Nesoenas mayeri (Prévost & Knip, 1843)

Hume, Julian Pender, 2011, 3124, Zootaxa 3124, pp. 1-62 : 8-12

Pink Pigeon Nesoenas mayeri (Prévost & Knip, 1843)

Columba mayeri Prévost & Knip 1843, p. 113 , pl. 60; Jouanin 1965, p. 975.

Columba meyeri: Schlegel and Pollen, 1868: 111 , pl. 36.

Carpophaga meyeri: Gray, 1849 , vol. ii: 45.

Peristera mayeri: Gray, 1854 , vol. iii., App: 24.

Trocaza meyeri: Bonaparte, 1854 , vol. ii: 45.

Trocaza meijeri: Pollen, 1863 , vol.i: 318.

Nesoenas mayeri: Salvadori, 1893 , vol. XXI p.327; Peters, 1937, vol. 3, p.74; Baptista et al. 1997, 4, 132, pl. 6; Cheke, 2005, p.294.

Streptopelia mayeri: Johnson et al., 2001 , 118(4), p.883.

Holotype: The authorship status of Nesoenas mayeri is confusing. The first description of N. mayeri was written by Prévost in Knip (1843: 113), which included a coloured illustration by Madame Knip, née Pauline de Courcelles (Knip 1843: Pl.60), this being the second volume that she illustrated on pigeons. Florent Prévost had written the text for the second volume, with Knip as senior author. Prévost had adopted the name from a manuscript description by one M. Marchel, who also owned the specimen, and Knip had illustrated the same bird; thus there was a specimen from which the painting and description were taken. However, this specimen no longer exists or remains unrecognised as the type. In earlier editions of the work, Knip’s entitlement to the role of senior author is riddled with irregularities (see Dickinson et al. 2010), but as the type specimen is no longer extant and Knip had obviously illustrated the same bird described by Prévost, following Jouanin (1962) and Dickinson et al (2010), the correct citation should be: Nesoenas mayeri Prévost & Knip in Knip, Les Pigeons, 1843.

Measurements: See Appendix 1.

Type locality: Mauritius, Mascarenes.

Distribution: Mauritius, Mascarenes.

Etymology: Jobling (1991) erroneously stated that the species is named after August Franz Joseph Carl Mayer (1787-1865), German anatomist and collector. The species was not mentioned at all in a later edition ( Jobling 2009). Prévost (1843:113) clearly stated that the species was named in honour of M. Gustave Mayer, details regarding whom I have not managed to trace. M. Gustave Mayer was based on Mauritius and presumably sent the first specimen of N. mayeri to M. Marchal; he also supplied some brief notes on the birds’ habits.

Referred fossil material: Subfossil material collected by Etienne Thirioux from unspecified caves in the valleys of Le Pouce and elsewhere, Mauritius. Possibly associated individuals: UMZC 583, including anterior portion of cranium, humerus (R) and (Lp), ulna (Rp), tibiotarsus (L) and (Rp), tarsometatarsus (R); and UMZC 582, including anterior portion of sternum, humerus (L) and (Rd), femur (R) and (L), tibiotarsus (Ld), and tarsometatarsus (L). Unassociated elements: sternum: four sterna MNHN u/r, largest and best preserved anterior fragment lacking incisura intercostalis intact and trabecula lateralis, rostrum sterni complete, but carina sterni lacking apex carinae; second best preserved specimen has right incisura intercostalis and proximal trabecula lateralis, but lacks carina sterni; third specimen has complete rostrum sterni and apex carinae almost complete; the last is an anterior fragment with complete rostrum sterni; UMZC 581 (anterior fragment); UMZC 581 (anterior fragment); UMZC 584 (anterior fragment); UMZC 584 (anterior fragment); UMZC 413D (anterior fragment); MNHN MAD 6988; MNHN MAD 6987 (anterior fragment); MNHN MAD6990 (anterior fragment); coracoid: UMZC 581 (R); UMZC 584 (R); UMZC 584 (L); MNHN MAD7142 (R); MNHN MAD7142 (R); MNHN MAD7142 (R); MNHN MAD7142 (R); MNHN MAD7142 (R); MNHN MAD7142 (L); MNHN MAD7142 (L); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (R); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (L); MNHN MAD6935 (Lp); humerus: UMZC 581 (R); UMZC 596 (L); UMZC 596 (L); UMZC 596 (L); UMZC 596 (L); UMZC 596 (L); UMZC 596 (R); UMZC 596 (R); UMZC 584 (L); MNHN MAD6994 (R); MNHN MAD6811 (R); MNHN MAD6808 (L); MNHN MAD7006 (L); MNHN 6802 (L); MNHN MAD7019 (L); MNHN MAD6997 (L); MNHN MAD6998 (L); MNHN MAD6991 (L); MNHN MAD6815 (R); MNHN u/r (R); MNHN MAD6995 (L); MNHN MAD7016 (L); MNHN MAD7002 (R); MNHN MAD6814 (R); MNHN MAD6999 (R); MNHN MAD7014 (R); MNHN MAD7004 (L); MNHN u/r (Lp); MNHN u/r (Lp); MNHN MAD7009 (Lp); MNHN MAD6993 (Rd); MNHN MAD7010 (Rd); MNHN MAD7012 (Rd); ulna: UMZC 584 (Rd); UMZC 413.D (R); UMZC 413.D (R); UMZC 584 (R); UMZC 584 (Ld); UMZC 583 (Rp); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/r (Lp); carpometacarpus: UMZC 582 (L); UMZC 584 (R); UMZC 583 (Rp); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6936 (R); MNHN MAD6908 (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); pelvis: UMZC u/r; UMZC 8; UMZC 584 fragment; UMZC 584 fragment; UMZC 584 fragment; femur: UMZC 584 (R); UMZC 584 (R); UMZC 584 (R); UMZC 584 (L); UMZC 584 (L); MNHN MAD7037 (R); MNHN MAD7039 (R); MNHN MAD7026 (R); MNHN 7025 (R); MNHN MAD7032 (R); MNHN MAD7028 (R); MNHN MAD7027 (R); MNHN MAD7034 (R); MNHN MAD7042 (L); MNHN MAD7036 (L); MNHN MAD7035 (Ld); MNHN MAD7044 (Ld); MNHN MAD7033 (Ld); MNHN MAD6895 (L); MNHN MAD6896 (Lp); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (L); MNHN MAD u/c (R); MNHN MAD u/c (R); MNHN MAD u/c (R); tibiotarsus: UMZC 584 (L); UMZC 584 (L); UMZC 584 (L); UMZC 584 (L); UMZC 584 (L); UMZC 584 (R); UMZC 584 (R); tarsometatarsus: UMZC u/r; UMZC 584 (L); UMZC 584 (L); UMZC 584 (R); UMZC 584 (R); UMZC 584 (Rd); UMZC 583 (R); MNHN MAD6871 (R); MNHN MAD6873 (L); MNHN MAD6865 (L); MNHN MAD6875(L); MNHN MAD6868 (R); MNHN MAD6858 (L); MNHN MAD6870 (R); MNHN MAD6879 (L); MNHN MAD6856 (L); MNHN MAD6877 (L); MNHN MAD6878 (R); MNHN MAD6859 (L); MNHN MAD6861 (L); MNHN MAD6872 (R); MNHN MAD6876 (R); MNHN u/r (R); MNHN u/r (R); MNHN u/r (R); MNHN u/r (R); MNHN u/r (R); MNHN u/r (R); MNHN u/r (L); MNHN u/r (L); MNHN u/r (L); MNHN u/ r (L); MNHN u/r (Rd); MNHN u/r (Rd).

Mare aux Songes marsh subfossil material collected by Théodore Sauzier in 1889 ( Newton and Gadow, 1893): sternum UMZC 413D (almost complete lacking anterior end); UMZC 413D (posterior fragment); ulna UMZC 413D (R); UMZC 413D (R); tarsometatarsus UMZC 413D (L).

Diagnosis: As for genus.

Description and comparison: The largest Mauritian species of Nesoenas ( Fig.2). Differs from all other Mascarene columbids and introduced Columba livia by the following suite of characters: cranium rounded with large orbits, os lacrimale wide, small indistinct fonticuli orbitocraniales, processus postorbitalis comparatively sharp and pointed, rounded and distinct condylus occipitalis, without medial groove running anterioposterior across ala parasphenoidalis complex ( Figs. 3a, 3b); sternum, comparatively reduced compared with other similar-sized pigeons, e.g. C. livia , no constriction between incisura lateralis, fenestra medialis absent or minute, foramen pneumaticum shallow, pila coracoidea square-shaped, carina sterni reduced ( Figs. 4a, 4b); humerus, robust with straight shaft, crista bicipitalis less right-angled from shaft; radius morphologically similar to other Mauritian columbids only longer, comparatively gracile compared with C. livia , sulcus tendinosa less excavated and extends further proximad; femur, distinct latero-medial curve at proximal end, shaft comparatively robust, tibiotarsus, pronounced ridge directed medially on epicondylus medialis; tarsometatarsus, shaft comparatively straight, crista lateralis hypotarsi prominent, directed plantad, not proximad.

Remarks: Considering the rapidity with which all other Mascarene pigeons became extinct following human arrival, it is surprising that N. mayeri has survived. One factor that may have contributed to this is an apparent seasonal inedibility due to its flesh becoming poisonous at certain times of year, described in many early accounts as well as in recent documentation (e.g. Meinertzagen 1912; Jones 1987). Such was the interest in its poisonous nature that virtually no other information was recorded about the biology of N. mayeri until the late 20 th century ( Cheke & Hume 2008). This phenomenon was also attributed to the Echo Parakeet Psittacula echo ( Cheke 1987; Hume 2007) and to some extent to Alectroenas nitidissima . Frequent discussion of this phenomenon ( Snouckaert 1935; Guérin 1940 -53; McKelvey 1976; Cheke 1987; Jones 1987; Temple in Jones 1987; Staub 1993) has failed to provide conclusive evidence as to its cause in N. mayeri .

The first mention of poisonous pigeons on Mauritius was made by Reyer Cornelisz, who remained on Mauritius for three months in 1602 (Begin en de voortgangh 1646: 30) and wrote a journal during his stay. The red-tailed birds he mentions could be either N. mayeri or A. nitidissima :

In this country [of Mauritius] occur……doves, some of which have red tails (by eating which many of the crew were made sick)……[ Strickland & Melville 1848: 125].

The account of Steven van der Hagen, who visited Mauritius in 1607 (Begin ende Voorgang 1646) is more specific, accurately describing N. mayeri . First, he discounts any suggestion that fish were responsible for poisoning-- -his crew having eaten plenty with no ill effects---but then goes on to query pigeons as the cause:

It [the illness] consists in a prostration or dejection felt in all the limbs, but once the illness is over, one is all the better for having it. Others have blamed this illness on the pigeons (these have red plumage on their bodies as well as their tails) which one eats there; nor can this either be true, for some of those who had eaten pigeon were not ill, and those who had been ill ate much pigeon after they had recovered, and found the flesh very good [ Barnwell 1948: 18].

By the 18th century, the supposedly poisonous nature of N. mayeri was widely known (see Cheke & Hume

2008). Desjardins ( Oustalet 1897: 72) even remarked on a technique for avoiding ill effects:

This bird [ N. mayeri ] is rare; it is ensured that the head should be cut off at once after it is killed, or otherwise it poisons [my translation].

Desjardins presumably meant bleeding the birds by removal of their heads, as the flesh would be bitter unless this procedure was carried out (see Jones 1987). Jean-François Charpentier de Cossigny, who was a military combat engineer working for the French East India Company, in 1755 sent a letter to René-Antoine Réaumur, a member of the Académie des Sciences in Paris ( Cossigny 1732 -55), describing the symptoms in some detail (see Cheke 1987 for full translation). He clearly distinguished N. mayeri from A. nitidissima , but was unable to determine the source of the poison.

In 1755, Cossigny wrote another letter to Réaumur (see Cheke 1987 for full translation), who had specifically asked Cossigny to resolve the poisonous pigeon phenomenon. He dissected a pink pigeon and indicated that digestion of a nightshade was responsible for making the flesh toxic. True nightshades ( Solanaceae ) do not occur in the Mascarenes, but two species of plants on which both Pink Pigeons and the Echo Parakeet feed, the native fandamane Aphloia theiformis (Flacourtiaceae) and endemic fangame Stillingia lineata ssp. lineata (Euphorbiaceae) , have been suggested as the source ( McKelvey 1976; Temple in Jones 1987; Jones 1987). Due to their present rarity, experimental consumption of Pink Pigeons obviously cannot be undertaken: thus this phenomenon has yet to be resolved.

Present status: The historic record of the Pink Pigeon is one of a gradually declining population, presumably as a result of human hunting, predation from introduced animals, and deforestation. Desjardins noted that it was rare in 1830 (see above), and Edward Newton similarly remarked on the scarcity of the species 30 years later ( Newton 1861). The decline continued into the 20th century. Meinertzhagen (1912) stated that a few remained in the south-west but were increasing, whereas Guérin (1940 -53) spoke of their imminent extinction in 1940. However, it appears that a small, stable population of c.50-60 birds remained in the south-west Black River Gorges area until about 1970. A sequence of natural disasters and increased deforestation in the pigeons’ remaining strongholds then reduced the population to as few as twenty birds by the mid 1970s and even fewer by the beginning of the 1980s (see Jones 1987 for details). Although the species then appeared to be doomed to extinction, intensive conservation management has since resulted in the population increasing to 300+ today (see Jones in Cheke & Hume 2008). Unfortunately N. mayeri is still vulnerable to introduced animals, particularly cats and rats, diseases are a continuing concern ( Swinnerton et al. 2005a,b), and supplementary feeding remains important to the species’ survival ( Edmunds et al. 2008), which would also make it difficult to determine anything about its poisonous qualities.