Delpinoa, n.s. 46 : 85-93. 2004

Federico Delpino's scientific thought and the birth of modern biology in Europe

1 2 G. A LIOTTA , A. A LIOTTA

1Dipartimento di Scienze della Vita, Seconda Università degli Studi di Napoli, Via Vivaldi 43, 81100 Caserta, ; 2Facoltà di Medicina e Chirurgia, Seconda Università degli Studi di Napoli, Via Arena 4, 81100 Caserta, Italy. [email protected]

Abstract. The authors discuss the figure of Riassunto. Gli autori presentano la figura di Federico Delpino (1833-1905), an outstanding Federico Delpino (1833-1905), illustre esponente Italian botanist of the XIX century, focusing on della botanica italiana dell'Ottocento, evidenzian - his cooperation with the major scientists of his do i suoi rapporti scientifici con altre eminenti time, among whom . Moreover, figure del suo tempo, tra cui Charles Darwin. they underline the crucial influence that Federico Inoltre, evidenziano l'influenza che il pensiero Delpino’s scientific thought and innovative stud - scientifico e gli studi innovativi di Federico ies on floral biology and plant systematic had Delpino, rivolti specialmente alla biologia fiorale e over the development of botanical sciences in the alla sistematica vegetale, hanno avuto sullo svilup - XIX century and on the birth of modern plant po delle scienze botaniche nell'Ottocento e sulla biology. nascita della moderna biologia vegetale. Key words : Charles Darwin, Federico Delpino, History of biology, Plant biology,

INTRODUCTION Jacques Rousseau’s books and those by illu - minist philosophers showed a renewed interest History of biology is marked by the in nature. It was a century characterized by longevity of its issues and it commands respect collections of plants, animals and minerals. because of the enormous diversity of life, in was still linked to medicine, thus terms of space (all the continents), time (since European physicians-naturalists mainly 3,8 billions of years ago), dimension (ranging searched for therapeutic properties of plants, from viruses to whales) and habitat (air, soil, particularly of those introduced after the great water). For centuries, biological phenomena voyages of exploration. Naturalists also have been classified in two main scientific dis - focused on classification of plants, following ciplines: medicine and natural history, corre - criteria that could facilitate their recognition sponding to modern natural sciences. (M AYR 1990) . The Greeks had already praised harmony of The Swedish scientist Charles Linnaeus nature and many of the current scientific issues gave the most important contribution to this (ex: embryology, systematic) were known to field. He proposed a new classification of Aristotle, who stood as a point of reference plants, according to a logic division, based on until the scientific revolution of the XVI cen - the presence or absence of the flower and on its tury. Since then, the western civilisation has characteristics. Linnaeus developed his con - been studying the rules of physics, but no other cept of species affirming that: we count as natural aspect was as reluctant to reveal its many species as the different shapes created by rules as diversity of organisms. God. Then, as other scientists of his time, he The XVIII century was the golden age of thought that the number of living things had natural history, that was enriched by the histor - been established at the moment of Creation ical voyages of James Cook, Louis Antoine de and their classification would reveal the divine Bouganville and Philibert Commerson. Jean pattern of creation. There was a dispute in France about the consider the cultural references, the method he validity of the Linnaean System that, having a applied and his cultural heritage. rigid scheme and being based on floral charac - ters, placed species in different classes, although they were similar, according to other ACADEMIC BIOGRAPHY characters (for example, a group such as gramineous plants was divided in seven class - Federico Delpino was born on December es). Moreover, observations made by relevant 27, 1833 in Chiavari, in the outskirts of , scientists of that period arose doubts about an Italy. He had a weak health and he was com - immutable number of species. This debate pelled by his mother, since early childhood, to involved scientists such as the French Georges spend most of the time outdoor, in a still exist - Louis Buffon, Bernard de Jussieu and Jean ing small garden, that became the place of his Baptiste Lamarck, the Germans Joseph early observations on swarms of ants and Gottlieb Kolreuter, Johann Wolfgang Goethe insects surrounding the flowers. After the High and Gottfried R. Treviranus, the English School at Chiavari, he attended the first Course Joseph Banks, president of the London Royal of Mathematics at University of Genoa, but he Academic Society and the Italian Domenico quit after the first year. Cirillo, physician and botanist at University of Referring to his childhood and youth, in , relevant personality of the Neapolitan 1864 Delpino wrote a note on a botany text - Enlightment and president of the Legislative book: “Studium vegetabilium puer meditabar Commission for the Neapolitan Republic of inconscius, adolescens adgrediebar ardentis - 1799 ( ALIOTTA & A LIOTTA 2001). sime. Sortes adversae me ad aliena rapuerunt.” The botanist Treviranus and the botanist- (“As a child, I unconsciously thought about zoologist Lamarck, independently from each studying plants. As a young man, I was deeply other, introduced the term “biology” in two fascinated by it. Fate brought me to different papers, both published in 1802. The first meant things.”). biology as philosophy of living nature, while After the disappointment for the experience the latter referred to a wider acception, beyond at the University, his parents allowed him a morphology and systematics, typical of natural long journey to the far east, where he collected history, rather concerning the study of func - plants, especially in the Dardanelli strait. tional processes of organisms with an holistic When he got back, his honesty drove him to view of nature ( TREVIRANUS 1802-1822; find a job, so that he could be economically LAMARCK 1802) . In this scenario, at the begin - independent from the family. He found a job at ning of XIX century, there were outstanding the custom office of the Ministry of Finance, in biological discoveries: the cellular theory, the Turin. Despite his weaknesses, Delpino could origin of species, sexual reproduction of find the time and the way to deepen his knowl - plants, hereditariness of characters. edge about literature and philosophy and to Federico Delpino is the brightest figure of commit himself to the study of plants. the Italian botany in the XIX century. The most In 1865, when the capital city of the king - innovative aspect of Delpino’s scientific dom was transferred from Turin to Florence, research is the institution of a new discipline, Delpino, apart from his duties, could live in a plant biology, that should collect and describe place suitable to his ambitions; he got access to relations established among living things, the resources of the Florence Botanical Garden among plants and between plants and animals, and Botanical Museum; he could also read the in order to understand functions such as repro - books of the Webbian library. In 1867, profes - duction, dissemination, survival and self sor proposed Delpino as an defence (B ORZÌ 1905; C ATALANO 1955) . adjunct professor, because he had appreciated Our aim is to introduce him with a brief his skills; so, Delpino officially started his biography, illustrating his activity in the scien - career. Four years later, in 1871, he became tific context of his time. In particular, we will professor of natural sciences at Vallombrosa

86 Royal Institute of Forestry, but he basically place of honour in the history of pollination, taught botany. While he was there, Delpino along with Camerarius, Koelreuter, Sprengel conceived the idea of a voyage around the and Darwin. This history began in 1694, when world, so he boarded the warship “Garibaldi” the German botanist CAMERARIUS (1694) as a naturalist. The ship had been set up for the coined the term and showed that pollination of educational voyage of the prince Tommaso di the stamen is necessary to the formation of Savoia. Unfortunately, Delpino was forced to seeds. Thus, stamen and pistils are “the sexual quit the circumnavigation because the ship organs of the flower”. These studies were not lacked basic scientific equipment. He returned continued until 1761, when Koelreuter discov - to homeland after visiting Brazil and collecting ered the different types of pollination, under - plants in the surroundings of Rio de Janeiro. At lining the role of insects and dichogamy, the the end of 1875, after a concourse, Federico floral structures and strategies that prevent Delpino became full professor of botany at self-pollination ( KOELREUTER 1761). University of Genoa, where he continued his In 1793, Konrad SPRENGEL , in his work studies on floral biology and published a book “The secret taken from nature about structure about phyllotaxis (DELPINO 1883). In 1884, he and pollination of flowers”, described accu - moved to the University of Bologna and his rately the pollination by insects and fields of interest were myrmecophilous struc - dichogamy, demonstrating the luring function tures and systematics. of corolla, that floral structure favours cross- Ten years later, he moved to Naples where pollination in certain plants, and the mutual he continued researches about pollination, the adaptation of the flowers and of their pollina - key point in his scientific work, which defines tors. His conclusion was: “…it seems that the field of interest for plant biology. In 1899 nature prevents flowers from self-pollination”. he started the journal “Il Bollettino dell’Orto Sprengel’s results were ignored until 1862, Botanico della Real Università di Napoli”. In when Charles DARWIN (1862) published “On 1903, in occasion of his 70 th birthday, the various contrivances by which British and for - University and his colleagues tributed him eign orchids are fertilized by insects”, explain - honours and gave him a medal. While he was ing results not in the terms of Sprengel’s teleo - in Naples, Delpino became Dean of the logical interpretation, but according to his the - Faculty of Science, “Accademico dei Lincei” ory on the origin of the species, published and President of the Italian Society of Botany. three years earlier ( DARWIN 1859). He died in Naples on May 14, 1905 and Inspired by the works of Sprengel and was buried in the cemetery of Poggioreale, in Darwin, Delpino, in the spring of 1865, made the famous people’s yard. cross examinations on the mechanism of polli - nation in orchids and asclepiads. Experiments brought him to discover a type of pollination RESEARCHES (1865-1905) by insects that he describes in this way: “The aggregation of pollens in masses involves Federico Delpino’s scientific thought is tes - astonishing contrivances among Asclepiadeae, tified by 492 papers and 90 of those represent just like those Darwin reported about the the better known part of Delpino’s production, orchids; indeed the masses of pollens produced which is about floral biology, systematic and by the stems are held in the trunk of imenotters defining plant biology. Other writings are frag - (purple xylocopa), when they visit flowers to mentary productions and concern all fields of suck out . Imenotters contribute to cross- botany ( GEREMICCA 1908). pollination when they visit other flowers”. Delpino agrees with a famous Darwin’s Floral biology sentence: “No organism is self-pollinated for - Delpino gave relevant original contribu - ever”, and he shows that cross-pollination, or tions in this field, that were acknowledged in dichogamy, is not prevented by hermaphro - the international community, giving him a ditism, because obstacles of different nature,

87 such as a different ripening of pollens and ordinary, shining, metal and dark. Delpino did eggs. In the course of evolution, hermaphro - not limit himself to classifying or defining dite flowers followed rather than preceded uni - colours, he also determined their range of vis - sexual ones, in order to arrange an economy of ibility and found that white is the most recog - energies and materials; this could be consid - nizable colour on a green background. White is ered as random, according to Darwin’s theory, followed by yellow, red, bright red, purple and while Delpino considers it a natural design, light blue. He observed that the range of chro - occurring whenever difficult environmental matic power in a meadow is different than the conditions require a system different than one found in a wheat field in bloom or in a bare eterogamy, such as autorganic reproduction. field with ashy or yellowish colours, because, DELPINO (1868) reviewed observations by in this case, the grades of red are predominant other botanists, collected from many textbooks and are followed by light red, purple, light of physiology, that represented the core of his blue, white and yellow. He also noticed plants scientific production. He classified and with yellow flowers were usually able to over - denominated different systems of pollen trans - come those with white flowers, elevating the portation, partly referring to an existing dic - stamen further and by growing bigger flowers. tionary, partly upgrading it; Delpino estab - Then, a peculiar phenomenon occurs; if one lished the definition of anemophilous and looks at a meadow from up above, from the top zoophilous plants, the latter divided into ento - of a tower for example, he mostly sees white mophila, ornitophila, malacophila. flowers; instead, while observing from the At the same time, the way opened by height of a man, species with yellow flowers Darwin and Delpino was followed by the most are in clear majority. important naturalists of the time and it allowed It is interesting to report what Delpino says Delpino to exchange information, comments about the blossoming of ivy, because it testifies and personal interpretations with the most his insect knowledge also: “In the exact time influential German scientists, Fritz, Hermann, of vining, ivy is in bloom. If some creeper cov - Muller, Friederich Hildebrand, and with ers up walls with a luxurious vegetation, one Darwin himself. The latter gave most attention can explain such an abundant and “mellifera” to works and criticism of the Italian botanist blossoming, that provides an extraordinary and was rather cautious in criticizing his teleo - show, from morning to evening and for many logical view. In the paper “The effects of cross days, luring an astonishing number of insects. and self fertilisation in the vegetable kingdom” There are bees and other insects. They are pol - (London 1876), Darwin reported references linating insects of second line. Many wasps are and results gathered by Delpino in “Ulteriori also lured, to catch other insects rather than osservazioni sulla dicogamia”, in particular collecting nectar. Blossoms are also an exclu - those about anemophilous syndromes and flo - sive benefit for diptera and silfidi.” Also: ral , while the most important European “Many years ago, at my house in Chiavari, I Schools focused their attention on Italian stood by a wall covered up with ivy and for researchers ( ALIPPI CAPPELLETTI 1996; many days, from dawn to dusk, I stared at the PANCALDI 1983, 1984). fascinating show of its blossoming. I wanted to It is worth mentioning Delpino’s observa - deepen my knowledge about genera and tions and interpretations of colours, scents and species of silfidi; I collected so many samples, structures of flowers, that reveal the intriguing that I achieved my goals beyond any expecta - interactions between those and pollinating tion. The most abundant and recklessly active insects, giving new explanations. He found pollinating insect was Helophilus florens , fol - that the colour of flowers must be different lowed by Erystalis tenax , then by genera such than green, so that pollinating insects can as Chrysotoxum , Volucella , Syrphus , Paragus , quickly detect and visit flowers. Delpino made Callicera and Eumerus .” ( GEREMICCA 1908). a very interesting biological classification of Moreover, Delpino did not miss to notice colours, by distributing them in four classes: the succession of two or more colours within

88 the same flower and, with clever observations, The importance of floral biology in classifica - he determined that a change of colour indicat - tion ed the right time for pollinating insects to visit As years went by, Delpino started to think the flower. about a complete reform of the classification Once established the value of chromatic of plants, according to biological criteria, function, he clarified the luring function of philogenetic method and morphology. Delpino coloured parts in the flowers, in order to per - considered this method as “non plus ultra” of form dichogamy. For example, it is worth perfection, because all the methods used since mentioning the acute interpretation of the dark then followed morphological criteria only, purple colour taken by flowers of wild carrot resulting to be weak and insufficient. As an (Daucus carota L.) in the middle section, expression and application of these new prin - called umbrella. That colour, detaching itself ciples, Delpino published, between 1888 and from white of inflorescences, is useful to make 1894, a collection of lectures entitled the umbrella of carrots easily recognizable for “Applicazione di nuovi criteri per la classifi - the bees and to divert them from other white cazione delle piante”. They were outstanding ombrelliferes ( DELPINO 1865). from any point of view, specially for the clever Delpino also defined 45 types of floral findings in the field of systematic botany scents, with direct observations, and distrib - (DELPINO 1888-1897). Among those, it is rele - uted them in five classes (of which a transla - vant the lecture read on March 19, 1889 at tion is here attempted): delicate, aromatic, car - Academy of Sciences of the University of pological, smelly, and nauseous Bologna, where he underlined: “Systematic These scents act in three different ways: botanists make a serious mistake by associat - apatic, luring pollinating insects in general; ing Taxaceae with the genus Ginkgo , when the sympathetic, attracting special kinds of polli - latter differs from both Taxaceae and nating insects, and non-sympathetic ones, Conifers”. This exclusion and the institution of keeping away small insects, unable to perform a new family, Ginkgoacaceae, stood as a mem - cross-pollination. Flower scent lures insects orable decision in history of taxonomy more than colours do, and some bees and lepi - (SAVELLI 1965). dopterists provide evidence for this, visiting Botanical geography also became an issue flowers that release scent only at night. of great importance to Delpino, always under Scents and colours lure pollinating insects the influence of biological investigation; asso - to flowers with an attracting function enabled. ciated with Morphology and Paleonthology, it Delpino detects eleven types of lure and, concurs to the foundation of history of evolu - among those, nectar, lymph and edible pollen tion in the Plant Kingdom and it is the ultimate represent the most important sources of nour - achievement of his botanical studies. These ishment ( GEREMICCA 1908). exact concepts inspired the following papers: He supports Aristotle’s theory about the “Studi di geografia botanica secondo un nuovo behaviour of bees while visiting flowers; it indirizzo”; “Rapporto fra la evoluzione e la affirms that bees only visit one kind of flower distribuzione geografica delle Ranuncolacee” during each trip. In this way, bees save time e “Comparazione biologica di due flore and labour during their visits, because flowers estreme, artica ed antartica” ( DELPINO 1898, of different species deal with different species 1899b, 1900). of insects; thus, it is easier and more effective Delpino, considering triple pentacyclic flo - for the bee to visit one species at time. Delpino ral architecture of many monocotyledon plants gave a finalistic explanation to this phenome - as the result of a long evolution, assigns to non: “The mutual adaptation among flowers them the value of archetypical and prototypi - and their pollinators reveals, with logical evi - cal. He also thought that monocot plants, hav - dence, the existence of a preconscious and ing a structure that he called policyclic, to dis - intelligent principle concurring in the develop - tinguish it from triple pentacyclic, derived ment of organisms”. from forms of dicot plants: “If one thinks that

89 Ranuncolaceae, Ninfaceae, Botumaceae, self, Hildebrand’s writings and my own; all Idrocarideae and Alismaceae are fresh water those works concerned about a single branch plants, it is confirmed the hypothesis that the of biology: floral biology. Although the rela - division between dicotyledon and monocotyle - tive unavailability of that material (one hun - don plants took place in flooded fields, in dred times that amount, today) we had, since ancient geological ages when the proportion then, a quite adequate understanding to define between emerged lands and flooded lands was a new scientific field; it shows in the following different than today”. passages, that are worth mentioning because Among the classifications of that period, the book I quoted is hard to find: “…phitolo - the one formulated by Engler in 1892 was the gists in their issues on botany, could not detect closest to his own, for biological and philoge - phenomena related to external vital functions netical aspects; this classification considered of plants; they neglected them, or associated two types of monocot plants: those with a vari - them to phenomena of internal life (p. 4)”. able number of floral cycles and organs within “Plants find endless source of nourishment single cycles, and those with trimerous and in the elements of soil and air; for being air a pentacyclic flowers. However, Delpino under - mobile element in itself, it was thought that, lined all the weak points of this theory, that because plants could not get to nourishment, it concerned the distribution of families in class - would get to plants”. es and showed that his monophyletic scheme “Another important need is getting away was closer to the truth than the one proposed from its foes. Nature…provided outstanding by Engler. He concluded: “We both started remedies…by releasing bitter, disgusting and from the same phylogenetic and biological poisonous juices or stingy smells that make principles; it shows that we applied a different them respected and avoided by the majority of explanation (and application) of those princi - animals.” And about cross-fertilisation, he ples. Further observations and considerations adds : “most of the flowers are traps with dif - would probably tell which theory is closest to ferent and wonderful mechanisms, that give the truth.” ( GEREMICCA 1908; DELPINO 1898, the role of pollinators to insects, unconscious - 1899b). ly dedicating themselves to the work of trans - porting pollens from one flower to another”. Definition and limits of plant biology “…And about the biological role of dis - This field represents the most important semination and distribution of plants, it seems legacy of Delpino’s scientific thought, and it is to succeed with a better rate among plants worth to make a summary of what he wrote in rather than animals, that are even able to move. Naples in 1899 on the first issue of the The vital principle controls weather agents and “Bulletin of the Botanic Garden” ( DELPINO imposes dissemination to the winds; it con - 1899a): verts stamen of some clematidi, anemoni, dri - “In 1867 it was proposed the institution of adi, etc. in feathery pappus. When calyx of sin - a new branch of phitology, destined to investi - genesie and of some valerians, surrounding the gate vital functions of plants; for that disci - seeds of cotton, of apocinee, asclepiadee and pline, I suggested the name of Plant Biology” epilobi of poplars with soft hair…”. (Pensieri sulla biologia vegetale, etc.; vol. “…reviewing those phenomena of biologi - XXV del “Nuovo Cimento”, Pisa, 1867); cal origin, I tried to define the limits of biolo - “Material for observations was not abundant, gy and of the materials it is made of. consisting mainly of Crist. Corr. Sprengel’s “…Morphology must be subordinated to biol - work entitled“Stealing a secret from nature: ogy. Without the support of biology, what is the structure and fertilization of flowers”, pub - morphology if not a mere and sterile contem - lished in 1793, the Issue released by C. Darwin plation of shapes and metamorphosis, that on “Apparecchi della fecondazione nelle lacks of concepts, meanings and spirit? What orchidee nostrane (sic!) ed esotiche” and other is simple morphology if not a measure of our few memories published by C. Darwin him - ignorance? But, if both disciplines support

90 each other, they represent an interesting scien - “I tend to agree with these concepts. I think tific compound. So, human mind can elevate that two new terms should have been coined: itself to the understanding of concepts that endobiology instead of physiology and exobi - have been developing during the evolution of ology instead of biology. This is opposed by vegetal organisms. According to this point of the fact that the term physiology has been used view, biology, considered as a separate branch for centuries by medical and naturalist of natural sciences, finds its own way and ful - schools.” fils its special mission. Being detached from “As we have to use the term physiology, physiology, that concerns internal life, biology although it has a conventional value, we develops within a different field.” should give a conventional value to the term The passages reported above show our biology too, and use it as an abbreviation of view of biology back in 1867. Three main exobiology.” types of biological phenomena were already “Warming and many others have used the considered; those about protecting the organ - term ecology lately…” ism, those about cross fertilization and finally “I should criticize Warming for replacing those about dissemination. Later on, there the term biology with ecology lately. I really were more papers about biological aspects. don’t like this word. In 1881, I analysed the This scientific field grew considerably. use of this word in my “Prolegomenon to “Fourteen years later, in 1881, I got back to Biology” and I rejected it. I don’t even like this the matter; after reviewing recent published term when it applies to exobiology of human papers, I proposed about limits and contents of beings “who build up roofs and houses, while plant biology; this is what we did with an arti - thinking of economic rules”. It might be said cle entitled “Basics of Plant Biology; prole - that this term is backed up by Haeckel’s gomenon”. This work was meant to be a trea - authority (1886). He just mentions that in a tise of plant biology, that classifies all the very brief footnote on page eight of his known biological phenomena according to Generelle Morphologie der Organismen. Let’s their functions. quote this along with the literature that legiti - “Defining the field of plant biology was mated the use of the term biology; as long as easy; as long as we don’t start from the posi - naturalists don’t realize that organisms are tion stated in the prolegomenon.” three dimensional beings, therefore they have a “It is better to consider each single organ - centre and a periphery, thus a central life and a ism or individual, no matter if unicellular or peripheral life. Only then, we will rationally made of millions of cells, a three-dimensional adopt the terms endobiology and exobiology, being as mathematician would call it, as it banning physiology and biology from the sci - expands in three dimensions, a solid body entific dictionary.” indeed. Because of its own nature, it needs a centre and a periphery, with central organs or Although most of Delpino’s plant biology parts and peripheral organs or parts. From the is included in ecology now, his researches on geometrical point of view, this is very much pollination haven’t lost their appeal. The true; after reflection, it is also true from an extraordinary variety of relationships between organic point of view, even though the organic flowers and their visitors demand the attention centre rarely corresponds to the geometrical of ecologists even today, and leads to new centre. As life applies to both central and findings ( FAEGRI & V AN DER PIJL 1979; PACINI peripheral organs, it follows that, although life 1988). should be considered as a unique thing, we Delpino’s scientific thought is part of the so should consider two types of life, an internal or called “Biocomplexity”, a term that scientists central one and an external or peripheral one. recently coined to conceptualise the vast vari - The first case is covered by physiology, a term ety of relations occurring among organisms accepted by all the scholars. The proposed and between these and the environment. Its term biology might cover the second case.” field of interest is wider than that of ecology; it

91 is about mutual relations among soil, bacteria, entific methodology” in biology is still open plants, herbivores, carnivores, geological and and scholars of “philosophy of ecology” and climatic factors ( TAYLOR & H AILA 2001; biocomplexity agree that both experimental COLWELL 2001). method and observative-comparative methods About the criticism that Delpino received are relevant. from fellow scientists because none of his Both methods are about collecting data and works was supported by observations at the observation plays a key role. microscopy, it is proper to state, as Catalano As Shopenhauer stated, a creative mind is did ( CATALANO 1955), that back to the days able “to think about something that no one had when Delpino defined the basis of his Plant thought of, by looking at something everyone Biology, these basis were provided by evi - can see” ( MAYR 1990). Imagination, then, is dence that could be observed “directly” with the most important prerequisite to “scientific the naked eye. Indeed, the debate on the “sci - progress”.

LITERATURE CITED

ALIOTTA G., A LIOTTA A. 2001. Domenico DELPINO F. 1868. Ulteriori osservazioni e Cirillo, medico e botanico dell’Illuminismo considerazioni sulla Dicogamia nel Regno italiano. Il Policlinico, Sez. Prat. 108: 181-187. Vegetale. Atti della Soc. Ital. di Sc. Natur. ALIPPI CAPPELLETTI M. 1996. Federico Milano. 1868, 11 pp. 265-352; 1869, 12 pp. Delpino, memorie di biologia vegetale. Giunti 21-141, e 179-235; 1873, 14, pp. 151-349; Gruppo Editoriale, Firenze. 1874, 17, pp. 266-407. BORZÌ A. 1905. Federico Delpino, discorso DELPINO F. 1883. Teoria generale della commemorativo. Nuovo Giorn. Bot. Ital., n.s. fillotassi. Atti d. R. Univ. Di Genova, 4: 1-345. 22: 417 – 439. DELPINO F. 1888-1897. Applicazione di CAMERARIUS R.J. 1694. De sexu plantarum nuovi criteri per la classificazione delle piante. epistola. Tubingen Akademischer. Mem. R. Acc. Sc. Bologna. I, s. IV, 9, 1888, CATALANO G. 1955. La biologia vegetale pp. 221-243; II s. IV, 10, 1889, pp. 43-73; III, del nostro tempo (Ricordando Federico ivi, pp. 565-599; IV, s. V, 1, 1890, pp. 253-278; Delpino nel 50° anniversario della morte). V, s. V, 3, 1892, pp. 217-244; VI, s. V, 6, 1896- Delpinoa VIII: 1 – 31. 97, pp. 83-116. COLWELL R.R. 2001. Biocomplexity. DELPINO F. 1898. Studi di geografia botani - Encyclopedia of Life Sciences. Macmillan ca secondo un nuovo indirizzo. Mem. d. R. Publishers Ltd, Nature Publishing Group. Acc. Scienze, Bologna, s. 7: 329-358. www.els.net. DELPINO F. 1899a. Definizioni e limiti della DARWIN C. 1859. On the origin of species biologia vegetale. Bollettino dell’Orto Bota- by means of natural selection on the preserva - nico della Regia Università di Napoli, 1: 5-23. tion of favored races in the struggle for life. DELPINO F. 1899b. Rapporti tra l’evoluzio - Murray, London. ne e la distribuzione geografica delle Ranunco- DARWIN C. 1862. On the various contrivan - lacee. Mem. d. R. Acc. Scienze, Bologna, s. V, ces by which British and foreign Orchids are 8: 17-66. fertilised by insects. Murray, London. DELPINO F. 1900. Comparazione biologica DELPINO F. 1865. Relazione sull’apparec - di due flore estreme, artica ed antartica. Mem. chio della fecondazione nelle Asclepiadee. d. R. Acc. Scienze, Bologna, s. V, 8: 527-564. Aggiuntevi alcune considerazioni sulle cause FAEGRI K., V AN DER PIJL L. 1979. Principles finali e sulla teoria di Carlo Darwin. Gazzetta of pollination ecology. Oxford Press. Medica, s. II, 15, pp. 372-374, 382-384. 390- GEREMICCA M. 1908. L’opera botanica di 391. 398-400. Federico Delpino esposta criticamente.

92 Bollettino della Società dei Naturalisti in smo, la corrispondenza tra C. Darwin e F. Del- Napoli. XXI: 8-200. pino. Ed. CLUEB, Bologna. KOELREUTER G. 1761. Vorlaufige SAVELLI R. 1965. L’orto botanico di Nechrichte von einigen das Geschlecht der Bologna. In: Giacomini V., Merola A. (Eds). Pflanzen betreffenden Versuchen und Orti Botanici delle Università Italiane. Beobachtungen. Leipzig. Biblioteca dell’Orto Botanico di Napoli. LAMARCK J.B. 1802. Recherches sur l’orga - SPRENGEL C.K. 1793. Das entdeckte nisation des corps vivants. Paris. Geheimniss der Natur, im Bau und in der MAYR E. 1990. Storia del Pensiero Befruchtung der Blumen. Berlin. Biologico. Ed. Bollati-Boringhieri, Torino. TAYLOR P.J., H AILA Y. 2001. Philosophy of PACINI E. 1988. Biologia della riproduzio - ecology. Encyclopedia of Life Sciences. ne, 100 anni di ricerche botaniche in Italia. In: Macmillan Publishers Ltd, Nature Publishing F. Pedrotti (Ed.). Centenario della Società Group. www.els.net. Botanica Italiana. pp. 345-354. Milano. TREVIRANUS G.R. 1802-1822. Biologie PANCALDI G. 1983. Darwin in Italia. Ed. oder Philosophie der Lebenden Natur fur CLUEB, Bologna. Natur-forscher und Aertze. voll. 6, Gottinga. PANCALDI G. 1984. Teleologia e Darwini-

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