Nightingale Reed-warbler

_Conopoderas luscinia nijoi_ Yamashina, Tori, 10, 1940, p. 674.

(Type locality, Agiguan.)

_Conopoderas luscinia nijoi_ Hand-list Japanese Birds, 3d ed.,

1942, p. 196 (Agiguan).

_Acrocephalus luscinia luscinia_ Mayr (part), Birds Southwest

Pacific, 1945, p. 294 (Agiguan).

_Geographic range._--Micronesia: Mariana Islands--Agiguan.

_Characters._--Adult: Resembles _A. l. luscinia_, but with shorter

bill. Yamashina (1940:674) describes the birds as, "upper parts

much less rusty in colour and the flanks and bellies are darker

and more brownish than those of the specimens from Almagan and

Saipan."

_Measurements._--Yamashina (1940:674) gives the measurements of

five adult birds from Agiguan as: exposed culmen 27-29, bill from

nostril 17.0-20.0; as compared with 27 adult birds from Almagan

and Saipan as: exposed culmen 30-34, bill from nostril 21.2-24.5.

_Remarks._--No specimens have been examined by me. The island of Agiguan

is a very small one lying offshore from Tinian and not far from Saipan,

where _A. l. luscinia_ occurs. _A. l. nijoi_ is given tentative

recognition, on the basis of the measurements of the five adult

specimens given by Yamashina. These indicate that the population has a

distinctly shorter bill.

_Evolutionary history of Acrocephalus luscinia._--The species of

_Acrocephalus_ in Micronesia and Polynesia have received several

taxonomic treatments. In regard to the Micronesian forms, Quoy and

Gaimard called the population at Guam _Thryothorus_ while Kittlitz

called the population in the Carolines, _Sylvia_. Evidently to emphasize

the distinctness of these two birds, Reichenbach in 1850 renamed the

bird in the Marianas as _Hybristes_ and the bird in the Carolines as

_Eparnetes_. The birds were later placed in the genus, _Tatare_, by

Hartlaub, Gray, Sharpe and other workers. Gray also used the name,

_Calamodyta_, for the bird in the Carolines. The generic term,

_Calamoherpe_, was employed also by a number of workers for the Caroline

population. Sharpe (1883:525) placed the reed-warblers in the family

Timelidae and retained the name, _Tatare_, for the Micronesian and

Polynesian forms. In distinguishing _Acrocephalus_ from _Tatare_ he has

the following to say of _Acrocephalus_: "besides having a much shorter

bill, possesses a very much more pointed wing, the distance between the

primaries and the secondaries being much more than the length of the

hind toe and claw; whereas in _Tatare_ the wing is much more obtuse, and

the distance between the primaries and the secondaries is less than the

length of the hind toe and claw." More recent authors have followed

Sharpe using the generic name, _Conopoderas_ (= _Tatare_, old name

preoccupied). However, Tristram (1883:38-46) regarded the separation of

these oceanic forms from _Acrocephalus_ as a taxonomic error. He said

that this is "one of the very few links (the others being the solitary

_Hirundo tahitica_ and the Merulae) between the avifauna of Oceania and

our own; and it has a much wider range east and west than either of the

other links, extending from the Carolines in the east to the Marquesas

in the west." Mayr has pointed out (orally to the writer) that the

separation of the Oceanic reed-warblers from _Acrocephalus_ is an

unnatural one, although it is perfectly true that the extreme members

(_A. caffra_ and _A. l. luscinia_) have a very long bill, but forms with

shorter bills like _A. l. syrinx_ point to the close affinity between

the continental species and these insular birds. This has also been

noted by Hartert (1898:58). Mayr (_in litt._) comments that "There is no

difference between _Acrocephalus_ and _Conopoderas_ in regard to the

wing formula, provided that we compare the Polynesian species with the

tropical forms of _Acrocephalus_ (such as _toxopei_ and _cervinus_). The

character mentioned by Sharpe is very artificial and merely indicates

the difference in the wing between a migrant of the temperate zone and a

resident of the tropics. There is no denying that some of the warblers

of eastern Polynesia are no longer reed-warblers but have become

dwellers of trees and bushes. However, this same tendency prevails among

some of the unquestionable species of _Acrocephalus_ (_scirpaceus_ and

_palustris_) and at any rate a slight change in habits is not sufficient

for generic separation." Earlier, Mayr (1942b:169) used _Conopoderas_ as

one of the several genera that is based on "morphologically distinct

geographic forms." The degree of modification that has occurred in these

oceanic reed-warblers, would, if the birds were in a continental area,

undoubtedly be considered worthy of specific or even generic rank by

some authors; however, as Mayr (1942b:162) points out, "the majority of

well-isolated subspecies have all the characters of good species and are

indeed considered to be such by the more conservative systematists."

Owing to their differentiation, the Micronesian and Polynesian

reed-warblers might not be considered by some ornithologists as

belonging to a single superspecies; however, all evidence seems to point

to the origin of this group by a single invasion from Asia."

Tristram (1883:41) was the first worker to recognize the relationship of

the Micronesian and Polynesian reed-warblers to the continental forms,

when he placed them within the genus _Acrocephalus_. Rothschild (1893:2)

further stated, "_Tatare_ cannot be separated generically from

_Acrocephalus_." In discussing the status of the Hawaiian species, _A.

familiaris_, Hartert (1898:58) also follows this treatment. Bryan

(1941:187) also comments on the relationship of the "miller" birds of

Laysan and Nihoa to species at Guam, Christmas and other islands of the

Pacific.

The reed-warblers of Polynesia and Micronesia represent an ancient

invasion from Asia. The continental form, _Acrocephalus arundinaceous_,

is apparently closest to the ancestral stock of these oceanic birds.

This species resembles the oceanic populations in size, general

coloring, shape of bill, and wing and tail structure. Some of the

continental races of this species have a shorter first primary which is

similar to that in the oceanic forms. How rapid the spread was of the

reed warbler through the large insular area that it now occupies is

unknown. _A. syrinx_ of Micronesia has a shortened wing and some

populations have a long bill. Species in Polynesia have stronger wings

than the one in Micronesia, but have become differentiated in other

ways, as, for example, by the long bill of _A. caffra_ and the small

size of _A. aequinoctialis_. In addition, call notes have become varied,

as noted by Chapin (in Mayr, 1942b:54). Also certain of the

reed-warblers have become bush and tree-living birds. The Hawaiian birds

are reduced in size and have become tree-living in a manner similar to

that of other Polynesian species. These modifications of the

reed-warblers of the Oceanic area appear, according to Murphy and

Mathews (1929), to indicate their long-time residence in the islands, as

compared with subspecies of _A. arundinaceous_ that are found in

Melanesia. The latter birds, which are not ancestral to the Polynesian

birds, resemble closely their Asiatic ancestors and have also retained

their swamp-living habits. This would seem to indicate that the birds in

Melanesia may be of more recent occurrence. Stresemann (1939b:324)

presents a map of the distribution of _A. arundinaceous_ in southeastern

Asia and adjacent islands. The original stock came from a point in

China, north of Indochina, spreading to the Philippines and to Celebes,

from where it reached the Solomons and New Guinea via the Lesser Sundas

and Australia.

[Illustration: FIG. 15. Geographic distribution of _Acrocephalus_ in the

Pacific area and routes of its dispersal. (1) _A. arundinaceus_; (2) _A.

luscinia_; (3) ranges of _A. atypha_, _A. caffra_, and _A. vaughani_;

(4) _A. aequinoctialis_; (5) _A. familiaris_.]

The path of invasion of Oceania by the reed-warbler is pictured in

figure 15. Probably the birds became established in Micronesia by an

invasion from the Bonins, where _A. arundinaceus orientalis_ is known to

occur today. From the Marianas and Carolines, the birds spread to

Polynesia; _A. l. rehsei_ of the Gilbert Islands (Nauru) might well be a

connecting link. Possibly, the Hawaiian birds came as a separate

invasion _via_ the Volcano and Bonin islands or through the Micronesia

Chain, or through the Line and Christmas islands from the south. It

seems evident, however, that owing to their geographic proximity and

comparative structural similarity, the species in Hawaii is closest to

_A. luscinia_ of Micronesia. The absence of reed-warblers from the

western Carolines and Palaus seems to reduce the possibility of an

invasion from the Philippine region. However, reed-warblers are absent

from the Marshall and the northern Gilbert islands, where there is

undoubtedly suitable habitat for their occurrence. Possibly these

islands were once occupied by the birds but they were eliminated by

natural causes or by man and his land uses.

=Rhipidura rufifrons uraniae= Oustalet

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