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Quest for the origin of Heliamphora heterodoxa. Report of May 2018 expedition to Venezuela with carnivorous flora occurrences in the base areas of Ptari-tepui

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In May 2018 I had the opportunity to co-organize an independent expedition to the areas surrounding the massif of Ptari-tepui and Sororopan-tepui in Estado Bolívar, Venezuela. My group led by a team of Pemón guides from the community of Santa Teresita de Kavanayen succeeded in reaching the location of the first scientific expeditions to the massif of Ptari-tepui led by W. Phelps in February 1944 and J. A. Steyermark in November 1944, including re-discovering the locus classicus of Heliamphora heterodoxa which had not been accurately located ever since the original publication of the taxon in 1951 (McPherson et al. 2011).
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183Volume 48 December 2019
Quest for the origin of
Heliamphora heterodoxa
.
Report of May 2018 expedition to Venezuela with
carnivorous flora occurrences in the base areas of Ptari-tepui
Mateusz Wrazidlo • Institute of Fundamentals of Machinery Design • Faculty of Mechanical
Engineering • Silesian University of Technology • Konarskiego 18a • 44-100 Gliwice • Poland
• matwraz@gmail.com
Introduction
In May 2018 I had the opportunity to co-organize an independent expedition to the areas sur-
rounding the massif of Ptari-tepui and Sororopan-tepui in Estado Bolívar, Venezuela. My group led
by a team of Pemón guides from the community of Santa Teresita de Kavanayen succeeded in reach-
ing the location of the first scientif ic expeditions to the massif of Ptari-tepui led by W. Phelps in
February 1944 and J. A. Steyermark in November 1944, including re-discovering the locus classicus
of Heliamphora heterodoxa which had not been accurately located ever since the original publica-
tion of the taxon in 1951 (McPherson et al. 2011).
Report of the expedition
The expedition started in the town of Santa Elena de Uairen in Estado Bolívar, Venezuela where
our group assembled on May 2nd, 2018. The following day on May 3rd we drove north on the Tron-
cal 10 Road, carried out observations of Heliamphora and other carnivorous plant populations in
the savannas by the roadside, around the marshlands in close vicinities of Rio Aponguao, and then
continued southwest to the Pemón village of Santa Teresita de Kavanayen. Along the way we made
short stops to visit the most popular touristic highlights of the Gran Sabana, such as the Jasper Creek
(Quebrada de Jaspe), Salto Kama Meru, and the famous tepui viewpoint, Mirador el Oso. After
reaching Kavanayen a short meeting was organized with the leader of the village and members of
the Parupa scientific station and plans were discussed for the upcoming days.
In the following morning on May 4th the team, led by guides from Kavanayen, advanced west-
wards and started the approach towards the southwestern slopes of Ptari-tepui. The first day of
trekking was spent on crossing the savanna to a lower forest campsite called Kavanaru (being an
Amerindian name of the cock-of-the-rock bird, Rupicola rupicola). The savanna had a notable pres-
ence of carnivorous flora typical to the Gran Sabana region, including populations of Drosera felix,
Drosera roraimae, Catopsis berteroniana, Utricularia subulata, Utricularia hispida, and Brocchi-
nia reducta. In addition, Utricularia olivacea, Drosera kaieteurensis, and Genlisea sp. plants were
found around a rest spot by a waterfall called Iwarakaru Meru approximately halfway through the
trail (Fig. 1). Within the perimeter of camp Kavanaru, several Utricularia jamesoniana plants were
spotted growing on mossy tree trunks.
The objective on May 5th was to reach Punto Phelps – a small cave campsite on the southwestern
foothill of Ptari-tepui, named after the American ornithologist William H. Phelps who was the first
to lead a scientific expedition in that area in February 1944 (Zimmer & Phelps 1944). The trail
became significantly steeper and more difficult. Before reaching the campsite, the group traversed
an elevated plateau area connecting the slopes of Ptari-tepui and Sororopan-tepui and passed by a
number of scattered patches of typical, highland moist savanna vegetation divided by Clusia sp. and
Bonnetia sessilis shrublands and savanna-forest mosaic vegetation (Fig. 2). This particular location
184 Carnivorous Plant Newsletter
reminded of the areas where some of Heliamphora heterodoxa collections had been made in No-
vember 1944 and described by J. A. Steyermark in his 1951 Fieldiana record (Steyermark 1951). No
Heliamphora plants were spotted, yet the climb provided the first encounters with members of flora
typical to higher elevations in the Pantepui – plants such as Brocchinia acuminata, Orectanthe scep-
trum, or Stegolepis ptaritepuiensis.
A particularly interesting area
was found around a rest spot by the
Iwore Meru waterfall, where the
vegetation became a lower cloud
forest characterized by the presence
of a significant number of Bromeli-
ads, Orchids, Aroids, and members
of the Rapateaceae family, such as
Saxofridericia regalis.
After reaching Punto Phelps, a
memorial plaque was found left by
W. Phelps inside the cave campsite
commemorating his visit to the spot
in February 1944. I noticed an in-
scription on the plaque made proba-
bly with a nail or some other impro-
Figure 1: Highlights of the trail to camp Kavanaru: 1) Catopsis berteroniana; 2) View over
Ptari-tepui from Iwarakaru Meru; 3) Drosera kaieteurensis; 4) Utricularia olivacea.
Figure 2: Ptari-tepui seen from one of the savanna
patches on the plateau area interlinking to the slopes
of Sororopan-tepui.
185Volume 48 December 2019
vised tool, saying: “Julian Steyermark, 10 Noviembre
1944”. That inscription served as a clear evidence
that the area was indeed the location of Steyermark’s
original Heliamphora heterodoxa collections as it cor-
responded to the dates put in the herbarium specimens
(Fig. 3).
May 6th was spent on a strenuous climb of the
southwestern slope of Ptari-tepui with the objective
of reaching as high as possible, making observations
focused on carnivorous plant populations encountered
along the way. The trail became increasingly steep,
muddy and difficult to cross. Several parts required
nearly vertical climbing using tree branches and roots
to get around steep rocky surfaces and narrow, muddy
ridges. When an exposed portion of the sandstone wall
was reached at approx. 1800 m a.s.l., a large population
of Heliamphora collina was found growing attached
to the rock wall accompanied by Utricularia alpina
and Drosera roraimae plants (Fig. 3). This observation
confirmed that Heliamphora collina is indeed present
on the cliffs of Ptari-tepui, as it had been suspected by
some other researchers (McPherson et al. 2011).
Figure 3: Highlights of the upper part of the Ptari-tepui trail: 1) Southwestern wall of Ptari-
tepui seen from Punto Phelps camp; 2) Memorial plaque left in the cave at Punto Phelps
camp, with J. A. Steyermark’s “vandalized” inscription; 3) Sandstone wall habitat with H.
collina; 4) H. collina population growing on the rock wall.
Figure 4: Heliamphora collina found
among shrub vegetation on the
slopes of Ptari-tepui. Note the
close morphological similarity to
Heliamphora folliculata leaves.
186 Carnivorous Plant Newsletter
Advancing further up the trail several other species of carnivorous plants were encountered,
namely Drosera arenicola, Utricularia quelchii, Utricularia amethystina, and Brocchinia reducta.
More high-tepui species were becoming present, such as Maguireothamnus speciosus, Brocchinia
tatei, Brocchinia steyermarkii, Pterozonium ferns, Elaphoglossum wurdackii, Ledothamnus sp. or
Connellia sp. More Heliamphora collina plants could be found growing around small patches of
vegetation on exposed rock faces and around shrubs (Fig. 4).
The trail ended on the surface of one of the large sandstone boulders which were common in the
slope areas. The summit of the boulder was partially bare rock, and in some parts thick shrub vegeta-
tion growing all around the slopes (Fig. 5), which directly matched the description of the location
left by J. A. Steyermark on his Heliamphora heterodoxa type collections: “matted on mossy exposed
top of big boulder; (…), Ptari-tepui, Bonnetia roraimae forest on southwest-facing shoulder, altitude
2000-2200 meters” (Steyermark 1951). Among the shrubs a significant amount of Heliamphora
plants were found, which after closer observations were identified as a hybrid swarm of two species,
Heliamphora collina and Heliamphora purpurascens.
Due to deteriorating weather conditions the group retreated back to Punto Phelps after spending
less than an hour on the top of the boulder and continued down to Kavanaru camp the same day.
The expedition to Ptari-tepui slopes ended on May 7th when the group returned from camp Ka-
vanaru to Kavanayen. Following days were dedicated to exploring various locations in the Gran Sa-
Figure 5: Vegetation on the top of the sandstone boulder location: 1) Bonnetia roraimae
shrub vegetation with Stegolepis, with Mona-tepui visible in the background; 2) Sandstone
boulders on the southwestern talus slope of Ptari-tepui, looking SE; 3) Shrub vegetation on
the boulder, looking NW; 4) Heliamphora collina x purpurascens growing among shrubs
on the top of the boulder.
187Volume 48 December 2019
bana, in the vicinities of Kavanayen and Parupa scientific station. Various locations were visited, for
instance forests and savannas around Salto Aponguao and Toron Meru. The expedition concluded
with a trekking from Uroy-Uaray to the community of Wuarapata, near the base of Tramen-tepui.
Acknowledgements: Members of the community of Santa Teresita de Kavanayen in Estado Bolívar,
Venezuela, especially Yoeli, Carlos, and Rosaura Chani are thanked for accompanying and provid-
ing guidance during the fieldwork that was carried out by the author on Ptari-tepui and its vicini-
ties. Izabela Stachowicz is thanked for co-organizing the expedition. Paolo and Mauro Costa are
thanked for providing useful insights about the areas around Kavanayen and Ptari-tepui. Andreas
Fleischmann is thanked for helpful comments.
References
McPherson, S., Wistuba, A., Fleischmann, A., and Nerz, J. 2011: Sarraceniaceae of South America.
Redfern Natural History Productions Ltd.
Steyermark, J.A. 1951. Sarraceniaceae. In: Contributions to the flora of Venezuela. Botanical explo-
ration in Venezuela - 1. Fieldiana: Botany 28(1): 239-242.
Zimmer, J.T., and Phelps, W.H. 1944. New species and subspecies of birds from Venezuela. 1.
American Museum novitates no. 1270: 10-11.
... The wider angle, roughly the mean estimated in uncrowded conditions (Figure 6), leads to evenly spaced pitchers in the mature rosette ( Figure 7A, B). Mature rosettes typically comprise five active pitchers (e.g., Butschi et al., 1989;Wistuba et al., 2005;Wrazidlo, 2019;Golos, 2020), and we added a sixth pitcher to demonstrate how a new pitcher would overlap with an older inactive pitcher ( Figure 7B). Furthermore, we envisioned how a clonal cluster of rosettes emerging from the rhizomes of a single plant could fill a confined space, such as a depression in the rock substrate ( Figure 7C). ...
Article
Premise The evolution of carnivorous pitcher traps across multiple angiosperm lineages represents a classic example of morphological convergence. Nevertheless, no comparative study to‐date has examined pitcher evolution from a quantitative morphometric perspective. Methods In the present study, we used comparative morphometric approaches to quantify the shape space occupied by Heliamphora pitchers and to trace evolutionary trajectories through this space to examine patterns of divergence and convergence within the genus. We also investigated pitcher development, and, how the packing of pitchers is affected by crowding, a common condition in their natural environments. Results Our results showed that Heliamphora pitchers have diverged along three main axes in morphospace: (1) pitcher curvature; (2) nectar spoon elaboration; and (3) pitcher stoutness. Both curvature and stoutness are correlated with pitcher size, suggesting structural constraints in pitcher morphological evolution. Among the four traits (curvature, spoon elaboration, stoutness, and size), all but curvature lacked phylogenetic signal and showed marked convergence across the phylogeny. We also observed tighter packing of pitchers in crowded conditions, and this effect was most pronounced in curved, slender pitchers. Conclusions Overall, our study demonstrates that diversification and convergent evolution of carnivory‐related traits extends to finer evolutionary timescales, reinforcing the notion that ecological specialization may not necessarily be an evolutionary dead end.
Sarraceniaceae. In: Contributions to the flora of Venezuela. Botanical exploration in Venezuela -1
  • J A Steyermark
Steyermark, J.A. 1951. Sarraceniaceae. In: Contributions to the flora of Venezuela. Botanical exploration in Venezuela -1. Fieldiana: Botany 28(1): 239-242.
New species and subspecies of birds from Venezuela. 1. American Museum novitates no
  • J T Zimmer
  • W H Phelps
Zimmer, J.T., and Phelps, W.H. 1944. New species and subspecies of birds from Venezuela. 1. American Museum novitates no. 1270: 10-11.