Origin of animal multicellularity:

precursors, causes, consequences—the

rstb.royalsocietypublishing.org choanoflagellate/sponge transition,
neurogenesis and the Cambrian explosion
Opinion piece Thomas Cavalier-Smith
Cite this article: Cavalier-Smith T. 2017 Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK
Origin of animal multicellularity: precursors,
Evolving multicellularity is easy, especially in phototrophs and osmotrophs
causes, consequences—the choanoflagellate/ whose multicells feed like unicells. Evolving animals was much harder and
sponge transition, neurogenesis and the unique; probably only one pathway via benthic ‘zoophytes’ with pelagic
Cambrian explosion. Phil. Trans. R. Soc. B 372: ciliated larvae allowed trophic continuity from phagocytic protozoa to gut-
20150476. endowed animals. Choanoflagellate protozoa produced sponges. Converting
http://dx.doi.org/10.1098/rstb.2015.0476 sponge flask cells mediating larval settling to synaptically controlled nemato-
cysts arguably made Cnidaria. I replace Haeckel’s gastraea theory by a
sponge/coelenterate/bilaterian pathway: Placozoa, hydrozoan diploblasty
Accepted: 5 September 2016 and ctenophores were secondary; stem anthozoan developmental mutations
arguably independently generated coelomate bilateria and ctenophores.
One contribution of 17 to a theme issue I emphasize animal origin’s conceptual aspects (selective, developmental)
related to feeding modes, cell structure, phylogeny of related protozoa,
‘Evo-devo in the genomics era, and the origins
sequence evidence, ecology and palaeontology. Epithelia and connective
of morphological diversity’. tissue could evolve only by compensating for dramatically lower feeding effi-
ciency that differentiation into non-choanocytes entails. Consequentially,
Subject Areas: larger bodies enabled filtering more water for bacterial food and harbouring
cellular biology, evolution, microbiology, photosynthetic bacteria, together adding more food than cell differentiation
sacrificed. A hypothetical presponge of sessile triploblastic sheets (connec-
palaeontology, developmental biology
tive tissue sandwiched between two choanocyte epithelia) evolved oogamy
through selection for larger dispersive ciliated larvae to accelerate benthic
Keywords: trophic competence and overgrowing protozoan competitors. Extinct Vendo-
epithelial origin, placozoa, nematocyst, zoa might be elaborations of this organismal grade with choanocyte-bearing
selective advantages of cell differentiation, epithelia, before poriferan water channels and cnidarian gut/nematocysts/
synapses evolved.
vendozoa, origin of bilateria
This article is part of the themed issue ‘Evo-devo in the genomics era,
and the origins of morphological diversity’.
Author for correspondence:
Thomas Cavalier-Smith
e-mail: tom.cavalier-smith@zoo.ox.ac.uk

1. Introduction: unicells to multicells (and vice versa)

Unicells vastly outnumber multicells and are far more important for the biosphere
in biogeochemical recycling. Bacteria and protists greatly exceed vertebrates in
different kinds of organism too. Lamarck thought unicells so evolutionarily
recent that they had not yet had time to inexorably become multicellular. Not
so; they existed billions of years longer than complex multicells and may outlive
them. There are hordes of excellent unicellular niches; multicellularity is often
selectively disadvantageous. Yeasts evolved multiply from multicellular filamen-
tous ancestors; Myxozoa are parasitic unicells that evolved from animals with
nervous systems (early-branching Cnidaria), losing epithelia, connective tissue,
nerves and 70% of genes as useless, only their multicellular spores keeping nema-
tocysts [1]. So how and why do some lineages become multicellular? Evolving
multicellularity is mechanistically extremely simple. Every unicell group has a
cellular and mutational potential to do so given a selective advantage.

& 2016 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.
 Multicellularity evolves in two ways. Naked cells, as in ani- them with eumetazoa, but doubted the homology of choanocy- 2
mals and slime moulds, evolve glue to stick together. Walled tes and choanoflagellates. For over a century, opinion ebbed

rstb.royalsocietypublishing.org
cells modify wall biogenesis to inhibit the final split that nor- and flowed between these contradictory views, until sequence
mally makes separate unicells, so daughters remain joined. trees proved that sponges are related to other animals and
The ease of blocking that split allowed almost every group choanoflagellates are the closest protozoan relatives of animals
of bacteria, fungi and plants (and many chromists) to evolve [1,7–9]. Ultrastructurally, collars of both consist of a circlet of
multicellular walled filaments, more rarely two-dimensional microvilli crosslinked by a mucus mesh into an extremely effec-
sheets, most rarely three-dimensional tissues. Tissues require tive bacterial filter; trapped bacteria are moved down to the cell
more geometric control of daughter wall orientation, as in body for phagocytosis [10]. Unaggregated microvilli are pre-
embryophyte green plants and chromist brown algae; both sent generally on the cell body; choanoflagellate microvilli
can grow longer than blue whales. Evolving tissues is selec- exemplify a broader class of narrrow cell extensions (filodigits
tively harmful to many walled multicells whose filaments [11]) supported by a tight actin-filament bundle that probably

Phil. Trans. R. Soc. B 372: 20150476

are best for reproductive success. Almost all multicells retain evolved from ancestral opisthokont filopodia in the common
unicellular phases (eggs, sperm, zygotes), so adhesion is tem- ancestor of holozoa, the clade comprising animals, choano-
porally controlled and developmentally reversible—except flagellates and Filosporidia (figure 1). Fascin that crosslinks
for purely clonal vegetatively propagating plants or ‘colonial’ filodigit actin filaments, villin at their base, signalling protein
invertebrates (evolutionarily transient) the only organisms Vav-1 at their tips, myosin X (transporting proteins to their
that are never unicellular. tip), and other functionally related proteins, all originated in
Merely joining daughter cells together suffices to create effi- the holozoan last common ancestor [13], as did several other
cient multicellular phototrophs or osmotrophic saprotrophs myosins [14]; that strongly substantiates the conceptual distinc-
because their essential trophic features remain intracellular. tion of filodigits from more generalised filopodia of deeper
A bacterial or protist phototroph can easily become multicellu- branching protozoa that lack them as well as filodigits being
lar while maintaining the same way of feeding and identical the key morphological synapomorphy for holozoa [11]. Filo-
cell functions. An algal filament feeds (on light, H2O, CO2 digits, absent from other protists, were presumably present in
and minerals) just as does a single cell; so does a saprotrophic the immediate ancestors of the first stem choanoflagellates
bacterial or fungal filament. However, a phagotrophic amoeba from which animals almost certainly evolved; they apparently
could not aggregate into a multicellular body and still locomote evolved at the same time as cadherins that might initially
and feed the same way. Nor could most other protozoa. Many have been involved in holozoan biology long before animals
amoebae have become multicellular, but only temporarily recruited them for epithelial cell adhesion [15]. Of choano-
for spore dispersal, not feeding. Aggregative multicellularity flagellate orders, the mostly surface-attached Craspedida of
has produced multispore fruiting bodies numerous times more primitive morphology and feeding mode are excellent
in fundamentally different protist lineages (dictyostelids in models for stem choanoflagellates also, except for having lost
Amoebozoa, Guttulinopsis in Cercozoa, in Ciliophora [2]), so some key animal precursor proteins that remain in more distant
is evolutionarily easy. That is because their multicellular protists; planktonic Acanthoecida whose collar filodigits
phases are non-trophic; they evolved purely for efficient manipulate secreted silica strips into elaborate loricas (enabling
aerial spore dispersal, free of conflict between need to a novel filter feeding mode) are highly derived, not directly rel-
feed and to aggregate; spores still function as unicells. Thus, evant to animal origins [10].
Dictyostelium [3] is irrelevant for properly understanding Among extant animals, only sponges could have evolved
animal multicellularity origins. directly from protozoa without changing feeding mode. The
key problems in understanding animal origins are therefore
how and why sponges evolved from a craspedid-like stem
choanoflagellate and later generated all other animals. I
2. Uniqueness of animal multicellularity attempt to explain both after briefly outlining enabling proto-
If evolving multicellularity is mechanistically so easy for zoan innovations. I shall emphasize simple conceptual
bacteria and protists, then why did animals evolve only aspects of the choanoflagellate/animal transition, often over-
once? Primarily, because it is selectively immensely harder looked but more important than discovering extra protozoan
for organisms that feed by swallowing others or bits of them genes suitable as precursors to animal functions. Such ances-
(a purely eukaryotic propensity) to switch from intracellular tral features exist in both choanoflagellates and more distant
phagocytosis, as in amoebae or ciliates, to eating with a multi- protozoan relatives of substantially different cell structures
cellular mouth and gut, whose cells have novel functions and and feeding mode [12].
structures absent in their unicellular ancestors. Animal feeding In the light of site-heterogeneous trees using 187 protein
is effective only if novel cell types cooperate at a higher organ- sequences [16,17], figure 1 summarizes the major eukaryote
izational level; most give up the ability to feed or reproduce, clades and key steps in eukaryote cell evolution that paved
huge selective disadvantages not easily overcome. the way for later innovations that generated animals. Like choa-
In 1866, James-Clark discovered choanoflagellate protozoa noflagellates, Filosporidia (next most distant animal cousins)
and their feeding on bacteria trapped by a collar surrounding belong to the protozoan phylum Choanozoa that ancestrally
their undulating cilium that generates the water current that evolved a swimming mode with a single posterior cilium
draws them towards it. He noted that sponge collar cells (choa- (i.e. opisthokont—‘posterior oar’ in Greek) like archetypal
nocytes) have the same structure and feeding method, correctly animal sperm or fungal zoospores that evolved by modify-
suggesting that sponges evolved from a choanoflagellate [4]. ing ancestral opisthokont cell structure [11,16]. Immediate
Often sponges were thought unrelated to other animals, outgroups to opisthokonts are successively more distant
being classified in Protista by Haeckel and Protozoa by Kent branching predominantly biciliate lineages of phylum Sulcozoa
[5]. Schulze [6] argued that sponge spermatogenesis allied that typically move not by swimming but by gliding on surfaces
 Bilateria 3
scotokaryotes
Coelenterata
anus, coelom collar

rstb.royalsocietypublishing.org
Placozoa tentacles/synapses
Porifera FUNGI
ANIMALIA Choanoflagellatea
Microsporidia
epithelia, ECM
eggs, sperm Rozellidea
Filosporidia
collar filter Cristidiscoidea
cadherins filodigits
pseudopodia
Choanozoa
anterior cilium lost pseudopodia
Apusomonadida Amoebozoa

Phil. Trans. R. Soc. B 372: 20150476

opisthokonts
Breviatea Diphylleida
catenins pseudopodia
integrins? Mantamonas
pseudopodia dorsal pellicle
myosin II
Sulcozoa Planomonadida
ciliary gliding
Malawimonas corticates
PLANTAE CHROMISTA
Neolouka
groove chloroplast
groove
Metamonada
4 cilia/centrioles Colponema
anaerobes cortical alveoli
ventral ciliary vane novel cytochrome c biogenesis
Eolouka cytopharynx
Jakobids (e.g. Reclinomonas) Eozoa
+ Tsukubamonas dorsal
4 cilia/centrioles ciliary vane Euglenozoa
Percolozoa
e.g. Naegleria eruptive ventral feeding ciliary paraxonemal rods
pseudopodia groove
Figure 1. Cell structure divergence in phagotrophic non-amoeboid flagellates provided the basis for evolving animals, fungi, plants and chromists. Pseudopodia
evolved secondarily, myosin II providing the basis for pseudopodia in animals, Amoebozoa (and Percolozoa) and muscles. Chloroplasts, originating when the plant
ancestor enslaved and modified undigested cyanobacteria, were transferred laterally (red arrow) to make chromists (e.g. brown seaweeds, diatoms, dinoflagellates)
whose ancestor modified an enslaved undigested red alga. The most basic eukaryote structural dichotomy contrasts Euglenozoa (parallel centrioles; cilia with para-
xonemal rods; cytopharynx for feeding) and excavates (Percolozoa, Eolouka, Neolouka: orthogonal centrioles: no paraxonemal rods; feeding by phagocytosing prey
drawn into a ventral groove by posterior ciliary currents). The pre-animal lineage lost excavate groove-feeding by evolving ventral ciliary gliding locomotion to
generate Sulcozoa, protozoa with a dorsal proteinaceous pellicle (blue). Irrespective of whether the eukaryote tree is rooted within the protozoan subkingdom
Eozoa as shown (most likely) or beside Eolouka-like Reclinomonas with the most primitive mitochondria, the immediate ancestors of animals (Choanozoa)
arose by loss of the anterior cilium and sulcozoan dorsal pellicle to make opisthokonts (in red) with a radically simplified, more radially symmetric, microtubular
cytoskeleton. Long actin-supported filodigits arose in the ancestor of Filosporidia and choanoflagellates and became a circlet of microvilli to make the choanofla-
gellate/sponge collar for catching bacteria. Filosporidia comprise Filasterea, Ichthyosporea, Corallochytrea [12]. The four derived kingdoms (e.g. ANIMALIA, PLANTAE)
are shown in upper case; all taxa in lower case belong to the basal eukaryotic kingdom Protozoa.

by ciliary surface motility propelling one semi-rigid cilium and modifications, but radically simplified and made more
feed by emitting newly evolved, bacteria-grabbing, branching symmetric during the origin of the opisthokont body plan
pseudopodia from the cell’s ventral ciliary groove [11,16]. by anterior ciliary loss, possibly in association with a
Sulcozoan flagellates clearly could not have retained their protochoanoflagellate feeding mode [11].
characteristic locomotory or feeding modes had they evolved Knowing the structure and evolutionary potential of the
glue to stick together as a multicellular organism; such mutants closest relatives and ancestors of animals (figure 1) and that
would necessarily quickly starve to death. Nor could their opisthokont cells were radically simplified compared with
immediate ancestors—three successive groups of swimming, their ancestors does not directly explain animal origins, but
not gliding flagellates (i.e. Neolouka, Eolouka, Percolozoa) col- helps distinguish central from peripheral aspects of the pro-
lectively called excavates because their ventral groove looks cess and avoid pitfalls from erroneous assumptions about
more obviously scooped out [11,17]. The groove phagocytoses ancestors. Most things we inherit from our unicellular ances-
prey propelled therein by both cilia, the posterior often having tors evolved before the excavate/Euglenozoa split. Only a
one or two lateral vanes to increase its thrust. Their ancient few arose within the scotokaryote clade that embraces opistho-
groove-supporting asymmetric cytoskeleton, with five distinct konts, Amoebozoa, Sulcozoa and Neolouka, and is sister to
microtubular ciliary roots and many characteristic filaments, the cytologically substantially different plant/chromist clade
was inherited by Sulcozoa, initially with diverse minor (Corticata) [17].
 Integrins and associated molecules used for epithelial cell epithelia and connective tissue cells embedded separately 4
adhesion to extracellular matrix (ECM) were secondarily lost in a gelatinous mesohyl. Did epithelia evolve first or did

rstb.royalsocietypublishing.org
by choanoflagellates and fungi; without full genomes for the epithelia and mesenchyme coevolve?
deepest branching Sulcozoa, the exact point of origin is Four different ways of making multicellular choanofla-
unclear (figure 1): though not yet known for branches gellates exist. Many become ‘colonial’ sessile organisms by
before Breviatea, integrins might have arisen earlier with sco- evolving thin extracellular stalks that join cells together to
tokaryote pseudopodia, for mediating reversible adhesion to form branched tree-like structures analogous to corals or
the substratum and/or pseudopodial actin bundle attach- plants [5,10]. Other flagellate groups also evolved multicellular
ment/assembly via talin/vinculin that certainly evolved sessile lineages with branching stalks; many heterotrophic, e.g.
earlier [12], at least prior to Amoebozoa. If, instead, integrins biciliate bicoecids (heterokont chromists), pseudodendromo-
help actin attachment to sulcozoan dorsal pellicles, they poss- nads (heterokont chromists), sessile ciliates (e.g. Carchesium,
ibly arose one node earlier. Determining intracellular Zoothamnion); some algal, e.g. chrysophyte Dinobryon. Mucila-

Phil. Trans. R. Soc. B 372: 20150476

distribution and functions in early Sulcozoa would clarify ginous multicellular branching structures are formed by
the integrin adhesion system’s original functions; as genomes Rhipidodendron (cercozoan chromists) or Phalansterium (uniciliate
are known only from very simplified and derived Amoeboza Amoebozoa). As no branching protists evolved a multicellular
lacking integrins, they are also needed for early diverging tissue, similar ‘colonial’ choanoflagellates are probably not
Amoebozoa with more complex extracellular coats/thecae directly relevant to animal origins. Nonetheless, they show that
[18] that might involve integrins. Though lacking typical various linked flagellates can still feed in the same way as
integrins, Dictyostelium has a b-integrin-like adhesion protein when unicellular, and their frequency suggests that branching
[19] and its multicellular prefruiting ’slug’ evolved ahaerens- stalks advantageously enable them to sweep prey from a much
like junctions involving preexisting actin-filament-binding larger water volume than can one sessile cell. Filtering more
catenins [20] (convergently with independently evolved water by a different sessile body form is, I argue, the selective
animal adhaerens junctions) but unlike sponges and other advantage that made sponges.
animals could not recruit cadherins as they only evolved More rarely, choanoflagellate multicells arise by linking
later (with filidigits in ancestral holozoa) [12]. adjacent cells by their collar microvilli as in Proterospongia
On present evidence, excavates and Sulcozoa, successive choanojuncta, but I doubt this had a potential to yield a
ancestors of Choanozoa, never evolved multicellularity, nor sponge. Sponge collars also join laterally often by a second
did any Choanozoa except choanoflagellates whose unique mucus mesh to achieve 100% removal of suspended bacteria
cell structure and feeding mode preadapt them for evolving [22], showing intercellular cooperation efficacy.
multicellularity. Therefore, discovery in non-choanoflagellate The loricate Diaphanoeca sphaerica, where cells often clump
Choanozoa and Sulcozoa of integrins and of cadherins, in hollow balls with cilia pointing inwards [23], exemplifies
and synaptic proteins and other neural channel proteins in a third multicell type incapable of progressing to a tissue.
choanoflagellates and filosporidia [15,21], does not explain Comparing this with a sponge choanocyte chamber [24] was
how animals originated. It tells us (unsurprisingly) that pre- misleadingly superficial as Diaphanoeca, like other loricates
existing proteins were recruited for the job and diversified (Acanthoecida), are tiny cells suspended within a much larger
by gene duplication and divergence (standard for any sub- lorica of siliceous strips porous to water currents carrying
stantial innovation) but not why these protozoa failed to prey. Aggregating porous loricas by connecting longitudinal
become animals and only one lineage did. We must identify strips allows colonial feeding despite cilia pointing inwards,
selective forces that make it impossible for most protists to as the collar outer surface that traps food still faces outwards.
evolve a body with a gut and explain why only one lineage Water and bacteria can pass through the lorica mesh or wide
ever did. I contend that it was not the presence of potential interlorica spaces, so feeding mode is unchanged compared
glue molecules, but the rare ability of choanoflagellate cells with unicells; cell bodies are not in contact so could not
to stick together yet still feed as before that made stem choa- evolve into an epithelium to make a sponge. Acanthoecida
noflagellates our ancestors. Inability to do this would are necessarily an evolutionary dead end.
strongly select against similar aggregative mutations in Non-loricates (Craspedida) never aggregate with cilium
other groups. facing inwards like sponges as that would suicidally stop
collar-based feeding. Sphaeroeca is a multicellular planktonic
craspedid whose colonies are hollow balls with a surface
3. Choanoflagellate and flagellate cell monolayer, associated by cell bodies not collars, analo-
gous to the alga Volvox that Hardy [25] invoked as a
multicellularity potential animal ancestor because of its simple feeding. The
In choanoflagellate colonies, every cell can feed. To become a craspedid Salpingoeca rosetta reversibly makes little multicel-
sponge, the majority must abandon feeding as collar cells, lular balls, a capacity influenced by bacteria [26]. Numerous
lowering feeding potential dramatically. A sponge could other flagellates, e.g. chrysophyte chromists, evolved similar
evolve only if a body were made where reduction in feeding free-swimming multicell balls. These would be incapable of
capacity caused by a lower ratio of feeding to non-feeding progressing towards a multilayered Haeckelian gastraea,
cells was more than compensated by an indirect increase in because gastrulation-like internalizing cells would prevent
feeding or survival efficiency. For understanding animal their feeding, without immediate benefit, and thus be
origins, the key problem is not how cells evolved a capacity strongly disadvantageous. However, by settling on stable
to stick together (trivial)—or even why—but defining the surfaces as sessile filterers, they would encounter new selec-
selective forces that promoted the fundamental differen- tive forces favouring cell differentiation, enabling animal
tiation between sponge feeding cells (choanocytes) and origin. Sponges evolved thus from a craspedid-like stem
non-feeding cells and between cells that stick together as choanoflagellate.
 5
(i) (v) (vi)

rstb.royalsocietypublishing.org
(iv)
B mesohyl

(ii)
(iii)
collar choanocytes
pinacocytes
choanocytes
(vii) zygote sperm

cleavage +

Phil. Trans. R. Soc. B 372: 20150476

egg mesenchyme

settling/maturation
pelagic dispersive larva epithelium
presponge life cycle sessile benthic adult
Figure 2. Evolution of an archetypal animal, a presponge (vii), from a stem choanoflagellate (i – ii,v) prior to integrin loss by crown choanoflagellates. Choano-
flagellates feed by catching bacteria (B) drawn by ciliary water currents (i, arrows) to their collar filters; the cell body phagocytoses them (ii). Extant craspedid
choanoflagellates may be unicells (i,ii) or daughter cells may stick together by branched stalks (iii) or collar microvilli (iv) to make sessile multicells or via cell bodies
to make planktonic swimming balls of cells (v). The first animal could simply have evolved (horizontal black arrow) by such a ball of cells joined laterally by
cadherins settling onto a rockface (cross-hatched), differentiating non-ciliated pinacocytes for attachment and for support secreting extracellular mesohyl (turquoise)
by both cell types and attached to them via pre-existing integrins (vi). This simplest presponge presumbly budded off ciliated swimming balls for dispersal (blue
arrow), and probably had to evolve nutrient transfer from choanocytes to pinacocytes. (vii) Competition for filtering larger water volumes led to larger, stronger,
three-layer (prototriploblastic) feeding laminas with mesenchyme cells specializing in ECM secretion sandwiched between choanocyte epithelia. Larger laminas led to
divergent selection for large eggs capable of rapid cleavage and more numerous smaller sperm, both originally differentiated from choanocytes (rightmost blue
arrows). As size increased, the pluripotent nonciliated mesenchyme cells differentiated into proliferative stem cells (archaeocytes: thenceforth the usual precursors of
eggs, choanocytes continuing to generate sperm) and terminally differentiated cells (lophocytes) secreting collagen fibres to increase mechanical strength.

The primary dichotomy between uniciliate choanocyte and

4. Evolving a triploblastic presponge non-ciliate pinacocyte is also mirrored by that between sperm
Willmer emphasized the basic dichotomy between ciliated epi- and egg. Therefore, part of the same gene switches needed for
thelial and non-ciliated, amoeboid, connective tissue cells as somatic differentiation could also be used to differentiate
fundamental to animal development [27]. Figure 2 summar- gametes. Once a three-layered structure with just two somatic
izes a potential pathway by which a stem choanoflagellate cell types evolved, presponges could become quite large
lineage, initially a standard swimming ball of choanocytes, (compared with choanoflagellate unicells); selection for rapid
could transform into a sessile precursor of sponges by evolving establishment of a large embryo would strongly favour
comparable somatic cell differentiation to anchor itself to a oogamy (large egg and numerous small sperm) by modifying
rock. The new cell type was a basal non-ciliate anchoring cell choanocytes, presumably hermaphrodite. The animal bauplan
that secreted ECM—effectively a basal pinacocyte. An ECM was in place once a selective force for ever-larger filtering struc-
of mucopolysaccharide and collagen would form a supportive tures built from two dissimilar cell types existed: two germ
mesohyl skeleton between two monolayer sheets of choano- line and two soma cell types. Accidental fragments could also
cytes—the ancestral choanoderm. The selective advantage of reproduce vegetatively as choanocytes retained pluripotency
this novel three-layer structure would be filtering food from a [28]. There was no necessary sacrifice of reproductive potential
much larger volume of water, just as branching colonial choa- as in Dictyostelium dead stalk cells.
noflagellates do. ECM support would allow a much larger Another selective advantage of evolving mesenchyme and
structure that could overtop simple branched choanoflagellates massive tissues perhaps gave extra impetus to early animal
with choanocytes only. This could have increased food caught evolution. Mucilage easily harbours bacterial symbionts poten-
by choanocytes more than enough to compensate a presponge tially able to provide enough extra food to repay a presponge
for loss of filter-feeding capacity by basal pinacocytes and ECM several times over the trophic and reproductive costs of non-
secretion costs. If so, selection for taller, wider multicellular feeding cells. Cultivating cyanobacteria in ECM mucilage
filters processing larger volumes of seawater would immedia- would make the photophagotrophic consortium an extremely
tely unavoidably ensue. Flow hydrodynamics for maximizing effective competitor with merely branched choanocyte-only
catch and architectural principles maximizing support and colonial choanoflagellates. Lichen fungi can survive solely by
filter area would impose novel selective forces yielding similar cultivating cyanobacteria; a presponge could be even better
structures to bivalve mollusc gills. Pinacocytes would develop off, being also a phagotroph able to grow far faster than a
contractile actomyosin and surface adhesion analogously to an lichen in bacteria-rich water. Great Barrier Reef sponges
amoeba to spread flattened extensions and cell contacts over 1–2 m high are often red through being packed with cyanobac-
the holdfast portion of the sessile lamina with least cost. teria whose biomass is greater than that of the sponge cells.
They retained a capacity for phagocytosis, thus providing a Lake Baikal giant freshwater sponge tissues cultivate green
primitive immune system by digesting potentially invasive algae. Both habitats are oligotrophic, making internal algae
bacteria for which mesohyl was a nice habitat and food.