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when they are clearly more expert than
I am. It helps to remind myself of this
advice.
The other best advice that I received
was also from Andrew Cockburn. As an
undergraduate, I worked summers as
one of the research assistants on the
‘superb fairywren project’. We were each
responsible for monitoring a subset of
breeding territories, and I was surprised
that Andrew monitored some himself,
on top of all his other responsibilities. He
told me that, if he didn’t continue regular
fieldwork, it would negatively affect his
ability to think critically about the system
and form new ideas. While I’m still
relatively junior, it is already becoming
clear how hard it is to find the time to
stay at the coalface of fieldwork. Keeping
this advice in mind, I’m trying very hard to
not become a visitor to my own projects.
What do you think are the big
questions to be answered next in your
field? The capacity for innovation, social
learning and culture is vitally important
in our success as a species, facilitating
colonisation of almost every habitat on
Earth. Yet, while once thought to be
almost exclusively human, research
over the last two decades has extended
the reach of culture. Cultural traits have
been identified across a multitude of
species and behavioural domains, from
migration routes in ungulates and song in
passerine birds to foraging behaviour in
killer whales. Experimental studies have
demonstrated the capacity for culture in
yet more species.
We are now at an inflection point in
the study of animal culture where we can
move to asking about the consequences
of these cultures for the evolution and
ecology of the animals embedded in
them. This leads to so many thrilling
possible questions, but three ‘big ones’
immediately spring to mind. First, what is
the influence of culture on the evolution
of cognition and associated life history
traits? Second, can we disentangle
relative contributions of genetic
inheritance and cultural inheritance,
and when do we observe gene–culture
co-evolution? Third (and my favourite),
what role do social learning and cultural
inheritance of behaviour play in shaping
locally adaptive responses to changing
environments?
Which aspect of science, your field or
in general, do you wish the general
R1070 Current Biology 31, R1065–R1071, September 27, 2021 © 2021 Elsevier Inc.
public knew more about? That the
study of animal behaviour matters for
conservation. I do a lot of my work in
urban areas, studying species that have
been highly successful at adapting
to human-induced change. People
passing by often ask me why I bother to
study the behaviour of these ‘common
birds’ — is it just for fun? But proxies for
cognition such as relative brain size are
predictors of urban success across a
wide range of species. By studying the
mechanisms that allow some species to
exhibit such behavioural flexibility, we
can learn from these ‘success stories’
for species management more generally.
Recent work by myself and
colleagues has shown that animal
cultures also matter for conservation.
If skills and behaviours are held
in populations as socially learnt
knowledge, they are both dynamic
and vulnerable. For example, vital
knowledge may be lost if local
populations are extirpated or older
individuals are selectively removed.
Conversely, cultures can potentially
change very quickly in some species,
with adaptive behaviours emerging
in response to new opportunities via
the spread of innovation. These ideas
are intuitive to many people, but for
others they require a fundamental shift
in the way they view animals and their
behaviour.
If you had not made it as a scientist,
what would you have become? Is
crazy parrot lady a possibility? No?
Well, I also studied environmental and
international laws, so I suppose I would
have fallen back on that. Towards the
end of my undergraduate degrees, I was
deciding between pursuing a PhD in
behavioural ecology or moving towards
a career in policy. At the time, I felt
strongly that conservation policy work
was important and interesting, but I just
couldn’t resist the siren call (and fun) of
pure research. I’ve been lucky enough
to dabble occasionally in conservation
policy in a scientific advisory capacity,
but to be honest I think I would have
made a pretty mediocre lobbyist — so
maybe it’s all for the best.
Cognitive and Cultural Ecology Lab, Max
Planck Institute of Animal Behaviour,
Radolfzell, Germany, and Edward Grey
Institute, Department of Zoology, University of
Oxford, Oxford, UK.
E-mail: laplin@ab.mpg.de
Quick guide
Salinella
Erik Tihelka1,2 and Chenyang Cai1,2,*
What makes Salinella interesting?
Advances in genomics have largely
clarified the status of most unusual
animals that could not be easily placed
on the animal tree of life, such as
Buddenbrockia, Chaetognatha, Myxozoa
and Xenoturbella. In some ways, Salinella
is one of the last truly mysterious
animals — mostly because nobody has
seen it since the 19th century.
So, what exactly is Salinella? Nobody
knows for sure. Salinella salve was
described in 1891 by the German
zoologist Johannes Frenzel. Frenzel
meticulously illustrated the taxon in his
1892 book on the microscopic fauna
of Argentina (Figure 1). It appears
to be sausage-shaped, bilaterally
symmetrical, possessing a gut starting
with a mouth and ending with an anus,
both surrounded by elongate cilia, a
dorsal and a ventral side, and its body
wall composed of a single layer of cells
ciliated on both sides (Figure 1). This
suite of characters excludes it from
all known animal groups. Frenzel saw
Salinella as bridging the gap between
unicellular eukaryotes and complex
animals. However, given some of its
derived characters such as bilateral
symmetry and a through gut, it is equally
possible that Salinella occupies a much
more derived position on the animal tree
of life and lost some features secondarily.
Given persisting controversies over the
basal branching order within Metazoa,
Salinella may seem as relevant to the
discourse on animal phylogeny today as
it was almost 130 years ago.
Where was it collected? Frenzel
received soil from the salt flats around
Río Cuarto, in the south of the Province
of Córdoba, Argentina, in 1890 or 1891.
He placed it in an aquarium, added
freshwater, creating an approximately
2% saline solution, and did what many
of us have done in our laboratories —
forgot about his material. When he came
back weeks later, he noticed, to his great
surprise, approximately 2-mm-long
vermiform organisms attached to the
aquarium walls and substrate. Rearing

Magazine
Figure 1. Some of the only published drawings of Salinella salve.
Dorsal (A), ventral (B), and lateral (C,D) views. Illustration by J. Frenzel and published in Archiv für
Naturgeschichte (1892). Scale bar: 200 µm.
experiments revealed that S. salve
reproduced asexually by transverse
fission and probable unicellular juveniles
were described. Frenzel also observed
conjoined adults, but their subsequent
fate could not be tracked.
What have biologists since Frenzel
thought of Salinella? Salinella
combines some animal traits, such as
a through gut, with features not known
in any living metazoan, such as a body
wall consisting of a single cell-layer
body wall and lack of interior cells. It
thus ignited a long-lived controversy
in zoological circles. Opinions have
ranged from treating Salinella as a
species of uncertain placement within
kingdom Metazoa or a member of a
separate animal phylum to outright
dismissal. In agreement with some of his
contemporaries, Richard Blackwelder
erected the phylum Monoblastozoa in
1963 with Salinella as its only member,
highlighting its single-celled body wall as
a key diagnostic feature. Some biology
textbooks continue to list Monoblastozoa
as an enigmatic animal phylum of
uncertain affinity. Others have postulated
that Salinella could have been a
misidentified ciliate or a fruit of Frenzel’s
imagination. Yet, Frenzel addressed at
length its differences from protozoans
and also described numerous other
taxa that remain uncontroversial to date.
Thus, Salinella is the sole representative
of the only metazoan phylum whose very
existence is disputed.
Has Salinella been re-discovered
since? Frenzel was never able to revisit
Argentina and look for Salinella again,
he fell to his death on October 21st,
1897 at lake Müggelsee near Berlin
during biological fieldwork. He never
specified Salinella’s exact type locality,
and it may now be lost given that large
parts of the Córdoba Province have
been converted to arable land. Recent
attempts at recollecting Salinella were
made by the German malacologist
Michael Schrödl and colleagues, who
collected soil samples from saline lakes
across Argentina and Chile and treated
them following Frenzel’s protocol, but to
no avail.
Where is the type material deposited?
Much like the animal itself, the type
material of Salinella cannot be traced.
Current Biology 31, R1065–R1071, September 27, 2021 R1071
ll
Frenzel’s documents suggest that he
took his collection to Germany after
leaving Argentina in late 1891. None
of it is held at the National University
of Córdoba today. However, nothing is
known of the collection’s subsequent
whereabouts. It is unclear if Frenzel
ever prepared type material of S. salve.
He explicitly wrote that the animals
disintegrate completely upon contact
with preservatives.
Does Salinella deserve a place in
biology textbooks? Speaking from
a strictly nomenclatural point of view,
the absence of traceable type material
and lack of recent specimens does
not invalidate the taxonomic status of
S. salve. Nonetheless, S. salve qualifies
for the status of species inquirenda,
i.e. a species of doubtful or disputed
identity. Admittedly, many microscopic
invertebrates are known from a few
specimens or collections and many
species described over a century
ago have not been recollected for a
long time. Ultimately, proving that an
animal doesn’t exist is more difficult
than the opposite. Yet, it is not entirely
out of question that future surveys of
Argentina’s salt pans, perhaps with the
aid of metabarcoding, could shed new
light on the Salinella enigma.
Where can I find out more?
Blackwelder, R.E. (1963). Classification of the Animal
Kingdom (Southern Illinois University Press).
Brusca, R.C., and Brusca, G.J. (1990). Invertebrates
(Sinauer Associates).
Dunning, H. (2012). Gone missing, circa 1892. The
Scientist 26, 84.
Frenzel, J. (1891). Das Mesozoon: Salinella. Biol.
Zbl. 11, 577–581.
Frenzel, J. (1892a). A multicellular infusorian-like
animal. Ann. Mag. Nat. Hist. Ser. 6 49, 109–111
[translated from Zool. Anz. 367, 230–233 (1891)].
Frenzel, J. (1892b). Untersuchungen über die
mikroskopische Fauna Argentiniens. Salinella
salve nov. gen. nov. spec. Ein vielzelliges,
infusorienartiges Tier (Mesozoon). Arch.
Naturgesch. 58, 66–96.
Giribet, G., and Edgecombe, G.D. (2020). The
Invertebrate Tree of Life (Princeton University
Press).
Kowalczuk, I.-S. (1987). Prof. Dr. Johannes
Frenzel (1858 bis 1897) — Der Begründer
der fischereilischen Forschungsstätte in
Berlin-Friedrichshagen. Z. Binnenfischerei 34,
320–325.
1
State Key Laboratory of Palaeobiology and
Stratigraphy, Nanjing Institute of Geology and
Palaeontology, and Centre for Excellence in
Life and Paleoenvironment, Chinese Academy
of Sciences, Nanjing 210008, China. 2School
of Earth Sciences, University of Bristol, Life
Sciences Building, Tyndall Avenue, Bristol
BS8 1TQ, UK.
*E-mail: cycai@nigpas.ac.cn