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HARVARD UNIVERSITY

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LIBRARY

OF THE

Museum of Comparative Zoology

AVEWJ WO), WOlVI I ee

LIBRARY

ULI 14 1968

POSTILLA
PEABODY MUSEUM
YALE UNIVERSITY

NUMBER 125. 16 SEPT. 1968

THE RELATIONSHIPS OF THE
“WREN-THRUSH"”, ZELEDONIA
CORONATA RIDGWAY

CHARLES G. SIBLEY

POSTILLA

Published by the Peabody Museum of Natural History, Yale University

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Syntheses and other theoretical papers based on research are also
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Editors: Jeanne E. Remington and Nancy A. Ahlstrom

Postilla is published at frequent but irregular intervals. Manuscripts,
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should be directed to:

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Lists of the publications of the Museum are available from the above
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in exchange for relevant publications of other scientific institutions
anywhere in the world.

THE RELATIONSHIPS OF THE “WREN-THRUSH”,
ZELEDONIA CORONATA RIDGWAY

CHARLES G. SIBLEY htt Ld) pt 1) Oe

LI
Department of Biology and BRARY

Peabody Museum of Natural History

Yale University ULI 14 1968

HARVARD
UNIVERSITy,

ABSTRACT

Comparisons of the electrophoretic pattern of the egg-white
proteins of Zeledonia coronata with the patterns of most groups
of passerine birds indicate that this species is not a thrush, but a
“nine-primaried oscine”, probably allied most closely to, and
possibly a member of, the wood warblers (Parulidae of Wetmore,
1960). Investigation of the taxonomic history of Zeledonia sug-
gests that an early combination of bias, errors, and incomplete
comparative studies conspired to divert the attention of later in-
vestigators from the ample anatomical evidence indicating the true
relationships of Zeledonia.

POSTILLA 125: 12 p. 16 SEPTEMBER 1968.

2 POSTILLA

THE TAXONOMIC HISTORY OF ZELEDONIA: 1888-1905

On November 23, 1888, Senor Anastasio Alfaro, the Director of
the National Museum of Costa Rica, collected a small passerine
bird at an elevation of 2660 meters, high on the slopes of the
Volcan de Poas in Costa Rica. Apparently the bird reminded him
of a small thrush for he wrote the generic name Catharus on the
label. This specimen was sent to Robert Ridgway who selected it
as the type of a new genus and species, Zeledonia coronata (Ridg-
way, 1889), dedicated to his friend, Senor José C. Zeledon.

In the original description Ridgway clearly was influenced by
Sr. Alfaro’s opinion that the bird was a thrush because he began
the description of Zeledonia by noting that it was “somewhat like
Catharus . ” and, later, mentioned that “the loose-webbed
rectrices with finely acuminate points, as well as the loosely-
webbed remiges, slender bill, and long-booted tarsi with sharp
posterior edge remind one of Catharus gracilirostris, to which
genus Mr. Alfaro, the collector, had referred the bird.’”’ However,
Ridgway also noted that “the coloration of the head suggests that
of Basileuterus coronatus. ...”

Ridgway found the Zeledonia had only 10 rectrices and 18
remiges while typical thrushes have 12 and 19 respectively. He
noted, however, that Catharus gracilirostris has only 18 remiges,
apparently not counting the relatively short outer primary in this
species. Ridgway also compared Zeledonia with Scytalopus (Rhino-
cryptidae) and Xenicus and noted that the tarsi of Zeledonia
were “faintly scutellate’, although described as “booted” and
hence thrush-like, earlier in the paper.

Although the collector’s designation of the specimen as a Cath-
arus apparently provided the original suggestion that Zeledonia
was a thrush, Ridgway did not assign Zeledonia to any family in
the original description. Instead he remarked (1889: 537) that
“this remarkable new genus is so peculiar in its characters that I
am in much doubt as to which family it belongs.”

In a footnote to the original description Ridgway (1889:538)
reported that F. A. Lucas had undertaken a study of the skeleton
of Zeledonia and that Lucas had informed him “that so far as his
investigations have gone they show that Zeledonia is not related to

THE RELATIONSHIPS OF ZELEDONIA CORONATA 3

Catharus. ...° This was all that was ever published as a result
of Lucas’ study.

Salvin and Godman (1888-97: 248) compared Zeledonia with
Basileuterus, Xenicus and Scytalopus and concluded that it was
probably not related to any of these genera and that “the position
of this genus must remain in abeyance pending a full examination
of its internal structure.”

The oscinine affinities of Zeledonia were established by Pycraft
(1901) who examined the syrinx and wing muscles. Pycraft noted,
however, that “its exact position had yet to be determined.”

Sharpe (1903: 183), acting upon Pycraft’s discovery that Zele-
donia was oscinine but apparently also influenced by the early bias
toward Catharus, placed Zeledonia between Catharus and Sialia.
In a footnote, however, he stated that “the correct position of this
genus is still unknown.”

In 1905 Pycraft published the results of a more extensive study
of the anatomy of Zeledonia that was undertaken “to decide, if
possible, whether the position assigned to Zeledonia by Dr. Sharpe,
on the evidence of external characters, was at least approximately
correct. ...” Pycraft examined the external morphology of the
bill and legs, the pterylography, the wing and thigh muscles, the
syrinx, and the skeleton. His paper appears to be a reasonably
definitive study until it is realized that the comparisons were made
primarily with various thrushes. A few comparisons with sylviids
were noted and other groups are mentioned but there is no evidence
that extensive comparisons with many passerine groups were
undertaken. Throughout the paper Pycraft lamented the lack of
comparative material and the unsatisfactory condition of the only
available anatomical specimen of Zeledonia. Only skins of most of
the thrush genera were available to him and he noted (p. 3) that
“these have proved to be of no help whatever in the matter.”

A RE-EXAMINATION OF PYCRAFT’S EVIDENCE

Because Pycraft’s paper is the basis for all subsequent opinions
concerning the affinities of Zeledonia it is important to examine it
in some detail. In the following section Pycraft’s principal points
are considered, accompanied by comments on certain comparisons
I have made.

4 POSTILLA

BILL AND LEGS

Pycraft (1905: 4) noted that “the nostrils are covered by a
membranous operculum having the form of a triangle. So far this
operculum appears to be unique.” The shape of the ventral margin
of the nasal operculum in Zeledonia, as noted b Pycraft, is con-
vex with a sharply inflected angle at the point of maximum con-
vexity, thus producing the triangular shape noted by Pycraft. This
contrasts markedly with the concave or straight ventral margin
in many thrushes. I have examined alcoholic specimens of Turdus
migratorius, Sialia sialis, Hylocichla ustulata, Catharus frantzii,
Saxicola rubetra, Erythropygia leucophrys and Erithacus rubecula.
In all except Erithacus the ventral margin of the nasal operculum
is straight or concave. However, in Erithacus there is a noticeable
angle, which produces a slightly triangular shape. The effect is to
provide an operculum that nearly occludes the nostril, as in
Zeledonia.

An examination of nasal opercula in the wood warblers reveals
a parallel situation. The margin is straight or concave in Dendroica
magnolia, Vermivora ruficapilla, Wilsonia pusilla, Setophaga ruti-
cilla, Helmitheros vermivora, Oporornis formosus, Geothlypis ae-
quinoctialis, Seiurus aurocapillus and Icteria virens. In the genus
Basileuterus, however, there is a tendency to develop a triangular
shape. The operculum of B. culicivorus has a decided angle, less
than in Zeledonia but of the same appearance. B. leucoblepharus
also has an obviously triangular operculum while in B. rufifrons
there is but a slight indication of the angle.

It is tempting to suggest that the similarity in dorsal coloration
between Zeledonia and Basileuterus coronatus, plus the similarity
in the nasal opercula of the two genera, indicates that Zeledonia is
closer to Basileuterus than to other paruline genera. However, the
existence of the triangular operculum in Erithacus suggests that
the shape of this structure is probably correlated with feeding on
the ground and perhaps with using the bill to dig and probe. Thus
convergence alone may have produced the similar opercular shapes
and they may indicate nothing about genetic relationships.

The podotheca was described by Pycraft (p. 7-8) and “found
to be formed by the fusion of four separate scutes, traces of which
can be distinctly seen.” Pycraft went on to discuss the uncertain

THE RELATIONSHIPS OF ZELEDONIA CORONATA 5

taxonomic value of tarsal scutellation and came to no firm con-
clusion as to its meaning in Zeledonia.

The margins of the tarsal scutes can be seen in the alcoholic
specimen available to me and Pycraft’s description is verified. The
tarsus is not truly “booted” but there has been a considerable
degree of fusion between the scutes. Presumably this is an adaptive
response to life on the ground in the wet, densely vegetated habitat
of Zeledonia.

PTERYLOGRAPHY

Pycraft described the feather tracts of his specimen and con-
firmed that Zeledonia has nine functional primaries and a vesti-
gial tenth. He noted (p. 6) that “according to the usual ornitho-
logical custom, this wing would be considered to have but 9
primaries.”

In a summary of the “pterylosis of Zeledonia and of the Tur-
didae in general” Pycraft (p. 9) presented the following incon-
clusive remarks:

In its pterylological characters Zeledonia, so far as I have
been able to discover, agrees more nearly with the Turdidae
than with any other group.

But what are, precisely, the pterylological characters of
the Turdidae? Unfortunately, owing to lack of material, I
cannot at present say, nor can I find any scientific contribution
to the subject. So much, however, seems apparent, that the
Turdidae, as a group, present certain common characters,
which may be regarded as distinctly Turdine. It is possible,
however, that these distinctions, which are of a somewhat
subtle description, will break down when the pterylosis of
the Timaliidae (revised), Pycnonotidae, Alaudidae, Motacil-
lidae, Mniotiltidae, and Sylviidae — of Dr. Sharpe’s ‘Hand-

list?’ — come to be studied. These several “families” will,
I believe, prove to be more closely related than has been
supposed.

These statements reveal only that Pycraft was unable to come
to any conclusions at all concerning the significance of pterylosis.

6 POSTILLA

MYOLOGY

‘Nothing of importance appears to be derivable from the study
of the myology of Zeledonia.” Following this remark Pycraft
commented upon the “typically passerine” wing and thigh muscles
and noted various differences and similarities between Zeledonia
and other genera, mostly thrushes.

OSTEOLOGY

Careful study of the skull, sternum, and shoulder-girdle
of Zeledonia leaves little doubt but that this bird must be
regarded as one of the Turdidae. The skull, however, pre-
sents one or two relatively important features, which may,
perhaps, be regarded as primitive characters. .. . The Thrush-
like characters of the skull are to be found in the form of the
tympanic cavity and of the palate. These points of common
resemblance, it must be remarked, by no means leap to the
eyes on a first examination, nevertheless they are real. They
seem to indicate that Zeledonia should be regarded as a primi-
tive Thrush, in the wide sense of the word. ...

Following this statement Pycraft (p. 14-22) described the tympanic
region, palate, and other features of the skeleton of Zeledonia.
Unfortunately, as for other characters, Pycraft mentioned only a
few genera, mainly thrushes, with which he compared Zeledonia.
In spite of his seemingly confident introductory statement that
Zeledonia is a thrush he recorded a series of differences between
the skeleton of Zeledonia and those of the thrushes he used in his
comparisons. In each instance Pycraft designated the condition
in Zeledonia as “primitive” or “specialized” to explain the differ-
ences from the thrushes.

It is difficult to judge the validity of Pycraft’s statements with-
out assembling the same specimens he used and checking each
point. However, it seems clear that he examined material from
only a few groups and that most of the osteological characters of
Zeledonia that he designated as thrush-like are either of wide dis-
tribution in the Passeres or actually differ from the condition in
the thrushes. For example, Pycraft laid particular emphasis upon
the tympanic cavity as being thrush-like in Zeledonia. He began
by noting the condition in Menura and stated (p. 19) that “this

THE RELATIONSHIPS OF ZELEDONIA CORONATA 7

type appears in a large number of widely different groups, and
the fact may be regarded as an additional indication of its primi-
tive character.”

The turdiform birds, Pycraft went on to say, “appear to have
departed from this type along two different lines.” These are rep-
resented by Sialia as the “ground-type” and by Erithacus, Saxicola
and the Sylviidae as the second type. Zeledonia is described as
having the tympanic region of the skull “specialized in one re-
spect” and “indeed the peculiar features of this region of the skull
appear to have resulted from a modification of the type of tympanic
region in Sialia, not as it is in adult life, but as it appears during
its earlier stages of development.” Then, in what must be regarded
as the death blow to his argument that these characters indicate
family level relationships, Pycraft wrote (p. 20): “The skulls of
Anthus and Motacilla, it should be remembered, resemble Sialia
in the form of the tympanic cavity.”

The palatal evidence fares no better. Pycraft described (p. 21-
22) and figured (pl. II) the palatal region of Zeledonia and noted
that “the style-shaped maxillo-palatines of Zeledonia represent an
undoubtedly specialized condition. In the typical Turdiform palate
... these structures are larger, spoon-shaped, and inflated at the
free end to form a kind of pocket. ... The linear form seen in
Zeledonia is obviously a degenerate condition of a maxillo-palatine
of the type found in Erithacus or Sialia, for example.”

Pycraft apparently did not compare the maxillo-palatines of
Zeledonia with those of the nine-primaried oscines. I have com-
pared the palatal regions of the paruline genera Seiurus, Geothly-
pis, Vermivora and Helmitheros, the turdine genera Hylocichla and
Sialia and the illustration of Zeledonia in Pycraft’s paper (pl. II,
7). It seems clear that the “linear form” of the maxillo-palatines
in Zeledonia agrees with the condition in the wood warblers and,
as noted by Pycraft (p. 22), contrasts markedly with the “larger,
spoon-shaped, and inflated” maxillo-palatines of the thrushes.

In his Summary (p. 22-23) Pycraft stated: “as to the precise
position of Zeledonia I regret that I can say nothing definite
until I have had an opportunity of examining much more material
than is procurable at present. ... The specimen submitted to me

. was so much damaged that reliable data on many questions
concerning the soft parts were impossible. ... Nevertheless, it

8 POSTILLA

seems to me that there can be no doubt about the Turdine affinities
of Zeledonia. .. . Its nearest allies seem to be among the Sialiinae.”

EVIDENCE AND OPINIONS AFTER 1905

Pycraft’s uncertain evidence was apparently ignored thereafter but
his conclusion was accepted without further debate. Ridgway,
however, went even further. He placed Zeledonia in the Turdidae
in the manuscript of Part IV of The Birds of North and Middle
America (1907: 69-72), which was written before Pycraft’s 1905
paper appeared; the publication of Pycraft’s results prompted
Ridgway (1907: 885) to prepare an addendum in which he de-
scribed the new family Zeledoniidae. His reason for this move was
based on the combination of “nine obvious primaries . . . and
only ten rectrices! This necessitates the removal of the genus from
the Turdidae, and there being no other group into which it can
be properly fitted, I propose a new family, Zeledoniidae, for its
accommodation. ...”

From this point on, Zeledonia was either placed with the
thrushes (Carriker, 1910; Ripley, 1952, 1964; Blake, 1958; Mayr
and Amadon, 1951; Beecher, 1953) or the family Zeledoniidae
was recognized (Hellmayr, 1934; Eisenmann, 1955; Wetmore,
1960; Slud, 1964).

The only expressed doubts seem to be those of Mayr and Ama-
don (1951: 18) who noted that the ““Turdinae include a number
of aberrant genera that ... may even be wrongly placed with this
subfamily. ... Among these difficult genera [is] ... Zeledonia ...
thought by Sharpe and by Pycraft (1905) to be an aberrant
this

Beecher (1953) discovered additional differences between Zele-
donia and the thrushes and similarities to the wood warblers but
he left Zeledonia in the Turdidae. In his study of the jaw muscles
of the oscines Beecher found (p. 281) that the typical thrushes
have a bifid M. pseudotemporalis superficialis (—M6) with parallel
muscle fibers. Zeledonia, in marked contrast, has a trifid M6 com-
posed of pinnately arranged fibers. Other groups with a “trifid
pinnate M6” include the vireos and the paruline warblers, accord-
ing to Beecher’s descriptions and illustrations (p. 305-307).

THE RELATIONSHIPS OF ZELEDONIA CORONATA 9

THE EGG-WHITE PROTEIN EVIDENCE

The first known nest of Zeledonia coronata was found by Mr.
James H. Hunt in April 1968 during his study of the species in
Costa Rica. Mr. Hunt collected the first egg that was laid and for-
warded the egg white via Mr. William Buskirk to Dr. George H.
Lowery, Jr. at Louisiana State University. Dr. Lowery sent the
specimen to me and a study of the egg-white proteins of Z.
coronata was carried out using starch gel electrophoresis. The tech-
nique is described in Sibley, Corbin and Haavie (in press).

Comparisons have been made with the electrophoretic patterns
of approximately 650 species representing 60 of the 70 passerine
families recognized by Wetmore (1960). These include 46 species
of thrushes, 47 species of emberizine finches, 18 species of tan-
agers, 17 species of wood warblers, 30 species of troupials and
similar numbers of species in other passerine groups. The com-
plete data will be presented elsewhere (Sibley, in press).

On the basis of these extensive comparisons it is clear that the
egg-white proteins of Zeledonia are electrophoretically indis-
tinguishable from those of the “nine-primaried oscines” and that
they differ from those of the thrushes. The “‘nine-primaried oscines”
and Zeledonia have a pattern in which there is a relatively slow
component 18, a fast, usually faint, pre-albumin, and an absence
of the “ovomucoid” fraction. The thrushes have a strong “ovomu-
coid” fraction migrating between the ovalbumin and the ovotrans-
ferrins and a tendency for the ovalbumin and component 18 to
migrate relatively fast. Figure 1 illustrates these similarities and
differences.

It is not possible, from the egg-white data alone, to determine
to which of the several groups of “nine-primaried” oscines Zele-
donia should be assigned. In its morphology, however, it seems
closest to the wood warblers. The allocation of Zeledonia within the
nine-primaried assemblage is part of a larger problem that is
considered in detail elsewhere (Sibley, in press).

10 POSTILLA

ACKNOWLEDGMENTS

I am grateful to Mr. James H. Hunt, to Mr. William Buskirk, and
to Dr. George H. Lowery, Jr., for providing the specimen of egg
white of Zeledonia coronata. It was due to Dr. Lowery’s interest
and encouragement that the study of Zeledonia was undertaken.

Dr. Kendall W. Corbin, Mr. Jon E. Ahlquist and Dr. N. Philip
Ashmole have provided advice and assistance during the study of
Zeledonia and in the preparation of the manuscript. Dr. Richard
L. Zusi permitted me to examine the type and other specimens of
Zeledonia coronata in the collection of the U.S. National Museum.
Lois Robertson assisted with the laboratory work and Georgette
Lewis typed the manuscript.

FIGURE 1. Starch gel electrophoretic patterns of the egg-white proteins
of a wren (Troglodytes), several thrushes (Hylocichla, Phoenicurus,
Turdus), the wren-warbler (Zeledonia), and _ several ‘“nine-primaried
oscines” (Dendroica, Thraupis, Habia, Carpodacus, Richmondena, and
Agelaius).

The application slots (A) are indicated by the white rectangles at the
left ends of the patterns. Component 18 (C-18) is the first band to the
right (anodal) of the application slot. The next anodal bands, usually two
or three, are the ovoconalbumins or ovotransferrins (Tr). To the right
of the conalbumins in the thrushes are two heavily staining areas, in the
nine-primaried oscines only one. The first of these bands in the thrushes
is possibly an “ovomucoid” (Om), the second one (most anodal) is
probably the ovalbumin (Ov). In Zeledonia and the nine-primaried oscines
the heavily stained area is thought to be the ovalbumin, and the ovomucoid
is assumed to be absent. However, in these there is a faint pre-albumin (Pre)
which is absent in the thrushes.

THE RELATIONSHIPS OF ZELEDONIA CORONATA 1]

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Troglodytes aedon

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Hylocichla mustelina

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Turdus libonyanus
Turdus falklandii
&
Zeledonia coronata
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AaG=15: Tr. Ov Pre

12 POSTILLA

LITERATURE CITED

Beecher, W. J., 1953. A phylogeny of the oscines. Auk 70: 270-333.

Blake, E. R., 1958. Birds of Volcan de Chiriqui, Panama. Fieldiana,
Zoology 36: 499-577.

Carriker, M. A., Jr., 1910. An annotated list of the birds of Costa Rica
including Cocos Island. Ann. Carnegie Mus. 6: 314-915.

Eisenmann, E., 1955. The species of Middle American birds. Trans. Linn.
Soc. NEY #2 12128:

Hellmayr, C. E., 1934. Catalogue of birds of the Americas and adjacent
islands. Part VII. Zool. Series, Field Mus. 13, publ. 330. 531 p.
Mayr, E. and D. Amadon, 1951. A classification of recent birds. Amer.

Mus. Novitates 1496: 42 p.

Pycraft, W. P., 1901. [Remarks upon a specimen... of Zeledonia.| Brit.
Orn. Club, Bull. 11: 12-13.

—— 1905. On the systematic position of Zeledonia coronata, with
some observations on the position of the Turdidae. Ibis 47: 1-24.

Ridgway, R., 1889. Notes on Costa Rican birds, with descriptions of seven
new species and subspecies and one new genus. Proc. U.S. Nat. Mus.
11: 537-546.

1907. The birds of North and Middle America. U.S. Nat. Mus.
Bull. 50, Part IV. 973 p.

Ripley, S. D., 1952. The thrushes. Postilla 13. 48 p.

1964. Subfamily Turdinae. Jn E. Mayr and R. A. Paynter, Jr.
[ed.] Check-list of birds of the world. Vol. X. Mus. Comp. Zool.,
Cambridge, Mass. 502 p.

Salvin, O. and F. D. Godman, 1888-97. Biologia Centrali-Americana. Aves.
Vol. Il. Taylor and Francis, London. 598 p.

Sharpe, R. B., 1903. A hand-list of the genera and species of birds. Vol. IV.
Trustees, British Mus., London.

Sibley, C. G., (in press). A comparative study of the egg-white proteins of
passerine birds. Peabody Mus. Nat. Hist., Bull.

Sibley, C. G., K. W. Corbin and J. H. Haavie (in press). The relationships
of the flamingos as indicated by the egg-white proteins and_he-
moglobins. Condor.

Slud, P., 1964. The birds of Costa Rica. Amer. Mus. Nat. Hist., Bull. 128.
430 p.

Wetmore, A., 1960. A classification for the birds of the world. Smithson.
Misc. Coll. 139 (11): 1-37.

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Any issue of Postilla will be copyrighted by Peabody Museum of

Natural History only if its author specifically reque

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