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Close-up of yellow tube sponge Aplysina insularis
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Stephen C. Meyer Philosopher of Science
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The Myth of Precambrian Sponges

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Sponges are sessile marine invertebrates that are considered to be the most basal and most “primitive” branch of the multicellular animals (Metazoa). Therefore, evolutionists would expect to find such sponges as the earliest animals in the fossil record. Also, immunological evidence (Wilkinson 1984) and especially molecular clock data placed the origin of sponges long before the Cambrian and even before the Ediacaran era (Peterson et al. 2004Sperling et al. 2007Sperling et al. 2010Erwin et al. 2011Cunningham et al. 2017a). Consequently, the alleged discovery of sponge-like fossils from layers prior to the Cambrian explosion, which gave rise to all the more complex animal phyla, was welcomed by evolutionary biologists as clear confirmation of Darwin’s theory. When the branching order of reconstructed phylogenetic relationships agrees well with the stratigraphic order of appearance in the fossil record, evolutionary biologists speak of so-called stratigraphic fit. The existence of Precambrian fossil sponges was accepted as established fact in many textbooks and academic articles (e.g., Debrenne & Reitner 2001Reitner 2005Budd 2008Müller et al. 2009). A perfect example is Carrera & Botting (2008), who confidently stated that “it is fairly clear that sponges possess a long record back to the Proterozoic, represented in the Ediacara fauna.” The evidence seemed convincing enough that even most critics of Darwinian evolution acknowledged the existence of sponges prior to the Cambrian explosion (e.g., Meyer 2013Evolution News 2016).

I will show that this concession was premature and much too generous. Let’s see how good or rather how embarrassingly bad the case for Precambrian sponges really is and have an exhaustive look at all the potential candidates in alphabetical order.

Ausia, Hahn & Plug, 1985

Ausia fenestrata is known from a few specimens from the Ediacaran of Namibia and Russia. These about 5 cm small fossils look like a triangular bag with numerous ovate depressions or openings in it. It has been considered to be a cnidarian sea pen by its original describers (Hahn & Plug 1985), or a halkieriid stem mollusk (Dzik 2011), or a chordate tunicate (ascidian) (Fedonkin et al. 2007, 2008, 2012), or a sponge-like animal related to the Cambrian Archaeocyatha or even as a true sponge (Fedonkin 1996, McMenamin 1998: 38-39). This shows that the preserved characters are totally insufficient for any definite placement. Unsurprisingly, Antcliffe et al. (2014) considered Ausia as failing to meet the diagnostic criteria of sponges, and Muscente (2015) commented that “such interpretations of aspiculate organisms are inherently subjective, and the affinities of the fossils remain disputed.” Interestingly, another genus and species that has been attributed to the same family Ausiidae, Burykhia hunti from the Ediacaran of Russia, was considered by its describers (Fedonkin et al. 2012) as a possible tunicate, but no possible relation to sponges was even mentioned.

Coronacollina, Clites et al., 2012

Coronacollina acula is a very strange Ediacaran organism described by Clites et al. (2012) from Australia. The fossil is a triradial cone-like mound with four 37 cm long radial spicules, which has been considered as resembling the Cambrian demosponge genus ChoiaCoronacollina has been called the oldest organism with a skeleton and in this way considered as a precursor to the Cambrian explosion (UC Riverside 2012). Serezhnikova (2014) accepted the similarity with Choia and stated that “records of Coronacollina supported the Precambrian origin of sponges and their ability for biomineralization in the Late Proterozoic.” In their revision of Ediacaran tri-radial body plans, Hall et al. (2018) excluded Coronacollina from the otherwise monophyletic group Trilobozoa or Triradialomorpha, but did not propose any other relationships. Muscente et al. (2015) refuted any structural similarity of the spicules with those of sponges. Cunningham et al. (2017a)observed that there is not even a definite association with the putative spicules and concluded: “As such, neither Palaeophragmodictya nor Coronacollina are considered to reflect poriferans, or even metazoans, on the basis of current evidence.” None of the recent exhaustive revisions of putative Precambrian sponges still considered Coronacollina worthy of discussion (Antcliffe et al. 2014Botting & Muir 2017).

Cucullus, Steiner, 1994 (= Doushantuospongia, Li in Ding et al., 1996)

Cucullus fraudulentus is a bag-like or tube-like fossil and represents the most abundant and largest organism from the Neoproterozoic Miaohe biota of the Doushatuo Formation in China. It was described by Steiner (1994: 125) as a putative cyanobacterial colony or mega-alga of uncertain affinity. Chen et al. (1994) suggested that their new species, Sinospongia hubeiensis, later been synonymized with Cucullus fraudulentus, is a poriferan. Therefore, Li et al. (1996) and Hu (1997) attributed Cucullus to demosponges, while Steiner & Reitner (2001) still considered them as microbial colonies. Due to poor preservation, its morphology and affinities remained unclear. Thus Xiao et al. (2002) still could not decide if it was a sponge or a siphonous green alga. Finks et al. (2004) rejected an affinity of Cucullus with sponges in their revised edition of the prestigious Treatise on Invertebrate PaleontologyXiao & Dong (2006) considered Cucullus to be a protoarenicolid, which these authors do not identify as an animal but as dasycladacean algae. When new and better preserved material became available from the Doushantuo Formation in China, Wang & Wang (2011)“resurrected” an affinity to demosponges based on the observation of assumed organic walls formed by non-mineralized spongin fibers. Antcliffe et al. (2014) disagreed and concluded: 

There is however, no unequivocal morphological evidence to support assignment to the Porifera. … Furthermore, no spicules have been described in association with Cucullus. We conclude that assignment to the Porifera is highly speculative and that Cucullus fails the diagnosis test as the possibility that the specimens are microbialites seems to us much more likely.

Let’s repeat this in plain English: Cucullus is not a sponge but rather of microbial origin. Serezhnikova (2014)also found that “the interpretation of Cucullus and Sinospongia as sponges and their comparison with Vaveliksia are difficult to support.” Muscente (2015) basically agreed with that skeptical position.

Eocyathispongia, Yin et al., 2015

Eocyathispongia qiania was described by Yin et al. (2015) after a single specimen from the Early Ediacaran Doushantuo Formation in China, as a sponge grade fossil with cellular resolution. Of course, it was immediately celebrated in the popular media as the discovery of the oldest sponge (Yirka 2015). Cunningham et al. (2017a) remarked: 

It represents the most plausible report of a sponge from the Precambrian. However, more analyses and specimens are needed to test this hypothesis. In particular, high-resolution tomographic analysis of the walls of the specimen could reveal whether pores are present, and therefore whether Eocyathispongia could have functioned as a sponge.

Cunningham et al. (2017b) commented, “Eocyathispongia is considered to be one of the strongest candidates for a Precambrian sponge. However, although it could be a sponge, it has no convincing sponge apomorphies such as pores or spicules, just a generalized sponge gestalt.” Bottjer et al. (2019) rather suggested that “although it contains no characters that are exclusive to crown group sponges …, if not a stem group sponge it could be an extinct organism between the last common ancestor of metazoans and the last common ancestor of living sponges.” However, Botting & Muir (2018) in their review of early sponge evolution mention a severe problem for the sponge hypothesis:

Eocyathispongia is an extremely interesting fossil, but several features are not easily compatible with sponge biology. The morphology is problematic for a sponge, especially a very small one, as it appears to actively minimise the available surface area for incurrent pores, on which it would depend for feeding.

In my view this tiny fossil, which is only about a single millimeter in size, represents just another vase-shaped encysting protist from the Doushantuo layers (Li et al. 2008).

Namapoikia, Wood et al., 2002

Namapoikia rietoogensis is a calcified reef-building organism of up to one meter size, discovered in the Ediacaran Nama group of southern Namibia. In the original description its affinities have been considered to be with either cnidarians or sponges (Wood et al. 2002). Zhuralev et al. (2012) agreed that Namapoikia is “only of a cnidarian or poriferan grade of organisation” Antcliffe et al. (2014) clarified in their large revision of alleged Precambrian sponges that “diagnostic poriferan morphological characteristics are lacking” and that “such a structure could arrive from the calcification of microbial colonies.” Wood & Curtis (2015) still considered it to be “of uncertain affinity,” though resembling “chaetetid sponges or simple colonial cnidarians.” Cunningham et al. (2017a) said that Namapoikia “possesses no characters diagnostic of any particular eukaryotic group.” Wood & Penny (2018) finally placed its “probable affinity within total-group poriferans,” mainly based on similarities in growth pattern and (inferred!) biomineralization. They concluded that “such an interpretation confirms the presence of poriferans, with calcareous skeletons, in the terminal Ediacaran.” However, Tang et al. (2019)remained unconvinced and offered the qualification that “Namapoikia has been interpreted as an encrusting poriferan, but more work is needed to confirm that it is a calcified encrusting sponge rather than a microbial structure,” which strongly suggests that Namapoikia still fails to meet the diagnostic criteria required by Antcliffe et al. (2014).

Curiously, Wikipedia claims that Zhuralev et al. (2015) instead proposed an affinity with lophophorate bilaterian animals or their stem group. However, this is total nonsense and factually incorrect as this paper does not even remotely make such a claim. This example shows how utterly unreliable Wikipedia is as source for scientifically accurate information.

Otavia, Brain et al., 2012

Otavia antiqua is an irregular spheroid microfossil found in 760-550 million year old layers in Namibia, of which some predate the Ediacaran and even the assumed neoproterozoic snowball earth period called Cryogenian. It was described as a sponge-like fossil by Brain et al. (2012), and again celebrated by the press as “Namibia sponge fossils are world’s first animals” (AFP 2012Gess 2012). Antcliffe et al. (2014) rejected its identification as a sponge and remarked that “alternative hypotheses have not been sufficiently explored and/or have been rejected without sufficient reason.” They even raised severe doubts that Otavia is a genuine fossil of biogenic origin at all, or maybe of microbial origin rather than an animal. Antcliffe et al. concluded that “no features are presented that are diagnostic of sponges. We interpret these objects as calciphosphate grains that have been pitted by sediment reworking.” This sounds much less spectacular and thus did not hit the news headlines. Even the more recent textbook by Jain (2016: page 7)Fundamentals of Invertebrate Palaeontology, ignored the refutation and still teaches gullible students the obsolete story that Otavia is a calcareous sponge and the oldest animal.

Palaeophragmodictya, Gehling & Rigby, 1996

The type species Palaeophragmodictya reticulata was described by Gehling & Rigby (1996) as “long expected sponges from the Neoproterozoic Ediacara fauna of South Australia.” The very title of their work shows how eagerly Darwinists longed for an empirical confirmation of their theoretical expectations. The sponge affinity was accepted by most other authors (e.g., Seilacher 1999, Finks et al. 2004, McCall 2006). About ten years later a second species P. spinosa was discovered by Serezhnikova (2007) in the White Sea region of Russia and demolished these expectations again. She did not identify Palaeophragmodictya as sponge at all, but recognized that it is just a holdfast of an unknown sessile organism (again affirmed by Serezhnikova 2014) that might have been a cnidarian. Nevertheless, the story was apparently too nice to be spoiled by stupid facts, so that the sponge interpretation was retained in many recent articles (e.g., Maloof et al. 2010Brain et al. 2012Dohrmann et al. 2013Stearley 2013). But the truth could not be ignored forever. Antcliffe et al. (2014) 

agree with Serezhnikova (2007a) that some of the material is likely to represent the holdfast of other Ediacaran-age organisms, we question whether these specimens represent sponges of any kind. … No compelling arguments are presented that Palaeophragmodictya should be considered separately from other disc-like structures of Ediacaran age.

Furthermore, Antcliffe et al. present extensive evidence and arguments that these fossils indeed represent holdfasts of the typical Ediacaran frond-like organisms. Cunningham et al. (2017a) acknowledged that Palaeophragmodictya “is perhaps the most widely recognized candidate for a sponge within the Ediacaran macrobiota” but “the taxon may be more readily interpreted as decayed attachment discs from an organism of uncertain affinity.” Likewise, Botting & Muir (2018) remarked that this fossil taxon was a misidentified holdfast of an Ediacaran frond and “bears little resemblance to extant or early fossil sponges.” Finally, the only potential sponge character in Palaeophragmodictya and some other alleged Precambrian sponges, the presence of apparent skeletal nets, has been discredited by a new study by Luzhnaya & Ivantsov (2019), who documented such structures in the characteristic Ediacaran fronds, which were never considered as sponges and certainly are not because they have no openings for water circulation.

Rugoconites, Glaessner & Wade, 1966

Rugoconites enigmaticus is another circular organism from the Ediacaran biota of Australia described by Glaessner & Wade (1966)Gehling & Rigby (1996) mentioned that Rugoconites has some characters in common with Palaeophragmodictya, but recognized that “the lack of evidence of a spicular framework prevents a clear assignment of Rugoconites to the poriferans.” Some authors considered them to be jellyfish (Cloud & Gessner 1982, Sepkoski 2002), while others considered them as possible sponges (Darroch et al. 2018). Nevertheless, the consensus among specialists today generally tends to classify Rugoconites as member of the Ediacaran monophylum Trilobozoa or Triradialomorpha with a tri-radial body plan (Ivantsov & Fedonkin 2002Xiao & Laflamme 2009Hall et al. 2018).

Sinospongia, Chen, 1992 (= Niuganmafeia, Chen, 1991)

The genus Sinospongia is known from two species, S. typica and S. chenjunyuani, from the Miaohe biota of the Doushantuo Formation in China (a third species Sinospongia hubeiensis was synonymized with Cucullus fraudulentus, see above). It was considered as mega-algae by Steiner (1994), as microbial colonies by Steiner & Reitner (2001), and as Protoarenicolidae by Xiao & Dong (2006), which these authors do not identify as animals but as dasycladacean algae with holdfasts. Xiao et al. (2002) suggested three possible interpretations: “sponge, cnidarian, or siphonous green alga,” because “no convincing sponge spicules have been found in association with these fossils, and alternative algal interpretations are possible” and “alternative interpretations, such as a cnidarian-grade organism or a siphonous green alga (especially of the order Dasycladales), should be entertained.” As already mentioned above, Serezhnikova (2014) concluded, “Therefore, the interpretation of Cucullus and Sinospongia as sponges and their comparison with Vaveliksia are difficult to support.” The more recent revisions of alleged Precambrian sponges by Antcliffe et al. (2014)Muscente et al. (2015), and Botting & Muir (2018) did not even bother to further discuss Sinospongia as a putative sponge.

Thectardis, Clapham et al., 2004

Thectardis avalonensis is a conical organism of about 9 cm length and 3 cm diameter, known from numerous specimens from the Ediacaran Mistaken Point locality in Newfoundland (Clapham et al. 2004). It is believed to have been attached with its pointed end to the bacterial mat sea floor as a suspension feeding “mat sticker,” but it lacks any visible holdfast structures. In the original description no affinity to sponges or any other group was suggested. Sperling et al. (2011) proposed a most likely attribution to sponges, only based on the habitat beneath the photic zone (which precludes photosynthesis) and because its body plan would be consistent with the hydrodynamics of the sponge water-canal system. This is a very weak level of argumentation for the far-reaching conclusion that “the recognition of sponges in the Mistaken Point biota provides some of the earliest body fossil evidence for this group, which must have ranged through the Ediacaran based on biomarkers, molecular clocks, and their position on the metazoan tree of life, in spite of their sparse macroscopic fossil record.” Consequently, Antcliffe et al. (2014) correctly pointed out that such poor indirect criteria could at best exclude certain affinities, but not confirm any. They concluded that “the interpetation of Thectardis avalonensisas a fossil sponge is therefore highly problematic.” They even elaborated that the fossil might rather be just the misinterpreted remains of the decay process of an unknown larger organism. Muscente et al. (2015)concluded “Thectardis from Newfoundland … have been interpreted as sponges based on inferences regarding soft tissue anatomy, biomechanics, and taphonomy. However, such interpretations of aspiculate organisms are inherently subjective, and the affinities of the fossils remain disputed.” Christian paleontologist Ralph Stearley (2013) still affirmed that “the conical fossil Thectardis from the Avalon assemblage is also probably a sponge,” but this was just in a book review. Serezhnikova (2014) agreed that the general body plan of Thectardis is similar to the sponge-like archaeocyaths, but offered the qualification that “the affinity of the Precambrian taxa to Porifera is limited by a lack of data on their possible filtering structures.” Liu & Conliffe (2015) remarked that “until more informative specimens are found, this taxon will likely remain of uncertain biological affinity.” Botting & Muir (2018) remarked that “although it is theoretically possible that some alleged Ediacaran Biota sponges (e.g., the featureless triangle-shaped object Thectardis Clapham et al., 2004; Sperling et al., 2011) are in fact genuine, there is no evidence to that effect.” Let that sink in: NO EVIDENCE! There seems to be a pattern here: desperate attempts to fulfill Darwinian expectations in the absence of any convincing evidence.

Vaveliksia, Fedonkin, 1983

A 3-10 cm large sac-shaped organism of radial symmetry with a perforated body wall and an attachment disc, the type species Vaveliksia velikanovi was described by Fedonkin (1983) from the Ediacaran of Ukraine. A second species, Vaveliksia vana, was described from the Vendian of the White Sea region in Russia by Ivantsov et al. (2004). They concluded: “From the above observations and assumptions, one may propose that Vaveliksia has the same level of organization as archaeocyaths or sponges.” McCall (2006) suggested “possible coelenterate and hydrozoa affinities” instead. Serezhnikova (2014), who co-described the second species of Vaveliksia, listed it as “problematic lower metazoans” with a “level of organization of parazoa (?).” Antcliffe et al. (2014) mentioned the possibility that Vaveliksia could rather be an agglutinated amoebozoan, thus a colonial protist, and concluded that “Vaveliksia vana lacks any definitive poriferan characteristics and fails the characters and diagnosis criteria.

Malloof et al. (2010) reported unnamed ellipsoidal fossils as sponge-grade metazoans from the Neoproterozoic of Australia. Paleontologist Chris Nedin (2010) was not convinced and commented on his respected Ediacaran blog that “Proterozoic Sponges Claim Doesn’t Hold Water.” Well, blogs don’t count in science, but Antcliffe et al. (2014) came to the same conclusion and remarked that 

no characteristics that are distinctive for sponges are presented by the authors. … These fossils have no morphology that is diagnostic of sponges and should in our view be more readily ascribed to the calcimicrobes that abound at these localities.

Finally, Wallace et al. (2014) described unnamed chambered structures from Cryogenian reefs, for which they pose the question of whether they could be the oldest sponge-grade organisms. However, they concluded: 

The closest morphological analogues for the structures are: a) some types of reef-dwelling sponges; and b) some complex microbialites from Archean and Paleoproterozoic carbonates. The structures lack spicules and ostia found in sponges, ruling out a true Poriferan origin. However, it is plausible that they are proto-sponges, sponge-grade organisms, or complex microbial precursors to sponge-grade organisms. Whatever their affinity, we suggest these structures record a significant evolutionary event on the path towards organic complexity.

It is very clear from their conclusions that these fossils also fail all three criteria of Antecliffe et al. (2014) for an identification as sponges and thus have to be considered as problematic fossils of uncertain affinity, but more likely of microbial origin.

That was the last candidate. But what about more indirect evidence from alleged sponge embryos, sponge needles, and sponge biomarkers? Here comes the story of their demise.

The Sponge Embryos That Aren’t

Phosphatized microfossils from the 609-million-year old Doushantuo Formation in China, which were originally interpreted as colonial green algae, were later re-interpreted as alleged animals (Xiao et al. 1998) and especially sponge embryos and sponge larvae (Li et al. 1998a1998b, Chen et al. 1999, Chen et al. 2000Xiao & Knoll 2000Xiao et al. 2000, Chien et al. 2001, Chen 2012). This interpretation was immediately disputed by other Chinese researchers (Zhang et al. 1998, Xue et al. 1999), who alternatively identified these fossils as collapsed acritarch protists, thus not sponges at all but unicellular organisms. Even though Cao & Zhu (2001) disagreed with this, they acknowledged that “whether or not they are larvae of sponges, it is not determined as yet.” Hagadorn et al. (2006) showed that “the absence of epithelialization is consistent only with a stem-metazoan affinity for Doushantuo embryos,” thus rejecting any sponge affinities.

Bailey et al. (2007) suggested that the alleged embryos are giant actually vacuolate sulphur bacteria close to the recent genus Thiomargarita. However, this was convincingly rejected by Donoghue (2007)Xiao et al. (2007)Yin et al. (2007)Cunningham et al. (2012), and Igisu et al. (2014), even though these authors did not all agree on the embryo nature of the fossils.

After careful synchrotron-tomographic studies, which included a team of the most eminent paleontologists like Philip Donoghue and Stefan Bengtson, Huldtgren et al. (2011) concluded that the alleged embryos are neither metazoans nor embryos but just encysting eukaryotic protists (also see Butterfield 2011, who commented that “wherever the Doushantuo fossils eventually end up, it will clearly not be within ‘crown-group’ Metazoa,” and Kaplan 2011, whose comments are titled, “Enigmatic fossils are neither animals nor bacteria”). Xiao et al. (2012) readily disagreed in a response to this article, but were again refuted by Huldtgren et al. (2012). But then, in a study by Chen et al. (2014), co-authored by Shuhai Xiao, the authors came to the same result that the “available evidence also indicates that the Doushantuo fossils are unlikely crown-group animals.”

Even the alleged and much celebrated oldest bilaterian animal Vernanimalcula from the Doushantuo Formation was debunked by Bengtson et al. (2012) in an article titled “A merciful death for the earliest bilaterian,” in which the authors came to the scathing conclusion that “there is no evidential basis for interpreting Vernanimalcula as an animal.”

Yin et al. (2016) presented new evidence in the form of meroblastic cleavage patterns for the identification of Doushantuo fossils as metazoan embryos, but acknowledged that “their phylogenetic affinity cannot be established.” Just recently, Yin et al. (2019) described gastrulation-like cell division in the fossil Caveasphaera from this locality, which they said foreshadows animal-like embryology. However, it was not considered as a sponge or even a metazoan by these authors. Carl Zimmer (2019) commented in the New York Times that these “balls of cells may be the oldest animal embryos — or something else entirely.

Cunningham et al. (2017b) reviewed all the published evidence from the Wen’an biota of the Doushantuo Formation and concluded that “although the Weng’an Biota includes forms that could be animals, none can currently be assigned to this group with confidence.” If there are not even definite animals, there can be no sponge embryos either.

Independent of the doubtful nature of the Doushantuo “embryos,” recently a multicellular organism was described as a microfossil from the Ediacaran Nyborg Formation in Norway (Agić et al. 2019). It was named Cyathinema digermulense and the authors considered it as sharing “characteristics with extant and fossil groups including red algae and their fossils, demosponge larvae and putative sponge fossils, colonial protists, and nematophytes.” Even though sponge affinities are not ruled out, they have not been demonstrated either.

Sponge Needles in a Haystack

Even if all the body fossils discussed above fail to establish the presence of sponges prior to the Cambrian era, maybe we could at least find their most durable parts as microfossils: sclerotized needles, so-called spicules, that form the skeleton of sponges and are made of silica or calcite.

Indeed, several works described alleged sponge spicules from Precambrian deposits, mainly in China (Dunn 1964, Tang et al. 1978, Ding et al. 1985, Zhao et al. 1988, Allison & Awramik 1989Brasier 1992Steiner et al. 1993Gehling & Rigby 1996Brasier et al. 1997Li et al. 1998aTiwari et al. 2000Xiao et al. 2000Du & Wang 2012, and Du et al. 2015). Even elaborate scenarios for the presumed evolution of sponge skeletons in the Proterozoic have been proposed based on this evidence (Müller et al. 2007).

Steiner et al. (1993) questioned the Doushantuo spicules and Gehling & Rigby (1996) mentioned that “of the many reported spicules from Proterozoic sediments most have proven to be volcanic shards or other inorganic crystals (Pickett, 1983).”

Zhou et al. (1998) considered the spicule-like structures from the Doushantuo Formation described by Li et al. (1998) as nothing but pseudo-fossils, while Zhang et al. (1998) considered them as detached spines of collapsed acritarch protists. Cao & Zhu (2001) disagreed and remarked that based on the observation of extant specimens they “tend to the interpretation of the sponges as monaxial spicules.”

Yin et al. (2001) showed that the alleged sponge spicules from the Doushantuo Formation in China are “indistinguishable from coexistent diagenetic crystals,” thus inorganic artefacts rather than fossil remains. They found that “the evidence for a sponge spicule interpretation of the Doushantuo spicular structures are at best ambiguous at present,” but strangely added the inconsistent disclaimer (likely for political reasons) that “despite our initial questioning the proposed interpretation of Doushantuo spicular structures as demosponge microscleres, we do not deny that sponges spicules do exist in Doushantuo cherts.”

Antcliffe et al. (2014) reviewed all the published evidence for alleged sponge spicules from the Precambrian and dismissed all of them, mostly as abiogenic artifacts (e.g., the once oldest, widely accepted hexactinellid spicules from Mongolia, which were shown to be cruciform arsenopyrite crystals by EDX analysis). They found that “the earliest reliable sponge fossils are hexact spicules from Iran dated to c. 535 Ma.”

Muscente et al. (2015) used the most modern analytical techniques like scanning electron microscopy and synchrotron nanotomography to decisively assess the veracity of assumed Precambrian sponge needles, especially from the Doushantuo Formation. They found that their “new data invalidate the oldest and only Precambrian demosponges with mineralized spicules.” The highlights section in their article says it all:

  • We characterize spicule-like structures hosted in Ediacaran phosphatized fossils.
  • Their compositions, mineralogies and ultrastructures are inconsistent with spicules.
  • They may be microbial strands, axial filaments or carbon-molded acicular crystals.
  • Data invalidate oldest and only Precambrian demosponges with mineralized spicules.
  • The results affirm no unequivocal biomineralizing sponges occur in the Precambrian.

Finally, Botting & Muir (2018) agreed in the most recent review of early sponge evolution that “there are no definite records of Precambrian sponges: isolated hexactine-like spicules may instead be derived from radiolarians.” And Tang et al. (2019) likewise stated that “the oldest spicules are Cambrian in age,” but hypothesized based on a new Cambrian fossil that potential Ediacaran sponges might have lacked biomineralized spicules (also see Tang 2019). So, when alleged Precambrian spicules were found they were naturally considered as evidence for Darwinian evolution, and now since they are consistently lacking, Darwinian evolution is invoked to explain their absence. Whatever the evidence says, Darwinists always claim victory.

The latest claim was made recently by Chang et al. (2019), who reported alleged sponge spicules from the Ediacaran-Cambrian boundary of the Yanjiahe Formation in China. However, in the actual article they only say that “monaxon spicules and spicule-like structures in the lower Yanjiahe Formation might putatively be interpreted as demosponge remains,” thus acknowledging a high degree of uncertainty. They also acknowledged that their interpretation of the monaxon spicules would be incongruent with the evolutionary scenario of Botting & Muir (2018), who suggested a hexactine-based ground plan. In the Yanjiahe Formation non-monaxon spicules only appear just beneath the Lower Cambrian Protohertzina anabarites zone (see their Figure 6).

The fact is this: Despite the multiple premature claims of success, genuine sponge needles from the Precambrian era thus not only have proved to be elusive like the proverbial needles in a haystack but indeed seem to be non-existent.

Fossil Fat Fails

Based on chemical analyses of sediments it has been suggested that fossil steroids (24-isopropylcholestane and 26-methylstigmastane) are lipid biomarkers that provide indirect evidence for the existence of demosponges in the Precambrian (McCaffrey et al. 1994, Moldowan et al. 1994, Love et al. 2009Sperling et al. 2010). Antcliffe (2013) questioned the evidence for sponge biomarkers because 24‐isopropylcholestane is also produced by marine algae or their diagenetic alteration, and claimed that “it seems more likely that these compounds represent algal biochemical evolution at a time when algal burial occurred in great quantity with well-known coeval algal fossils but no sponge fossils.” This was reasonably rejected by Love & Summons (2015), and even very recent studies by Gold et al. (2016a)Gold et al. (2016b)Brocks et al. (2017)Sperling & Stockey (2018), and Zumberge et al. (2018) still considered this so-called sponge biomarker hypothesis to be validated by the most up-to-date evidence. The popular science media reported “Sponges on ancient ocean floors 100 million years before Cambrian period” (UC Riverside 2018). However, Nettersheim et al. (2019)found these putative typical sponge biomarkers to be common among unicellular organisms (Rhizaria) and concluded that “negating these hydrocarbons as sponge biomarkers, our study places the oldest evidence for animals closer to the Cambrian Explosion.” Ooops!

Thorough Revision

In a thorough revision of all twenty potential Precambrian sponge fossils, including most of the taxa discussed above (except Eocyathispongia) as well as all the other unnamed candidates for sponge-grade metazoans, Antcliffe et al. (2014) came to the conclusion that “that no Precambrian fossil candidate yet satisfies all three of these criteria to be a reliable sponge fossil.” The authors suggested that molecular clocks should be recalibrated accordingly to a Lower Cambrian age of sponges and metazoans (such a recalibration is just a euphemism for using a fudge factor to get the desired outcome). Muscente et al. (2015) confirmed this result that “no unequivocal sponge fossils occur below the Ediacaran–Cambrian boundary.” Another recent study about the early history of sponges (Botting & Muir 2018) also found that “there are no definite records of Precambrian sponges.” Just a year later, Nettersheim et al. (2019) refuted the only remaining biomarker evidence (see above), so that all the empirical evidence for Precambrian sponges had finally evaporated. Maybe new fossil finds will come up with something better, but considering the track record so far, we probably don’t have to hold our breath.

Zero Evidence

We can safely conclude that, contrary to common misconception, there exists zero compelling evidence for the existence of any genuine Precambrian fossil sponges. Unambiguous sponges only appear in the Cambrian explosion together with the other animal phyla. The oldest reliable fossil record for sponges is represented by siliceous spicules from the basal Cambrian of Iran (Antcliffe et al. 2014) and China (Chang et al. 2017Chang et al. 2019) (the latter are slightly older, just below the Protohertzina anabarites zone). With an age of about 535 million years these are even two million years younger than the oldest trace fossil evidence for crown-group arthropods like trilobites recently dated to 537 million years ago (Daley et al. 2018). The first complete body fossils of sponges only appear even somewhat later in the Lowermost Cambrian (Steiner et al. 1993Yuan et al. 2002Flügel & Singh 2003). Only after 525 million years ago sponges became more common (Antcliffe et al. 2014), and crown group demosponges do not appear before 515 million years (Botting et al. 2015).

Having sponges appear after arthropods is not only a very bad stratigraphic fit, but indeed rather a temporal paradox like in many other cases (e.g., the oldest tetrapods and birds). If a good stratigraphic fit is considered to be confirmation for Darwin’s theory, then a poor stratigraphic fit (as well as a mismatch between molecular clocks and fossil record) has to count as conflicting evidence, even if evolutionists can fudge bold ad hoc explanations (like 200-million-year-long ghost lineages, Sperling et al. 2010) to accommodate and explain away such unwelcome data.

Design Theorist Vindicated

Interestingly, the most comprehensive revision of the early fossil record of sponges, by Antcliffe et al. (2014), came to a conclusion very similar to that of intelligent design proponent Stephen C. Meyer in his seminal book Darwin’s Doubt. Here is what they said in their conclusion: “The Cambrian explosion was an evolutionary event of great magnitude and closely connected to the origin of animals.” Science deniers like Jerry Coyne, Donald Prothero, and Nick Matzke, who downplayed the Cambrian explosion in their polemical reviews of Meyer’s book, should read the actual specialists to learn about the significance and abruptness of the origin of animal body plans in the Cambrian explosion, including the body plan of the most primitive animals, sponges.

References:

  • Agić H, Högström AES, Moczydłowska M, Jensen S, Palacios T, Meinhold G, Ebbestad JOR, Taylor WL & Høyberget M 2019. Organically-preserved multicellular eukaryote from the early Ediacaran Nyborg Formation, Arctic Norway. Scientific Reports 9, 14659, 1–12. DOI: 10.1038/s41598-019-50650-x.
  • Allison CW & Awramik SM 1989. Organic-walled microfossils from earliest Cambrian or latest proterozoic Tindir Group rocks, Northwest Canada. Precambrian Research 43(4), 253–294. DOI: 10.1016/0301-9268(89)90060-0.
  • AFP 2012. Namibia sponge fossils are world’s first animals. Phys.org Febr. 6, 2012.
  • Antcliffe JB 2013. Questioning the evidence of organic compounds called sponge biomarkers. Palaeontology 56(5), 917–925. DOI: 10.1111/pala.12030.
  • Antcliffe JB, Callow RHT, Brasier MD 2014. Giving the early fossil record of sponges a squeeze. Biological Reviews 89(4), 972–1004. DOI: 10.1111/brv.12090.
  • Bailey JV, Joye SB, Kalanetra KM, Flood BE & Corsetti FA 2007. Evidence of giant sulfur bacteria in Neoproterozoic phosphorites. Nature 445, 198–201. DOI: 10.1038/nature05457.
  • Bengtson S, Cunningham JA, Yin C & Donoghue PCJ 2012. A merciful death for the “earliest bilaterian,” Vernanimalcula. Evolution & Development 14(5), 421–427. DOI: 10.1111/j.1525-142X.2012.00562.x.
  • Botting JP, Cárdenas P & Peel JS 2015. A crown-group demosponge from the early Cambrian Sirius Passet biota, North Greenland. Palaeontology 58(1), 35–43. DOI: 10.1111/pala.12133.
  • Botting JP & Muir LA 2018. Early sponge evolution: A review and phylogenetic framework. Palaeoworld27(1), 1–29. DOI: 10.1016/j.palwor.2017.07.001.
  • Bottjer DJ, Yin Z, Zhao F & Zhu M 2019. Comparative taphonomy and phylogenetic signal of phosphatized Weng’an and Kuanchuanpu Biotas. Precambrian Research 105408. DOI: 10.1016/j.precamres.2019.105408.
  • Brain CK, Prave AR, Hoffmann KH, Fallick AE, Botha A, Herd DA, Sturrock C, Young I, Condon DJ & Allison SG 2012. The first animals: ca. 760-million-year-old sponge-like fossils from Namibia. South African Journal of Science 108(1/2), 658, 1–8. DOI: 10.4102/sajs.v108i1/2.658.
  • Brasier M 1992. Nutrient-enriched waters and the early skeletal fossil record. Journal of the Geological Society 149, 621–629. DOI: 10.1144/gsjgs.149.4.0621.
  • Brasier M, Green O & Shields G 1997. Ediacarian sponge spicule clusters from southwestern Mongolia and the origins of the Cambrian fauna. Geology 25(4), 303–306. DOI: 10.1130/0091-7613(1997)025<0303:ESSCFS>2.3.CO;2.
  • Brocks JJ, Jarrett AJM, Sirantoine E, Hallmann C, Hoshino Y & Liyanage T 2017. The rise of algae in Cryogenian oceans and the emergence of animals. Nature 548, 578–581. DOI: 10.1038/nature23457.
  • Budd GE 2008. The earliest fossil record of the animals and its significance. Philosophical Transactions of the Royal Society B 363(1496), 1425–1434. DOI: 10.1098/rstb.2007.2232.
  • Butterfield NJ 2011. Terminal Developments in Ediacaran Embryology. Science 334, 1655–1656. DOI: 10.1126/science.1216125.
  • Cao F & Zhu S 2001. New evidences of sponge fossils in the Weng’an Biota. Acta geologica Sinica75(3), 289. [In Chinese] [ResearchGate]
  • Carrera MG & Botting JP 2008. Evolutionary History of Cambrian Spiculate Sponges: Implications for the Cambrian Evolutionary Fauna. Palaios 23(3), 124–138. DOI: 10.2110/palo.2006.p06-089r.
  • Chang S, Feng Q, Clausen S & Zhang L 2017. Sponge spicules from the lower Cambrian in the Yanjiahe Formation, South China: the earliest biomineralizing sponge record. Palaeogeography, Palaeoclimatology, Palaeoecology 474, 36–44. DOI: 10.1016/j.palaeo.2016.06.032.
  • Chang S, Zhang L, Clausen S, Bottjer DJ & Feng Q 2019. The Ediacaran-Cambrian rise of siliceous sponges and development of modern oceanic ecosystems. Precambrian Research 333, 105438, 1–16. DOI: 10.1016/j.precamres.2019.105438.
  • Chen J-Y 2012. Evolutionary Scenario of the Early History of the Animal Kingdom: Evidence from Precambrian (Ediacaran) Weng’an and Early Cambrian Maotianshan Biotas, China. pp. 239-379 in: Talent JA (ed.) Earth and Life. Springer, Dordrecht. DOI: 10.1007/978-90-481-3428-1_10.
  • Chen JY, Li CW, Chien P, Zhou G-Q, Gao F 1999. Wen’an Biota: A Light Casting on the Precambrian World. Paper presented to The Origin of Animal Body Plans and Their Fossil Records, Kunming, China, June 20–26, 1999.
  • Chen J-Y, Oliveri P, Li CW, Zhou G-Q, Gao F, Hagadorn JW, Peterson KJ & Davidson EH 2000. Precambrian animal diversity: Putative phosphatized embryos from the Doushantuo Formation of China. PNAS 97(9), 4457–4462. DOI: 10.1073/pnas.97.9.4457.
  • Chen L, Xiao S, Pang K, Zhou C & Yuan X 2014. Cell differentiation and germ–soma separation in Ediacaran animal embryo-like fossils. Nature 516, 238–241. DOI: 10.1038/nature13766.
  • Chen ME, Xiao ZZ & Yuan XL 1994. A new assemblage of megafossils: Miaohe biota from Upper Sinian Doushantuo Formation, Yangtze Gorges. Acta Palaeontologica Sinica 33, 391–403.
  • Chien P, Chen JY, Li CW, Leung F 2001. SEM Observation of Precambrian Sponge Embryos from Southern China, Revealing Ultrastructures including Yolk Granules, Secretion Granules, Cytoskeleton, and Nuclei. Paper presented to the North American Paleontological Convention, University of California, Berkeley, June 26 – July 1, 2001.
  • Clapham ME, Narbonne GM, Gehling JG, Greentree C & Anderson MM 2004. Thectardis avalonensis: A New Ediacaran Fossil from the Mistaken Point Biota, Newfoundland. Journal of Paleontology 78(6), 1031–1036. DOI: 10.1666/0022-3360(2004)078<1031:TAANEF>2.0.CO;2.
  • Clites EC, Droser ML & Gehling JG 2012. The advent of hard-part structural support among the Ediacara biota: Ediacaran harbinger of a Cambrian mode of body construction. Geology 40(4), 307–310. DOI: 10.1130/G32828.1.
  • Cunningham JA, Thomas C-W, Bengtson S, Marone F, Stampanoni M, Turner FR, Bailey JV, Raff RA, Raff EC & Donoghue PCJ 2012. Experimental taphonomy of giant sulphur bacteria: implications for the interpretation of the embryo-like Ediacaran Doushantuo fossils. Proceedings of the Royal Society B 279, 1857–1864. DOI: 10.1098/rspb.2011.2064.
  • Cunningham JA, Liu AG, Bengtson S, Donoghue PCJ 2017a. The origin of animals: Can molecular clocks and the fossil record be reconciled? BioEssays 39(1), 1–12. DOI: 10.1002/bies.201600120.
  • Cunningham JA, Vargas K, Yin Z, Bengtson S & Donoghue PCJ 2017b. The Weng’an Biota (Doushantuo Formation): an Ediacaran window on soft-bodied and multicellular microorganisms. Journal of the Geological Society 174, 793–802. DOI: 10.1144/jgs2016-142.
  • Daley AC, Antcliffe JB, Drage HB, Pates S 2018. Early fossil record of Euarthropoda and the Cambrian Explosion. PNAS 115(21), 5323–5331. DOI: 10.1073/pnas.1719962115.
  • Darroch SAF, Smith EF, Laflamme M & Erwin DH 2018. Ediacaran Extinction and Cambrian Explosion. Trends in Ecology & Evolution 33(9), 653–663. DOI: 10.1016/j.tree.2018.06.003.
  • Debrenne F & Reitner J 2001. Sponges, Cnidarians, and Ctenophores. pp. 301–325 in: Zhuravlev AY & Riding R (eds) The Ecology of the Cambrian Radiation. Columbia University Press, New York. DOI: 10.23689/fidgeo-2696.
  • Ding Q-X, Xing Y-S & Chen Y-Y 1985. Metazoa and trace fossils. pp. 115–119 in: Zhao Z et al. (eds) Biostratigraphy of the Yangtze Gorge Area. 1: Sinian. Geological Publishing House, Beijing, 143 pp.
  • Ding L, Li Y, Hu X, Xiao Y, Su C & Huang J 1996. Sinian Miaohe Biota. Geological Publishing House, Beijing, 221 pp.
  • Dohrmann M, Vargas S, Janussen D, Collins AG & Worheide G 2013. Molecular paleobiology of early-branching animals: integrating DNA and fossils elucidates the evolutionary history of hexactinellid sponges. Paleobiology 39(1), 95–108. DOI: 10.1666/0094-8373-39.1.95.
  • Donoghue PCJ 2007. Embryonic identity crisis. Nature 445, 155–156. DOI: 10.1038/nature05520.
  • Du W & Wang XL 2012. Hexactinellid sponge spicules in Neoproterozoic dolostone from South China. Paleontological Research 16(3), 199–207. DOI: 10.2517/1342-8144-16.3.199.
  • Du W, Wang XL & Komiya T 2015. Potential Ediacaran sponge gemmules from the Yangtze Gorges area in South China. Gondwana Research 28(3), 1246–1254. DOI: 10.1016/j.gr.2014.08.012.
  • Dunn PR 1964. Triact spicules in proterozoic rocks of the Northern Territory of Australia. Journal of the Geological Society of Australia 11(2), 195–197. DOI: 10.1080/00167616408728571.
  • Dzik J 2011. Possible Ediacaran ancestry of the halkieriids. In: Johnston PA & Johnston KJ (eds) International Conference on the Cambrian Explosion – Proceedings. Palaeontographica Canadiana 31, 205–218. [PDF of conference abstract].
  • Erwin DH, Laflamme M, Tweedt SM, Sperling EA, Pisani D, Peterson KJ 2011. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334, 1091–1097. DOI: 10.1126/science.1206375.
  • Evolution News 2016. New Precambrian Embryos Are Equivocal at Best. Evolution News August 18, 2016.
  • Fedonkin MA 1983. Non-skeletal fauna of Podolia, Dniester River valley. pp. 128–139 in: Velikanov VA, Asseeva EA & Fedonkin MA (eds) The Vendian of the Ukraine. Naukova Dumka, Kiev. [In Russian]
  • Fedonkin MA 1996. Ausia as an ancestor of archeocyathans, and other sponge-like organisms. pp. 90-91 in: Enigmatic Organisms in Phylogeny and Evolution. Abstracts. Paleontological Institute, Russian Academy of Sciences, Moscow. [In Russian]
  • Fedonkin MA, Gehling JG, Grey K Narbonne GM & Vickers-Rich P 2007. The Rise of Animals: Evolution and Diversification of the Kingdom Animalia. Johns Hopkins University Press, 326 pp.
  • Fedonkin MA, Vickers-Rich P, Swalla BJ, Trusler P & Hall M 2008. A Neoproterozoic chordate with possible affinity to the ascidians: New fossil evidence from the Vendian of the White Sea, Russia and its evolutionary and ecological implications”. HPF-07 Rise and Fall of the Ediacaran (Vendian) biota. International Geological Congress – Oslo 2008.
  • Fedonkin MA, Vickers-Rich P, Swalla BJ, Trusler P & Hall M 2012. A New Metazoan from the Vendian of the White Sea, Russia, with Possible Affinities to the Ascidians. Paleontological Journal 46(1), 1–11. DOI: 10.1134/S0031030112010042.
  • Finks RM, Reid REH & Rigby JK 2004. Porifera. pp. 9-173 + 872 in Kaesler RL (ed.) Treatise on Invertebrate Paleontology, Part E (revised), Porifera, 3. The Geological Society of America and University of Kansas, Boulder and Lawrence, Kansas, 902 pp.
  • Flügel E & Singh IB 2003. Stromatoporoid-grade and other sponge fossils from the upper Krol Formation of the Lesser Himalaya (India) Implications for the biotic evolution around the Precambrian-Cambrian boundary interval. Facies 49, 351–372. DOI: 10.1007/s10347-003-0038-6.
  • Gehling JG & Rigby JK 1996. Long Expected Sponges from the Neoproterozoic Ediacara Fauna of South Australia. Journal of Paleontology 70(2), 185–195. DOI: 10.1017/S0022336000023283.
  • Gess RW 2012. The oldest animal fossils. South African Journal of Science 108(1/2), 1064, 1–2. DOI: 10.4102/sajs.v108i1/2.1064.
  • Glaessner ML & Wade M 1966. The late Precambrian fossils from Ediacara, South Australia. Palaeontology 9, 599–628. [BHL]
  • Gold DA, Grabenstatter J, de Mendoza A, Riesgo A, Ruiz-Trillo I & Summons RE 2016a. Sterol and genomic analyses validate the sponge biomarker hypothesis. PNAS 113(10), 2684–2689. DOI: 10.1073/pnas.1512614113.
  • Gold DA, O’Reilly SS, Luo G, Briggs DEG & Summons RE 2016b. Prospects for Sterane Preservation in Sponge Fossils from Museum Collections and the Utility of Sponge Biomarkers for Molecular Clocks. Bulletin of the Peabody Museum of Natural History 57(2), 181–189. DOI: 10.3374/014.057.0208.
  • Hagadorn JW et al. 2006. Cellular and Subcellular Structure of Neoproterozoic Animal Embryos. Science 314(5797), 291–294. DOI: 10.1126/science.1133129.
  • Hahn G & Pflug HD 1985. Polypenartige Organismen aus dem Jung-Präkambrium (Nama-Gruppe) von Namibia. Geologica et Palaeontologica 19, 1–13.
  • Hall CMS, Droser ML, Clites EC & Gehling JG 2018. The short-lived but successful tri-radial body plan: a view from the Ediacaran of Australia. Australian Journal of Earth SciencesDOI: 10.1080/08120099.2018.1472666.
  • Hu XS 1997. A study on macrofossils ecostratigraphy in Doushantuo Formation at the Late Sinian in the eastern Yangtze Gorges. Journal of Xi’an College of Geology 19, 9–13. [In Chinese with English abstract]
  • Huldtgren T, Cunnigham JA, Yin C, Stampanoni M, Marone F, Donoghue PCJ & Bengtson S 2011. Fossilized Nuclei and Germination Structures Identify Ediacaran “Animal Embryos” as Encysting Protists. Science 334(6063), 1696–1699. DOI: 10.1126/science.1209537.
  • Huldtgren T, Cunnigham JA, Yin C, Stampanoni M, Marone F, Donoghue PCJ & Bengtson S 2012. Response to Comment on “Fossilized Nuclei and Germination Structures Identify Ediacaran ‘Animal Embryos’ as Encysting Protists”. Science 335(6073), 1169d (plus 3 pages Addendum). DOI: 10.1126/science.1219076.
  • Igisu M, Komiya T, Kawashima M, Nakashima S, Ueno Y, Han J, Shu D, Li Y, Guo J, Maruxama S & Takai K 2014. FTIR microspectroscopy of Ediacaran phosphatized microfossils from the Doushantuo Formation, Weng’an, South China. Gondwana Research 25, 1120–1138. DOI: 10.1016/j.gr.2013.05.002.
  • Ivantsov AYU, Fedonkin MA 2002. Conulariid-like fossil from the Vendian of Russia: a metazoan clade across the Proterozoic/Palaeozoic boundary. Palaeontology 45(6), 1219–1229. DOI: 10.1111/1475-4983.00283.
  • Ivantsov AY, Malakhovskaya YE & Serezhnikova EA 2004. Some problematic fossils from the Vendian of the southeastern White Sea Region. Paleontological Journal 38(1), 1–9. [PDF]
  • Jain S 2016. Fundamentals of Invertebrate Palaeontology. Springer, 405 pp.
  • Kaplan M 2011. Enigmatic fossils are neither animals nor bacteria. NatureDOI: 10.1038/nature.2011.9714.
  • Li C-W, Chen J-Y, Hua T-E 1998a. Precambrian sponges with cellular structures. Science 279, 879–82. DOI: 10.1126/science.279.5352.879.
  • Li C-W, Chen J-Y, Hua T-E 1998b. Interpreting Late Precambrian microfossils: Response. Science 282, 1783a. DOI: 10.1126/science.282.5395.1781r.
  • Li Y, Ding LF & Huang JC 1996. Eumetazoa and Porifera. pp. 96–119 in: Ding LF et al. (eds) Sinian Miaohe Biota. Geological Publishing House, Beijing.
  • Li Y, Guo J, Zhang X, Zhang W, Liu Y, Yang W, Li Y, Liu L & Shu D 2008. Vase-shaped microfossils from the Ediacaran Weng’an biota, Guizhou, South China. Gondwana Research 14(1–2), 263-268. DOI: 10.1016/j.gr.2007.10.002.
  • Liu AG & Conliffe J 2015. The Ediacaran fossils of the Avalon Peninsula. GAC Newfoundland and Labrador Section – 2015 Fall Field Trip, 62 pp. [PDF]
  • Love GD et al. 2009. Fossil steroids record the appearance of Demospongiae during the Cryogenian period. Nature 457, 718–721. DOI: 10.1038/nature07673.
  • Love GD & Summons RE 2015. The molecular record of Cryogenian sponges – a response to Antcliffe (2013). Palaeontology 58(6), 1131–1136. DOI: 10.1111/pala.12196.
  • Luzhnaya (Serezhnikova) EA & Ivantsov AY 2019. Skeletal Nets of the Ediacaran Fronds.Paleontological Journal 53(7), 667–675. DOI: 10.1134/S0031030119070050.
  • Maloof A et al. 2010. Possible animal-body fossils in pre-Marinoan limestones from South Australia. Nature Geoscience 3, 653–659. DOI: 10.1038/ngeo934.
  • McCaffrey MA, Moldowan JM, Lipton PA, Summons RE, Peters KE, Jeganathan A & Watt DS 1994. Paleoenvironmental Implications of novel C30 steranes in Precambrian to Cenozoic age petroleum and bitumen. Geochimica et Cosmochimica Acta 58(1), 529-532. DOI: 10.1016/0016-7037(94)90481-2.
  • McCall GJH 2006: The Vendian (Ediacaran) in the geological record: enigmas in geology’s prelude to the Cambrian explosion. Earth-Science Reviews 77, 1–229. DOI: 10.1016/j.earscirev.2005.08.004.
  • McMenamin MAS 1998. The Garden of Ediacara: Discovering the First Complex Life. Columbia University Press, 295 pp.
  • Meyer SC 2013. Darwin’s Doubt. HarperOne, viii+498 pp.
  • Moldowan JM, Dahl J, Jacobsen SR, Huizinger BJ, McCaffrey MA & Summons RE 1994. Molecular Fossil Evidence for Late Proterozoic-Early Paleozoic Environments. Terra Nova 6 (abstract supplement 3), 6.
  • Müller WEG, Li J, Schröder HC, Qiao L & Wang X 2007. The unique skeleton of siliceous sponges (Porifera; Hexactinellida and Demospongiae) that evolved first from the Urmetazoa during the Proterozoic: a review. Biogeosciences 4, 219–232. DOI: 10.5194/bg-4-219-2007. [PDF]
  • Müller WEG, Wang X & Schröder HC 2009. Paleoclimate and Evolution: Emergence of Sponges During the Neoproterozoic. pp. 55–77 in: Müller WEG & Grachev MA (eds) Biosilica in Evolution, Morphogenesis, and Nanobiotechnology. Progress in Molecular and Subcellular Biology, vol 47. Springer, Berlin, Heidelberg. DOI: 10.1007/978-3-540-88552-8_3.
  • Muscente AD, Michel FM, Dale JG, Xiao S 2015. Assessing the veracity of Precambrian ‘sponge’ fossils using in situ nanoscale analytical techniques. Precambrian Research 263, 142–156. DOI: 10.1016/j.precamres.2015.03.010.
  • Nedin C 2010. Proterozoic Sponges Claim Doesn’t Hold Water. Ediacaran September 19, 2010.
  • Nettersheim BJ et al. 2019. Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals. Nature Ecology & Evolution 3, 577–581. DOI: 10.1038/s41559-019-0806-5.
  • Peterson KJ, Lyons JB, Nowak KS, Takacs CM, Wargo MJ & McPeek MA 2004. Estimating metazoan divergence times with a molecular clock. PNAS 101(17), 6536–6541. DOI: 10.1073/pnas.0401670101.
  • Pickett JW 1983. An annotated bibliography and review of Aus­tralian fossil sponges. Association of Australasian Palaeontologists Memoir 1, 93–120.
  • Reitner J 2005. Precambrian-Cambrian sponge critical intervall – Insights in old animals. 75. Jahrestagung der Paläontologischen Gesellschaft Graz, Österreich 27. August – 2. September 2005. Berichte des Institutes für Erdwissenschaften der Karl-Franzens-Universität Graz 10, 105–106. [PDF]
  • Seilacher A 1999. Biomat-related Lifestyles in the Precambrian. Palaios 14(1), 86–93. DOI: 10.2307/3515363. [Link]
  • Sepkoski Jr JJ (edited by Jablonski D & Foote M) 2002. A compendium of fossil marine animal genera. Bulletins of American Paleontology 363(1): 1–560. [Online]
  • Serezhnikova EA 2007. Palaeophragmodictya spinosa sp. nov., a bilateral benthic organism from the Vendian of the Southeastern White Sea Region. Paleontological Journal 41, 360–369. DOI: 10.1134/S0031030107040028.
  • Serezhnikova EA 2014. Skeletogenesis in Problematic Late Proterozoic Lower Metazoa. Paleontological Journal 48(14),1457–1472. DOI: 10.1134/S0031030114140123.
  • Sperling EA & Stockey RG 2018. The Temporal and Environmental Context of Early Animal Evolution: Considering All the Ingredients of an “Explosion”. Integrative and Comparative Biology 58(4), 605–622. DOI: 10.1093/icb/icy088.
  • Sperling EA, Robinson JM, Pisani D, Peterson KJ 2010. Where’s the glass? Biomarkers, molecular clocks, and microRNAs suggest a 200-Myr missing Precambrian fossil record of siliceous sponge spicules. Geobiology 8, 24–26. DOI: 10.1111/j.1472-4669.2009.00225.x.
  • Sperling EA, Pisani D & Peterson KJ 2007. Poriferan paraphyly and its implications for Precambrian palaeobiology. In: Vickers-Rich P & Komarower P (eds) The Rise and Fall of the Ediacaran Biota. Geological Society, London, Special Publications 286, 355–368. DOI: 10.1144/SP286.25.
  • Sperling EA, Peterson KJ & LaFlamme M 2011. Rangeomorphs, Thectardis (Porifera?) and dissolved organic carbon in the Ediacaran Ocean. Geobiology 9(1), 24–33. DOI: 10.1111/j.1472-4669.2010.00259.x.
  • Stearley R 2013. The Cambrian Explosion: How Much Bang for the Buck? Perspectives on Science and Christian Faith 65(4), 245-257. [PDF]
  • Steiner M & Reitner J 2001. Evidence of organic structures in Ediacara-type fossils and associated microbial mats. Geology 29(12), 1119–1122. DOI: 10.1130/0091-7613(2001)029<1119:EOOSIE>2.0.CO;2
  • Steiner M, Mehl D, Reitner J, Erdtmann B-D 1993. Oldest entirely preserved sponges and other fossils from the Lowermost Cambrian and a new facies reconstruction of the Yangtze platform (China). Berliner geowissenschaftliche Abhandlungen 9, 293–329.
  • Steiner M 1994. Die neoproterozoischen Megaalgen Südchinas. Berliner geowissenschaftliche Abhandlungen 15, 1–146.
  • Tang Q 2019. How to find early sponge animal fossils? Nature Ecology & Evolution Community.
  • Tang Q, Wan B, Yuan X, Muscente AD & Xiao S 2019. Spiculogenesis and biomineralization in early sponge animals. Nature Communications 10, 3348, 1–10. DOI: 10.1038/s41467-019-11297-4.
  • Tang T, Zhang J & Jiang X 1978. Discovery and significance of the Late Sinian fauna from Western Hunan and Hubei. Acta Stratigraphica Sinica 2, 32–45. [In Chinese]
  • Tiwari M, Pant CC & Tewari VC 2000. Neoproterozoic sponge spicules and organic-walled microfossils from the Gangolihat Dolomite, Lesser Himalaya, India. Current Science 79(5), 651–654. [ResearchGate]
  • UC Riverside 2012. Oldest organism with skeleton discovered in Australia. Science Daily March 8, 2012.
  • UC Riverside 2018. Sponges on ancient ocean floors 100 million years before Cambrian period. Science Daily October 15, 2018.
  • Wallace MW, Hood AvS, Woon EMS, Hoffmann KH & Reed CP 2014. Enigmatic chambered structures in Cryogenian reefs: the oldest sponge-grade organisms? Precambrian Research 255, 109–123. DOI: 10.1016/j.precamres.2014.09.020.
  • Wang Y & Wang X 2011. New observations on Cucullus Steiner from the Neoproterozoic Doushantuo Formation of Guizhou, South China. Lethaia 44(3), 275–286. DOI: 10.1111/j.1502-3931.2010.00241.x.
  • Wilkinson CR 1984. Immunological evidence for the Precambrian origin of bacterial symbioses in marine sponges. Proceedings of the Royal Society B 220(1221), 509–517. DOI: 10.1098/rspb.1984.0017.
  • Wood RA, Grotzinger JP & Dickson JAD 2002. Proterozoic modular biomineralized metazoan from the Nama Group, Namibia. Science 296(5577), 2383–2386. DOI: 10.1126/science.1071599.
  • Wood R & Curtis A 2015. Extensive metazoan reefs from the Ediacaran Nama Group, Namibia: the rise of benthic suspension feeding. Geobiology 13(2), 112–122. DOI: 10.1111/gbi.12122.
  • Wood R & Penny A. 2018 Substrate growth dynamics and biomineralization of an Ediacaran encrusting poriferan. Proceedings of the Royal Society B 285, 20171938, 1–8. DOI: 10.1098/rspb.2017.1938.
  • Xiao S & Dong L 2006. On the morphological and ecological history of Proterozoic macroalgae. Chapter 3 pp. 57–90 in: Xiao S & Kaufman AJ (eds) Neoproterozoic Geobiology and Paleobiology. Topics in Geobiology, vol 27. Springer, Dordrecht. DOI: 10.1007/1-4020-5202-2_3.
  • Xiao S & Knoll AH 2000. Phosphatized Animal Embryos from the Neoproterozoic Doushantuo Formation at Weng’an, Guizhou, South China. Journal of Paleontology 74(5), 767–788. DOI: 10.1017/S002233600003300X.
  • Xiao S & Laflamme M 2008. On the eve of animal radiation: phylogeny, ecology and evolution of the Ediacara biota. Trends in Ecology and Evolution 24(1), 31–40. DOI: 10.1016/j.tree.2008.07.015.
  • Xiao S, Zhang Y & Knoll AH 1998. Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorite. Nature 391, 553–558. DOI: 10.1038/35318.
  • Xiao S, Yuan X & Knoll AH 2000. Eumetazoan fossils in terminal Proterozoic phosphorites? PNAS97(25), 13684–13689. DOI: 10.1073/pnas.250491697.
  • Xiao S, Yuan X, Steiner M & Knoll AH 2002. Macroscopic carbonaceous compressions in a terminal Proterozoic shale: A systematic reassessment of the Miaohe biota, south China. Journal of Paleontology76(2), 347–376. DOI: 10.1017/S0022336000041743.
  • Xiao S, Zhou C & Yuan X 2007. Undressing and redressing Ediacaran embryos. Nature 446, E9-E10 (Bailey et al. reply E10-E11). DOI: 10.1038/nature05753.
  • Xiao S, Knoll AH, Schiffbauer JD, Zhou C & Yuan X 2012. Comment on “Fossilized nuclei and germination structures identify Ediacaran ‘animal embryos’ as encysting protists”. Science 335(6073), 1169c. DOI: 10.1126/science.1218814.
  • Xue Y, Zhou C & Tang T 1999. “Animal Embryos”, a Misinterpretation of Neoproterozoic Microfossils. Acta Micropalaeontologica Sinica 16(1), 1–4.
  • Yin L, Xiao S & Yuan X 2001. New observations on spicule-like structures from Doushantuo phosphorites at Weng’an, Guizhou Province. Chinese Science Bulletin 46(21), 1828–1832. DOI: 10.1007/BF02900561.
  • Yin L, Zhu M, Knoll AH, Yuan X, Zhang J & Hu J 2007. Doushantuo embryos preserved inside diapause egg cysts. Nature 446, 661–663. DOI: 10.1038/nature05682.
  • Yin Z, Thu M, Davidson EH, Bottjer DJ, Zhao F & Taffereau P 2015. Sponge grade body fossil with cellular resolution dating 60 Myr before the Cambrian. PNAS 112(12), E1453–E1460. DOI: 10.1073/pnas.1414577112.
  • Yin Z, Zhu M, Bottjer DJ, Zhao F & Tafforeau P 2016. Meroblastic cleavage identifies some Ediacaran Doushantuo (China) embryo-like fossils as metazoans. Geology 44(9), 735–738. DOI: 10.1130/G38262.1.
  • Yin Z, Vargas K, Cunningham J, Bengtson S, Zhu M, Marone F & Donoghue P 2019. The Early Ediacaran Caveasphaera Foreshadows the Evolutionary Origin of Animal-like Embryology. Current Biology 29, 1–8. DOI: 10.1016/j.cub.2019.10.057.
  • Yirka B 2015. Oldest known sponge found in China. Phys.org March 10, 2015.
  • Yuan X, Xiao S, Parsley RL, Zjou C, Chen Z & Hu J 2002. Towering sponges in an Early Cambrian Lagerstätte: Disparity between nonbilaterian and bilaterian epifaunal tierers at the Neoproterozoic-Cambrian transition. Geology 30(4), 363–366. DOI: 10.1130/0091-7613(2002)030<0363:TSIAEC>2.0.CO;2.
  • Zhang Y, Yuan X, Yin L 1998. Interpreting Late Precambrian microfossils. Science 282, 1783. DOI: 10.1126/science.282.5395.1781r.
  • Zhao Z, Xing Y, Ding Q, Liu G, Zhao Y, Zhang S, Meng X, Yin C, Ning B & Han P 1988. The Sinian System of Hubei. China University of Geosciences Press, Wuhan, 205 pp.
  • Zhou CM Yuan XL & Xue YS 1998. Sponge spicule-like pseudofossils from the Neoproterozoic Doushantuo Formation in Weng’an, Guizhou, China. Acta Micropalaeontologica Sinica 15, 380–384.
  • Zhuralev AY, Liñán E, Gámez Vintaned JA, Debrenne F, Fedorov AB 2012. New finds of skeletal fossils in the terminal Neoproterozoic of the Siberian Platform and Spain. Acta Palaeontologica Polonica 57(1), 205–224. DOI: 10.4202/app.2010.0074.
  • Zhuravlev AY, Wood RA & Penny AM 2015. Ediacaran skeletal metazoan interpreted as a lophophorate. Proceedings of the Royal Society B 282(1818), 20151860, 1–10. DOI: 10.1098/rspb.2015.1860.
  • Zimmer C 2019. Is This the First Fossil of an Embryo? New York Times Nov. 27, 2019.
  • Zumberge JA, Love GD, Cárdenas P, Sperling EA, Gunasekera S, Rohrssen M, Grosjean E, Grotzinger JP & Summons RE 2018. Demosponge steroid biomarker 26-methylstigmastane provides evidence for Neoproterozoic animals. Nature Ecology & Evolution 2, 1709–1717. DOI: 10.1038/ s41559-018-0676-2.

Photo: Living Guantanamo sponge, by Timothy W. Brown / Public domain.