This Fossil Friday we will have a look at three new insect orders that have been described in the past decade from almost 100 million year old mid-Cretaceous Burmese amber, but later turned out to be highly problematic and likely no new orders at all.

Alienoptera

The new insect order Alienoptera was erected by Bai et al. (2016) for strange roach-like insects with short scale-like forewings, which were considered “an evolutionary dead end in the roach-mantis transition zone/” A second species was described by Kočárek (2019), who considered the insects of this order as diurnal predators of small invertebrates. Sendi et al. (2020) could show that some alienopterids were nectar-feeding pollinators. I co-authored an extensive study by Vršanský et al. (2018), which suggested that alienopterids are just a derived subgroup of umenocoleoid roachoids (Vršanský 2003, Nel et al. 2014) that were adapted for a Batesian mimicry of wasps and ants. Wipfler et al. (2019) supported the interpretation as predators but rejected the mimicry hypothesis suggested by Vršanský et al. (2018) and Hinkelman (2020) and again supported by Luo et al. (2022). Alienoptera was considered as an invalid subjective junior synonym of the order Blattaria, and Alienopteridae a second family within Umenocoleoidea, by Vršanský et al. (2018, 2021) and Sendi et al. (2020). Luo et al. (2022) agreed that “Alienopteridae should not be ranked as a separate order” but suggested the name Alienoptera for an unranked clade that includes Umenocoleidae and Alienopteridae and represents the sister group of praying mantids (Mantodea). This makes Alienoptera equivalent to and co-extensive with the superfamily Umenocoleoidea (Vršanský et al. 2021), but not formally a synonym because the international rules for zoological nomenclature do not apply to taxa above the family-group. Nel et al. (2014) had introduced the new family taxon Ponopterixidae for umenocoleoid roachoids, because they agreed with Kirejtshuk et al. (2013) that the holotype of Umenocoleus sinuatus is rather a primitive beetle than a roachoid. This was refuted by Luo et al. (2021), who studied new material that clearly suggests a dictyopteran (roachoid) affinity of Umenocoleus as sister group of Alienopteridae, so that Ponopterixidae is an invalid junior synonym of Umenocoleidae.

Aethiocarenodea

The new insect order Aethiocarenodea was described by Poinar & Brown (2017), who did not find any evidence for relationship with other known orders. This discovery was even celebrated as newest insect order in the Guinness Book of World Records 2017. Based on new material, we could show (Vršanský et al. 2018), that Aethiocarenus burmanicus (featured this Fossil Friday) is just a nymph of a new species of Alienopteridae, as was already suggested by Hörnig et al. (2017) and later confirmed by Luo et al. (2022). Therefore, the supposed new order Aethiocarenodea is yet another invalid subjective junior synonym of Blattaria, and Aethiocarenidae a synonym of Alienopteridae.

Tarachoptera

Finally, the latest new insect order Tarachoptera was described by Mey et al. (2017), who placed this new order in the stem-group of Amphiesmenoptera (caddisflies and butterflies), because they allegedly differ in nearly all aspects from caddisflies (Trichoptera) and butterflies (Lepidoptera). This is surprising, because the authors describe the presence of scales on the wings of Tarachoptera, which happens to be the most distinctive derived character that distinguishes butterflies from caddisflies, and even is the character that inspired the scientific name of butterflies (Lepidoptera = scaly wings). The erection of the new order and its placement is at odds with usual systematic reasoning, and Tarachoptera more likely just represents a very basal group of butterflies. Indeed, Mey et al. (2018a) no longer mentioned the order Tarachoptera (except in the references), but classified these insects as a family Tarachocelidae with uncertain position within Amphiesmenoptera, which is the clade formed by caddisflies, moths, and butterflies. Mey et al. (2018b, 2020), Wichard & Mey (2021), Wichard (2022), and Mey & Wichard (2023) again used the order Tarachoptera in their description of further new taxa. None of the multiple studies on Tarachoptera co-authored by Wolfram Mey, Wilfried Wichard, and their colleagues presented any proper phylogenetic analysis to support the position of Tarachoptera as distinct order.

A Flawed Study

However, an independent group of mostly Chinese scientists (Wang et al. 2022) recently claimed that Tarachoptera, Trichoptera, and Lepidoptera all shared wing scales as a primitive ground plan condition, and presented a cladistic analysis, which suggested that Tarachoptera forms the sister-group of a Trichoptera + Lepidoptera clade. However, their cladistic analysis is tarnished by errors. For example these authors listed the presence of hairy pulvilli, as an allegedly non-homoplastic character (character 117) that supports an exclusion of Tarachoptera from a clade formed by Trichoptera and Lepidoptera, even though they themselves write in their supplementary information that this character is present in the most basal tarachopteran genus Tarachocelis. Furthermore, they support this topology with the absence of a jugate wing coupling (character 82) in Tarachoptera, even though this is also absent in many Lepidoptera and some Trichoptera. Indeed, the diversity of very different wing coupling mechanisms within Trichoptera and within Lepidoptera makes a homology and common ground plan highly unlikely. Stocks (2010) therefore concluded that “the caddisfly suborders Annulipalpia and Integripalpia (Trichoptera) each have evolved a wing coupling apparatus, with at least three systems having evolved within the suborder Annulipalpia”. This leaves only a single obscure character (character 45: posteromedian corner of lateral cervical sclerite protruding towards the prosternum or not) in support of an outgroup phylogenetic position of Tarachoptera, instead of the above mentioned obvious position as most basal Lepidoptera, which is arguably supported by all four wings with scales. Interestingly, Mey & Wichard (2023) explicitly state that Tarachoptera shares the character “lower posterior corner of laterocervicale produced towards prosternum” as an amphiesmenopteran synapomorphy with Trichoptera and Lepidoptera. This would refute the last support for a position of Tarachoptera basal of the Trichoptera-Lepidoptera split.

According to a colleague (pers. comm. July 2023), who is also a renowned expert on aquatic insects but for obvious reasons prefers not to be quoted by me with his name, the study by Wang et al. (2022) is indeed very flawed. For example, it also describes from Burmese amber the new family Lepidochlamidae as alleged most basal caddisflies with scales on the hind wings. According to this colleague, the wing venation of the holotype specimen of Lepidochlamus nodosa clearly suggests an attribution to the derived clade Leptoceroidea, which is deeply subordinated and not representative for basal caddisflies. All basal caddisflies have hairy wings, without any scales, while scales are only found as convergence — wait for it — in the derived Leptoceridae (Huxley & Barnard 1988) and some other derived subgroups (Robertson & Holzenthal 2008). Incidentally, the specimen later described as first Tarachoptera was figured as “first leptocerid caddisly (Trichoptera: Leptoceridae) in Burmese amber” in a book chapter by Ross et al. (2010) (quoted by Wichard 2022). Robertson & Holzenthal (2008) unambiguously clarify that “although superficially similar, the scales of Lepidoptera and Trichoptera are not homologous.” A similar convergence of scaled wings is also found in the butterfly-like Mesozoic lacewing family Kalligrammatidae (Labandeira et al. 2016). Wang et al.’s case for a clade of scale-winged insects does not stand to scrutiny. As we have seen, the case for a distinct insect order Tarachoptera is also built on a very shaky foundation of highly incongruent (homoplastic) similarities.

Another Flawed Study

Like the study of Wang et al. (2022), a more recent paper by Mey & Wichard (2023: fig. 10) featured a tree, which resolves Tarachoptera outside of the Trichoptera+Lepidoptera clade, but the authors propose an even more distant relationship with these Amphiesmenoptera sensu stricto. However, these authors once more did not provide a proper cladistic analysis but only proposed a manually constructed tree as a theoretical model to interpret the data. The authors explicitly admit:

It is not a new hypothesis, but rather a theoretical construct that aims at providing some guidance into the interpretation of known and unknown fossil taxa. … A good and comprehensive knowledge about the richness of the fossil record is a prerequisite for assigning characters and taxonomic names into the cladogram, knowledge, which we do not have at this stage of research.

Their tree and its labelling shows a confusion about the terminology of phylogenetics, even from a mainstream point of view. For example the authors say:

According to the model in Fig. 10, Trichoptera and Lepidoptera do not have a strict sister-group relationship. Their proper sister-groups have still to be found or defined. The same applies to the Amphiesmenoptera in its traditional sense of being restricted to Trichoptera and Lepidoptera. The sister group of Amphiesmenoptera is unknown, but some of the known fossils might be ascribed to this clade. The Protomeropina represent the most basal group in the amphiesmenopteran clade and include the oldest fossils assigned to this taxon. 

The Tarachoptera are grouped in a Tarachoptera clade, distantly related to Trichoptera and Lepidoptera, which is expressed in the placement of this order in a somewhat remote branch of the cladogram.

Sorry, but this is meaningless, and shows that the authors don’t know how technical terms like sister-group and stem-group are properly defined in phylogenetics or cladistics (see Budd & Jensen 2000: fig. 1). Sister-groups are simply the most closely related pair of living taxa, so that there is no dispute among experts that Trichoptera and Lepidoptera are sister-groups. In their figured tree (fig. 10) the authors strangely labelled unnamed branches as “sistergroup” and “stemgroup” to Tarachoptera, Trichoptera, and Lepidoptera respectively, and obviously mean unknown extinct taxa. However, in cladistic terminology a fossil sister-group of a taxon simply is a derived member of its stem-group. A stem-group is defined as the group of all extinct taxa that are closer related to a living group than to its living sister-group, but not nested within the living group. What their tree really implies is simply that the authors consider Tarachoptera as a more basal member of the amphiesmenopteran stem-group than the extinct Protomeropina. But they do not justify this with any actual data or arguments. Protomeropina are are an enigmatic group of Paleozoic holometabolans of highly disputed relationship, which have been suggested to be related to either Mecoptera, or Trichoptera, or Amphiesmemoptera, or Antliophora. But the authors do not even discuss their own placement of Protomeropina and just cite a twenty year old book on caddisflies (Wiggins 2004), even though newer paleoentomological studies by actual experts have proposed very different relationships for Protomeropina (e.g., Nel et al. 2007, Sukatcheva & Aristov 2013, Bashkuev & Sukatsheva 2021, Poschmann & Nel 2021). In summary, the paper by Mey & Wichard (2023) is plagued by various shortcomings and cannot be considered as authoritative. Until a proper phylogenetic analysis supports a position of Tarachoptera outside the stem-group of any living order, its status as new insect order remains dubious and doubtful. If they should turn out to be closer related to Lepidoptera as I suppose, their morphology would hardly be distinct enough from primitive moths to warrant a separate ordinal status. 

An Inflation of Orders

It is tempting to suppose that the recent inflation of new insect orders from Burmese amber might have more to do with public relation issues than with realties of nature. Nevertheless, a proper analysis can well justify the erection of new orders from this locality. An example is the new order Permopsocida by Huang et al. (2016), which I happened to co-author, but which was just an elevation in rank for a taxon previously established as suborder by Tillyard (1926). This new rank was based on its recovered phylogenetic position as sister group to thrips and hemipterans, which was independently confirmed by Yoshizawa & Lienhard (2016). Other new insect orders co-authored by me include the neuropteran order Schwickertoptera (Martins-Neto et al. 2007: 329) and the mayfly-related chimera-wings Coxoplectoptera (Staniczek et al. 2011), both from the Lower Cretaceous Crato Formation of Brazil. Coxoplectopterans are likely stem-group mayflies and thus could as well be included as suborder or the order Ephemeroptera, but the morphology of the adults and larvae is so strikingly unique and distinct from mayflies that we decided that a separate ordinal status is more appropriate. So, my issue is not the erection of new higher taxa when justified by the evidence, but the premature and weakly supported inflation of higher taxa caused by the publish-or-perish peer pressure in modern science.

Evidence for Design

Apart from this more general critique, are there any implications from these amber insects for intelligent design theory? You bet! For example the ubiquitous phenomenon of convergence (beetle-like Umenocoleoidea, wing scales and different wing coupling mechanisms in various insect orders) and the fantastic design of Batesian mimicry, which are not plausibly explained by an unguided evolutionary process, but arguably well explained by intelligent design.

References

• Bai M, Beutel RG, Klass K-D, Zhang W, Yang X & Wipfler B 2016. †Alienoptera — A new insect order in the roach-mantodean twilight zone. Gondwana Research 39, 317–326. DOI: https://doi.org/10.1016/j.gr.2016.02.002
• Bashkuev AS & Sukatsheva ID 2021. New species of Kamopanorpa Martynov from the Permian of South Siberia with comments on the systematic position of Microptysmatidae (Protomeropina = Permotrichoptera). Palaeoentomology 4(5), 453–461. DOI: https://doi.org/10.11646/palaeoentomology.4.5.11
• Budd GE & Jensen S 2000. A critical reappraisal of the fossil record of the bilaterian phyla. Biological Reviews 75(2), 253–295. DOI: https://doi.org/10.1111/j.1469-185x.1999.tb00046.x
• Hinkelman J 2020. Earliest behavioral mimicry and possible food begging in a Mesozoic alienopterid pollinator. Biologia 75(1), 83–92. DOI: https://doi.org/10.2478/s11756-019-00278-z
• Hörnig MK, Haug JT & Haug C 2017. An exceptionally preserved 110 million years old praying mantis provides new insights into the predatory behavior of early mantodeans. PeerJ 5: e3605, 1–19. DOI: https://doi.org/10.7717/peerj.3605
• Huang D-Y, Bechly G, Nel P et al. 2016. New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria). Scientific Reports 6: 23004, 1–9. DOI: https://doi.org/10.1038/srep23004
• Huxley J & Barnard PC 1988. Wing-scales of Pseudoleptocerus chirindensis Kimmins (Trichoptera: Leptoceridae). Zoological Journal of the Linnean Society 92(3), 285–312. DOI: https://doi.org/10.1111/j.1096-3642.1988.tb01514.x
• Kirejtshuk AG, Poschmann M, Prokop J, Garrouste R & Nel A 2013. Evolution of the elytral venation and structural adaptations in the oldest Palaeozoic beetles (Insecta: Coleoptera: Tshekardocoleidae). Journal of Systematic Palaeontology 12(5), 575–600. DOI: https://doi.org/10.1080/14772019.2013.821530
• Kočárek P 2019. Alienopterella stigmatica gen. et sp. nov.: the second known species and specimen of Alienoptera extends knowledge about this Cretaceous order (Insecta: Polyneoptera). Journal of Systematic Palaeontology 17(6), 491–499. DOI: https://doi.org/10.1080/14772019.2018.1440440
• Labandeira CC, Yang Q, Santiago-Blay JA et al. 2016 The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies.
  Proceedings of the Royal Society B 283(1824): 20152893, 1–9. DOI: https://doi.org/10.1098/rspb.2015.2893
• Luo C-H, Beutel RG, Thomson UR, Zheng D-R, Li J-H, Zhao X-Y, Zhang H-C & Wang B 2021. Beetle or roach: systematic position of the enigmatic Umenocoleidae based on new material from Zhonggou Formation in Jiuquan, Northwest China, and a morphocladistic analysis. Palaeoworld 31(1), 121–130. DOI: https://doi.org/10.1016/j.palwor.2021.01.003
• Luo C, Beutel RG, Engel MS, Liang K, Li L, Li J, Xu C, Vršanský P, Jarzembowski E & Wang B 2022. Life history and evolution of the enigmatic Cretaceous–Eocene Alienopteridae: A critical review. Earth-Science Reviews 225: 103914, 1–18. DOI: https://doi.org/10.1016/j.earscirev.2021.103914
• Martins-Neto RG, Heads S & Bechly G 2007. Neuropterida: snakeflies, dobsonflies and lacewings. Chapter 11.16, pp. 328–340 in: Martill DM, Bechly G & Loveridge RF (eds). The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press, Cambridge (UK), xvi + 625 pp. https://books.google.de/books?hl=de&id=5ZBh_-QvX2MC
• Mey W & Wichard W 2023. Tarachoptera: The extinct and enigmatic cousins of Trichoptera and Lepidoptera, with descriptions of two new species. Contributions to Entomology 73(2), 137–146. DOI: https://doi.org/10.3897/contrib.entomol.73.e110233
• Mey W, Wichard W, Müller P & Wang B 2017. The blueprint of the Amphiesmenoptera – Tarachoptera, a new order of insects from Burmese amber (Insecta, Amphiesmenoptera). Fossil Record 20(2), 129–145. DOI: https://doi.org/10.5194/fr-20-129-2017
• Mey W, Wichard W, Ross E & Ross A 2018a. On the systematic position of a highly derived amphiesmenopteran insect from Burmese amber (Insecta, Amphiesmenoptera). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 107(2-3), 249–254. DOI: https://doi.org/10.1017/S1755691017000330
• Mey W, Wichard W, Müller P, Ross E & Ross A 2018b. New taxa of Tarachoptera from Burmese amber (Insecta, Amphiesmenoptera). Cretaceous Research 90, 154–162. DOI: https://doi.org/10.1016/j.cretres.2018.04.006
• Mey W, Wichard W, Müller P & Wang Bo 2020. Descriptions of two new species of Tarachoptera from Burmese amber (Insecta, Amphiesmenoptera: Tarachoptera). Beiträge zur Entomologie = Contributions to Entomology 70(1), 181–188 Seiten. DOI: https://doi.org/10.21248/contrib.entomol.70.1.181-188
• Nel A, Roques P, Nel P, Prokop J & Steyer JS 2007. The earliest holometabolous insect from the Carboniferous: a “crucial” innovation with delayed success (Insecta Protomeropina Protomeropidae). Annales de la Société Entomologique de France 43(3), 349–355. DOI: https://doi.org/10.1080/00379271.2007.10697531
• Nel A, Prokop J, Grandcolas P, Garrouste R, Lapeyrie J, Legendre F, Anisyutkin LN & Kirejtshuk AG 2014. The beetle-like Palaeozoic and Mesozoic roachoids of the so-called “umenocoleoid” lineage (Dictyoptera: Ponopterixidae fam. nov.). Comptes Rendus Palevol 13(7), 545–554. DOI: https://doi.org/10.1016/j.crpv.2014.05.002
• Poinar G Jr & Brown AE 2017. An exotic insect Aethiocarenus burmanicus gen. et sp. nov. (Aethiocarenodea ord. nov., Aethiocarenidae fam. nov.) from mid-Cretaceous Myanmar amber. Cretaceous Research 72, 100–104. DOI: https://doi.org/10.1016/j.cretres.2016.12.011
• Poschmann MJ & Nel A 2021. The first Permian scorpionfly from Germany (Insecta, Panorpida: Protomeropidae). Palaeoentomology 4(3), 231–236. DOI: https://doi.org/10.11646/palaeoentomology.4.3.10
• Robertson DR & Holzenthal RW 2008. Two New Species and a New Record of Protoptila from Bolivia (Trichoptera: Glossosomatidae: Protoptilinae). Annals of the Entomological Society of America 101(3), 465–473. DOI: https://doi.org/10.1603/0013-8746(2008)101[465:TNSAAN]2.0.CO;2
• Ross A, Mellish C, York P & Crighton B 2010. Burmese Amber. pp. 208–235 in: Penney D (ed.). Biodiversity of fossils in amber from the major world deposits. Siri Scientific Press, Rochdale (UK), 304 pp. https://siriscientificpress.co.uk/products/biodiversity-of-fossils-in-amber-from-the-major-world-deposits
• Sendi H, Hinkelman J, Vršanská L, Kúdelova T, Kúdela M, Zuber M, van de Kamp T & Vršanský P 2020. Roach nectarivory, gymnosperm and earliest flower pollination evidence from Cretaceous ambers. Biologia 75(10), 1613–1630. DOI: https://doi.org/10.2478/s11756-019-00412-x
• Staniczek AH, Bechly G & Godunko RJ 2011. Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera). Insect Systematics & Evolution 42, 101–138. DOI: https://doi.org/10.1163/187631211X578406
• Stocks IC 2010. Comparative and functional morphology of wing coupling structures in Trichoptera: Annulipalpia. Journal of Morphology 271(2), 152–168. DOI: https://doi.org/10.1002/jmor.10788
• Sukacheva ID & Aristov DS 2013. New Caddisflies of the Suborder Protomeropina (Insecta: Trichoptera) from the Permian of Russia. Paleontological Journal 47(7), 741–751. https://www.researchgate.net/publication/258402993
• Tillyard RJ 1926. Kansas Permian insects; Part 8, the order Copeognatha. American Journal of Science 11(64), 314–349.
• Vršanský P 2003. Umenocoleoidea – an amazing lineage of aberrant insects (Insecta, Blattaria). AMBA Projekty 7(1), 1–32.
• Vršanský P, Bechly G, Zhang Q et al. 2018. Batesian insect-insect mimicry-related explosive radiation of ancient alienopterid cockroaches. Biologia 73(10), 987–1006. DOI: https://doi.org/10.2478/s11756-018-0117-3
• Vršanský P, Sendi H, Hinkelman J & Hain M 2021. Alienopterix Mlynský et al., 2018 complex in North Myanmar amber supports Umenocoleoidea/ae status. Biologia 76(8), 2207–2224. DOI: https://doi.org/10.1007/s11756-021-00689-x
• Wang J, Zhang W, Engel MS, Sheng X, Shih C & Ren D 2022. Early evolution of wing scales prior to the rise of moths and butterflies. Current Biology 32(17), 3808–3814. DOI: https://doi.org/10.1016/j.cub.2022.06.086
• Wichard W 2022. Tarachoptera, eine neue fossile Insekten-Ordnung im Burma-Bernstein. pp. 68–73. https://www.researchgate.net/publication/373093673
• Wichard W & Mey W 2021. Order Tarachoptera in mid-Cretaceous Burmese amber (Insecta, Amphiesmenoptera). Cretaceous Research 119: 104681. DOI: https://doi.org/10.1016/j.cretres.2020.104681
• Wiggins G 2004. Caddisflies: The Underwater Architects. University of Toronto Press, Toronto (Canada), 304 pp. DOI: https://doi.org/10.3138/9781442623590
• Wipfler B, Kočárek P, Richter A, Boudinot B, Bai M & Beutel RG 2019. Structural features and life habits of †Alienoptera (Polyneoptera, Dictyoptera, Insecta). Palaeoentomology 2(5), 465–473. DOI: https://doi.org/10.11646/palaeoentomology.2.5.10
• Yoshizawa K & Lienhard C 2016. Bridging the gap between chewing and sucking in the hemipteroid insects:new insights from Cretaceous amber. Zootaxa 4079(2), 229–245. DOI: https://doi.org/10.11646/zootaxa.4079.2.5