Psittacula echo ( Newton & Newton, 1876 )

HUME, JULIAN PENDER, 2007, Reappraisal of the parrots (Aves: Psittacidae) from the Mascarene Islands, with comments on their ecology, morphology, and affinities, Zootaxa 1513 (1), pp. 1-76 : 21-22

Echo Parakeet Psittacula echo ( Newton & Newton, 1876) .

Perroquet vert, Cossigny, 1732: 168 –96,205–82, 305–16., in Trieze letters de Cossigny á Reaumer. Recueil trimestriel de documents et travaux inédits pour servir á l’histoire des Mascareignes françaises. [Original not seen.]

Perruche; La Motte 1754, 1756., in Cheke 1987: 45. Mascarene Island Birds. Cambridge University Press. [Original not seen]

Palaeornis echo Newton & Newton, 1876: 284 ; Rothschild 1907: 68.

Palaeornis eques ; Salvadori, 1891: 442; Meinertzhagen, 1912: 94.

Psittacula echo ; Peters, 1937: 243.

Measurements: See Appendix 3.

Type locality: Mauritius, Mascarenes.

Distribution: Mauritius, Mascarenes.

Etymology: From Latin echo , meaning a woodnymph.

Syntypes: UMZC18 / Psi /67/k/1-4. Collected on Mauritius, 2 immature males, Vacoa, 12 Apr and male and female on 31

Dec 1860 by C.E.Banks, Bois Sec, Oct 1873.

Referred material: Despite this species being known from at least 34 museum skins ( Cheke 1987), their comparative use is limited therefore they are not included here. Trunk skeleton including sternum, pelvis, femora, and coracoids removed from spirit specimen of captive bird BMNH S/2000.44.1. Subfossil material collected by E. Thirioux, Le Pouce, Mauritius and elsewhere: rostrum MNHN MAU528; mandible MNHN MAU599; MNHN MAU509; MNHN 760; MNHN MAU602; MNHN MAU573 (juv); palatine MNHN MAU593(L); sternum UMZC 599(p); UMZC 599(p); UMZC 599(p); UMZC 599(p); UMZC 600(p); coracoid UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(R); UMZC 600(L); MNHN MAU534(L); MNHN MAU516(L); MNHN MAU589(R); MNHN MAU578(R); MNHN MAU543(R); MNHN MAU504(L); MNHN MAU577(R); MNHN MAU545(R); MNHN MAU554(R); MNHN MAU588(R); humerus UMZC 600(L); UMZC 600(Lp); MNHN u/c(L); MNHN u/c(R); MNHN u/c(L); MNHN MAU505(L); MNHN MAU512(L); MNHN MAU586(R); MNHN MAU506(L); MNHN MAU517(R); MNHN MAU552(R); MNHN MAU498(L); MNHN MAU551(R); MNHN MAU7017(L); MNHN MAD7000(L); MNHN u/c(L); ulna UMZC 600(L); UMZC 600(R); MNHN MAU547(R); MNHN MAU495(R); MNHN MAU555(L); MNHN MAD7189(L); MNHN MAD7200(L); MNHN MAD7188(R); MNHN MAD7195(L); radius UMZC 600(L); carpometacarpus MNHN MAU500; MNHN MAU560; femur MNHN u/c(R); MNHN u/c(L); MNHN MAU570(L); MNHN MAU548(L); tibiotarsus MNHN u/c(L); MNHN u/c(L); MNHN MAU511(L); MNHN MAU601(R); MNHN MAU535(Rd); MNHN MAU529(Ld); MNHN MAU761(Rd); tarsometatarsus UMZC 594(Lp) (juv); UMZC 594(Lp) (juv); UMZC 594(Lp) (juv); UMZC 594(Lp) (juv); UMZC 594(Lp) (juv); UMZC 594(Lp) (juv); UMZC 594(Ld); UMZC 594(Ls); UMZC (Rp); UMZC (Rp); UMZC (Rp); UMZC 577(Rs); UMZC 593(Rs); UMZC 593(Rs); UMZC 593(Lp); UMZC 593(R); UMZC 593(L); MNHN u/c(L); MNHN 560(L); MNHN 582(L); MNHN 597(L); MNHN 546(R); MNHN 507(R); MNHN 579(R); MNHN 596(L); MNHN 758(R); MNHN 556(R); MNHN MAD6880(L); MNHN MAD6864(L); MNHN MAD6888(L) (juv).

Diagnosis: Apart from its distinctly smaller size, Psittacula echo may be distinguished from P. bensoni by the following suite of characters. Sternum; spina externa weakly bifurcated. Coracoid; lateral margin of angulus medialis pointed; proximal to facies articularis clavicularis, a small circular deeply excavated foramen present; processus procoracoideus short with a shallow cotyla scapularis.Humerus: shorter condylus dorsalis,

distal end mediolaterally less expanded and fossa olecrani shallow.Tarsometatarsus: impressiones retinaculi extensorii less defined. Despite molecular evidence for a sister relationship between P. echo and the Indian P. krameri (borealis) ( Groombridge et al. 2004), my morphological analysis indicates that P. echo is more closely related to P. eupatria than to P. krameri .

Description and comparison: See Appendix 2c, tables 1–11.

Remarks: This species, the last surviving Mascarene parrot, was reduced to fewer than 12 birds by the 1980s ( Cheke 1987; Jones 1999), but has now recovered to 250+ after conservation efforts (Cheke and Hume in prep.). La Motte in 1754 & 1756 described their former abundance:

One eats here [in Mauritius] a good number of long-tailed green parrots called perruches whose flesh is black and very good. A hunter can kill three or four dozen in a day. There is a time of year when these birds eat a seed that makes their flesh bitter and even dangerous [ Cheke 1987: 45].

Unsurprisingly, the Echo Parakeet is very rare in skeletal collections and its osteology is known mainly from fossil remains. It is easily distinguishable from all other Mauritian parrots by size, all elements being much smaller and less robust. It is intermediate in size between P. krameri and P. eupatria . The Echo Parakeet is the smallest Mauritian parrot and the one most frequently found in cave fossil deposits, where both adult and juveniles are represented. Groombridge et al. (2004) concluded from a DNA analysis that P. echo is derived from the Indian Ring-necked Parakeet Psittacula krameri borealis , as opposed to the nominate subspecies P. k. krameri in Africa ( Smith 1975). As no species of Psittacula occurs on Madagascar, it is reasonable to assume that P. krameri populated Africa via the Middle East and not the Indian Ocean islands.

The Echo Parakeet has been afforded species-level status, being stockier than P. krameri with shorter, more rounded wings, and a broader shorter tail ( Jones 1987; Forshaw 1989; Low 1994; Juniper & Parr 1998), and has been reported to measure 25% larger in weight and body size than P. krameri ( Low 1994) . The size difference in the live bird is also particularly noticeable in the bill ( Juniper & Parr 1998; Hume pers. obs.). Most available skeletal elements, when compared to its closest relative Psittacula krameri borealis ( Groombridge et al. 2004) , are similar in size except the hindlimb elements being 4.8% – 6.8% larger and the sternum 6.4% smaller. According to Glenny (1957), parrots generally have limited powers of flight compared to many other bird groups, but many are agile, swift fliers. Psittacula echo is an adept but not a long distance flier, and can rapidly maneuver between small openings in the forest canopy making clear observation of the birds difficult (pers. obs.). The reduction of the sternum thus appears to have had little affect on its flight capabilities. A similar proportional reduction in the sternum is found in the endemic Mauritian Pink Pigeon Nesoenas mayeri , when compared to similar-sized columbids (Hume unpubl.), also without any noticeable reduction in flight capabilities (Hume pers.obs.).

The Echo Parakeet feeds on buds, emergent shoots, leaves, flowers, berries, seeds, twigs and bark/sap, never descending on the ground to forage ( Jones 1987; Jones & Owaddally 1988; Forshaw 1989). This behaviour is in direct contrast to P.krameri , which regularly feeds on the ground ( Smith 1979) and is a bird of open areas, not dense forests ( Forshaw, 1989). If the other species of parrot already established on the Mascarenes were adapted to a ground niche, the most recently arrived may have had to adjust its habits accordingly; therefore, the different morphology of P.echo may be linked to an arboreal habitat adjustment ( Jones 1987).