Ge Nature
YW) Conservation

Nature Conservation 56: 119-126 (2024)
DOI: 10.3897/natureconservation.56.132036

Lost species, neglected taxonomy, and the role of natural history
collections and synonymization in the identification of the World's
forgotten biodiversity

Spartaco Gippoliti'®, Simone Farina2®, Franco Andreone?®

1 IUCN/SSC Primate Specialist Group, I-00198 Roma, Italy

2 Museo di Storia Naturale dell’Universita di Pisa, I-56011 Calci, Italy

3 Museo Regionale di Scienze Naturali, I-10123 Torino, Italy

Corresponding author: Franco Andreone (franco.andreone@regione.piemonte.it; franco.andreone@gmail.com)

OPEN Qaccess

Academic editor: Klaus Henle
Received: 13 July 2024
Accepted: 7 September 2024
Published: 2 October 2024

ZooBank: https://zoobank.
org/4DB6682A-635D-49C0-BF9C-
80E9B13E07BC

Citation: Gippoliti S, Farina S, Andreone
F (2024) Lost species, neglected
taxonomy, and the role of natural history
collections and synonymization in the
identification of the World's forgotten
biodiversity. Nature Conservation 56:
119-126. https://doi.org/10.3897/
natureconservation.96.132036

Copyright: © Spartaco Gippoliti et al.

This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).

Abstract

A recent paper highlighted the need for increased attention towards species that have
vanished from scientific literature and proposed a new nomenclature in view of needed
conservation actions, based on the time interval since their disappearance. Here, we
propose to implement this concept by showcasing a taxonomic perspective that is often
overlooked in conservation studies. In recent decades, research on vertebrates showed
that, in several cases, species considered as undescribed were, in fact, first discovered
and already named a long time ago. Quite often they fell into oblivion while being con-
sidered synonyms of other taxa until they were re-evaluated in subsequent taxonomic
revisions. In this context, we are proposing the concept of “long-lost synonymized” spe-
cies, asking for greater attention to the discipline of taxonomy, the relevance of spec-
imen-based taxonomy and the value of natural history collections within conservation
and academic communities.

Key words: Conservation, long-lost synonymized species, natural history museums, tax-
onomy, vertebrates

In a recent paper, Long and Rodriguez (2023) highlighted the cases of species
that have dropped off science’s radar and proposed a new nomenclature to
guide conservation actions. They stated that species absent for at least 50
years should be termed “long-lost” species, those missing for more than 10
years as “lost species”, and those not found for 5-10 years as “missing spe-
cies”. However, the authors omitted to address taxonomy in their contribution,
a fundamental scientific discipline that itself can also be considered missing,
threatened or lost. It is a well-known fact that the ranks of taxonomists are
shrinking all around the world, and natural history museums (NHMs), where
many of them usually work, are suffering a parallel decline in support (Pearson
et al. 2011; Engel et al. 2021; Lobl et al. 2023).

In our opinion, unambiguously identified taxa are crucial to many biological
disciplines, especially conservation biology, as they represent the first compul-
sory steps to enable the confirmation or rejection of any scientific study that

119

Spartaco Gippoliti et al.: Lost species and neglected taxonomy

refers to them (Gippoliti and Amori 2002; Dubois 2003). More properly, Long
and Rodriguez (2023) focused on distinct and poorly known vertebrate taxa.
Obviously, no one can highlight and conserve species that are not yet known
to science and/or that are not formally described, and for this taxonomy, once
more, plays a crucial role. As a well-known fact, species must be formally de-
scribed prior to being subjected to red list assessments, as undescribed and/or
candidate species simply do not exist in terms of conservation priorities. This
is why taxonomic actions are often, albeit informally, encouraged (i.e., Zachos
et al. 2014).

We here report some cases regarding vertebrates, as they are better known
and easier to deal with in terms of nomenclature and taxonomy. As a remark-
able example we may cite amphibians, recently confirmed as the most threat-
ened terrestrial vertebrates, with more than 41% of described species reported
on IUCN Red Lists (8761 according to AmphibiaWeb 2024), but, at the same
time, with many new species discovered and constantly being described (Lu-
edtke et al. 2023). In fact, we believe that there is another category of “long-
lost” species that are often overlooked by conservation literature: the syn-
onymized (and often forgotten) taxa. Indeed, the 20" century was dominated
by an overly synthetic and “lumping” approach to the study of biological diver-
sity that resulted in a significant decrease in the number of recognized species,
especially in vertebrates. Several nominal species were forgotten because they
were synonymized or because they were moved to the sometimes-neglected
sub-specific rank (Burgin et al. 2018). Such an approach concerns mainly the
relatively well-known vertebrates, for which red lists are already available. For
invertebrates the situation is quite different, with species often known from just
a single specimen, the so-called “singletons” (Wells et al. 2019), and with red
lists, with just a few exceptions (Neubert et al. 2019), missing.

As a consequence, the “taxonomic inertia”, a term used to describe the per-
sistence of incomplete knowledge of biodiversity due to a lack of taxonomic
interest during most of the 20° Century (Gippoliti et al. 2017), propagated the
remarkable prevalence and persistence of taxonomic errors. So far, the sub-
species category has lost much of its scientific credibility and therefore many
subspecies were de facto forgotten by conservation biology (Gippoliti et al.
2018). Subspecies are often not even considered worthy of conservation atten-
tion except in a few cases, mostly involving charismatic animals such as the
mountain gorilla Gorilla beringei beringei Matschie 1903 (Gippoliti and Amori
2007). Among mammals, other than the well-known example of primates (Ry-
lands and Mittermeier 2014), even among supposedly well-known groups, such
as felids, there are overlooked species and resurrection of old synonyms, such
as in the case of the Neotropical genus Leopardus (Carnivora, Felidae) with the
recent re-validation of L. guttulus Hensel 1872 (do Nascimento and Feijo 2017).
African canids are also little known from a taxonomic perspective. For exam-
ple, the newly identified lineage of the African wolf Canis anthus F. Cuvier 1820
(Rueness et al. 2011) probably includes several distinct species (Allen 1939)
that were later considered subspecies of a phylogenetically distinct species,
the golden jackal C. aureus Linnaeus 1758 (Ellerman and Morrison-Scott 1951)
and that consequently disappeared from science’s radar for more than half a
century. This seems to be the case for the recently observed wolf in Danakil
that has been preliminarily assigned to C. mengesi Noack 1897 (Gippoliti and

Nature Conservation 56: 119-126 (2024), DOI: 10.3897/natureconservation.56.132036 120

Spartaco Gippoliti et al.: Lost species and neglected taxonomy

Lupi 2020), a long-lost synonymized taxon. New World deer of the tribe Odocoil-
eini offer a clear example of the effect of taxonomic inertia. At the end of the
20" century most authors recognized a total of 12 species in six genera (Corbet
and Hill 1986). Nowadays, after a revival of interest in their taxonomy and the
contribution of molecular techniques, the number of accepted genera is 10 with
around 20 species (Gutiérrez et al. 2017; Morales-Donoso et al. 2023; Barrio et
al. 2024). Interestingly, the 40% increase in recognized species in about forty
years is almost completely due re-evaluation of synonyms (the only exception
being the newly discovered Pudella carlae Barrio et al. 2024). Bisbalus citus Os-
good 1912 (Artiodactyla; Cervidae) is another good example of a “long-lost syn-
onymized” species. In fact, the genus Bisbalus is based on Mazama americana
cita, a taxon that has been overlooked, as most other subspecies, for almost a
century (Sandoval et al. 2024).

The importance of a revisionary and taxonomic working activity is even more
evident in other speciose vertebrate groups, like amphibians and reptiles. In
these cases, the intensification of field surveys and the use of integrative tools
(i.e., biomolecular, acoustics, etc.) often leads to the reevaluation of the fauna
of understudied geographic areas. Of course, the list of these works would be-
come almost endless and is one reason for the continued description of new
herpetological species (i.e., Kohler et al. 2005). As an example, in a recent work
that described 24 new species in the genus Mantidactylus (Anura; Mantellidae)
from Madagascar, a detailed comparison of historical museum types allowed
the re-validation of four old taxonomic names that were formerly synonymized
(and forgotten) for more than one century (Scherz et al. 2022). The availability
of new techniques to extract genetic and genomic data from historical muse-
um vouchers also led to the development of the exciting field of “Museomics”
that offers a further hope for filling a gap between natural history museums and
biodiversity researchers (Fong et al. 2023). Furthermore, incorporating avail-
able names by generating molecular data for types of synonymized species
is definitely feasible meanwhile and should be a standard protocol when de-
scribing new species using proper taxonomy. This would help in bringing these
names back into scientific process and helping to avoid nomenclatorial chaos.

It is also well-known that the historical vouchers may be the only evidence
for distinct and probably already extinct species, as is the case with the reptile
Lygodactylus neglectus Ceriaco and Passos 2023 found in the herpetological
collections of the Museu Nacional, Rio de Janeiro (Brazil) (Ceriaco and Pas-
sos 2023). First evidence for new species may originate directly from muse-
um-based revisions, as in the case of the Somalian mole-rat Fukomys ilariae
Gippoliti and Amori 2011 in the collection of the Museo Civico di Zoologia in
Rome (Gippoliti and Amori 2011) or, more recently, the two new species of Neo-
tropical small-eared Cryptotis shrews discovered at the Smithsonian National
Museum of Natural History (Woodman 2023). In other cases, museum spec-
imens offer definitive evidence regarding the former distributions of species
and open new taxonomic questions, as is the case of the recently verified Med-
iterranean occurrence in historical times of the small-eye hammerhead Sphyr-
na tudes (Valenciennes 1822) (Carcharhiniformes; Sphyrnidae) confirmed by
the (re)description of the specimen preserved in the NHM of Pisa University
(Collareta and Farina 2023). This species is today reported only from the At-
lantic coast of South America, from Venezuela to Uruguay (Pollom et al. 2020).

Nature Conservation 56: 119-126 (2024), DOI: 10.3897/natureconservation.56.132036 121

Spartaco Gippoliti et al.: Lost species and neglected taxonomy

Hammerhead sharks have declined in the Mediterranean Sea by more than
99.99% since the late 19* century (Ferretti et al. 2008) and the reappraisal of
historical sphyrnid specimens from natural history collections could help to
clarify the taxonomic identity and the true extent of species diversity, and its
decline, in the Mediterranean Basin.

Finally, as reported by Wheeler (2024), we underline here the central role
of natural history collections in systematics, and the pivotal role of NHMs as
centers for taxonomic research and preservation of voucher specimens, thus
reflecting species diversity as fully as possible, even of the most obscure tax-
onomic groups on Earth (Suarez and Tsutsui 2004; Fisher 2019). It is therefore
crucial that taxonomists and museum communities persist, and are fully inte-
grated, into worldwide efforts to map the biodiversity of the planet, including
the specific mission to rediscover and conserve lost and long-forgotten spe-
cies. Only with this in mind is it possible to re-align historical collections with
the new role of biodiversity assessment. And, obviously, taxonomy plays an
important role in resurrecting lost species, most of which are lost in museums.
It is time that international bodies, such as IUCN, recognize that biodiversity
conservation needs to be fully embedded in a wider vision that is fully aware
of the crucial role of botanical and zoological collections and of the people
that care for them and study them. Success relies on a future with a stronger
collaboration between the conservation and taxonomy communities. This will
certainly help in developing conservation actions that are urgently needed.

Acknowledgements

We are greatly indebted to George Beccaloni and Quentin D. Wheeler for having
read and improved this paper through their comments an earlier draft. We also
greatly thank Babu Saddam, Cristian Pertegal, Arni Sholihah, two anonymous
reviewers, and Klaus Henle for having provided useful comments and integra-
tions to our contribution.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

No funding was reported.

Author contributions

Conceptualization: FA, SF.

Author ORCIDs

Spartaco Gippoliti © https://orcid.org/0000-0001-5025-7216
Simone Farina ® https://orcid.org/0000-0002-6109-6961
Franco Andreone ® https://orcid.org/0000-0001-9809-5818

Nature Conservation 56: 119-126 (2024), DOI: 10.3897/natureconservation.56.132036 122

Spartaco Gippoliti et al.: Lost species and neglected taxonomy

Data availability

All of the data that support the findings of this study are available in the main text.

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