Travel Nature – Manu 5 day
This immense 1.5 million-hectare park has successive tiers of vegetation rising from 150 to 4,200 metres above sea-level. The tropical forest in the lower tiers contains an unrivalled variety of animal and plant species. Some 850 species of birds have been identified and rare species such as the giant otter and the giant armadillo have found refuge there. Jaguars are often sighted in the park.
Travel Nature – Manu 5 day
Travel Nature – Manu Biosphere 5 days
- Length: 5Days /4 Nights
- Type of service: Private and Group .
- Location: Southern Peru, Madre de Dios Department, Manu Tour, Manu National Park, Tours, Peruvian Amazon
- Activities: Manu Tour, Flora & Fauna, Lake Salvador , Otorongo Lake , parrot clay-lick
- Altitude: 600 – 4,000 m.a.s.l.
- Best time to visit: March – December
- Departure: All Year
- Minimum of participants: 2
- Maximum of participants: 10
- Price per person: USD
Manu Tours Biosphere 5 days .
Travel Nature Day 01: Cusco – Cloud Forest Manu Park – Pilcopata .
We pick up from your hotel very early in the morning from 5:30 a.m. at 6:00 a.m. in our comfortable and private bus. On the way, we visit the funerary towers or Chullpas de Ninamarca at 3750 meters above sea level and a brief visit to Paucartambo, a folkloric town, a colonial bridge. We ascend to the upper area of Paucartambo, the Acjanaco sector. (4000 m.s.n.m) This is the point of entry to the Manu National Park. Then we descend through the mysterious cloud forest, which hosts a great variety of flora and fauna, full of beautiful orchids, heliconias and primitive ferns. We took a walk to give you the opportunity to witness the Cock of the Rocks (Rupícola Peruviana) in the ritual of mating. There are also possibilities to observe Trogones (Trogon sp.), Quetzales (Pharomachrus sp.) In addition, much more and if we are lucky we can observe Spectacled Bear (Tremantus Ornatos). In the afternoon, we arrive at a village Pilcopata at 550 meters above sea level in our typical lodge that is outside the town with private bathrooms and showers where we will spend the night.
Travel Nature Day 02: Pilcopata – Atalaya Port -Boca Manu Cocha Otorongo Manu Park .
After the delicious breakfast we continue our trip 45 minutes by bus to watchtower on the way we observe coca plantations and fruit trees and orchids, we embark on motorized boat by the Madre, On the way we can observe a great variety of birds, such as herons, Kingfisher and always the presence of the vultures of which the most spectacular is the condor of the jungle, the king of the vultures (Sarcoramphus papa) and the Jaguar (Pantera Onca) we have a stop at Boca Manu (280 m.s.n.m) and then a short stop at the rangers stations . The boat trip continues and one begins to realize why Manu is so famous for its wildlife. On the river, there are groups of turtles, white alligators (Cayman cocodrylus) or perhaps some ronsocos (Hydrochoerus, hydrochaeris) and many more. In the afternoon we arrive at our safari camp in Cocha Otorongo (250 m.s.n.m) (double beds inside platforms) shortly after we organize a walk to visit the lake and the observation tower of 30 meters from where we have the opportunity to observe the river giant otters, turtles and birds. Here in the Manu the animals have never suffered hunting persecution by men. Night in safari camping on platforms with roof double beds with mosquito net. Showers and shared bathrooms.
Travel Nature Day 03: Cocha Otorongo – Cocha Salvador Manu Biosphere .
The group visits Cocha Salvador today, which is 30 minutes away from Cocha Otorongo. Today they explore walking in the virgin primary forest; we visit the lake to paddle silently on a catamaran that gives us the chance to observe the river wolves again and a variety of strange birds, the sultana (Porphyrula Martinica or Garza Agami). Agamia) and monkeys of different species are almost certainly observed 9 species. We later walk on the main trails with the guide to learn the operation and secrets of the tropical forest. In the afternoon we return to our camp in Cocha Otorongo. On platforms with a roof Beds with mosquito net Shared showers and baths available Optional night walk.
Travel Nature Day 04: Cocha Otorongo – Biosphere to Boca Manu .
Very early in the morning, the group will visit the MACAW CLAY LICK is in Quebrada salvadorcillo then return by the river MANU OBESERVING THE BIODIVERCITY TO BOCA MANU brief visit to Cocha Brasco to be able observe giant otters. NIGHT IN LODGE. With bathrooms and showers.
Travel Nature Day 05: Boca Manu Rainforest Lodge to Cusco .
We continue our return by boat on the Alto Madre de Dios River. On the way, we have again the option of seeing many birds and likewise with very lucky animals. This day is very sad because we have to leave our wonderful jungle full of mysteries and return by boat to Atalaya then we travel by bus to Cusco on the way we can see the Cloud Forest again, with immense variety of flora and fauna. In addition, arrive at the city at 8.30 to 9 p.m.
Fin de los servicios turisticos con Amazon-Travels.com
Included In The Travel Nature 5 days
- Motorboat transportation;
- Private vehicle land transportation for jungle trips;
- Entrance fee to the Reserve ;
- A professional cook,
- Meals: 4 breakfasts, 5 lunches, 4 dinners. and drinking water (Please note: vegetarian option available upon request for no extra cost!);
- Accommodation: 4 nights in jungle lodges;
- First aid kit;
- Radio communications for jungle trips;
- Rubber boots.
Not included in the Travel Nature 5 days:
- Flights and airport departure taxes;
- Travel insurance;
- Vaccinations for jungle trips;
- Breakfast on the first day and dinner on the last day;
- Tips to local staff.
What to take with you on the Travel Nature 5 days:
- Mosquito repellent (DEET 35 recommended as a MINIMUM!!),
- Original passport for jungle trips,
- Small backpack,
- Long sleeved cotton shirts (preferably green coloured),
- Long cotton trousers,
- Cotton long socks (to be put into your trousers),
- Comfortable walking shoes,
- Sandals or light shoes,
- Rain gear (e.g. rain poncho),
- Binoculars (also available for rent),
- Camera and charger,
- Plastic bags to be used for clothes and camera,
- Hat as a protection against the sun and/or rain,
- Small towel,
- Toilet paper,
- Antibacterial gel,
- Sun cream,
- Flashlight (with spare bulb and batteries),
- Water bottle (1 litre as a minimum),
- Pocket money (soles) to buy some beverages and souvenirs, as well as to tip.
Fin de los servicios turisticos con Amazon-Travels.com
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Travel Nature – Manu 5 day
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The goal of rapid biological and social inventories is to catalyze effective action for conservation in threatened regions of high biological diversity and uniqueness.
During rapid biological inventories, scientific teams focus primarily on groups of organisms that indicate habitat type and condition and that can be surveyed quickly and accurately. These inventories do not attempt to produce an exhaustive list of species or higher taxa. Rather, the rapid surveys 1) identify the important biological communities in the site or region of interest, and 2) determine whether these communities are of outstanding quality and significance in a regional or global context.
During social asset inventories, scientists and local communities collaborate to identify patterns of social organization and opportunities for capacity building. The teams use participant observation and semi-structured interviews to evaluate quickly the assets of these communities that can serve as points of engagement for long-term participation in conservation. In-country scientists are central to the field teams. The experience of local experts is crucial for understanding areas with little or no history of scientific exploration. After the inventories, protection of natural communities and engagement of social networks rely on initiatives from host-country scientists and conservationists.
Once these rapid inventories have been completed (typically within a month), the teams relay the survey information to local and international decisionmakers who set priorities and guide conservation action in the host country.
Region : The 216,005 hectares of intact forest in the Zona Reservada Megantoni (ZRM)
are situated along the eastern slopes of the Andes, in the department of Cusco (province of Convención, district of Echarate) in the central part of the Urubamba valley. The terrain is steep and spectacular, crossing different altitudinal gradients ranging from deep, humid canyons to the highland grasses of the puna, with forests growing on a heterogeneous mix of uplifted rocks, steep slopes, jagged mountain ridges, and middle-elevation tablelands.
Two steep mountain ranges traverse stretches of the Zona Reservada, descending from east to west. In the southwestern corner, the Río Urubamba bisects one of these ranges, creating the mythical canyon, Pongo de Maenique. Three of the Urubamba’s tributaries—the Río Timpía and the Río Ticumpinía from the north and the Río Yoyato on the southern limit—run haphazardly through the deep valleys in the Zona Reservada, carving a path among the towering ridges above them.
We surveyed three sites between 650-2,350 m. Although lowland forests harbor many more species, higher elevations tend to support more endemic species and species with restricted ranges. We chose the most inaccessible and isolated sites possible.
Kapiromashi Camp (bamboo in Machiguenga): This was the only inventory site in a large river valley. Our camp was situated in a regenerating landslide, along a small creek about 200 m from the Río Ticumpinía. The Río Ticumpinía, one of the largest rivers in the ZRM, reaches widths of 150 m or more during the rainy season. Similar to other areas in Megantoni, bamboo is pervasive at this site.
We surveyed forests growing at elevations between 650-1,200 m.
Katarompanaki Camp (Clusia in Machiguenga): At the heart of Zona Reservada
Megantoni, several massive tablelands rise between two tributaries of the
Río Ticumpinía. These tablelands are obvious on satellite images and do not appear in either Parque Nacional Manu or the Vilcabamba conservation complex. Our second campsite was on the highest of these tablelands, and we explored both this higher tier and another platform 400 m below it. This campsite was christened Katarompanaki for the Clusia tree species that dominates the canopy on the top tier of the tablelands. At this camp we surveyed elevations between 1,300-2,000 m.
Tinkanari Camp (tree fern in Machiguenga): Our third inventory site was in the
eastern corner of the Zona Reservada, close to its border with Parque Nacional Manu. Throughout the Andes and in parts of the Zona Reservada, this elevation contains some of the steepest slopes. This site was atypically flat, however, with water pooling in several places in the forest, and even forming a small (20-m diameter) blackwater pond that was not visible on the satellite image. The headwaters of the Río Timpía and the Río Manu originate several hundred meters above this site, and our trails crossed dozens of small creeks with moss-covered rocks. At this camp, we surveyed between the elevations of 2,100 and 2,400 m.
The biological communities in Zona Reservada Megantoni are an interesting mix of
species from north and south, east and west. Prior to our fieldwork, we expected to find a mix of components from the adjacent protected areas, Parque Nacional Manu and Cordillera Vilcabamba. The avifauna fit our expectations, and was a mix of these areas, but the other organisms were more closely related to communities in Manu, and some species occur exclusively in Megantoni. During our three-week field survey, we found more than 60 species new to science (more than 20 were orchids)—which is extraordinary. Habitat diversity in the Zona Reserva is extremely high.
Plants: The team registered more than 1,400 species, and we estimate that 3,000-4,500 plant species occur in the entire Zona Reservada, including lowland forest and puna species. In just 15 days, we found a surprising number of species new to science: 25 to 35. Great habitat diversity exists in the region and several plant species have very restricted ranges, confined to a certain type of soil or bedrock; these conditions may in part drive speciation. Orchids and ferns are especially diverse in the Zona Reservada and represent one quarter of all the plant species observed. Approximately one fifth of the flowering orchids we found were new to science (20 of 116 species).
Dung beetles: The team registered 71 of the 120 estimated dung beetles for the Zona Reservada. We found very few species in more than one site (and when we did, the species abundance was much greater in one site than the other). Species richness is exceptionally high in the region, even more so than in similar elevations in the Valle Kosñipata (Parque Nacional Manu). The two highest elevations we surveyed had great abundance of large dung beetles, which are more vulnerable to extinction. Secondary forests and bamboo forests had fewer species. Many of the species found have restricted elevational (and probably
geographic) ranges and are most likely endemic to the region. Some of the
(continued) Pharaeires species found were just recently described for science, some are rare, and some are new to science. In ecological terms the larger species are especially important because they recycle waste, control parasites, and disperse seeds.
Fish: In the Río Ticumpinía and numerous smaller creeks, the team registered 22 fish species. We estimate that the ichthyofauna in Zona Reservada Megantoni exceeds 70 species, the majority living in the waters of lowland forests (< 700 m) not visited during this inventory. Some of the highland species (Astroblepus and Trichomycterus) appear endemic to the area, with unique morphological adaptations to the turbulent waters of the region. All sampled aquatic habitats are in an excellent state of conservation, free of the introduced rainbow trout (Oncorhynchus mykiss) that has displaced (and in some cases, driven to extinction) native fauna in other sites in the Peruvian Andes.
Reptiles and amphibians: The herpetologist team registered 32 amphibian species
(anurans) and 19 reptiles (9 lizards and 10 snakes) in three inventory sites between 700 and 2200 m. Based on previous inventories along the same altitudinal transect in the Valle Kosñipata (Parque Nacional Manu), we estimate 50-60 amphibians occur within Zona Reservada Megantoni. We found some species in unexpected elevations (Phrynopus lower than expected and Epipedobates macero higher) and some outside of their expected geographic ranges (e.g., Syncope further south, Liophis problematicus further north). Zona Reservada Megantoni shares some of the herpetofauna with neighboring Parque Nacional Manu, but more than a fifth of the species we recorded are unique to Megantoni. We found 12 species new to science (7 amphibians, 4 lizards, and 1 snake).
Birds: The ornithologist team registered 378 species in the three inventory sites. Including species from unvisited habitats (lowland tropical forest, high montane forest, and puna) and migratory species, we estimate 600 bird species occur within Zona Reservada Megantoni. The avifauna was a mix of species from the central Peruvian Andes, some only recorded west of Cordillera Vilcabamba, and species from the Bolivian Yungas, some only recorded from Puno or on the eastern side of Parque Nacional Manu. Protecting this area would preserve the remarkably high densities of guans and macaws we observed during this inventory. In other parts of Peru, hunting of large birds, like guans and tinamous, has seriously reduced their abundance. Even in our first camp (Kapiromashi), we found signs of hunting and guans were notably scarcer. Extremely rare and local species such as Black Tinamou (Tinamus osgoodi), Scimitar-winged Piha (Lipaugus uropygialis) and the Selva Cacique (Cacicus koepckeae), which are vulnerable to extinction
(Birdlife International) and inhabit few sites worldwide, would be protected (continued) in Megantoni.
Mammals: Of the 46 expected species, the team registered 32 large and medium mammal species (belonging to 7 orders and 17 families) during the inventory. Five of these species are considered endangered and 12 are considered potentially threatened according to the Convention on International Trade in Endangered Species (CITES). In the three sites we found a large number of tracks and other signs of the spectacled bear (Tremarctos ornatus), indicating the presence of healthy populations and further stressing the importance of protecting the Megantoni corridor. The Zona Reservada Megantoni is likely an extremely important corridor for other migrating species, such as Panthera onca and Puma concolor. Conservation targets include mammals listed on CITES, Appendix I: Tremarctos ornatus, Panthera onca, Leopardus pardalis, Lontra longicaudis and
Priodontes maximus; and on CITES Appendix II: Myrmecophaga tridactyla,
Dinomys branickii, Herpailurus yagouaroundi, Puma concolor, Tapirus terrestris,
Alouatta seniculus, Cebus albifrons, Cebus apella, Lagothrix lagothricha, Tayassu pecari and Pecari tajacu.
There are 38 native communities representing four distinct ethnicities in the upper and lower Urubamba river basins, north and south of Megantoni. The Machiguenga, Ashaninka, Yine Yami, and Nanti have lived in these forests for thousands of years hunting, fishing, and cultivating their small farms. For many of them, their spiritual roots are centered in Megantoni, especially in the turbulent waters of Pongo de Maenique—the sacred place where spirits travel between this world and the next, and where the world was created. Twenty-two years ago, the indigenous people of the region formed an alliance with CEDIA to promote effective natural resource management and protect their land, its biodiversity, and the center of their spiritual world. South of Megantoni, more than 150,000 colonist settlers live in the Alto Urubamba drainage.
Along both sides of the lower Urubamba there is substantial deforestation, with larger slash and burn plots obvious on the satellite image, and evidence of colonization disappearing only at the boundary of the proposed reserve. Upriver of the Pongo de Maenique, and along the Río Yoyato on the southern side of the proposed Zona Reservada, the colonization threat from higher in the Andes is even greater, with the canyon appearing to provide at least a partial barrier to deforestation. In addition to habitat destruction, uncontrolled hunting within ZRM could threaten much of its fauna. We observed evidence of hunting impacts in our first camp, Kapiromashi.
In 1988, CEDIA (Centro del Desarrollo del Indígena Amazónico) and COMARU
Reservada Megantoni : (Consejo Machiguenga del Río Urubamba) appealed to the Ministry of Agriculture
to declare Megantoni a protected area (210,000 ha). In 1992, they prepared a technical document calling for the creation of a strictly protected area in
Megantoni, “Santuario Nacional Machiguenga Megantoni.” In 1998, INRENA passed responsibility to the Dirección Regional Agraria de Cusco (Regional Agricultural Office in Cusco) to produce information about species listed by CITES, and describe the lands neighboring the proposed protected area.
Between 1997 and 1998, the Inca Region, now known as the Cusco Regional Government, assembled local institutions to form a sustainable development plan for the entire Lower Urubamba drainage. This assembled groups strongly urged completing all pending studies before officially declaring Zona Reservada Megantoni a protected area.
In March 2004, 16 years after CEDIA began its work to protect the area, the government passed Ministerial Resolution Number 0243-2004-AG creating Zona Reservada Megantoni and incorporating it into the National System of Natural Protected Areas (SINANPE).
(a temporary designation with limited protection).
On August 11, 2004, Supreme Decree Number 030-2004-AG created the new
Santuario Nacional Megantoni based on the technical documents prepared by CEDIA, incorporating our findings. The Santuario is now an essential component of Peru’s extensive, protected biological corridor that starts in Vilcabamba, crosses Manu and Bahuaja-Sonene and then continues into Bolivia.
Principal 01 Zona Reservada Megantoni should be granted the strongest protection status recommendations possible to conserve its valuable cultural and biological resources, including species for protection and potentially endemic to Megantoni’s mountains, and to maintain the extremely
management important corridor between Parque Nacional Manu and Cordillera Vilcambamba. [Note: This recommendation has already been implemented, with the creation of the Santurio Nacional Megantoni as this report was being finalized.]
A. Within the new conservation area, we propose the following zoning recommendations:
i. Protect the area where indigenous people in voluntary isolation live, for their exclusive use (Uso Exclusivo) ii. Create a special use (Uso Especial) area for the indigenous people living in
Sababantiari that allows them to continue their traditional use of the forest. In this area, we recommend implementing a participatory community program, to monitor the impact of hunting, and if necessary, to manage hunting practices accordingly.
iii. The isolated puna habitat in the southeastern corner of Megantoni (see map above) should be strictly protected. Because it is isolated from the more extensive and interconnected puna habitat in other parts of Megantoni and PN Manu, it could harbor endemic and rare species.
iv. Promote and ensure possibilities to research intact puna habitats along the Zona Reservada’s southern border; these studies could eventually help restore and manage degraded puna in nearby areas.
v. Promote a low-impact tourism zone around Pongo de Maenique and other possible entrance points (e.g., north of the Estrella highway) to benefit neighboring communities (see map).
02 Promote the completion of the physical and legal land titling in the areas next to the Zona Reservada Megantoni.
03 Prevent public works or infrastructure construction within the fragile Zona Reservada.
04 Develop engaging and effective ways for neighboring populations to participate in the protection and management of the new protected area.
Long-term There are very few pristine areas like Zona Reservada Megantoni connecting conservation benefits puna to lowland tropical forest. These types of continuous corridors not only
contain an impressive richness of both endemic species and species of restricted altitudinal range, but they are extremely important for fauna, especially when considering global issues of climate change and deforestation.
Zona Reservada Megantoni represents a unique opportunity to expand some of the most globally important biological and cultural reserves: the Parque Nacional Manu and the protected areas of the Cordillera Vilcabamba. Immediate protection of these approximately 200,000 ha would provide an intact, forested link between two tremendously important national parks, making the total effective area of protection double the size of each individual park (a total area of more than 2.6 million ha).
Elevating the status of the Zona Reservada to Santuario Nacional Megantoni would ensure protection of thousands of species, prevent advancing deforestation, and create the only secure and intact corridor for animals migrating between Manu and Vilcabamba. The forests of Santuario Nacional Megantoni will also support and provide shelter to the Machiguenga, Ashaninka, Yine Yami and Nanti (Kugapakori) people. These indigenous people have lived in Megantoni’s forests and valleys for thousands of years and today they survive cultivating root crops, and hunting in the traditional manner of their ancestors. A strictly protected area would also preserve their cultural heritage.
Megantoni is a critical piece of the conservation puzzle in southeastern Peru. Seated on the eastern slopes of the Peruvian Andes, it fits snugly between two of the largest protected areas in Peru: Parque Nacional Manu (1.7 million hectares) and the conservation complex in Cordillera Vilcabamba (Reserva Comunal Machiguenga, Parque Nacional Otishi, Reserva Comunal Ashaninka: total area 709,347 hectares).
With 216,005 hectares Megantoni may appear small compared to its neighbors, but in rugged terrain spanning 500-4,000 meters in elevation, along steep slopes marked by massive landslides, in waters flowing through deep river gorges, on jagged mountain ridges and in nearly impenetrable patches of native bamboo, the wilds of Megantoni harbor an astonishing diversity of life. Conservative estimates place Megantoni’s plant diversity between 3,000-4,500 species, indicating that its forests may contain almost a quarter of the plant species in Peru. Many birds and mammals threatened elsewhere in Peru and South America find refuge here, and endemic species abound, around 20% of the frogs and fishes living in Megantoni do not occur anywhere else in the world.
According to the mythology of the traditional inhabitants of the region — the Machiguenga, Ashaninka, Nanti, and Yine Yami (Figure 12) — the abundant flora and fauna are protected by Tasorinshi Maeni, the spectacled bear (Tremarctos ornatus, Figure 11B). Indigenous peoples have lived in these forest valleys for millennia by cultivating root crops and hunting with bows and arrows, and their lives and fates are intimately linked to Megantoni’s wildlife and forests.
Megantoni offers the unique opportunity to link two biodiversity giants, securing protection not only to the diverse biological and cultural communities of Megantoni, but to a continuous expanse of more than 2.6 million hectares. Ill-planned colonization from the south, and gas exploration and deforestation in the north threaten the Megantoni corridor. This one-time chance to preserve intact one of the richest portions of the world depends on the fast action and long-term vision of Megantoni’s local inhabitants, its supporting organizations, and the Peruvian government.
Overview of Results
Before setting foot in the forests of Zona Reservada Megantoni (ZRM) on the eastern slopes of the Andes, we knew that our rapid inventory would focus on some of the most diverse biological communities on the planet. The Andes shelter nearly 15% of the world’s plant diversity and almost 20% of the world’s terrestrial vertebrates (~3,200 species). These mountain ranges are known not only for their species richness but also for their unique and undescribed taxa: close to half of the Andean flora and fauna is considered endemic, i.e., occurring nowhere else on Earth.
Megantoni fits the Andean mold. During our rapid inventory of its forests in April-May 2004, we catalogued nearly 2,000 species: many endemic to the region, several threatened or vulnerable in other parts of their range, and 60-80 new to science. Herpetologists found 7 new species of frogs; ichthyologists discovered endemic fishes clinging to rocks in turbulent streams; entomologists uncovered at least 30 new species of dung beetles; and botanists catalogued 1,400 plant species, including more than 400 species of orchids and ferns, with some 25 species new to science. Animals threatened in other parts of South America— including spectacled bears, tapirs, and jaguars—commonly roam the Megantoni landscape. Game birds, such as guans and tinamous, are remarkably abundant.
In the following sections we summarize the principal results of our rapid inventory within ZRM. We highlight the new species discovered in Megantoni and, for known species, the range extensions we documented during the inventory. Starting from the lowest site and moving uphill, we describe our findings at the three inventory sites, integrating information from all organisms sampled. Finally, we outline the region’s assets, and the threats to its biological and cultural riches.
NEW SPECIES AND RANGE EXTENSIONS
Before our inventory, Megantoni was almost entirely unknown to scientists, and during our rapid inventory, we encountered many of the species we suspected would live here. However, some of our finds were entirely unexpected (Table 1). For every 100 plant species we recorded, 2 are probably species new to science; for every 10 dung beetles, 1 to 4 are probably new; for every 10 fishes, 1 or 2
are probably new; for every 10 amphibians or 10 reptiles, 2 are probably new. For a 15-day inventory, these are impressive numbers and hint at the species richness that remains to be documented in the wilds of Megantoni.
Table 1. Species richness (observed and estimated) and the number of species potentially new to science for each group sampled during the 25 April-13 May 2004 rapid inventory in Zona Reservada Megantoni, Peru. Missing records are represented with a dash (–).
Number of Species
Observed Estimated Potentially
Plants 1,400 3,000-4,000 25-35
Dung Beetles 71 120 10-30
Fishes 22 70 3-5
Amphibians 32 55 7
Reptiles 19 – 5
Birds 378 600 –
Mammals 32 45 –
We discovered almost all of the potentially new species at our two higher-elevation campsites, with the exception of 1 Osteocephalus frog and ~8 new dung beetles that were found in the lowlands. For plants, the bulk of potentially new species are orchids; preliminary impressions suggest that perhaps 20 of the 116 fertile orchids collected are new to science (see Flora and Vegetation, Figure 6). Based on digital photographs we took in the field, specialists have tentatively identified 9 additional plant species, from 9 different families, as new to science.
Many of the 22 fish species we registered during the inventory are endemic to Megantoni. In particular, some species in the families Astroblepidae and Trichomycteridae have almost certainly undergone speciation within the isolated watersheds of Megantoni. At least 3 species we collected are new to science, including Cetopsis sp. (Figure 8G), Chaetostoma sp. B (Figure 8A), and Astroblepus sp. C (Figure 8D). Several species within the Trichomycteridae are potentially new as well.
We encountered 51 species of amphibians and reptiles. Slightly more than 20% are new to science: 7 anurans, 4 lizards, and 1 snake. The new amphibian species include an Osteocephalus (Figure 9E), a Phrynopus, at least 1 new species of Eleutherodactylus, a Centrolene (Figure 9H), a Colostethus, a Gastrotheca (Figure 9F), and a Syncope. We also discovered a new species of snake (Taeniophallus, Figure 9D) on the mid-elevation slopes and 4 new species of lizards (Alopoglossus [Figure 9C], Euspondylus, Neusticurus, and Proctoporus) living on the isolated tablelands in the heart of Megantoni.
Our inventory in Megantoni registered some species that were previously known only from areas more than 500 km away, as well as some species at much higher or lower elevations than previously recorded. Other groups are so poorly known for the rest of the region (e.g., dung beetles, fishes) that more data need to be collected before we can draw conclusions about endemism or range extensions.
For plants, amphibians, reptiles, birds, and mammals, we can compare some of our Megantoni findings to records from other sites in Peru and elsewhere in South America. As we continue to examine our collections and to research published reports from other sites, we expect to uncover even more geographic and elevational range extensions within the biological communities of Megantoni.
For plants, several collections in Megantoni extend the known ranges of species hundreds of kilometers farther south. At our low-elevation campsite, Kapiromashi, we registered Wercklea ferox (Malvaceae) for the first time in southern Peru. At the two higher campsites, we found Ceroxylon parvifrons (Arecaceae), Tapeinostemon zamoranum (Gentianaceae, Figure 4B), Sarcopera anomala (Marcgraviaceae), Macleania floribunda (Ericaceae), Miconia condylata
(Melastomataceae), and Peltastes peruvianus (Apocynaceae, Figure 4D), all previously known only from northern Peru.
Our collection of Heliconia robusta (Heliconiaceae) fills a large gap in the knowledge of its distribution. Known mostly from Bolivia, it has been collected only a handful of times in Peru, always in sites north of Megantoni. This Heliconia, with triangular green and red bracts and yellow flowers, dominated parts of the naturally disturbed forest around Kapiromashi.
Amphibian and reptiles
Our inventory increased the known geographic and elevational distributions of several species and even a few genera. At Megantoni we noted the southernmost distributional record in Peru for Syncope, and the lowest elevation recorded for Phrynopus and Telmatobius. We also registered an apparently undescribed species of Neusticurus, recorded previously from Santa Rosa (~800 m asl), in the Inambari basin, Puno Department, some 230 km southeast of Megantoni.
At Kapiromashi we found Epipedobates macero (Figure 9G), a rare poison dart frog restricted to the Río Purús in Brazil, Parque Nacional Manu, and the rivers in the Urubamba valley. This record extends its elevational range to 800 m from the previous record of 350 m where the species was described in Manu. While sampling in the leaf litter, we discovered a small Phrynopus cf. bagrecito, known from higher elevations in Manu but never before reported from such low elevations (~2,200 m asl).
We encountered bird species outside their published elevational ranges at every inventory site. Our records in Megantoni extend distributional limits for some species farther south, for others farther north. Three birds deserve special mention: Scimitar-winged Piha (Lipaugus uropygialis, Figure 10D), Selva Cacique (Cacicus koepckeae, Figure 10E), and Black Tinamou (Tinamus osgoodi, Figure 10C). All three species are remarkably abundant in parts of Megantoni, although exceedingly uncommon worldwide. Our records substantially increase our understanding of the distribution of these rare birds.
Our record of Scimitar-winged Piha is the second for Peru; it was previously known only from Abra Marancunca in Puno Department. From Puno, the species occurs eastward along the humid Bolivian yungas to Cochabamba Department (Bryce et al., in press). Our record is a range extension of more than 500 km to the northwest and suggests the species may occur along other mountain ranges in Cusco and Puno Departments, such as within Parque Nacional Manu.
At Kapiromashi we registered Selva Cacique, a species described from Balta, Ucayali Department, by Lowery and O’Neill in 1965, and essentially unknown until rediscovered by Gerhart near Timpía (Schulenberg et al. 2000; Gerhart, 2004; Figure 1, A13). Ours is one of a handful of sightings, and the highest elevational record for the species.
Black Tinamou has a highly spotty distribution in the Andes, with scattered records from Colombia, Ecuador, Peru, and Bolivia. This species was common at our two high elevation sites, and our records fill one of the many large gaps in our knowledge of its distribution.
We observed a group of four brown capuchin
(Cebus apella) individuals at an elevation of 1,760 m. This record is 260 m higher than the elevational range reported by Emmons and Feer (1999).
FINDINGS AT EACH INVENTORY SITE
During our 15-day inventory, we explored three sites ranging from 650 to 2,400 m in elevation, all different from one another in topography, geology, and species composition. As expected, we encountered the greatest species richness in our low-elevation site, Kapiromashi, (Table 2), and, as we climbed higher, we recorded more endemics and more new species. Collectively our findings sketch a preliminary picture of a highly diverse and heterogeneous region, where habitat types vary on scales small enough that one can walk through stunted, epiphyte-laden forests on rock outcrops, to tall forests on fertile soils, in less than an hour.
In the following sections we present a summary of our major findings, focusing on each inventory site, rather than on individual taxonomic groups as in the Technical Report (page 171). Although our inventory covered only a subset of the topographic and geological diversity in Megantoni, we believe that our inventory sites are representative of other areas within ZRM, and that the differences among them are representative of larger-scale patterns within the region.
Table 2. Species richness at each campsite, for all organisms sampled in the Zona Reservada Megantoni, Peru
Organism Kapiromashi Katarompanaki Tinkanari
Plants ~650-800 ~300-450 ~300-450
Dung Beetles 41 32 14
Fishes 17 3 5
Herpetofauna 20 19 16
Birds 243 102 140
Mammals 19 10 11
Lower mountain slopes (Kapiromashi, 650-1,200 m)
At this site in the Río Ticumpinía valley, we camped 200 m from the main river channel and explored the forested slopes on either side of the river, the large river island, the river itself, and several of its tributaries. Recent landslides, and forests regenerating on old landslides, are obvious features of the landscape. They reminded us that the area is geologically active and that natural disturbance to biological communities is frequent throughout the region. Lower-elevation sites exist in the ZRM (~500 m). Kapiromashi (650-1,200 m), however, was the lowest elevation we sampled.
We found the highest species richness for all organisms here (Table 2). Lowland and upland species overlapped at this site: species more typical of lower elevations reached their upper elevational limits, and upland species occurred at atypically low elevations, presumably because of the humidity trapped within the narrow river valley. In comparison to other sampled groups, we recorded few lowland species of fishes. Enormous waterfalls separate this part of the Río Ticumpinía from the Bajo Urubamba and presumably prevent most lowland fishes from reaching this site.
Numerous patches of large-stemmed bamboo (Guadua sp., Poaceae; known locally as paca) irregularly occur throughout Megantoni, and are especially dense in Kapiromashi. In bamboo patches, the species richness of plants, dung beetles, birds, and mammals is markedly depressed compared to that of patches free of bamboo. Clumps of bamboo, however, can harbor species that have evolved to specialize on this habitat. Such species include at least 1 amphibian (Dendrobates biolat, expected but not encountered during the inventory) and nearly 20 birds (17 recorded during the inventory).
We encountered a small patch of 8-9 plantains and old hunting trails on the southern slope of the valley, indicating that the inhabitants of Sababantiari, a community one day of travel downriver, likely hunt in this area. The near absence of several mammal species, including both species of peccaries (Tayassu pecari, Pecari tajacu) and several large primates (Alouatta seniculus, Lagothrix lagothricha), may reflect either large-scale seasonal migrations or overhunting in the area. Gamebirds, principally guans (Cracidae), were less common at this site than at the other two and, when sighted, appeared more apprehensive about our presence than the almost tame individuals spotted at our higherelevation sites. Despite local hunting, we recorded healthy populations of large carnivores (jaguar, Panthera onca) and large ungulates (tapirs, Tapirus terrestris).
Mid-elevation tablelands (Katarompanaki, 1,350-2,000 m)
Only 12 km east of Kapiromashi, broad tablelands rise between two tributaries of the Río Ticumpinía. Our second campsite was on the highest of these tablelands, and we explored both this higher tier and another platform 400 m below it. Radically different vegetation grows on each tier: on the higher platform shortstatured, low-diversity vegetation grows on hard acidic rock; the lower tier supports taller, higher-diversity forest on much richer soils. We observed differences in composition and richness between the two platforms in all organisms. Richness was greater on the lower platform; in many groups, a more specialized community lives on the upper platform.
Specialization was most obvious in fishes. Fast-flowing streams feed the dramatic waterfalls that pour over the sheer edge of the tablelands into the river gorges below. Few fish species live in these streams, but the three endemics we registered during the inventory are abundant and uniquely adapted to the turbulent waters, using their adhesive mouths to cling to rocks, and their ventral muscles to pull their bodies upstream against the current.
As did the specialists in fishes and dung beetles, herpetologists found few species but many endemics. Nutrient-poor forests are generally unfavorable habitats for amphibians and reptiles, and on the upper platform the team found only 16 species: 8 anurans, 3 lizards, and 3 snakes. Nearly half, however, are species potentially new to science—3 lizards (Euspondylus, Neusticurus, Proctoporus) and 3 frogs (Centrolene, Eleutherodactylus, Syncope)—suggesting that these isolated tablelands could drive evolution in fishes, frogs, lizards, and dung beetles in similar ways.
Plant diversity—concentrated in trees and shrubs in Kapiromashi and on the lower platform of Katarompanaki—shifted to smaller lifeforms on the upper platform at Katarompanaki. Here, the highest richness was concentrated in epiphytes and trunk climbers, particularly orchids and ferns. Of the 275 fertile specimens on the tablelands, a quarter were orchids, including about 15 species new to science. In other areas of Peru (e.g., Cordillera del Cóndor, Cordillera Azul), stunted forests support a suite of specialized bird species, but we did not encounter these elfin-forest specialists at Katarompanaki. Ornithologists documented only moderate numbers of bird species at this site, although the densities of game birds, particularly the typically rare Black Tinamou, were extraordinarily high.
We found numerous signs of spectacled bear (Tremarctos ornatus) in the stunted forest, including trails, dens, and discarded and half-eaten palm stems. Our Machiguenga guides estimated that bears were in the area three months prior to our visit, confirming other research that suggests these animals travel widely through large territories, tracking seasonal fluctuations in food abundance.
On the lower platform, species richness in all groups was much higher, although researchers spent less time investigating this area. Most notable were the abundance of fruiting trees and the extraordinary densities of woolly monkeys (Lagothrix lagothricha) feeding on them, including an enormous group of 28 individuals.
We believe humans have never visited this site before. Reaching the tablelands without a helicopter appears nearly impossible.
Mid-elevation slopes (Tinkanari, 2,100-2,400 m)
Our third inventory site was in the western corner of the Zona Reservada, close to its junction with Parque Nacional Manu (Figure 3B). The headwaters of the Río Timpía and the Río Manu originate several hundred meters above this site, and our trails crossed dozens of small creeks with moss-covered rocks (Figure 3K). This site was atypically flat, however, with water pooling in several places in the forest and forming boggy areas.
As at Katarompanaki camp, we distinguished two forest types at this site. A tall forest on richer soils dominates 90% of the area and surrounds a neatly delimited area (~0.5 km2) of stunted shrub forest growing on a much harder acidic rock. The stunted shrub forest was obvious from the air and was similar to other outcrops on acidic rock seen during the overflights of the Zona Reservada.
Signs of spectacled bear were common and widespread in the stunted forest and ranged from trails and dens to recent food remains and fresh scat. Spectacled bears were one of the most abundant mammals we recorded in Megantoni, second only to woolly monkeys. Moreover, our Megantoni surveys recorded the highest relative density of spectacled bear reported in any Peruvian inventory.
Again, game birds were abundant and tame, including Sickle-winged Guan (Chamaepetes goudotii), Wattled Guan (Aburria aburri), and Andean Guan (Penelope montagnii). At this site, ornithologists photographed Scimitar-winged Piha (see Range Extensions, above) and tape-recorded calls and a flight display. We believe this flight display has never been witnessed before.
We found several new species and range extensions for plants at this site. Ferns dominated these forests (Figure 5) with high richness (~30 species/100 m2) and high densities, especially of tree ferns (~2,000 individuals/ha). As in the Katarompanaki tablelands, species richness was concentrated in epiphytes rather than trees and shrubs.
Amphibians and reptiles showed patterns of diversity parallel to those of fishes, as they did at Katarompanaki. Species richness was limited overall, but several novelties and endemics dominated the community. Ichthyologists found high fish densities in all streams sampled, registering 5 species of fishes, including 2 Astroblepus not found at Katarompanaki. Herpetologists recorded 10 species of anurans, 2 lizards, and 4 snakes. One of the most notable records, Atelopus erythropus, previously was known only from the holotype and populations in the Kosñipata valley. The largest of all frogs found at this site was an arboreal marsupial frog, Gastrotheca sp. (Figure 9F), similar to G. testudinea (W. Duellman, pers. comm.). Gastrotheca sp. was nearly ubiquitous—males sang from the canopies in almost every habitat—and this species is almost certainly new to science.
In contrast to the biological communities, the social landscape was well known before our inventory. For more than two decades, CEDIA and other organizations have engaged in participatory work with many communities in the region, and their efforts, in conjunction with the long-term vision of many of the native inhabitants, inspired the proposal for a protected area in Megantoni.
To date, CEDIA’s efforts have focused largely on the traditional inhabitants of the region— the Machiguenga, Ashaninka, Yine Yami, and Nanti.
However, two distinct cultural groups live in the area surrounding Zona Reservada Megantoni: native populations living in communities and colonists living in rural settlements (see Figure 1). These groups are coarsely separated within the landscape. The bulk of the native peoples live north rather than south of Megantoni (12,000 vs. 4,000 people) and inversely, most colonists live south rather than north of ZRM (150,000 vs. 800 people). Native peoples practice subsistence agriculture and have lived in these forests for millennia, while colonists are more recent arrivals, and typically practice larger-scale commercial agriculture. A large part of the long-term success of a protected area in Megantoni will rely on stabilizing the agricultural frontier, particularly in the south, and engaging both native inhabitants and colonists in the protection and management of the region.
CONSERVATION RISKS AND OPPORTUNITIES
The conservation landscape we propose for Megantoni will provide long-term, strong protection for a biologically and culturally rich region, and is an unparalleled opportunity to
01 Protect unique flora and fauna, including the 60-80 species new to science found in Megantoni,
02 Link two large protected areas, ascending from the Amazonian plain in Parque Nacional Manu to the Andean slopes in the Cordillera Vilcabamba,
03 Preserve a landscape sheltering uncontacted indigenous communities, living in the extreme northeastern corner of Megantoni, and
04 Work with neighboring communities in designing ecologically compatible activites (including wellmanaged ecotourism) that will reinforce the protection of Megantoni.
The isolation and ruggedness of Megantoni, the collective knowledge of its inhabitants, and the biological and cultural riches within its borders are enormous assets for conservation in the region. Here we detail several of the most striking and general conservation assets within ZRM, although undoubtedly many more exist.
Several particularly well-preserved and unique habitats exist within ZRM. Elsewhere in the Peruvian Andes, high-altitude grasslands (puna) experience intensive land use, overgrazing, and overburning, and upland streams are populated with invasive, exotic rainbow trout that have decimated native fishes. Megantoni provides an opportunity to preserve the full richness of this intact mountain flora and fauna and could provide a living reference for restoration efforts in degraded grasslands and aquatic habitats in nearby areas.
Traditional knowledge/cultural richness
These forests are intimately familiar to the
Machiguenga, Nanti, Ashaninka, and Yine Yami. Collectively, these groups safeguard a wealth of traditional knowledge—an understanding of animal movements and behaviors, seasonal fluctuations in weather and resources, favorable planting times and ecologically sensitive cultivation methods— providing the closest approximation to a communal almanac for the region.
Pongo de Maenique
The rough waters and life-threatening whirlpools and rapids of the Pongo de Maenique are a spiritual center for the traditional inhabitants of the region, and separate the Alto Urubamba from the Bajo Urubamba. Although now navigable, for centuries the Pongo shielded the Bajo Urubamba from development or colonization.
Today the Pongo remains an asset, continuing to play a deep spiritual role in the lives of traditional inhabitants of the region, and providing spectacular ecotourism opportunities for native communities.
The forested ridges and valleys of Zona Reservada Megantoni are difficult to reach—they require three full days of travel from Cusco, via planes, boats, and trails—and their isolation has spared Megantoni the deforestation common in many parts of the Andes.
Among the major threats to Zona Reservada Megantoni are the following:
Rampant, ill-planned colonization
Colonists have settled on steep, landslide-prone slopes. Conservation-compatible cultivation is impossible in these areas. Typically, colonists move from one unsuitable area to the next, barely eking out an existence, and deforesting vast areas in the process.
Natural gas pipeline development The extraction of hydrocarbons is perhaps the largest threat to ZRM, as the Camisea gas operation lies just ~40 km north of the Zona Reservada. Natural gas extraction in the area has already forced native communities to leave their traditional lands, and the next few years may see increased exploration for gas deposits along the Bajo Urubamba.
Enforcing forestry laws is nearly impossible in such a remote area, and illegal loggers have extracted timber from areas to the north of ZRM (e.g., Reserva del Estado a Favor de los Grupos Étnicos Kugapakori-Nahua).
OVERVIEW OF INVENTORY SITES
Zona Reservada Megantoni is an intact wilderness corridor of 216,005 ha on the eastern slopes of the Peruvian Andes, widest at the eastern end along its broad border with Parque Nacional Manu, and tapering to a narrow wedge at its western limit where it joins with the Vilcabamba conservation complex (Reservas Comunales Machiguenga and Ashaninka, and Parque Nacional Otishi, see Figure 1). Elevation decreases along a similar westward trajectory. From higher-altitude grasslands (up to 4,000 m) restricted to the southeastern end of the area, the landscape descends in a spectacular series of sharp ridges and rugged slopes until it reaches the river valley bottoms (500+ m) of the lowlands in the west.
In the southwestern corner, the Río Urubamba bisects a large ridge, creating the mythical Pongo de Maenique canyon and exposing clay licks used by Military Macaws (Ara militaris) and spider monkeys (Ateles sp.). Three of the Urubamba’s tributaries—the Río Timpía and the Río Ticumpinía in the north, and the Río Yoyato along the southern boundary—originate within the Zona Reservada, as do the headwaters of the Río Manu (see Figures 1,2).
Much of Zona Reservada Megantoni is covered in patches of live and dead bamboo: Guadua species (Figure 3E) at lower elevations and Chusquea species and their allies at higher elevations. In some places the bamboo creates a nearly impenetrable, monodominant stand, whereas in others the bamboo species is draped on and around several tree species, usually only a small subset of the diversity in the surrounding forest.
Pronounced patchiness characterizes the entire Zona Reservada Megantoni. Over short distances—as small as several hundred meters—habitats can change from stunted shrub forests growing on exposed acidic rocks, to forests growing on richer soils with a canopy taller by nearly tenfold, with little or no overlap in species composition between the two areas.
Since the Zona Reservada contains extraordinarily high habitat
heterogeneity, both horizontally, at small spatial scales, and vertically, along an altitudinal gradient, our goal in selecting biological inventory sites was to sample the habitat diversity to the greatest extent possible.
SITES VISITED BY THE BIOLOGICAL TEAM
We combined our observations from the November
2003 overflight and our interpretations of Landsat TM+ images (bands 4, 5, 3, and 8 panchromatic) to select inventory sites at different elevations, trying to include access to a range of altitudes and habitats at a single site (see Figures 2, 3). Because of the rugged terrain, access to highland sites in Megantoni is challenging, and in many cases impossible. For our two higher-elevation inventory sites, the advance team—who cut trails and prepared camp before the inventory—gained access to the sites by descending a cable from a hovering helicopter.
During the rapid biological inventory of Zona Reservada Megantoni from 25 April to 13 May 2004, the inventory team surveyed three sites spanning a 1,700-m altitudinal gradient, starting from 650 m above sea level (asl) and reaching 2,350 m. Below we describe these three sites in more detail and include information on a fourth site that was visited only by the advance trail-cutting team. Each site name, in Machiguenga and chosen by the Machiguenga guides accompanying us on the inventory, represents an obvious and dominant feature of the vegetation.
Kapiromashi (12°09’43.8”S 72°34’27.8”W, ~760-1,200 m asl, 25-29 April 2004)
This was the first site we visited, and the only one in a large river valley (Figures 3A, 3C). Our camp was situated in a regenerating landslide, along a small (5-m-wide) transient stream about 200 m from its junction with the Río Ticumpinía. Although the Río Ticumpinía measured ~40 m across during our stay, it is one of the largest rivers in Zona Reservada Megantoni and can span 150 m or more when it is fully charged with water.
Our Machiguenga guides from Timpía, a community 28 km to the northwest at the junction of the Río Timpía and the Río Urubamba, had never visited this site. However, we encountered a small patch of 8-9 plantains and old hunting trails on the southern slope of the valley, indicating that the inhabitants of Sababantiari, a community one day of travel downriver, likely hunt in this area.
Over four days we explored more than 12 km of trail on either side of the Río Ticumpinía valley, often walking for more than 0.5 km along the rock-strewn, sandy beaches to reach one of the few places where we could cross the river. One additional trail traversed a large island formed where the river diverges and rejoins itself 1.5 km downriver.
Our trail system reached the crest of the southern ridge around 1,100 m asl. Although the ridge on the opposite side of the river appeared to extend to at least 1,500 m asl, we could not reach areas above 1,200 m asl on this higher ridge. Clouds typically moved from the south over the lower ridge and settled against the northern slope, forming a cloud bank around 1,100 m asl. In general, the area contains exceptionally humid forest. Nonetheless, while we were in the field no large downpours occurred, several streams dried up and drought stress was evident in orchids on the northern slope.
Kapiromashi means “much bamboo” in Machiguenga and is the word our local guides used to describe the impressive patches of Guadua bamboo (Figure 3E) that dot slopes on both sides of the river, as well as the river island. All trails contained at least one patch of Guadua bamboo, and several traversed upwards of 80% bamboo-dominated forest (pacal). We found evidence of natural disturbance on most of the trails, often walking through a time series of forest in different stages of recovery from old and new landslides, with more mature forest marked by larger-sized tree stems and their greater epiphyte loads. Underlying this matrix of disturbance is a mosaic of limestone-derived and more acidic soils sometimes separated only by tens of meters. Several plant species are restricted to only one of these soil types.
Judging from our overflight of the area and the satellite images, this area is likely representative of the habitat along the Río Timpía inhabited by the voluntarily isolated Nanti people (Figure 12E).
Katarompanaki (12°11’13.8”S 72°28’13.9”W,
~1,300-2,000 m asl, 2-7 May 2004) At the heart of Zona Reservada Megantoni, several massive tablelands rise between two tributaries of the Río Ticumpinía (Figure 3A). These tablelands are obvious on satellite images and do not appear to occur in either Parque Nacional Manu or the Vilcabamba conservation complex. Our second campsite was on the highest of these tablelands, and we explored both this higher tier and another platform 400 m below it. This campsite was christened Katarompanaki for the Clusia tree species (Figure 3G) that dominates the canopy on the top tier of the tablelands.
Although from the air the area appears to be a flat, slowly ascending surface, on the ground the surface is uneven and crisscrossed by a network of small streams that carve deeply into areas of softer substrate. At each stream crossing, the trails descended and rose sharply. The two largest streams (10-20 m wide), one on each tier, consisted of enormous, entire slabs of rock, and were composed of such a hard substrate that scratching the surface, even with plant clippers, was nearly impossible.
Radically different vegetation grows on each tier. On the higher platform short-statured, low-diversity vegetation grows on hard acidic rock. The lower tier supports taller, higher-diversity forest on much richer soils. Because of the slow rates of decomposition, the forest floor on the upper tier is a treacherous tangle of roots and fallen trees, distinctly spongy, and sprinkled with large holes more than 1 m deep. We found little evidence of mineral soil, although a humic layer is present. On the lower tier, the forest is more productive and the richer clay soils support several fruiting species and a substantial mammal fauna.
Traveling between the two tiers was difficult, with a frighteningly vertical descent in some spots. Once on the second tier, the trail passed below a spectacular waterfall pouring over the lip of the first tier, the water cascading past 40 m of vertical rock to crash directly onto the second tier below.
On the few cloud-free days, the southern edge of the top platform granted researchers spectacular views of a string of triangular slabs known as Vivian formations to the west (Figure 3F), a jumble of steep ridges to the south, several jagged peaks to the east and, across the expanse of short-statured forest of the platform to the north, a sheer rock wall rising from the other side of the river valley. The river island of Kapiromashi camp— a mere 12 km to the west—was distinctly visible from the southwestern corner of the higher platform.
During our six days at this site, we experienced several localized downpours, with intense steady rain in one spot, and blue skies 1.5 km away.
We found no evidence that humans had ever visited this site and the density of woolly monkeys (Lagothrix lagothricha, Figure 11C) in the lower-tier forest was noticeably high.
Tinkanari (12°15’30.4”S 72°05’41.2”W, ~2,1002,350 m asl, 9-13 May 2004)
Our third inventory site was in the western corner of the Zona Reservada, close to its junction with Parque Nacional Manu (Figure 3B). Throughout the Andes and in parts of the Zona Reservada, this elevation contains some of the steepest slopes. This site was atypically flat, however, with water pooling in several places in the forest, and even forming a small (20-m-diameter) blackwater pond that was not visible on the satellite image.
The headwaters of the Río Timpía and the Río Manu originate several hundred meters above this site, and our trails crossed dozens of small creeks with mosscovered rocks (Figure 3K). A creek formed by a recent landslide (huayco), the largest waterway at this site, provided us with a window on the complicated geology of the area. Walking upward along the landslide, on rocks that were still free of moss, we observed different strata within the exposed rocks, alternating hard sandstone with other layers of substrate, including shale, and even carbon.
The streams in the area often descended stepwise, with a flat stretch and a steep descent followed by another flat stretch. Our working hypothesis is that the flat stretches reflect softer substrates that erode quickly, or alluvium, followed by harder sandstone, and then the next layer of softer substrate.
As at Katarompanaki camp, we distinguished two forest types at this site. A tall forest on richer soils dominates 90% of the area and surrounds a neatly delimited area (~0.5 km2) of stunted shrub forest growing on a much harder acidic rock. One forest type abutted the other, with no transition.
The stunted shrub forest was obvious from the air and was similar to other outcrops on acidic rock seen during the overflights of the Zona Reservada. The lower portion of the shrub forest was even shorter, dominated by terrestrial orchids and a thin-stemmed Clusia sp. In addition to our cut trails, a grid of spectacled bear trails traversed the stunted forest.
More than ten species of tree ferns, or tinkanari (Figure 5A), dominated the higher forest, in addition to several species and relatives of Chusquea bamboo.
Shakariveni (12°13’08.9”S 72°27’09.1”W, ~960 m asl, 13-19 April 2004)
About 13 km east of Kapiromashi camp, and directly below the large tablelands of Katarompanaki camp, the advance team established a campsite at the junction of the Río Shakariveni and a small tributary. From here, they spent six days exploring the region, hoping to reach the tablelands above. During their unsuccessful efforts to reach higher ground the team observed several vertebrates that are included in the appendices. Close to their campsite, the team encountered an abandoned farm plot (chacra), potentially cleared by colonists entering the Zona Reservada from the south. This area closely resembles Kapiromashi camp in its matrix of forest containing large areas dominated by Guadua bamboo patches, as well as in the successional flora along the rocky riverbed.
OVERFLIGHT OF ZONA RESERVADA MEGANTONI
Authors: Corine Vriesendorp and Robin Foster
ZONA RESERVADA MEGANTONI
Situated on the eastern slopes of the Andes, the rugged, spectacular terrain of Zona Reservada Megantoni ranges from deep, humid canyons to moist, high-altitude puna grasslands. Formed during the geological turmoil associated with the uplift of the Andes, the forests within Megantoni grow on a heterogeneous mix of uplifted rocks, steep slopes, jagged mountain ridges, and flat tablelands ranging in elevation from 500 to 4,000+ m.
Two steep ridges traverse stretches of the Zona Reservada, descending from east to west. The Río Urubamba bisects one of these ridges in the southwestern corner, creating the Pongo de Maenique river gorge (see Figures 2, 13). The Urubamba’s tributaries (principally the Río Timpía, the Río Ticumpinía, and the Río Yoyato) run haphazardly through the deep valleys in the Zona Reservada, carving a path between the towering ridges above them.
Along both sides of the lower Urubamba, deforestation is substantial, with larger slash-and-burn plots obvious on the satellite image, and evidence of colonization disappearing only at the boundary of the reserve (Figure 1). Upriver from the Pongo de Maenique, and along the Río Yoyato on the southern side of the Zona Reservada, the colonization threat from higher in the Andes seems even greater. The river gorge appears to provide at least a partial barrier to deforestation.
On 3 November 2003, a team of scientists from
CEDIA, INRENA, CIMA, PETT and The Field Museum flew by helicopter over the rugged terrain of Zona Reservada Megantoni. The flight route traversed an impressive altitudinal gradient, starting from the lowlands (300+ m asl) in the northwestern corner of the ZRM, crossing over expansive table mountains and isolated ridges (1,000-2,000 m asl) near the center, and reaching the highlands (4,000+ m asl) in the southeastern edge. Below we complement the satellite images with our observations from the overflight, focusing specifically on obvious changes in vegetation and habitat within the area.
From the Pluspetrol base in Malvinas, we followed the meandering Río Urubamba upriver to a narrow tongue of steep ridges extending in a long line on either side of the Pongo de Maenique. A striking contrast exists between the northern and southern faces of these ridges. The northern faces usually are covered in immense, scrambling patches of bamboo (Guadua spp.) and the southern faces in vegetation of much higher diversity. The nearly complete cover of bamboo on the northern faces of these ridges suggests that a catastrophic disturbance, such as a massive fire or an earthquake, might have cleared competing vegetation and promoted bamboo colonization in the past.
On some of the northern faces, open patches suggest that tall forests have collapsed under the weight of the Guadua bamboo, creating a mixture of Iriartea deltoidea (Arecaceae), Triplaris americana (Polygonaceae), and Cecropia sp. (Cecropiaceae) crowns amidst the bamboo tangles. Even where tall forest is present, the understory appears to be dominated by bamboo. At higher elevations, the Guadua bamboo is confined to small, disturbed areas and is eventually replaced by Chusquea and other small bamboo species.
In contrast, a much more diverse vegetation grows on the southern faces of the ridge, sporadically interrupted by stunted forest on quartzite outcrops, and by vast landslides colonized by a suite of fast-growing species. The ridgetops separating the two faces often support monodominant patches of forest, likely reflecting the poor growing conditions on these exposed, older soils.
Farther eastward, the ridge is interrupted by a string of triangular slabs known as Vivian formations (Figure 3F), with slopes gently rising toward their apex on one side and abruptly falling along a sheer rock face on the other. On their slopes, Vivians support a variety of stunted vegetation, and sometimes are covered in bamboo. After nearly 30 km the Vivians disappear and are replaced by a series of expansive tablelands.
From the flat tops of the tablelands, dramatic waterfalls pour over sandstone cliffs into the river gorge below. Vegetation on the tablelands is variable, usually dominated by atypically short Dictyocaryum lamarckianum palms mixed with other stunted trees. At least one shelf of the tablelands is dominated by monocarpic Tachigali (Fabaceae) trees, both alive and recently dead. Steep slopes on the higher mesas at
1,500-2,000 m asl are dotted with Alzatea (Alzateaceae) trees often mixed with tall treeferns (an ideal habitat for Andean Cock-of-the-Rock, Rupicola peruviana).
From the mesas we descended to the river confluence of two tributaries of the Río Timpía, passing through narrower and deeper valleys with steep slopes. Along the isolated but broad valley of the largest tributary of the Timpía we observed 10-15 small plots with plantains and thatched-roof shelters, confirming the previously suspected presence of voluntarily isolated groups of Nanti in this area.
Cliffs and steep slopes with landslides continue up to the highest point in the southeastern corner of the reserve, where the ridgetops are more gently sloping. The trees are shorter, twisted, and covered with lichens, giving way to the high-altitude puna grasslands intermixed with patches of shrubby forest composed principally of Polylepis (Rosaceae) and Gynoxys (Asteraceae). The puna is dotted with scattered tarns and has a mix of giant Puya bromeliads and other herbaceous flora, along with the grass cover.
Although we heard reports of this area being used for grazing, we did not see cattle paths from the air. We did see evidence of recent fires, with blackened stems dotting several ridge tops, but we believe the fires to be natural. Cove forests along small highland streams appear to act as natural firebreaks.
From here, we descended along the southern edge of the ZRM, flying over successively lower crests on our return to Malvinas.
FLORA AND VEGETATION
Participants/Authors: Corine Vriesendorp, Hamilton Beltrán,
Robin Foster, Norma Salinas
Conservation targets: Hyperdiverse Andean plant families, especially orchids and ferns, along an altitudinal gradient from lowland forest to puna; small populations of timber trees at lower elevations (Cedrela fissilis [cedro]; Cedrelinga cateniformis [tornillo]); large tracts of bamboo-dominated forest; pristine expanses of high-altitude grasslands; stunted shrub forests on acidic rock outcrops; and the more than 25 plant species endemic to Zona Reservada Megantoni
Before setting foot in the forests of Zona Reservada Megantoni (ZRM), we knew that our rapid inventory would focus on some of the most diverse plant communities on the planet. Considered “the global epicenter of biodiversity” (Myers et al. 2000), the tropical Andes shelter nearly 15% of the world’s plant diversity within their slopes, peaks, and isolated valleys. Moreover, close to half of the Andean flora is likely endemic, i.e., occurs nowhere else in the world.
Andean forests are still poorly understood from a floristic standpoint, and our botanical knowledge of the distribution, composition, and dynamics of these dauntingly diverse forests remains rudimentary. During this inventory, our closest points of comparison were the protected areas adjacent to Megantoni, Parque Nacional Manu to the east, and the Cordillera Vilcabamba
(Parque Nacional Otishi, Reserva Comunal Machiguenga, Reserva Comunal Ashaninka) to the west.
Although Manu is one of the best-studied sites in South America (Wilson and Sandoval 1996), most research has focused on elevations lower than those at any site within Zona Reservada Megantoni (500-4,000 m asl). Botanists have collected in the Kosñipata valley in Manu from 2,600 to 3,600 m asl and have generated a preliminary list of the flora (Cano et al. 1995). Recently, Miles Silman, N. Salinas, and colleagues have established several 1-ha tree inventories from lowlands to treeline within the Kosñipata valley of Manu (7003,400 m asl). These plots are more comparable to our inventory sites than are floristic studies from Cocha Cashu in Manu (Foster 1990). To the west of ZRM, there is some sampling overlap between our inventory sites (650 m, 1,700 m, 2,200 m asl) and the rapid inventory in the Cordillera Vilcabamba (1,000 m, 2,050 m, 3,350 m asl; Boyle 2001).
Finally, to the north of ZRM, scientists working with the Smithsonian Institution documented an intact, highly diverse forest mixed with bamboo as part of the biodiversity surveys and environmental impact assessments for the Camisea natural gas extraction project (Holst 2001, Dallmeier and Alonso 1997). These forests in the lower Urubamba valley are on hills lower and drier than those in Megantoni, and forests similar to these are protected in Reserva Kugapakori-Nahua, which abuts ZRM on its northeastern border.
To characterize plant communities at each inventory site, the botanical team explored as many habitats as possible. We used a combination of general collections, quantitative sampling in transects, and field observations to generate a preliminary list of the flora (Appendix 1).
During our three weeks in the field, we collected 838 fertile specimens now deposited in the Herbario Vargas in Cusco (CUZ), the Museum of
Natural History in Lima (USM), and The Field Museum (F). R. Foster and N. Salinas took approximately 2,500 photographic vouchers of plants.
C. Vriesendorp inventoried understory plants (1-10 cm dbh) in ten transects: three in Kapiromashi, four in Katarompanaki, and three in Tinkanari, for a total of 1,000 stems. Understory transects varied in area but were standardized by the number of stems, following the method of Foster et al. (http://www.fieldmuseum.org/rbi). All members of the botanical team catalogued plants of all life forms, from canopy emergents and shrubs to herbs and epiphytes. In addition to making general collections, N. Salinas (Orchidaceae) and H. Beltrán (Asteraceae and Gesneriaceae) focused on their families of expertise at each site.
FLORISTIC RICHNESS AND COMPOSITION
Conservative estimates of vascular plant diversity for the eastern Andean slopes of Peru range from 7,00010,000 species, suggesting that forests in these areas may contain half or more of the plant species in Peru (Young 1991). Based on our field observations and collections at the three inventory sites, we generated a preliminary species list of ~1,400 species for Zona Reservada Megantoni (Appendix 1). Using preliminary lists from similar elevations in the Cordillera Vilcabamba to the west (Alonso et al. 2001) and Parque
Nacional Manu to the east (Cano et al. 1995, Foster 1990), we estimate a total flora of 3,000-4,000 species for the 215,006 ha of Megantoni. This is necessarily a broad approximation as our quick survey covered only a subset (650-2,350 m) of the full elevational range (500-4,000+ m) within the Zona Reservada.
As in other forests on the slopes of the eastern Andes, floristic richness within Megantoni is extremely high. In ZRM, we documented an astonishing diversity of orchids and ferns, particularly at the two higherelevations sites of Katarompanaki and Tinkanari. These two plant groups dominated the flora and contained at least a quarter of the species we observed in the field (Pteridophyta, 190 species; Orchidaceae, ~210 species; Appendix 1). Ferns are commonly encountered in montane habitats; however, the diversity and abundance of ferns in Megantoni were particularly high. Of the 118 genera reported for Peru (Tryon and Stolze 1994), we found representatives of nearly half (~ 55) in Megantoni.
Of the 116 fertile orchid collections, we suspect that 20 represent species new to science (see Figure 6). The number of new orchid species still awaiting discovery may be even higher, given that the majority of the orchids we observed in the field were sterile or in fruit (and therefore effectively sterile for orchid taxonomists). Moreover, we were unable to take comprehensive samples of tree canopies where orchid abundance and diversity are usually highest. For that reason, the number of new orchid species we observed was even more remarkable.
Compared to the floras of other lower- and middle-elevation sites on the Andean slopes, certain families and genera were notably rich in species. We observed high numbers of Rubiaceae (92),
Melastomataceae (64), Asteraceae (53), Araceae (52), Fabaceae (sensu lato, 52), and Piperaceae (49) across all three sites. At the generic level, we encountered 33 species each of Psychotria (Rubiaceae) and Miconia
(Melastomataceae), 25 species of Peperomia (at least 10 species at each site), 24 species of Piper (Piperaceae), and at least 15 species each for Pleurothallis and Maxillaria (Orchidaceae, Figures 6C, 6E, 6F, 6G, 6I, 6S, 6T, 6X, 6Y, 6Z, 6AA, 6HH, 6JJ, 6KK, 6LL, 6NN).
We found high species richness in Anthurium (24) and Philodendron (18), both genera in the Araceae, a principally epiphytic and typically species-rich family at higher elevations. Species richness of Elaphoglossum ferns (more than 15, Figure 5H) was astonishing at Tinkanari (2,100-2,350 m); this campsite may be one of the global centers of diversity for the genus. At this same elevation, we recorded sympatric populations of at least 10 species of tree ferns (mostly in the genus Cyathea, Figures 5A, 5B, 5K), as well as 8 species of bamboo (Chusquea and close relatives).
We encountered fewer species and individuals of palms (Arecaceae; 23 species) than we expected, but lower-elevation sites in Megantoni probably support larger populations and more species. For Bromeliaceae, a principally epiphytic family, the area supports several abundant species, but with the exception of Guzmania (15 species) it does not seem especially species rich.
VEGETATION TYPES AND HABITAT DIVERSITY
In contrast to nearby Amazonian forests, where broad floristic similarity can been found over thousands of kilometers (Pitman et al. 2001), Andean forests are floristically heterogeneous at almost any spatial scale— from satellite images, to helicopter overflights, to short hikes on the ground. Even forests at similar elevations typically exhibit differences in composition and structure. Much of this heterogeneity derives from the rugged and varied topography, microclimatic changes along elevational gradients, disturbance from landslides, and dramatic, small-scale variation in substrate.
However, our understanding of how these factors interact to determine plant community composition remains limited.
Our inventory sites spanned 650-2,350 m in elevation. We were not able to sample sites at either the lowest (500-650 m) or the highest (2,350-4,000 m) elevations that make up some 20% of the Zona Reservada, but we believe that the sites we visited are representative of plant communities across a large proportion of the Zona Reservada.
Lower mountain slopes (Kapiromashi, 650-1,200 m, 26-30 April 2004)
Our first campsite was situated adjacent to the Río Ticumpinía. We explored the forests dominated by patches of bamboo on the steeply ascending slopes on either side of the river valley. Overflights of the region suggest that similar plant communities grow along the Río Timpía on the eastern side of the Zona Reservada. One of the largest rivers in the region, the Ticumpinía is fast-flowing and dynamic, changing course rapidly enough that our 2001 satellite image was already outdated. During our visit in the late rainy season, the river levels were unexpectedly low, exposing a broad floodplain (Figure 3C). We suspect that these forests receive 5-6 m of rain per year, with no significant dry periods. The high humidity, further exaggerated by the narrowness of the valley, may explain why we found several species typical of higher elevations at this site.
A key feature of vegetation here and elsewhere at lower elevations in the Zona Reservada is scattered stands of stout Guadua bamboo (Figure 3E). Although the factors influencing the distribution of bamboo patches across the landscape are poorly understood, these stands are a continuation of the Guadua patches that dominate vast stretches of southwestern Amazonia. All trails at this site crossed Guadua patches, ranging from isolated clumps of bamboo to tangles covering several kilometers. Within larger patches of bamboo, species richness of plants was markedly reduced, and in some places downright depauperate. Transect data reveal that the understory plant community growing in areas with bamboo is approximately half as rich as that growing nearby, outside Guadua-dominated forest (29 vs. 57 species).
Typically, bamboo stems were interspersed with a mixture of palms (Socratea exorrhiza, Iriartea deltoidea; Arecaceae) and secondary forest species (Cestrum sp., Solanaceae; Neea sp., Nyctaginaceae;
Triplaris sp., Polygonaceae; Perebea guianensis,
Moraceae; and spiny lianas of Uncaria tomentosa, Rubiaceae, the medicinal plant uña de gato, or cat’s claw). Thin-stemmed shrubs and suffrutescent herbs dominated the understory, including Begonia parviflora (Begoniaceae), a Sanchezia sp. (Acanthaceae) with bright red bracts, and Psychotria viridis (Rubiaceae), an ingredient of the hallucinogen ayahuasca. Less frequently, we encountered species more typical of mature forest, including Guarea spp. (Meliaceae) and at least three species of Lauraceae.
Farther upslope in this area (above 800 m asl), we explored areas without Guadua bamboo and encountered more exciting plant assemblages, with a higher diversity of trees and shrubs. The plant community here was a mix of species typical of higher elevations, species typical of the lowlands, and secondary-forest species colonizing local disturbances. Because of the high frequency of landslides and treefall gaps, we found few undisturbed sites and few true dominants in the plant community.
Within the more humid valleys, several species grow below their known altitudinal ranges. Below 1,000 m we encountered Bocconia frutescens (Papaveraceae), which grows elsewhere above 1,700 m, and Maxillaria alpestris (Orchidaceae), an orchid known from 1,800-2,700 m at Machu Picchu.
Canopy trees here were larger than those growing within the Guadua stands, and more likely to be covered in trunk climbers. Several large tree species (dbh > 30 cm) typical of lower-elevation sites occurred here, including natural rubber, Hevea guianensis (Euphorbiaceae); two important but infrequent tropical timber trees, Cedrela fissilis (Meliaceae) and
Cedrelinga cateniformis (Fabaceae s.l.); Poulsenia armata (Moraceae); Dussia sp. and Enterolobium sp. (Fabaceae s.l.); and several species of Ficus (Moraceae). We observed few palm species. At surprisingly low densities, we encountered Socratea exorrhiza, Iriartea deltoidea, Oenocarpus bataua, Wettinia maynensis, and a few species of Geonoma, but we did not observe any species of the Bactris or Euterpe palms that typically co-occur with these species in lowland sites.
Like those of the overstory, understory communities contained a mix of secondary-forest species and species more typical of mature forest. We encountered 57 species in a 100-stem understory transect, and the most “common” species, Henriettella sp. (Melastomataceae), made up only 6% of the stems. Other common understory species included Perebea guianensis (Moraceae, 5%), Miconia bubalina (Melastomataceae, 5%), and Tapirira guianensis
(Anacardiaceae, 4%). In the same transect, we registered 20 different families. Four families harbored the bulk of the species diversity: Lauraceae (7 species), Fabaceae sensu lato and Rubiaceae (6 species each), and Melastomataceae (5 species). In some areas, the shrubs
Psychotria caerulea and Psychotria ramiflora (Rubiaceae) were locally dominant. In species richness this non-Guadua forest seems similar to other areas of wet Andean foothills in southern Peru and Bolivia, higher than much of the central Amazon, but not as rich as the flora of northern Peru and Ecuador.
River floodplain and islands
An obvious successional flora grows along the edge of the Río Ticumpinía and on the river island near our campsite, similar to riverside plant communities throughout lowland southeastern Peru (e.g., Madre de Dios). Clumps of Tessaria integrifolia (Asteraceae), Gynerium sagittatum (Poaceae), and Calliandra angustifolia (Fabaceae s.l.) grew closest to the river, followed by an overstory of Ochroma pyramidale (Bombacaceae), Cecropia multiflora (Cecropiaceae), and Triplaris americana (Polygonaceae). Behind these taxa, or sometimes interspersed with them, we frequently encountered trees of Guettarda crispiflora (Rubiaceae) and Inga adenophylla (Fabaceae s.l.). At the center of the river island, a slightly depressed, wetter area supported an herb layer including species of Mikania
(Asteraceae), Costus (Costaceae), and Renealmia (Zingiberaceae).
In the beds of the larger streams thrives a low-diversity assemblage of colonizing species, including Tovaria pendula (Tovariaceae), three species of Urera (U. caracasana, U. baccifera, U. laciniata; Urticaceae),
Acalypha diversifolia (Euphorbiaceae), scrambling Phytolacca rivinoides (Phytolaccaceae) and Mikania micrantha (Asteraceae), a spiny Wercklea ferox shrub (Malvaceae), and dense patches of Banara guianensis treelets (Flacourtiaceae). In the second-growth overstory alongside the stream, we found several abundant species important for vertebrate frugivores. They included Inga adenophylla (Fabaceae), an Allophyllus sp. (Sapindaceae), four species of Piper (Piperaceae), and a large-leaved Guarea (Meliaceae).
Large Ladenbergia (Rubiaceae) trees— a species obvious from afar with its broadly ovate leaves and panicles of dried, dehisced capsules—and Triplaris (Polygonaceae) trees, protected by fierce Pseudomyrmex ants, dominate the less frequently disturbed forest along the streams. Below their canopy, we commonly encountered an understory flora of Sanchezia sp.
(Acanthaceae), Psychotria caerulea (Rubiaceae),
Macrocnemum roseum (Rubiaceae), and Hoffmannia spp. (Rubiaceae). Also, we observed extensive understory populations of Heliconia robusta (Heliconiaceae), a rarely collected species, known in Peru from only a handful of collections.
(Katarompanaki, 1,350-2,000 m, 2-7 May 2004) From the Río Ticumpinía valley we flew via helicopter to an isolated, two-tiered tableland near the center of ZRM. From the air we saw stunted vegetation on the top tier, and much taller, closed-canopy forest on the lower tier. On the ground we found that extremely hard rock underlay the vegetation growing on the top tier, and the stunted size of the free-standing plants likely reflects the limited nutrient availability and poor growing conditions of this substrate. Our camp was centered in the stunted vegetation on the top tier, and we spent most of our time sampling these plant communities. Our last two days were dedicated to exploring the lower tier.
Stunted forests on rock outcrops are common in Megantoni. Although these communities have a consistent appearance and forest structure when seen from the air, floristic composition appears to vary substantially from one to the other, with different species dominating each hard-rock surface. This variation may reflect biogeographic barriers to dispersal among sites with similar geology, random assembly of communities, different geochemistry, or even microendemism caused by recent speciation on isolated substrates.
A wall of clouds forms an almost permanent bank on the southern edge of the top tier of the platform, and both the density and the diversity of epiphytic climbers are higher at this site than at the first campsite. Whereas on the lower mountain slopes plant diversity is concentrated in trees and shrubs, at this higher site the bulk of the diversity shifts to epiphytic and herbaceous plants, especially on the top tier of the tablelands.
Orchids illustrate this shift dramatically.
The ~120 species of Orchidaceae recorded across both tiers account for nearly a quarter of the plant diversity at this site. Moreover, of the 66 species we found in bloom, at least 17 do not resemble any known species and are likely new to science (Figure 6).
Below we focus on the floristic differences and similarities between the two tiers, characterizing the vegetation on each, and comparing the vegetation on these two tiers of the middle-elevation tablelands to our inventory sites on the lower slopes (Kapiromashi) and somewhat higher middle-elevation slopes (Tinkanari).
Upper tier (1,760-2,000 m)
The top tier of the tableland resembles a tilted platform, which slopes upward toward the southeast and rises over 200 m in elevation from its lowest to its highest point. As the elevation on the upper platform increases, the plant community decreases in stature and in diversity. This change is perhaps best exemplified by the distribution of Dictyocaryum lamarckianum (Arecaceae) palms. At lower elevations on the upper tier, D. lamarckianum is one of the dominant trees. But as elevation increases, the population thins out and individuals are shorter. At upper elevations on the platform, the low-diversity assemblage does not include any D. lamarckianum palms.
Our 100-stem understory transect at lower elevations on the upper tier (~1,760 m) contained 28 species, compared to only 13 species at the upper end of the platform (~2,000 m). However, diversity in both transects was lower than would be expected for either of these elevations, presumably because the extremely hard (and likely acidic) rock substrates limit the number of species able to colonize this site. As a point of comparison, an understory transect at the higher Tinkanari campsite (~2,200 m) registered 32 species.
On the lower reaches of the platform, the canopy was ~15-20 m tall, and three tree species dominated the overstory and understory: Alzatea verticillata (Alzateaceae) and a large-leaved
Clusia sp. (Clusiaceae, Figure 3G) growing alongside D. lamarckianum palms. Filling in the gaps among individuals of these three species was a mix of shortstatured trees in the families Melastomataceae, Rubiaceae, and Euphorbiaceae (four species each); at least three species of tree ferns; and an occasional small palm (Euterpe precatoria, Geonoma spp.; Arecaceae).
On the upper reaches of the platform, where lightning strikes probably cull out tall trees, vegetation was much shorter: the canopy was ~2 m high, with a few “emergents” reaching 4 m. Three species were overwhelmingly abundant here, including Weinmannia sp. (Cunoniaceae), Cybianthus sp. (Myrsinaceae), and a small-leaved Clusia sp. A few species of Chusquea-like bamboos with thin, floppy stems occurred infrequently at this elevation, often supported by other stems. Silvery Ceroxylon parvifrons palms, a preferred food of spectacled bears (Tremarctos ornatus, Figure 11B), were scattered throughout. Many of these short palms had obvious chew marks on their stems. The uprooted stems of others were discarded, with the soft interior nearly entirely consumed.
Lower tier (1,350-1,600)
To gain access to the lower tier, we followed a trail 5 km northeast of camp, crossing half a dozen streams and ending in a spectacularly steep descent of nearly 250 m. On the lower tier, patches of closed-canopy forest grew interspersed with patches of secondary growth. Here the canopy was 30-40 m tall, with several emergent trees extending another 10 m above it.
Our only trail on the lower tier skirted the steep rock face below the upper tier, passing one large waterfall and crisscrossing a large stream. We examined the plant communities along both sides of the stream and studied the flora along the banks from within the stream itself, walking as far up- and downstream as possible.
Alongside the stream, we encountered a flora principally composed of species from lower elevations; every once in a while we were surprised to find a species typically occurring at much higher elevations. Many of the species had been recorded at the Kapiromashi campsite, nearly 500 m lower than this site, including species such as Guettarda crispiflora (Rubiaceae) and Banara guianensis (Flacourtiaceae). Within the canopies of streamside trees, we spotted the large red flowers of Mucuna rostrata (Fabaceae), a species known from lowland floodplains in Peru. This elevation may be the highest recorded for this species. Also growing alongside the stream were species more typical of higher elevations, such as an abundant flowering Turpinia tree (Staphyleaceae).
Fruiting plants were remarkably abundant on the forested terraces on either side of the stream, and species with large fruits important for frugivores were especially well represented (see Mammals). Walking around the 1.5-km loop, we encountered fruits of Caryocar amygdaliforme (Caryocaraceae), three species of Ficus (Moraceae), two species of Myrtaceae (probably Eugenia), Tabernaemontana sananho (Apocynaceae), two Psychotria spp. and one
Faramea sp. (Rubiaceae), at least four species of Melastomataceae, a softball-sized Cucurbitaceae, and an Anomospermum sp. (Menispermaceae).
On these forested terraces, we encountered several trees more than 1 m in diameter, including 2 legumes (Parkia sp., Dussia sp.; Fabaceae), 3 species of Ficus (Moraceae), and at least 1 species of Pouteria (Sapotaceae). For our two 100-stem understory transects on either side of the stream, we recorded ~70% species overlap and nearly equivalent species richness (50 and 46 species). One transect was dominated by Iriartea deltoidea (Arecaceae, 9%), a palm species that is often the most common tree in Amazonian tree plots (Pitman et al. 2001), and the other by a Croton sp. (Euphorbiaceae, 8%) with a single gland on the petiole. In addition to these 2 species, both transects contained nearly equivalent numbers of Protium (Burseraceae), Coussarea (Rubiaceae), Mollinedia (Monimiaceae), and Chrysochlamys (Clusiaceae). Several dominants in the understory at this site, including Urera baccifera (Urticaceae) and Pourouma guianensis (Cecropiaceae), were present in similar abundances in the Kapiromashi understory transects. More than half of the species belonged to five families: Lauraceae (8 species), Rubiaceae (7 species), Melastomataceae (6 species),
Myrtaceae (3 species), and Chloranthaceae (2 species).
Middle-elevation slopes (Tinkanari, 2,100-2,400 m, 9-14 May 2004)
From the middle-elevation tablelands, we flew across 41 km of ridges and valleys to reach the eastern edge of the Zona Reservada, adjacent to Parque Nacional Manu. The headwaters of the Río Timpía and the Río Manu originate on these slopes, and most of the streams contained fast-flowing, oxygenated water.
Slopes at this elevation are usually precipitous, and views across the valley revealed several sheer cliffs and sharp inclines on most of the facing slopes. However, this site was uncharacteristically flat. In one depressed area, a small, stagnant, blackwater pond had formed, and all of the forest we explored had only a gentle slope.
As at the middle-elevation tablelands, two main forest types occur at this site. A tall, closed-canopy forest is the principal vegetation in the area and surrounds small, isolated patches of stunted shrub forest growing on shallow soils over hard rock near the edge of the escarpment that drops to the valley below. These two forest types share only 10% of their species, despite abutting each other.
Tall forest dominates the vegetation at this site. The canopy ranges from 30 to 40 m tall, with some emergent trees surpassing 50 m. The tree community is not diverse and is dominated by a few species. Along most trails, Calatola costaricensis (Icacinaceae) accounts for a quarter of the trees in the subcanopy and its large, hard seeds litter the ground. Growing in the understory alongside Calatola was a mix of two species of Hedyosmum (Chloranthaceae) and tree ferns or bamboo
(Chusquea). The high canopy is dominated by trees of Hyeronima sp. (Euphorbiaceae), Heliocarpus cf. americanus (Tiliaceae), Weinmannia sp. (Cunoniaceae), Elaeagia sp. (Rubiaceae), Ficus spp. (Moraceae), and many huge, broad-crowned Sapium (Euphorbiaceae) of a species none of us had seen before (and which we were unable to collect). We found few large (> 80 cm diameter) individuals of Podocarpus oleifolius (Podocarpaceae), Juglans neotropica (Juglandaceae), and
Cedrela montana (Meliaceae). Alnus acuminata (Betulaceae) and Morus insignis (Moraceae), genera typical of northern temperate forests, are frequent colonizers of landslide disturbances.
The dominant shrubs are Mollinedia sp. (Monimiaceae) and an Oreopanax sp. (Araliaceae), while Pilea spp. (Urticaceae) are the most conspicuous terrestrial herbs. We found abundant root parasites
Corynaea crassa (Balanophoraceae) growing on Hedyosmum roots, but not exclusively.
Ferns are an important and conspicuous element of these forests (Figure 5). In a 5 x 25 m transect we counted 30 species of ferns and their allies (Pteridophyta). Pteridophyta also dominated the epiphyte community, and trees supported an average of 10 epiphytic fern species per trunk. Tree ferns (mostly Cyathea; Figures 5A, 5B, 5K) are superabundant and diverse in the understory at this site. Extrapolating from a 150 x 1 m transect, tree fern densities at this site could reach 2,000 individuals per ha. We commonly encountered 5 to 6 species of tree ferns; 3 or 4 others specialize on particular habitats and occur infrequently.
Tree ferns were most common in intact forest with a high canopy, even if the understory received substantial amounts of light. In contrast, the common large Chusquea bamboo (with stems ~10 cm diameter) formed extensive solid stands principally in areas with few high canopy trees. Tree fern and bamboo populations rarely co-occur, suggesting that bamboo may invade areas where the canopy has been disturbed (e.g., after a violent windstorm), but as trees recover and begin to shade the bamboo, tree ferns can gradually recolonize the area and ultimately replace the bamboo.
Stunted shrub forest
Shrub forest covers a 0.5-km2 area on the exposed southwestern face of these middle-elevation slopes and is distinctly visible from the air. On the upslope portions of the shrub forest, the tallest stems nearly reach 6 m. The plant community decreases in stature and changes in composition as the slope descends. On the upslope portions, the forest appears orange from afar, thanks to orange-leaved species in the genera Graffenrieda (Melastomataceae), Clethra (Clethraceae), Clusia,
Weinmannia (Cunoniaceae), Styrax (Styracaceae), and
Cybianthus (Myrsinaceae). Several small species of Chusquea bamboo grow haphazardly on this upper portion. Although many genera are shared with the stunted forest growing on the Katarompanaki tablelands, most species are distinct.
Further downslope, the Graffenrieda is still present, albeit shorter in stature, but most of the other upslope dominants disappear. The plant community here is much shorter, averaging 1.5 m. Several terrestrial orchids are common this area, including Gomphichis plantaginifolia and Erythrodes sp., mixed with Blechnum ferns and a small-leaved clonal Clusia that dominates the vegetation, along with three species of less common Ilex (Aquifoliaceae) and a stiff-leaved Miconia (Melastomataceae), among others. The small wax palm present at Katarompanaki, Ceroxylon parvifrons, occurs here as well, where spectacled bears also consume it.
ORCHIDS (Norma Salinas)
The Orchidaceae is one of the most diverse flowering plant families in the world, with 25,000 to 35,000 species. Individuals vary broadly in size, ranging from almost tiny epiphytes to shrubs.
The eastern slopes of the Andes—from Colombia to Bolivia—support a high diversity of orchids, with many endemic species. During the last 30 years in Peru, few plant collectors have focused on orchids, and there are probably many orchids species still awaiting discovery. In a rapid inventory of Cordillera del Cóndor, 26 of the 40 orchid species were new to science (Foster and Beltrán 1997). More recent studies have resulted in hundreds of new records for Peru, and have shifted the known centers of diversity for many genera, including Lycaste, Kefersteinia and Stenia, from Ecuador and Colombia to Peru.
Not surprisingly, we encountered a rich orchid community in Megantoni during our inventory, in almost every sampled habitat (Figure 6). In a little over two weeks, we found 116 species of fertile orchids, and ~80 sterile species. Our estimates for sterile species are conservative, as many species of the subtribe Pleurothallidinae are easily confused without flowers. Nor do these estimates include the subtribe Oncidiinae, a family that flowers during a different season, as do several other subtribes.
We suspect 20 of the 116 fertile species may be new to science. Additionally, various species are new records for Peru, including Elleanthus hirtzii, previously known only from Ecuador. The Zona Reservada is pristine, and we found healthy and large orchid populations. Of the flowering species, 90% were epiphytic, and 10% terrestrial. A few species were lithophytes, growing on rocks or cliff faces.
Species in the genera Maxillaria, Epidendrum, Lepanthes, Platystele, Pleurothallis and Stelis represented the majority of the fertile orchids observed during the inventory. We registered several species in rare genera, including Baskervilla, a genus with fewer than ten species, and distributed from Nicaragua to Peru and Brazil. Additionally, we found a species of Brachionidium, a genus that is poorly represented in
Peru even though it ranges from Costa Rica to Bolivia.
All sites visited during the inventory (from 760 to 2,350 m asl) displayed high orchid species richness. Of all the flowering plants at each site, in Kapiromashi (~760-1,200m) 7% were orchids, in Katarompanaki (~1,300-2,000 m) 24% were orchids, and in Tinkanari (~2,100-2,350 m) 11% were orchids.
In several genera we observed hints of incipient speciation. For example, we found two species of Sobralia that closely resembled S. virginalis and S. dichotoma. However, upon closer inspection, morphological differences on the lip of both species are large enough to suggest that the specimens from Megantoni are either in the later stages of speciation, or already distinct species. Similarly, we observed variability in form and color of many species of Maxillaria, along with high species richness in this genus.
Of the fertile orchid species, few are shared with other orchid-rich areas in Peru (e.g., Machu Picchu, Manu, Vilcabamba). A few species are restricted to small areas, or threatened in other areas of Peru, but present large healthy populations in the ZRM. For example, both Masdevallia picturata (Figures 6A, 6H) and Maxillaria striata (Figure 6TT) are considered threatened in Parque Nacional Manu, yet appear abundant in Megantoni.
Also, an Otoglossum sp. found abundantly at the
Tinkanari campsite was found only occasionally in PN Manu at 2,500-2,600 m asl. A species recently described from Machu Picchu, Prosthechea farfanii, also has large populations in Megantoni. These data suggest that other sites in ZRM may harbor populations of orchids that are suffering declines in other areas of the Andes, including perhaps Masdevallia davisii, a species with critically low numbers of individuals.
NEW SPECIES, RARITIES, AND RANGE EXTENSIONS
Although most of the plant species we collected during the inventory are still unidentified, some already have been confirmed as new species, or substantial range extensions for described species. As more species are identified, or additional new species are confirmed, we will update our plant list at http://www.fieldmuseum.org/rbi/. We include collection numbers for each potential new species or range extension, as a reference to collections housed at the Vargas Herbarium in Cusco (NS, Norma
Salinas) or the Museum of Natural History in Lima (HB, Hamilton Beltrán).
The bulk of potentially new species are orchids; most come from the higher-elevation campsites. Preliminary revisions in the Vargas Herbarium in Cusco of collections from Peru, Bolivia, and Ecuador suggest that perhaps 20 of the 116 fertile orchid collections may be new to science (see Orchids, Figure 6), a remarkable number for a three-week inventory. Based on digital photographs we took in the field, specialists have tentatively identified 9 additional plant species as new to science. All are from our two higher-elevation campsites.
On the upper tier of the middle-elevation tablelands at Katarompanaki campsite (1,300-2,000 m), we encountered potential new species in the following genera: Psammisia (Ericaceae, NS6931; Figure 4A), Schwartzia (Marcgraviaceae, NS6880; Figure 4F),
Trichilia (Meliaceae, NS6788), and Macrocarpaea (Gentianaceae, NS6869). At Tinkanari, our highestelevation site on the middle-elevation slopes, we found several potential new species, including an Acanthaceae with lilac-colored flowers (NS7198), a Sphaeradenia (Cyclanthaceae, NS7184), a Gesneriaceae with a big, pedunculate fruit (HB5950, Figure 4C), a Hilleria cf. sp. with bright orange flowers (Phytolaccaceae, NS7237;
Figure 4E), and a Tropaeolum sp. (Tropaeolaceae, NS7235).
Several collections in Megantoni extend the known ranges of species hundreds of kilometers farther south. One is from our low-elevation campsite, Kapiromashi, where we registered Wercklea ferox
(Malvaceae, NS6735) for the first time in southern Peru.
At Katarompanaki, we found Ceroxylon parvifrons
(Arecaceae, NS7037), Tapeinostemon zamoranum
(Gentianaceae, NS6857; Figure 4B), Sarcopera anomala
(Marcgraviaceae, NS6881) and Macleania floribunda
(Ericaceae, NS6939). At Tinkanari, we encountered
Miconia condylata (Melastomataceae, NS7211) and
Peltastes peruvianus (Apocynaceae, NS7273; Figure
4D), both previously known only from northern Peru.
Our collection of Heliconia robusta (Heliconiaceae, NS6600) fills a large gap in its distribution. This Heliconia, with triangular green and red bracts and yellow flowers, dominated parts of the naturally disturbed forest around our low-elevation campsite, Kapiromashi. Known mostly from Bolivia, it has been collected only a handful of times in Peru and was overlooked in the Catalogue of the Flowering Plants and Gymnosperms of Peru (Brako and Zarucchi 1993).
Two species encountered at the higher-elevation sites are first collections for Peruvian forests. Although seen and reported in Huanuco and Puno, our Tinkanari collection of Spirotheca rosea (Bombacaceae, NS7128; Figure 4G) is the first specimen for any Peruvian herbarium. Another first specimen for Peru, Guzmania globosa (Bromeliaceae, NS6808; Figure 4H) grew in small patches on the upper tier of the Katarompanaki tablelands. This species was previously known from Ecuador and photographed, but not collected, in the rapid biological inventory of Cordillera Azul (Alverson et al. 2001).
THREATS, OPPORTUNITIES, AND RECOMMENDATIONS
Zona Reservada Megantoni connects two important conservation areas: Parque Nacional Manu and the Vilcabamba conservation complex (Parque Nacional
Otishi and Reservas Comunales Ashaninka and Machiguenga, see Figure 1). We recommend the highest level of protection for the valleys, slopes, mesas, ridges, and high-altitude grasslands that span the elevational gradient of more than 3,500 m within ZRM. Intact elevational transects are rare in the tropical Andes, and protecting ZRM is urgent. Natural gas is being extracted to the north, and colonization threatens from the south. If ZRM is not protected, a rare opportunity to link two large protected areas and to protect more than 2.6 million ha will be lost.
Judging from our observations from the overflight, the inventory, and satellite images, we recognize several particularly well-preserved and unique habitats within ZRM. Wet high-altitude grasslands (puna) experience intense land use, overgrazing, and overburning in other areas of Peru. Compared to Parque Nacional Manu and other areas of the eastern Andean slopes, from the air Megantoni appears to contain possibly the least disturbed extensions of high-altitude grassland in Peru. Protecting ZRM provides an opportunity to preserve the full richness of this intact mountain flora and could provide a living reference for restoration efforts in degraded grasslands nearby.
The expansive mid-elevation tablelands, including Katarompanaki camp where we found more than 15 orchid species new to science, are a geological formation that appears to occur only in Megantoni, and not in the neighboring Cordillera Vilcabamba conservation complex or Parque Nacional Manu. Protecting Megantoni will safeguard these unique landscape features and will protect a site important for orchid populations. Orchids are protected under CITES regulations in Peru, and formal protection of Megantoni will impede unauthorized orchid collecting.
The importance of Megantoni as a conservation area does not rest solely on its role as a pristine biological corridor, but also reflects the endemic species that occur within its boundaries. We estimate a flora of ~3,000-4,500 species for ZRM. We know some of these plant species are shared with neighboring Manu and Vilcabamba. However, our knowledge of plant communities at all three of these sites is too limited to calculate exact numbers of species unique to each area. As a preliminary indication, the 25-35 species potentially new to science imply high levels of plant endemism within ZRM (see Figures 4, 6). These potential new species, discovered during 15 days of plant surveys, suggest that 1-2% of all the plant species projected to occur in Zona Reservada Megantoni are not currently known from adjacent protected areas or any other site in the world. Additional surveys may uncover some of these undescribed species in neighboring Manu or Vilcabamba. However, given the number of floristic novelties found during the rapid inventory, future inventories in Megantoni are likely to uncover additional endemic species in ZRM.
DUNG BEETLES (Coleoptera: Scarabaeidae: Scarabaeinae)
Participant/Author: Trond Larsen
Conservation targets: Large dung beetle species, susceptible to local extinctions and functionally important for dispersing seeds, controlling mammalian parasites, and recycling nutrients (especially Deltochilum, Dichotomius, Coprophanaeus, Phanaeus, and Oxysternon); several rare and restricted-range species (including at least ten species new to science); healthy populations of medium-sized and large mammals, especially monkeys, that provide essential dung resources; intact habitats that support distinct dung beetle communities sensitive to habitat degradation
Dung beetles (subfamily Scarabaeinae) are diverse and abundant, and their diversity often mirrors broader patterns within the community (Spector and Forsyth 1998). Since they depend on mammal dung for food and reproduction, dung beetle populations often reflect mammal biomass, and by extension, hunting intensity. Moreover, dung beetles show high betadiversity across habitat types and are sensitive to many kinds of disturbance, including logging, hunting, and most types of habitat degradation (Hanski 1989; Halffter et al. 1992). Dung beetles also play an important role in ecosystem functioning. By burying vertebrate dung, beetles recycle plant nutrients, disperse seeds, and reduce infestation of mammals by parasites (Mittal 1993; Andresen 1999).
To my knowledge, no one has published a study of dung beetle communities in the Peruvian Andes. Between 1998 and 2003, I sampled dung beetles at several sites in southeastern Peru, both in Amazonian and Andean forests on the eastern side of Zona Reservada Megantoni. The dung beetle diversity in several of the lowland sites (the Río Palma Real area, Los Amigos Biological Station, and Cocha Cashu Biological Station) is among the highest known in the world, with over 100 dung beetle species at a single site. In the Kosñipata valley, adjacent to ZRM, I found that dung beetle diversity decreases with increasing elevation. Many of these dung beetle species show restricted ranges and many remain undescribed.
To sample dung beetle communities, I used a combination of baited pitfall traps and unbaited flight intercept traps. Each pitfall trap consisted of two stacked 16-oz (473-ml) plastic cups buried in the ground with the top rim flush against the soil surface. I filled the top cup halfway with water and a small amount of detergent to reduce surface tension. For each dung-baited trap, I wrapped ~20 g of human dung in nylon tulle and suspended the bait above the cups by tying it to a short stick pushed into the ground. Traps were standardized with human dung because it was readily available and is among the most attractive types of dung to most species of dung beetles (Howden and Nealis 1975). To prevent beetles from landing on the bait and to protect the trap from sun and rain, I covered the bait and the cups with a large leaf. I collected the samples every 24 hours, usually for a period of four days, although a few traps were set for only two days. This trapping method and trapping period usually provides relatively complete and quantitative descriptions of the diversity, composition, and relative abundances of the beetle community.
Within each of the four sites (Kapiromashi, Katarompanaki upper and lower platform, Tinkanari), pitfall traps were placed along as many trails and habitats as possible, and spaced at least 50 m apart. I installed at least ten traps in primary forest at each site, and as many traps as possible in additional habitats. I replaced dung baits every two days.
Since many generalist and specialist species of dung beetles use other food resources, I also set pitfall traps baited with rotting fruit (primarily banana), rotting fungus, dead fish, and dead insects. I placed up to three traps with each of these bait types in each of the four sites, spacing them at least 50 m apart.
To sample dung beetle species not attracted to any of these bait types, I set flight intercept traps to catch beetles passively without any bait by stretching a rectangular sheet of dark green nylon mesh (1.5 x 1 m) between two sticks, and placing trays of soapy water beneath the mesh, to catch beetles flying into the mesh (Figure 7C). One or two flight intercept traps were placed at each site.
I identified and counted beetles the same day they were collected, preserved voucher specimens in alcohol, and deposited these specimens in the Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos in Lima, Peru, and at Princeton University in Princeton, New Jersey, USA. Additional specimens will eventually be deposited in the U.S. National
Museum of Natural History of the Smithsonian Institution in Washington, D.C.
I recorded 71 species and 3,623 individuals of dung beetles during 15 days of sampling in Zona Reservada Megantoni. Judging from my collections from Madre de Dios, I estimate that ~10-35 of the dung beetle species are new to science. Using EstimateS (Colwell 1997), a software program that predicts species diversity based on sampling effort, I evaluated the efficacy of my sampling during the inventory. Although additional sampling would register more species, in two weeks I managed to sample the majority of dung beetle species at the four inventory sites. Extrapolating from my dung beetle research in Manu and other Peruvian sites, I estimate that additional sampling would register ~120 species of dung beetles for the entire Zona Reservada.
Figure 14. Dung beetle species abundance distribution for all sites in the Zona Reservada Megantoni. Rarer species shown at left.
Dung beetle communities in Megantoni contained an unusually high number of rare species (Appendix 2, Fig. 14). Twelve species were trapped only once, and an additional five species were trapped twice, suggesting that these species are naturally rare or near the edge of their distributional limits. Several species, such as Coprophanaeus larseni and a new species of Eurysternus, appear to be genuinely rare throughout their range. As a point of comparison, the most common species, Ontherus howdeni, was represented by 446 individuals.
At this lowest site, I sampled dung beetles in primary forest, secondary forest, Guadua bamboo, and a wide, dry gravel riverbed. Of the 41 species I found at this site, 39 were encountered in primary forest, 23 in bamboo, and 20 in secondary forest. Only 4 species were trapped in the riverbed. Beetle abundance was highest in primary forest, followed in descending order by secondary forest, bamboo, and riverbed. I captured 16 species in only one habitat type. Two species of Canthidium were found only in flight intercept traps and may specialize on an unusual food or microhabitat. I trapped one individual of Coprophanaeus larseni in a carrion-baited pitfall trap in upper primary forest.
This species appears to be very rare and was recently described on the basis of just three specimens.
Katarompanaki lower platform
This lower platform contained mostly tall, mature forest very distinct from the vegetation of the upper platform. I sampled only in primary forest at this site and found 30 species. Beetle abundance here was just slightly lower than in the primary forest of Kapiromashi. I captured 3 species (2 Canthidium spp., 1 Ateuchus sp.) only in flight intercept traps, and these species may specialize on unknown resources. I captured one individual of a species of Bdelyrus in a fruit-baited trap. This dung beetle genus is poorly represented in museum collections, probably because of its unusual diet. Some Bdelyrus species may specialize on the detritus that collects in bromeliads or in tangled lianas, and other species have been attracted to rotting fungi.
Katarompanaki upper platform The upper platform was characterized by unusual, stunted vegetation with low tree diversity growing on hard acidic rock with little or no soil and a thick humus layer. This site had only ten dung beetle species and low beetle abundance. Two of the species (Deltochilum sp. nov. aff. barbipes and Uroxys sp. 6) were not found in the lower platform and appear to be new to science.
At the highest site, I sampled beetles in tall primary forest (~15-25 m tall), intermediate primary forest (~5-15 m tall), short forest/open shrub (~0-5 m tall), Chusquea bamboo, and young regrowth forest along a landslide. Thirteen of the 14 species found at this site were in tall primary forest, 8 in secondary forest, 5 in intermediate-height primary forest, 5 in bamboo, and 3 in short forest/open shrub. Abundance was highest in tall primary forest, followed by bamboo, secondary, intermediate, and short forest. I found four species in only one habitat type: 3 in tall primary forest and 1 in short forest/open scrub. Two of these were species of Canthon, a genus rarely found in this elevational range, and both species appear to be new to science. This site contained a higher proportion of nocturnal species (64%) than the lower sites. Although I sampled several species at carrion and in flight intercept traps, these same species were also attracted to dung. I did not collect any dung beetle species at fruit or fungus traps at this site.
Community patterns across habitats and sites Across the four sites, species richness and abundance decreased with increasing elevation, with the exception of the upper platform at Katarompanaki, which exhibited lower species richness and abundance than Tinkanari (Table 3). This pattern likely reflects the distinct, stunted vegetation and low mammal biomass on the upper platform of the Katarompanaki tablelands. Within each site, species richness and abundance varied greatly among habitat types (Table 4), with several general trends. Tall primary forest (~15-25 m tall) always contained the highest diversity and abundance of dung beetles, followed by intermediate-height forest (~5-15 m tall), secondary forest, and bamboo, and finally by scrub and open habitats (~0-5 m tall).
Species composition varied greatly among sites, and across habitats within sites (Table 4). Most species (80%) showed restricted elevational ranges of 300 m or less. The sites closest in elevation shared the most species; the most widely separated sites shared only one species. Similarity indices (Sorenson abundance and MorisitaHorn) among all sites were very low. When species did occur at more than one site, they were typically abundant at one site yet represented by one or a few individuals at another, suggesting that they may have been collected near the limits of their range at one of the sites. Within sites, habitats most similar in forest height, forest structure, and soil seemed to have the most similar species composition of dung beetles. Although larger species of dung beetles are often less abundant than smaller species, Zona Reservada Megantoni contains uncommonly high abundances of large species such as Dichotomius planicollis, D. diabolicus, D. prietoi, Phanaeus meleagris, P. cambeforti, and Oxysternon conspicillatum.
The areas we visited were almost completely pristine. Human disturbances within the reserve could harm dung beetle populations. At the lowest-elevation site, Kapiromashi, we found evidence of past hunting activity (old hunting trails, planted bananas) and observed fewer mammals, particularly monkeys, than expected. Although dung beetles were most abundant at this site, abundance standardized by sampling effort (21.9 individuals/trap) was lower than I expected at 850 m asl and was only slightly higher than the beetle abundance at the Katarompanaki lower platform (19.3 individuals/trap). Natural disturbance regimes
also affected dung beetles. In Kapiromashi and Tinkanari I found much lower beetle diversity and abundance in secondary forest than in primary forest (Table 4). Areas colonized by bamboo had much lower beetle diversity and abundance than primary forest.
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