Hyperepilithics—An overlooked life form of vascular plants on tropical vertical rock walls

Abstract

Vascular epiphytes are a characteristic life form in many tropical regions and often occur growing on bare rocks. South America has the highest diversity. Here, we describe a neglected life form: hyperepilithics adapted and restricted to growing on vertical (inclination above 70°) and bare rock walls without having roots intruding the substrate. Hyperepilithics are in particular present on Brazilian inselbergs and dominated by highly specialized Bromeliaceae, mainly of the genera Stigmatodon , Tillandsia and Alcantarea , whereas Orchidaceae surprisingly has a low representation. An overview of this habitat, the life form hyperepilithics and a comparison with similar paleotropical habitats (mainly inselbergs in Western/Eastern Africa and India) are provided. Attention is drawn to hyperepilithics as a most promising and not yet exploited source for a sustainable urban ‘vertical gardening’, for example in tropical megacities.

Full text

Austral Ecology. 2023;00:1–9. wileyonlinelibrary.com/journal/aec
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© 2023 Ecological Society of Australia.
INTRODUCTION
Vascular epiphytes grow on trunks and branches of other plants without parasitizing them and are even often highly
specialist to particular host plants (Zotz,2013 ). Epilithic plants grow on rock surface and are usually treated under the
term ‘epiphyte’ without being epiphytic (Porembski,2007). Tree bark and rocks have a limited availability and storage
capability of water (Porembski,2007; Zotz,2013), requiring similar adaptations to overcome to similar stresses (Givnish
et al.,2007). Bromeliaceae, Orchidaceae and ferns have many lithophyte taxa and show high diversity in Neotropical
ecosystems (de Paula et al.,2020; Zotz,2013). This suggests that most current epiphytes may have evolved from litho-
phytes or terrestrial predecessors – or vice versa (Zotz,2013).
On tropical inselbergs, not vertical bare rock surfaces are colonized by epilithic vascular plants often forming
mats. Only few studies have addressed the diversity and ecology of mat communities (e.g. de Paula et al.,2016),
however, without taking into account the plants of vertical rock walls. Thus, we present an overview of this over-
looked habitat and the life form hyperepilithics and provide a comparison with similar paleotropical habitats.
We expect hyperepilithics to provide a promising source for a sustainable urban ‘vertical gardening’ in tropical
megacities.
RESEARCH NOTE
Hyperepilithics— An overlooked life form of vascular plants
on tropical vertical rock walls
Dayvid R.Couto1 | StefanPorembski2 | WilhelmBarthlott3 |
Luiza F. A.de Paula4
Received: 28 November 2022
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Revised: 25 April 2023
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Accepted: 27 April 2023
DO I: 10.1111/ae c.13 352
1Instituto Nacional da Mata Atlântica
(INMA), Santa Teresa, ES, Brazil
2Department of Botany, Institute of
Biosciences, University of Rostock,
Rostock, Germany
3Nees- Institut für Biodiversität der
Pflanzen, Universität Bonn, Bonn,
Germany
4Departamento de Genética, Ecologia e
Evolução, Universidade Federal de Minas
Gerais, Belo Horizonte, Minas Gerais,
Brazil
Correspondence
Dayvid R. Couto, Instituto Nacional da
Mata Atlântica (INMA), Avenida José
Ruschi, 4, Centro, 29650- 000, Santa
Teresa, ES, Brazil.
Email: dayvidcouto@hotmail.com
Luiza F. A. de Paula, Departamento
de Genética, Ecologia e Evolução,
Universidade Federal de Minas Gerais,
Avenida Antônio Carlos 6627, 31270- 901,
Belo Horizonte, MG, Brazil.
Email: luizafap@ufmg.br and luizafap@
gmail.com
Abstract
Vascular epiphytes are a characteristic life form in many tropical regions and
often occur growing on bare rocks. South America has the highest diversity.
Here, we describe a neglected life form: hyperepilithics adapted and restricted
to growing on vertical (inclination above 70°) and bare rock walls without hav-
ing roots intruding the substrate. Hyperepilithics are in particular present on
Brazilian inselbergs and dominated by highly specialized Bromeliaceae, mainly
of the genera Stigmatodon, Tillandsia and Alcantarea, whereas Orchidaceae
surprisingly has a low representation. An overview of this habitat, the life form
hyperepilithics and a comparison with similar paleotropical habitats (mainly in-
selbergs in Western/Eastern Africa and India) are provided. Attention is drawn
to hyperepilithics as a most promising and not yet exploited source for a sustain-
able urban ‘vertical gardening’, for example in tropical megacities.
KEYWORDS
Bromeliaceae, cliff ecology, epilithic, inselberg, rock outcrops, urban vertical gardening

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COUTO et al .
VERTICAL ROCK WALLS: A NEGLECTED
HABITAT FOR VASCULAR PLANTS
Vertical or almost vertical rock walls (VRWs) form a distinct habitat type
in inselbergs with an inclination between 70 and 90° (see Figure1). They
represent important elements of inselbergs, as they can reach a height
of several hundred meters. The access to them is very difficult and that
imposes great physical challenges for scientists. In the Neotropics, VRWs
are especially common on Brazilian inselbergs in the semi- arid Caatinga
(north- eastern region) and in the Atlantic Forest (south- eastern region).
The Atlantic Forest VRWs are mainly present on lowland, dome- shaped
outcrops also known as sugarloaves sensu de Paula et al.(2016, 2020).
For instance, VRWs are present in the iconic Sugarloaf Mountain in Rio
de Janeiro, south- eastern Brazil (Figure2). They are also present on the
inselbergs in the Amazon Forest (e.g. Villa et al.,2018) and in Colombia
(e.g. Aristizábal- Botero et al.,2020), but we have little information about the
VRW flora in these regions. VRWs can also occur in karst formations in the
Paleotropics (e.g. karsts of East Asia, Wang et al.,2019) and Neotropics
(e.g. Brazilian karst, Bystriakova et al., 2019), as well as in Neotropical
quartzite and sandstone rock outcrops (e.g. Tepuis from Guiana Shield,
Riina et al.,2019).
HYPEREPILITHICS: DEFINITION AND
CHARA CTE RIZ ATION
We propose the new term ‘hyperepilithics’ for vascular plants that grow
on vertical or almost vertical (inclination over 70°) rock walls (Figure3).
Although VRWs are extensively covered by lichens and cyanobacteria
(Porembski, 2007), only relatively few vascular plants grow exclusively
on this habitat (hyperepilithic specialist), and they never appear on more
gentle slopes. On the contrary, some species are not restricted to VRWs,
but are commonly found on slopes with 70– 90° inclination, which we call
facultative hyperepilithic. A notable example of a facultative hyperepilithic
FIGURE 1 Schematic overview of habitat types on inselbergs with emphasis on vertical rock walls. Credits: Gabriela Ferreira.

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HYPEREPILITHICS PLANTS ON VERTICAL ROCK WALLS
bromeliad is Encholirium horridum (subfamily Pitcairnioideae), that occur
in the Atlantic Forest inselbergs (de Paula et al.,2016). Other examples
of facultative hyperepilithic species are some orchids (e.g. Brassavola tu-
berculata, Cattleya lobata and Pseudolaelia dutrae), cacti (e.g. Rhipsalis
cereoides and Coleocephalocereus uminensis) and lycophytes (e.g.
Selaginella convoluta and S. sellowii). Finally, we highlight that many spe-
cies can accidentally occur on VRWs, as they preferentially grow on flat
to somewhat inclined (<40°) slopes, represented by example of Araceae,
Cactaceae and Orchidaceae.
NEOTROPICAL HYPEREPILITHICS: TAXONOMIC
GROUPS AND ADAPTATIONS
The centre of diversity for hyperepilithics lies undoubtedly in South
America in the inselbergs of south- eastern Brazil, a hotspot of lithophytes
plants (de Paula et al.,2020). In this region, hyperepilithic bromeliads stand
out, especially the genera belonging to the subfamily Tillandsioideae.
Within the hyperepilithic specialist members, the most remarkable ex-
ample is Stigmatodon, with most species being restricted to VRWs
(Couto et al.,2022). Equally important are species of Tillandsia subge-
nus Anoplophytum (e.g. T. araujei, T. castelensis and T. grazielae), which
generally occur in sympatry with Stigmatodon species (see Figure3b) on
large VRWs of lowland inselbergs. Furthermore, some species in the gen-
era Alcantarea (e.g. A . acuminatifolia, A. cero sa and A. robertokautskyi)
and Vriesea (e.g. V. saundersii and V. botafogensis) are also examples of
hyperepilithic specialists. Of these genera, Stigmatodon and Alcantarea
are almost exclusively epilithic (but see exceptions in Couto et al.,2022),
while Vriesea and Tillandsia are highly important in epiphytic communities
in Neotropical Rainforests (Zotz,2013). On the contrary, Orchidaceae gen-
era, such as Bulbophyllum, Cattleya and Epidendrum (similarly important
as lithophytes and epiphytes), preferentially grow on flat to somewhat in-
clined slopes in inselbergs, and are not notable in VRWs.
All hyperepilithic species possess relevant adaptations for survival on
VRWs. They are able to secure themselves firmly to the bare rock so as to
persist for long time spans and to survive long periods of drought. However,
there appear to be no specific morphological traits that are unique to hy-
perepilithics, many adaptations are similar to epiphytic plants, for example
FIGURE 2 Vertical rock walls with hyperepilithic species in the iconic Sugar Loaf Mt in
Rio de Janeiro, south- eastern Brazil. Photograph credits: Luana P. Mauad.

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COUTO et al .
CAM, succulence, presence of a velamen radicum and clonal reproduc-
tion (Zotz,2013). Mainly in Stigmatodon and Tillandsia, we note that they:
(i) are small or medium in size (ca. 5– 110 cm tall when flowering; Couto
et al., 2022; Tardivo, 2002) and display water- impounding tanks or non-
impounding; and (ii) have xeromorphic leaves that are densely lepidote on
both surfaces, which in many cases, are completely covered with a dense
layer of cinereous trichomes, giving the leaf blade a silvery colour. On the
contrary, larger hyperepilithic bromeliads (ca. 200– 370 cm tall when flow-
ering; Versieux & Wanderley, 2015), such as species of Alcantarea (see
Figure3c), comprise large water- storing tanks, that can accumulate con-
siderable amounts of water per rosette (Lehmann et al.,2022), and usually
comprise mesomorphic leaves covered sparsely with trichomes. It remains
difficult to understand how old, large and heavy Alcantarea species attach
themselves to VRWs.
Another remarkable characteristic is that, generally, hyperepilithic bro-
meliads possess prostrate stems that either grow strictly upwards (e.g.
FIGURE 3 Hyperepilithic Bromeliaceae on inselbergs in south- eastern Brazil. (a)
Stigmatodon goniorachis population growing on a vertical rock wall (VRW) (ca. 90° inclination)
in an Atlantic Forest inselberg in Rio de Janeiro city, note the stems attached to the bare rock.
(b) Tillandsia araujei (right) and Stigmatodon francae (left) on a VRW in Rio de Janeiro city.
(c) Alcantarea robertokautskyi growing on a VRW in an inland inselberg, municipality of Serra,
Espírito Santo state. (Photographs: (a,b) by D.R. Couto and (c) by E. Leme).

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HYPEREPILITHICS PLANTS ON VERTICAL ROCK WALLS
various species of Stigmatodon, Figure3a, and Alcantarea) or downwards
(e.g. Tillandsia araujei, Figure3b). Consequently, hyperepilithics slowly
creep across the vertical slopes. Further studies are necessary to better
understand the particularities of hyperepilithics.
HYPEREPILITHICS ON
PALEOTROPICAL INSELBERGS
In the Paleotropics, VRWs in dome- shaped inselbergs are frequent in
Africa, for example around Idanre, Nigeria (Figure4a), around Ambalavao,
Madagascar (Figure4c), and in India, near Bangalore (Figure4b). In this
region, VRWs are usually not colonized by vascular plants. Only very occa-
sionally mat- forming, desiccation- tolerant Cyperaceae, such as Afrotrilepis
pilosa in West Africa, and Coleochloa setifera in East Africa and Madag ascar
(Porembski,2007), can be observed as colonizers (Figure4c). These spe-
cies, however, primarily grow on flat to slightly inclined rocky slopes.
FIGURE 4 Dome- shaped inselbergs with vertical rock walls (VRWs) in the paleotropics,
usually without hyperepilithic plants. (a) VRW in Africa (around Idanre, Nigeria). (b) VRW in
India (near Bhubaneswar). (c) Rare case of Coleochloa setifera (Cyperaceae) inhabiting VRW
on a Malagasy inselberg (northern Madagascar). (Photographs: S. Porembski).

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COUTO et al .
HYPEREPILITHICS ON NEOTROPICAL
TEPUIS AND PALEOTROPICAL ASIAN
KARST FORMATIONS
Tepuis are considered a centre of richness and endemism of the neo-
tropical flora (Riina et al.,2019). In the Pantepui, two main different rocky
plant habitats are recognized: (i) the more or less flat summits that have
been extensively studied in recent decades (e.g. Riina et al.,2019) and
(ii) the vertical cliffs, that due to the difficult accessibility, almost nothing
is known about its flora (Huber & Rull,2019). It is possible that hyperepi-
lithic species inhabit these vertical cliffs, considered here as VRWs, for
example members of the Bromeliaceae. In the Pantepui, Bromeliaceae
represent ancestral lineages within the family, such as Lindmania and
Brochinia (Givnish et al., 2007). Another example is Pitcairnia paten-
tiora, a bromeliad endemic to the Pantepui and sister group to other
lineages that widely diversified in the inselbergs of south- eastern Brazil
(Givnish et al.,2007). Therefore, it would not be surprising to find hyper-
epilithic bromeliads (and perhaps other taxa) growing on VRWs of the
Pantepui.
In the Paleotropics, vertical Karst habitats seem to occur mainly in
southwestern China. There, the towers karst (fenglin) form large vertical
walls, with towers over 100 m tall and a mean slope angle of 75° (Tang &
Day,2000). In Southeast Asia, karst occurs mainly in Indonesia, Thailand
and Vietnam, we expect that hyperepithics plants may occur in these verti-
cal karst walls. However, published information is absent.
HYPEREPILITHICS AS A NEW SOURCE FOR
FUTURE SUSTAINABLE URBAN VERTICAL
GARDENING
The demand for ‘green nature’ and gardens, not only for aesthetical rea-
sons but also with regards to environmental aspects of CO2 fixation and
temperature control in our modern (mega- ) cities, dominated by bare con-
crete architecture, has led to the concept of ‘vertical gardening’. One of the
pioneers of vertical gardens was Roberto Burle Marx (Montero,2001), but
since then, the concept has worldwide gained great public attention (e.g.
Davis et al.,2016; Lotfi et al.,2020).
Usually, climbers or hanging plants in particular containers are attached
or integrated in building facades. The maintenance of these gardens is still
a challenge concerning water and nutrient supply for plants. Hyperepilithics
could open a completely new dimension for vertical gardens in the humid
tropics: The plants can grow directly on the vertical concrete wall without
continuous horticultural care or even containers. This could reveal new and
unexpected aspects for ‘future cities’.
The best candidates will probably be hyperepilithics plants, from Brazilian
inselbergs. According to our observations of natural vertical gardens, the
hyperepilithic Bromeliads are most promising for urban vertical gardening:
for example Alcantarea acuminatifolia, A. cerosa, A. robertokautskyi, A. sim-
plicisticha, Stigmatodon costae, S. euclidianus, S. goniorachis, Tillandsia
araujei, T. gardneri, T. graomogolensis, T. nuptialis, T. reclinata, Vriesea
saundersii and V. botafogensis. Possibly, some other families could be in-
teresting, for example epiphytic Cacti belonging to the genera Rhipsalis,
Epiphyllum and Hylocereus.
The selection of suitable clones and breeding work will generate op-
timized candidates. On the contrary, architects and building constructors
in cooperation with horticulturists have to develop optimized concrete

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HYPEREPILITHICS PLANTS ON VERTICAL ROCK WALLS
architectural surfaces for the vertical gardening of these plants. It is a new
challenge, but already the spatial dimension of concrete vertical walls in
megacities like Rio de Janeiro, São Paulo or Singapore justifies high efforts
for a sustainable green urban future.
CONCLUSIONS, CONSERVATION ASPECTS AND
FUTURE PERSPECTIVES
In this study, we demonstrate that VRWs represent a unique habitat type,
colonized by hyperepilithic plants, represented mainly by Tillandsioid bro-
meliads. Even though VRWs are common features on inselbergs, charac-
teristic species of this habitat are endangered due to increasing human
activities. In particular, mining and the destruction and degradation of the
flanking vegetation in the surrounding region for agriculture and urban de-
velopment (Porembski et al.,2016) are particularly damaging to VRW spe-
cies. Also, despite the difficult access to VRWs, recreational activities such
as rock climbing can result in damage to the vascular hyperepilithic flora
(Couto et al.,2022) as occurs for non- vascular plants (Clark & Hessl,2015).
Thus, climbers and mountaineering societies should be aware of this spe-
cialized plant group in the tropics and avoid opening new routes in regions
rich in hyperepilithics. Future studies should concentrate on the phyloge-
netic (but see Couto et al.,2022) and functional aspects of hyperepilithic
plants to understand why some of them are exclusive to VRWs while oth-
ers are more variable in their habitat preferences. In addition, inventories
and phytosociological data could be accessed using drones (see Lehmann
et al.,2022) and by climbing techniques that enable sampling of VRWs,
which were already described for cliff communities (Larson et al.,2000). In
these respects, special attention should be paid to regions where data on
the hyperepilithic flora is very limited, such as Amazonian inselbergs from
northern Brazil and Colombia, the quartzite and sandstone rock outcrops of
the Tepuis from Guiana Shield and the tropical Asian Karst formations. We
also see a great potential of using hyperepilithics on vertical urban gardens.
Finally, we hope to draw attention of researchers working on inselberg veg-
etation, as well as on other rocky type ecosystems, on the existence of the
life form and encourage further studies on hyperepilithics in other parts of
the world.
AUTHOR CONTRIBUTIONS
Dayvid Rodrigues Couto: Conceptualization (lead); data curation (lead);
investigation (lead); writing – original draft (lead); writing – review and ed-
iting (equal). Stefan Porembski: Conceptualization (lead); data curation
(equal); investigation (lead); supervision (lead); writing – original draft (lead);
writing – review and editing (equal). Wilhelm Barthlott: Conceptualization
(equal); data curation (equal); investigation (equal); writing – review and
editing (equal). Luiza de Paula: Conceptualization (lead); data curation
(lead); investigation (lead); supervision (lead); writing – original draft (lead);
writing – review and editing (equal).
ACKNO WLE DGE MENTS
Vertical rock walls were first noticed by S. Porembski and W. Barthlott
years ago, and their own fieldwork on inselbergs in both tropical and tem-
perate regions has been necessary to fully understand the uniqueness
of vertical rock walls as a habitat type. We would like to thank Doug S.
Larson (Guelph), Mandar Datar and Aparna Watve (both Pune) for in-
sightful discussions, which draw our attention to cliff ecosystems. This re-
sulted in our description of this new inselberg habitat type. We express

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COUTO et al .
our gratitude to Peggy Fiedler for English proofreading. D.R. Couto thanks
a research grant by the Nacional Council for Scientific and Technological
Development (CNPq, Programa de Capacitação Institucional— PCI/
INMA— 317789/2021- 0) of the Brazilian Ministry of Science, Technology
and Innovation (MCTI), and L.F.A. de Paula thanks the grants from
Conselho Nacional de Desenvolvimento Científico e Tecnológico- CNPq
(150683/2022- 7) and from Coordenação de Aperfeiçoamento de Pessoal
de Nível Superior- CAPES (88887.569558/2020- 00).
CONFLICT OF INTEREST STATEMENT
There is no conflict of interest to report.
DATA AVAILABILITY STATEMENT
Data will be available on request.
ORCID
Dayvid R. Couto https://orcid.org/0000-0002-9563-8001
Stefan Porembski https://orcid.org/0000-0003-2889-7174
Wilhelm Barthlott https://orcid.org/0000-0002-1990-1912
Luiza F. A. de Paula https://orcid.org/0000-0002-3818-7363
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How to cite this article:
Couto, D.R., Porembski, S.,
Barthlott, W. & de Paula,
L.F. A. ( 2 023)
Hyperepilithics— An
overlooked life form of
vascular plants on tropical
vertical rock walls. Austral
Ecology, 00, 1–9. Available
from: ht tps: //d oi.o r g/10.1111/
aec.13352