Editing Rodrigues solitaire (section)

==Behaviour and ecology==

{{multiple image
|align = left
|total_width = 300
|image1 = Pezophaps wing bones.jpg
|alt1 = Illustration of an assemblage of solitaire wing-bones
|caption1 = Wing bones, including carpal knobs (87–90) in the middle right
|image2 = Pezophaps limb bones.jpg
|caption2 = Limb bones, two with healed fractures (135–136) lower right
|alt2 = Limb bones
}}

Observations of the Rodrigues solitaire in life indicate that they were highly [[territory (animal)|territorial]]. They presumably settled disputes by striking each other with their wings; to aid this purpose, they used the knobs on their wrists.<ref>{{cite journal |last= Rand |first= A. L. |year= 1954 |title= On the Spurs on Birds' Wings |journal= The Wilson Bulletin |volume= 66 |issue= 2 |pages= 127–134 |jstor= 4158290 |url= http://sora.unm.edu/node/127344 }}</ref> Fractures in their wing bones also indicate that they were used in combat.<ref name="Livezey1993"/> It has also been suggested that these fractures may have been the result of a hereditary bone disease rather than battle-injuries.<ref>{{cite journal |doi= 10.2307/2405692 |last= Amadon |first= D. |date=1951 |title= Inbreeding and Disease |journal= Evolution |volume= 5 |issue= 4 |pages= 417 |jstor= 2405692 }}</ref> But in all extant birds where carpal spurs and knobs are present, these are used as weapons without exceptions. Though some dodo bones have been found with healed fractures, it had weak pectoral muscles and more reduced wings in comparison with the Rodrigues solitaire. Since Rodrigues receives less rainfall and has more seasonal variation than Mauritius, which would have affected the availability of resources on the island, the Rodrigues solitaire would have more reason to evolve aggressive territorial behaviour.<ref name="HumeSteel2013"/> Several accounts state that they also defended themselves with a powerful bite.<ref name="Fuller2001"/>

In addition to their use as weapons, both sexes of the Rodrigues solitaire also used their wings for communication. The wings could create low-frequency sounds for communicating with mates, or to warn rivals, but it is unknown exactly how this sound was created. The sound could be heard 200 yards (182 m) away, and this may therefore be the size of the territory of an individual. Other species of birds (e.g., [[club-winged manakin]]) are also known to use their wings to create sounds that attract mates or mark their territory.<ref name="HumeSteel2013"/>

In 1869, the Newton brothers suggested that the carpal knobs may have been formed through continuous injuries, as they resemble diseased bone.<ref name="NewtonNewton1869"/> It has also been claimed that the carpal knobs were instead formed due to a hereditary disease caused by inbreeding. This was dismissed in a 2013 study, since such lesions would likely not occur only in a specific part of the skeleton, but would appear in any growing bone tissue. If such a disease was due to inbreeding, it would also be present in other isolated island bird populations, but it is not. The authors instead suggested that the wing bones contained [[metaplastic]] tissue able to form the knob. This development was either in response to continuous impacts during combat, or to hormones released when individuals paired up and acquired territories. It appears a male which had long held a territory would possess especially large carpal knobs, and that their mates would have such developments as well, only smaller.<ref name="HumeSteel2013"/>

{{multiple image
|align = right
|total_width = 300
|image1 = Pezophaps sternum.jpg
|alt1 = Male and female Rodrigues solitaire sterna
|caption1 = [[sternum|Sterna]] of a female (above) and male from below
|image2 = Pezophaps vertebrae and foot.jpg
|caption2 = Vertebrae and left foot
|alt2 = 
}}

Some evidence, including their large size and the fact that tropical and [[frugivorous]] birds have slower growth rates, indicates that the Rodrigues solitaire may have had a protracted development period. Based on mass estimates, it has been suggested the male could reach the age of 28, and the female 17.<ref name="Livezey1993"/> The French economist [[Pierre-André d'Héguerty]], writing about his time on the island around 1735, stated that a captive Rodrigues solitaire (which he described as having a melancholic appearance) would always walk in the same line until running out of space, and then return.<ref name="ChekeHume2008p167">{{cite book| last1= Cheke| first1= A. S.| last2= Hume| first2= J. P.| year= 2008| title= Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues| publisher= T. & A. D. Poyser| location= New Haven and London| isbn= 978-0-7136-6544-4| pages= 167–168}}</ref> The species may have lived primarily in the island's woodlands, rather than on the shores.<ref name="Fuller2001"/>

Many other of the endemic species of Rodrigues became extinct after the arrival of humans, so the [[ecosystem]] of the island is heavily damaged. Before humans arrived, forests covered the island entirely, but very little remains today due to deforestation. The Rodrigues solitaire lived alongside other recently extinct birds such as the [[Rodrigues rail]], the [[Rodrigues parrot]], [[Newton's parakeet]], the [[Rodrigues starling]], the [[Rodrigues scops owl]], the [[Rodrigues night heron]], and the [[Rodrigues pigeon]]. Extinct reptiles include the [[domed Rodrigues giant tortoise]], the [[saddle-backed Rodrigues giant tortoise]], and the [[Rodrigues day gecko]].<ref>{{cite book| last1= Cheke| first1= A. S.| last2= Hume| first2= J. P.| year= 2008| title= Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues| publisher= T. & A. D. Poyser| location= New Haven and London| isbn= 978-0-7136-6544-4| pages= 49–52}}</ref>

===Diet===

{{multiple image
|align = left
|total_width = 300
|image1 = Pezophaps pelvis and gizzard stone.jpg
|alt1 = Gizzard stone, and pelvic and wishbones
|caption1 = Pelvis of a female (1) and male (2), [[furcula]] (3) and [[gizzard stone]] (4–6)
|image2 = Rodrigues Solitaire male pelvis.jpg
|caption2 = Male pelvis from below and [[coccyx]]
|alt2 = Map of human settlement on Rodrigues
}}

Leguat stated that the Rodrigues solitaire fed on [[date (fruit)|dates]], whereas Tafforet mentioned seeds and leaves. No other accounts mention diet.<ref name="Fuller2001"/> It has been suggested it ate [[latan palm]] fruits, for which it competed with the now extinct ''[[Cylindraspis]]'' tortoises. It is not known how the young were fed, but related pigeons provide [[crop milk]].<ref name="ChekeHume2008p45"/> The risings on the crop of the female may have covered glands that produced the crop milk. If the theory is correct, the birds may have practiced a division of labour, where the female stayed and fed the young crop milk, while the male collected food in the crop and delivered it to the female. It has been suggested that the maximum size attained by the solitaire and the dodo was limited by the amount of crop milk they were able to produce for their young during early growth.<ref>{{cite journal |doi= 10.1093/auk/122.3.1003 |last1= Storer |first1= Robert W. |year= 2005 |title= A possible connection between crop milk and the maximum size attainable by flightless pigeons |journal= The Auk |volume= 122 |issue= 3 |pages= 1003–1004 |doi-access= free }} {{free access}}</ref>

Several contemporary accounts state that the Rodrigues solitaire used [[gizzard stone]]s. Dodos also did this, which may imply a similar diet.<ref name="ChekeHume2008p45"/> Leguat described the stones in the following passage, mentioning that Rodrigues solitaires refused to feed in captivity:

{{blockquote|Tho' these Birds will sometimes very familiarly come up near enough to one, when we do not run after them, yet they will never grow Tame. As soon as they are caught they shed Tears without Crying, and refuse all sustenance till they die.
We find in the Gizzards of both Male and Female, a brown Stone, of the bigness of a Hen's Egg, 'tis somewhat rough, flat on one side and round on the other, heavy and hard. We believe this Stone was there when they were hatched, for let them be never so young, you meet with it always. They never have but one of 'em, and besides, the Passage from the Craw to the Gizard is so narrow, that a like Mass of half Bigness cou'd not pass. It serv'd to whet our Knives better than any other Stone Whatsoever.<ref name="Rothschild1907p177"/>}}

In 1877 three stones were found in a cavern on Rodrigues, each near a Rodrigues solitaire skeleton, and were inferred to be the gizzard stones mentioned by Leguat. One of the stones was examined and found to be [[dolerite]]: somewhat rough, hard and heavy, {{circa|{{convert|50|g|frac=4|abbr=on}}}}, but hardly flat on one side as described by Leguat. This could be due to its association with a young individual.<ref name="NewtonClark1879"/> Although Leguat asserted that the bird hatched with the gizzard stone already inside, in reality adults most likely fed the stones to their hatchlings.<ref name="ChekeHume2008p45"/>

===Reproduction===

[[File:Pezophaps solitaria recreation.jpg|thumb|right|[[Photo collage]] restoration of a nesting female and a male in their environment<ref>{{cite journal |doi= 10.1080/08912963.2014.954569 |url= https://www.researchgate.net/publication/266260711 |title= Digital reconstruction of Rodrigues Solitaire (Pezophaps solitaria) (Aves: Columbidae) physical appearance based on early descriptive observation and other evidence |journal= Historical Biology |volume= 28 |issue= 3 |pages= 1–17 |year= 2014 |last1= Rodríguez-Pontes |first1= M. N. A.|s2cid= 86229916 }}</ref>|alt=Greyish bird]]
The most detailed account of the reproductive habits of the Rodrigues solitaire is Leguat's. He described mating and nesting as follows:

{{blockquote|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. The Male and Female both cover it in their turns, and the young is not hatch'd till at seven Weeks' end: All the while they are sitting upon it, or are bringing up their young one, which is not able to provide itself in several Months, they will not suffer any other Bird of their Species to come within two hundred Yards round of the Place; But what is very singular, is, the Males will never drive away the Females, only when he perceives one he makes a noise with his Wings to call to the Female, and she drives the unwelcome Stranger away, not leaving it till 'tis without her Bounds. The Female do's the same as to the Males, whom she leaves to the Male, and he drives them away. We have observ'd this several Times, and I affirm it to be true.
The Combats between them on this occasion last sometimes pretty long, because the Stranger only turns about, and do's not fly directly from the Nest. However, the others do not forsake it till they have quite driven it out of their Limits. After these Birds have rais'd their young One, and left it to itself, they are always together, which the other Birds are not, and tho' they happen to mingle with other Birds of the same Species, these two Companions never disunite. We have often remark'd, that some Days after the young leaves the Nest, a Company of thirty or forty brings another young one to it, and the now fledg'd Bird, with its Father and Mother joyning with the Band, march to some bye Place. We frequently follow'd them, and found that afterwards the old ones went each their way alone, or in Couples, and left the two young ones together, which we call'd a Marriage.<ref name="Rothschild1907p177"/>}}

The [[clutch (eggs)|clutch]] was described as consisting of a single egg; given the bird's large size, this led to proposals that the solitaire was [[K-selected]], which means it produced a low number of [[altricial]] offspring, which required extensive parental care until maturity. The gathering of unrelated juveniles suggests that they formed [[crèche (zoology)|crèches]], which may have followed foraging adults as part of the learning process.<ref name="Livezey1993"/> A study of subfossil remains found that the carpal knob only developed after the bird reached skeletal maturity.<ref name="HumeSteel2013"/>

{{multiple image
|align = right
|total_width = 300
|image1 = Pezophaps solitaria.jpg
|alt1 = Illustration of the skeletons of a small female and large male solitaire
|caption1 = Skeletons of a female and male collected in 1874
|image2 = Pezophaps.jpg
|caption2 = Skulls of a male and female in several views
|alt2 = Skull of male (1–3) and female (4–5) Rodrigues solitaires
}}

Tafforet's account confirms Leguat's description of reproductive behaviour, adding that Rodrigues solitaires would even attack humans approaching their chicks:
{{blockquote|They do not fly at all, having no feathers to their wings, but they flap them, and make a great noise with their wings when angry, and the noise is something like thunder in the distance. They only ly, as I am led to suppose, but once in the year, and only one egg. Not that I have seen their eggs, for I have not been able to discover where they lay. But I have never seen but one little one alone with them, and, if any one tried to approach it, they would bite him very severely. These birds live on seeds and leaves of trees, which they pick up on the ground. They have a gizzard larger than the fist, and what is surprising is that there is found in it a stone of the size of a henn's egg, of oval shape, a little flattened, although this animal cannot swallow anything larger than a small cherry-stone. I have eaten them: they are tolerably well tasted.<ref name="Rothschild1907p177"/>}}
The size difference between sexes has led to the suggestion that the Rodrigues solitaire was not monogamous as stated by Leguat, and that this deeply religious man attributed the trait to the bird for moral reasons.<ref name="NewtonClark1879"/> It has been proposed that it was instead [[Polygyny in animals|polygynous]], and the wing-rattling behaviour described for males suggests [[lek-mating]], where males gather for competitive [[mating display]].<ref name="Livezey1993">{{cite journal |doi= 10.1111/j.1469-7998.1993.tb02686.x |last= Livezey |first= B. C. |year= 1993 |title= An Ecomorphological Review of the Dodo (''Raphus cucullatus'') and Solitaire (''Pezophaps solitaria''), Flightless Columbiformes of the Mascarene Islands |journal= Journal of Zoology |volume= 230 |issue= 2 |pages= 247–292 }}</ref> However, size dimorphism does occur in some monogamous birds; most other pigeons are monogamous as well.<ref name="HumeWalters2012p137"/> A 2015 article proposed that males invited females into their territories as secondary mates, which would result in the resident female acting aggressively towards the newcomer. Similar behaviour is seen in species that practice resource-defence polygyny. The territories probably provided all the food the birds needed in addition to acting as breeding-areas, and there was probably intense competition for favourable territories. The fact that Rodrigues island shrank by 90% at the end of the [[Pleistocene]] may also have contributed to such competition over territories, and thereby furthered sexual dimorphism.<ref>{{cite journal|doi=10.1111/ibi.12329|first1=Ingvar|last1=Byrkjedal|first2=Gaute|last2=Grønstøl|first3=Terje|last3=Lislevand|title=Possible resource-defence polygyny in the extinct Rodrigues Solitaire ''Pezophaps solitaria'' (Columbidae: Raphini|journal=Ibis|volume=158|issue=1|pages=199–201|date=January 2016|doi-access=free}} {{free access}}</ref>