The role of zoological gardens in the multiple origins of Italian Primatology

The scientific role of zoological gardens has received scanty attention even in cases, such as Italian Primatology, where they seemed to have played a considerable role, especially in the field of behavioral studies. Notably, the relationship between zoos and primatology in Italy is much longer than the 40 years of life of official Primatology. The history of this relationship and connections between zoos and primatologists can reveal not only much about zoo philosophy but also about the origin of Italian Primatology. Among the various aspects that deserve attention is the history of increased interest toward issues such as primate conservation and welfare.


Introduction
Modern zoological gardens (but not aquaria) were, compared to other European countries, a relatively recent acquisition for Italy.The history of Italian zoos is not well known, therefore, even less is known about the relationship of Zoos and scientific research.It should be noted that after death zoo primates were often utilized for anatomical studies (see De Stefano, 1984-1985).However, the focus was almost invariably for evolutionary comparison with Homo sapiens.There was little interest in the order Primates per se.
While the history of zoological gardens is receiving increasing attention within a general interest toward human/animals relationship (Serna, 2017), not the same can be said of the contribution of zoos to the development of biological sciences.In the present paper, a brief overview on the multiple origin of Primatology in Italy is presented and relationships with zoological gardens are highlighted.I note that primatological research often involved and developed in collaboration with zoos.

Michele Lessona
In 1889 Michele Lessona (1823Lessona ( -1894)), Zoology docent at Turin University and one of the major Italian supporter of Darwin's theories, described an ideal orangutan Pongo sp.zoo exhibit, certainly based on his observations of some young great apes at the Royal Zoological Garden of Turin.He proposed the building of a large greenhouse with living trees, shrubs, running water, poles and bars to allow arboreal locomotion typical of the species.Lessona also cited the need to maintain other individuals of compatible animal species together with orangutans, evidently recognizing the necessity of both a physical and social enriched habitat for captive apes (Lessona, 1889;Gippoliti, 2000).Lessona was ahead of his time for his concern regarding the behavioral needs of great apes, and currently welfare research is a growing field of 'applied primatology' both in Italy and abroad (cfr.Sandri et al., 2017) that will not discussed further in this essay.

Gioacchino Leo Sera
Gioacchino Leo Sera (Rome 1878-Florence 1960) (Fig. 1) is considered a pioneer of primatological research, in fact he published in Pavia the first ever journal dedicated to Primates; the short-lived Giornale per la Morfologia dell' Uomo e dei Primati founded in 1917.Sera was a strong supporter of a polyphyletic origins of 'Homo sapiens' and of a similarly extravagant theory of an aquatic habitat for extinct giant lemurs (Sera, 1935) and Neotropical Primates (Sera, 1938).Yet Sera's research program included an extensive study of most of the primatological collections found in the major Italian Zoological and Anthropological museums (Genoa foremost, but also Turin, Milan, Pisa, Florence, Pavia, and Rome) (Sera, 1918).Furthermore, in one of his primatological papers (Sera, 1923) on the relationship between skeleton characters and locomotion in primates, he particularly acknowledged the help of Theodor Knottnerus-Meyer (1876-1936) (Fig. 2), then director of the Giardino Zoologico in Rome «che gli facilitò le frequenti e lunghe visite al Giardino Zoologico di Roma, di cui tanto giovamento ha tratto questo lavoro» (Sera, 1923, 5).The author cites his observations of two geladas (Theropitecus gelada), in Rome noting their scarce arboreal capabilities.Knottnerus-Meyer deserved the merit to have developed in Rome the 'Monkeys Village' that housed a troop of rhesus macaques living in an open air exhibit year-round (Fig. 3).
In his book, Nel Giardino Zoologico.Osservazione e Studi (Knottnerus-Meyer, 1925), which was translated in several languages the author included his behavioral observations on these macaques and formulated the hypothesis that grooming had a mainly hygienic function.Sera was greatly helped also by another zoo biologist, Oscar de Beaux (Gippoliti, 2006a), who was a fine mammal taxonomist and at the time was curator of vertebrates at the Genoa Natural History Museum.De Beaux contributed several papers to Sera's Giornale and greatly helped him furnishing taxonomically well-identified specimens.Before the Great War de Beaux had served for three years as zoologist in the famous Hagenbeck's Tierpark near Hamburg.Among his many primatological papers there is one regarding the penis morphology in the genus Macaca based on individuals that he studied living at the zoo and that later preserved as museum specimens (de Beaux, 1917): this study exemplifies how zoos and museums could collaborate for advancing comparative primate biology.

Brunetto Chiarelli
But it was only in the 1960s that a vigorous re-birth of primatology occurred in Italy.Brunetto Chiarelli received the first chair of Primatology in Italy at Turin University.He profitted from the attention given to research by the local zoological garden, and in 1962 a Primatological Centre of the University of Turin inside the zoo was established.This Primatological Centre was perhaps the first one in Europe (Torchio, 1963).Chiarelli recalled that «Thanks to the help of its owner Mr Arduino Terni and its director Prof. Alula Taibel I also create laboratory of Primatology in the Zoological Garden of Parco Michelotti, in homology with the Primate Center I started the previous years at the Zoological Garden of Rome» (Chiarelli, 2006).I was unable to find any further reference to a formal Primate Center in Rome, but by 1961 Chiarelli had published on Nature the chromosomes of orangutan utilizing the pair living in the Giardino Zoologico in Rome (Chiarelli, 1961).We now known that the male Carlo (Fig. 4) was a Sumatran orangutan Pongo abeli while the female Tuta belonged to the Borneo species Pongo pygmaeus (Gippoliti and D'Alessandro, 2013).Chiarelli's main interest was in karyological research and the rich taxonomic assemblages of zoos was a great opportunity for him.Other than a number of foreign zoos, Turin, Rome and Naples zoological gardens furnished valuable materials to his research (Chiarelli, 1962).Chiarelli soon understood the importance of taxonomy for modern primatology, and tried to integrate chromosome research on zoo primates with taxonomy (Chiarelli, 1972).Yet the absence of true taxonomists in Italian museums and a general neglect of the discipline led to severe criticism (Herskovitz, 1973).Several zoos and researchers contributed to a review paper on captive Catharrine Primate hybrids, an issue of great interest to Chiarelli (Chiarelli, 1973) that is regaining importance because of its potential role in speciation (Zinner et al., 2011).
Turin Zoo was also the place where the first behavioral studies on Primates were done in Italy (Giacoma, 1983;Giacoma and Messeri, 1992) before the zoo had to close in 1988.But the more organized attempts were done some years later in Rome.

The official birth of Primatology in Rome
Two different behavioral projects developed contemporaneously inside the «Giardino Zoologico di Roma».One project, in collaboration with the Institute of Comparative Psychology of the National Research Council (CNR) saw the creation of a research unit to study the behavior and cognitive development of capuchin monkeys belonging to genus Sapajus which at that time were housed at the zoo (Antinucci and Visalberghi, 1986).In the second project, an intensive research program was developed on the Japanese macaque (Macaca fuscata) colony.This colony arrived in Rome in 1977 directly from Takasakiyama, Japan, as a result of selection process of a Japanese commission (Fig. 5).The Japanese macaque colony occupied a renovated 'Villaggio delle Scimmie' where the previous one existed before 1935 on which grooming research was among the issues investigated for nearly 40 years.In 1981 a special issue of the journal Antropologia Contemporanea, edited in Florence by Chiarelli, included the results of a first interdisciplinary scientific campaign done on the Rome macaque colony (Chiarelli, 1981).Subsequently, behavioral studies on the macaque colony proceeded under the leadership of Alfonso Troisi (later at Rome II University) and Gabriele Schino (now at CNR) (Fig. 6).Other species were studied in a more opportunistic way in Rome (Visalberghi, 1984;Troisi and Schino, 1987).The time was ripe for the birth in Rome, on 22 march 1982, of the Associazione Primatologica Italiana (API), the Italian Primatological Association (De Stefano, 1985).While the CNR Primatological Centre was (and still is) managed separately from the zoo, the colony of Japanese macaques offers a unique Italian example of long-term collaboration between researchers and a zoo that, despite no specific funding, is able to produce abundant high quality research (for a review see Majolo et al., 2005; see also Tiddi et al., 2017).In the first decade following the official birth of primatology in Italy, Visalberghi (1988) recorded only three zoos involved with behavioral research, Turin, Rome and Cavriglia.Cavriglia was a municipal 'faunistic park' that received a small group of Japanese macaque from Rome (Camperio Ciani et al., 1984), but never had a true scientific direction and it is now closed.Despite this small number of ethological research groups operating in Zoos, Visalberghi (1988) found that for the first six Italian Primatological Meetings, almost 20% of the total communications were based on data gathered in Italian zoos.It is notable that this percentalge was about double that of communications in the International Primatological Meetings of a similar time period.
In the last two decades two other zoos have had long-term links with primatological research; the Pistoia Zoological Garden and the Parco Natura Viva near Verona.In Pistoia, the main target species was Lemur catta studied by researchers of the University of Pisa (Palagi et al., 2002;2005;Palagi, 2009), while a wider number of species, belonging to both haplorrhines and strepsirrhines, is subject to behavioral research by universities and the zoo's scientific staff at the Parco Natura Viva (e.g.Huffman et al., 2010;Regaiolli et al., 2018).Finally, Parco Natura Viva, Cavriglia and the Rome Zoological Garden (including the Museum of Zoology) were more than once hosts of the meetings of the Italian Primatological Association (API).

Perspectives for a deeper collaboration between zoos and primatologists
Although Primates in zoos are recognized as a valuable educational resource for primary and secondary schools (e.g.Falchetti et al., 1999), it seems that in the present university courses rarely the importance of zoological gardens as research and educational facilities is emphasized, and it is fairly common to find harsh criticism against zoos even among newly graduated biologists and naturalists (Gippoliti, pers. observ.).Although a specific investigation is lacking, available evidence suggests that most primatologists recognize figures such as Jane Goodall and Diane Fossey as their «iconic» masters and model of field primatologists, overly neglecting the history of Primatology and the opportunities offered by captive settings.It is a duty for zoological gardens to pay greater attention to their history also in relation with the development of scientific research and theories.When possible, zoo personal should be allowed to teach primate management, welfare and scientific potential of zoo primates in university courses.As showed by the present paper, primatology advanced not only through scientific interest at universities but also thanks to the wide knowledge of zoo staff and their interest to increase scientific knowledge and animal's welfare.Researchers and students should be made aware that zoo design and management evolve constantly as a result of development in medicine science and behavior studies (cfr.Gippoliti, 2006c).Contrary to what is usually believed today, concern for the survival of wild species and the ethical problems around human mistreatment of wild animals is well embodied within the history of Italian zoological gardens (Gippoliti, 2021).The already cited Oscar de Beaux (Florence 1879-Torre Pellice 1955) wrote a booklet titled Etica Biologica (de Beaux, 1930) just two years before opening a small zoo in Genoa Nervi.The mission of the Nervi Zoo included «education and zoophilia», recognizing the potential of natural history museums, zoos and botanical gardens in communicating not only knowledge, but also respect toward the diverse life forms (biodiversity).It should be not overlooked that primatology benefitted from some of the few scientifically managed zoos in Italy; one (Turin) directed by Prof Alula Taibel (1892-1984) that was the most prolific zoo-biologist in Italy (Gippoliti, 2019) and the second (Rome) directed by Prof Ermanno Bronzini (La Spezia 1914-Rome 2004), a leader in zoo cultural development and a follower of de Beaux's biological ethics (Gippoliti, 2021).
A relatively little-known chapter is represented by the contribution Italian zoos had furnished in changing the attitude toward non-human primates in the society.Papers on primates and their cognitive capacities appeared on the journal of the Giardino Zoologico in Rome well in the 60's of the twenty century (Chiarelli, 1960;Mangili, 1962) (Fig. 7) while two decades later a book on primates was published by Giusto Benedetti (Vittorio Veneto 1943-Turin 2011), then vice-director of the Turin Zoo (Benedetti, 1979).Important births, such as that of the orangutan Petronilla in Rome in 1970 (Fig. 8) were amply utilized by new conservation associations such as the World Wide Fund for Nature -WWF Italy Appeal, that received a strong support by several zoos, foremost those of Naples and Rome (Gippoliti, 2021).It is urgently required that Italian zoos, Primatologists and Zoologists became aware of their history.They need to better disseminate knowledge regarding their past and current role, with a special attention to promoting the conservation of biodiversity that is based on sound scientifically principles.Certainly they should avoid any resemblance of a neo-colonialism approach in tropical countries and a bias toward only a few charismatic wild species at the expense apparently less spectacular species and of ecological (and social) eco-systems.
Acknowledgement -I wish to thank Gabriele Schino for help during the preparation of this note.Fabio Di Vincenzo, Roscoe Stanyon and two anonymous referees furnished valuable suggestions to a first version of the ms.

Fig 2 .
Fig 2. Theodor Knottnerus-Meyer with the first chimpanzee of the Giardino Zoologico in Rome and a keeper (1911).

Fig 4 .
Fig 4. Male Sumatran orangutan Carlo.The history of Primatology is also the history of individual primates and of their care-givers.

Fig 5 .
Fig 5. Prof Bronzini seated among the Japanese commission looking for a new home for a colony of Japanese macaques.First on the left is Guglielmo Mangili, naturalist of the Giardino Zoologico (Rome 6 April 1974).

Fig 8 .
Fig 8. Petronilla on the cover of one of the first issues of the WWF Italy bulletin.