Mammal taxonomy without taxonomists: A reply to Zachos and Lovari

Abstract

Ontological and epistemological properties of the Phylogenetic Species Concept (PSC) as applied in recent mammalian taxonomic works are redefined and defended against criticisms raised by Zachos and Lovari (2013), which we find inapplicable to taxonomy because they relate more to the field of population biology. We summarize the negative impacts of the polytypic species concept for conservation and evolutionary biology, with emphasis on Rhinocerotidae. The priority need to embrace and strengthen museum-based taxonomic research is emphasized. The reply of Zachos and Lovari (2013) to our previous opinion pa-per (Gippoliti and Groves, 2012) provides us with the opportunity to expand our argument.

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Hystrix, the Italian Journal of Mammalogy
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Commentary
Mammal taxonomy without taxonomists: a reply to Zachos and Lovari
Spartaco Gippolitia,∗
, Fenton P.D. Cotterillb, Colin P. Grovesc
aViale Liegi 48A, 00198 Roma, Italy
bAEON – Africa Earth Observatory Network, Geoecodynamics Research Hub, c/o Department of Botany and Zoology, University of Stellenbosch, Stellenbosch 7602, South Africa
cSchool of Archaeology & Anthropology, Australian National University, Canberra, Australia
Keywords:
Rhinocerotidae
subspecies
conservation
polytypic species concept
Article history:
Received: 20 April 2013
Accepted: 24 April 2013
Abstract
Ontological and epistemological properties of the Phylogenetic Species Concept (PSC) as applied
in recent mammalian taxonomic works are redefined and defended against criticisms raised by
Zachos and Lovari (2013), which we find inapplicable to taxonomy because they relate more to the
field of population biology. We summarize the negative impacts of the polytypic species concept
for conservation and evolutionary biology, with emphasis on Rhinocerotidae. The priority need to
embrace and strengthen museum-based taxonomic research is emphasized.
The reply of Zachos and Lovari (2013) to our previous opinion pa-
per (Gippoliti and Groves, 2012) provides us with the opportunity to
expand our argument.
In 2011 three putative “subspecies” of the family Rhinocerotidae
were declared officially Extinct by IUCN: the western black rhinoceros
Diceros bicornis longipes Zukowsky, 1949; the Nile white rhinoceros
Ceratotherium simus cottoni (Lydekker, 1908) (with a handful of in-
dividuals surviving in captivity); and the Indochinese Javan Rhino-
ceros Rhinoceros sondaicus annamiticus Heude, 1892. We contend
that Zachos and Lovari’s reply (2013), and other similar papers (Heller
et al., 2013; Zachos et al., 2013a,b), are missing the core point. The
taxonomic approach they are defending has some responsibility for the
demise of so many unique forms of rhinoceros in Africa and Asia. The
raising of cottoni to specific level by Groves et al. (2010) is based more
on new data (and we should add real interest in a too long forgotten
question) than a shift in the species definition adopted. The Nile white
rhinoceros deserves species status on any or all species criteria, and
we fear that a demonstration of this would be apparent if interbreeding
with the southern species will be attempted as a last hope to “save” it.
Although even more poorly known due to their historic rarity, and the
consequent scarcity of study material, the same predicament may well
apply to Diceros bicornis longipes and Rhinoceros sondaicus annam-
iticus. Here knowledge of the palaeoclimatological and geomorpho-
logical history of both continents (Cotterill, 2003a; Whitmore, 1987;
Kingston, 2007; Senut et al., 2009), together with comparative phylo-
geographic studies of extant biogeographic patterns, strongly suggest
that these now extinct rhinoceros taxa actually constituted distinct spe-
cies. If we look at recent proposals of giraffe taxonomy, for instance
(Brown et al., 2007; Groves and Grubb, 2011), all recognize specific
status for the West-central African taxa that occurred sympatrically
with Diceros longipes and remained isolated from eastern populations.
This possibly related to an important barrier: Lake Megachad (Has-
sanin et al., 2007). We are not proposing a formal taxonomic revision,
which of course would require detailed research; but we would like
to draw attention to how – as exemplified by Roberts’ Lechwe Kobus
robertsi and Upemba Lechwe Kobus anselli (Cotterill, 2003b, 2005)
∗Corresponding author
Email address: spartacolobus@hotmail.com (Spartaco Gippoliti)
– a taxon becomes nearly invisible to conservation when it is “only a
subspecies”.
That said, we note that the cases cited by Zachos and Lovari (2013)
actually contradict, not support, their fear (Zachos et al., 2013a) of the
PSC as applied by Groves and Grubb (2011). Their Fig. 1 clearly shows
that species in the Cervus elaphus complex form monophyletic clades,
exactly as maintained by Groves and Grubb (2011) on the basis of com-
mon morphological characters. The fact that monophyletic lineages ex-
ist within each of these three clades (some of them picked out by roun-
ded squares in Zachos and Lovari’s (2013) Fig. 1) is irrelevant: Zachos
and Lovari appear to have forgotten that the species, however defined,
is a population concept (“a population or aggregation of populations” –
Groves and Grubb 2011:1). And indeed this fundamental ontological
nature of the species category was emphasized on the opening page
of Groves and Grubb 2011:1: ”a lineage . . . evolving separately from
others and with its own evolutionary role and tendencies” (Simpson,
1961), which is operationalized by the PSC (Cotterill, 2003b, 2005;
Groves and Grubb, 2011).
We emphasize that the concept of evolutionary species is a cent-
ral ontological foundation of phylogenetic systematics; the species-
as-lineage concept of the ESC (Evolutionary Species Concept) struc-
tures discovery operations – using the PSC and relevant evidence – to
characterize those lineages that qualify as individuated species. The
realm of organismal more-making , the tokogenic processes that create
populations, occupies the hierarchical level below that where species
evolve, persist and go extinct (Hennig, 1966; Frost and Kluge, 1994).
Zachos and Lovari are broadly correct that events testifying to the cir-
cumstances of lineage scission can be challenging to elucidate – but
this ontology accommodates such apparent conflicts in evidence (de
Queiroz, 2007; Mayden, 2013). At its most fundamental, this is the
nature of biodiversity resplendent in the one tree shaped by the gene-
alogical history of life on earth, and it evolved independently of our
ability to discover its topology. Alongside the realities of hypothesis-
driven biological research, this singularity obviates the credibility of
dividing or lumping organisms into artificial groups (Darwin, 1859;
O’Hara, 1988; Brooks and McLennan, 2002; Naomi, 2011; Wiley and
Lieberman, 2011; Mayden, 2013).
Again, this failure to understand the ontological context of how the
PSC is applied in the lineage framework of the ESC (cf. Cotterill
2003b, 2005) is revealed in their Fig. 2 and text (Zachos and Lovari,
Hystrix, the Italian Journal of Mammalogy ISSN 1825-5272 29th April 2013
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Hystrix, It. J. Mamm. (2013) — online first
2013), which focus on recent population fragmentation – with the In-
dian tiger Panthera tigris tigris as an example – and its possible genetic
consequences. Previously Zachos et al. (2013a) had heavily criticized
a three-species arrangement of tigers (Groves and Mazák, 2006) res-
ulting from the complex history of the Sunda Shelf and its sequence
of separation from and reconnection with mainland Asia in the Qua-
ternary (Michaux, 2010). Indian tigers may likewise present genetic
and morphological substructuring over their formerly vast range, and
this is as likely to be maintained by contemporary ecological processes
as by drift, as has been shown for boreal grey wolves (Musiani et al.,
2007). If today’s remnant isolates are discovered to be strongly differ-
entiated genetically, it is likely to be initially a legacy of this previous
substructuring, even if drift later modifies populations, and it is import-
ant that such processes should be considered in conservation strategy.
Unfortunately, too many translocations of wildlife have had detrimental
effects on the conservation of biodiversity by introducing alien genetic
material (Champagnon et al., 2012) because they are misled as a con-
sequence of prevalent taxonomic views based on the Biological Species
Concept (BSC) (Gippoliti, 2004).
We further argue that setting draconian restrictions on the evid-
ence judged sufficient to characterize species underestimates cryptic
diversity, especially where such actions seek to reduce “taxonomic in-
flation” (Bernardo, 2011). The argument that “If an independent evol-
utionary history, i.e. largely separate gene pools, is what ultimately
makes a species, then conclusive nuclear genetic or mitochondrial and
nuclear genetic data is what should be required from geneticists be-
fore splitting a species” (Zachos et al. 2013a: 5) violates the Precau-
tionary Principle in cases of some real species. Even a surfeit of gene
trees can still fail to discover some recently diverged species (Arnold
et al., 2013). These include lineages that (1) were founded by hybrid-
ization, (2) have experienced introgression, and/or (3) retain ancestral
polymorphisms (Norris and Hull, 2011).
In summary, criticism of excessive use of the diagnosability criterion
is unwarranted, not least for Groves and Grubb (2011), who maintained
subspecific taxa against PSC criteria.
We share the concern against non-peer-reviewed taxonomic de-
cisions (cf. Kaiser et al. 2013) but we maintain the importance of mu-
seum based taxonomy to offer solutions even with limited specimens
at hand – this Precautionary Principle sets the rule for little-known and
threatened taxa. Here, we emphasize that the overwhelming advant-
age that the PSC has over any other species concept is its testability;
the accumulation of further evidence can falsify a taxonomic statement
under the PSC. This is a requirement for a credible scientific proposi-
tion (Bernardo, 2011). As an historical example from before the advent
of the BSC, we highlight the pioneering revisions of Lorenzo Camer-
ano (1856-1917), director of the Zoological Museum at Turin Univer-
sity. Camerano stands out for introducing mathematical methods in
taxonomy and ecology (cf. Cohen 1994) and for studying species vari-
ability with large museum samples, in particular skulls, in his studies
of Rupicapra and Capra ibex taxonomy. In one of his last papers he
lamented the abuse of the subspecies category in several bovid species
(Camerano 1916: 4). In a little-known paper, he (Camerano, 1917)
challenged the validity of the taxonomic arrangement of the Iberian
ibexes proposed by Cabrera (1911: one species and four subspecies);
he proposed, on the basis of published data, and the study of two further
specimens in the Turin Museum, the existence of two distinct species,
Capra pyrenaica and Capra hispanica, with three subspecies possibly
of ancient hybrid origin (Camerano, 1917). This little-appreciated pa-
per (but see García-González 2011) offers an excellent example of a
phylogenetic approach to classical taxonomy, and is all the more per-
tinent today to inform conservation policies of threatened mammal pop-
ulations.
It is important to reconcile with the circumstances that the BSC
(Mayr, 1963) and the Genetic Species Concept (Baker and Bradley,
2006) are constrained by operational strictures as to where a species
“appears” in the process of lineage individuation. It is indeed unclear
how the BSC can actually discover lineages in the framework of phylo-
genetic systematics. Only the ESC is free of these operational strictures
(Frost and Kluge, 1994; Mayden, 2013). The poverty of operational-
ism and its primal failure of systematics was refuted over forty years
ago (Hull, 1968; Johnson, 1970; Frost and Kluge, 1994):
...the operational point of view depends for what success it
does have on the very element of science which it is designed
to eliminate. Operationism was intended as a cathartic to
purge physics of all non-empirical wastes, but it proved to
be so strong that the viscera were eliminated as well. (Hull
1968: 440).
An extreme version of operationalism has appeared in the Differential
Fitness Species Concept (DFSC) of Hausdorf (2011) – unfortunately
lauded by Frankham et al. (2002) in a recent publication – which has
tried to avoid the species-are-lineages ontology. This stricken attempt
to exhume a sterile research program removes the concept of the pop-
ulation from the definition of the species category; the DFSC strips
science of the reality of population biology, and ignores the hierarch-
ical organization of biodiversity, represented in the singular species tree
that systematics is challenged to discover. We are unconvinced by this
strange attempt to overturn a fundamental foundation of evolutionary
biology. Ultimately, all these attempts to dismiss the PSC (Frankham et
al., 2002; Heller et al., 2013; Zachos et al., 2013a,b; Zachos and Lovari,
2013) are crippled by operationalism. It is unfortunate that this ret-
rogressive perspective is held up as the way forward for conservation
policy, considering that the bioGenesis research strategy founded on
phylogenetic systematics (Donoghue et al., 2009; Hendry et al, 2010)
was developed expressly to guide policy formulated by the Conference
of the Parties (COP) to the Biodiversity Convention.
In the light of the consolidation of phylogenetic systematics and its
demonstrable progress toward discovering the singular Tree of Life
(Bernardo, 2011; Wiley and Lieberman, 2011; Mayden, 2013), all other
pretenses at dividing or lumping organisms into artificial taxonomic
groups can no longer be defended, at least on scientific grounds. Con-
servation costs of taxonomic mistakes can be extreme. We cited these
historical examples to stress the need to integrate old and new method-
ologies for the study of mammal diversity – before it is also too late, and
more taxa meet the same plight exemplified by Ceratotherium cottoni.
Especially in Western Europe, demands for knowledge, and improve-
ments in technology (Moodley and Bruford, 2007; Guschanski et al.,
2013), highlight the practicalities, and especially the urgency to restore
natural history museums and taxonomists into the epicenter of taxo-
nomic research (sensu Wheeler 2008; Cotterill and Foissner 2010). We
therefore fully support the view that “the headquarters of taxonomic
efforts are natural history museums, which need unrelenting support
in their fundamental goals to maintain present collections, accumulate
new ones, and support alpha-taxonomic and revisionary studies now
and into the future” (Reeder et al., 2007).
References
Arnold B., Corbett-Detig R.B., Hartl D., Bomblies K., 2013. RADseq underestimates di-
versity and introduces genealogical biases due to nonrandom haplotype sampling. Mo-
lecular Ecology (Online First). doi:10.1111/mec.12276
Baker R.J., Bradley R.D., 2006. Speciation in mammals and the genetic species concept. J.
Mammal. 87: 643–662.
Bernardo J., 2011. A critical appraisal of the meaning and diagnosability of cryptic evol-
utionary diversity, and its implications for conservation in the face of climate change.
In: Hodkinson T.R., Jones M.B., Waldren S., Parnell J.A.N. (Eds.) Climate Change,
Ecology and Systematics. Cambridge University Press: The Systematics Association.
380–438.
Brooks D.R., McLennan D.A., 2002. The Nature of Diversity: An evolutionary voyage of
discovery. The University of Chicago Press, Chicago.
Brown D.M., Brenneman R.A., Koepfli K.-P., Míla B., Georgiadis N.J., Louis E.E.L. jr.,
Grether G.F., Jacobs D.K., Wayne R.K., 2007. Extensive population genetic structure in
the giraffe. BMC Biol. 5: 57.
Cabrera A., 1911. The subspecies of the Spanish ibex. Proc. Zool. Soc. London 66: 963–
977.
Camerano L., 1916. I caratteri del cranio, della colorazione e delle corna nella distin-
zione dei Camosci in specie e sottospecie. Rivista di Antropologia (Roma) 20: 3–14.
[In Italian]
Camerano L., 1917. Contributo allo studio degli stambecchi iberici. Boll. Mus. Zool. Anat.
Comp. R. Univ. Torino 32: 1–30. [In Italian]
Champagnon J., Elmberg J., Guillemain M., Gauthier-ClercM., Lebreton J.-D., 2012. Con-
specifics can be aliens too: a review of effects of restocking practices in vertebrates. J.
Nature Conserv. 20: 231–241.
Cohen J.E., 1994. Lorenzo Camerano’s contribution to earlyfood web theory. Lecture Notes
in Biomathematics, 100: 351–359.
2

Mammal Taxonomy and the Phylogenetic Species Concept
Cotterill F.P.D., 2003a. Geomorphological influences on vicariant evolution in some
African mammals in the Zambesi Basin: some lessons for conservation. In: Plowman
A. (Ed.) Proceedings of the ecology and conservation of mini-antelopes: An Interna-
tional Symposium on Duiker and Dwarf Antelope in Africa. Filander Verlag, Füürth.
11–58.
Cotterill F.P.D., 2003b. Species concepts and the real diversity of antelopes. In: Plowman A.
(Ed.) Proceedings of the ecology and conservation of mini-antelopes: An International
Symposium on Duiker and Dwarf Antelope in Africa. Filander Verlag, Füürth. 59-118.
Cotterill F.P.D., 2005. The Upemba lechwe Kobus anselli: an antelope new to science em-
phasizes the conservation importance of Katanga, Democratic Republic of Congo. J.
Zool., Lond. 265: 113–132.
Cotterill F.P.D., Foissner W., 2010. A pervasive denigration of Natural History misconstrues
how biodiversity inventories and taxonomy underpin scientific knowledge. Biodiv. Con-
serv. 19: 291–303.
Darwin C.R., 1859. The Origin of Species. First Edition. John Murray, London.
de Queiroz K., 2007. Species concepts and species delimitation. Syst. Biol. 56: 879–886.
Donoghue M.J., Yahara T., Conti E., Cracraft J., Crandall K.A, Faith D.P., Häuser C.,
Hendry A.P., Joly C., Kogure K., 2009. bioGENESIS: providing an evolutionary frame-
work for biodiversity science. DIVERSITAS Report No. 6.
Hendry A.P., Lohmann L.G., Conti E., Cracraft J., Crandall K.A, Faith D.P., Häuser C.,
Häuser C., Joly C.A., Kogure K., Larigauderie A., Magallón S., Moritz C., Tillier S.,
Zardoya R. Prieur-Richard A.-H., Walther B.A., Yahara T., Donoghue M.J., 2010. Evol-
utionary biology in biodiversity science, conservation, and policy: a call to action. Evol-
ution 64: 1517–1528.
Frankham R., Ballou J.D., Dudash M.R., Eldridge M.D.B., Fenster C.B., Lacy R.C.,
Mendelson J.R. III, Porton I.J., Ralls K., Ryder O.A., 2002. Implications of different
species concepts for conserving biodiversity. Biol. Conserv. 153: 25–31.
Frost D.R., Kluge A.G., 1994. A consideration of epistemology in systematic biology, with
special reference to species. Cladistics 10: 259–294.
García-González R., 2011. Elementos para una filogeografía de la Cabra Montés ibérica
(Capra pyrenaica Schinz, 1838). Pirineo, Rev. Ecol. Mont. 166: 87–122.
Gippoliti S., 2004. Captive breeding and conservation of the European mammal diversity.
Hystrix 15(1): 35–53.
Gippoliti S., Groves C.P., 2012. “Taxonomic inflation” in the historical context of mamma-
logy and conservation. Hystrix 23(2): 6–9.
Groves C.P., 2013. The nature of species: A rejoinder to Zachos et al. Mamm. Biol. 78:
7–9.
Groves C.P., Fernando P., Robovský J., 2010. The sixth rhino: a taxonomic re-assessment
of the Critically Endangered northern white rhinoceros. PLoS ONE 5(4): e9703. doi:
10.1371/journal.pone.0009703
Groves C., Grubb P., 2011. Ungulate Taxonomy. The Johns Hopkins University Press, Bal-
timore.
Groves C., Mazák J., 2006. A taxonomicrevision of the tigers (Panthera tigris) of Southeast
Asia. Mamm. Biol. 71: 268–287.
Guschanski K., Krause J., Sawyer S., Valente L. M., Bailey S., Finstermeier K., Gilissen
E., Sonet G., Nagy Z.T., Lenglet G., Mayer F., Savolainen V., 2013. Next-Generation
Museomics disentangles one of the largest primate radiations. Syst. Biol. (Advance Ac-
cess) doi:10.1093/sysbio/syt018
Gutiérrez E.E., Helgen K.M., 2013. Outdated taxonomy blocks conservation. Nature 495:
314.
Heller R., Frandsen P., Lorenzen E.D., Siegismund H.R., 2013. Are there really twice as
many bovid species as wethought? Syst. Biol. 62(3): 490–493. doi:10.1093/sysbio/syt004
Hassanin A., Ropiquet A., Gourmand A.-L., Chardonnet B., Rigoulet J., 2007. Mithocon-
drial DNA variability in Giraffa camelopardalis: consequences for taxonomy, phylo-
geography and conservation of giraffes in West and central Africa. C.R. Biologies 330:
265–274.
Hausdorf B., 2011. Progress toward a general species concept. Evolution 65: 923–931.
Hennig W., 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, Illinois.
Hull D.L., 1968. The operational imperative: sense and nonsense in operationism. Syst.
Zool. 17: 438–457.
Kaiser H., Crother B.I., Kelly C.M.R., Luiselli L., O’Shea M., Ota H., Passos P., Schleip
W.D., Wüster W., 2013. Best practices: In the 21th Century, taxonomic decisions in
herpetology are acceptable only when supported by a body of evidence and published
via peer-review. Herpet. Rev. 44: 8–23.
Kingston J.D., 2007. Shifting adaptive landscapes: progress and challenges in reconstruct-
ing early hominid environments. Amer. J. Phys. Anthrop. 134: 20–58.
Johnson L., 1970. Rainbow’s end: the quest for an optimal taxonomy. Syst. Zool. 19: 203–
239.
Mayden R.L., 2013. Species, trees, characters, and concepts: ongoing issues, diverse ideo-
logies, and a time for reflection and change.In: Pavlinov I.Ya. (Ed.) The Species Problem
– Ongoing Issues. InTech. 171–191.
Mayr E., 1963. Animal Species and Evolution. Belknap Press, Harvard.
Michaux B., 2010. Biogeology of Wallacea: geotectonic models, areas of endemism, and
natural biogeographical units. Biol. J. Linnean Soc. 101: 193–212.
Moodley Y., Bruford M.W., 2007. Molecular biogeography: towards an integrated frame-
work for conserving Pan-African biodiversity. PLoS ONE 2(5): e454. doi:10.1371/journal.
pone.0000454
Musiani M., Leonard J.A., Cluff H.D., Gates C., Mariani S., Paquet P.C., Vilàs C., Wayne
R.K., 2007. Differentiation of tundra/taiga and boreal coniferous forestwolves: genetics,
coat colour and association with migratory caribou. Mol. Ecol. 16: 4149–4170.
Naomi S.-I., 2011. On the integrated frameworks of species concepts: Mayden’s hierarchy
of species concepts and de Queiroz’s unified concept of species. Journal of Zoological
Systematics and Evolutionary Research 49: 177–184.
Norris R.D., Hull P.M., 2011. The temporal dimension of marine speciation. Evol. Ecol.
26: 393–415.
O’Hara R.J., 1988. Homage to Clio, or, toward an historical philosophy for evolutionary
biology. Syst. Zool 37: 142–155.
Reeder D.M., Helgen K.M., Wilson D.E., 2007. Global trends and biases in new mammal
species discoveries. Occ. Papers Mus. Texas Tech. Univ. 269: 1–35.
Senut B., Pickford M., Ségalen L., 2009. Neogene desertification of Africa. Comptes Ren-
dus Geoscience, 341(8–9): 591–602.
Sharma R., Stuckas H., Bhaskar R., Khan I., Goyal S.P., Tiedemann R., 2011. Genetically
distinct population of Bengal tiger (Panthera tigris tigris) in Terai Arc Landscape (TAL)
of India. Mamm. Biol. 76: 484–490.
Simpson G.G., 1961. Principles of Animal Taxonomy. Columbia University Press, New
York.
Wheeler Q.D. (Ed.), 2008. The New Taxonomy. Systematics Association Special Volume
Series. CRC Press, Boca Raton, FL.
Whitmore T.C. (Ed.), 1987. Biogeographical Evolution of the Malay Archipelago. Claren-
don Press, Oxford.
Wiley E.O., Lieberman B.S., 2011. Phylogenetics: theory and practice of phylogenetic sys-
tematics. Second Edition. Wiley-Blackwell.
Zachos F.E., Apollonio M., Bärmann E.V., Festa-Bianchet M., Göhlich U., Habel J.C., Har-
ing E., Kruckenhauser L., Lovari S., McDevitt A.D., Pertoldi C., Rössner G.E., Sánchez-
Villagra M.R., Scandura M., Suchentrunk F., 2013a. Species inflation and taxonomic
artefacts – A critical comment on recent trends in mammalian classification. Mamm.
Biol. 78: 1–6.
Zachos F.E., Clutton-Brock T.H., Festa-Bianchet M., Lovari S., Macdonald D.W., Schaller
G.B., 2013b. Species splitting puts conservation at risk. Nature 494: 35.
Zachos F.E., Lovari S., 2013. Taxonomic inflation and the poverty of the Phylogenetic Spe-
cies Concept – a reply to Gippoliti and Groves. Hystrix (Online First) 10.4404/hystrix-
24.1-8849
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