# It Don't Mean a Thing, if It Ain't Got That Wing:
## Aphids in the Arts, or the Invisible Made Visible

Sibylle Orlandi, Yvan Rahbé

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Sibylle Orlandi, Yvan Rahbé. It Don't Mean a Thing, if It Ain't Got That Wing: Aphids in the Arts, or the Invisible Made Visible. 2024, 10.15454/Z3BOPV. hal-05026512

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# It Don't Mean a Thing, if It Ain't Got That Wing: Aphids in the Arts, or the Invisible Made Visible

Sibylle Orlandi<fnref n="1" /> and Yvan Rahbé<fnref n="2" />*

**Simple Summary:** Insects are among the most prominent and diverse animal groups interacting with humans. How they are perceived by human societies, and how artists mediate this interaction, is a singular object of reflection. The relationship has already been analyzed for the most visible/em- blematic of insect taxa, but many others remain seemingly hidden. With a case study of aphids, we will show that invisibility itself, and many other traits of “tiny insects”, have been the subject of reflection for a cohort of artists across human history. We will briefly scan a series of examples on the interaction between nature (aphids, insects), art (artists and scientists) and, finally, human soci- eties from before ancient Greece through to the internet era. Aristotle, the Brueghel's, Joris Hoef- naeghel, Maria Sibylla Merian, Victor Hugo, Simone de Beauvoir and other artists have all been somewhat inspired by such *insignificant* insects and their peculiarities (primarily their reproduction, but also feeding habits or their minute size). However, in the times of the microbiota, of insects' decline and of the sixth extinction, one might actually question the link between size and signifi- cance.

**Abstract:** (1) Background: Insects have been frequently used by many artists throughout history to illustrate a series of well-defined symbols of human cultural traits, and these insects were usually holometabolous members of the class. Many others were, nonetheless, chosen to represent more diverse feelings and intentions but were never reviewed until now. Our purpose is to unbury this material and analyze why and how they were used by artists, and by which artists. (2) Methods: A database has been built from different French, Dutch, English and internationally accessible sources, resulting in more than 200 mentions of either aphids (our case study) or "equivalent" underexposed insects. This DB includes community inputs, is freely accessible and was used to analyze semanti- cally, aesthetically and symbolically the resulting corpus. (3) Results: In contrast to our initial intu- itive feelings, extensive references to aphids do exist in (all) the arts, proving that artists were sen- sitive to their presence from at least the seventeenth century onwards. Graphical arts and literature account for the most abundant occurrences (>80 each) and all the arts were concerned, albeit at low levels (3 mentions for architecture, 6 for music). The three main concepts associated with aphids are undeniably reproduction (including parthenogenesis, an astonishingly well-known feature), mi- nuteness and villainy (in the form of parasitism). (4) Conclusions: This picture gives a quite varied and equivocal "personality" to our selected taxon, rendering mademoiselle aphid, la *pucerone* as Charles Bonnet would have said a very modern insect character.

**Keywords:** art; aphids; insects; painting; graphical arts; literature; movies; theatre; music; photography

**Citation:** Orlandi, S., Rahbé, Y. It Don't Mean a Thing, if It Ain't Got That Wing: Aphids in the Arts, or the Invisible Made Visible.

Academic Editor(s):

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**Copyright:** © 2022 by the authors. Submitted for possible open access publication under the terms and conditions of the Creative Commons Attribution (CC BY) license (https://creativecommons.org/licenses/by/4.0/).

<footnote n="1">¹ Univ Nantes, LAMO Littératures Modernes et Antiques, Censive Bldg, Chemin de la Censive du Tertre BP 81227, CEDEX 3, 44312 Nantes, France; sibylle.orlandi@univ-nantes.fr</footnote>

<footnote n="2">² INRAE, Univ Lyon, INSA Lyon, CNRS, UCBL, MAP UMR5240, 69621 Villeurbanne, France</footnote>

<footnote>* Correspondence: yvan.rahbe@inrae.fr; Tel.: +33-675179032</footnote>
# 1. Introduction

The story all began in the Pinacoteca Ambrosiana of Milan in 2017, when one of the authors was amazed to discover that an aphid, the crescent lily aphid, had indeed been painted in the seventeenth century on a bouquet of flowers, an example of the then flourishing genre of the Dutch Renaissance—flower still lifes. This was incredible because it was unexpected (never before seen despite years of both visiting museums and studying insect science), surprising (“would never think this possible”), and, of course, almost invisible (only detected by chance because of a crowded room). The painter, far from being obscure, was Brueghel the Elder, nicknamed velvet Brueghel, one of the most valued artists of his time. This experience has been briefly described [1,2], including the painting and species identification of the magnified aphid ([2, Fig.1]Figure 1). However, this triggered a more comprehensive experience culminating in the present publication reflecting academic ideas on the reasons causing an artist to embrace unlikely and unexpected subjects. It may be noted, as we will see later, that the topic is a declination of a wider questioning in the history of arts, that of the *detail*, which has been brilliantly investigated by Arasse, for example [3].

“Insects in arts” is a subject of interest for many entomologists [4,5] and some humanities scholars who have produced inspiring books [6,7]. Overall, the topic has generated more than 120 texts, in all the arts, including exhibitions and, therefore, is of general and open interest [8-10]. A quotation from Pablo Picasso exemplifies the impressions of insects themselves as artwork, and symbolizes a rather general fascination for their (bio)diversity of forms: *I don't know what it is for, I don't know where it comes from, but I understand what the artist wanted to do* [10]. However, a careful parsing of the natural objects of such representations show that they are taxonomically heavily biased: they are almost exclusively showing, or referring to, the four major holometabolous orders (Lepidoptera, Coleoptera, Hymenoptera and Diptera), with the addition of the intermediate Odonata (dragon- and damselflies) and the paurometabolous Orthoptera (grasshoppers, crickets etc.). All the other orders (Hemiptera, Blattodea, Thysanoptera...) are rarely represented, estimated at considerably less than 2% in Western painting, for example [11]. A notable exception is an interest of scholars and artists represented by a specific analysis of Neuroptera in the arts, approximately one decade ago [12].

As will be seen below, the main symbolic attributes linked to the principal insects represented in paintings, for example, are related to life and death (putrefaction, death, afterlife, metamorphosis...) or to beauty (sky, lightness) and the senses (mainly sight and touch, with compound eyes and antennas). We will see that this is quite different from the representative attributes of “non-canonical” insects, such as aphids or other under-represented taxa. We feel that such minority-linked diversity is one of the main interests of this work. It must be acknowledged, however, when comparing our work with previous accounts, that we intentionally cover all arts and all time periods, creating, ipso-facto, a “diversity bias” unlinked to our insects but instead related to our loose corpus selection methodology. One other bias should be mentioned, and it will also be treated as an aesthetic constraint: the size issue in its more general sense, or “the relative size of subjects” in its more technical declination: all insects included in this study (aphids and affiliates) are actually in the lower size range, even within the insect class. Therefore, their inclusion in artworks often induces severe constraints (how to draw *to scale*—the scale of a human, a flower, a canonical insect) that maybe interfere drastically with the artistic impact of such representations (how to say more with less). Such “art de la litote” (i.e., “art of understatement”) is actually consubstantial to our subject.

Although uncommon in the first accounts of human art, insects are represented in Paleolithic art, the earliest attested record to-date being that of the Chauvet cave in southern France, where a two-times six-legged animal may represent an ant-like double individual (Figure 1a, ≈31,000 years BP). The most famous ones, however, are the “the buprestid” of Arcy-sur-Cure (FRA; ≈15,000 years BP), the grasshopper of Enlène [13] (FRA; ≈13,000 years BP) and the stag fly larvae of Rosenberg-Klein Scheuer (GER; ≈14,000 years
BP). The latter two owe part of their celebrity to their morphological precision, it being
possible to describe their models at the genus level (Figure 1C,D). The stag fly is also of
high and rare interest in that it shows an immature insect, and it probably transcribes into
the persisting present-day feeding rituals of reindeer arctic communities [14].

<figure><img src="image_4.png" /><figcaption>**Figure 1.** Four examples of prehistoric insect representations, among which are the putative two earliest (A,B). (A): The Chauvet cave insect-like drawings [15] of the “red-signs panel”; (B): The Arcy-sur-Cure carved wood ground beetle (formerly “buprestid”); (C): the stone carving of stag fly maggot from Rosenberg-Klein Scheuer; (D): the Enlène grasshopper [13]. See text for dating and additional comments. © and credits: Adapted from https://archeologie.culture.gouv.fr/chau-vet/fr/salle-bauges/notice/panneau-signes-rouges (visited on 31/01/2023), Copyright 2023, Ministry of Culture, courtesy C. Fritz (A); photography YR of “buprestid” reproduction (Beetles: Extraordinary Insects Exhibit 2019, Lyon, Copyright Musée des Confluences and Ministry of culture; https://archeologie.culture.gouv.fr/fr/archeologie-fiche-site/arcy-sur-cure-et-de-saint-more, visited on 31/01/2023) (B); Wikipedia creative commons (C,D).</figcaption></figure>

Lastly, due to methodological reasons in corpus identification, a few words should
be added on the appearance of the nominal aphid/aphides. A thorough examination of
the etymology of both the word and the biological object (the family Aphididae) is clearly
beyond the scope of this article, and actually is “still-to-do”. It was coined by Linné in his
*Systema naturae* opus in Latin but from probable Greek roots [16]. However, it is unrelated
to both its Latin and Greek predecessors, which are Aristotle’s *ψυλλίδα/psyllida* {62} and
Pliny’s pulex, both with a great deal of taxonomic inaccuracy (an acceptable modern syn-
onym being, in lay language and for both terms, “tiny sucker”). This situation implies
that, before the seventeenth century, it remains very problematic to trace “the subject” of
our research with confidence. This is probably a very common situation for many insects,
but not for all: identifying bees or butterflies in ancient sources is not as problematic and
induces additional biases that may seem trivial but should be mentioned explicitly.

**2. Materials and Methods**

All occurrences of the corpus have been accumulated since 2017 in a relational data-
base comprising 17 fields that are exportable for content and structure and made freely
available at data.inrae.fr (https://doi.org/10.15454/Z3BOPV, accessed on 21/11/22).
Sources were from Google images and Google Art and Culture sections
(https://artsandculture.google.com, accessed on 21/11/22) searched with the following
items: *aphid(s)* (i.e., *aphid* and *aphids*), *puceron(s)*, *pulgon(es)*, *bladluis*, *blattlaus*. Occasion-
ally the (key)words **アブラムシ** (aburamushi, Japanese) and **حشرة المئة** (al-manna insect,
Arabic) were used. They were crossed with the keyword *art* (AND *art*), or one of the 8 art
category keywords (see below). In addition to those Google databases, we also systemat-
ically retrieved, with the same simple keyword strategy, the painting and art databases:
*joconde* (https://www.pop.culture.gouv.fr, accessed on 21/11/22), *rkd* (rkd.nl), *gallica*
(https://gallica.bnf.fr, accessed on 21/11/22), initial catalogue of *manuscrits enluminés*
(http://initiale.irht.cnrs.fr, accessed on 21/11/22), British library manuscripts and illum-
nated manuscripts bases (https://www.bl.uk/manuscripts; https://www.bl.uk/cata-
logues/illuminatedmanuscripts, accessed on 21/11/22), the scala archive
(http://www.scalarchives.com, accessed on 21/11/22), and the discog and imdb databases (https://www.discogs.com; https://www.imdb.com, accessed on 21/11/22) for music and movies. During our quest, some non-aphid items were collected and were marginally included in the database according to this simple color code: green were fully functional aphid occurrences; blue items were hemipteran quotes of equivalent interest for our work (e.g., see section on cochineal red); brown items were completing the same scheme for other insect taxa (i.e., responding to *classical* insect use in arts, see Introduction); finally, red items were rejected items (for example, when recognition of the insect required time and was ultimately negative) that were kept in the DB for tracing and sometimes for general artistic interest but not used either in the analysis or in the statistics. This strategy and categorization are the reasons why not all items selected are aphids; the status of *generic insect* or *unidentified insect* is likewise treated loosely. When insects could reasonably be treated as non-classical species, they might be classified as green/aphids (see Figures 4A, 9B and 10A) or even factually missing, (as in Figure 5C, for example).

With regards to paintings, several museums were visited and searched extensively in *praesentia* for target subjects (according to the genre and periods/countries, then selecting all still life paintings that included plants or flowers): the Pinacoteca Ambrosiana of Milan (Ita), The Museums of Fine Arts in Clermont-Ferrand (MARQ), Colmar, Dijon, Gray (Baron Martin museum), Lyon (http://collections.mba-lyon.fr and its gigapixel collection https://www.mba-lyon.fr/sites/mba/files/gigapixel/mignon/mignon.html, accessed on 21/11/22) and Paris (Le Louvre; https://collections.louvre.fr, accessed on 21/11/22). Additionally, for all graphical arts, several non-Google private databases were searched: pixel.com, fineartamerica.com, beaux-arts.com, alamy.com, getty.com, pinterest.com.

For the literature, searches included only French language sources and were based on Gallica at the French National library (gallica.bnf.fr, accessed on 21/11/22) and the lexicographic base Frantext (www.frantext.fr, accessed on 21/11/22).

Our database fields were either informative (e.g., author of artwork) or of statistical use (century, art category, etc.), or, finally, of analytical assistance, collecting symbolic, artistic or semantic ideas on the occurrence. They may be fully examined in the excel file deposited at https://doi.org/10.15454/Z3BOPV (accessed on 21/11/22). Art categories were basic and consisted of: Painting (graphical arts), Sculpture (3D art objects), Music (including lyrics/titles), Theatre (performing arts), Literature (poetry and all writing forms), Architecture, Cinema (including any dynamic 2D arts, except videos), Photography (all still 2D arts). Comics and Videos were included but did not reveal any significant items (note that "insect" musical videos were reviewed recently [17] but contained no mention of aphid or affiliate insects). Photography posed a specific boundary issue, since it revealed numerous potential items that were not selected due to both lack of space and lack of specific relevance to the subject. The literature, too, posed similar questions (not every [aphid] that was written about or photographed was taken as a "valid artistic occurrence"). It is, therefore, impossible to objectively (scientifically) delineate our corpus and, together with the ultra-vast source of potential occurrences, we have to accept these inevitable limits to the objectivity of our study. Corpus items are indicated in the present manuscript as bracketed integers {62}.

### 3. Results

Aphids are true insects of the hemimetabolous group, displaying incomplete metamorphosis, i.e., the juveniles resemble adults (in contrast to the holometabolous clade assembling Lepidoptera (*butterflies*), Coleoptera (*beetles*), Diptera (*flies*), etc.). Nevertheless, aphids also display a series of biological characteristics differing from even closely related species (within the Hemiptera order, for example, comprising cicadas, whiteflies, scales or psyllids). Such characteristics are listed in Table 1, and often constitute biological peculiarities (left column) warranting specific studies and establishing aphids as an alternative model insect species [18]. Two of these traits, feeding habits and virus transmission, are also responsible for the economic impact of aphids and, hence, their higher social
visibility. A partial measure of this is the scientific effort invested in aphids as a group: in 2020, and worldwide, aphids represented ≈10% of the insect bibliography per year, while the whole Hemiptera amounted to 18% (WoS®, values stable for the last five years). However, this scientific analysis of the impact of this insect family (Aphididae) is only part of the game, and our quest within the artists' universe should unravel a hidden part of that same game (human ideas). When choosing a subject, be it partial or of minor importance in the finished work, the artist is exerting an act of free will by which he or she says something about themselves, about their art and, finally, about the interaction of the subject (the insect, the aphid) with the human society of their time.

<table><caption>**Table 1.** Biological attributes of aphids among insects and other scientific model organisms.</caption><thead><tr><th>Main Attributes</th><th>Other</th></tr></thead><tbody><tr><td><ul><li>Aphid Life cycles:<ul><li>Host alternation</li><li>Sex polyphenism</li><li>Parthenogenesis <sup>1</sup>, clonality (cyclic parthenogenesis)</li></ul></li><li>Symbiosis: bacterial vertical transmission</li><li>Symbiosis: mutualism ant-aphid associations</li><li>High R (reproduction)/ r strategy organism <sup>2</sup></li><li>Piercing-sucking/ Plant sap parasitism</li><li>Virus transmission (plant)</li></ul></td><td><ul><li>Viviparity</li><li>Telescopic generations</li><li>Sociality in aphids</li><li>Defensive behaviors</li><li>Size</li><li>Gall formation</li><li>Wing polyphenism <sup>3</sup></li></ul></td></tr></tbody></table>

Examining the more than 250 elements under analysis, we will first give a very short statistical description of the corpus. As will be seen just below, in spite of our aphid-centered perspective, not all target items are describing an aphid (see methodology, with the associated color coding of aphids in green). Figure 2 splits our data into (i) insect types (aphids, green, accounting for more than 80% of the items), (ii) art category, and (iii) dating of artwork, grouped by century. A split graph of art x century shows that several more populated associations appear, with XVIIth century painting (the Dutch influence), the XIX century's literature (a methodological bias of the Gallica database) or a contemporary focus on graphical and plastic arts.

<figure><img src="image_5.png" /><figcaption>**Figure 2.** Statistical split of the corpus, between art types (main, left), time periods (main, right), insect types (far left inlet: aphids in green, Hemiptera in blue, other out-of-scope insects in red). The central graph is a heat-plot crossing the left and right categories (arts plotted vs. time).</figcaption></figure>

<footnote n="1">¹ parthenogenesis is the ability to reproduce as female-only lineages (for a hitherto sexual organism);</footnote>

<footnote n="2">² an r strategy organism shows both high growth rates/population sizes and a high variability in such population levels in time (antonym K strategy);</footnote>

<footnote n="3">³ polyphenism is the faculty to adopt distinct phenotypes with a single genotype (e.g., wing and no wing, sex and no sex).</footnote>
Before addressing the core of the aphid subject, we will illustrate how our quest was sometimes oriented towards the analysis of slightly out-of-corpus material (Figure 3). The Hemiptera constitute a family with biological features occasionally congruent with those of aphids (size, parasitism) but quite different from those of the canonical insects used in arts. “La chinche” (the bug, Figure 3A) is a remarkable account of the zoological interests of both scholars and the powerful during the Siglo de oro (manuscript dated ≈1570) and it is a perfect description of “a villain”, as we will see below with aphids. It may be noted that, from the textual description and name, the species is a *Cimex* sp. bedbug; however, its drawing is anything but precise (as seen with its green color), exemplifying that neither naturalism nor science had started to widely invest in insect subjects at that time (some examples, such as the Cocharelli codex, being exceptional in this respect; see later and Figure S1). The second example is even more emblematic, as it relates to one of the most materially important interactions of insects with the arts: the industry of cochineal red in the XVI century, and well before, is a perfect example of the “gift” contributed by (some) insects to artists at the expense of their small bodies [19,20]. Additionally, this example may be linked to another aphid species, the galls and honeydew of which were used in the confection of pigments for material dyes. A.F de Réaumur reported in 1737, in one of the first true entomological treatises (he also extensively wrote on aphids), that “the Turks were mixing two parts of “Bazgendges” (*Baizongia* sp) with one part of cochineal” to produce their red dye mixture [21]. It is noteworthy that the honeydew and the proper (and chemically very specific) pigments of many species of aphids and scales [19] have contributed to the general history of humanity to a degree which is probably not acknowledged properly in school textbooks (Figure 3B). One wonders what would have been human culture without the color red (although other reds, animal or mineral, did also contribute [22]). Moreover, what would have been human worship and spirituality without the biblical manna or Egyptian honey (i.e., insect-collected sugars)? Finally, and closer to our times, certain Homopteran insects did attract considerable artistic interest, as exemplified by the decorative strips drawn by the famous art nouveau artist Emile-Allain Séguy (Figure 3C). These assemblages were actually drawn from insect collections, in this case the Fulgoridae/lanternflies, with a daunting naturalistic precision. The size and colored phenotypes of these Hemiptera actually categorize them as closer to Lepidoptera in the inspiration palette used for insects in arts. They are, therefore, true outliers in the inspiration typology that will be developed later for aphids, and not really influential until recently [10] due to their mostly tropical distribution. Notably, the graphical precision of some representations (e.g., Figure 3C) might be qualified as being of “taxonomic grade” (although a fabric/wallpaper painter, the art-nouveau author EAS was also reported to paint from museum specimens) and may presently be the only representations of that particular insect species available on the web (the scientific database being free of digital specimens: see *Lycorma imperialis* in the Flow DB for fulgorids https://flow.hemiptera-databases.org/, accessed on 21/11/21).

<figure><img src="image_6.png" /></figure>

<figure><img src="image_7.png" /></figure>

<figure><img src="image_8.png" /></figure>
**Figure 3.** Remarkable relatives, non-aphid hemipteran: la Chinche, the cochineal red and Les Fulgores, from (A): the Chinche (bedbug) entry in the Bestiary of Don Juan of Austria; (B): Jose Antonio de Alzate y Ramirez, Memoria sobre la naturalesa cultivo y beneficion de la grana (Mexico City, 1777, Newberry Library, Chicago, Edward E. Ayer Manuscript Collection (Ayer MS1031)), quoted in Phipps, E., 2010. Cochineal red: the art history of a color. Metropolitan Museum of Art; Yale University Press; and, finally, (C): Emile-Allain Séguy's *Insectes* (1929) [046, 173, 141]. Personal photo of facsimile, original in monastery of Santa María de la Vid (Burgos) (A); cropped from published material © Newberry Library freed of rights (B) [19]; from web catalog, https://bibliotheques-spezialisees.paris.fr, free of rights, accessed on 10/10/2021, Copyright Ville de Paris Bibliothèque Forney (C) [23].

### 3.1. *The Size Issue*

The more obvious problem when dealing with aphids in the graphical arts is their realistic insertion into the composition. The problem of scale is challenging and requires specific treatment. This is the only group of arts with such a problem: sculpture, literature or music do not face the same technical obstacle when dealing with such tiny subjects.

#### 3.1.1. The Brueghel Aphids: Aesthetics of Details in Classical Painting

The Brueghelian aphid that appears in the "Bouquet of Flowers with Jewels, Coins and Shells" (1606, oil on copper, 65 x 45 cm; [2], {2}) was the first specimen in our corpus. Its presence in the Pinacoteca Ambrosiana is explained by the contacts made by the painter Jan Brueghel the Elder, known as "velvet", in Milan at the end of the sixteenth century. What do such flowers tell us, and what is this aphid doing, half-hidden in the dark? There is, in the painter's undertaking, an unmistakable taste for collecting: the bouquets bear witness to an extraordinary attention to the living, in its plant variations. Dozens of species of flowers rub shoulders, in a touch-and-feel that disregards both the rhythm of the seasons and the laws of gravity: if each petal is reproduced with the greatest care, if each element (flower, insect, small object) manifests, on the part of the artist, a concern for precision of a scientific nature, the composition itself depends on the pictorial choices in terms of color and form—in other words, the balance is above all aesthetic, and it would be difficult to recompose such architecture in a vase.

As for our other aphids (Figure 4), they seem to fit perfectly into the Brueghelian project of inventorying the world, especially the microcosm. However, the precision with which the insects are drawn is less exact than that of the first instance [2], with no possibility of species identification nor even the certainty that all insects can be identified as aphids (Figure 4A could be alternatively seen as a small stylized beetle). The entry of insects into art coincided with the invention of the category: until the end of the Middle Ages, little was known about "small animals", and snails, mice, mussels and flies were readily lumped together. It was not until the Renaissance that entomology was born as a discipline and the insect family appeared. It was also during the Renaissance that the fly was invited into paintings: Daniel Arasse [3] sees in this the emergence of modern artistic awareness, because the presence of a useless insect in places that are, to say the least, unusual (the cover of a book, the breast of Christ) makes painting visible as painting.

The same cannot be said of Brueghel's aphid: because it is part of a network, a constellation of insects, and because it appears in a natural environment—even if this nature is "dead"—it is more of an encyclopedic temptation. It can also be seen as an invitation to the viewer: come closer, come as close as possible, because the grain of this work rivals the grain of the world, and there is not even the smallest, the most insignificant, the most undesirable of the beings that inhabit this earth that I cannot accurately represent. The detail here expresses a double requirement and a double curiosity—entomological curiosity and curiosity about the possibilities of the pictorial gesture; a requirement of fidelity ("I paint as it is"), a requirement of exhaustiveness ("I paint because it is").
<figure><img src="image_9.png" /></figure>

**Figure 4.** The Brueghel aphids. Extraordinary repeated "aphid" occurrences among one of the most famous Dutch artist families, from Jan the Elder ("velvet Breughel", (A-C)) to Abraham the grandson (D); the relative sizes of aphids in the composition are so exceptionally small, as compared to other insects usually pictured in the Flemish workshops of that time, that it is difficult to believe that they are not both (i) related to each other and (ii) pursuing a specific intention in this very artist (family); (A): *the flower crown* by Jan Brueghel II the Younger (© Louvre Museum in Musée Baron Martin at Gray, Bourgogne FR; part of the MNR set), (B): flowers in a vase by Jan Brueghel I the Elder (© the royal museum of fine arts Antwerp BE, visible in augmented reality at Google arts and culture), (C): *bouquet in a clay vase* by Jan Brueghel I the Elder (© national gallery in London, UK; visible in HD at https://www.nationalgallery.org.uk/paintings/jan-brueghel-the-elder-bouquet-in-a-clay-vase); (D): flower garland, by Abraham Brueghel (© Galleria Nazionale d'Arte Antica, Roma IT) (190, 098, 104, 201). © Cropped in public photos from Le Louvre and others mentioned, with permission as *low-res* and *cropping* reproductions.

Other artwork of the Dutch XVIIth century represents aphids in our corpus, including a flower composition with the Holy Family, by Brueghel again, including many insects and an aphid in the crown {94}, clearly refuting the hypothesis of the exclusion of insects from religious subjects (Figure 4A and {94}). Other instances are by Abraham Mignon {101}, Ambrosius Bosschaert {218} and Bartholomeus van Winghen {219}. Even some negative items (items rejected as being aphids after close examination) merit scrutiny under our present analysis, as is the case of the artwork "La belle Carafe" (Abraham Mignon, the beautiful carafe, Le Louvre, {194}). An interesting feature of this piece is that it belonged to Louis the 14th, one of the most powerful kings of France, indicating, as noted above for the religious themes, that having power and owning or buying a work full of insects was not contradictory.

Finally, the punch line should be left to the artist himself, mirror in the mirror:

> "Il faudrait voir sur chaque objet que tout détail est aventure", translating as *We should see in each object that every detail is an adventure* (Guillevic, 1939. *Terrraqué*).

### 3.1.2. Si Parva Licet Componere Magnis (Virgil): Of Size and Men

This frequent motto in aphid science (literally: *If it be allowable to compare small things with great*) is difficult to track in the arts; it is, however, the source of a singular aesthetics, as we will see just below. One of the great specialists of aphids in the British XIXth century, GB Buckton [16], was citing a French scientist {202} and the quote summarizes well the situation where the small animal (and its science) is eager to compete with its bigger relatives, and of course, above all, Man.

Professor Huxley quotes the following words of the enthusiastic savant Duvau:
> "J'ai souvent pensé qu'on pourrait dans l'histoire des sciences désigner les époques par les principales découvertes. Par exemple, 1665 serait l'époque de la Gravitation, 1789 l'époque de la méthode naturelle en Botanique, et, "si parva licet componere magnis" les années 1740 à 1750 seraient l'époque des Pucerons."

(Translating as: "I have often thought that one could, in the history of science, designate the epochs by their principal discoveries. For example, 1665 would be the epoch of
*Gravitation*, 1789 *the epoch of the natural method in botany, and,*" si parva licet com-
ponere magnis " *the years 1740 to 1750 would be the age of aphids.*")

After such opinions, no apology will be needed for setting forth those discoveries
and ascertained facts which are connected with this division of the Hemiptera [16].

This raising of aphids to a level of interest competing with the "great subjects" of human intellect is present in some [16] instances of the collected corpus, mainly within the sections dealing with the science of aphids (3.3, {32}, etc.), or with the special reproduction of this insect family (3.2). This magnification trend is also present in Maurice Carême's poem devoted to an aphid ("an aphid was thinking", see later, 3.5) and it culminates in the entire short-story devoted to the nascent "humaphid" hybrid species (pucerhom) in "Des pucerons et des hommes", a children's novel by J.&C. Held {135}. The introduction sets up the argument of the story with "Trees, forests and people are worried and wondering: what exactly is the role of the aphid in the universe? (translation)".

It is also very tempting to recall here the aphorism of the philosopher Bachelard, in his *Poetics of Space* essay [24], which serves as the best example of the size paradox in the representation of aphids.

The minuscule, a narrow gate, opens up an entire world.

"Ainsi le minuscule, porte étroite s'il en est, ouvre un monde. Le détail d'une chose peut être le signe d'un monde nouveau, d'un monde qui comme tous les mondes, contient les attributs de la grandeur. Gaston Bachelard, la *Poétique de l'espace* English translation cited in Hugh Raffles' « *Insectopedia* » [25]"

There are only a couple of steps from philosophy to poetical aesthetics, often crossed
by haiku writers when sketching insects: the mirror effect renders the size paradox very
efficiently (Issa, with dragonflies below) and the night/day alternance obviates the invisibility of the tiny animals (Seishi, with ants). Intriguingly, only one translated Japanese haiku was found to quote an aphid, effectively playing with the modest size of the companion but in a burlesque rather than an aesthetical fashion, an otherwise common feature in haiku poetry. Nevertheless, we think that aphids have most likely been used more extensively in Japanese literature and painting and that the paucity of their occurrence here is probably a heavy language sampling bias.

The distant mountains
in the eyes
of the dragonfly
Issa

In daylight I saw
an ant
It haunts me tonight
Seishi

Slipping into my kimono
The snow aphid
Must feel very lonely
Murakoshi Kaseki

Cited in Coyaud M. 1978: *Shadowless Ants, the Book of Haiku* (Phebus publ., Paris, in French)

3.1.3. The Hidden World

Although being small is an ideal way of hiding, this is not a usual trend for aphids in
paintings: most examples in the corpus are "visible when noticed", which is not the case,
for example, for spiders, often hidden in dark fields of the artwork and just visible by
fragments, usually of striped abdomen and parts of the legs. Some appearances in paint-
ings (Bosschaert, {218}) seem to expose specimens in dark or faintly contrasted environ-
ments, leading to the trivial question: why draw and then hide "a character"? Hide-and-
seek is a common theme in painting, Arcimboldo being a master in that regard, but the
size and light parameters add an additional constraint for the reading of such an object,
and there is no simple answer to this aesthetic oddity. In modern photography, the hide-
and-seek characteristic is a renewed commonplace feature ("topos") of aphid representa-
tion, often related to color mimetism {31}.

3.1.4. The Untold and the Undrawn

Beyond dissimulation is the crossing of the barrier of traits and forms. Sometimes, no
recognizable detail is present, but the larger-scale aphid-object is clearly recognizable
(impressionistically inferred), running onto its rosy stem. With the help of light, the lower
limit of drawing is easily crossed, and many instances are encountered where small white
dots, reflections on a shiny abdomen, do represent aphids. This typically occurs in a wide
format "Stone Cartouche Surrounded by Flowers" {219} by Bartholomeus van Winghen,
where two falling roses, at the bottom side of the bouquet, support the frenetic march of
a colony.

Interestingly, it is possible to find a literary counterpart of these "unobservable/un-drawn" aphids, with the perspective aphid pictured by Victor Hugo {152} who uses "red and white aphids, which were a herd of oxen, were roaring in a meadow to the right (...) other blue and red aphids (...)" to represent the distant colored dots, barely visible but recognizable animals: oxen and ... men (blue and red are actually later described as soldiers in a distant field). In addition to the effect of distance and size, the use of the aphid metaphor renders the insignificance of all these living beings, set together in a single fate: aphids per se, oxen for food, and men for death.

From death to birth, aphids cycle in a singular way.

3.2. *The Reproductive Outlaw*

3.2.1. Reproduction and Parthenogenesis

One of the most exciting features in the scientific research on aphids is the presence
of cyclic parthenogenesis (Table 1), a trait featuring the alternation of sexual reproduction
(the norm) and asexual clonal reproduction (the outlaw). Biologically, this coincides with
host-plant alternation (the cycle between a perennial winter host and a deciduous sum-
mer host, usually an annual plant) and those two features, combined with viviparity,
probably gave aphids their main bio-evolutionary advantage over even close hemipteran
lineages. Although not a specific feature of aphids, this element of aphid biology is largely
acknowledged in a wide cultural context (61 languages in Wikipedia, more than 10 men-
tions of aphids in the English page on parthenogenesis, including proper identification of
the discoverer Bonnet). Parthenogenesis was discovered [26], but not named [27], by
Charles Bonnet, a Swiss scholar and follower of AF de Réaumur, who could be regarded
as one of the first aphidologists [21]. Bonnet deserves a specific place in this review, as he
acted both as a scientist and as an artist, drawing first and in precise detail the birth of an
aphid from its mother (Figure 5A; "Without Having Learned to Draw" [26]). The cited
book [26] is actually much more structured as a present-day scientific article, with a
strictly factual collection of results, a true material and methods section, a series of tables
of numbers and figure legends, and even an explicit mention of reproducibility issues
("My main purpose in publishing them has been to give others the opportunity to verify
them and push them further", preface xiv). In our opinion, the impact of this citation was
immense in the European Enlightenment: our Corpus registers 7 mentions of Bonnet, in-
cluding those by Voltaire {157}, Delacroix {80}, Buckton {202} and, later, Simone De Beau-
voir {66}). Delacroix's quote is interesting as being explicitly ironical, if not contemptuous
(ending with "What chapter to add to the one on the uselessness of scientists and espe-
cially of aphids" {80} relating, however, to the fact that Bonnet lost his sight because of his
close scrutiny of aphids). Beauvoir's reference (Figure 5B) to the work of Bonnet is indirect
(female clonality in animals), as is the mention by JJ Rousseau (Figure 5D; clonality in
botanics). Aphids are rallied in Beauvoir's "*The Second Sex*" as living examples of the pos-
sible uselessness of males in some animal societies, breaking down the myths of human
(masculinized) societies with facts from the biological world (*Facts and Myths* being the
 subtitle of her first volume).

Although statistically marginal in biology, the parthenogenetic (female)/viviparous
model of nature is ultimately quite a representative feature of the living world (mother
aphid/mother nature/mother earth), making it an emblematic outlaw. It may be retrieved
in documentaries or in movies, such as Richard Attenborough's *Virgin Births* {234} or in
the digital short *Aphids* (2010, {229}), where the trailer includes "Everything in nature is a
story within a story within a story", which perfectly fits the matriochka paradigm
illustrated in Figure 5C. This matriochka example points to another outstanding representation of maternity in the arts, namely the special representation of (virgin) Mary in the cult of Saint Anne (the mother of Mary). This most interesting topos in late medieval central Europe is characteristically depicting, in sculpture or painting, the three characters of Anne, mother of Mary, mother of Jesus, in a specific and non-realistic descending scale (Figure 1C). In the late period of this cult, the mother of Anne eventually completed the parentage, and the cult was named Anna Selbdritt (Ger), Anna Samotrzecia (Pol, {82}), Anna Meterza (Ita) or Anne Trinitaire (Fra, {1}) in different parts of Europe. This flourishing cult was judged to be too anarchic in the constitution of the holy family, and too female in its matrilineal genesis, finally resulting in its abolition at the Trento council in ≈1550 by redefinition of hyperdulia (marital cult) and replaced by the Joseph/Mary/Jesus holy family cult, conforming much more to the standard reproductive models. Although, of course, never linked to animal (or aphid) parthenogenesis by theological canons, the link between parthenogenesis (i.e., the reality of male-less reproduction in metazoa) and the Immaculate Conception appears to be legitimate. It is still a matter of cryptotheological debate, as exemplified by the chapter on human parthenogenesis in the Wikipedia entry [28]. Moreover, the parallel with aphid "matriochka-like" reproduction was introduced in an art blog, Quadrivia, a few years ago (Figure 1C, inlet), while the cult of Ste Anna was eminently documented by numerous scholars, including Annie Cloulas-Brousseau [29,30].

<figure><img src="image_10.png" /><figcaption>**Figure 5.** The reproductive outlaw. Aphids as a first documented example and paradigmatic model of animal parthenogenesis (and viviparity). (A): Charles Bonnet's own drawing of a laying virginous female, published in 1745 (note accompanying ladybird, a subliminal clue to aphid death following its birth?) [26], (B): Simone de Beauvoir's first citation on aphid reproduction in "The Second Sex" (Vol.1 *Facts and Myths*), (C): Anna Selbdritt depiction in sculpture form shows the three characteristic scales, from eye to shoulder doubled by red lines/dots. (D): Jean-Jacques Rousseau's *Botanics*; © personal copy (A) and detail photo in Warsaw Museum of Fine Arts; with permission (C), personal translation (B) and cropped in Gallica © Bnf (D).</figcaption></figure>

### 3.2.2. It Don't Mean a Thing, if It Ain't Got Those Wings [31]: Wing Polyphenism

What is the standard representation of an aphid in a layperson's mind? Very probably an insect with numerous progeny/colony (i.e., virginoparous morph), very probably green (i.e., leaf mimetic, although variant rose morphs are frequent), and possibly an insect with no wings (i.e., apterae morph). This feature list is typical of a source of variation which is called polyphenism: individuals with a single genome (clones) may give birth to variant morphs displaying drastically different properties:

(i) We have already cited *sexual dimorphism*, the ability to produce either females or males (which is totally absent from artistic representation, parthenogenesis having fully captured the representation).

(ii) *Color dimorphism*, actually under classical (mendelian) genetic control, is merely anecdotal, except regarding the mechanisms of genetic and metabolic control of color in aphids, which is linked to very specific pigments called aphins. That color morphs
do exist in aphids is related to the fact that they are able to synthesize and easily modulate their pigments' spectrum, a trait common in plants but rather sparsely distributed in animals, which are poor organic chemists. This trait has been treated already in the contribution of aphids to the materiality of different arts (aka the origin of pigments).

(iii) *Wing dimorphism*, which is the most frequently encountered polyphenism, is controlled by crowding and food quality. "Bad conditions induce migration" constitutes, simply drawn, the evolutionary constraint maintaining this biological trait. What about artists then? Voilà: nothing, or almost nothing. In our > 220 item corpus, we can count less than 10 winged aphids, most of them represented here (Figures 6A, 7A,D, 8A, 9B, 12A and S1C). Of course, large differences between art forms are to be expected, with literature underscoring such forms while photography should be more exact in the quantitative representation of wings in aphids.

Wings or not wings? Does it really matter? Probably yes, as this “option” is really linked to the artistic genre in action: photography more often figures alae aphids (Figure S1C), as is the realistic or scientific representation (Figures 6A and 7A,D), which is considered here as a true branch of the arts. In those contexts, aphids are represented as they are, with all their biological variety, and not as “we think” they are. Scientists, scientific illustrators and animal photographers are all artists with a natural science knowledge far beyond the average level, while most other artists develop an artistic project which is a personal projection first. While the latter group seems to look at aphids as exclusively wingless, and this is a majority rule for aphids, the former group is more flexible in its choice, and even tends to reformulate aphids within the standard boundaries of insect representation (and insects do have wings after all).

Therefore, for the majority of artists, we can state that *"those wings"* seem to be paradigmatic of insects, but that aphids are mostly seen as unwinged outliers. Again, formerly for size or for sex, and now for wings, aphids genuinely occupy a specific aesthetic niche.

### 3.3. Sapienza e Rinascimento

As time and knowledge progressed, so the intimacy and familiarity with the world of insects increased in the minds of artists. Starting more extensively in the XVth century in Europe [11], painters gradually represented insects, and part of their diversity, in their works. This trend was related to a new genre in Western painting, a subdivision of still lifes that adopted flowers, bouquets or vases as the object of interest and as the subject of the technical skills of an artist. This shift away from portrait, battle or landscape motives induced, in part, a correlated interest of the painters towards other living beings, i.e., other animals associated with the flower "ecosystem" (a term, of course, anachronic here): the entrance of flowers within the homes of artists and their influential sponsors was obviously accompanied by their arthropod companions [6,7], following a powerful hitchhiking movement.

#### 3.3.1. The Margins of Specimen Logic: Lilliput vs. Hercules

A return to classical antiquity and a return to nature is a coupled signature of an artistic renaissance, as opposed to medieval symbolism and the corset of the religious canon. In Holland, this artistic movement resulted in novel artistic genres [6], such as the flower declination within the still life. Representing animals also started to flourish in parallel with the birth of the curiosity cabinet—wunderkammer, studiolo, etc.—which allowed the animal specimen to enter the artist's workshop. While insects naturally followed this route, they ended up with emblematic artistic objects such as the *Hercules* stagbeetles of Dürer and Hoefnagel, which are among the best-known insect figurations in
history. The latter artist actually specialized and almost created a modern way to look at insects through a *specimen logic*, described by Janice Neri [8,32,33]. In this context, aphids are relatively rarely represented, except as a random taxonomic sampling (Figure 6A, green arrow). Interestingly, here, they are surrounded by their most common antagonistic mates—ladybugs, syrphids, etc. A further step in “ecological imagery” is given by the other insect portraitist of late renaissance Netherlands: Anna Maria Sibylla Merian [32]. This famous entomology painter did select aphid subjects in many instances [85,86,87], as a result not because of their taxonomic selection but due to their ecological importance, and are always accompanied by their trophic guild, most commonly hoverflies and lady-bugs (Figure 6B red arrows). As opposed to the deliberately impressive impact of Durer and Hoefnagel’s extreme specimens, the intention is obviously oriented towards the mi-nute and the pedagogic: care is directed not on the act and skill of painting (insect-object: Hercules) but on the insect-subject (Lilliput).

To terminate this parsing of aphids within the worldwide paintings of insects, we chose to group below two pictures that exemplify a frequent representation of aphids in their ecological context, either facing their predators (Figure 6C) or their symbiotic partners (ants, Figure 6D). This example (Figure 6C) is also a tribute to the treatment of natural subjects in Japanese culture (see also the haiku quotes above). We have not found any drawing of aphids in Japanese paintings so far, including in web-available samplings of Kitagawa’s “Crawling Creature” book [33]. We are convinced that this situation is due to heavy bias in localizing items on the web or in specialized and indexed databases. Japan’s naturalistic sensitivity is actually irrigating both arts and insects, as described specifically in a chapter of [5]. As illustrated by the puzzling popularity of the French entomologist JH Fabre [8,34] in Japan, insects are familiar to the majority. This is reflected in popular culture by the unique appearance of an aphid hero-character in Japanese web-art (Figure 6C right). Similarly, a major manga artist Osamu Tezuka is an entomology fan, as his self-portrait testifies (Figure C bottom right), and he invented a woman-insect heroine of fuzzy aphid/silverfish identity [35].

<figure><img src="image_11.png" /></figure>

**Figure 6.** The birth of naturalism. From early European "specialists" (Hoefnagel and Merian) to modern artists from very different horizons (social network artist, manga, educational pamphlet painting) and cultures (Japan, USA), aphids have always represented a "visible minority" subject, often associated with their ecological food web partners (ladybirds, syrphids, parasites or ants). (A). drawing from Joris Hoefnagel ≈1580 ("Ten Insects..." among which an aphid (green arrow) and many competitors (red arrows); from Washington DC Natl. Gallery of Arts and RKD database); (B): painting by Maria Sibylla Merian ≈ 1690 ("Red Columbine with Aphid..."; from St Petersburg Russian academy and RKD Database); (C): Aburamushi by web-artist Xquid © (https://www.deviart.com/xquid/ accessed 4/05/2020), and manga star artist Ozamu Tesuka, drawn as an entomologist by himself (bottom right inlet, ©Tezuka Productions) Tezuka's *Insect Woman* was interestingly analyzed as an aphid-silverfish hybrid [35]; (D): Insect-filled fragile ecosystems do include aphid imagery—with ant mates, top inlet with arrows—(Central Park biodiversity: color pencil, Barrett Klein 1998 © Reprinted/adapted with permission from BK [36]).

### 3.3.2. The Naturalist and the Scientific
Having mentioned Maria Sibilla Merian, precursor of this evolution, we logically move on to science and group together a series of representations of aphids in the joint context of science and art. It is art by methodology and science by objectives. It is also art through the effect it has on us. One may say that this movement encompasses naturalism and is not restricted to the graphical arts. The best example of this is the just-cited Jean-Henri Fabre. The last chapter (chapter X, 1903) of his major opus contains a long description of the pistachio aphid (76). Why it is art is obvious in the quotation below, and why it is science probably needs a bit more effort to comprehend.

It is on these plant miseries that the enormous cornicular edifices (i.e., galls) are founded; and still the leaf is not used in its entirety, but only one of its leaflets, in short, a point, a nothing. Exploited by the aphid, this nothingness acquires singular energy. First, it is welded to the tip of the twig and becomes one with it so that it remains on the tree when the leaves fall, and with them the other galls; then it provokes an influx of sap comparable to that of the pumpkin stalk feeding its pumpkin. The very small generates the huge...

Jean-Henri Fabre (1879–1907). Le puceron du térébinthe (*the pistachio aphid*), in *“Souvenirs entomologiques”* » (Entomological memories)

Apart from writing, the other main pedagogic art is that of drawing and making pictures. This art "section" is the principal provider of aphid imaging; however, we have only cited in our corpus a couple of representatives of aphid science paintings. The interest of these references is that the scientific authors are mentioned as well as the authors of the selected exquisite drawings (Figure 7), as exemplified by the work of R.E. Snodgrass (Figure 7B), whose first commitments were actually as an illustrator (his Wikipedia entry is well worth reading).

<figure><img src="image_12.png" /><figcaption>**Figure 7.** Science illustrated. Four time-scattered scientific representations of aphids (1745–present). (A): Charles Bonnet (publ. 1745), the father of aphid parthenogenesis (*Aphis solanella* Theobald, 1914, the spindle aphid (?), [26]). (B): George B. Buckton (publ. 1881), the grandfather of British aphid fauna (*Macrosiphum rosae* (Linnaeus, 1758), the rose aphid, [16]). (C): Robert Snodgrass (publ. 1935), the incredible author of Principles of Insect Morphology (*Aphis poni* De Geer, 1773, the apple aphid). (D): Keith Wheeler (*Microlophium carnosum* Buckton (1876), the nettle aphid); species names are the current accepted name of the insect (aphid species file: http://aphid.speciesfile.org, accessed on 21/11/22).</figcaption></figure>

### 3.3.3. The Microscopic Aphid

"Getting closer" is the usual means of "knowing better" in the naturalistic approach to the graphic arts, including photography. At every technical progressive step in the evolution of "optics sl", artists became involved in the appropriation and diffusion of novelties among a wide public audience (think of pointillism and the perception of color). Just before the invention of the microscope, the "limits of sight" were a well-known frontier for artists and writers. The "ciron" (acari/mite, a common cheese mite) was famous among artists and scholars, for example, Blaise Pascal and Victor Hugo (44), as the smallest visible living being. The latter author mixed mites and aphids in a religiously cosmic poem juxtaposing the smallest (arthropods and worms) and the largest (stars and suns) to celebrate
"the big one". With the advent of the microscope, many revisited this frontier and exemplified their findings with numerous specimens of aphids drawn through the magnifying glass, from the earliest Englishman Robert Hook (255; see Figure 4.8 reproduced in {Neri, 2008 #194} from R. Hook's "Micrographia"} through to German and, later, Japanese books and widely-distributed works (Figure 8 A,B). Other bursts of knowledge regarding the hitherto invisible also occurred later, with the advent of the electron microscope in the second half of the XXth century, revealing numerous, and slightly terrifying, visions of aphids and other insect heads. These were mostly exposed in "art" and photographic databases rather than in art exhibitions *per se* (21), but they represent an interesting analogy with a true artistic genre: the portrait. Actually, insects are almost absent in this genre, but they are seen, exceptionally, in modern art, for example, through a series of "en pied" paintings of many insects by Bernard Buffet (MARQ museum, Clermont-Ferrand). There are no aphids in this series, nor anywhere in our corpus, but there is a fly (254), out of the many variations painted by the same artist, which also included a dragonfly and many butterflies—typical insects represented in an atypical form.

Finally, we may cite here, within the global field of the microscope, many literary citations using the artifice of other optical instruments (telescope, Figure 8C) or aeronautical devices (meteorological balloons describing countless aphid swarms by the famous explorer William Beebe, Figure 8D), allowing the observation of minute insects in huge numbers and from a long distance. Summing up their impressions is an alternative way to say more about these unimpressive individuals.

<figure><img src="image_13.png" /></figure>

<figure><img src="image_14.png" /></figure>

<figure><img src="image_15.png" /></figure>

<figure><img src="image_16.png" /></figure>

**Figure 8.** The microscopic aphid. (A): Picture book for children (and later encyclopedia) from Friedrich Justin Bertuch (1792) {4}, a German industrialist; the page features at least five aphid specimens as well as magnified microscopic water arthropods; aphids are characterized as plant parasites with both puncturing stylets and trumpet-like cornicles, (B): Kenbikyo Mushi No Zu ("Illustrations of Microscopic Insects"), an illustrated treatise of insects drawn from microscope magnified views (Japan 1860) (6), (C): literary version of the microscope, working as a telescope. Famous quote from the encyclopaedist Diderot comparing humans to two-legged aphids, when observed from a separate universe; here quoted by Victor Hugo in *La Légende des siècles* (1877) "... from where we can see this globe on which we live so proudly, big at most as a pumpkin, and from where we might observe, through a telescope, the infinite multitude of gaits of the two-legged aphids we call men." (D): Mention of aphids in *Raffles Insectopedia* [25], as scientifically-attested swarms of microscopic insects (204,212,213).

### 3.3.4. The Decorative Aphid Exception

Usually, aphids are not considered for their aesthetic value, in contrast to the archetypal beautiful insects: butterflies. The use of aphids, or similar lay insects, in the decorative arts (jewellery, ceramics, furniture, etc.) would be unexpected. For this very reason, their occurrence is interesting and subject to different kinds of driving forces. Scale insects were found in native American *Mimbres* pottery (Figure 9A), together with a vast array of animal representations (not shown). This probably results from cultural sensitivity to natural subjects, an ecologically-driven taste of naturalism. The choice of this "species" may be compared with the knowledge related to cochineal red production in this area (Figure 3B), but further historical analysis is needed to make the link between the two items. Other
occurrences of aphids or non-canonical insects in the decorative arts are presented in Fig-
ure 9, with a peculiar figure of a single and lonely insect representing newly acquired
"free citizenship" after the French Revolution (Figure 9B). This insect is often figured to-
gether with one, or a few, "worker citizens" en route to their new life {208).

Other rare examples are those featured on XIXth century vessels from central/north-
ern French factories, which introduced oriental-like motives, including insects frequently
seen in Japanese or Chinese landscape paintings. Figure 9C presents both conventional
(butterflies) and non-conventional insects. Finally, the decorative usage of insects in phi-
lately is common, but not normally aphids. Figure 9D is a good illustration of an excep-
tion, with a nice birch aphid photograph, which is probably one of the more efficient
means of presenting aphid morphology to a large public (although it is restricted to Ice-
land).

<figure><img src="image_17.png" /></figure>

**Figure 9.** The (rare) decorative aphids. (A): Like many animals, sap-sucking Hemiptera, probably scale insects {105}, were used in decorative pottery of the *Mimbres* classical era (1000–1130), Mogollon culture of northern Mexico. Copyright permission granted by JL Zeddam and Peabody museum. (B): During the late French revolution, a rare though recurrent motif of Limoges ceramic vessels figures a single (undetermined) insect of small size (human scale) frequently associated with freedom and revolutionary symbols {208}; in this instance, the priest on the right declaims "I announce the happiness of France". (C): "Saladin-Levesque" plate featuring two uncommon insect motives among the three outer ones (in a common three animals and three plants design); red arrow indicates a topical insect of oriental design, the green arrow a putative aphid or rare occurrence in this context, potentially recognized by its cornicular appendices {223}. B,C copyright through the Joconde database. (D): Birkis-protalùs, i.e., the downy birch aphid *Euceraphis punctipennis*, on an Icelandic stamp (Nordic insect series).

Concluding this section is the realization that aphids, as with all other insects, are
biological objects differing profoundly from what is empirically known and felt about hu-
man societies and their domestic animal associates (mostly mammals). Thus, throughout
the preceding section(s), insects are perceived through the scope of human knowledge,
including its technological trappings (microscopes, foreign journeys or long and painful
drawing sessions, for example). In the subsequent sections most representations are
based, in contrast, on the feelings and artistic emotions generated by aphids, either nega-
tive (3.4) or positive (3.5).

3.4. *The Evil*

Aphids are fundamentally and ecologically plant parasites. From the advent of neo-
lithic culture, most humans have been primarily consumers of cultivated plant products.
It is, therefore, not surprising that a dominant representation of these insects is as that of
an evil thief of human labor. The villainous aphid is a recurrent figure used by the French
writer Victor Hugo {53, 153}, as illustrated in the Figure 10C poem, but it is also apparent
through the use of serial metaphysical metaphors by the Russian author Chekhov.

Die Blattlaus vernichtet die Pflanzen, der Rost Metall und die Lüge die Siele

(*the aphid destructs plants, rust metal, and lie does so with the soul*)
<figure><img src="image_18.png" /></figure>

**Figure 11.** Aphids and performing arts. Three records (© Discog) and a web-theatre using aphids or aphid-related elements in the title: (A): vinyl single dated 1976 (Schwamp, Aphids In The Hall, CA, USA), (B): Hum's Aphids album dated 1998 (Champaign, IL, USA), (C): Theatre blog entitled "Café des pucerons" (Aphids Café; Twitter and Blogspot {57}), (D): digital record by Polish artist Duy Ge-bord—anagram of situationist Guy Debord—entitled Mildew, including a series of aphid titles: *pineapple gall adelgid, plant sap, aphid and balsam woolly adelgid* {64}.

### 3.5. *The Good, Rebirth of the Small*

#### 3.5.1. Empathy for the Evil

The XXIst century introduced a clear shift in the perception of insects, including aphids (formerly considered as a pest species). This shift probably started in the popular graphic arts, whose artists started to pay attention to all kind of animals. Through different approaches, such as children's illustrations (Figure 12A) or the traditional fine arts (Figure 12B), aphids were regarded as desirable, funny or interesting partner species.

In literature, as well, examples of affectionate glances are available here and there (*Pierre Senges*, see below), in sharp contrast to the works produced over the years and centuries before.

Si fragiles signifiait sensibles, on aimerait imaginer des pucerons des bois profondément affectés par des nuances, celles du matin, celles du soir, attentifs et douillets à la manière du petit Marcel Proust: observateurs parce que toujours malades, la crainte du rhume rendant particulièrement réceptif au moindre brin d'air—des insectes qui ont choisi, somme toute, de ne pas mener des existences de brutes. *Fragments de Lichtenberg*, Pierre Senges 2008. Translation: «If fragile means sensitive, we might picture tiny insects (aphids in original text) that are profoundly affected by the least nuance, such as the nuances of morning or the nuances of evening, as attentive and cozy to these details as little Marcel Proust: they observe everything because they're always sick, their fear of catching a cold makes them particularly sensitive to the slightest breeze—insects that have decided, in the end, not to lead the lives of simple brutes»*. (Fragments of Lichtenberg, transl.: Gregory Flanders) [37].

<figure><img src="image_19.png" /></figure>

**Figure 12.** (re-)Birth of the cool: “coolness” as the new aphid paradigm. Contrasting with late XXth century (Figure 11 note plurals and no direct imaging), aphids are now regarded both as individuals and as cool guys. (A,C). Goppel's *Anna Aphid* (2005), children's cartoon book {11}, (B): Trijnie
Goudriaan's *Bladluizen* fragment (2003) with permission www.trijniegoudriaan.nl, (C): T. Langham's aphid cartoon (2015), (D): Christy Rupp's parasitized aphid sculpture with permission (2002).

### 3.5.2. The Aphid Gang Is Cool

Pushing the picture one step further, situations of predation and parasitism are depicted as completely non-conflictual, organizing peaceful gatherings of aphids with loving and friendly ladybirds (Figure 12C; prudent aphids staying out-of-reach though). Alternatively, aphids are depicted as the victims of deadly parasites (Figure 12D) and, overall, the ecology of aphids is presented to the young general public in a very different context from that used by M.S. Merian (Figure 6), although ideologically and pedagogically quite similar.

### 3.5.3. The Lone Revolutionary Insect

Another context showing a high degree of empathy with a "small unnamed insect" (that we sometimes considered as aphids when they had no other possible taxonomic identification) is apparent and worth mentioning. We only discovered this topos in the context of post-revolutionary ceramic productions (Figure 9B): a clear revolutionary context related to factory workers or to converted priests (preaching revolution) and exposing a single flying insect that both accompanies the main heroic character and illustrates its lonely and tiny (humble) status. This representation was not frequent but sufficiently repeated—see the three occurrences in {208}—to be highly symbolic of the times and their glorification of the underprivileged, literally speaking.

### 3.5.4. The Theological Aphid: From Lesser to Maurice Carême

From empathy to sanctification {207; Christian Lesser, 1742 "*Theology of Insects or Demonstration of the Perfections of God*"}, the way was straightforward. One instance of global empathy towards insects is given by a Parisian theatre play of the late XIXth century where all the characters were attributed to insects (Figure 2A: The King Carrot, 1872 {165}). The play was a success, thanks to both the music (Offenbach) and the costumes (Thomas, inspired by the famous zoomorphic illustrator Grandville), and it was brilliantly analyzed in [38]. The aphid character (another topos of French XIXth century theatre) is obviously depicted with empathy and as a quiet dew-producing animal, in contrast to the neighboring noisy bumblebee, for example (Figure 2A right and left). The archetypal empathetic depiction of aphids comes later, however, with a poem dedicated to the humble insect by M. Carême, deploying the fully anthropomorphic figure of the man = aphid and its generous and invisible creator(s) (Figure 12B). Maurice Carême, in "Le Puceron", seems to unburden his text of any virulence: it is not a question of blaming man for his presumption but rather of unfolding a joyfully incongruous hypothesis, a "what if"—what if the aphid were to meditate? If there is a slight criticism of anthropocentrism, it is there in the background, like a reminiscence or a wink. The heart of the poem lies elsewhere, in the speculative tranquillity. The form, which borrows from the fable (the very first lines are reminiscent of Jean de La Fontaine's incipits), allows for this deployment. There is no narrative as such, as the overall movement is dedicated to the thoughts of the protagonist. By a fair return (i.e., by a new analogical transfer), we are invited to take advantage of this wisdom and to accept the relative character of our judgements. Ladybirds, aphids, chances or plagues, it all depends on the perspective adopted, it all depends on the enunciation. Who speaks, who goes there? What remains of this poem, what settles in, what inhabits the readers, is a voice: Carême attributes to the aphid a sensitivity, a rhythm, an attention of its own. Although not a priori the ideal candidate, and perhaps precisely because it is not, our aphid becomes a delicate knot: a form of tenderness is at work, a trend we find, with more or less surprise, in other texts in our corpus (notably in Pierre Senges' *Fragments de Lichtenberg*; see above, § 3.5).
One may group together, within this empathetic vision, two opposite representations of aphids, either as an endangered species saved by opportune geometric constraints on their two-spotted predators (Figure 12C) or as ironically dangerous predators “loose in the garden” of the exquisite Gary “caustic” Carlson (Figure 13D).

<figure><img src="image_20.png" /></figure>

**Figure 13.** The humble (and pious) aphid. The small is fundamentally harmless: bless (or make fun of) the tiny. (A): Aphid character and costume in an all-insect theatre play in XIXth century Paris (1872, an Offenbach-Sardou Opéra Bouffe: *The King Carrot*, [165]); (B): Maurice Carême dedicated poem: *The Aphid* (publ. 2014). (C): Harper’s *The Last Aphid* (1981); (D): Gary Larson’s *Henry! Aphids...* (date unknown), probably a discrete tribute to Dr Seuss’ “*Quick, Henry, a Flit*”.... [184, see WikiMedia’s entry for cartoonist Theodor Seuss Geisel] see https://thefarside.net/i/165457f0 (accessed on 25 09 2022). $$ (© farside asked again, as for Carême’s poem -because it is a complete citation).

Looking backwards through Figure 13 as a whole may give a heteroclite vision of this *empathic* evolution, but the sense of a dissimulated smile that prevails after careful scrutiny of the four citations seems to us a real indicator of the unity of feelings provoked by such artistic creations, typical of some modernity initiated in the XIXth century. We sense it, therefore, it must be true...

## 4. Conclusions

Arriving at the end of this essay, a first impression concerning the representation of aphids and associated hemipterans is that they develop within a universe of much less constrained symbolic boundaries than many more “conventional” insects, despite the rarity of their occurrences. Related to this, their specific attributes (Table 1) are actually not universally known, and their three main features comprise either trivial patterns (**size**, piercing-sucking **parasitism**) or not (**parthenogenesis** and possibly viviparity). These characteristics do imprint their relationship on mankind, starting “from the beginning” (the two first features) or from the late seventeenth century, where the knowledge of their peculiar and impressive biology percolated through from scientific personalities to artists and the educated, and even the general public.

In these two regards (the minoritarian and the historical), one may argue that aphids are, at least partly, attached to the modern section of the arts. Could we state this in contrast to butterflies, for example, which are not modern subjects for art? In our opinion, this is most visible in the last “pot-pourri” (*aka* unclassifiable) collection of aphid representations, such as those appearing in supplementary Figure S1. In this last compilation, (Figure S1 A–H), our target animals do illustrate, in turn, colonialism, suicide behavior and post-apocalyptic heroic fantasy; the reader should normally be able to recognize each one.

However, other features of their relationship to humans may also be qualified as classical or mainstream. This is the case with their inability to influence the subconscious realms of the human mind: as far as can be found through specific studies on the subject, no aphids have ever been reported in human dreams (at least as appears in the extravert manifestations of “insect dreams” in movies or literature [39]). The “tiny suckers” *psylla* and *pulex* of the classics, therefore, still have a whole continent and a harsh ecosystem to conquer.
In composite homage to Duke Ellington and to Gary Larson (Once a Cartoonist, For-
ever a Wingless Insect).

**Supplementary Materials:** The following supporting information can be downloaded at:
www.mdpi.com/xxx/s1, Supplementary Figure S1: The special aphids.

**Author Contributions:** Conceptualization, Y.R. and S.O.; methodology, Y.R.; corpus investigation and resources, S.O. and Y.R.; data curation, Y.R.; writing—original draft preparation, Y.R.; writing—review and editing, Y.R. and S.O.; visualization, Y.R. All authors have read and agreed to the published version of the manuscript. Terms after the CRediT taxonomy.

**Funding:** This research received no external funding for completion but was funded by ANR, grant number ANR-21-CE20-0001-01(ArtiCute) for the APC.

**Institutional Review Board Statement:** *Not applicable.*

**Informed Consent Statement:** *Not applicable.*

**Data Availability Statement:** Data upon which the analysis was drawn are made available through the institutional INRAE (agronomy research France) data repository at data.inrae.fr. See doi and more details (versions) are in Appendix section below.

**Acknowledgments:** Warmest regards to Duke Ellington for his valued contribution to the art and science of jazz and swing, anagram of the most glorious invention of insects. Edwige Keller-Rahbé is warmly thanked for her contributions to this and earlier versions of this work. We are much indebted to the INRAE (Plant Health Dept) special interest group BAPOA (aka aphid biology group, including associated organisms), who contributed to the elaboration of the studied corpus, in particular, but not exclusively: Baptiste Monsion, Charles-Antoine Dedryver, Denis Tagu, Estelle Postic, Etienne Herrbach, Jean-Louis Zeddam, Jean-Luc Gatti, Marilyne Uzest, Nathalie Boissot, Gérard Febvay and Sophie Muller. We also thank Delphine Montagne, Marcel Dicke and Barrett Klein for their contributions and for interesting correspondence on the topics of insects and arts. We thank art history colleagues for their kind answers to our requests, among which Claudia Rabel (CNRS Orléans), $. We also thank Valérie James for her English language revision of the penultimate version of the manuscript.

**Conflicts of Interest:** The authors declare no conflict of interest.

## Appendix

The original database containing all the corpus' elements is available as two files, one pdf and one excel file, on INRAE's institutional data repository (data.inrae.fr). Version 1 corresponds to this paper's submission process/publication. The previous and further version will be available upon actualization of the base. https://doi.org/10.15454/Z3BOPV.

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