Early Portuguese natural history collections regarded Brazil as one of its main interests, influenced by the kingdom’s political authorities and naturalists’ own fascination. Two unique and almost identical copies of the same work on Brazilian butterflies, “Lepidoptera Fluminensis”, with specimens predating Brazil’s independence, are held today at the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa (MUHNAC), in Portugal, and in the private collection of António Bonchristiano in Brazil. Both books were prepared using a poorly known and now extinct technique, lepidochromy, sometimes labeled as “nature printing”. Here we present a concise history on lepidochromy, as well as the history of these books from conception until the present day, including their conservation status, and a taxonomic review of their content. We suggest that the authorship of the books should be attributed to two authors, Friar José Mariano da Conceição Veloso and Father Francisco Solano, while previous works had only identified the former as the possible author. As the authors used Linnaeus’ “Systema Naturae” as a major reference for the species’ identification in “Lepidoptera Fluminensis”, here we provide a taxonomic update for all species based on current classification of Lepidoptera. Originally regarded as having representatives of 169 different taxa, “Lepidoptera Fluminensis” represents specimens of 145 taxa currently recognized, from nine families. We also present a brief note on the importance of Friar Veloso’s work as an early scholar of Brazil’s natural history and the relevance of “Lepidoptera Fluminensis ”as both an artwork and a naturalistic endeavor.

Eighteenth century, history of natural history, Natural History collections, naturalists, Specimen preparation techniques

Modern natural history studies are one of the most important outcomes of the Enlightenment period. During the eighteenth century new methods, approaches and techniques were developed to help the study of the surrounding nature and to contribute to the ongoing imperial agendas of most European nations. While many of these methods and techniques are the same as those commonly used today, such as taxidermy/skin preparations, fluid preservation, herborization of plant specimens, or pinning of insects, other rarer techniques appear odd to modern-day scientists, such as the case of “fish-herbaria” (Bauchot 1976; Péquignot 2006; Ceríaco and Marques 2011; Ceríaco et al. 2023), lepidochromy (Cowan 1968; Orousset 2008; Péru 2016; Mandrij 2024), or the preservation of butterflies between mica sheets (Salmon et al. 2000; Romano 2023). In the second half of the eighteenth century, different preservation methods were adopted and implemented by naturalists working in the Portuguese Empire, especially in the Brazilian territory, which was then the most important Portuguese colonial possession and one of the main interests of both naturalists and political authorities (Ceríaco 2021). Due to such entanglement of political and scientific interests, the birth of Portuguese and Brazilian natural history studies is deeply connected. These entanglements also led to a considerable movement of people, knowledge and collections between Portugal and Brazil. While the number of extant specimens in Portuguese and Brazilian collections that predate Brazilian independence (1822) is considerably small (LMPC, unpublished data) some of the surviving specimens are particularly interesting. That is the case of two copies of a book on Brazilian butterflies entitled “Lepidoptera Fluminensis”, prepared using the lepidochromy technique and currently held by the historical archives of the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa (MUHNAC/AHMB) and in the private collection of António Bonchristiano in São Paulo, São Paulo, Brazil, herein the “Lisbon copy” and “Bonchristiano copy”, respectively. Interestingly, this preparation technique has never been mentioned in any of the “naturalist instructions” published in Portugal from the eighteenth century to late twentieth century (Ceríaco and Marques 2025). This rare book represents the oldest available work on Brazilian Lepidoptera and possibly one of the few survivors of an eighteenth century scientific work using the lepidochromy technique in the world.

The first known uses of this technique are those of the Dutch painter and naturalist Otto Marseus van Schrieck (1613–1678), who pressed butterfly wings on the canvas of some of his naturalist artistic paintings (Mandrij 2021). The British artist John Peele (1716–1770) in his “The art of drawing and painting in water-colours…” presented a small chapter on the “Manner of making the impressions of any butterfly, in a minute in all their colours” (Peele 1735). The scales, and consequently the color and patterns of the butterfly wings, would then become part of the painting, at a time when reproducing the color and brightness of butterfly wings was almost impossible by other means. The technique was sometimes labelled as “nature printing”. However, the first reference to the use of such technique for scientific purposes was presented by the British naturalist George Edwards (1694–1773), who in the last pages of the second part of his book “A natural history of birds” (Edwards 1747) briefly explained the technique. The same author would make reference to the technique again in the book “Essays upon Natural History and other miscellaneous subjects”, where he presented an essay titled “A receipt for taking the figures of butterflies on thin gummed paper” (Edwards 1770). The publication of Edwards’ (1770) “receipt” was shortly followed by another publication, this time by the French clergyman and naturalist Jean-Baptist-Gabriel-Alexandre Rozier (1743–1824) titled “Manière de fixer sur le papier les ailes des papillons, et de les représenter au naturel” (Rozier 1771). Orousset (2008) attributes to Rozier the priority of the description of the method, even if Rozier’s paper was published one year later than the “receipt” by Edwards. Rozier attributes the discovery of this technique to an unnamed French friar who observed that, after a butterfly had landed on a recently varnished barometer, its scales were fixed to the varnish. Both Rozier’s “technique” and this anecdotal story were later mentioned in the article about butterflies on the fifth tome of the Panckoucke’s “Encyclopédie méthodique” (see Péru 2016; for a detailed description of the contents of the “Encyclopédie” see Evenhuis 2003) . Still in the eighteenth century, other European authors would reference similar techniques, such as Ernst and Engramelle (1779) and Köster (1793). The library of the Natural History Museum of London has a small collection of lepidochromy from the English Moses Harris (1730–1788) which dates circa 1766, while the library of the Museo Nacional de Ciencias Naturales in Madrid also has a manuscript entitled “Collectio insectorum pictorum divisa in genera et species iuxta Car. Dom. Carolum Linaeum comparate cum ordine metamorphosean” by the Dutch naturalist Johannes le Francq van Berkhey (1729–1812), representing some of the earliest known examples of the technique still extant.

In the early nineteenth century, the technique continued to be used in Europe (Orousset 2008; Péru 2016), with authors like the French entomologist Jean-Baptiste Godart (1775–1825: 280–281) dedicating some pages of his book “Histoire naturelle des Lépidoptères ou Papillons de France” to explain the “Manière d’imprimer les Papillons” (Godart 1821). As shown by Péru (2004; 2016), the technique gained considerable popularity in France in the second half of the nineteenth century, with several contributions being published on the topic (Poulin 1876, 1899; Watilliaux 1879 [see Orousset 2008]; Sériziat 1892). It was also at this time that the technique became known as lepidochromy. Some nineteenth-century specimens prepared using lepidochromy are still extant, such as the 292 specimens prepared by Just Veillat (1813–1866) and reported by Péru (2004), the specimens prepared by Pierre Chrétien (1846–1934) which are still housed in the entomological collections of the Muséum national d’Histoire naturelle, Paris, or even the tomes of some works such as “Les Papillons” by Depuiset (1877), “Les Papillons de France ”by Rothschild (1880) and the “Tableaux analytiques des Lépidoptères de la faune franco-rhénane” by André and Lucas (1899–1916) (Orousset 2008; Péru 2016). This seems also to be the case with the collection of 66 nature prints made from 39 butterflies collected in the Shire Valley (East Africa) by English Anglican missionary Horace Waller (1833–1896), currently in the collections of the Smithsonian Institution, Washington D.C.

In the early twentieth century, several works relying on the use of lepidochromy were still being published around the world, such as the case of the two volumes “As Nature shows them. Moths and Butterflies of the United States East of the Rocky Mountains” by the North American naturalist Sherman Foote Denton (1856–1937; Denton 1900). Denton’s (1900) frontispiece proudly mentions that the work is illustrated “With over 400 photographic illustrations in the text and many transfers of species from life”. Other late nineteenth to early twentieth century examples of the use of lepidochromy for scientific purposes are the work of the Martinican naturalist Théophile Raymond (date of birth unknown–1922) on Venezuelan butterflies (González et al. 2021) or “The Pressed Specimens of Butterflies and Moths”, a work by the Japanese author Yasushi Nawa (1857–1926). However, during the course of the twentieth century, the technique became increasingly rarer and was eventually abandoned by entomologists. One of the latest known works that resorted to lepidochromy was that of British entomologists Lionel Gilbert Ollyett “L.G.O.” Woodhouse (1888–1965) and George Morrison Reid Henry (1891–1983), who published a book on the butterfly fauna of Ceylon (Woodhouse and Henry 1942). From then on, the technique seems to have vanished from common practice and only the recent interest in the topic, brought about by several historians of art and science, has resurrected the topic (Ceríaco 2021; González et al. 2021; Mandrij 2021, 2024).

Besides its rarity, the “oddity” of the lepidochromy preparations, and the fact that it may constitute one of the oldest entomological works dealing with South American lepidopterans, the history of “Lepidoptera Fluminensis” is still not properly known. This work was completely forgotten by entomologists and historians of science until the dawn of the twenty-first century. Faria (1999) was, to our knowledge, the first to mention this work, attributing its authorship to the Brazilian friar José Mariano da Conceição Veloso (1742–1811). However, Faria (1999) did not know the whereabouts of the work, and assumed it remained unpublished. Pataca (2006) alluded to the presence of the Lisbon copy in MUHNAC, also identifying it as a work authored by Veloso. Independently and at around the same time of the works of Pataca, during the research for his PhD thesis, the first author also located the Lisbon copy in MUHNAC, and provided some comments about the technique and the authorship, also attributing it to Veloso (Ceríaco 2014). Felismino (2014) provided a brief account on the Lisbon copy in a catalogue of the historical books of MUHNAC. The most recent iteration about the history of “Lepidoptera Fluminensis” was provided by Costa et al. (2019), who agreed with the previous authors, also attributing its authorship to Veloso. In the meantime, a second copy of the book surfaced in an auction held at the New York Antiquarian Book Fair in April 2013 by the hands of the natural history specialized bookseller Antiquariaat Junk from Amsterdam, with an auction price of 117.000 USD (Schierenberg 2013). By chance, the first author was attending the fair and saw the copy, and in a brief glimpse was able to see that it was mostly identical to that of the copy held in MUHNAC. The copy ended up being sold to the Brazilian collector Antonio Bonchristiano (1967–present). Due to its rarity and scientific importance, here we provide a detailed description of both copies, a history of its creation and authors, as well as a taxonomic identification of all the represented taxa.

The books are similar to each other, but with a few noteworthy differences both in their external format/binding and contents. The Lisbon copy has a luxurious reddish leather binding and gold engravings in its front cover depicting the Portuguese crown (Fig. 1).

Figure 1. 

Red-leather cover of the Lisbon copy. Source: AHMB Res. 3.

It measures 22 × 17.5 cm and has 172 pages depicting different Lepidoptera taxa. Each of the taxa is classified following the Linnean system and depicted in both dorsal and ventral views. As attested by ex-libris stamps, the book was originally from the library of the Royal Cabinet of Natural History of Ajuda, the most important and active natural history collection in eighteenth-century Portugal, where all of the specimens collected in the then extensive Portuguese overseas empire were deposited to be studied by a team of naturalists (Ceríaco 2021). The book, together with hundreds of other specimens, herbaria, books, and manuscripts, was “requested” and transported to the Muséum national d’Histoire naturelle in Paris by the French naturalist Étienne Geoffroy Saint-Hilaire (1772–1844) when he visited the Ajuda collections in 1808 during the French invasions of Portugal, but was returned to Lisbon in 1814 (Bocage 1862; Daget and Saldanha 1989). The frontispiece of the book presents a lavishly illustrated dedication (Fig. 2) to Queen Maria I (1734–1816) signed solely with an “S.” (most likely Francisco Solano, see below), with the following Latin inscription:

Figure 2. 

Frontispiece of the Lisbon copy, with a dedication to Queen Maria I. Source: AHMB Res. 3.

“Omnium rerum naturalium in Brasilia creatarum Dominæ Divæ Lusitanorum Augustæ MARIÆ I.^æ Lepidopterarum suorum Omne pulchrum Præfectura Fluminensis S.”

[Translation: Of all natural things created in Brazil, to the Lady Diva of the Lusitanians, the August Maria I, all the beauty of her butterflies of the Fluminense state, S]

The following page of the manuscript (Fig. 3) bears an adapted quote from page 109 of John Ray’s (1710) “Historia Insectorum”:

Figure 3. 

First page of the Lisbon copy (after the frontispiece) with a passage from John Ray’s Historia Insectorum. Source: AHMB Res. 3 / PT-MUL-RMJBA-TC-02.

“Quæri fortasse à nonnullis potest, Quis Papilionum usus sit? Respondeo, ad ornatum Universi, e ut hominibus spectaculo sint: Ad rura illustrandæ velut tot bracteae inservientes. Quis enim eximiam earum pulchritudinem e varietatem contemplans mira voluptate non afficiatur? Quis tot colorum et artifici penicillo depictas curiosis oculis intuens, divinae artis vestigia eis impressa non agnoscat et miretur? Ray. Histor. Insect. de Papil. pag. 109.”

[Translation: Some can however ask: what is the utility of butterflies? I answer that they adorn the Universe and as a spectacle for men. To adorn the fields as leaves of gold. Who can contemplate their eximious beauty and variety without being impressed with admirable passion? Who, looking with curious eyes at the elegance of so many colors from these sketches of nature, devised by nature’s own genius, and painted by the artist’s brush, does not acknowledge and wonder at the traces of the divine art imprinted upon them?].

The passage is a eulogy of the beauty and utility of butterflies and an oath to natural theology, a philosophical trend that defended that the study of nature was a way to praise God (Ceríaco and Brigola 2014). Comparing it to Ray’s original quote, the author cut a segment, namely “… et schematum elegantias naturae ipsius ingenio excogitatas...” between “Quis to colorum” and “et artifici…”. After these introductory passages, a total of 172 plates with butterflies and moths are represented at 1:1 size. The first six plates present larger animals, with their respective dorsal and ventral views on different plates, but all subsequent pages present smaller animals and have both their views depicted in the same plate. Given the nature of the technique required the use of a complete animal for each illustration, a total of 338 lepidopterans are thus depicted in the book. While the wings are prepared as lepidochromy, the bodies are painted in watercolor (Fig. 4). As noted by Costa et al. (2019), plates 43, 45, 72 and 115 are an exception, as the wings are also painted and don’t have traces of scales. The species are classified according to Linnaeus’ (1758) 10^th edition of the “Systema Naturae” which divided them into the following classes: Equites Troiani (14 species), Equites Achivi (17 species), Heliconii (24), Danai Candidi (31 species), Danai Festivi (8 species), Nymphales Gemmati (16 species), Nymphales Phalerati (36 species), Plebei Urbicolae (8 species) and Plebei Rurales (14 species). The taxonomic coverage and implications of these collections will be further explored in the Taxonomic accounts section below. Similarly to the frontispiece page, all the pages depicting lepidopterans are bordered by a line of red ink and gold leaf (Fig. 5).

Figure 4. 

Magnified views of the wings and body parts of some individuals represented in the Lisbon copy. Note the wing scales in the wings and surrounding areas. A, B 1 – Papilio Equites Troes Dardanus [= Morpho menelaus coeruleus (Perry, 1810)]; C, D 55 – Papilio Heliconii Pindo [= Eueides vibilia vibilia (Godart, 1819)]; E 49 – Papilio Heliconii Saphus [= species ignota]. Photos by Martim Baptista.

Figure 5. 

Gold leaf and red paint in the frame of one of the book’s pages. Photo by João Ferrand de Almeida.

The Bonchristiano copy is overall similar to the Lisbon one with only a few slight differences. While the Lisbon copy depicts the Portuguese coat of arms in the cover, this copy depicts what is assumed to be the coat of arms of Brazil’s Viceroy (Fig. 6). The frontispiece is also different, as instead of being dedicated to Queen Maria I, it is dedicated to the Viceroy of Brazil, Luis Vasconcelos de Sousa, and the explicit indication that the book was offered by Father Francisco Solano (Fig. 7). The frontispiece bears the following dedication:

Figure 6. 

Red-leather cover of the Bonchristiano copy.

Figure 7. 

Frontispiece of the Bonchristiano copy, with a dedication to the Viceroy of Brazil.

“Ill^mo ac Proest^mo Pro-Regi Immortali Vasconcellio, Per manus Fr. Francisci Solano erga Claustru patria que ejus beneficioru memoris omne suum pulchrum in tesseram gratitudinis Lepidoptera Fluminensia O. D. D.”

[Translation: To the immortal Vasconcelos, distinguished and notable Viceroy. By the hands of Father Francisco Solano, representing the Cloister and the Motherland, remembering its benefits, and everything that within is beautiful. As a testimony of gratitude. Lepidoptera Fluminensia. Offers, Devout, Dedicate].

Unlike the Lisbon copy, the following page has a quote in French from Rozier (1771) (Fig. 8), which reads:

Figure 8. 

First page of the Bonchristiano copy (after the frontispiece) with a passage from Rozier (1771).

“Ceux-là, fiers de leur être, et d’un eclat nouveau s’élancent dans les airs, sortant de leur tombeau. Un Dieu, de ses bienfaits prodigue sans mesure, semble avoir pris plaisir à former leur parure, Comme si la beauté ce don si précieux, Ainsi qu’à nos regards, étoit cher à ses yeux. Leurs armes, leurs combats, leurs travaux, leur génie, Le courage des uns, des autres l’industrie, Leurs moeurs, leurs passions, leurs amours, leurs plaisirs, Pourroient seuls occuper mes utiles loisirs.”

[Translation: Those, proud of their being, with a new burst soar into the air, coming out of their tomb. A God, of his prodigal benefits without measure, seems to have taken pleasure in forming their adornment, As if beauty this precious gift, As well as to our eyes, were dear to his eyes. Their weapons, their fights, their labors, their genius, The courage of some, of others industry, Their morals, their passions, their loves, their pleasures, Can alone occupy my useful leisure.].

This page is followed by an additional one which is exactly the same as that of the Lisbon copy presenting John Ray’s citation. After these introductory passages, and similarly to the Lisbon copy, a total of 172 different lepidopteran plates are included, with specimens represented at 1:1 size. Larger animals present dorsal and ventral views on different pages, but smaller animals have both their views depicted on the same page. The species are classified according to Linnaeus’ (1758) 10^th edition of the “Systema Naturae” classification, divided then into the following classes: Equites Troiani (14 species), Equites Achivi (17 species), Heliconii (24 species), Danai Candidi (31 species), Danai Festivi (8 species), Nymphales Gemmati (16 species), Nymphales Phalerati (36 species), Plebei Urbicolae (8 species) and Plebei Rurales (14 species). There are some differences in the species used between the two versions, as well as the sex represented, which will be further described in the Taxonomic diversity section (see below).

Interestingly, and unlike the Lisbon copy, Bonchristiano’s copy had a more complex custodial history. The book appeared in April 2013 at the New York Antiquarian Book fair and was later acquired by Antonio Bonchristiano. It is assumed that the book was originally offered to the Viceroy, but its whereabouts between that offer and 2013 are unknown, as well as how it came into the hands of the bookseller.

Overall, the Lisbon copy presents a relatively good conservation state. However, some of the pages show a few conservation problems around the edges, due to the different characteristics of the materials used and the environmental conditions to which the book has been subjected over the decades. In the areas where the gold leaf is applied, it is possible to see that some pages are torn and fractured (Fig. 9), especially in the inner area of the pages (the one closest to the book’s spine), an area where more tension is applied to the paper when it is opened. These tears and fractures may have occurred due to a combination of several factors: in order for the gold leaf to be applied to the paper, an adhesive must be used first. The adhesive utilized was probably a natural one, like hide glue, fish glue, egg white, gesso or gum Arabic. These glues (with the exception of gum arabic) are strong but not very flexible, and just as the gold leaf, they don’t react to variations in humidity. Unlike these materials, paper reacts greatly to variations in humidity, shrinking and stretching. The difference between the materials’ reactions to variations in relative humidity meant that the paper ended up breaking in this area. In addition, on the pages where these edges are in the worst condition, it is possible to see small black spots, both in the areas where the gold leaf would have been (which already has gaps and is incomplete) (Fig. 10) and on the reverse side of the sheet, where these spots are in a line on the area where the gold leaf is applied on the opposite side (Fig. 11). These small black spots are a sign of biological attack by fungi, which prove the differences in relative humidity to which the book has been subjected over the years, and may also have contributed to the weakening of this area of the paper, leading to its breakage. As well as feeding on the paper, these fungi may have been feeding on the glue used to fix the gold leaf.

Figure 9. 

Tears and fractures in the gold leaf frame of different pages. Photo by Sofia Perestrelo.

Figure 10. 

Gaps in the gold leaf of the frame. Photo by Sofia Perestrelo.

Figure 11. 

Gaps and traces of biological attack by fungus in the frame area of the pages. Photo by Sofia Perestrelo.

For these reasons, and to prevent damage from progressing or worsening, it is important that the book be stored in a place with temperatures between 16°C and 19°C and relative humidity between 45% and 60%, avoiding fluctuations in these parameters (Elkin and Norris 2019). This way, we can avoid the presence of microorganisms and prevent the different materials from reacting to environmental conditions, so that the pages remain stable.

The authorship of the book has been attributed to the Brazilian friar José Mariano da Conceição Veloso (1742–1811) by several authors (Faria 1999; Pataca 2006; Ceríaco 2014, 2021; Felismino 2014; Costa et al. 2019; Ceríaco et al. 2023) although Bocage (1862) did not note any authorship. Veloso is considered one of the forefathers of Brazilian natural history studies, especially due to his works “Flora Fluminensis” (Bediaga and Lima 2015) and “Ichthyologia Fluminensis” (Ceríaco et al. 2023). Born in São José del-Rei, state of Minas Gerais, on 14 October 1742, Veloso initiated his ecclesiastic career in the convent of São Boaventura de Macucu (today Itaboraí, Rio de Janeiro state) in 1761, moving to the convent of Santo Antônio in Rio de Janeiro four years later to study philosophy and theology (Nunes and Brigola 1999). In 1771, Veloso was sent from Rio de Janeiro to the São Paulo convent as a missionary, and from there instructed to teach geometry to the soldiers stationed in the indigenous village of São Miguel, nowadays São Paulo (Bediaga and Lima 2015). Still in São Paulo, in July 1781, Veloso was ordered by Martim Lopes Lopo de Saldanha (birth and death dates unknown), then governor of the captaincy of São Paulo, to collect natural history specimens to be shipped to Rio de Janeiro, and from there to Lisbon (Bediaga and Lima 2015). After returning to Rio de Janeiro in the early 1780s, Veloso was elected “Master” (Professor) of the natural history classes being held in the Santo Antônio Convent in 1786 (Gama 1869). There, Veloso started to amass a rich and diverse collection of natural history specimens and an herbarium (Nunes and Brigola 1999). Given his experience and interest in the study of Brazilian natural history, in 1783 Veloso was hired by Luis Vasconcelos e Sousa (1742–1809), viceroy of Brazil from 1779 to 1790, to conduct the first official botanical survey of the state of Rio de Janeiro and collect natural history specimens to be shipped to Lisbon (Gama 1869; Nunes and Brigola 1999). For seven years (1783–1790), Veloso conducted botanical and zoological expeditions in the state of Rio accompanied by two other friars, Anastacio de Santa Ignez (birth and death dates unknown), who was responsible for penning down the description of the collected plants, and Francisco Solano (1743–1818), acting as illustrator and preparator, among a larger team of military and other logistical personnel (Bediaga and Lima 2015; Pataca 2019; Ceríaco et al. 2023).

During these expeditions, Veloso and his team collected thousands of botanical and zoological specimens that were prepared and then sent to Lisbon (Nunes and Brigola 1999; Ceríaco 2021; Ceríaco et al. 2023). These collections included the herbaria and illustrations that gave birth to the “Flora Fluminensis”, only published posthumously in 1825 (Carauta 1969), as well as the fish-herbaria which were the base of the never published “Ichthyologia Fluminensis” (Ceríaco et al. 2023). According to Bediaga and Lima (2015), not much is known about the itineraries taken by Veloso and his team during his collecting trips between 1783 and 1790, but these appear to have been circumscribed to the areas between present-day Rio de Janeiro and São Paulo, with references to Ilha Grande, Santos, Serra do Mar, Paraíba River valley, Paranapiacaba mountains and Paraty (Pastore et al. 2021).

Pastore et al. (2021) attempted to correlate the terminology used by Veloso in the “Flora Fluminensis” with geographical localities. Several toponyms such as Pagum Cunha [=Cunha], Praedium Boavista [=Church of São José da Boa Vista], Regnii Praedii Sanctae Crucis [=Santa Cruz], Pharmacopolitanis [=Paraty], etc., or the terms “mediterraneis”, “campis”, “apricis”, “transalpinis”, “silvis”, “maritimis”, etc., used to describe the vegetation, were combined with the phenological data provided along with the descriptions (in months of the year). It can be concluded that the term “mediterraneis” used by Veloso probably refers to the captaincy of São Paulo, and more specifically the municipality of Cunha, state of São Paulo. The term “campis apricis” was probably used to describe typical Cerrado vegetation (possibly the first descriptive term for this type of vegetation in Brazil). The term “transalpinis” is another term for the municipality of Cunha, and would designate the region after the top of the mountain on the way from Paraty to Cunha, the border area between the states of São Paulo and Rio de Janeiro, known locally as the turn of the mountain. The terms “silvis” and “maritimis” were generally used to describe a typical Atlantic Forest landscape, with dense forests along the Brazilian coast.

In 1790, Veloso left Brazil for Portugal, accompanying Luis Vasconcelos de Sousa, who had finished his duties as viceroy. Veloso brought to Lisbon all his natural history collections to be deposited in the collections of the Royal Cabinet of Natural History of Ajuda. The relationship between Veloso and the Ajuda cabinet was already well established, as many of the collections he amassed since the mid-1770s were sent there. In a report dated from 1797 and provided by Veloso to Rodrigo de Sousa Coutinho (1755–1812), at the time the acting prime minister of Portugal, the friar lists some of the most important collections brought with him to Lisbon and delivered to the Royal Cabinet of Ajuda:

“... eleven volumes in folio, of which two with descriptions, one collection of all the fresh and saltwater fishes prepared using a new method, as also another with marine and terrestrial insects, and finally, one collection of butterflies pressed by the colored dye with which the membranes of their wings are covered, a work so rare and valuable that the supplicant [authors’ observation: Veloso himself] attests that no such work exists in any other royal cabinets in Europe: everything is now in the Your Highness museum, where this was delivered as soon as himself arrived”.

Original manuscript in Arquivo Histórico Ultramarino (AHU), Lisbon, Portugal (AHU-CU-Reino, Cx 30, Pasta 7), transcribed in Nunes and Brigola (1999), Ceríaco (2021) and translated in Ceríaco et al. (2023).

The eleven volumes in folio are certainly the manuscripts and illustrations of the “Flora Fluminensis”, whereas the collections of fishes and marine and terrestrial insects “prepared using a new method” refer to fish herbaria specimens recently studied by Ceríaco et al. (2023). The collection of butterflies refers to the Lisbon copy of the “Lepidoptera Fluminensis”. This is further attested by the fact that in a complete inventory of the Royal Cabinet of Ajuda collections, compiled by the then interim director Alexandre Rodrigues Ferreira (1756–1815) in 1794, the book is listed under the entomology section of the library as “Vellozo / Frei José Marianno da Conceição Vellozo / Lepidoptera fluminensis (…) 1 Vol. in 1°” (Ferreira 1794).

Given such dates, the butterfly book had to be prepared before 1790, and it is safe to assume that it partly resulted from the expeditions made by Veloso and his team in the Rio de Janeiro and São Paulo regions. The citation of Rozier (1771) also shows that the book had to be prepared after 1771. However, the emergence of the Bonchristiano copy has brought additional elements that seem to contradict the general assumption of Veloso’s sole authorship of “Lepidoptera Fluminensis” (Ceríaco 2014, 2021; Felismino 2014; Costa et al. 2019; Ceríaco et al. 2023). While Ferreira’s (1794) inventory of the Royal Cabinet of Ajuda identifies Veloso as the author of the book, and Veloso’s report to Rodrigo de Sousa Coutinho establishes a direct link between himself and “Lepidoptera Fluminensis”, the Bonchristiano copy frontispiece explicitly states that the book was presented to the Viceroy by Francisco Solano (see Fig. 5), not mentioning Veloso at any point. Given that both copies are almost identical, it is safe to assume that they had the same author. There is, however, a vague reference to Solano in the Lisbon copy frontispiece. The eulogy to Queen Maria I ends up with a single “S.” (see Fig. 2). Costa et al. (2019) interpreted this “S” as an abbreviation of “Salve”, an effusive eulogy, but it is most likely just an abbreviation of “Solano”. The same authors excluded Solano from the list of potential authors of “Lepidoptera Fluminensis” given that “the historiography only associates him with the botanical drawings of “Florae Fluminensis” and because the floral decoration of the frontispiece reflects a lack of experience” (Costa et al. 2019). It is important to note that Costa et al. (2019) did not know about the existence of the Bonchristiano copy, and were thus unaware of the presence of Solano’s name in the frontispiece of that copy. In light of these apparent contradicting evidences regarding the authorship of “Lepidoptera Fluminensis”, it is likely that the work was the result of a collaboration between both Veloso and Solano, the first being responsible for the scientific direction and taxonomic identification of the species, while the latter was the main person responsible for the lepidochromy preparations and overall illustrations on the book.

Both books present a taxonomic organization that follows Linnaeus’ organization of the order Lepidoptera in the 10^th edition of “Systema Naturae” (Linnaeus 1758), dividing the butterflies (Papilio) species into five main groups and corresponding subdivisions, namely Equites Troiani (18 plates), Equites Achivi (17 plates), Heliconii (26 plates), Danai Candidi (31 plates), Danai Festivi (8 plates), Nymphales Gemmati (16 plates), Nymphales Phalerati (36 plates), Plebei Urbicolae (8 plates) and Plebei Rurales (14 plates). Each of the divisions begins with a cover page, and then the taxonomic arrangement is always presented at the top of each plate as a string of taxonomic categories (see for example plate 1, “Lepidoptera Papilio Equites Troes [=Troiani] Dardanus”, Fig. 13C, D), starting in the broadest categories – Ins.[ecta] – and ending with the specific name – Dardanus – in a very similar way as to how taxonomic accounts were presented in Linnaeus’ works.

Linnaeus (1758) taxonomic arrangement of Lepidoptera divided it into three main groups, namely Papilio, Sphinx and Phalaena. The Papilio were characterized by having the “antennae pointed towards the heavier bodies, often club-headed”, with “Wings (of the sitting ones) erect and converging upwards”, and “flying during daytime”. Sphinx were those with the “antennae thicker in the middle and tapered at both ends, subprismatic”, with “wings deflected”, and “flying mostly during the evening”. Finally, the Phalaena having “the setaceous antennae gradually tapering from the base to the tip”, the “wings often depressed when the animal is sitting”, and “flying at night”. Papilio was further divided into six subsequent groups (“phalanges”), which themselves were internally divided (Fig. 12). Thus Papilio was divided into the Equites, which were characterized by “the wings are longer at the tips than at the base; the antennae are often threadlike”, with the subdivision Equites Trojani (“chest with red spots (often black)”) and Equites Achivi (“unstained chest, an ocellus at the anal angle [of the hind wings]”); the Heliconii with “narrow, very straight wings: elongated forewings; very short hindwings”; the Danai, with “the most perfect wings” and its subdivisions Danai Candidi (“with white wings”) and Danai Festivi (“with colorful wings”); the Nymphales with their “jagged wings” divided into Nymphales Gemmati (“with spotted wings”) and Nymphales Phalerati (“with spotless wings”); the Plebeji, those in which “the larva is often contracted”, further divided in Plebeji Rurales (“with wings marked by dark spots”) and Plebeji Urbicolae (“wings often spotted with translucent spots”) and, finally, the Barbari, which were simply “added as a corollary, not related to the orders”. Linnaeus (1758) Papilio contained a total of 192 different species (7 Plebeji Urbicolae, 13 Danai Festivi, 15 Heliconii, 17 Plebeji Rurales, 17 Equites Trojani, 19 Danai Candidi, 23 Equites Achivi, 23 Nymphales Gemmati, 24 Barbari, and 34 Nymphales Phalerati. All groupings are present in “Lepidoptera Fluminensis”, except for Barbari.

Figure 12. 

Organization of Papilio as proposed by Linnaeus (1758).

Figure 13. 

A, B Equites Troiani frontispiece; C, D plate 1 – Equites Troes Dardanus [= Morpho menelaus coeruleus (Perry, 1810)]; E, F plate 2 – Equites Troes Dardanus [= Morpho menelaus coeruleus (Perry, 1810)], from the Lisbon and Bonchristiano copy respectively.

Except for the first three plates, those depicting P. Equites Troes Dardanus (Fig. 13, plate 1 and 2, female individuals of Morpho menelaus coeruleus (Perry, 1810)), P. Equites Troes Archilocus (Fig. 14, plate 3 and 4, male individuals of Morpho menelaus coeruleus (Perry, 1810)) and P. Equites Troes Acamas (Fig. 14E, F, 15A, B, plate 5 and 6, male individuals of Morpho hercules (Dalman, 1823)), the authors classified each plate with a different Latin name, and it is safe to assume that of the 172 plates, they recognized 169 different taxa. This is a slight overestimation of the actual number of represented taxa, as in several cases the male and female individuals of the same species were understood as different taxa by the authors and given a different Latin name (see Table 1 and Taxonomic diversity section below).

Figure 14. 

A, B plate 3 – Equites Troes Archilocus [= Morpho menelaus coeruleus (Perry, 1810)]; C, D plate 4 – Equites Troes Archilocus [= Morpho menelaus coeruleus (Perry, 1810)]; E, F plate 5 – Equites Troes Acamas [= Iphimedea hercules hercules (Dalman, 1823)]; from the Lisbon and Bonchristiano copy respectively.

Figure 15. 

A, B plate 6 – Equites Troes Acamas [= Morpho hercules hercules (Dalman, 1823)]; C plate 7 – Equites Troes Pandarus [= Parides agavus (Drury, 1782)], D plate 7 – Equites Troes Pandarus [= Parides bunichus bunichus (Hübner, [1821])]; E plate 8 – Equites Troes Adrastus [= Parides anchises nephalion (Godart, 1819)], F plate 8 – Equites Troes Adrastus [= Eurytides (Mimoides) lysithous rurik (Eschscholtz, 1821)]; from the Lisbon and Bonchristiano copy respectively.