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Rodrigues solitaire

The Rodrigues solitaire (Pezophaps solitaria) was a large, extinct flightless bird endemic to the Mascarene island of Rodrigues in the western Indian Ocean, closely related to the dodo (Raphus cucullatus) within the pigeon and dove family Columbidae.^[1]^[2] This species, part of the subfamily Raphinae, with males up to 90 cm (35 in) tall and females up to 70 cm (28 in) tall, had a robust, terrestrial build adapted to island life, including short legs, a slightly hooked beak, and reduced wings incapable of flight.^[3]^[1]^[4] First documented by French Huguenot settler François Leguat in 1691 during a brief colony attempt on Rodrigues, the bird was described in his 1708 travelogue Voyage et aventures de François Leguat as a solitary, non-flocking species with striking sexual dimorphism: males were predominantly grayish-brown, while females exhibited lighter blonde or brown plumage.^[3] Skeletal remains, including around 5,000 fossils discovered since the 19th century, have confirmed these accounts and revealed unique adaptations, such as bony "musket ball"-like knobs (up to 3.3 cm in diameter) on the wings of adult males, covered in hardened skin and used as defensive clubs in territorial disputes.^[5] These structures, absent or smaller in females, developed primarily after males secured mates and territories, producing a loud rattling or thunderous sound during confrontations.^[5] The Rodrigues solitaire inhabited dry lowland forests and possibly scrublands on Rodrigues, where it foraged terrestrially for hard seeds, fallen fruits, and leaves, laying a single large egg about 10 cm long in simple nests of palm leaves.^[1]^[3] Genetically, it forms a clade with the dodo and the Nicobar pigeon (Caloenas nicobarica), its closest living relative, supporting an evolutionary history of island colonization and flightlessness among Mascarene columbids.^[2] Human activities drove the species to extinction in the late 18th century, with the last confirmed sightings in 1761 and no reports after 1778; sailors and settlers hunted it extensively for food, while introduced goats, cats, and rats devastated its habitat and preyed on eggs.^[1]^[6] Classified as Extinct by the IUCN Red List, the Rodrigues solitaire exemplifies the rapid biodiversity loss on isolated islands following European contact.^[1]

Taxonomy and Evolution

Classification

The Rodrigues solitaire is classified within the kingdom Animalia, phylum Chordata, class Aves, order Columbiformes, family Columbidae, subfamily Raphinae, subtribe †Raphina, genus Pezophaps, and species P. solitaria.^[7]^[1]^[8] The species was first formally described in 1789 by Johann Friedrich Gmelin, who named it Didus solitarius based on historical accounts from Rodrigues Island, placing it initially near the dodo (Raphus cucullatus) due to superficial similarities in descriptions of flightlessness and size.^[7]^[9] In 1848, Hugh Edwin Strickland established the genus Pezophaps (from Greek pezos meaning "walking" and phaps meaning "pigeon") specifically for this species, using subfossil remains to confirm its distinct columbiform affinities and reclassify it away from the dodo genus.^[10] A 2024 taxonomic review confirmed this classification and proposed the subtribe †Raphina to unite Pezophaps and Raphus as sister taxa within Raphinae.^[8] Historical synonyms include Didus solitarius (the original binomial), Pezophaps minor (a junior synonym), and Pezophaps solitarius (a variant spelling), reflecting early uncertainties in its placement.^[7] It was sometimes misclassified as a close relative of the dodo under Didus or even as a type of rail, but subfossil evidence solidified its position within Columbidae.^[10] The Rodrigues solitaire shares a close relation with the dodo and the living Nicobar pigeon (Caloenas nicobarica), forming part of the Raphinae radiation.^[7]

Phylogenetic Relationships and Evolution

The Rodrigues solitaire (Pezophaps solitaria) belongs to the family Columbidae and forms a monophyletic clade with the extinct dodo (Raphus cucullatus), as confirmed by ancient DNA analysis of mitochondrial cytochrome b sequences extracted from subfossil bones.^[11] This study positions the solitaire and dodo as sister taxa within the pigeons, with their closest living relative being the Nicobar pigeon (Caloenas nicobarica), an Indo-Pacific species, based on shared genetic markers that place them basal to other columbid lineages such as the crowned pigeons (Goura spp.).^[11] Subsequent mitogenomic analyses using complete mitochondrial genomes from solitaire subfossils further refine this phylogeny, confirming the solitaire-dodo clade's placement within an Indo-Pacific radiation of pigeons and highlighting minimal genetic divergence from flying ancestors despite morphological extremes.^[12] Evolutionary divergence of the solitaire lineage from continental pigeon ancestors occurred during the Miocene, with molecular clock estimates indicating separation from the Caloenas lineage approximately 18 million years ago (95% CI: 13.6–23.2 million years ago), predating the volcanic formation of the Mascarene Islands by millions of years and suggesting initial dispersal via oceanic stepping-stones from Southeast Asia.^[12] Subfossil evidence, primarily from late Holocene deposits (ca. 4,000–200 years before present) in Rodrigues caves, documents the solitaire's presence on the island until human arrival around 1,300 years ago, with no earlier fossils indicating a protracted island-specific evolution.^[12] Flightlessness in the solitaire evolved convergently from flying pigeon stock, involving reductions in wing bone length and sternal keel depth compared to volant relatives, adaptations that minimized energy costs in a predator-free environment but retained robust forelimb musculature.^[13] A distinctive adaptation was the development of carpal knobs—bony exostoses on the carpometacarpus—forming late in ontogeny after skeletal maturity and serving as combat weapons for territorial disputes, unique among columbids and linked to intraspecific aggression in dense island populations.^[14] Recent modeling integrates population dynamics with game theory to explain heightened territoriality and aggression in the solitaire, attributing these traits to severe habitat contraction on Rodrigues, where post-Pleistocene sea-level rise and erosion reduced island area by up to 90% over 20,000 years, far exceeding losses on neighboring Mauritius and favoring aggressive "hawk" strategies for resource defense below critical population thresholds.^[15] This contrasts with the dodo's more passive behavior, underscoring how insular habitat dynamics drove divergent evolutionary pathways within the clade.^[15]

Physical Description

Morphology and Size

The Rodrigues solitaire (Pezophaps solitaria) exhibited pronounced sexual size dimorphism, with males reaching heights of 75.7–90 cm and weights up to 28 kg, while females measured 63.8–70 cm in height and approximately 17 kg in weight.^[16]^[17] This dimorphism, among the greatest recorded in carinate birds, is evident in subfossil remains, where male skeletal elements consistently exceed female dimensions by about 40%.^[17] Key morphological features included a long neck and elongated legs adapted for browsing vegetation at various heights, with subfossil tibiotarsi disproportionately lengthened relative to other columbids.^[17] The wings were vestigial and reduced, featuring prominent carpal knobs on the carpometacarpus that reached depths of up to 32.9 mm in males, far exceeding the 7.6 mm maximum in females.^[18] The bill was robust and hooked, resembling that of its relative the dodo (Raphus cucullatus) but shorter and more slender, measuring around 25 mm in length based on skull reconstructions.^[16]^[17] Skeletal analyses of subfossils reveal adaptations supporting a heavy body mass, including robust leg elements with greater relative shaft widths than in flying pigeons, enabling terrestrial locomotion.^[17] Bone ratios, such as the elongated tibiotarsi paired with shorter tarsometatarsi and digits, indicate an upright posture that facilitated stability and reach in foraging.^[17]

Plumage and Adaptations

The plumage of the Rodrigues solitaire (Pezophaps solitaria) was described as predominantly grey-brown, with males exhibiting a darker brown-grey coloration overall compared to females, which were paler and sometimes featured lighter, fair-haired tones on parts of the body along with white chest feathers. The feathers had a hair-like, plush texture, and sparse iridescence was noted on the neck, distinguishing it from the more vibrant displays seen in mainland pigeons. Subdued grey tones on the underparts provided a paler contrast to the darker back, contributing to an overall cryptic appearance suited to the island's sparse vegetation. The tail was notably short and rounded, resembling a horse's crupper with scarcely any protruding feathers, which aligned with the bird's flightless morphology and emphasized its ground-dwelling adaptations. This rounded hindpart lacked the elongated or iridescent features typical of flying columbids, further reducing visibility in the understory. Key adaptations included a short, sharp, incurved bill hooked at the tip like that of a turkey, measuring approximately 1 inch in length and suited for processing tough vegetation through its robust structure. The wings bore prominent carpal knobs—bony outgrowths on the carpometacarpus, larger in males and comparable in size to a musket ball—which were covered in thick skin for protection during use as weapons in territorial combat. These knobs formed post-maturity and enabled audible signaling and physical defense, reflecting the bird's solitary, territorial lifestyle where subdued plumage likely aided in intraspecific crypsis rather than predator avoidance in the absence of natural threats. Males were generally larger than females, enhancing dimorphism in these combat-related features.

Ecology and Behavior

Habitat and Diet

The Rodrigues solitaire (Pezophaps solitaria) was endemic to Rodrigues Island, part of the Mascarene archipelago in the western Indian Ocean, where it occupied dry forests and coastal scrublands on the island's volcanic terrain.^[1] These habitats featured native vegetation adapted to the island's limited freshwater availability and rugged landscape, providing suitable cover and food resources for the flightless bird.^[1] The bird's diet primarily consisted of plant matter, including hard seeds, fallen fruits, and leaves from native species such as Latania palms (Latania loddigesii and L. verschaffeltii) and Pandanus screwpines (Pandanus spp.).^[19]^[16] Historical accounts from François Leguat, who observed the species in 1691–1693, described it feeding on dates and other fruits, while later reports by Julien Tafforet in 1725 noted consumption of seeds and foliage.^[19] To process this tough, fibrous vegetation, the solitaire ingested gastroliths—smooth stones held in its muscular gizzard to grind food—similar to its relative, the dodo; such stones were later collected from sites associated with solitaire remains after the bird's extinction.^[20] As a ground-dwelling species, the Rodrigues solitaire foraged by browsing on the forest floor in a solitary or paired manner, relying on its strong legs to navigate the understory for accessible plant material without the need for flight.^[1] This behavior aligned with its terrestrial adaptations and the availability of fallen fruits and seeds in its preferred habitats.^[16] Historical accounts describe the Rodrigues solitaire as forming monogamous pairs that remained together for life and bred within defended territories approximately 200 meters in radius, though modern analyses suggest possible resource-defence polygyny.^[3]^[21] These pairs constructed simple ground nests from heaps of palm leaves amid vegetation, which rendered the eggs and chicks particularly vulnerable to predation.^[3] Each breeding season, the female laid a single large egg, comparable in size to that of a goose, which both parents incubated alternately for about seven weeks (Leguat 1708), though modern estimates based on egg size suggest around 37 days.^[21]^[20] Following hatching, the helpless chick required several months of intensive parental care, during which the young bird remained dependent on the adults for feeding and protection.^[3] Leguat described a behavior where groups of 30 to 40 birds would bring another young to a pair, after which the adults would leave the two juveniles in an unoccupied territory; this has been interpreted as possible formation of small juvenile crèches rather than large communal rearing groups.^[3]^[21] This cooperative system likely enhanced chick survival in the resource-limited island environment, supported in part by the birds' fruit-based diet that provided essential nutrients for egg production and chick growth.^[3] Socially, the Rodrigues solitaire exhibited a territorial structure, with low population densities on the isolated island of Rodrigues contributing to its "solitary" reputation despite the paired and communal breeding behaviors.^[3] Both sexes defended their territories vigorously against same-sex intruders, while opposite-sex encounters involved courtship displays; males particularly employed their prominent carpal knobs—bony wing outgrowths—to deliver blows during aggressive territorial disputes.^[22] These knobs, unique among columbiforms, served as both weapons and signaling devices, producing rattling sounds when wings were whirled in warning postures.^[22]

Discovery and Human Impact

Historical Accounts and Descriptions

The Rodrigues solitaire was first documented in detail by François Leguat, leader of a group of French Huguenot refugees marooned on Rodrigues from 1691 to 1693. In his 1708 publication Voyages et Avantures de François Leguat & de ses Compagnons, Leguat described the bird—named for its solitary habits—as typically occurring in pairs that defended fixed territories aggressively, using their wings to buffet intruders with a knob-like bony projection that produced a rattling sound audible from afar. He portrayed it as a large, flightless species resembling a goose in size, with greyish-brown plumage in males, paler variations in females, a long neck, small useless wings, and a hooked beak; the birds were easily hunted by foot due to their slow gait and lack of flight, providing substantial meat that weighed up to 45 pounds when in season from March to September. Leguat's account emphasized the solitaire's behavioral traits, including lifelong monogamous pairing after raising a single large egg in a nest of palm leaves, with both parents sharing incubation for about seven weeks and the young becoming independent after several months. He noted their familiarity with humans yet refusal to be tamed, often shedding tears and starving in captivity, and highlighted their diet of fruits like dates from latanier palms. The book included one of the earliest illustrations of the living bird, depicting it alongside Rodrigues Island's landscape, which served as a primary visual reference for later naturalists. Additionally, Leguat recorded that the solitaires ingested gizzard stones—smooth, egg-sized pebbles retained in their stomachs to aid digestion—which island inhabitants collected from the birds to sharpen knives. Eighteenth-century observations by sailors and settlers reinforced Leguat's descriptions, portraying the solitaire as a readily available food source amid growing human pressures on the island. A key account came from Julien Tafforet, a French mariner from Réunion dispatched to assess Rodrigues for settlement and marooned there from 1725 to 1726; in his report, he described the solitaire as a hefty bird weighing 40 to 50 pounds, easily caught by hand or simple pursuit in open terrain, and suitable for provisioning ships or small groups. Tafforet echoed reports of its territorial aggression and scarcity in flocks, noting that pairs were widespread but vulnerable to overhunting by visitors. No live Rodrigues solitaires were ever transported to Europe, with all records limited to textual and illustrative accounts from the island itself, as the birds' size and fragility prevented successful shipment.^[22]

Fossil Evidence and Modern Research

Subfossil bones of the Rodrigues solitaire were first discovered in 1786 by French explorers led by Captain de Labistour, who found six bones encrusted in stalagmite within a cave, likely the Grande Caverne on the Plaine Corail, along with a tortoise bone; five of these were sent to the Muséum National d'Histoire Naturelle in Paris, while one went to the University Museum of Zoology in Cambridge. These early finds provided initial osteological evidence of the species, though limited in quantity. Subsequent explorations in the 19th century expanded collections significantly. Major excavations occurred in 1874 during a Royal Society expedition led by Henry H. Slater, who unearthed thousands of subfossil bones from caves and coastal dunes across the Plaine Corail region, including two nearly complete associated skeletons. These efforts built on earlier 19th-century collections by naturalists such as Edward Newton and George Jenner, who targeted solitaire remains in similar sites during the 1860s and 1870s. The resulting specimens, numbering in the thousands, are now housed in institutions like the Natural History Museum in London, enabling detailed anatomical studies. Modern osteological analyses of these subfossils have confirmed pronounced sexual dimorphism in the Rodrigues solitaire, with males exhibiting significantly larger body sizes and robust wing structures compared to females, potentially the most extreme among carinate birds.^[17] A 2024 study by Rijsdijk et al. utilized subfossil data to model population dynamics and paleoecology, linking the species' aggressive traits—such as reinforced wing knobs for territorial combat—to habitat contraction from post-glacial sea level rise on Rodrigues.^[13] In September 2025, Colossal Biosciences announced the generation of a 10-fold coverage genome from Rodrigues solitaire subfossils, providing genetic insights for de-extinction efforts related to its close relative, the dodo.^[23]

Extinction

Timeline and Causes

The Rodrigues solitaire (Pezophaps solitaria) was reported as common across much of Rodrigues Island in 1733, but experienced a rapid population decline in the following decades. By 1761, it persisted only as a rare species in isolated areas, with no confirmed sightings thereafter, and the bird is considered to have become extinct around the 1760s. The absence of subsequent records confirms its disappearance by the late 18th century.^[1] The primary causes of the solitaire's extinction were anthropogenic, beginning with direct hunting by French settlers, tortoise traders, and shipwrecked sailors who targeted the flightless birds for food due to their ease of capture and palatable meat. The intensive tortoise trade, which removed an estimated 280,000 individuals between 1735 and 1771, indirectly impacted the ecosystem by removing key species that maintained habitat structure and seed dispersal. Habitat destruction from clearance for agriculture, shipbuilding, and fuel, along with fires, further accelerated the decline and loss of the bird's native woodland environment. Introduced predators, including cats (arriving around the mid-18th century), pigs, and rats (introduced earlier but with delayed impact), preyed on eggs, chicks, and adults, while goats contributed through competition for resources and vegetation disturbance; cats in particular are noted for preventing successful breeding.^[1]^[24]

Conservation Legacy

The Rodrigues solitaire, alongside its relative the dodo, has become an iconic symbol of human-induced extinction, exemplifying the rapid loss of island biodiversity due to hunting and invasive species introduced by settlers.^[8] This pairing in historical accounts underscored the vulnerability of isolated ecosystems. The solitaire's story, though less popularized than the dodo's, reinforced emerging ideas about the irreversibility of extinction and the ethical responsibilities of colonization-era exploitation.^[25]^[26] In modern conservation, the solitaire's extinction has informed restoration initiatives on Rodrigues Island, where organizations like the Mauritian Wildlife Foundation (MWF) have led efforts to replant native forests since the early 2000s. These projects, including the restoration of reserves like Anse Quitor starting in 2009, aim to rehabilitate habitats degraded by similar historical pressures, fostering ecosystem recovery and preventing further losses among surviving endemics.^[27] Studies on rewilding potential draw lessons from the solitaire's former range, evaluating how restored forests could support analogous species and enhance island resilience.^[28] Genetic research following the extraction of ancient DNA in 2002 has fueled discussions on de-extinction, with the solitaire's mitochondrial genome sequenced to explore cloning feasibility using related pigeons as proxies.^[29] Recent advancements, including a 10-fold genome assembly by Colossal Biosciences in 2025, highlight the solitaire's role in understanding dodo-like traits, though practical de-extinction remains exploratory and not advanced.^[30] The International Union for Conservation of Nature (IUCN), through BirdLife International, cites the solitaire as a cautionary case for island endemics, stressing the ongoing threats from invasives and habitat loss to inform global strategies for protecting over 95% of at-risk island species.^[1]^[31]

References

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Rodrigues Solitaire Pezophaps Solitaria Species Factsheet
The Rodrigues Solitaire is extinct, with no mature individuals remaining. It was hunted to extinction in the 18th century, likely around the 1760s.Missing: facts | Show results with:facts
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The mysterious Spotted Green Pigeon and its relation to the Dodo ...
Jul 16, 2014 · More recent genetic analyses have revealed that the Dodo from Mauritius and the closely related Rodrigues Solitaire (Pezophaps solitaria), both ...
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Dodos and Solitaires | American Scientist
The raphid bird of Rodriguez, known generally today as the Solitaire, or scientifically Pezophaps solitaria (Gmelin), is in some ways the best known of the ...
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Angry bird's unique wing weapon revealed - Phys.org
May 28, 2013 · Natural History Museum scientists have discovered that the ball was in fact a deadly weapon, used by the angry bird to defend territory and its mate.
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Rodrigues Solitaire - Pezophaps solitaria - Birds of the World
Reported in 1761, but had become extinct by 1778 (2 ...Missing: facts | Show results with:facts
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https://birdsoftheworld.org/bow/species/rodsol2/cur/introduction
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[8]
Complete mitochondrial genomes of living and extinct pigeons ...
Oct 26, 2016 · This work also suggested that the dodo and solitaire lineages diverged 18–36 Million years ago (Mya), during the late Oligocene [13]. This date ...
[9]
Article Sea level rise and the evolution of aggression on islands
Nov 15, 2024 · ... Rodrigues solitaire to examine divergent pathways in the evolution of aggression. ... aggressiveness in the solitaire and island size reduction ...
[10]
Fight club: a unique weapon in the wing of the solitaire, Pezophaps ...
May 20, 2013 · The solitaire (Columbidae; Pezophaps solitaria) of Rodrigues was an extinct giant flightless pigeon and the sister taxon to the dodo ...<|control11|><|separator|>
[11]
https://www.researchgate.net/publication/11489398_Flight_of_the_Dodo
[12]
(PDF) Digital reconstruction of Rodrigues Solitaire (Pezophaps ...
Sep 29, 2014 · Rodrigues Solitaire (Pezophapssolitaria) is an extinct (eighteenth century) flightless bird endemic to Rodrigues Island (Mascarene Islands) ...
[13]
An ecomorphological review of the dodo (Raphus cucullatus) and ...
This paper describes a morphological study of the dodo Raphus cucullatus and solitaire Pezophaps solitaria extinct, flightless Columbiformes of the ...Missing: Rodrigues source
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A unique weapon in the wing of the solitaire, Pezophaps solitaria ...
The solitaire (Columbidae; Pezophaps solitaria) of Rodrigues was an extinct giant flightless pigeon and the sister taxon to the dodo (Columbidae; ...
[15]
The Dodo and the Tambalacoque Tree: An Obligate Mutualism ...
Aug 5, 2025 · ... Rodrigues solitaire fed on seeds,. including whole palm fruits (Latania spp.), and foliage. (Hachisuka 1953, Cheke 1985). This kind of diet is.
[16]
Mauritius extinct endemic bird species - Oiseaux-Birds
The Dodo became a rare bird and the last one was killed in 1681. The Rodrigues Solitaire (Pezophaps solitaria) was endemic to Rodrigues Island, E of Madagascar ...
[17]
Dodos and Solitaires (Raphidae) | Encyclopedia.com
Compared with the dodo, the Rodrigues solitaire was taller and more slender, with a smaller head and a slighter bill, metacarpal wing spurs, and largely ...Missing: structure | Show results with:structure
[18]
(PDF) Possible resource-defence polygyny in the extinct Rodrigues ...
The polygyny threshold model predicts that monogamous and secondary females on average settle at the same time and have similar reproductive success. This is ...
[19]
None
Nothing is retrieved...<|control11|><|separator|>
[20]
None
### Summary of Rodrigues Solitaire (Pezophaps solitaria) from Phelsuma 21b.pdf
[21]
systematics and nomenclature of the Dodo and the Solitaire (Aves
Aug 16, 2024 · Abstract. The Dodo and its extinct sister species, the Solitaire, are iconic exemplars of the destructive capabilities of humanity.Missing: timeline | Show results with:timeline
[22]
As Dead As A Dodo – How The Dodo Became Extinct | Bird Spot
The last confirmed sighting of a dodo was in 1662, recorded by Dutch sailor Volkert Evertsz, who had been shipwrecked on Amber Island, a small islet off ...
[23]
The dodo bird: The real facts about this icon of extinction
It had a large gizzard stone to help with digestion, which suggests it ate a diet of fallen fruits and seeds. CT scans of its braincase have revealed it had a ...Missing: habitat | Show results with:habitat
[24]
Restoration of Anse Quitor Nature Reserve (Rodrigues) - Project Detail
Mauritian Wildlife Foundation (Rodrigues Branch). Country: Mauritius. Area Of ... Start Date: 11/2009. End Date: 11/2011. Status: Satisfactorily Completed ...
[25]
Mauritian Wildlife Foundation
Our hands-on conservation projects are carried out in Mauritius including the offshore islets and Rodrigues. We work closely with local and international ...Projects · What is Friends of Mauritian... · Contact us · Ile aux AigrettesMissing: timeline | Show results with:timeline<|separator|>
[26]
[PDF] Flight of the Dodo
The evolutionary history of the dodo is very poorly understood. Like many avian island en- demics, a high degree of morphological change.Missing: et | Show results with:et
[27]
Reviving the Dodo - Colossal Biosciences
There are varying accounts of the last official sighting, but in 2003, David Roberts and Andrew Solow used a formula to place the dodo's final date of ...<|control11|><|separator|>
[28]
Managing invasive species on islands can curb global extinction rates
Oct 25, 2017 · Managing invasive species could benefit 95 per cent of Endangered and Critically Endangered amphibians, birds and mammals that live on islands.