This art icle was downloaded by: [ OARE Consort ium ] On: 28 January 2015, At : 21: 52 Publisher: Taylor & Francis I nform a Lt d Regist ered in England and Wales Regist ered Num ber: 1072954 Regist ered office: Mort im er House, 37- 41 Mort im er St reet , London W1T 3JH, UK Historical Biology: An International Journal of Paleobiology Publicat ion det ails, including inst ruct ions f or aut hors and subscript ion inf ormat ion: ht t p: / / www. t andf online. com/ loi/ ghbi20 Digital reconstruction of Rodrigues Solitaire (Pezophaps solitaria) (Aves: Columbidae) physical appearance based on early descriptive observation and other evidence ab Mart ín A. Rodríguez-Pont es a Laborat orio de Et ología, Ecología y Evolución, Inst it ut o de Invest igaciones Clement e Est able, Minist erio de Educación y Cult ura, Avenida It alia 3318, CP 11600, Mont evideo, Uruguay Click for updates b Depart ament o de Ciencias Biológicas, Cent ro Regional de Prof esores del Cent ro (ANEPCFE), Independencia y 24 de Abril, Ciudad de Florida, Uruguay Published online: 29 Sep 2014. To cite this article: Mart ín A. Rodríguez-Pont es (2014): Digit al reconst ruct ion of Rodrigues Solit aire (Pezophaps solit aria) (Aves: Columbidae) physical appearance based on early descript ive observat ion and ot her evidence, Hist orical Biology: An Int ernat ional Journal of Paleobiology, DOI: 10. 1080/ 08912963. 2014. 954569 To link to this article: ht t p: / / dx. doi. org/ 10. 1080/ 08912963. 2014. 954569 PLEASE SCROLL DOWN FOR ARTI CLE Taylor & Francis m akes every effort t o ensure t he accuracy of all t he inform at ion ( t he “ Cont ent ” ) cont ained in t he publicat ions on our plat form . However, Taylor & Francis, our agent s, and our licensors m ake no represent at ions or warrant ies what soever as t o t he accuracy, com plet eness, or suit abilit y for any purpose of t he Cont ent . Any opinions and views expressed in t his publicat ion are t he opinions and views of t he aut hors, and are not t he views of or endorsed by Taylor & Francis. 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Term s & Condit ions of access and use can be found at ht t p: / / www.t andfonline.com / page/ t erm s- and- condit ions Historical Biology, 2014 http://dx.doi.org/10.1080/08912963.2014.954569 Digital reconstruction of Rodrigues Solitaire (Pezophaps solitaria) (Aves: Columbidae) physical appearance based on early descriptive observation and other evidence Martı́n A. Rodrı́guez-Pontes* Laboratorio de Etologı́a, Ecologı́a y Evolución, Instituto de Investigaciones Clemente Estable, Ministerio de Educación y Cultura, Avenida Italia 3318, CP 11600 Montevideo, Uruguay; Departamento de Ciencias Biológicas, Centro Regional de Profesores del Centro (ANEP-CFE), Independencia y 24 de Abril, Ciudad de Florida, Uruguay Downloaded by [OARE Consortium] at 21:52 28 January 2015 (Received 7 May 2014; accepted 11 August 2014) Rodrigues Solitaire (Pezophaps solitaria) is an extinct (eighteenth century) flightless bird endemic to Rodrigues Island (Mascarene Islands) and a sister taxon of iconic extinct Mauritian Dodo (Raphus cucullatus) (Columbiformes, Raphinae). Although numerous sub-fossilised bones have been recovered, no soft-tissue samples or taxidermically prepared specimens of Pezophaps solitaria are known to exist. The objective of this study was to produce accurate detailed digital reconstructions of Pezophaps solitaria based on early descriptions, contemporary and post-contemporary paintings, academic literature and images of mounted skeletons. Reconstructed images of Pezophaps solitaria generated illustrate the use of an established approach to visualise the appearance of this bird just before human influence and extinction. Digital simulation of photographic quality of males, females, juveniles and eggs of Pezophaps solitaria was obtained for the first time in this study. Inferred plumage crypsis in the Solitaire can be suggested to be superfluous in that their habitat was free of predators. There were a relatively large number of birds seen on the island by contemporary visitors, and thus it can be suggested cryptic colouration might be part of an adaptive strategy for intraspecific competition. Keywords: Solitaire; Pezophaps solitaria; zoo-archaeology; digital technology; virtual recreation Introduction Rodrigues Solitaire Pezophaps solitaria (Gmelin, 1788) was a flightless bird endemic to Rodrigues Island (one of the smallest Mascarene Islands east of Madagascar) (Cheke and Hume 2008; Parish 2013; Hume et al. 2014). Pezophaps solitaria was most closely related to the wellknown bird driven to extinction by anthropogenic factors: the Dodo Raphus cucullatus (Linnaeus, 1758) of neighbouring Mauritius Island (Fuller 2002; Shapiro et al. 2002; Hume 2006; Turvey and Cheke 2008; Parish 2013). The Dodo became extinct in the seventeenth century and the Solitaire in the eighteenth century. Both species were eradicated due to overhunting and introduction of invasive predators (Hume 2006; Cheke and Hume 2008; Parish 2013; Hume et al. 2014). They were the only two representatives of the subfamily Raphinae, now placed in the family of doves and pigeons (Columbidae). The recent discovery of extinct Viti Levu Giant Pigeon Natunaornis gigoura (Worthy, 2001) from the late Quaternary in Fiji indicated that flightless gigantism in insular Columbiformes was not restricted to the Mascarenes. The appearance and behaviour of the Solitaire were described in some detail by three contemporary visitors (Franc ois Leguat, Julien Tafforet and Gennes de La Chanceliére) to Rodrigues Island and to a lesser extent by at least two visitors to larger Mascarenes Islands *Email: martinangel@adinet.com.uy q 2014 Taylor & Francis (Pierre-André D’Heguerty and Jacques-Thomas de Jonchée) (Rothschild 1907; Cheke and Hume 2008; Parish 2013). Many compilations of illustrations of both ‘didines’ during the intervening four centuries reveal a much greater historical interest in picturing the Mauritian Dodo than the Rodrigues Solitaire (Owen 1879; Hume 2006; Hume et al. 2006; Parish 2013). The Dodo was a subject of more interest, celebrity and fascination than the Solitaire both in academic circles and popular culture (Hume 2006; Turvey and Cheke 2008; Parish 2013; Hume et al. 2014). Many extant Dodos were exported to Europe and at least one portrait of a living bird was painted (Parish 2013). In addition, there were a considerable number of postcontemporary paintings, outlines and even recent digital reconstructions of the Dodo (Rothschild 1907; Hume 2006; Hume et al. 2006; Parish 2012a, 2012b, 2012c, 2013). In contrast, only Leguat (1708 in Leguat 1891; Rothschild 1907; Parish 2012a, 2012b) in having observed living specimens provided the very few reliable contemporary illustrations of Solitaires. Only one such illustration shows the bird in some detail (Figure 1), whereas many very small drawings of Solitaires are included in the maps of Rodrigues Island. Many postcontemporary illustrations of this bird, e.g. by Guéneau de Montbeillard and by Émile Oustalet (in Parish 2013) are very inaccurate representations of Leguat’s work Downloaded by [OARE Consortium] at 21:52 28 January 2015 2 M.A. Rodrı́guez-Pontes Figure 1. (Colour online) From left to right and from top to bottom: the contemporary drawing of Pezophaps solitaria made by Leguat (1708 in Leguat 1891; Rothschild 1907), and some pictorial representations showing the evolution of the aspects of reconstructions of Solitaire. The post-contemporary drawings by Guéneau de Montbeillard (1803, pl. xxxiii in Parish 2012b) and Émile Outstalet (1874, p. 408 in Parish 2012b) are modified reproductions of Leguat’s original work. In contrast, post-contemporary paintings by Frederick William Frohawk, in Rothschild (1907, pl. xxiii in Parish 2012b), J. Hume (NMH Picture Library, w Julian Pender Hume in Parish 2012b) and J.C. Parish (w Jolyon C. Parish in Parish 2012b) are based on additional and more detailed evidence and analysis. (Figure 1). On the contrary, many post-contemporary paintings and outlines of Solitaires based both on skeletal reconstructions and early descriptions have been made since the nineteenth century (Rothschild 1907; Parish 2012a, 2012b, 2013; Hume and Steel 2013), but far fewer than those of the Dodo. Detailed digital photographquality reconstructions of Pezophaps solitaria showing its appearance in life could contribute to a better understanding of its ecology, behaviour and systematics. In addition, due to the lack of stuffed specimens, paintings or scientific illustrations of living Solitaires, virtual zooarchaeological digital simulations will likely enable refinement in understanding contemporary descriptions, paintings and osteological relics (Bawaya 2010; Betts et al. 2011). There is controversy and much evidence of misinterpretation in existing contemporary illustrations of Dodos (Hume 2006; Hume et al. 2006; Parish 2012a, 2013) that have in turn contributed to an incorrect understanding of many of its physical attributes (plumage textures, intraspecific colour variation, as well as colour tone of beak and foot). This, however, is not the case for Solitaires in which there remains only one contemporary drawing of a living bird in some detail (Leguat 1708 in Leguat 1891; Rothschild 1907). Only a few post-contemporary pictorial representations of the Solitaire derived from skeletal Historical Biology evidence (Parish 2012a, 2012b, 2013; Rothschild 1907; Figure 1) were found to be useful in guiding research in this study. The pictorial shortcomings and descriptions of the Dodo and the Solitaire noted above may contribute to inaccuracies in understanding the Columbiformes. One objective of this study was to verify the accuracy of the earliest descriptions of the Solitaire and potentially help clarify morphological trends within the Columbid clade. A better understanding of physical appearance of this extinct species may provide new insight into its ecological and behavioural adaptation on Rodrigues Island. Therefore, the principal aim of this study was to digitally reconstruct the appearance in life of Pezophaps solitaria as individuals before human influence and extinction. Downloaded by [OARE Consortium] at 21:52 28 January 2015 Materials and methods Digital images of Pezophaps solitaria were obtained by incorporating information from generally distinct types of resources. The first type includes scientific and scholarly literature as well as descriptions by early naturalists, e.g. translated reports from naturalists of the seventeenth and eighteenth centuries. Egg size shown was estimated following mathematical models (Rahn et al. 1975). The second type of resource includes recreational outlines and paintings of Solitairies. All images of Solitaire listed in compilations were from Rothschild (1907) or Parish (2013). The following were the images analysed in this study (Figure 1): . Contemporary drawing of Pezophaps solitaria made by Leguat (1708 in Leguat 1891; Rothschild 1907) and reproductions of this image (in Parish 2013, p. 67). . Post-contemporary painting by Frederick William Frohawk, in Rothschild (1907, pl. xxiii in Parish 2013). . Post-contemporary painting by J. Hume (NMH Picture Library, w Julian Pender Hume, in Parish 2013). . Post-contemporary painting by J.C. Parish (w Jolyon C. Parish, in Parish 2013). Photographs and illustrations of mounted skeletons (Figure 2) constitute another resource type. The following were the images used for analysis and reconstruction: . Female RCSHM/Aves 707 (Hunterian Museum of the Royal College of Surgeons, London). . Male RCSHM/Aves 706 (Hunterian Museum of the Royal College of Surgeons, London). . Female NHMUK A3506 (Owen 1879, pl. IV) (Scientific illustration). . Male NHMUK A3505 (Owen 1879, pl. IV) (Scientific illustration). . Z.DT.648 (National Museums of Scotland). 3 . A skeleton (q 2013 Simon J. Tonge) on public display on Rodrigues Island. . Skeleton of Pezophaps solitaria, Museum of Zoology, Cambridge (Oliver 1891 in Leguat 1891: frontispiece). In order to maintain consistency between resulting images and information, digital drawings as well as skeletal outline overlays of Solitaires were made directly from these skeletal images. Additional skeletal imagery information (Owen 1879; Parish 2013; Hume et al. 2014) was used and found to be complementary. As part of reconstruction of pre-human influence Solitairies, prehuman influence landscapes of Rodrigues Island were also constructed. Landscapes constructed, however, provide inferred visual appearances of an ancestral stage of Rodrigues Island but may lack accurate floristic detail. Although the general appearance of the island is likely to be correct, the reconstructions do not necessarily reflect specific plant varieties present on Rodrigues Island before human influence. A considerable number of photographs of landscapes, vegetation formations as well as floral and faunal elements were digitally exported and edited for landscape design. Generation of reconstructions involved image processing and final design. All compiled data were edited by image simulation, and artwork was done using software including Corel Draw w, Adobe Photoshop w, Adobe Illustrator w and Gimp w. Solitaire plumage was reconstructed by export and editing of many photographs of various presumed similarly coloured Columbidae species. A preliminary cephalic comparison was made using data from photographs of cladistically related genera (Pezophaps, Raphus, Caloenas, Natunaornis, Goura and Didunculus) (Worthy 2001; Shapiro et al. 2002). Natunaornis was partially recreated by guesswork using the only known premaxillar bone available (Worthy 2001). Raphus was digitally recreated from the analysis of numerous reconstructions (Parish 2013) as was Didunculus from diverse photographs. Quotes included in figures throughout the text (including citations from early descriptions) were added to indicate assumptions and evidence supporting the images shown (Ottino 2003). Results and discussion The ‘Contemporary descriptions of Rodrigues Solitaire (seventeenth to eighteenth centuries)’ section provides an overview of contemporary descriptions of the Solitaire used in this study. This is followed by a detailed discussion of observed or reported morphological and behavioural features. Finally, a general description of Rodrigues Island before human influence is outlined. A summary of reconstructed image production is provided followed by final comments. Downloaded by [OARE Consortium] at 21:52 28 January 2015 4 M.A. Rodrı́guez-Pontes Figure 2. (Colour online) Mounted skeletons of Pezophaps solitaria. From left to right and from top to bottom: male RCSHM/Aves 706 and female RCSHM/Aves 707 (q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London), Z.DT.648 (q National Museums of Scotland), Museum of Zoology of Cambridge (Oliver 1891 in Leguat 1891, frontispiece), female NHMUK A3506 and male NHMUK A3505. Contemporary descriptions of Rodrigues Solitaire (seventeenth to eighteenth centuries) This insular bird was described by Leguat (1708 in Leguat 1891, Rothschild 1907) as follows: Of all the birds on the island the most remarkable is that which goes by the name of the solitary, because it is very seldom seen in company, though’ there are abundance of them. The feathers of males are of a brown-gray color, and the feet and beak are like a turkey’s, but a little more crooked. They have scarcely any tail, but their hind-part covered with feathers is roundish, like the crupper [rump] of a horse; they are taller than turkeys. Their neck is straight and a little longer in proportion to that of a turkey’s when it lifts up its head. Its eyes are black and lively, and its head without comb or cop. They never fly, their wings are too little to support the weight of their bodies; they serve only to beat themselves, and flutter when they call one another. They will whirl about twenty or thirty times together on the same side, over a period of four or five minutes. The motion of their wings doing this makes a noise very like that of a rattle; and one may hear it two hundred paces off. The bone of their wing grows greater towards the extremity, and forms a little round mass under the feathers, as big as a musket ball. This and its beak are the chief defenses of this bird. It is very hard to catch it in the woods but easy in open places. Because we can run faster than they do, we can sometimes approach them without much trouble. They are extremely fat from March to September, and taste admirably well, especially Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology 5 while they are young. Some of the males weigh forty-five pounds. the tortoise, the making a pie out of of this bird fat was found so tough that one could not enjoy it. The females are wonderfully beautiful, some fair, some brown; I call them fair, because they are the color of fair hair. They have a sort of peak, like a widow’s upon their breasts [beaks], which is of a dun color. No one feather is straggling from the other all over their bodies, they being very careful to adjust themselves, and make them all even with their beaks. The feathers on their thighs are round like shells at the end, and have an agreeable effect being very thick there. They have two risings on their craws [crop] and the feathers are whiter than the rest, which livelily represents the fine neck of a beautiful woman [‘un beau ƒein de femme’ in the original French version; Parish 2012d]. They walk with so much stateliness and good grace that one cannot help admiring them and loving them; by which means their fine mein often saves their lives. Pierre-André D’Heguerty described the Solitaire, but probably describes individuals that kept in captivity on another of the Mascarenes Islands (Cheke and Hume 2008): Tafforet (1726 in Cheke and Hume 2008, Parish 2013) gave the following descriptive data: Together, these various descriptions indicate that the Solitaire was a stocky, flightless bird larger than turkey or swan. Although the Solitaire’s colouration is treated in more detail in the next sections, its plumage seems to have been predominantly light grey, with a darker brownish colouration on the back. The female can be inferred to be white on the chest. Presumably, these birds also had a few bold tones present on the dorsal surface of the neck region (Leguat: ‘fair hair’). The beak was incurved, short, sturdy and hooked at the tip similar to the beak of a turkey. Its irises were black and also had a black ‘velvet’ caruncular ridge just above the beak. Its size was similar to that of a swan or a turkey, and its tail was atrophied. Their wings were stunted with feathers (e.g. rectrices and tectrices) apparently somewhat reduced. Although their wings were not airworthy, they served in resolving territorial conflicts and fighting in that the frontal portion of wing bone was like a ‘musket ball’ for beating loudly or for defence. These descriptive data are taken into account in subsequent sections for detailed comparative analyses with those obtained from other categories of resource types. The solitaire is a large bird, which weighs about forty or fifty pounds. They have an extremely large head with a manner of headband/frontlet to the face that one would say is black velvet. Their feathers are neither feathers nor hair; they are of light gray color, with a little black on their backs. Strutting proudly about either alone or in pairs, they preen their plumage or fur with their beak and keep themselves very clean. Their toes are furnished with hard scales and can run quickly among rocks. They are well adapted among the rocks where a man, however agile, can hardly catch them. They have a very short beak of about an inch in length, and which is sharp. They, nevertheless, do not attempt to hurt anyone. An exception is when they find someone before them, and then when hard pressed will try to bite him. They have a small stump of a wing, which has a sort of bullet at its extremity to serve for defense. Jacques-Thomas de Jonchée (1729 in Parish 2013) referred to the Rodrigues Solitaire as: ‘There are birds which one names solitaires, flamingos, parrots, turtledoves, wood pigeons, blackbirds, and bats, from which an excellent fat is drawn.’ Cheke and Hume (2008) and Parish (2013) have considered the possibility that Solitaires described here were captive specimens and were likely exported along with tortoises from Rodrigues Island. Significantly, Gennes de la Chanceliére (1733 in Parish 2013) reported: Our people are said to have seen kids and a great number of birds of various species together; two bird species, among the others, were larger by a third than the stoutest turkey. These large birds appeared, nevertheless, quite young still with down on the neck and the head. They had wings with only little covering, and without having the tail formed. Three sailors also said that they had seen two others of the same species, as stout as the largest ostrich. The smaller birds had heads nearly like that for an ostrich, but with feet similar to those of turkeys. Their feet were unlike those of the ostrich, which are forked and split in the shape of foot of a hind. These two birds, when skinned, had an inch of fat on the body. Unlike the delicate flesh of One finds there also birds of various types which one often takes after chase, & among others the Solitaires which almost do not have plumes on the wings; this bird [is] larger than a swan, with a sad countenance; tamed [probably meaning ‘imprisoned’], one always sees it walking in the same line as long as it has space, and retrogressing in the same way without deviation. When one makes the opening of it, one usually finds there the Bézoards which one does value, & which are useful in medicine. Rodrigues Solitaire: general morphology Available literature indicates that the Solitaire had a large body size but with a considerable reduction in the pectoral region and wings, as well as short tarsometatarsi and digits. These body-size reductions were in contrast with its disproportionately long tibiotarsi (Owen 1879; Rothschild 1907; Livezey 1993). The latter would be an apomorphic character among Columbiformes because they were generally of compact morphology with short legs, a small beak and small head (Baptista et al. 2009). Single paedomorphic characters (which were prominent in Raphus cucullatus, e.g. pectoral underdevelopment) and peramorphic structure of trunks, skulls and pelvic appendages were also reported for Pezophaps solitaria 6 M.A. Rodrı́guez-Pontes Downloaded by [OARE Consortium] at 21:52 28 January 2015 (Strickland and Melvilla 1848 in Livezey 1993). The unusual sexual size dimorphism in Pezophaps solitaria (Figures 2, 6, 7 and 9) was considered one of the greatest among carinate birds (Rothschild 1907; Livezey 1993; Dickinson 2003). Body mass and size measurements were estimated to be 28 kg (90 cm) and 17 kg (70 cm) for male and female, respectively (Livezey 1993). This agrees with early descriptions noted previously in which the Solitaire size was compared with that of turkeys and swans. Extant Columbiformes, however, had enormous size and weight variation ranging from 31 g in Columbina passerina to 2000 g in Goura (Dunning 1992 in Baptista et al. 2009). The insular genera Raphus, Pezophaps and Natunaornis were thus exceptional in size for this family. Plumage and skin Some discordance in plumage colouration is evident in the literature. Grey and brown (Leguat 1708 in Leguat 1891; Rothschild 1907) and also predominantly light grey (Tafforet 1726 in Cheke and Hume 2008; Parish 2013) have been reported for the plumage of the Solitaire. Intraspecific variation in plumage colour (from albino to dark) observable comparing paintings of Raphus cucullatus was not reported for the Solitaire. Tafforet (1726 in Cheke and Hume 2008) observed that Pezophaps solitaria fed on seeds and leaves of trees. Accordingly, this flightless bird only could have been occasionally frugivorous taking fallen fruits (e.g. pandans and palms; Figure 8). The seed-eating Columbidae were often seen having grey and brownish colours but with occasional iridescent tones around the neck or breast region. The predominantly fruit-eating members of this family tend to be more colourful with greens, purples, oranges and reds (Baptista et al. 1992, 2009; Gibbs et al. 2010; Wells J and Wells A 2001; Lack 2003). The exceptionally colourful character of the seed-eaters Goura and Caloenas suggests that these genera represent an apomorphy compared with other members of the clade (e.g. Raphus, Pezophaps and Didunculus). These later genera all show plesiomorphic grey-brownish plumage tones. As for the Dodo, there is a considerable variation in the appearance of the Solitaire comparing various paintings, drawings and outlines. Frohawk’s painting shows an exaggerated caruncular ridge, as noted by Parish (2013), e.g. whereas other representations do not show such an exaggerated structure (Figure 1). Such discrepancy is also evident when describing the bird’s plumage, as in the case of older post-contemporary pictorial recreations. Some paintings show the bird as having a brownish plumage, whereas in others only grey predominates (Figure 1). The typical combination of shades of grey and brown in forest seed-eating pigeons (e.g. Patagioenas and Zenaida) has not been discussed previously for Pezophaps. Recent scientifically based analyses of the Solitaire, however, provided a new insight in determining its likely plumage colour (Hume 2013; Parish 2013). A presumed adult plumage with characteristics of juvenility (paedomorfism) was observed in the Solitaire, although not as prominently as in the Dodo (Strickland and Melvilla 1848 in Livezey 1993). This characteristic agrees with Tafforet (1726 in Cheke and Hume 2008): ‘neither feathers nor hair’. This hair-like plumage might have had characteristics more like the plush feathery texture covering many Columbiformes (e.g. Streptopelia) rather than the horsehair-like plumage appearance of Casuarius (Struthioniformes) or Apteryx (Apterygiformes). Moreover, this texture must have been present in the cephalic, neck and in parts of ventral surfaces of the bird (Figures 3, 4 and 7– 9). Columbinae may be sexually monomorphic or dimorphic in which the male is frequently larger and more colourful (Baptista et al. 1992, 2009; Wells J and Wells A 2001; Lack 2003; Gibbs et al. 2010) (Figures 5 and 6). It must be acknowledged that sexual dichromatism in pigeons can vary from a slight to markedly distinct dimorphic tones between the sexes. Differences in colouration are usually confined to breast, head or neck regions (Goodwin 1960). This variation is consistent with early descriptions of the living Solitaire discussed previously. All columbiform species have a ring of bare skin around their eyes (‘orbital ring’) (Baptista et al. 2009). Orbital rings and hindlimbs can be blue, red, white or yellow (Baptista et al. 1992, 2009; Wells J and Wells A 2001; Lack 2003; Gibbs et al. 2010). In addition, feathered tarsi are also found as a general feature among Columbiformes (Baptista et al. 2009), and so would be expected to have been present in the Solitaire also (Figures 7 –9). In summary, it can be inferred that the plumage of Pezophaps solitaria was mostly grey, dorsally brownish with darker tones on the back, and with males being more colourful than paler females (Figures 7 – 9). It can be hypothesised from Leguat (1708 in Leguat 1891; Rothschild 1907): ‘ . . . they are the color of fair hair’. There is a possible comparison that can be made between the silkiness of a women’s hair and aspects of the sparse iridescent plumage on the Solitaire’s neck (Figure 4). Both black-eyed sexes had a conspicuous black caruncular ridge at the base of their beak (Figures 3 and 4) and were more prominent in males as is usually the case in pigeons. It should be considered that this structure is very likely overly represented in several paintings of this bird (Parish 2013; Figure 1). The digital recreation of Pezophaps solitaria plumage shown (Figures 7 – 9) is based only on typical tones and textures found in Columbiformes order. Solitaire is most generally considered seed-eating in its forest habitat and is most usually described as having grey and brown plumage tones. Neotropical seed-eating Picazuro Pigeon Patagioenas picazuro, the Eared Pigeon Zenaida auriculata Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology 7 Figure 3. (Colour online) Cephalic reconstructions (q 2014 M.A. Rodrı́guez-Pontes) from different specimens of Solitaire. From left to right and from top to bottom: derived artwork from a skeleton (q 2013 Simon J. Tonge) in a public exhibit on Rodrigues Island, from an individual of the Museum of Zoology of Cambridge (Oliver 1891 in Leguat 1891, frontispiece), male NHMUK A3505 and female NHMUK A3506 (Owen 1879, pl. IV), male RCSHM/Aves 706 and female RCSHM/Aves 707 (original q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London), and Z.DT.648 (original q National Museums of Scotland). and the Collared Dove Streptopelia decaocto were considered to have suitable sources of plumage colour and texture variation for the reconstruction of Solitaire’s plumage. Further discussion involving plumage and behaviour is given in the ‘Behavioural aspects’ section. Behavioural aspects The Solitaire had a strongly territorial (Figures 9 and 10) and aggressive behaviour (Hume and Steel 2013). The prominent wing carpal knob (Figures 6 – 9) was greater in males and was used as a weapon by either sex in very audible territorial combats (Leguat 1708 in Leguat 1891; Rothschild 1907; Tafforet 1726 in Hume and Steel 2013). Hume and Steel (2013) proposed that limited resource availability on the small Rodrigues Island generated evolutionary pressure for this strong territorial behaviour. Accordingly, the plumage of both male and female Solitaires would have been unnecessarily cryptic in the pre-human influence forest habitat and with no known predators (Figures 8 – 11). As noted previously, many forest seed-eating pigeons have similar crypticism. An interesting evolutionary hypothesis posing a link between plumage and behaviour can be envisaged. Plumage crypticism in Solitaire could still be considered as a remaining plesiomorphy to avoid predation. However, this crypticism could have resulted from or have been retained in response to adaptive pressure to secure limited resources also reflected by the strong territorial behaviour of this species. By hiding their presence from other individuals, not to mention males hiding females from other males, the adaptation might Downloaded by [OARE Consortium] at 21:52 28 January 2015 8 M.A. Rodrı́guez-Pontes Figure 4. (Colour online) Rodrigues Solitaire: a detail of the skull of a skeleton (q 2013 Simon J. Tonge) on public exhibit on Rodrigues Island, and its reconstruction (q 2014 M.A. Rodrı́guez-Pontes). Ramosmania rodriguessi (q 2011 C.T. Johansson, Wikipedia) appears in the background. Centre right: a reconstructed leg; bottom: detail of a reconstructed adult male (q 2014 M.A. Rodrı́guez-Pontes) showing different plumage tones and textures, sparse iridescence on the dorsal neck, and the carpal knob (derived artwork from original skeletal image RCSHM/Aves 706 q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London). have enabled more effective intraspecific competition for limited environmental resources. Nesting and egg When these birds build their nests, they choose a clean place, gather together some palm-leaves for that purpose, and heap them up a foot and a half high from the ground, on which they sit. They never lay but one egg, which is much bigger than that of a goose. (Leguat 1708 in Leguat 1891; Rothschild 1907) (Figure 6) The eggs in Columbidae are usually white or lightly coloured. Rahn et al. (1975) found a strong correlation between female body size and egg mass. They analysed data from 45 species of Columbiformes (8 Pteroclidae and 37 Columbidae), the following regression equation developed in that study was used to generate a realistic digital recreation (shape, size and colour) of the Solitaire’s egg. W ¼ a £ B b: where W is the egg weight in grams, B is the female body weight in grams (estimated to be 17000 for Pezophaps solitaria), a is the proportionally constant (0.206 in Columbiformes) and b is the regression line slope Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology 9 Figure 5. (Colour online) Skeletons of the specimens analysed in this study, and their respective overlays and reconstructions (q 2014 M.A. Rodrı́guez-Pontes). From left to right: derived artwork from original skeletal images of a specimen maintained in the Museum of Zoology of Cambridge (Oliver 1891, in Leguat 1891: frontispiece), male NHMUK A3505 and female NHMUK A3506 (Owen 1879, pl. IV), male RCSHM/Aves 706 and female RCSHM/Aves 707 (original q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London), and Z.DT.648 (original q National Museums of Scotland). (logarithmic) for Columbiformes, 0.759. W ¼ 0:206 £ 17; 0000:759 ¼ 334:8 g ðPezophaps solitariaÞ: Log W ¼ log a þ B log b; Log 334:8 ¼ log 0:206 þ 0:759; Log 17; 000 ¼ 2:52 ðPezophaps solitariaÞ: Standard error of estimate calculated for log W ¼ 0.101; confidence: 68%; limits of the value of log W (CL) ¼ 1.26; correlation coefficient (r) ¼ 0.94 (Rahn et al. 1975). Rahn et al. (1975) determined the range and mean values of egg weight for Columbiformes at 100 g body weight to be 35 – 2000 and 313 g, respectively. Although body weights of the Mauritian Dodo, the Rodrigues Solitaire and of the Viti Levu Giant Pigeon (Natunaornis gigoura) (Worthy 2001) could have exceeded the range of applicability of the equation’s components a and b, the mathematic results obtained here for Pezophaps solitaria suggest that the egg could have had at least three times the mass (ca. 150 – 334.8 g) of that of Goura cristata (ca. 50 g), the largest living Columbidae (Rahn et al. 1975; Frith 1982; Saunders et al. 1984). Downloaded by [OARE Consortium] at 21:52 28 January 2015 10 M.A. Rodrı́guez-Pontes Figure 6. (Colour online) A detailed view of mounted skeletons of Solitaires and reconstructions. Reconstructions in the centre and on the right are mounted in what appear to be an unrealistic stance. From left to right: derived artwork from originals images of skeletons maintained in the Museum of Zoology of Cambridge (Oliver 1891 in Leguat 1891, frontispiece), male NHMUK A3505 and female NHMUK A3506 (Owen 1879, pl. IV). According to Leguat (1708 in Leguat 1891; Rothschild 1907), the Solitaire’s egg was much larger than that of the Greylag Goose Anser anser (5 – 7 cm £ 8 – 10 cm, 150 – 220 g; Soames 1986). However, the assumption of similar egg densities between these species results in an overestimated size for the Solitaire’s egg. Its egg would be predicted to be as big as that of, e.g. the Greater Rhea Rhea americana (ca. a third larger and heavier bird than Pezophaps solitaria; egg: 9 £ 13.2 cm, 600 g; Rodrı́guezPontes, unpublished results). Thus, provisional estimation of the Pezophaps solitaria’s egg size (ca. 8 cm £ 12 cm; Figures 7 and 9) was based on a presumed higher density in relation to the egg of Anser anser, and on maximum egg weights found for Solitaire (334.8 g) and Goose (220 g). Although great variation (at least 50% larger) in both egg size and mass between avian populations of a given species may occur (Christians 2002), our estimated egg size for Pezophaps solitaria is reasonably larger than that of Anser anser in agreement with Leguat’s accounts. Rodrigues Island: pre-human environment The generation of accurate detailed images of the Solitaire in its natural pre-human environment is a primary objective in this work. Pictorial representation of 11 Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology Figure 7. (Colour online) Skeletons of Solitaires in presumed realistic stances and their reconstructions. From left to right: derived artwork from originals male RCSHM/Aves 706 and female RCSHM/Aves 707 (q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London), and Z.DT.648 (q National Museums of Scotland). biological systems such as the Solitaire in the habitat to which it was highly adapted can be more informative than simply listing relevant data. The observer generally assumes a uniform level of information in a given figure. If reconstructed images of Rodrigues Solitaire are intended to be realistic, then background landscapes in a given figure should be as well (Ottino 2003). Therefore, positioning of digital reconstructions of the Solitaire within what can be considered its likely former natural scenery was carefully developed as part of this study (Figures 4 and 8– 11). Rodrigues Island (17.7 km long, 8.45 km wide, 104 km2 surface area) is tropical and well known for its unique native fauna and flora. Antoine Valleau (1692 in Parish 2013) reported: . . . that there are plenty of tortoises and birds larger than the turkey [cocq’s d’Inde] called solitaires, which one takes by hand, and many fish, that he left there eight M.A. Rodrı́guez-Pontes Downloaded by [OARE Consortium] at 21:52 28 January 2015 12 Figure 8. (Colour online) Above (from left to right and from top to bottom; q 2014 M.A. Rodrı́guez-Pontes): a cephalic comparison between Didunculus (inspired artwork from an original image by U. Beichle), Goura, Caloenas, Raphus, Pezophaps and a presumed overlay of Natunaornis (inspired artwork from Worthy 2001, p. 781). Below (q 2014 M.A. Rodrı́guez-Pontes): a juvenile Solitaire feeding on Pandanus fallen fruit. Plants of Dombeya and Gouania, and extinct giant tortoises Cylindraspis are also included. passengers with grain and all kinds of seeds to see what the ground could produce. The pristine landscape of the island was described by Leguat (1708 in Leguat 1891, Rothschild 1907): we could hardly take our eyes off the mountains of which the island almost entirely consists. They are so richly spread with great and tall trees: the valleys are covered with Palm trees, Plantains, Ebonies and several other sort of trees. This ancient natural environment and its biodiversity have suffered from four centuries of exploitation, unsustainable agriculture, deforestation and introduction 13 Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology Figure 9. (Colour online) A male (top) and a nesting female (bottom) with an egg. Plants of Pandanus, Latania, Ramosmania and Gouania, and two giant tortoises Cylindraspis are also represented (q 2014 M.A. Rodrı́guez-Pontes; derived artwork from originals Male RCSHM/Aves 706 and Female RCSHM/Aves 707 q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London, Z. DT.648 q National Museums of Scotland and q 2011 C.T. Johansson, Wikipedia). of invasive biota. None of the native primary forests remain on the island today. The pre-human Rodrigues Island landscape has been characterised with having various types of tree forests, including those of other tropical to semi-dry tree species (Leguat 1891; Gade 1985; Cheke and Hume 2008; Hume 2013). Much of the island’s rich biodiversity has disappeared, including the loss of at least 20 bird species that are now presumed to be extinct. Downloaded by [OARE Consortium] at 21:52 28 January 2015 14 M.A. Rodrı́guez-Pontes The extinction of endemic terrestrial vertebrates includes two giant tortoise species (Cylindraspis vosmaeri and C. peltastes; Griffiths et al. 2013; Figures 8 –11). Today, there remain only three endemic representatives of pre-human influence terrestrial vertebrates: two passerines (Foudia flavicans and Acrocephalus rodericana) and the fruit bat Pteropus rodricensis (Hume 2013, Figures 10 and 11). Although much less is known about terrestrial invertebrate fauna, compared with vertebrates, significant endemisms have been reported (Griffiths et al. 2013). The native flora of Rodrigues comprises 133 indigenous plant species including five endemic genera. Those remaining at the present time includes 123 species, 37 of which are critically endangered of becoming extinct, e.g. Ramosmania rodriguesii (Figures 4 and 9), Dombeya rodriguesiana, Myoporum mauritianum and Gouania leguatii (Strahm 1989). Of the 25 species of Pteridophytes originally present on the island, including two endemic Selaginella sp., four are thought to have become extinct (Lorence 1976). The pandans, Pandanus heterocarpus and Pandanus verschaffeltii, as well as the now threatened palms, Latania verschaffeltii, Dictyosperma album and Hyophorbe verschaffeltii, may have been significant in the diet of Pezophaps solitaria (Strahm 1989; Figure 8). Semi-dry tropical forest vegetation matrices were constructed in our study to serve as a suitable background for the digital reconstruction of Pezophaps solitaria. Dictyosperma, Gouania, Pandanus, Ramosmania, and other flora and fauna of Rodrigues Island were included to improve landscape background. Ecological restoration of indigenous forest on this island was improved using image resources from the Nature Reserve of Grande Montagne (Figure 11). The landscapes developed do not reflect an exhaustive archaeological recreation of the former Rodrigues Island environment (Figures 8– 11). Digital synthesis of the Solitaire in its native landscape Although images resulting from digital recreations are dependent on technology and data sources used and subjectivity of the author (Ottino 2003; Bawaya 2010), the approach used to generate the living appearance of Pezophaps solitaria, as an individual in its natural environment before human influence is internally consistent with descriptive and anatomical data used. To minimise inaccuracies, plesiomorphic characters (or probable characteristics) were used that were widely distributed among Columbiformes (e.g. greyish skin and Figure 10. (Colour online) An adult and a young male Solitaire are preparing to fight, and a female stands nearby, in a reconstructed pre-human forest landscape on Rodrigues Island. Tall palms are included in which Dictyosperma flourish in the background. Among the Diospyros and Pandanus trees, many giant tortoises Cylindraspis are present and are in search of food. Plants of Gouania and Ramosmania are also present (q 2014 M.A. Rodrı́guez-Pontes derived artwork from the original landscape image of Rodrigues Island 2010, q Jean Mark Lim, Flickr). Technical detail in J.P. Hume artwork (Parish 2012b; Hume and Steel 2013) was useful for these reconstructions. Downloaded by [OARE Consortium] at 21:52 28 January 2015 Historical Biology 15 Figure 11. (Colour online) A digital simulation of a male Solitaire courting a female individual in a realistic picture of an ecological restoration of an indigenous forest on Rodrigues Island (Nature Reserve of Grande Montagne). Simulated giant tortoises Cylindraspis and flying foxes Pteropus are included in the background. q 2014 M.A. Rodrı́guez-Pontes derived artwork from original landscape image q 2006 B. Navez, Wikipedia, and from Male RCSHM/Aves 706 q Courtesy of the Hunterian Museum of the Royal College of Surgeons, London. beak tones, and white egg shell) (Baptista et al. 2009) in Solitaire reconstructions. It is acknowledged that although greater probability of accuracy was sought, the occurrence of still unknown apomorphies is possible. As in the case of the Solitaire’s reconstruction made by Parish (2013), tarsal scutellation and leg colour are guesswork inspired by those characters of the Dodo consistent with those reported in Columbiformes (Baptista et al. 2009; Figures 4 –7). All skeletal overlays and body reconstructions shown in Figures 3 –7 reveal some stance variation dependent on how realistically these skeletons were mounted. Recent evidence for a head-forward stance in moas (Dinornithiformes) (Worthy and Holdaway 2002) rather than that for an upright position as in older skeletal mountings was taken into consideration. Although a biomechanical analysis would be necessary to solve this issue, the possibility of an ‘alertness stance’ for many uprightly mounted skeletons of Solitaires was taken into account. Leguat (1708 in Leguat 1891; Rothschild 1907) referred the Solitaire as lifting its straight neck up. Subsequently, we considered that our most accurate reconstructions of Pezophaps solitaria were obtained from the RCSHM/Aves 707 (female) and RCSHM/Aves 706 (male) specimens’ images (Figure 7). Restoration of life-like appearance of specimen Z.DT.648 provided evidence for the possibility of its being a juvenile individual (presumed male) based relative body proportions, i.e. shorter neck, longer legs and a more stylised adult male trunk. Also, body size was more like that of a male than a female. A possible allometry involving – among others – these body parts between these three reconstructed specimens (male and female vs. juvenile) is shown in Figure 7. The striking ‘deformed pigeon’ appearance of the Solitaire is particularly evident considering the consistently ‘pigeon-like’ head generated during reconstructions done as part of this study. A considerable variation in cephalic morphology is seen comparing species of AfroEurasian and Australasian pigeons (Pereira et al. 2007 in Baptista et al. 2009). This is most evident comparing related members of the clade Columbiformes (Caloenas, Didunculus, Goura, Natunaornis, Pezophaps and Raphus), as shown in Figure 8. The Manumea Pigeon, Didunculus strigirostris, had traditionally been considered most 16 M.A. Rodrı́guez-Pontes closely related to Raphus and Pezophaps due to its hooked beak (Wendt 1982). Actually, results presented here further illustrate that reconstructions of Pezophaps have a morphology that can be considered as transitional between that of the Mauritian Dodo and that of many extant pigeons and doves. Recent molecular studies showed that the brilliant and colourful Nikobar Pigeon Caloenas nicobarica – despite its very different appearance – is the closest living relative of Raphus cucullatus and Pezophaps solitaria (Shapiro et al. 2002). Downloaded by [OARE Consortium] at 21:52 28 January 2015 Final remarks and conclusion This study provides a basis for further research for understanding the natural history and evolutionary adaptation of the Solitaire. Digital reconstructions of photographic image quality of males, females, juveniles and eggs of Pezophaps solitaria were produced based on observations made by early naturalist as well as essentially all scientific and scholarly literature and anatomical data available. In the absence of known natural predators for the Solitaire, it is suggested that plumage crypsis in this species may be part of an adaptive strategy for intraspecific competition. Acknowledgements Academic support from Prof. Fernando Costa-Carbonell (IIBCE, Laboratorio de Ecologı́a, Etologı́a y Evolución), many valuable suggestions made by two reviewers, useful comments and essential bibliography provided by Jolyon C. Parish and by Julian P. Hume as well as additional comments and help with the manuscript by Paul R. Gill are fully acknowledged. Indispensable osteological images for our research were provided by Sarah Pearson (Curator of the Hunterian Museum of the Royal College of Surgeons, London) or authorising their use in this article was done by Margaret Wilson (National Museums Scotland, Administrator). References Baptista L, Trail P, Horblit H. 1992. Family columbidae (pigeons and doves). In: del Hoyo J, Elliott A, Sargatal J, editors. Handbook of the birds of the world. Vol. 4. Barcelona: Lynx Edicions; p. 60 –243. Baptista LF, Martı́nez-Gómez E, Horblit HM. 2009. Darwin’s pigeons and the evolution of the Columbiforms: recapitulation of ancient genes. Acta Zool Mex (n.s.). 25(3):719–741. Bawaya M. 2010. Virtual archaeologists recreate parts of ancient worlds. Science. 327:140 –141. Betts MW, Maschner HDG, Schou CD, Schader R, Colmes J, Clement N, Smuin M. 2011. 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