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Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
1
Botanical observations from a threatened riverine lowland
forest in Aborlan, Palawan, Philippines
Jonah van Beijnen1,* and Edgar D. Jose2
1 Fins & Leaves, Oude Bennekomseweg 23, 6706 ER Wageningen, the Netherlands
2 Western Philippines University, Puerto Princesa City, Palawan, Philippines
*Correspondence: info@finsandleaves.org
ABSTRACT
This research provided a general overview of the vegetation
structure of the Talakaigan watershed, Aborlan, Palawan, Philippines,
with highlights on some ecological aspects of selected flora and intent
of providing urgently needed data supporting existing conservation
efforts in the area. Observations were carried during regular trekking
activities in the watershed and surrounding areas from 2009 to 2016.
Photographs were taken to facilitate species identification. Several
noteworthy observations are presented, including a new locality for
Begonia palawanensis, a short description of several new species of
Begonia and notes on a large population of the Critically Endangered
Orania paraguanensis, including details on the early life history of
these palm. A large number of anthropogenic disturbances were
observed in the area, including well-intended forestation and
development efforts by the local and provincial government. Since the
watershed does not hold any formal protective status, these
disturbances pose a serious threat to the future existence of this
unique watershed and some of the endemic species it contains. Our
findings support the call to declare the entire watershed as an official
protected area.
Keywords: floral inventory, lowland rainforest, conservation,
Southeast Asia, Begonia, Orania.
INTRODUCTION
Palawan is a large island province in the southwest of the Philippines,
northeast of Borneo. Because of the island’s low population density and the
fact that its forests contain relatively few valuable hardwood species, it has
been spared from massive deforestation that has plagued the rest of the
Philippines (Vitug 1993). Subsequently, the 450 km long island still has a
forest cover of approximately 50% (PCSDS 2015). In Southeast Asia forests
have been decimated and continue to disappear at a rapid and unprecedented
pace (Hughes 2018). The high forest cover in Palawan is thus exceptional, but
unconfirmed reports suggest that the island is quickly being deforested and
heading down the same road.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
2
Previous local bureaucratic barriers made biological research difficult
to conduct in the province and as a result, Palawan’s biodiversity and the
status of its forests are still poorly mapped and understood. However,
renewed interest in research and conservation from both the local and
provincial government has recently resulted in a number of new research
initiatives to map the island’s biodiversity. A collaborative framework of local
stakeholders have engaged in a research and conservation initiative that
focuses on the Cleopatra’s Needle Mountain Range in northern Palawan (van
Beijnen and Hoevenaars 2015; Vermeer et al. 2016). This initiative opened
the gateway for researchers to record and publish several new island records
including a very large freshwater fish (van Beijnen and Jose 2016),
rediscovered two lost amphibian species (Jose and van Beijnen 2017), and
described a number of ants and damselflies that are new to science (General
and Buenavente 2015; Villanueva et al. 2018). Moreover, recent botanical
expeditions in central and northern Palawan, resulted in the description of
several new Begonia species for the island (Hughes et al. 2010; 2011; 2018).
Other parts of Palawan, especially south of the island’s capital city of
Puerto Princesa, have been much less explored for biological research.
Updated records on the distribution, natural history and behavior of species
from these areas, particularly locally endemic ones, will be of great use to both
conservationists and policy makers. Especially given the paucity of
information on species in this region, baseline data is lacking and essential
for further studies and conservation, thereby empowering conservation
efforts.
One of the least explored areas in Palawan is the municipality of
Aborlan, located 69 km south of the island’s capital city. This municipality
covers 807.33 km2 and contains one small town, with most of the surrounding
countryside being covered by primary forest and secondary vegetation
(PCSDS 2015). This forest is part of the Victoria-Anipahan Mountain Range
that stretches from the municipality of Narra in the south up to Puerto
Princesa. In Aborlan, three rivers spring from this mountain range, of which
the Talakaigan River and its accompanying watershed contain some of the
least disturbed lowland riverine forest remaining on the island. The forested
area in this municipality is entirely listed as a Key Biodiversity Area in the
Philippines (Ambal et al. 2012) and it is part of the ancestral lands of the
Tagbanua tribe, who still depend on these forests for a large portion of their
livelihood (Sopsop and Buot Jr. 2011). However, these forests, including the
Talakaigan River watershed, largely remain unprotected and are in peril due
to the influx of outsiders that bring in unsustainable harvest practices of a
variety of forest products (Bernardo 2016; Sopsop and Buot Jr. 2011), and
the provincial government push for so-called economic development,
including a proposed hydropower (Tabuada 2015).
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
3
This research provided a general overview of the vegetation structure
of the Talakaigan watershed and to highlight some ecological aspects of
selected flora with the intent of providing urgently needed scientific data to
support existing conservation efforts in the area.
METHODS
As part of their weekly exercise routine, two biologists that are based
in Palawan made bimonthly treks through the Talakaigan watershed and
surrounding areas (Figures 1 and 2) between 2009 and 2016. The watershed
was crossed mostly at random, following the river upstream and several of its
side creeks uphill. All interesting encounters with different plant species were
photographed using a Canon 6D Camera with high-resolution capabilities
and GPS coordinates were recorded automatically. All photographs were
taken by the first author. Species identifications were verified through
consultation with taxonomic experts for each species group. The Co’s Digital
Flora of the Philippines online platform was extensively used for this purpose.
Ecological observations, including records on vegetation structure, were
noted down after each trip and stored for future reference. Based on the
observations of selected threatened and undescribed species, a family by
family, genus by genus approach were carried out to describe the sub-canopy
and understory of the study site and highlight the importance of protecting its
biodiversity. This sometimes required the team, during for example the
germination of palm seeds, to make follow up visits to specific areas in the
watershed within a short timeframe.
Figure 1. Map of the Philippines (inset), Aborlan and the location of the
Talakaigan watershed (yellow circle).
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
4
Figure 2. The upper reaches of Talakaigan River characterized by rocky
substrate, and shallow clear waters with lush vegetation along the banks.
As this research had been based on opportunistic non-professional
observations only, no specimens were hurt, mishandled, or collected for the
production of this research and besides photographs with GPS coordinates,
no actual measurements were taken.
RESULTS
The vegetation structure of the Talakaigan watershed was a
characteristic for riverine lowland forest in central and southern Palawan
(Figure 3); with the canopy along the river dominated by massive
Koompassia excelsia (Becc.), Dipterocarpus, and Ficus species. Some large
Agathis philippinensis (Warb.) are still present in the canopy layer, especially
on hill tops. The under canopy was characterized by a high density of Orania
paraguanensis (Becc.) and Barringtonia acutangula (L.), as well as some
Artocarpus species. The understory was dominated by Myrmeconauclea
strigosa (Korth.) at the riverbank, mixture of Etlingera spp., Pinanga spp.,
Caryota spp., Alocasia spp., Areca spp., Begonia spp., and a variety of other
species.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
5
Figure 3. Views of the steep riverbank (a) with assorted palm trees (b) that
characterize a large portion of the watershed of the Talakaigan River.
Arecaceae: Orania paraguanensis
A notably large population of O. paraguanensis (Becc.), consisting of
at least several hundred individuals, was encountered at the watershed. This
species seems to prefer the riverbank as they mainly occurred in close vicinity
of the river, where they grow predominantly on very sandy loam or sand,
sometimes mixed with limestone. Palm trees were observed flowering in June
and July, during the peak of the rainy season, with fruits maturing around
November (Figure 4). Trees carry several inflorescences, totaling most likely
over 100 fruits. The seeds were perfectly round, very hard, and approximately
5 cm in diameter (Figure 4c). When seeds mature, they fall off and, in most
cases, they ended up in the river because of the steeply inclining riverbank.
This method is presumably their main mode of dispersal, and this would
explain why the presence of these palm trees is concentrated around a small
number of larger rivers, underlining the importance of protecting upstream
populations.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
6
Figure 4. Different developmental stages of Orania paraguanensis: Sub-
adult tree (a), fruiting tree (b), freshly fallen seed (c), germinating seeds (d),
germinated and settled seedling (e), close-up of the trunk of a mature tree (f)
and close-up of the canopy of a mature tree (g).
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
7
Begoniaceae: Begonia palawanensis, Begonia mindorensis and
several new Begonia species
On March 20, 2015, about 5-km hike upstream along the watershed a
very small number of plants were found. Three non-flowering individuals of
Begonia palawanensis were observed. Plants measured about 10 to 15 cm tall
and this small size in combination with the absence of flowers likely indicates
that the plants were still immature. All observed plants were growing in loamy
clay, mixed with loose limestone. The plants were overhanging a steep bank
of a side-creek of the Talakaigan River (Figure 5), approximately 2 m from the
creek bed and roughly 150 m above sea level.
Figure 5. Immature Begonia palawanensis growing in-situ along a small
creek in Aborlan, Palawan. This is the first published in-situ photograph of
the species.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
8
Additionally, further upstream, one epiphytic individual that
somewhat seems to resemble B. mindorensis has been observed (Figure 6).
The plant is characterized by having long petioles (about 10-20 cm),
surrounded with reddish brown hairs. Leaves are medium-size, ovate-
lanceolate shape. We found the plant inhabiting a tree trunk shared with
dense fern-moss species. However, B. mindorensis is said to be restricted in
the forest floor. Apparently, our observation of the species could be the first
record of its epiphytic habit.
Figure 6. An undescribed epiphytic Begonia found in the area.
Besides B. mindorensis (Merr.), three other Begonia phenotypes had
been observed in this particular watershed and surrounding areas. The leaves
of this deciduous Begonia measure only 1 - 2 cm in diameter (Figure 7).
During the dry season leaves were absent and plants survived in tuberous
form.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
9
Figure 7. An undescribed miniature deciduous Begonia found in the area (a),
and a different flowering individual from the same area (b).
Zingiberaceae: Wurfbainia palawanensis
Another noteworthy finding was the endemic Wurfbainia
palawanensis (previously Amomum palawanense, Elmer) on July 13, 2015.
Several hundred individuals were found growing in a relatively open
secondary forest area on sandy clay. Some of these plants were flowering
(Figure 8). This rare, large-size ginger plants bear flowers sprouting from its
rhizomes. Our record is the first photograph and known in-situ season of its
inflorescence.
Figure 8. Inflorescence of Wurfbainia palawanensis.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
10
Araceae tribe Potheae: Pothos insignis
Several fruiting individuals of the liana Pothos insignis (Becc.) were
recorded on November 12, 2013. These fruiting individuals (Figure 9) were
part of a large group of plants that were found growing in the trunk of a large
unidentified tree just along the river. This climbing aroid was commonly
growing and climbing on large trees along the forest of the Talakaigan
watershed.
Figure 9. Fruits (a) and habit (b) of Pothos insignis.
Araceae: Homalomena palawanensis
On November 12, 2013, Homalomena palawanensis (Merr.) was also
recorded in the watershed (Figure 10). A small group of these semiaquatic
clump-forming perennials was found growing along the side of the river
where the water reached a depth of 15 – 20 cm. Aborlan, and most likely this
particular watershed, is the type locality of this species.
Miscellaneous recordings
Other plants that have been recorded in the watershed by this study
and that have been identified by consulted experts include Pothos
dolichophyllus (Merr.), Globba aurea (Elm.), Geocharis fusiformis (Ridl.),
Citrus macroptera (Mon.), Boesenbergia spp., Amischotolype marginata
(Hassk.), Leea spp., Psychotria spp., Alocasia culionensis (Engl.), Pinanga
curranii (Becc.), Caryota spp., Hoya spp., and Piper spp.
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
11
Figure 10. Semi-aquatic Homalomena palawanensis growing in shallow
water along the river.
DISCUSSION
Orania paraguanensis is endemic to Sabah and Palawan (Keim and
Dransfield 2012). It is classified as Critically Endangered by DENR
Administrative Order 2017-11 but has not been yet assessed for IUCN (2018).
Mature individuals can reach an average height of 9 to 12 m, with literature
references of up to 18 m and a trunk diameter of 20 – 30 cm (Keim and
Dransfield 2012). Almost all populations of O. paraguanensis in Palawan are
located on the east coast of the island, where higher rainfall occurs, with the
species dominating only a few kilometers from the seashore, where seeds can
be carried inland during high tide and strong winds. However, a small
number of individuals are found pretty far from the riverbanks and it could
be that the wild boar, large flying foxes, rodents (squirrels) and birds
(hornbills), of which Palawan harbors several species, disperse some of the
seeds as well. Dispersal by flying foxes has been recorded for Orania sylvicola,
a species that occurs throughout most of Southeast Asia and which has a very
similar seed size (Zona 2006). Seeds were observed to germinate rapidly after
landing on the forest floor, often within 1 or 2 weeks. The seeds germinate
remotely and the first structure to emerge from the seed is the cotyledonary
petiole (Meerow and Broschat 2017). The petiole will securely anchor the seed
at a depth of 20 – 30 cm after which it swells and develops the first root, and
eventually a single shoot. From this shoot, the first leaf will emerge and
seedlings add 2 or 3 more leaves in the two following months, before the start
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
12
of the dry season in January. It should be noted that the species has been
officially recorded for Palawan and the province of Sabah in Malaysia.
However, the presence of this species in Malaysia is based on a single
observation from 1961 (Keim and Dransfield 2012), with no other records
known. As such, it is safe to assume that the center of distribution for this
species is located in Palawan.
Since the original discovery of the species in 1910, there have been
only three recorded observations of B. palawanensis in Palawan; first in 1991
in Brooke’s Point (Hughes et al. 2010), approximately 100 km southeast from
the original type locality. Additionally in 2011 and 2014 two other
observations have been made in the vicinity of the Salakot waterfalls in Puerto
Princesa (Hughes et al. 2018). Besides B. mindorensis (Merr.) which is
restricted to the forest floor, three other Begonia phenotypes have been
observed in this particular watershed and surrounding areas. This includes a
still to be described minute deciduous Begonia (Hughes pers. comm.) that
limits its occurrence to some rocky outcrops along the river. A colleague,
Esquerion Prieto found a very similar Begonia (Figure 11) in the Inagawan
watershed, just 20 km north of the Talakaigan watershed. Actual specimens
and further research are needed to determine if this Begonia is just an
anomaly or if it represents a new species. More research is needed to
determine if they belong to the same species or if they show enough variation
to be classified as different species.
Figure 11. An undescribed Begonia from Inagawan watershed (a), and a
different flowering individual from the same area (b).
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
13
Wurfbainia palawanensis is somewhat uncommon Zingiberaceae,
and has only been recorded a few times (Nickrent et al. 2018), not because of
its rarity but more likely because it rarely flowers and is consequently often
overlooked in botanical surveys.
Pothos insignis is an uncommon aroid known only from Borneo and
Palawan (Boyce and Hay 2001) and this is the first publication of in-situ
photographs and inflorescence season of the species (Boyce pers. comm.). A
literature review confirmed only one observation of the species since its
original description: an observation that was made by Leonard Co in the same
area in 2011 (Nickrent et al. 2018).
The above observations considerably increase our insights in the
general vegetative structure of the watershed and the ecology and distribution
of several endemic and threatened species. To further update the botanical
record for this watershed, Leonard Co identified a number of other species
and genera in the watershed in 2011 that have not been published, and which
were not observed by this team. These species include: Aerva lanata (Schult.),
Pomatocalpa bicolor (J.J.Sm.), Ptyssiglottis spp., Arenga brevipes (Becc.),
Habenaria muricata (Barb. Rodr.), Rinorea spp., Jatropha gossypifolia (L.),
Mycetia javanica (Hook.), Trigonostemon villosus (Hook.) and Selaginella
spp. Photographs of these species are available from Nickrent et al. (2018).
Altogether, these recorded observations demonstrate that the study area has
interesting diversity, and includes a number of noteworthy species of
conservation importance. More studies are needed to compare the results of
this study with other sites to demonstrate how remarkable the diversity and
conditions of this primary forest are, but based on their years of experience
the authors believe this particular watershed clearly stands out.
Besides the noted species records and vegetation analysis, the team
observed a number of anthropogenic disturbances in the watershed and
surrounding areas and over the years, the severity of these disturbances has
increased. Most notably these include the well-intended forestation and
development efforts (Figures 12 and 13) by the local and provincial
government, land grabbing and consequent clearings for land speculation and
illegal quarrying activities, again highlighting the urgency for further study
and formal protection of the area. The clearing of primary vegetation to make
place for non-native trees with a higher economical value, like mahogany and
rubber, is a common development approach used by government agencies
across Palawan (CALG 2015), even in protected areas like national parks.
Although this “greening” policy stems from poorly guided but well-intended
efforts to sustain and improve the livelihoods of local residents, the impact of
this practice on the native biodiversity can be substantial. Most endemic
species, like the discussed O. paraguanensis and Begonia spp., have very
specific habitat preference and small tolerance range in terms of temperature,
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
14
humidity, and light influx (Işik 2011). When clearing the understory, the
increased influx of light, increases the temperature while decreasing the
humidity, and endemic species with their narrow niche are the first to
disappear (Işik 2011). With the extremely limited distribution range of many
Begonias in Palawan, this practice clearly has the potential to push these
species into extinction.
Figure 12. Well-intended clearing of the understory inside the Talakaigan
watershed to make place for non-native trees.
More importantly, the Talakaigan watershed and surrounding forests
are also vital to the survival of indigenous people, like the Tagbanua tribe that
are mainly dependent on the forests for their livelihood (Sopsop and Buot Jr.
2011). These indigenous people primarily use native plant species to sustain
their diet and livelihood and the practice of slowly replacing the original forest
composition with non-native species will certainly affect their livelihood
negatively. As such, it is highly recommended to the relevant government
agencies to reevaluate this counterproductive “greening” and “livelihood”
policy and modify it.
The livelihoods of indigenous people are more likely positively
impacted by officially protecting these last remaining lowland forest habitats
and by limiting the influx of informal settlers and other outsiders that often
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
15
employ less sustainable methods of extracting forest resources. Additionally,
when planting seedlings of tree species, it is highly recommended to solely use
native species that have germinated from seeds that have been sourced in the
same area. Species like Almaciga (A. philippinensis), assorted dipterocarps
and Ficus spp. are recommended as they play an important role in the
livelihood of indigenous people while also performing a key function in these
threatened forest ecosystems.
Figure 13. A public irrigation project in vicinity of the Aborlan River
watershed.
Our observations also give us the opportunity to make some
recommendations on the existing or proposed conservation status of the
discussed species. Orania paraguanensis seems to be much more common
than previously reported, which can probably be attributed to the limited
amount of fieldwork in Palawan by previous authors. Over the years, the team
has recorded the species in many coastal riverine areas along the east coast of
the island. At least three of these rivers are located in the newly established
protected area of the Cleopatra’s Needle Critical Habitat (Dasgupta 2017) in
central Palawan. However, the distribution of the species remains limited to
riverbanks of larger rivers in lowland areas on the east coast of the island, and
this specific habitat type is increasingly under pressure by anthropogenic land
conversions. Considering that the species occurs in an area over 5,000 km2,
but of less than 20,000 km2, and that the adult population most likely holds
over 2,500 individuals, based on IUCN criteria it is recommended to update
the status of this species to Vulnerable.
Begonia palawanensis has no longer been observed at its type locality
in Napsan, Puerto Princesa, and the continued occurrence of the population
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
16
from which the specimen that was collected in 1991 originated in Brook’s
Point, could not be verified despite visiting the area. This makes the
watershed of the Aborlan River and the forest around the Salakot waterfalls
in western Puerto Princesa the only two verified locations where the species
currently occurs. The lack of any formal protective status of both of these sites,
together with the large number of disturbances in the watershed, endanger
the future existence of this species. Additionally, the area surrounding the
Salakot waterfalls seems to have been opened up for development by the local
government and the recent influx of numerous settlers have resulted in
massive deforestation. With this extremely limited distribution range, the
small number of mature plants that have been observed and the large number
of threats to the species, the authors believe it is warranted to classify B.
palawanensis as Endangered, supporting the earlier proposal by Hughes et
al. (2010).
Despite the numerous treks made by the team around the entire
municipality, the miniature deciduous Begonia and the epiphytic Begonia
have both been recorded from a single location only. While the larger Begonia
with an acuminate apex has been observed in two small localities, all localities
spanning less than 1 km2 per site, and with only a handful of individual plants
observed. Considering this very limited distribution and adding the clearing
of the understory in the watershed, there is an urgent need of taxonomic and
ecological studies for the species. Results from these studies would be the
basis of recommendations for IUCN status. With our current knowledge of
these species, we argue recommending a data deficient status for each species
based on IUCN criteria. Wurfbainia palawanensis is currently listed as
Vulnerable and despite the fact that the species is uncommon throughout its
range, it has a wide distribution and few threats, thus its conservation status
is likely warranted. Lastly, the single observation of P. insignis and H.
palawanensis confirmed that both species are still present in the study site.
The lowland forest of Aborlan still holds several endemic and
threatened species and, as recorded by other studies, these forests continue
to play a vital role in the survival of the island’s indigenous people (Sopsop
and Buot Jr. 2011). With the many threats present, immediate action to
protect these forests is paramount or they, and the local indigenous people
who depend on them for their survival, will be lost forever. There seems to be
hope as the local government of Aborlan is planning to prioritize conservation
and tourism in their future development plan (WWF 2016). Furthermore,
with considerable support of the local government, a local conservation NGO
is in the process of developing a plan to protect 164.789 hectares of old growth
forest in the area (Magdayao 2018). With forests throughout Southeast Asia
disappearing at an alarming rate (Hughes 2018), support and funding for
these type of conservation projects and for more comprehensive, targeted
surveys to fully document community composition and prevalence of rare and
Van Beijnen and Jose: Endemic flora of Aborlan, Palawan
The Palawan Scientist, 12: xx-xx
© 2020, Western Philippines University
17
threatened species should be prioritized, as to hopefully safeguard Palawan’s
amazing forests for future generations.
ACKNOWLEDGEMENTS
We would like to thank Dr. Mark Hughes for identifying the Begonias
and Dr. Peter Boyce for the identification of the individual of Pothos insignis.
Gratitude is likewise due to Dr. Axel Poulsen and Rudolph Valentino A. Docot
for providing inputs on W. palawanensis, Dr. John Dransfield and Dr. Ary P.
Keim for inputs on the Orania section and Raab Bustamante for the
confirmation on H. palawanensis and his comments on the manuscript.
Thanks goes furthermore out to the Co’s Digital Flora of the Philippines
online community for providing the platform to communicate with the above
scientists. Dr. Pieter Pelser provided review and valuable inputs in the early
draft of the manuscript. Also, we would like to thank the two anonymous peer
reviewers. Finally, special thanks goes to our friend Hans Bonekamp for
joining the team on so many trips and for always bringing us home safely in
his four-wheel drive vehicle.
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ARTICLE INFO
Received: 23 October 2018
Revised: 03 September 2019
Accepted: 06 September 2019
Available online: 18 October 2019
Role of authors: JVB – conceptualized,
compiled and analyzed data, secured
expert opinions and helped in the species
identification, wrote the paper; EDJ - co-
developed the concept and wrote the
paper
