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Reintroducing apex
rsos.royalsocietypublishing.org
predators: the perils of
muddling guilds and
Comment
taxocenoses
Cite this article: Miranda EBP. 2018 Everton B. P. Miranda1,2
Reintroducing apex predators: the perils of 1 ONF Brasil Gestão Florestal, Cotriguaçu, MT, Brazil
muddling guilds and taxocenoses. R. Soc. open 2 Universidade do Estado de Mato Grosso, Alta Floresta, MT, Brazil
sci. 5: 180567.
http://dx.doi.org/10.1098/rsos.180567 EBPM, 0000-0003-2198-4742
Wolf & Ripple [1] make the case for ‘large carnivore’ conservation
through reintroductions. The selected species, however, were
neither necessarily apex predators nor carnivores, but a subset
Received: 14 April 2018 of the mammalian order Carnivora, including those ranging
Accepted: 7 June 2018 from insectivores–omnivores (sloth bear, Melursus ursinus [2,3];
sun bear, Helarctos malayanus [4]) to herbivores–omnivores
(Andean bear, Tremarctos ornatus [3,5]; American black bear, Ursus
americanus [6]), full omnivores (Asiatic black bear, Ursus thibetanus
[7,8]; brown bear, Ursus arctos [9,10]) and several hypercarnivores
Subject Category:
(jaguar, Panthera onca [11]; dhole, Cuon alpinus [12]; cheetah,
Biology (whole organism) Acinonyx jubatus [13]). At first, Wolf & Ripple lump together
mammalian carnivores (those in the order Carnivora, which
Subject Areas: may or may not be regular predators), apex predators (which
ecology occupy the top of food chains [14]) and any carnivore (which
primarily consumes other vertebrates). Further, they mixed guilds
(a group of species that exploit similar resources) with taxocenosis
(a group of sympatric species sharing a common phylogenetic
clade). Consequently, the authors provide a biased assessment
Author for correspondence: of reintroduction priorities based on conservation imperatives
Everton B. P. Miranda that are not restricted to any of these groups, but primarily
e-mail: mirandaebp@gmail.com associated with apex predators. This commentary explores the
development of this muddle, which I interpret to be mainly due
to a combination of inaccurate use of nomenclature and concepts.
Furthermore, the authors make some mistakes for a number of
proposed reintroduction sites that proper investigation would
have revealed to be inappropriate.
Mammalian carnivores are not alone as apex predators, as the
latter comprise a diverse set of terrestrial and aquatic vertebrates
on Earth [14], which include (but are not limited to) large raptors
and large, predatory reptiles. By equating apex predators with
Carnivora mammals, the authors ignore the diverse predator
species that have the same potential to: (i) perform predator-
related ecosystem functions [15,16]; (ii) create wildlife-viewing
tourism opportunities [15,17]; and (iii) reduce species extinction
risk by re-establishing extirpated populations and reversing range
The accompanying comment can be viewed at shrinkage. Furthermore, this diverse array of apex predators is
http://dx.doi.org/10.1098/rsos.172235.

2018 The Authors. Published by the Royal Society under the terms of the Creative Commons
Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted
use, provided the original author and source are credited.
 Downloaded from http://rsos.royalsocietypublishing.org/ on July 12, 2018

threatened by exactly the same extinction drivers as ‘large carnivores’, namely livestock predation
2
[16,18], perceived risk to humans [19,20], overhunting [21,22] and habitat loss. In temperate and boreal
regions, extant, large raptors may now perform reduced functions as top predators [23], while the role

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of large reptiles may be non-existent [24]. These latitudes contain most of the developed countries, and
therefore most of the ongoing scientific research, which helps explain the authors’ biased world view of
which species qualify as apex predators in ecosystems outside the Northern Hemisphere. Apex predators
outside the Northern Hemisphere are, however, characterized by high diversity.
Reptiles such as crocodilians and giant snakes are important predators of the land–water interface
on tropical and subtropical realms. Crocodilians—the largest living continental predators on Earth—can
control access of terrestrial vertebrates to vital resources such as water [25] and can inflict havoc and
significant mortality on terrestrial mammals, with concomitant effects that cascade down into entire
aquatic food webs [26]. Pythons and anacondas, on the other hand, prey on terrestrial vertebrates,
thus exerting top-down control over their populations [27,28], as do other apex predators [29], and
some mammalian Carnivora [30]. Given their lower metabolic rates, reptiles feed much less frequently
than do hypercarnivorous mammals [31], but this is compensated for by the extremely high biomass
of heterothermic carnivores [22,32]. Physical power is clearly not a problem, because pythons and
anacondas are capable of killing and eating ‘large carnivores’ such as bears and large cats [33,34]. Like
many mammalian carnivores, large, predatory reptiles also kill domestic livestock and are vulnerable to
the same sort of retaliatory persecution that affects ‘large carnivores’ [35,36]. Giant snakes regularly kill
domestic dogs in urban, peri-urban and rural landscapes in the tropics [19,37,38]—as do the dog-eating
leopards Panthera pardus of India [39]. The well-known preference that pythons show for eating canids
must be taken into account when planning a possible reintroduction of red wolves (Canis rufus) into
the Everglades National Park; else we risk losing them to Python molurus predation. Predatory reptiles
are indeed conspicuous elements of apex predator guilds throughout many regions, but frequently are
overlooked as the top predator that they are.
Raptors are the Earth’s largest aerial predators in tropical, subtropical and temperate landscapes.
As do large, predatory reptiles, giant raptors occasionally even snatch offspring of ‘large carnivores’
[40,41]. Extant, large-bodied raptors such as harpy eagles (Harpia harpyja), crowned eagles (Stephanoaetus
coronatus) and Philippine eagles (Pithecophaga jefferyi) perform the role of apex predators in the tropical
forest canopy [42–44], a habitat that is largely or completely inaccessible to large, terrestrial carnivores.
Giant raptors are therefore apex predators in the world’s most biodiverse terrestrial ecosystems [45,46].
In tropical forests and other biomes, giant raptors exert the same top-down roles [47–49] of the more
widely celebrated ‘large carnivores’. As for wolves, research has documented the cascading effects on
diverse trophic levels when apex aerial predators are missing. Clearly, the largest raptors play important
roles in maintaining balance in the vertebrate communities of forest canopies [49–51]. Trophic cascade
ecology is a research topic that has conspicuously focused on the Carnivora mammal taxocenosis. By
contrast, giant raptors have been largely ignored by Wolf & Ripple and by other research reviews on
apex predators.
While proposing species reintroductions, Wolf & Ripple provided a superficial examination of the
occurrence, habitat suitability and original distributions of ‘large carnivores’. For the sake of brevity, I
restrict my comments to their proposed reintroductions in the Neotropics. For example, the first of the
six sites suggested for jaguar reintroduction, La Payunia, is a barren scrubland that provides inadequate
habitat for jaguars, while the other five sites already harbour jaguar populations, thus obviating the
need for reintroduction there. Four of the last five sites, namely Campos Amazônicos [52], Guaporé
[53], Pacaás Novos [54] and Mapinguari [55], have published reports already confirming the presence of
jaguars, while the final site, Rio Novo, obviously must have jaguars as that protected area is embedded
within a vast region of continuous Amazon forest with a low human footprint. Cordillera Azul National
Park, where Wolf & Ripple propose the reintroduction of pumas (Puma concolor), already has a population
of pumas [56]. Similarly, I found weaknesses in the recommended reintroduction sites for Andean bears.
Running down that list of six sites, one finds (i) La Tatacoa and Cerro Saroche are arid, rocky landscapes
that, while they might support some bears as landscapes (as do Chaparri), are far from good habitat for
the species, and hence should not be prioritized; (ii) Serrania de Minas has an Andean bear population
discovered recently by local, government-supported researchers [57], and researchers found bears in
Pampa Hermosa in Peru [58]; (iii) Tinigua is at approximately 300 m.a.s.l. and offers only marginal habitat
for Andean bears, which tend to do best in cloud forests at 2500–3000 m.a.s.l. [59]; and (iv) the occurrence
of Andean bears in Panama is at best speculative [60], with the proposed site (Dárien) lacking sufficient
elevation [61] and perhaps receiving wandering bears into Panama from Los Katios in Colombia (Bernard
Peyton, personal communication).
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Therefore, all the reintroductions recommended for the Neotropics are either inappropriate or
3
mistaken. Global-scale studies commonly show weakness when one examines specific cases one by one,
but then one wonders how valid conclusions can be drawn from what appear to be such universally

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incorrect locations in the Neotropics.
When they state that ‘reintroduction efforts would lead to complete large carnivore guilds’, Wolf &
Ripple ignore many of the Earth’s most quintessential predators. Taxonomic chauvinism has long led
to a biased body of research that does not lead to a realistic model of the frequency and magnitude of
the ecological roles of different species. Their paper is another example of the lack of balance in global
analyses of predator ecology. Attention to predators that in fact have more varied feeding habits such
as omnivory as well as the need to include non-mammalian predators and the need for reliable global
databases on species presence are all essential to the study of ecology. Ignoring these factors results in
biased conclusions that are of questionable value in constructing and carrying out conservation policy in
this age of extremely scarce funding for conservation.
Data accessibility. This article has no additional data.
Competing interests. I have no competing interests.
Funding. I greatly appreciate the generous financial support of the following donors: Rufford Small Grants Foundation
(18743-1 and 23022-2), Rainforest Biodiversity Group, Idea Wild, The Mamont Scholars Program of the Explorer’s
Club Exploration Fund, Cleveland Metroparks Zoo and the SouthWild.com Conservation Travel System.
Acknowledgements. Luísa Genes started the discussion which resulted in this Comment, for which I am grateful. This
work received major logistical support from the Peugeot-ONF Brasil Carbon Sink Reforestation Project, based at
Fazenda São Nicolau in the Municipality of Cotriguaçu, Mato Grosso, Brazil. This ambitious project is an initiative of
Peugeot to fulfil directives of the Kyoto Protocol.

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