2001).
Lichens are also important agents of soil stabilization and may represent a
source of nitrogen and organic matter in some soils (Brodo et al. 2001, Jones 1988).
Lichen colonization in disturbed and sandy soils may also be important for controlling erosion and the sedimentation of water bodies. As early colonizers, lichens are
critical components of landslide recovery (Walker and Shiels 2013). In Puerto Rico,
the growth of Stereocaulon ramulosum in recently exposed soil resulting from
landslides or the construction and expansion of roads in mountainous areas of the
island may constitute an important biological buffer against this type of disturbance
(fig. 7a). Vast patches of different Cladonia species, like C. subradiata and the
endemic C. robusta, have been observed growing in the siliceous sands of Laguna
Tortuguero in Vega Baja and may contribute to reducing losses of sand via wind or
water movement (fig. 7b).
As in other tropical regions, most of the lichen species in forests in Puerto
Rico are epiphytic, classified either as foliicolous or corticolous. In areas of high
Figure 7—(A) Stereocaulon ramulosum growing in recently exposed soil along a road in El Yunque National Forest,
Puerto Rico; (B) Growth habit of Cladonia subradiata.
8
Lichens in Puerto Rico: An Ecosystem Approach
humidity with adequate light exposure (e.g., elfin woodlands in El Yunque National
Forest), mosses and liverworts grow faster than lichens and therefore colonize most
of the available epiphytic substrate area in these ecosystems. On the contrary, in
forests with lower humidity like the Northern Karst Belt and the Guánica State
Forest, lichens are possibly among the dominant epiphytes, at least in terms of the
number of species.
Lichens are believed to have an important role in water and nutrient cycles in
forest ecosystems (Beckett 1995, Green and Lange 1991, Zotz et al. 1998). For many
species, the lichen thallus is capable of storing up to 800 percent of its dry weight
in water. This water, which is initially absorbed by direct assimilation from water
vapor or rainwater, is released slowly to the environment, thereby influencing the
immediate microclimate. Tree-dwelling lichens absorb a significant amount of
nutrients from rainwater that passes through the canopy over the leaves, and differentially absorb minerals as water flows down to the soil (Brodo et al. 2001, Knops
et al. 1991). Fruticose lichens like Ramalina menziesii have been shown to significantly affect the interception of rainwater and the deposition of water and nutrients
in throughfall collected beneath the canopy in some temperate ecosystems (Knops
et al. 1996). These effects potentially influence the composition and concentration
of nutrients in underlying soils. Although it is likely for this kind of influence on
water and nutrient status in trees and soils to be considerable in some forest areas of
Puerto Rico (especially those where fruticose and foliose lichens are frequent), little
is known about this subject in tropical ecosystems, thus it represents a promising
research topic that deserves further investigation.
It is possible that some of the nitrogen fixed by cyanobacterial lichens leaches
from the lichen thallus or becomes available to neighboring plants when the lichen
dies (Brodo et al. 2001), hence contributing to soil fertilization and consequently
vegetation growth. Although quantitative estimates for this type of fertilization are
scarce, it is not unreasonable to suggest that areas with the highest diversity and
abundance of cyanobacterial lichens are the places where this mode of fertilization
is more likely. One of the places in Puerto Rico where this may happen is in El
Yunque National Forest, which has a relatively high diversity of species with cyanobacterial photobionts such as Stereocaulon, Peltigera, Dictyonema, and Sticta.
Some evidence appears to indicate that lichens are being used as a food source
by several animals and insects in Puerto Rico. For instance, the tracks of what
appears to be a snail possibly feeding on the thalli of several crustose species have
been observed on numerous leaf, bark, and other artificial surfaces in El Yunque
National Forest and other forests of the island1 (fig. 8). Some of these tracks may
1
Mercado-Díaz, J.A. 2014. Personal observation.
9
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 8—Lichen thallus with grazing “tracks” (reddish areas), which are evidence of the feeding
activities of a snail. This lichen is growing on an artiicial substrate (a car door).
belong to Scharammia alta, an endemic snail known from Luquillo and other
places around the island that presumably feeds on encrusting algae and lichens.2
Gastropod grazing and lichen growth have been negatively associated for years;
however, recent studies suggest that gastropods grazing on lichens may represent
important vectors for lichen dispersal (Bosh et al. 2011). On the other hand, the
frequent observation of pyrenocarpous lichens lacking their fruiting bodies
(perithecia) may represent important evidence of direct grazing by insects.
Lichens appear to be important nesting material for some bird species on
the island. For example, fragments of foliose species like Parmotrema and Hypotrachyna are found in almost every hummingbird nest (fig. 9). These pieces of
lichen are most likely part of a camouflage strategy by birds to evade predation of
nestlings. Insects may also be using lichens for camouflage; there is documentation
that the carapace of some tree-dwelling insects in Puerto Rico and Hispaniola
resembles the patterns of lichens growing on leaves or tree trunks.3 4
2
Robinson, D.G.; García-Díaz, W.; Fields, A.; Pérez, J., Tang, T. 2011. The terrestrial malacofauna of Puerto Rico and the U.S. Virgin Islands. Unpublished manuscript. Washington,
DC: U.S. Department of Agriculture, Animal and Plant Health Inspection Service, Plant
Protection and Quarantine. 107 p.
3
4
Mercado-Díaz, J.A. 2014. Personal observation.
Torres-Santana, C. 2014. Personal observation. Director, Arboretum Parque Doña Inés,
Fundación Luis Muñoz Marín, RR 2 Buzón #5, San Juan, PR 00926-9766.
10
José González Díaz
Lichens in Puerto Rico: An Ecosystem Approach
Figure 9—A hummingbird resting in a nest completely covered with foliose lichens.
There are at least 1,181 recognized species of lichenized fungi in Puerto Rico
(Mercado-Díaz 2009); however, a recently published work proposes that the island
may harbor around 1,600 species (Lücking et al. 2009a). Based on the checklist of
lichens and lichenicolous fungi of Puerto Rico (Mercado-Díaz 2009) and preliminary results from a tropical lichenology workshop in the island (Mercado-Díaz
2011), the real diversity of the island may reach about 2,000 species. This is almost
half of the species known for the whole continent of North America.5
Methods
There may be around
2,000 species of
lichens in Puerto Rico,
which represents
nearly half of the
species known for
the continent of North
America.
Study Areas
Most of the information presented in this report comes from intensive lichen
collecting activities conducted by Joel A. Mercado-Díaz between August 2011
and October 2012. Lichens were collected around the periphery of 10 × 10 m2
plots that were established for a study characterizing several biological and environmental parameters along an elevational gradient (fig. 10; González and Luce
2013; González et al. 2013, 2007; Gould et al. 2006; Medina et al. 2013; Ping et al.
2013). Lichens were sampled in 24 sites concentrated in northeastern Puerto Rico
(fig. 11). These plots are representative of eight forest types: elfin woodland, sierra
palm forest, palo colorado forest, tabonuco forest, lowland moist forest, dry forest,
mangrove forest and Pterocarpus forest (Gould et al. 2006). Surveys were made in
three replicate areas for each forest type (24 sites).
5
Lücking, R. 2013. Personal observation.
11
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 10—Sequence of eight forest types along a gradient of elevation, rainfall, and distance from
the coast in northeastern Puerto Rico.
Figure 11—Location of the 24 sampling sites in northeastern Puerto Rico where lichen collecting activities for this study were
completed.
12
Lichens in Puerto Rico: An Ecosystem Approach
Sampling Scheme
Corticolous lichens were collected in sampling sites following a nonquantitative
opportunistic sampling method (Cáceres et al. 2008). Because plots used for the present study are intentionally located far from widely used trails in areas of homogeneous forest cover representative of particular ecosystems (Gould et al. 2006), lichen
collection followed a slightly different approach than was used by Sipman (1996) and
Cáceres et al. (2008), in which lichen collecting efforts were concentrated on main
trails and occasionally at points within the forest. In this respect, information about
lichen communities presented here corresponds to forest ecosystems with continuous
canopy cover, which represent the vast majority of the forested land area of Puerto
Rico (Brandeis and Turner 2013). Nonetheless, to partly compensate for the apparent
underrepresentation of the lichens present in open areas in these forests, lichens were
also collected around canopy openings commonly encountered during collecting
activities. This additional information was used to supplement the descriptions of
lichen communities presented in subsequent sections.
Lichen surveys were similar in practice to the nonquantitative opportunistic
sampling strategy used by Cáceres et al. (2008). Trees were approached both randomly and subjectively depending on visual identification of conspicuous lichen
thalli. Each tree was inspected for 3 to 5 minutes and specimens were collected for
each lichen thallus recognized as a different species in each tree. From 20 to 60 trees
were inspected at each site and between 50 to 100 lichen samples were collected per
locality. Sampling was limited to first 0 to 2.5 m of each tree, and specimens were
also collected from woody vines and areal roots available within this range. Collecting efforts stopped after 10 minutes passed without encountering new species.
This “species saturation” time varied significantly between sites. For example, in
mangrove forests, most of the species were collected before 30 minutes had passed;
however, in palo colorado and tabonuco forests, 2 hours may have passed without
reaching species saturation. Specimens were taken to the Multipurpose Laboratory
at the International Institute of Tropical Forestry and identified using microscopic
techniques and spot tests (Brodo et al. 2001, Orange et al 2010). Additional identification of difficult taxa was done in laboratory facilities of the Field Museum, Chicago. Information about lichens for each forest type was organized and manipulated
in Microsoft Excel® tables.6 PC-ORD™ v. 5.32 (MjM Software Design 1999) was
used to generate ordinations based on species and genus composition (not presented
in this report) that helped in visualizing general patterns of community organization
and revealing factors that may explain variability between forest types.
The use of trade or irm names in this publication is for reader information and does not
imply endorsement by the U.S. Department of Agriculture of any product or service.
6
13
GENERAL TECHNICAL REPORT IITF-GTR-46
The diversity of thelotremoid lichens in a
forest can provide
valuable information
about its conservation
status.
14
Although nonquantitative opportunistic sampling may fail to detect rare,
inconspicuous, sterile, or cryptic species and therefore may result in an underestimation of the real lichen diversity in these forests (Cáceres et al. 2008), the use of
this approach in this study is justified mostly because our primary goal is to provide
a general view of the most conspicuous elements and characteristics of the lichen
flora of different forest types in Puerto Rico. Nevertheless, we strongly endorse
additional efforts that use more rigorous quantitative methods to estimate lichen
diversity in the island. Hopefully, this study will stimulate the formation of new
initiatives that will move further research in that direction.
Forest Health and Thelotremoid Lichens
Lichens could potentially provide valuable information about the conservation
status of tropical forests. A recent study found that the number of morphotypes
of taxa belonging to thelotremoid Graphidaceae is correlated with different
levels of ecological continuity in these forests (Rivas-Plata et al. 2008). An
easy-to-implement sampling protocol, in which thelotremoid lichens are used to
assess disturbance levels of tropical forest sites based on the Index of Ecological
Continuity (IEC), was also proposed (Rivas-Plata et al. 2008). In the context of
their method, IEC could be a measure of morphotype richness depending
on ecological continuity.
The method consists of establishing 500-m transects in forest areas to be studied. The number of thelotremoid morphotypes present in a number of trees along
the transect is recorded. Finally, an IEC is calculated for the forest area studied by
using the following formula: IEC = 100 × n/Nmax. In this formula, n = number of
morphotypes per site, and Nmax = maximum expected number of morphotypes per
site. There are 24 properly described thelotremoid morphotypes (fig. 12). Morphological characters of these morphotypes are described in table 1.
The most critical element of the IEC is the setting of Nmax, which has to be
adjusted to the forest ecosystem (Rivas-Plata et al. 2008). Although Rivas-Plata
et al. (2008) provided Nmax values for most of the forest conditions to be found
between an altitudinal range of 0 to 3500 m in tropical America, the information
was derived from a dataset from Costa Rica, a continental area not affected by the
mass elevation effect, a meteorological phenomenon that lowers altitudinal zones
in island ecosystems (Grubb 1971). Thus, the direct use of Rivas-Plata et al. (2008)
Nmax values in IEC calculations for forests in Puerto Rico would suppose an
underestimation of ecological continuity. Altogether, these observations highlight
the importance of making corrections to Nmax values for insular regions.
A list of the total number of thelotremoid morphotypes found in the three
sampled areas for each forest ecosystem treated in this study is presented at the
Lichens in Puerto Rico: An Ecosystem Approach
Figure 12—Morphotypes of corticolous thelotremoid lichens.
15
GENERAL TECHNICAL REPORT IITF-GTR-46
Table 1—Morphological characteristics of corticolous thelotremoid lichens (modiied from Rivas-Plata
et al. 2008)
Morphotype
Apothecia and reproductive structures
Thallusa
Genera
Chroodiscoid
Leprocarpoid
Cruentodiscoid
Gyrotremoid
Open with recurved lobules
Open with erect lobules
Open with erect lobules, disc pigmented
Open with recurved lobules, disc,
pigmented with concentric rings
Open with erect lobules
Open with erect lobules
Closed with a tiny pore
Immersed with small pore
Immersed with small pore
Prominent with wide pore, in section
with pale walls
Prominent with wide pore, pore with
inner ring
Prominent with wide pore and inner
“mouth”
Prominent with wide pore, in section
with black walls
Prominent with wide pore, pore with
“inger” (columella), in section with
black walls
Large and prominent with small pore,
pore with “inger” (columella), in
section with black walls
Large and prominent with small pore,
pore with “inger” (columella), in
section with black walls and pigment
Immersed with small pore and black
margin, pore with “inger” (columella),
in section with black walls
Prominent with wide pore and black
margin, pore illed with broad “stump”
(columella), in section with black walls
Prominent with wide pore, pore illed
with irregular structures, in section
with pale walls
Prominent with wide pore, pore illed
with irregular structures, in section
with black walls
Immersed with linear slit, slit illed with
irregular structures, in section with
black walls
Apothecia lacking, with isidia
Apothecia lacking, with schizidia
Apothecia lacking, with soralia
Smooth, ± shiny
Mealy, ± matte
Smooth, ± shiny
Smooth, ± shiny
Acanthotrema, Chapsa
Chapsa
Chapsa
Gyrotrema
Rough, with crystals
Rough, with crystals
Rough, with crystals
Rough, with crystals
Smooth, ± shiny
Smooth, ± shiny
Reimnitzia
Leucodecton glaucescens
Leucodecton
Leptotrema
Myriotrema (glaucopallens group)
Myriotrema
Smooth, ± shiny
Myriotrema, Thelotrema
Smooth, ± shiny
Thelotrema
Smooth, ± shiny
Ampliotrema, Ocellularia
Smooth, ± shiny
Ocellularia
Smooth, ± shiny
Ocellularia (praestans group)
Smooth, ± shiny
Ocellularia (rhodostroma group)
Smooth, ± shiny
Clandestinotrema
Smooth, ± shiny
Smooth, ± shiny
Melanotrema, Ocellularia,
Clandestinotrema,
Trinathotrema
Stegobolus (wrightii group)
Smooth, ± shiny
Stegobolus
Smooth, ± shiny
Redingeria, Stegobolus
Smooth, ± shiny
Smooth, ± shiny
Smooth, ± shiny
Myriotrema, Ocellularia
Stegobolus
Myriotrema, Ocellularia
Reimnitzioid
Glaucescentoid
Leucodectonoid
Leptotremoid
Myriotremoid
Glaucophaenoid
Annulotremoid
Thelotremoid
Ampliotremoid
Ocellularioid
Praestantoid
Rhodostromoid
Tenuitremoid
Melanotremoid
Pallidostegoboloid
Stegoboloid
Redingerioid
Isidiotremoid
Schizotremoid
Sorediotremoid
a
The symbol “±” is used here to mean “more or less.”
16
Lichens in Puerto Rico: An Ecosystem Approach
end of each “Lichen Community” subsection. This information should aid in the
correct determination of appropriate Nmax values to be used in future IEC calculations for Puerto Rican forests. Suitable Nmax values are fundamental for the
effective use of the methods proposed by Rivas-Plata et al. (2008) and simplified by
Mercado-Díaz et al. (N.d).7
Organization and Other Considerations
Each section describing forest types in this report has three main subsections.
The first, Environment, offers a general description of the physical environment
of each forest type. This includes information on mean annual temperature, mean
annual precipitation, and elevation range in which this type of forest is found
on the island. The second section, Vegetation, aims to briefly illustrate the main
aspects of vegetation composition and structure in these forests. This information
is important because vegetation surfaces ultimately provide the substrate on which
most of the lichen species present in the forest understory grow. The third section
is titled Lichen Communities; its main purpose is to describe general characteristics of lichen communities in these forests. Information presented in this section
include (1) observations about the abundance and richness of the different species
and genera; (2) descriptions of commonly found growth forms, genera and species,
(3) accounts of species that were found only in some of these forests; and (4) other
ecological observations concerning the interaction between lichen communities and
particular forest elements, e.g., lichen species occurrences in particular tree species.
Each section is supplemented with photographs that illustrate common views of
forest types and other vegetation attributes found in these forests.
Bound into the center of this volume is a set of species plates, which contain
images of the lichen species mentioned in the “Lichen Communities” subsection
of each forest type. The species names followed by an asterisk (*) indicate that the
photograph was taken in the field with a high-resolution digital camera. The rest
of the images were taken with a digital camera mounted on a stereomicroscope.
To detect the detailed morphological characters shown in these images, and consequently facilitate field identification, the use of a 10× hand lens is recommended. It
should be noted that photographs taken with the stereomicroscope are of herbarium
specimens. Specimens in herbarium conditions are dry and usually have less vivid
colors than species growing in their natural habitats; therefore, care should be taken
when using these images for identification purposes.
7
Mercado-Díaz, J.A.; Gould, W.A.; González, G.; Torres-Santana, C. [N.d]. Using lichens
as indicators of forest health in Puerto Rico. Manuscript in preparation. On ile with: J.A.
Mercado-Díaz, U.S. Department of Agriculture, Forest Service, International Institute of
Tropical Forestry, Jardin Botanico Sur, 1201 Calle Ceiba, San Juan, PR 00926-1119.
17
GENERAL TECHNICAL REPORT IITF-GTR-46
Note that although lichens were surveyed exhaustively at each site, the number
of species reported for each ecosystem is not thought to be definitive. Additional
lichen surveys in other areas will likely increase the number of species for a particular ecosystem. Species richness is reported with the main purpose of offering
a general view of the diversity of lichens that a person sampling a similar area of a
particular forest type might find. Likewise, the information related to the species
and genera found exclusively in a particular ecosystem is tentative. Subsequent field
surveys in other forests should likely reveal the presence of many of these species
and genera elsewhere.
Some information presented throughout this report concerning lichen species
occurrences in different areas of Puerto Rico and general notes about the island’s
lichen diversity are personal observations made by Joel A. Mercado-Díaz.
Forests
Puerto Rico has an extremely rich diversity of forests. This diversity could be
explained by the role that a number of environmental and historical factors have
had on the ecological processes shaping these ecosystems (Lugo 2005). There are
numerous schemes that have been used to describe these forests; however, many
of them lack a comprehensive description of their ecology (Lugo 2005).
The following sections aim to broaden the ecological knowledge of eight forest
ecosystems of Puerto Rico by presenting information on their abiotic characteristics, vegetation, and lichen communities. These ecosystems are elfin woodland,
palo colorado forest, sierra palm forest (fig. 13), tabonuco forest, lowland moist
forest, dry forest, mangrove forest, and Pterocarpus forest. Many ecological aspects
of these forests and the sampled sites have been described previously (González
et al. 2007, Gould et al. 2006) and are the object of ongoing ecological research
(González et al. 2013).
Elfin Woodland
Environment—
The elfin woodland forest type (fig. 14) occurs within the two wettest Holdridge life
zones of Puerto Rico, the lower montane wet and rain forests. The elfin woodland
has an approximate mean annual precipitation of 3908 mm and a mean annual temperature of 19.5 °C (Gould et al. 2006). This forest type occupies very little area in
Puerto Rico, occurring in a single crescent-shaped band on the windward faces of
the Luquillo Mountains (Miller and Lugo 2009). The approximate elevation of this
forest type within El Yunque National Forest is 1010 m (Gould et al. 2006). Owing
to the superabundance of water, soils are at field capacity much of the year and the
18
J.A. Mercado-Díaz
Lichens in Puerto Rico: An Ecosystem Approach
J.A. Mercado-Díaz
Figure 13—A ire ant walking over a gelatinous-foliose lichen of the genus Leptogium in a sierra
palm forest, El Yunque National Forest, Puerto Rico.
Figure 14—The understory of an elin woodland forest in El Yunque National Forest, Puerto Rico.
19
GENERAL TECHNICAL REPORT IITF-GTR-46
total annual runoff (3400 mm) is more than twice the annual rainfall input received
by most areas of the world (Miller and Lugo 2009). Because the elfin woodland
ecosystem of Puerto Rico lies entirely within El Yunque National Forest, the whole
forest type is legally protected.
J.A. Mercado-Díaz
Vegetation—
The elfin woodland is a closed broad-leaved evergreen forest that generally occurs
in exposed areas and ridges and is characterized by twisted, gnarled trees that are
less than 7 m tall, with small diameters, a large number of stems per unit area,
and extremely slow growth rates (Gould et al. 2006, Miller and Lugo 2009) (fig.
15). All trees have sclerophyllous leaves that tend to be grouped near the ends of
the branches (Miller and Lugo 2009). Common trees found in this forest include
Ocotea spathulata Mez., Eugenia borinquensis Britton, Tabebuia rigida Urb.,
Magnolia splendens Urb., and Clusia rosea Jacq.; whereas rare endemic tree
species include Ardisia luquillensis (Britton) Alain and Ilex obcordata Sw. var.
obcordata (Gould et al. 2006, Miller and Lugo 2009). This forest type is also
characterized by its superabundance of nonvascular epiphytes, particularly mosses
and liverworts, and the presence of lianas like the endemic Marcgravia sintenisii
Urb. and Mikania cordifolia (L. f.) Willd. (Gould et al. 2006, Lugo 2005). Refer to
table 2 for common names of plant species identified in this report.
Figure 15—Species of Clandestinotrema growing on an upright branch of a tree in an elin woodland
forest, El Yunque National Forest, Puerto Rico.
20
Lichens in Puerto Rico: An Ecosystem Approach
Table 2—Common names of plant species identiied in this report
Scientiic name
Common name in English
Acrostichum aureum
Alchornea latifolia
Ardisia luquillensis
Avicennia germinans
Bourreria succulenta
Bursera simaruba
Byrsonima wadsworthii
Cecropia schreberiana
Clusia rosea
Conocarpus erectus
Cordia borinquensis
Cyathea arborea
Cyrilla racemilora
Dacryodes excelsa
Erythroxylum brevipes
Eugenia bilora
Eugenia borinquensis
Faramea occidentalis
Henriettea squamulosum
Hippocratea volubilis
Hymenaea courbaril
Ilex obcordata var. obcordata
Inga laurina
Ipomoea triloba
Laguncularia racemosa
Magnolia splendens
Manilkara bidentata
Marcgravia sintenisii
Mikania cordifolia
Neea buxifolia
Ocotea leucoxylon
Ocotea spathulata
Paullinia pinnata
Prestoea acuminata var. montana
Pterocarpus oficinalis
Quadrella cynophallophora
Rhizophora mangle
Rondeletia portoricensis
Sloanea berteroana
Spathodea campanulata
Tabebuia rigida
Tetragastris balsamifera
Golden leather fern
Mountain marlberry
Black mangrove
Bodywood
Gumbo limbo
Pumpwood
Scotch attorney
Button mangrove
West Indian tree fern
Swamp titi
Candle tree
Brisselet
Blackrodwood
False coffee
Medicine vine
Stinkingtoe
Sacky sac bean
Little bell
White mangrove
Bulletwood
Shingleplant
Florida Keys hempvine
Saltwood
Loblolly sweetwood
Bread and cheese
Sierran palm
Dragonsblood tree
Jamaican caper
Red mangrove
Bullwood
African tulip tree
Common name in Spanish
(Puerto Rican usage)
Palmita del rio
Achiotillo
Mameyuelo
Mangle negro
Palo de vaca
Almácigo
Almendrillo
Yagrumo hembra
Cupey
Mangle boton
Capa cimarron
Helecho arboreo
Palo colorado
Tabonuco
Rocío
Hoja menuda
Guayabota de sierra
Cafeillo
Camasey jusillo
Bejuco prieto
Algarrobo
Cuero de sapo
Guama
Bejuquillo de puerco
Mangle blanco
Laurel sabino
Ausubo
Pegapalma
Guaco
Laurel geo
Cabalonga cimarrona
Bejuco de costilla
Palma de sierra
Palo de pollo
Burro prieto
Mangle rojo
Juan Tomás
Motillo
Tulipán africano
Roble de sierra
Masa
21
GENERAL TECHNICAL REPORT IITF-GTR-46
Lichen communities—
Elfin woodlands are characterized for having low species richness of corticolous
lichens if compared to other forest types in El Yunque National Forest. Only 33
species have been recorded in sampled areas of this forest type. Nonetheless, these
forests have the highest number of species within the genus Clandestinotrema, and
a good representation of taxa belonging to the genus Chapsa. As mentioned before,
the presence of Clandestinotrema species in Neotropical forests is positively correlated with elevation (Rivas-Plata et al. 2008), which agrees with the observations
reported here. Competition for space appears to be a major constraint for lichen
colonization in the elfin woodland as bryophytes are more adapted to the highmoisture conditions found in this ecosystem.
Although most of the lichen flora in this forest adopts a crustose growth habit,
it is not uncommon to find species with fruticose or squamulose growth forms, like
Cladia aggregata (plate 1A) and Cladonia spp., growing on tree branches in the
understory of this forest type. Foliose lichens are scarce and, if present, are most
likely cyanobacterial such as species of Leptogium. Dictyonema sericeum (plate
1B) and Cyphellostereum (plate 1C) are examples of a filamentous basidiolichens
present in these forests. Foliicoulous lichens are uncommon in the understory of
the elfin woodland forest and nearly all species to be found are close to canopy
openings.
Some of the most common lichen species found in this forest type include
Chapsa sp. A (plate 1D), Chapsa thallotrema (plates 1E and 1F), Clandestinotrema
leucomelaenum (plate 1G), Dictyonema sericeum, Graphis adpressa (plate 1H),
and Ocellularia rhodostroma (plate 2A). Several species have only been found
in sampled areas of this forest type. Some of these are Chapsa sp. A, Chapsa sp.
B (plate 2B), Cladia aggregata, Clandestinotrema analorenae (plate 2C), Clandestinotrema stylothecium (plate 2D), Fissurina crassilabra (plate 2E), Fissurina
incrustans (plate 2F), and Graphis adpressa.
At least six morphotypes of thelotremoid lichens have been found in elfin
woodland forests in El Yunque National Forest (Rivas-Plata et al. 2008). These are
the encountered morphotypes and their representative species:
Leprocarpoid: Chapsa dissuta
Sorediotermoid: Chapsa thallotrema
Tenuitremoid: Clandestinotrema analorenae
Melanotremoid: Clandestinotrema leucomelaenum, C. stylothecium
Rhodostromoid: Ocellularia rhodostroma
Thelotremoid: Thelotrema porinoides
22
Lichens in Puerto Rico: An Ecosystem Approach
An interesting fact from a lichenological perspective is the common sighting of
free-living Trentepohlia spp., a filamentous green algae hanging from tree branches
in the elfin woodland forest (fig. 3). Trentepohlia is the most common photobiont
participating in lichenological associations in tropical ecosystems and is recognized
by its filamentous growth habit and distinctive orange color, which is caused by the
presence of carotenoid pigments that mask the green pigments of chlorophyll. In
lichenized state, Trentepohlia loses its ability to produce carotenoids. Trentepohlia
filaments may also be found in other areas around the island, particularly under
humid and illuminated conditions.
Palo Colorado Forest
J.A. Mercado-Díaz
Environment—
The palo colorado forests in Puerto Rico (fig. 16) are considered “mountain wetlands” (Frangi 1983), and according to Holdridge’s system (Ewel and Whitmore
1973), this forest lies within the subtropical lower montane wet forest (Gould et al.
2006, Lugo 2005, Miller and Lugo 2009). It occurs in both the eastern and central
parts of the island up to the summits of some mountains that are above 1000 m
(Miller and Lugo 2009). The palo colorado forest has a high cloud cover, low illumination, extreme soil saturation, and a high water table (Lugo 2005). Mean annual
temperature is about 23 °C, and mean annual precipitation reaches about 2932 mm
(Gould et al. 2006). This forest covers only about 1.2 percent of the island’s total
Figure 16—The understory of a palo colorado forest in El Yunque National Forest, Puerto Rico.
23
GENERAL TECHNICAL REPORT IITF-GTR-46
surface area (Weaver 1987), but encompasses nearly 17 percent of the Luquillo
Mountains, and grows at elevations higher than 600 m. In El Yunque National
Forest, these forests are considered cloud forests precisely because cloud condensation commonly starts at this elevation (Lugo 2005).
J.A. Mercado-Díaz
Vegetation—
The palo colorado forest is mostly a closed broad-leaved evergreen forest that
gets its name from the presence of its common tree Cyrilla racemiflora L. (Gould
et al. 2006) (fig. 17). This forest corresponds to mature vegetation of the zonal
association in the subtropical lower montane wet forest (Miller and Lugo 2009).
It is a low-statured forest (3 to 9 m) and, in some places, trees are considerably
dispersed owing either to high area occupancy of large-diameter trees or because
environmental conditions limit their density (Lugo 2005). Although the palo
colorado forest is a closed forest, canopy openings allow light penetration. Clouds
continuously penetrate the understory, which is considerably open (Lugo 2005).
The palo colorado forest is poorer in plant species than the adjacent tabonuco
forests (Miller and Lugo 2009). Common tree species include Cyathea arborea (L.)
Sm., Prestoea acuminata (Willd.) H.E. Moore var. montana (Graham) A. Hend. &
G. Galeano, Ocotea spathulata, Alchornea latifolia Sw., among others (Gould et al.
2006, Lugo 2005). A study classifying vegetation in these forests found about 14
endemic tree species and six rare native species in the palo colorado forests of El
Yunque National Forest (Gould et al. 2006). Refer to table 2 for common names of
plant species.
A
B
Figure 17—Lichens are commonly found on aerial roots of trees in the palo colorado forest in El
Yunque National Forest. In this photo: (A) Coenogonium linkii, and (B) Myeloconis guyanensis.
24
Lichens in Puerto Rico: An Ecosystem Approach
Plates
Species photographs in plates are organized in the order in which
they are mentioned in the text. Species names followed by an asterisk (*) indicate that the photograph was taken in the ield with a
high-resolution digital camera. The remainder of the images were
taken with a digital camera mounted on a stereomicroscope.
25
Plate 1
(A) Cladia aggregata
(B) Dictyonema sericeum*
(growing on moss)
(C) Cyphyellostereum phyllogenum
(D) Chapsa sp. A
(E) Chapsa thallotrema*
(F) Chapsa thallotrema* (detail)
(G) Clandestinotrema leucomelaenum
(H) Graphis adpressa.
Plate 2
(A) Ocellularia rhodostroma
(B) Chapsa sp. B
(C) Clandestinotrema analorenae
(D) Clandestinotrema stylothecium
(E) Fissurina crassilabra
(F) Fissurina incrustans
(G) Coenogonium linkii*
(H) Herpothallon aurantiacolavum* (left)
and Herpothallon granulare* (right).
Plate 3
(A) Herpothallon aurantiacolavum
(B) Herpothallon granulare
(C) Dichosporidium nigrocinctum*
(D) Dichosporidium nigrocinctum
(fruiting bodies)
(E) Graphis rhizocola*
(F) Malmidea nigromarginata
(G) Myeloconis guyanensis*
(H) Ocellularia praestans*
Plate 4
(A) Ocellularia praestans (detail)
(B) Porina nucula
(C) Ampliotrema rimosum*
(D) Arthonia sp. (yellow margin)
(E) Chapsa alborosella
(F) Clandestinotrema cf. tenue
(G) Coenogonium nepalense
(H) Graphis dimidiata.
Plate 5
(A) Malmidea furfurosa
(B) Malmidea vinosa
(C) Paratopeliopsis caraibica
(D) Ocellularia aff. perforata
(E) Ocellularia cavata
(F) Ocellularia crocea
(G) Ocellularia dolichotata
(H) Ocellularia rhabdospora.
Plate 6
(A) Platythecium grammitis
(B) Sticta sp.*
(C) Leptogium azureum*
(D) Leptogium denticulatum*
(E) Graphis duplicata
(F) Porina subpungens*
(G) Thelotrema porinoides
(H) Arthonia complanata*.
Plate 7
(A) Graphis proserpens*
(B) Fissurina sp. nov.*
(C) Fissurina sp. nov. (detail)
(D) Megalotremis lateralis
(E) Pyrenula aff. fetivica*
(F) Sticta beauvoisii*
(G) Porina scabrida
(H) Mazosia endonigra.
26
Plate 8
(A) Herpothallon rubrocinctum*
(B) Phyllopsora buettneri
(C) Phyllopsora corallina*
(D) Usnea sp.*
(E) Stereocaulon ramulosum*
(F) Cladonia subradiata *
(G) Arthonia platygraphidea
(H) Arthothelium distendens.
Plate 9
(A) Ocellularia interposita
(B) Pyrenula macrocarpa
(C) Pyrenula massariospora*
(D) Heterodermia speciosa
(E) Acanthotrema alboisidiatum
(F) Arthonia aff. bessalis
(G) Arthothelium sp. nov
(H) Pseudochapsa dilatata.
Plate 10
(A) Chapsa elabens
(B) Eugeniella sp. nov
(C) Herpothalon aff. pustulata*
(D) Letrouitia vulpina
(E) Leucodecton compuctellum
(F) Malmidea amazonica
(G) Mazosia verrucosa
(H) Megalotremis infernalis.
Plate 11
(A) Ocellularia aff. cavata
(B) Ocellularia ascidiodea
(C) Ocellularia mordenii
(D) Borinquenotrema soredicarpum
(E) Porina americana
(F) Rhabdodiscus emersellus
(G) Wirthiotrema glaucopallens
(H) Coenogonium leprieurii*.
Plate 12
(A) Physcia atrostriata*
(B) Parmotrema praesorediosum*
(C) Dirinaria purpurascens
(D) Ramalina complanata*
(E) Ramalina peruviana*
(F) Cresponea melanocheiloides*
(G) Cresponea melanocheiloides (detail)
(H) Cryptothecia striata*.
Plate 13
(A) Diorygma poiteai*
(B) Diorygma poiteai (detail)
(C) Hemithecium balbisii*
(D) Hemithecium balbisii (detail)
(E) Graphis glaucescens*
(F) Graphis glaucescens (detail)
(G) Malmidea piperis*
(H) Malmidea piperis (detail).
Plate 14
(A) Opegrapha bonplandii
(B) Opegrapha dekeselii
(C) Phaeographis brasiliensis
(D) Porina distans*
(E) Porina distans (detail)
(F) Porina conspersa
(G) Pyrenula mamillana
(H) Fissurina subnitens (between
bark issures)*.
Plate 15
(A) Fissurina subnitens (detail)
(B) Anisomeridium subprostans
(C) Arthonia aff. pyrrholiza
(D) Arthonia bessalis
(E) Dyplolabia afzelii
(F) Letrouitia domingensis*
(G) Letrouitia domingensis (detail)
(H) Leucodecton bisporum*.
Plate 16
(A) Leucodecton bisporum (detail)
(B) Mazosia carnea
(C) Melanotrema platystomum
(D) Monoblastia borinquensis
(E) Porina tetracerae
(F) Psoroglaena cubensis
(G) Strigula phaea
(H) Trinathotrema stictideum.
Plate 17
(A) Physcia crispa
(B) Physcia sorediosa
(C) Pyxine berteriana
(D) Pyxine eschweileri
(E) Anisomeridium biforme
(F) Arthonia portoricensis
(G) Arthopyrenia majuscula
(H) Bactrospora denticulata.
Plate 18
(A) Graphis furcata
(B) Graphis tenella
(C) Opegrapha cf. varia
(D) Pyrenula quassiaecola
(E) Arthonia antillarum
(F) Arthonia caribaea
(G) Celothelium dominicanum
(H) Coenogonium aurantiacum.
Plate 19
(A) Coenogonium borinquense
(B) Enterographa multilocularis
(C) Fissurina tachygrapha
(D) Leucodecton occultum
(E) Opegrapha astraea
(F) Phaeographis inusta
(between lenticels)
(G) Phaeographis inusta (detail)
(H) Pyrenula thelomorpha.
Plate 20
(A) Ramonia rappii
(B) Ramonia valenzuelana
(C) Parmotrema endosulphureum
(D) Parmotrema endosulphureum
(isidia and yellow medulla)
(E) Physcia erumpens
(F) Pyrenula cerina*
(G) Pyrenula cerina (detail)
(H) Bactrospora myriadea.
Plate 21
(A) Enterographa aff. anguinella
(B) Pyrenula cocoes
(C) Pyrenula ochraceolava
(D) Pyrenula microcarpa
(E) Herpothallon minimum
(F) Porina curtula
(G) Arthonia aff. rubella
(H) Coenogonium dimorphicum.
Plate 22
(A) Coenogonium portoricense
(B) Crocynia gossypina
(C) Fissurina adscribens
(D) Graphis farinulenta
(E) Diorygma poiteai* (left) and
Graphis farinulenta* (right)
(F) Lecanographa lyncea
(G) Malmidea aff. coralliformis
(H) Schismatomma rappii.
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
R. Lücking
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 1
H
Plate 1—(A) Cladia aggregata; (B) Dictyonema sericeum* (growing on moss); (C) Cyphyellostereum phyllogenum; (D) Chapsa sp. A;
(E) Chapsa thallotrema*; (F) Chapsa thallotrema* (detail); (G) Clandestinotrema leucomelaenum; (H) Graphis adpressa.
27
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
R. Lücking
Plate 2
B
D
F
H
Plate 2—(A) Ocellularia rhodostroma; (B) Chapsa sp. B; (C) Clandestinotrema analorenae; (D) Clandestinotrema stylothecium;
(E) Fissurina crassilabra; (F) Fissurina incrustans; (G) Coenogonium linkii*; (H) Herpothallon aurantiacolavum* (left) and
Herpothallon granulare* (right).
28
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 3
H
Plate 3—(A) Herpothallon aurantiacolavum; (B) Herpothallon granulare; (C) Dichosporidium nigrocinctum*; (D) Dichosporidium
nigrocinctum (fruiting bodies); (E) Graphis rhizocola*; (F) Malmidea nigromarginata; (G) Myeloconis guyanensis*; (H) Ocellularia
praestans*.
29
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 4
B
D
F
H
Plate 4—(A) Ocellularia praestans (detail); (B) Porina nucula; (C) Ampliotrema rimosum*; (D) Arthonia sp. (yellow margin);
(E) Chapsa alborosella; (F) Clandestinotrema cf. tenue; (G) Coenogonium nepalense; (H) Graphis dimidiata.
30
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 5
H
Plate 5—(A) Malmidea furfurosa; (B) Malmidea vinosa; (C) Paratopeliopsis caraibica; (D) Ocellularia aff. perforata; (E) Ocellularia
cavata; (F) Ocellularia crocea; (G) Ocellularia dolichotata; (H) Ocellularia rhabdospora.
31
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 6
B
D
F
H
Plate 6—(A) Platythecium grammitis; (B) Sticta sp*; (C) Leptogium azureum*; (D) Leptogium denticulatum*; (E) Graphis duplicata;
(F) Porina subpungens*; (G) Thelotrema porinoides; (H) Arthonia complanata*.
32
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 7
H
Plate 7—(A) Graphis proserpens*; (B) Fissurina sp. nov.*; (C) Fissurina sp. nov. (detail); (D) Megalotremis lateralis; (E) Pyrenula aff.
fetivica*; (F) Sticta beauvoisii*; (G) Porina scabrida; (H) Mazosia endonigra.
33
E
G
J.A. Mercado-Díaz
A. Cuevas-Padró
C
R. Lücking
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 8
B
D
F
H
Plate 8—(A) Herpothallon rubrocinctum*; (B) Phyllopsora buettneri; (C) Phyllopsora corallina*; (D) Usnea sp.*; (E) Stereocaulon
ramulosum*; (F) Cladonia subradiata *; (G) Arthonia platygraphidea; (H) Arthothelium distendens.
34
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 9
H
Plate 9—(A) Ocellularia interposita; (B) Pyrenula macrocarpa; (C) Pyrenula massariospora*; (D) Heterodermia speciosa;
(E) Acanthotrema alboisidiatum; (F) Arthonia aff. bessalis; (G) Arthothelium sp. nov; (H) Pseudochapsa dilatata.
35
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 10
B
D
F
H
Plate 10—(A) Chapsa elabens; (B) Eugeniella sp. nov; (C) Herpothalon aff. pustulata*; (D) Letrouitia vulpina; (E) Leucodecton
compuctellum; (F) Malmidea amazonica; (G) Mazosia verrucosa; (H) Megalotremis infernalis.
36
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 11
H
Plate 11—(A) Ocellularia aff. cavata; (B) Ocellularia ascidiodea; (C) Ocellularia mordenii; (D) Borinquenotrema soredicarpum;
(E) Porina americana; (F) Rhabdodiscus emersellus; (G) Wirthiotrema glaucopallens; (H) Coenogonium leprieurii*.
37
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 12
B
D
F
H
Plate 12—(A) Physcia atrostriata*; (B) Parmotrema praesorediosum*; (C) Dirinaria purpurascens; (D) Ramalina complanata*;
(E) Ramalina peruviana*; (F) Cresponea melanocheiloides*; (G) Cresponea melanocheiloides (detail); (H) Cryptothecia striata*.
38
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 13
H
Plate 13—(A) Diorygma poiteai*; (B) Diorygma poiteai (detail); (C) Hemithecium balbisii*; (D) Hemithecium balbisii (detail);
(E) Graphis glaucescens*; (F) Graphis glaucescens (detail); (G) Malmidea piperis*; (H) Malmidea piperis (detail).
39
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 14
B
D
F
H
Plate 14—(A) Opegrapha bonplandii; (B) Opegrapha dekeselii; (C) Phaeographis brasiliensis; (D) Porina distans*; (E) Porina distans
(detail); (F) Porina conspersa; (G) Pyrenula mamillana; (H) Fissurina subnitens (between bark issures)*.
40
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 15
H
Plate 15—(A) Fissurina subnitens (detail); (B) Anisomeridium subprostans; (C) Arthonia aff. pyrrholiza; (D) Arthonia bessalis;
(E) Dyplolabia afzelii; (F) Letrouitia domingensis*; (G) Letrouitia domingensis (detail); (H) Leucodecton bisporum*.
41
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 16
B
D
F
H
Plate 16—(A) Leucodecton bisporum (detail); (B) Mazosia carnea; (C) Melanotrema platystomum; (D) Monoblastia borinquensis;
(E) Porina tetracerae; (F) Psoroglaena cubensis; (G) Strigula phaea; (H) Trinathotrema stictideum.
42
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 17
H
Plate 17—(A) Physcia crispa; (B) Physcia sorediosa; (C) Pyxine berteriana; (D) Pyxine eschweileri; (E) Anisomeridium biforme;
(F) Arthonia portoricensis; (G) Arthopyrenia majuscula; (H) Bactrospora denticulata.
43
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
R. Lücking
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 18
B
D
F
H
Plate 18—(A) Graphis furcata; (B) Graphis tenella; (C) Opegrapha cf. varia; (D) Pyrenula quassiaecola; (E) Arthonia antillarum;
(F) Arthonia caribaea; (G) Celothelium dominicanum; (H) Coenogonium aurantiacum.
44
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
J.A. Mercado-Díaz
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
R. Lücking
Plate 19
H
Plate 19—(A) Coenogonium borinquense; (B) Enterographa multilocularis; (C) Fissurina tachygrapha; (D) Leucodecton occultum;
(E) Opegrapha astraea; (F) Phaeographis inusta (between lenticels); (G) Phaeographis inusta (detail); (H) Pyrenula thelomorpha.
45
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 20
B
D
F
H
Plate 20—(A) Ramonia rappii; (B) Ramonia valenzuelana; (C) Parmotrema endosulphureum; (D) Parmotrema endosulphureum (isidia
and yellow medulla); (E) Physcia erumpens; (F) Pyrenula cerina*; (G) Pyrenula cerina (detail); (H) Bactrospora myriadea.
46
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
D
J.A. Mercado-Díaz
J.A. Mercado-Díaz
B
F
R. Lücking
E
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
Plate 21
H
Plate 21—(A) Enterographa aff. anguinella; (B) Pyrenula cocoes; (C) Pyrenula ochraceolava; (D) Pyrenula microcarpa;
(E) Herpothallon minimum; (F) Porina curtula; (G) Arthonia aff. rubella; (H) Coenogonium dimorphicum.
47
E
G
J.A. Mercado-Díaz
J.A. Mercado-Díaz
C
J.A. Mercado-Díaz
A
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
J.A. Mercado-Díaz
Plate 22
B
D
F
H
Plate 22—(A) Coenogonium portoricense; (B) Crocynia gossypina; (C) Fissurina adscribens; (D) Graphis farinulenta; (E) Diorygma
poiteai* (left) and Graphis farinulenta* (right); (F) Lecanographa lyncea; (G) Malmidea aff. coralliformis; (H) Schismatomma rappii.
48
Lichens in Puerto Rico: An Ecosystem Approach
Lichen communities—
Along with the tabonuco and lowland moist forests (see below), palo colorado
forests in El Yunque National Forest are perhaps one of the most lichen-diverse forest ecosystems of Puerto Rico. About 85 species have been found in sampled areas
of this forest type. Sampled areas of palo colorado forests resulted in the highest
number of species within the genera Ocellularia, Chapsa, Malmidea, Phyllopsora,
Rhabdodiscus, and Thelotrema. Species of Ampliotrema, Compositrema, and
Paratopeliopsis were found only in this forest type (Mercado-Díaz et al., in press).
Palo colorado forests are second to elfin woodland forests in the number of species
within the genus Clandestinotrema. This confirms that similar to other forests
in the Neotropics, species within this genus are limited to high-elevation forests
(Rivas-Plata et al. 2008).
Filamentous lichens are evident in palo colorado forests in El Yunque National
Forest, where species like Coenogonium linkii (plate 2G) are occasionally found at
the base of trees. Basidiolichens are also present in these forests, more commonly
the species Dictyonema sericeum (plate 1B). Foliose and fruticose lichens are
mostly absent under the shaded understory; only microfoliose species within the
genus Phyllopsora may be observed. Several species with byssoid growth habit
including Herpothallon aurantiacoflavum (plates 2H and 3A), H. granulare (plates
2H and 3B), and Dichosporidium nigrocinctum (plate 3, C and D) are frequently
found in these forests.
Common corticolous species in the palo colorado forest include Chapsa thallotrema (plate 1, E and F), Coenogonium linkii, Dichosporidium nigrocinctum,
Dictyonema sericeum, Graphis rhizocola (plate 3E), Herpothallon aurantiacoflavum, Malmidea nigromarginata (plate 3F), Myeloconis guyanensis (plate 3G),
Ocellularia praestans (plates 3H and 4A), and Porina nucula (plate 4B). Many of
these species can also be found in other ecosystem types in El Yunque National
Forest.
Several corticolous lichens have been found only in sampled areas of palo
colorado forests in El Yunque National Forest. Some of these include Ampliotrema
rimosum (plate 4C), Arthonia spp. (yellow margin) (plate 4D), Chapsa alborosella
(plate 4E), Clandestinotrema cf. tenue (plate 4F), Coenogonium nepalense (plate
4G), Graphis dimidiata (plate 4H), Malmidea furfurosa (plate 5A), M. vinosa (plate
5B), Paratopeliopsis caraibica (plate 5C), Ocellularia aff. perforata (plate 5D), O.
cavata (plate 5E), O. crocea (plate 5F), O. dolichotata (plate 5G), O. rhabdospora
(plate 5H), and Platythecium grammitis (plate 6A).
49
GENERAL TECHNICAL REPORT IITF-GTR-46
At least 11 morphotypes of thelotremoid lichens have been found in sampled
areas of palo colorado forests in El Yunque National Forest (Rivas-Plata et al.
2008). These are the encountered morphotypes and several representative species:
Ampliotremoid: Ampliotrema rimosum
Chroodiscoid: Chapsa alborosella, Astrochapsa platycarpella
Leprocarpoid: Chapsa dissuta, C. esslingeri
Melanotremoid: Clandestinotrema leucomelaenum
Tenuitremoid: Clandestinotrema cf. tenue
Myriotremoid: Paratopeliopsis caribica
Ocellularioid: Ocellularia crocea, O. perforata, O. tacarcunae
Rhodostromoid: Ocellularia cavata, O. rhodostroma, O. xanthostroma
Praestantoid: Ocellularia praestans, O. rhabdospora, O. dolichotata
Stegoboloid: Rhabdodiscus emersus, R. schizostomus, R. isidiiferus
Thelotremoid: Thelotrema porinoides, T. lepadodes
Sierra Palm Forest
J.A. Mercado-Díaz
Environment—
In Puerto Rico, the sierra palm forest ecosystem (palm break) is found mostly
within the lower montane wet forest life zone but might also reach areas classified
as subtropical wet and subtropical rain forest (fig. 18). This forest type consists of
nearly pure stands of Prestoea acuminata var. montana (Gould et al. 2006, Miller
and Lugo 2009). Palm forest stands have a mean annual temperature of 20.7 °C and
a mean annual precipitation of 3956 mm. These forests may cover nearly 11 percent
Figure 18—The understory of a sierra palm forest in El Yunque National Forest, Puerto Rico.
50
Lichens in Puerto Rico: An Ecosystem Approach
of the Luquillo Mountains (Gould et al. 2006, Miller and Lugo 2009, Weaver 2012).
Some of the characteristics of the “palm break” include its high dominance and a
sharp ecotone with adjacent associations (Miller and Lugo 2009). The sierra palm
forest is present in the 730 to 915 m altitudinal zone and can be found frequently
interspersed with palo colorado and tabonuco forests in El Yunque National Forest
(Miller and Lugo 2009).
J.A. Mercado-Díaz
Vegetation—
Plant communities of the sierra palm forest are mostly found in azonal, steeply
sloping sites dominated by P. acuminata var. montana and are characterized by
having a high abundance of epiphytes (Gould et al. 2006) (fig. 19). The development
of an understory in the sierra palm forest is poor and occasionally absent. Soils
are periodically covered by herbaceous vegetation and mosses (Lugo 2005). Five
endemic tree species with diameter at breast height > 2 cm are commonly found
in sierra palm forests in El Yunque National Forest, the most abundant being
Henriettea squamulosum (Cogn.) W.S. Judd, Eugenia borinquensis, and Cyathea
arborea (L.) Sm. (Gould et al. 2006). The abundance of Cecropia schreberiana
Figure 19—Patches of the common crustose lichen Chapsa thallotrema in El Yunque National Forest, growing on the trunk of
Prestoea acuminata var. montana in a sierra palm forest.
51
GENERAL TECHNICAL REPORT IITF-GTR-46
Miq. in the sierra palm forest is indicative of past hurricane disturbance (Gould et
al. 2006). When occurring in the tabonuco forest, P. acuminata var. montana does
not form any particular association in the forest canopy and is frequently found in
the understory and in microenvironments that are water-saturated (Lugo 2005).
See table 2 for common names of plant species.
Lichen communities—
Perhaps the most distinctive characteristic of the sierra palm forest in terms of its
lichen flora is that it has the lowest species diversity compared to the rest of the
forest types in El Yunque National Forest. Only 24 species have been found in
sampled areas of this forest type. Despite its low species diversity, lichens in the
sierra palm forest are usually more evident than in other forest types of El Yunque
National Forest owing to the frequent occurrence of foliose lichens on tree trunks.
For example, species in the genus Sticta (plate 6B) were mostly observed within
this forest type. Many species of Leptogium, such as L. azureum (plate 6C) and
L. denticulatum (plate 6D), are commonly found in the sierra palm forests. The
new species Thalloloma rubromarginatum has been found only in this forest type
(Mercado-Díaz et al., in press).
Most of the lichen flora in palm forests is crustose. Coenogonium linkii (plate
2G) is an example of filamentous species found in these forests. Byssoid growth
forms are represented by species of Herpothallon, like H. granulare (plates 2H
and 3B).
Some of the most commonly encountered lichen species of the sierra palm
forest are Chapsa thallotrema (plate 1, E and F), Coenogonium linkii, Graphis
duplicata (plate 6E), Herpothallon granulare, Leptogium azureum, Porina
subpungens (plate 6F), and Thelotrema porinoides (plate 6G). Several of these
species may also be found in other ecosystem types in El Yunque National Forest.
Only a few species have been found exclusively in sampled areas of this forest
type. Some of these include Arthonia complanata (plate 6H), Graphis proserpens
(plate 7A), Fissurina sp. nov. (plate 7, B and C), Megalotremis lateralis (plate 7D),
Pyrenula aff. fetivica (plate 7E), and Sticta beauvoisii (plate 7F).
Only two morphotypes of thelotremoid lichens have been found in sampled
areas of sierra palm forests in El Yunque National Forest (Rivas-Plata et al. 2008).
These are the encountered morphotypes and their representative species:
Sorediotremoid: Chapsa thallotrema
Thelotremoid: Thelotrema porinoides
It is interesting, from an ecological perspective, that most of the corticolous
lichens occurring in sierra palm forests grow on the trunks of tree species other
52
Lichens in Puerto Rico: An Ecosystem Approach
than P. acuminata var. montana, which is the dominant tree in this ecosystem.
Nonetheless, specimens of P. acuminata var. montana present in other forest types
of El Yunque National Forest apparently have higher species diversity on their
trunks than specimens occurring in pure sierra palm forest stands (table 3). A
possible explanation for this phenomenon is that trees with high species diversity
surrounding P. acuminata var. montana individuals in these other forests are facilitating the immigration of species to neighboring P. acuminata var. montana trees.
Table 3—Lichens that grow on Prestoea acuminata var. montana trees in
different forest types in El Yunque National Forest, Puerto Rico
Species
Chapsa thallotrema
Chapsa alborosella
Dichosporidium nigrocinctum
Graphis duplicata
Herpothallon rubrocinctum
Mazosia ocellata
Ocellularia sp.
Ocellularia sp. A
Porina scabrida
Pyrenula aspitea
Pyrenula sp. A
Sterile crustose
Thelotrema porinoides
Total
Elin
Palo colorado
Sierra palm
Tabonuco
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
1
3
3
10
Tabonuco Forest
Environment—
The tabonuco forest type occurs mainly in mid- to high-elevation areas of Puerto
Rico and its elevational range spans from 15 to 550 m (Brown et al. 1983) (fig. 20).
Nonetheless, in mountains like those in Tres Picachos State Forest in Jayuya, this
forest may be found at 800 m (Lugo 2005). This ecosystem is characterized by
having high tree-species diversity per unit area (Gould et al. 2006) and the presence
of tall broadleaf evergreen hardwood trees that commonly exhibit buttressed
trunks (Miller and Lugo 2009). It harbors a diverse epiphyte flora and many
species of lianas. Many of the plants present in the tabonuco forest exhibit drip-tip
leaves. It has a mean annual precipitation of 3060 mm per year and a mean annual
temperature of 23 °C (Gould et al. 2006). Water deficits in the tabonuco forest are
very small and soil moisture usually drops below field capacity (Miller and Lugo
2009).
53
Vegetation—
The tabonuco forest is a closed broad-leaved forest that
gets its name from its dominant tree, Dacryodes excelsa
Vahl (fig. 21). Forest patches of this ecosystem in El
Yunque National Forest could harbor about 40 tree species and about 86 plant species (Gould et al. 2006). These
forests form a complete canopy at about 20 m. Other
prominent species are Sloanea berteriana Choisy ex DC,
Manilkara bidentata (A. DC.) A. Chev, and Tetragastris
balsamifera (Sw.) Oken (Gould et al. 2006, Lugo and
Miller 2009). After disturbances, Cecropia schreberiana
invades open areas and rapidly closes the canopy. This
forest type includes several rare endemic species like
Cordia borinquensis Urb., Rondeletia portoricensis Krug.
& Urb., and Byrsonima wadsworthii Little (Gould et al.
2006). Along with the palo colorado forest, the tabonuco
forest has the highest number of native plant species per
Figure 20—The trunk and sap of Dacryodes excelsa in El
Yunque National Forest. This trunk is almost completely
unit area (Gould et al. 2006). Lugo (2005) reported that
covered with a greenish crustose lichen.
the tabonuco forest has high populations of “microscopic
fungi” colonizing leaf surfaces in the canopy. Most of these microscopic fungi are
possibly foliicolous lichens that have been observed growing along with liverworts
on the leaves of these forests. See table 2 for common names of plant species.
J.A. Mercado-Díaz
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 21—Mosaic of lichens growing on the lower trunk of a tree in a tabonuco forest. El Yunque
National Forest, Puerto Rico.
54
Lichens in Puerto Rico: An Ecosystem Approach
Lichen communities—
Tabonuco forests have one of the richest lichen biota in Puerto Rico, with at least
76 corticolous species recorded in sampled areas of this forest type in El Yunque
National Forest. Perhaps because D. excelsa is a dominant element in these forests,
it appears to be the tree species with the highest number of lichen taxa on its bark.
There are at least 27 known species growing on the lower trunk sections of D.
excelsa (see list below). Although not as conspicuous as D. excelsa, tree individuals
of Prestoea acuminata var. montana are often found with several lichen species
such as Dichosporidium nigrocinctum (plate 3, C and D), Porina scabrida (plate
7G), Graphis duplicata (plate 6E), and Mazosia endonigra (plate 7H). These forests
also possess the highest number of Porina and Herpothallon species for any forest
type discussed in this report. Species of Acanthotrema and Borinquenotrema have
only been found in this forest type (Mercado-Díaz et al., in press).
Lichens that grow on D. excelsa trees in El Yunque National Forest, Puerto Rico:
Acanthotrema alboisidiatum
Ocellularia cf. praestans
Arthonia aff. bessalis
Ocellularia mordenii
Bacidia sp. A
Ocellularia sp. D (sorediate)
Chapsa thallotrema
Ocellularia umbilicata
Dichosporidium nigrocinctum
Opegrapha sp. A
Herpothallon aurantiacoflavum
Phyllopsora sp. A
Herpothallon rubrocinctum
Phyllopsora buettneri
Leucodecton compunctelum
Porina (sterile)
Mazosia sp. A
Porina conspersa
Mazosia sp. B
Pyrenula macrocarpa
Mazosia sp. nov.
Pyrenula mastophoroides
Myeloconis guyanensis
Pyrenula fetivica
Myriotrema sp. B
With the exception of occasional foliose lichens in upper canopy layers, nearly
all lichen taxa documented in these forests are crustose. Several species with
byssoid growth habit are commonly found; e.g., D. nigrocinctum, Herpothallon
aurantiacoflavum (plates 2H and 3A), H. rubrocinctum (plate 8A), and H. granulare
(plates 2H and 3B). Squamulose species are represented by several species of Phylopsora, like P. buettneri (plate 8B), P. corallina (plate 8C), and the basidiolichen
Cyphellostereum (plate 1C). Fruticose lichens are absent in the understory, although
species of Usnea (plate 8D) can be found in light-exposed conditions. Dimorphic
species, like those in the genus Cladonia (e.g. Cladonia subradiata, plate 8F) are
expected to be found in open areas in these forests, growing directly on the soil or
on other exposed substrates; however, these have not yet been encountered.
55
GENERAL TECHNICAL REPORT IITF-GTR-46
Common lichen species within this forest type include Arthonia platygraphidea
(plate 8G), Arthothelium distendens, (plate 8H), Chapsa thallotrema (plate 2, E
and F), Ocellularia interposita (plate 9A), Mazosia endonigra (plate 7H), Pyrenula
macrocarpa (plate 9B), and Pyrenula massariospora (plate 9C). Some of these
species may also be found in other ecosystem types in El Yunque National Forest
and even in forests at lower elevations. Corticolous species of more exposed
conditions or occurring in the canopy include Thelotrema porinoides (plate 6G)
and Heterodermia speciosa (plate 9D). Several species have only been found
in sampled areas of this forest type; for example: Acanthotrema alboisidiatum
(plate 9E), Arthonia aff. bessalis (plate 9F), Arthothelium sp. nov. (plate 9G),
Pseudochapsa dilatata (plate 9H), Chapsa elabens (plate 10A), Eugeniella sp.
nov. (plate 10B), Herpothallon aff. pustulata (plate 10C), Letrouitia vulpina (plate
10D), Leucodecton compuctellum (plate 10E), Malmidea amazonica (plate 10F),
Mazosia verrucosa (plate 10G), Megalotremis infernalis (plate 10H), Ocellularia
aff. cavata (plate 11A), O. ascidioidea (plate 11B), O. mordenii (plate 11C),
Borinquenotrema soredicarpum (plate 11D), Porina americana (plate 11E), P.
scabrida, Rhabdodiscus emersellus (plate 11F), and Wirthiotrema glaucopallens
(plate 11G).
Tabonuco forests have the second highest diversity of thelotremoid lichens
documented in El Yunque National Forest (compared to palo colorado forests). In
this respect, at least nine morphotypes of thelotremoid lichens have been found
(Rivas-Plata et al. 2008). The most numerous genus of thelotremoid lichens in this
forest is Ocellularia with 12 species. These are the encountered morphotypes and
several representative species:
Leprocarpoid: Pseudochapsa dilatata
Sorediotremoid: Chapsa thallotrema
Leucodectonoid: Leucodecton compunctellum
Myriotremoid: Myriotrema sp. nov., Wirthiotrema glaucopallens
Ocellularioid: Ocellularia ascidiodea, O. perforata, O. umbilicata
Rhodostromoid: Ocellularia mordenii, O. aff. cavata
Praestantoid: Ocellularia cf. praestans, O. interposita
Stegoboloid: Rhabdodiscus emersus, R. emersellus
Thelotremoid: Thelotrema porinoides
56
Lichens in Puerto Rico: An Ecosystem Approach
Lowland Moist Forest
J.A. Mercado-Díaz
Environment—
The lowland moist forest occurs in the lowland subtropical moist Holdridge life
zone (Ewel and Whitmore 1973) (fig. 22). It covers more area than any of the other
life zones described by Ewel and Whitmore (1973). Mean annual rainfall in the
lowland moist forest varies from 1000 to 2200 mm and mean annual temperature
is about 27.5 °C (Gould et al. 2006, Miller and Lugo 2009). This forest type occurs
in a variable elevation range, from sea level to 671 m (Ewel and Whitmore 1973).
Throughout the world, lowland moist forests are among the most intensively
used life zones (Ewel and Whitmore 1973). In Puerto Rico, most of these forests
were deforested in the past mainly because their climatic conditions are favorable
for agricultural activities (Miller and Lugo 2009). Other forest ecosystems, like
Pterocarpus wetlands and mangrove swamps, could be classified as lowland moist
forests; however, their unique biological and environmental characteristics justify a
separate formal description (see further sections).
Figure 22—A tree of Spathodea campanulata, a fast-growing species commonly observed in lowland
moist forests in Puerto Rico.
Vegetation—
This forest is characterized by trees up to 20 m tall with rounded crowns (Miller and
Lugo 2009) (fig. 23). Dominant canopy vegetation includes several tree species such
as Manilkara bidentata, Ocotea leucoxylon (Sw.) De Laness. and Hymenaea courbaril L. (Gould et al. 2006). Faramea occidentalis (L.) A. Rich. and Inga laurina
57
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 23—Trunk of a tree in lowland moist forest. To the right, two thalli of Cryptothecia striata.
Río Piedras, Puerto Rico.
(Sw.) Willd. are common understory species (Gould et al. 2006). These forests also
have the highest mean number of nonnative plant species per unit area (Gould et
al. 2006). Epiphytes are common but rarely cover the surface of branches and trees
entirely (Miller and Lugo 2009). Many of the woody species are deciduous during the dry season (Miller and Lugo 2009). Because most of this forest type was
subjected to intensive human use in the past, it is extremely difficult to find natural
undisturbed stands.
Lichen communities—
Although most lowland moist forests in Puerto Rico have been subject to intense
human alteration, these areas may be among the forest ecosystems with the highest
number of lichen species on the island. At least 86 species of corticolous lichens
were identified in the understory of these forests. These forests also exhibit the
highest number of lichen genera represented in a forest type (37). Because lowland
moist forests occupy most of the island’s surface area (59 percent) (Miller and Lugo
2009), and include other interesting vegetation associations occurring in serpentineand limestone-derived soils not discussed in this report, lichen species richness of
these forests should be higher than that reported here.
58
Lichens in Puerto Rico: An Ecosystem Approach
The lichen biota of lowland moist forests in Puerto Rico is almost completely
dominated by crustose growth forms. Filamentous growth forms may occasionally
be found on the base of trunks, particularly the species Coenogonium leprieurii
(plate 11H) and Coenogonium linkii (plate 2G). Foliose growth forms are rarely
found in the shaded understory of these forests; however, several species such
as Physcia, Pyxine, and Parmotrema have been seen on fallen canopy branches,
indicating their presence in upper canopy layers. Physcia atrostriata (plate 12A),
Parmotrema praesorediosum (plate 12B), and Dirinaria purpurasens (plate 12C)
are commonly observed foliose lichens in marginal areas of these forests. Fruticose
lichens like the species Ramalina complanata (plate 12D) and Ramalina peruviana
(plate 12E) are occasionally observed in open areas of these forests.
Compared to other forest ecosystems presented here, lowland moist forests may
have the highest diversity of species in the genera Fissurina and Opegrapha. Commonly encountered species of these forests include Cresponea melanocheiloides
(plate 12, F and G), Cryptothecia striata (plate 12H), Diorygma poiteai (plate 13,
A and B), Hemithecium balbisii (plate 13, C and D), Graphis glaucescens (plate 13,
E and F), Malmidea piperis (plate 13, G and H), Opegrapha bonplandii (plate 14A),
Opegrapha dekeselii (plate 14B), Phaeographis brasiliensis (plate 14C), Porina
distans (plate 14, D and E), Porina conspersa (plate 14F), Pyrenula mamillana
(plate 14G), and Fissurina subnitens (plates 14H and 15A).
Species that were only found in these forests include Anisomeridium
subprostans (plate 15B), Arthonia aff. pyrrholiza (plate 15C), Arthonia bessalis
(plate 15D), Coenogonium leprieurii, Cresponea melanocheiloides, Dyplolabia
afzelii (plate 15E), Letrouitia dominguensis (plate 15, F and G), Leucodecton
bisporum (plates 15H and 16A), Mazosia carnea (plate 16B), Melanotrema
platystomum (plate 16C), Monoblastia borinquensis (plate 16D), Porina tetracerae
(plate 16E), Psoroglaena cubensis (plate 16F), Strigula phaea (plate 16G), and
Trinathotrema stictideum (plate 16H).
There are at least five morphotypes of thelotremoid lichens present in sampled
areas of lowland moist forests. These are the encountered morphotypes and their
representative species:
Leprocarpoid: Chapsa defecta
Sorediotremoid: Chapsa thallotrema
Leucodectonoid: Leucodecton bisporum
Melanotremoid: Melanotrema platystomum, Trinathotrema stictideum
Glaucophaenoid: Myriotrema erodens
59
GENERAL TECHNICAL REPORT IITF-GTR-46
Dry Forest
Environment—
Dry forests have the highest temperatures and lowest precipitation of all forest ecosystems in Puerto Rico and are found within the subtropical dry forest zone (Ewel
and Whitmore 1973) (fig. 24). These forests cover vast areas in southwestern Puerto
Rico, offshore islands like Mona, Culebra, Vieques, Caja de Muertos, Desecheo,
and smaller areas to the east around the vicinity of Fajardo (Lugo 2005, Miller and
Lugo 2009). Dry forests mostly occur along the coast and their elevation spans
from sea level to about 200 m (Lugo 2005). Because mean annual rainfall ranges
from 600 to 1100 mm, water scarcity is the predominant environmental condition
in these forests (Lugo 2005, Miller and Lugo 2009). Dry forest air temperatures can
be extremely variable during the day, occasionally reaching more than 30 °C (Lugo
2005). Most dry forests in Puerto Rico occur on calcareous substrates, and their
mean annual temperature is about 25 °C (Lugo 2005). Noncalcareous dry forests
cover a smaller area (about 1.2 percent of the island) and their mean annual temperature might be slightly higher than calcareous dry forests (27.5 °C) (Gould
et al. 2006, Lugo 2005).
J.A. Mercado-Díaz
Vegetation—
Vegetation of dry forest areas visited for this project is typical of dry forests occurring in noncalcareous substrates (fig. 25). Commonly encountered tree species
include Bursera simaruba (L.) Sarg., Bourreria succulenta Jacq., Quadrella
Figure 24—The understory of a dry forest. Former Naval Station Roosevelt Roads, Ceiba, Puerto
Rico.
60
cynophallophora (L.) Hutch., Erythroxylum brevipes
DC., Neea buxifolia (Hook. f.) Heimerl, and Eugenia
biflora (L.) DC. (Gould et al. 2006). Plant species
diversity per unit area was the highest (99 species per
300 m2) among several forest ecosystems (Gould et al.
2006). Evergreen species are predominant and there is
low dominance of deciduous species compared to dry
forests occurring on calcareous substrates (Lugo 2005).
In calcareous dry forests, soils accumulate little water
because of their rocky nature and shallowness, creating
arid conditions for plant growth (Lugo 2005). In this
respect, plants in noncalcareous dry forests exhibit
less extreme xeromorphic traits than their counterparts
in calcareous dry forests. Overall, vegetation of the
island’s dry forests tends to form a complete ground
cover and is mostly semideciduous on most soils
(Miller and Lugo 2009). See table 2 for common
names of plant species.
J.A. Mercado-Díaz
Lichens in Puerto Rico: An Ecosystem Approach
Figure 25—Trunk of a tree in a dry forest. Note the mosaic of
crustose lichens in the trunk. Former Naval Station Roosevelt
Roads, Ceiba, Puerto Rico.
Lichen communities—
A total of 77 species of corticolous lichens has been found in sampled areas of this
forest ecosystem. This suggests that, compared to other lowland forest ecosystems,
these forests are among the richest in terms of their corticolous lichen biota. Trees
of dry forests in the southwestern portion of the island have also been seen to contain high lichen cover and diversity. Southwestern dry forests have slightly different
vegetation composition owing in part to the calcareous nature of their substrate and
are therefore expected to vary in terms of their lichen composition. Although dry
forests on calcareous substrates were not sampled for this work, it is suspected that,
if taken into consideration, dry forests as a whole may contain the highest species
diversity for any lowland ecosystem of the island.
As in other forest ecosystems, dry forests are dominated by crustose growth
forms. Several foliose species are occasionally found, such as Physcia crispa
(plate 17A), Physcia sorediosa (plate 17B), Physcia atrostriata (plate 12A), Pyxine
berteriana (plate 17C), Pyxine eschweileri (plate 17D), and Parmotrema praesorediosum (plate 12B). Although fruticose species were not found, species such as
Ramalina complanata (plate 12D) have been observed in similar forests around the
region. Many lichens found in these forests are whitish or brightly colored (yellow,
orange, etc.). This is a common characteristic exhibited by lichens growing in dry
environments with high light exposure (Brodo et al. 2001).
61
GENERAL TECHNICAL REPORT IITF-GTR-46
Dry forests were found to have the highest diversity of species in the genera
Arthonia, Bactrospora, Physcia, and Pyrenula. Ramonia species were only found
in these forests. Some of the most common corticolous lichens found in these
forests are Anisomeridium biforme (plate 17E), Arthonia portoricensis (plate
17F), Arthopyrenia majuscula (plate 17G), Bactrospora denticulata (plate 17H),
Graphis furcata (plate 18A), Graphis tenella (plate 18B), Mazosia endonigra
(plate 7H), Opegrapha cf. varia (plate 18C), Porina nucula (plate 4B), Pyrenula
quassiaecola (plate 18D), and Arthonia antillarum (plate 18E). Among the species
that have only been found occurring in these forests are Arthonia caribaea (plate
18F), Celothelium dominicanum (plate 18G), Coenogonium aurantiacum (plate
18H), Coenogonium borinquense (plate 19A), Enterographa multilocularis (plate
19B), Fissurina tachygrapha (plate 19C), Graphis tenella, Leucodecton occultum
(plate 19D), Opegrapha astraea (plate 19E), Phaeographis inusta (plate 19, F and
G), Pyrenula telomorpha (plate 19H), Ramonia rappi (plate 20A), and Ramonia
valenzuelana (plate 20B).
Only two morphotypes of thelotremoid lichens were found in sampled areas
representative of dry forests. These are the encountered morphotypes and their
representative species:
Leucodectonoid: Leucodecton occultum
Sorediotremoid: Sorediate crustose lichen, possibly Chapsa
Mangrove Forest
Environment—
Mangrove forests grow near sea water and are found at lowland elevations (<5
m) within the lowland subtropical dry and moist Holdridge life zones (Ewel and
Whitmore 1973) (fig. 26). Mean annual temperature is 27.3 °C and mean annual
precipitation is 1414 mm (Gould et al. 2006). Mangrove forests are often referred to
as tidal “fringing forests” because they thrive in an ecotonal fringe along protected
tropical coasts, lagoons, bays, and offshore islands (Miller and Lugo 2009). Because
salt water can flow upstream from estuaries to rivers, mangroves may extend kilometers inland, as far as the presence of salt water occurs (Miller and Lugo 2009).
By 1975, about half of the mangrove forests of Puerto Rico had been destroyed by
agricultural activities and other human activities along the coast. In the last few
years, there has been a net gain in mangrove coverage as a result of their protected
status (Martinuzzi et al. 2009, Miller and Lugo 2009). Mangrove areas are the most
extensive estuarine forested wetlands. The largest (Bosque de Pinoñes) is just east
of San Juan in the municipalities of Carolina and Loíza (Miller and Lugo 2009).
62
J.A. Mercado-Díaz
Lichens in Puerto Rico: An Ecosystem Approach
Figure 26—View of the forest loor in a mangrove forest. Trees, saplings, and pneumatophores of
Avicennia germinans. Ceiba, Puerto Rico.
Vegetation—
There are four common mangroves species typical to Puerto Rico and the Caribbean: Rhizophora mangle L., Avicennia germinans (L.) L., Laguncularia racemosa
(L.) C.F. Gaertn., and Conocarpus erectus L. (fig. 27). These forests are typically
found as Rhizophora mangle–dominated stands on coastal and estuarine fringes
and in basins as pure or mixed stands of A. germinans and L. racemosa (Gould et
al. 2006). When found in the subtropical moist forest life zone, mangroves appear to
grow taller than in the subtropical dry life zone (Miller and Lugo 2009). Mangrove
forests have the lowest plant species diversity per unit area as compared to other
plant communities in Puerto Rico, although most of their flora is native (Gould et
al. 2006). Mangrove is a general term that can refer to numerous families, genera,
and species in totally different taxons (Miller and Lugo 2009). All plant species
occurring in mangrove forests are salt-tolerant, or “halophytes.” Mangroves provide
a safe wildlife habitat both on land and around their prop roots in water, dampen the
power of storm waves, and help stabilize land that normally would be eroded to the
sea. See table 2 for common names of plant species.
Lichen communities—
In terms of their lichen composition, mangrove forests along the coastline are perhaps the most species-poor forest ecosystems of the island. Only 18 lichen species
have been identified in sampled areas of these forests. However, if inland mangrove
63
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
Figure 27—Several trunks of young mangrove trees. Orange color patches belong to Pyrenula cerina, a common crustose lichen found
in these forests. Las Cabezas de San Juan Natural Reserve, Fajardo, Puerto Rico.
forest systems are included, species diversity increases substantially. For example,
one mangrove stand in Sabana Seca Naval Station alone has about 30 corticolous
lichen species. A possible explanation for the higher species diversity observed in
inland mangrove forests is that these forests are often close to other species-rich
forest ecosystems that may be actually serving as a source of new species to these
mangrove forests. For instance, the mangrove stand in the Sabana Seca Naval Station is only several meters away from a Pterocarpus forest.
Similar to other forest ecosystems, species with a crustose growth habit dominate in these ecosystems. Foliose species such as Parmotrema endosulphureum
(plate 20 C and D) and Physcia erumpens (plate 20E) are occasionaly found.
Fruticose growth forms are rare or absent in the understory of these forests.
The most conspicuous lichen element in these forests is the ubiquitous presence
of Pyrenula cerina (plate 20F), growing almost exclusively on the trunk and aereal
roots of R. mangle. Other species associated with P. cerina and commonly found in
64
Lichens in Puerto Rico: An Ecosystem Approach
these forests include Arthonia antillarum (plate 18E), Arthonia portoricensis (plate
17F), and Bactrospora myriadea (plate 20H). The species Enterographa aff. anguinella (plate 21A), Pyrenula cocoes (plate 21B), Pyrenula ochraceoflava (plate 21C),
Pyrenula cerina, and Pyrenula microcarpa (plate 21D) are commonly observed and
have only been found occurring in these forests. Species in the genus Graphis have
also been observed in mangrove forests.
No thelotremoid lichens were found during fieldwork for this study. Only
sorediotremoid morphotypes (Rivas-Plata et al. 2008) are expected to be found in
some mangrove forests of Puerto Rico. Although mangrove forests are not treated
in Rivas-Plata et al. (2008), if their methods were to be applied for estimating ecological continuity, it can be predicted that even old, undisturbed mangrove forests
around the island would be classified as “anthropogenic vegetation.” These facts
highlight the importance of identifying suitable bioindicators of ecological continuity for this type of forest.
Pterocarpus Forest
Environment—
Pterocarpus forests occur in relict patches within the lowland subtropical moist
Holdridge life zone (Ewel and Whitmore 1973) (fig. 28). They are mostly found
on undisturbed freshwater, seasonally flooded, noncalcareous alluvial substrates
(Gould et al. 2006). Pterocarpus wetlands commonly occur along the coast,
although a few populations are present along streams in northeastern Puerto
Rico (Alvarez-López 1990). Compared to mangrove and dry forests, Pterocarpus
forests have slightly lower temperatures (mean annual temperature = 26.4 °C) and
higher precipitation (mean annual precipitation = 1685 mm) (Gould et al. 2006).
These forests occur in the most heavily populated coastal plain and are subject to
disturbance from changes in the hydrologic regime, particularly those related to
urban development (Gould et al. 2006). Pterocarpus forests were likely much more
extensive on coastal plains and along riparian corridors prior to deforestation for
agricultural activities.
Vegetation—
Most Pterocarpus forests are considered palustrine forested swamps that usually
have more than 40 percent of their area in tree cover (Miller and Lugo 2009) (fig.
29). It is typically a monoculture of the tree species P. officinalis Jacq., which
occupies the overstory and the fern Acrostichum aureum L. in the understory.
Like mangrove forests, Pterocarpus forests have low plant species diversity, but
most of their flora is composed of native species (Gould et al. 2006). Pterocarpus
trees may occasionally be found among mangrove trees as part of floodplain forests
in riverine estuaries (Miller and Lugo 2009). Common lianas found here include
65
J.A. Mercado-Díaz
GENERAL TECHNICAL REPORT IITF-GTR-46
J.A. Mercado-Díaz
Figure 28—Trunk and buttressed roots of a Pterocarpus oficinalis tree. Sabana Seca Naval Station, Toa Baja, Puerto Rico.
66
Figure 29—Patches
of crustose lichens
growing on young
Pterocarpus oficinalis
trees. Sabana Seca
Naval Station, Toa
Baja, Puerto Rico.
Lichens in Puerto Rico: An Ecosystem Approach
Ipomoea triloba L., Paullinia pinnata L., and Hippocratea volubilis L. Freshwater
Pterocarpus communities exhibit great canopy heights, with trees reaching more
than 34 m (Gould et al. 2006). As a result of former agricultural activities and
other coastal development, Pterocarpus forests have been reduced to only a few
remnant stands. It is believed that in Puerto Rico these forests occupy no more than
5 percent of their original range, making them deserving of special protection status
(Miller and Lugo 2009).
Lichen communities—
Pterocarpus forests are almost entirely a monoculture of P. officinalis; therefore,
most of the corticolous lichens found in these forests grow on the trunk of this tree
species. Compared to other forest ecosystems, corticolous lichen species richness is
moderate, with about 59 species found in sampled areas of this forest type. Lichens
of Pterocarpus forests share some similarities with the lichen flora of lowland moist
forests, particularly at the genus level. This is not surprising, as both forest types
are found within the lowland subtropical moist Holdridge life zone (see above).
With the exception of fruticose lichens, most lichen growth forms are represented in these forests. As other forests, crustose growth forms dominate; however,
foliose lichens such as Parmotrema praesorediosum (plate 12B), Physcia erumpens
(plate 20E), and Physcia sorediosa (plate 17B), as well as squamulose species in
the genus Phyllopsora can be found occasionally. Species of filamentous lichens
such as Coenogonium linkii (plate 2G) can be frequently found on the base of tree
trunks. Dichosporidium nigrocinctum (plate 3, C and D), Herpothallon aurantiacoflavum (plates 2H and 3A), and H. minimum (plate 21E) are examples of byssoid
growth forms found sporadically in these forests.
Species that are commonly observed in these forests include Diorygma
poiteai (plate 13, A and B; plate 22E), Graphis glaucescens (plate 13, E and F),
Hemithecium balbisii (plate 13, C and D), Opegrapha dekeselii (plate 14B), Physcia
sorediosa (plate 17B), and Porina curtula (plate 21F). Several species have only
been found in this type of forest, e.g., Arthonia aff. rubella (plate 21G), Coenogonium dimorphicum (plate 21H), Coenogonium portoricense (plate 22A), Crocynia
gossypina (plate 22B), Fissurina adscribens (plate 22C), Graphis farinulenta (plate
22, D and E), Herpothallon minimum, Lecanographa lyncea (plate 22F), Malmidea
aff. coralliformis (plate 22G), and Schismatomma rappii (plate 22H). Pterocarpus
forests have the highest number of Coenogonium species recorded among all
sampled ecosystems.
67
GENERAL TECHNICAL REPORT IITF-GTR-46
Only two morphotypes of thelotremoid lichens were found in sampled areas
representative of Pterocarpus forests. These are the encountered morphotypes and
their representative species:
Sorediotremoid: Chapsa (sorediate)
Leprocarpoid: Chapsa dissuta
Concluding Remarks
Results from this study suggest that tabonuco and palo colorado forests of El
Yunque National Forest, as well as lowland moist forests, are the ecosystems
with the highest lichen diversity in Puerto Rico. Along with these ecosystems,
other forests on the island, such as the state forests of Maricao, Toro Negro, Tres
Picachos, and Carite, are expected to harbor the vast majority of lichens. Dry
forests are also species-rich and make a significant contribution to the island’s
lichen diversity. The least diverse forests of the island are most likely mangrove
and elfin woodland forests and those in highly human-disturbed areas, such as
forests in urban environments.
Some evidence appears to suggest that tabonuco, palo colorado, Pterocarpus,
and dry forests are areas harboring a high number of lichen species that may be
endemic. For instance, four species from Pterocarpus and dry forests mentioned in
this report (Coenogonium aurantiacum, C. borinquense, C. dimorphicum, and C.
portoricense) appear to be endemic (Mercado-Díaz et al. 2013). Several new species
from tabonuco, palo colorado, and sierra palm forests have been recently described
and are potentially endemic taxa (Mercado-Díaz et al., in press).
Table 4 summarizes the number of species, genera, and families of corticolous
lichens found in sampled areas of each forest type described in this report. In this
sense, these values should be regarded as relative measures of species richness; they
do not represent the total number of species found in these ecosystems. Although
limitations related to our sampling approach and other external factors may have
influenced the robustness of our estimates, we still consider this information valuable because it presents a rough idea of the diversity to be found in these forests.
An example of how limitations related to sampling may have limited our estimates
comes from the mangrove ecosystem. Namely, one of the mangrove forest plots was
considerably higher in species richness than the two other mangrove forest plots
sampled. It is thought that this higher-than-expected richness is associated with its
proximity to an adjacent Pterocarpus forest fragment. If this plot were excluded,
mangrove forests would total 17 species, 9 genera, and 7 families, less than half
of the richness reported for all taxonomic levels presented for mangrove forests in
table 4.
68
Lichens in Puerto Rico: An Ecosystem Approach
Table 4—Number of species, genera, and families of corticolous lichens occurring in sampled areas of
each forest type described in this study
Species
Genera
Families
Elin
woodland
Palo
colorado
Sierra
palm
Tabonuco
Dry
Lowland
moist
Mangrove
Pterocarpus
33
17
8
85
30
15
24
15
10
76
32
14
77
30
17
86
37
17
48
27
18
59
27
14
Acknowledgments
We thank Constance Carpenter and the International Institute of Tropical Forestry
(IITF) State and Private Forestry (S&PF) Program for their unconditional support
during the course of this project. Dr. Eugenio Santiago-Valentín, Professor of the
University of Puerto Rico (UPR) in Río Piedras and Director of the Herbarium of
the Botanical Garden (UPR), has supported Joel A. Mercado-Díaz’s work and has
provided resources and space for lichen specimens in the herbarium. Special thanks
to María Rivera, IITF biological science technician, and to Johanna Colón-López,
Carlos Castro, Benjamín Curet, Fernando “Madera” Ortiz, and Waldemar Alcobas
for their company on field trips during completion of this project. Jorge DíazMéndez provided assistance in graphic arts. We are grateful to Christian TorresSantana, Dr. André Aptroot, and Dr. Paul Bayman for their useful comments to
the report.
Funds sponsoring this report came from the IITF project “Characterizing lichen
communities along an elevational gradient in Puerto Rico: assessing their role as
indicators of forest health, biodiversity and microclimate,” led by Joel A. MercadoDíaz, William Gould, and the IITF S&PF and Forest Health Protection programs.
This work was done in collaboration with the University of Puerto Rico.
English Equivalents
When you have:
Millimeters (mm)
Meters (m)
Kilometers (km)
Hectares (ha)
Square kilometers (km2)
Degrees Celsius (°C)
Multiply by:
0.0394
3.28
621
2.47
.386
1.8 °C + 32
To get:
Inches
Feet
Miles
Acres
Square miles
Degrees Fahrenheit
69
GENERAL TECHNICAL REPORT IITF-GTR-46
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Glossary
Apothecia—Disk or cup-like ascoma of Ascomycota.
Ascocarp—An anatomically differentiated structure of asci (sac-like, sporebearing structure) and interascal hyphae enclosed within distinct layers of hyphae.
Ascomata—Fruiting bodies of Ascomycota.
Basidiolichen—Lichenized members of the Basidiomycota.
Basidiomata—Fruiting bodies of Basidiomycota.
Bioindicator—Species that can be used to monitor the health of an environment or
ecosystem.
Byssoid—Entirely composed of delicate, densely interwoven threads.
Cilia—Long-acute, multicellular hair-like outgrowths with the appearance of an
“eye lash,” generally originating from the margin or upper surface (but close to the
margin) of a thallus lobe or along the margin of apothecia.
Columella—A sterile central axis within a mature ascocarp.
Corticolous—Inhabiting the bark of trees or shrubs.
Cyphellae—Pores recessed into the lower thallus surface, surrounded by a pale
ring and lined throughout with loosely interwoven, non-gelatinized hyphae, originating from the medulla.
Crustose—General growth form where the entire thallus forms a crust.
Dimorphic—Having two growth forms.
Endolichenic—Living in close association with algal and fungal components inside
a lichen thalli.
Epiphytes—A plant or plant-like organism growing on another plant
Epiphytic—Growing on plants or plant like organisms, as an epiphyte.
Exciple—An anatomical term generally referring to any lateral layer surrounding,
embracing or enveloping an ascocarp.
Filamentous—Hair- or thread-like.
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GENERAL TECHNICAL REPORT IITF-GTR-46
Foliicolous—Growing on the surface of leaves of vascular plants
Foliose—General growth form of a “leaf-like” thallus.
Fruticose—General growth form where the thallus is three-dimensional, either
erect, pendulous (= pendent) or prostrate.
Gelatinous—Jelly-like, gel-like, with a consistency of a gel.
Isidia—A small (mostly 0.5 to 1 mm) asexual outgrowth of the thallus containing
both mycobiont and photobiont. Often with a granular, warty, pin- or finger-shaped
appearance.
Lichenization—Any complex organism composed of a fungus in symbiotic union
with an alga.
Lobule (pl. Lobules)—A small (possibly juvenile) lobe; flattened and usually
corticate on both the upper and lower side.
Morphotype—An informal group of taxa with similar or identical morphology.
Parasitism—A close association of organisms that is detrimental for one of the
symbionts.
Pendulous—Type of fruticose growth which is much stouter, not as finely divided,
and with vine-like branching pattern.
Photosymbiodeme—A lichen fungus forming morphologically and anatomically
identical (isomorphic) or different (heteromorphic) thalli with different photobionts.
Pyrenocarp—A globose to flask-shaped ascocarp opening with a pore.
Pyrenocarpous—With a structure and appearance similar to a pyrenocarp.
Rhizines—Root-like strands of closely agglutinated hyphae on the lower side of
a foliose thallus, usually thread-like to intricately branched, scant to more or less
numerous; attaching a foliose thallus to its substrate.
Soredia (pl. Soralia)—Microscopic groups of photobiont cells aggregated by
loosely interwoven hyphae, erupting from cracks or pores in the thallus surface
with a finely powdery to coarsely granular appearance.
Schizidium (pl. Schizidia)—A scaly propagule, formed by flaking off from the
thallus surface.
Squamule—A small, more or less complanate, scale-like thallus or thallus segment.
Squamulose—Forming squamules, referring to a growth form intermediate
between crustose and foliose thalli.
Symbiosis—Any form of cohabitation where two organisms form a close association, thus living together either to the benefit (mutualism) or detriment of one
another (parasitism) or without any apparent effect (commensalism).
Thallus (pl. Thalli)—The vegetative and assimilative body of both myco- and
photobiont.
Thelotremoid—Typically referring to a fruit wart commonly observed in species
of thelotremoid Graphidaceae where the thalline exciple and proper exciple are not
adherent.
76