The word suture is strongly engrained in the language we use for describing insects. Who would have thougt that we have been using it wrong for a long time. I know who, Snodgrass did!
Back in the 60s Snodgrass talked about "sutres" in a treatise about the cephalic capsule. Then in1963 when his last contribution was published posthumously —he had written multiple essays on topics of insect anatomy with the goal to assemble an entire enyclopedia. I wish he have had the time to complete the task. But that's the thing with humans. We eventually pass away and we scientists are pretty good at leaving unfinished businesses. As Glenn Richards (who wrote the foreword of that last publication) put it: "who besides Snodgrass could write Snodgrass's Encyclopedia?"
Anyway. About the word suture Snodgrass (1960) wrote:
"Scientific terms should express facts rather than perpetuate errors. When errors become chronic, however, they are hard to eradicate."
Robert Evans Snodgrass, 1963
And he insisted in 1963:
When thinking about this, one of the issues is —at least from my own experience in my early learning of insect anatomy— that we end up interpreting insects as if they were assembled in parts. What comes to mind are those cartoons where the armor of the knight assembles onto the person by parts and ends up being functional and protecting the person under it.
In reality, the entire exoskeleton of insects is continuous, constituted by hardened (sclerotized) plates connected by flexible membranes (conjunctivae), and marked by points of inflection that produce those folding lines that have been traditionally called sutures. These lines were originally interpreted the same way as the sutures of the vertebrate skull, where separate bone plates meet and immovably join together. The whole nomenclatural system for insect anatomy being historically based on vertebrates causes many issues. A friend of mine who is an artist and works at the library, understood the situation completely and immediately referred to Kafka's metamorphosis. That is a good way to understand what happens. It is as if parts of Gregor Samsa's skin got hardened, with just enough flexible bits in between to allow him to move around.
In anatomical terms, the connection between the bones of a skull and the sclerites of an insect (or any arthropod for that matter) is not at all the same condition. Sclerites, the hardened plates, do not start as separate units that grow closer together as the bones of the vertebrate skull, but rather each sclerotized plate is fully embedded and connected throughout the whole integument, which is secreted together, as a continuous unit, as it originates from the single layer epithelium (see Girón et al. 2023). Anyone who has seen an insect molt, a cicada for instance, can visualize this interconnection.
Snodgrass, argued this position in 1960 and 1963. Beutel et al. (2014) in their Insect morphology and phylogeny textbook support this view (at least in the introductory morphology section of the book). The thing is that most people, at least on this side of the world, learned insect anatomy and morphology from Snodgrass himself, but from Snodgrass in 1935, when he had not yet analyzed the issue as he did when writing his encyclopedia.
And so, there I was, with the biggest chance I have had of changing tradition for accuracy. Of course, my coauthor and coeditors were not really on board. It is very difficult to change when you have used the term with that meaning for 20, 30, 40 years or more, and you feel there is no need to change that, since "everyone knows" what is meant. I climbed up on this hill, decided to hold the fort. I even consulted with my anatomy buddy, István Mikó, who is in the same boat as me for accuracy of definitions for anatomical terms. And then beetles happened.
Beetles, my friends, are such amazing creatures. Their external anatomy is so highly modified from the general structure of insects, that there are beetle structures in the same regions of the body as in other insects, in the thorax in particular, for which the nomenclature has been fairly recently modified (Beutel and Haas 2000; Friedrich et al. 2009; Lawrence et al. 2010, 2011).
In case you didn't know it yet, beetles do not have a mesosternum or a metasternum in the strict sense. Regions of these plates have extended internally to form endoskeletal elements for muscle attachment. The most famous, the metendosternite or metafurca. Crowson had two entire treatments about it (Crowson 1938, 1944), but even before I leaned about Crowson's work on this, I read Velázquez de Castro (1998), since he studied this structure for a particular group of broad-nosed weevils.
The ventral plates of the beetle mesothorax and metathorax are termed mesoventrite and metaventrite nowadays (Beutel and Haas 2000; Friedrich et al. 2009; Lawrence et al. 2010, 2011). The name of the visible badominal plates is now ventrites, as the first two true sternites are usually not externally visible.
Crazy, right? The most incredible thing, is that even if these terms were updated about 25 years ago, and a lof of people have larned and corrected their usage, many still use the traditional mesosternum, metasternum, and sternite when talking about beetle anatomy. I guess it takes an extra effort and for sure a long time for everyone to catch up.
Going back to the word suture in insect anatomy, my latest realization is that the meaning of the word might just have changed, perhaps by usage, or because it has been interpreted wrong all along, and we keep perpetuating the error. I ended up getting a copy of the Henderson's dictionary of biological terms (Lawrence 2016), where the definition of suture even includes the arthropod exoskeleton as an example of the word!
The issue now is that beetles do have multiple parts that have actually fused or come together to be solidly fixed. For instance, the gular, notopleural, pleurosternal, and metakatepisternal, all seem to be sutures —the immovable connection between sclerites—. The "elytral suture" though, even if we have been calling it a "suture", is not it unless the elytra are actually fused together along the junction between the elytra.
I do think that the proper terminology should be adopted and used, just as people should stop using "mesosternum" for beetles. We'll see how long it takes for beetle researchers to catch up.
Beutel, R.G. and F. Haas. 2000. Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics 16(1): 103–141. https://doi.org/10.1006/clad.1999.0124
Beutel, R.G., Friedrich, F., Yang, X.-K. & Ge, S.-Q. (2014) Insect Morphology and Phylogeny Insect morphology and phylogeny: a textbook for students of entomology. De Gruyter. 516 pp. https://doi.org/10.1515/9783110264043
Crowson, R.A. (1938) The metendosternite in Coleoptera: a comparative study. Transactions of the Royal Entomological Society of London 87, 397–415. https://doi.org/10.1111/j.1365-2311.1938.tb00723.x
Crowson, R.A. (1944) Further studies on the metendosternite in Coleoptera. Transactions of the Royal Entomological Society of London 94, 273–310. https://doi.org/10.1111/j.1365-2311.1944.tb01220.x
Friedrich, F., B.D. Farrell, and R.G. Beutel. 2009. The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda). Cladistics 25: 1–37. https://doi.org/10.1111/j.1096-0031.2008.00233.x
Girón, J.C., Tarasov, S., González Montaña, L.A., Matentzoglu, N., Smith, A.D., Koch, M., Boudinot, B.E., Bouchard, P., Burks, R., Vogt, L., Yoder, M., Osumi-Sutherland, D., Friedrich, F., Beutel, R. & Mikó, I. (2023) Formalizing Invertebrate Morphological Data: A Descriptive Model for Cuticle-Based Skeleto-Muscular Systems, an Ontology for Insect Anatomy, and their Potential Applications in Biodiversity Research and Informatics. Systematic Biology, syad025. https://doi.org/10.1093/sysbio/syad025
Lawrence, E (2016) Henderson's dictionary of biological terms. Pearson Education Limited, United Kingdom. 683 pp.
Lawrence, J.F., Beutel, R.G., Leschen, R.A. & Ślipiński, A. (2010) Glossary of morphological terms. Handbuch der Zoologie/Handbook of Zoology. Band 4, 9–20. https://doi.org/10.1515/9783110911213.9
Lawrence, J.F., Ślipiński, A., Seago, A.E., Thayer, M.K., Newton, A.F. & Marvaldi, A.E. (2011) Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici 61, 1–217. https://doi.org/10.3161/000345411X576725
Snodgrass, R.E. (1935) Principles of Insect Morphology. Cornell University Press, 768 pp. https://www.jstor.org/stable/10.7591/j.ctv1nhm1j
Snodgrass, R.E. (1960) Facts and theories concerning the insect head. Smithsonian Miscellaneous Collections 142, 1–61. http://repository.si.edu/xmlui/handle/10088/22981
Snodgrass, R.E. (1963) A contribution toward an encyclopedia of insect anatomy. Smithsonian Miscellaneous Collections 146, 1–48. https://www.biodiversitylibrary.org/page/30259679
Velázquez de Castro, A.J. (1998) Morphology and taxonomy of the genus Sitona Germar, 1817. (I): the metendosternite (Coleoptera: Curculionidae). In: Proceedings of the XX International Congress of Entomology. Museo Regionale di Scienze Naturali, Torino, Italy, pp. 109–123.