THE COMP‘ARATWE MYOLGGY OF THE SYRINX
EN THE AVIAN FAMtLY TIMALIEDAE

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This panel" resorts on ‘."‘.€? swim": in three su‘ifa~'_i".-i..3s vithin the.
avian fiwily Tinaliiiie, The tirpes~ Oi ffiis stuiv was to se‘ernine
whether n:r not fhe syrinx is chzraeterisinIlly uwi.Jrfi throat sit
the family ani whither or not it is a useidl too in detaraninj the
$113110 :‘JI’I'EtlT relationship 01‘ bir-is. me specimens used in this sfufif
were collecfed in [lam iistrict, E.

The syrinx is loca‘ed at 0? near the Vronahial biTurcaFiin and
is uSIally associated with one or more pairs of svringeal miscles.

“he historv of svrinreil invas‘isations wws traced ‘roi r. .iCj
3' Asar i1 l”79 up to the Uresent century.

SYrinqeal nusnulifiure terminolo"v Hes leen coniisinv. A chart
showind muscle syhsqymy Dreesref “v ea“1ier workers is revised ind

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n this UlUPr.

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iei
Tue 3 Tim? in fihe Tifiqliidie is quite regular. Eijht intrinsic
muscles are present in the Tirloiiinae and in fihe Pellorneiiae
whereis seven is the 13131 nuwber in the Pomatorhininie. The
Turdoidinae and the Cinclwsowatinae have equally well developed

syrinqes. Lne ?ellorneisae syrinx is intermediate betWeen *~e

Pomitorhininae an? the T rdoiiinae syrin*es.

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MABTE” OF SCIEIPV

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Department of 7oolozy

1962

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VISTOYITiL TEVIEJ .............
P??;£NTamIO” CT DiTA
Naterials and Nethods ..
fiuscle Terminoloqy .....
Xuscle Synonymy Chart ..

Description of Jyrinveal

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ACKNO‘IIL 'E'JGEWCI‘ITS

I wish to express my thanks to Professor l. J. Wallace for
his guidance and help during the preparation of this paper. In
aidition I would like to express my appreciation to the other nemhers
of my committee, Dr. R. H. Baker and Dr. J. L. Locmwood, for their
helpful comments and suggestions.

I am expecially zrateful to Dr. Robert L. Flening of Kathmandu,
Nepal, for collectinq and sending me the specimens on which this

study is based.

INTKODJTTIOH

This paper presvnts a constrative study of the srrindeal 'usculature
from selected samples of Iimaliidae - a family of Asian and African
bahblers and lauqhinv thrushes. The purpose of this study was to
determine the variahility of the syrinx withiv one family as this has
a direct bearing on whether or not the syrinx may be used as an aid to
determine phylopcnetic relationships. I chose the Tinaliidae because it
is a large, well develop d and extremely diversified ”anily. Previously,
Forbes (1882) and “ayr (1931) had examined the subfamily Cinclosomatinae;
howeveg,the syrinx in the remainder of the fanily has not been studied.
The external morphology and especially the hahits (which include the
call iotes and various songs)_of members in the family are sometimes
surprisinsly different. Therefore it is important to see wdat variations,
if any, occur in supposedly conservative internal taxonomic characters
such as the syrinx.

are

The Tinaliidae is composed of several distinct arwups of birds.
Taxonomists are not in complete avreement as to the relationship of
these groups. Delacour and Vaurie (1957) are of the opinion that the
IPimaliidae should be divided into five subfamilies, one restricted to

Dhew Guinea and Australia, the others found either in the Oriental or
Ailrican rezions, or both. This investigation does not attempt to cover
tile entire familv, which includes some 280 species, hut reports on the
Erfirinx in three of the subfamilies. Eventually, it is hoped, a
COPHIilation of several smaller reports may lead to a better understanding
<13 the syrinx in the Family as a whole. This when combined with other

charwacters should shed liqht on the relationships within the Iinaliidae.

The syrinx, also called the lower larynx (Jnterkopf), is
peculiar to the Class Aves and it is the muscular control center
for the vocal sounds produced by birds. The syrinx is located, with
variations, at or hear the reqion where the two bronchial tubes
bifurcate from the trachea. There are three major variations. The
tracheo-bronchial syrinx is most commonly found in passerine birds
and incorporates both the caudal part of the trachea and the cephalic
portions 0” the two bronchial tubes. The bronchial syrinx, on the
other hand, involves only areas of the bronchial tubes and is found
in come cuckoos, Cuculidae (Berger, 1960) and nightjars, Caprimulgidae
(Garrod, 1873). The third type, the tracheal syrinx, forms just
cephalad to the bronchial bifurcation and is characteristic of some
New'World passerine families such as the woodcreevers (Dendrocolaptidae).
It should also be noted that not all birds posses a functional syrinx.
The New World vultures produce a erupting sound without the benefit of
syrinqeal muscular modifications (Wishinen,195”). Besides the svrinx
other modifications of t“e respiratory tract serve, in a few birds,
to amplify various sounds produced. The male chachalaca (Ortalis)
for example, has a coiled trachea extending as a loop ventral to the
‘ieart and this allows for its famous noise-making ability. It is also
knuown that the larynx serves to modify the sound in some birds
((30le, 19LS).
The syrinx itself is a modified area in the respiratorv tract
31nd usually is associated with one or more pairs of intrinsic and

eflctrinsic muscles. Muscles that connect the syrinx with other parts

of the body are called extrinsic while intrinsic huscles originate
and insert on the syrinx itself. Ten pairs of intrinsic svringeal
muscles have Ween described.

In the Passerifornes the characteristic syrinx consists partly
of a modification of the cazlal partion of the tracheal tube which
is often called the tracheal drum or the Trommel. For the most

"0

part the drum is fashioned tron the fision o? the three host posterior

U)

tracheal rings (Ruppell, 1933). tr,semann {1927-193h) indicated
that in some cases at least six rings may be fused. In most birds
it is the last three.

Also involved in the syrinx are the anterior bronchial semirinzs,
which unlike the tracheal rings, are not completely fused on the
median or interior surfaces. The first three semirints (ecunting
caudai from the trachea) are the most often modified and involved
with the svrinx. ‘t is on these rinis that most of the syringeal
muscles inSert.

Internally, cartilitinous components are important. The
pessulus, located at the bronchial bifurcation, is an anterior
projection which arises fron a f sion of the most posterior part of

the tracheal rings. Another prominent feature, in some syringes,

is the anterior cartiliuo te sor which usually articulates ventrally

 

with the first bronchial semiring and projects dorsally and medially.
There are four major membranes associated with the syrinx.

The semilunar or membrane semilunaris is attached to the "es ulus

 

and proj,cts anteriorly. It does not play an important part in the

-14-

sound prodlcti‘n (Hiskimin, 1951). The external and internal
tympaniform memhranes are the chief soxnf—vroducina oraans. The
internal menbrane, located medially and projecting oosteriorly
from the stsulus , is attached to the ends 0" the bronchial senirinzs.
The external tympaniforn membrane is attached laterally and extends
between the bronchial semirinqs. Sound is produced by the vibrations
of these membranes when air is forced out ”rom the lanes and air
sacs. The tension of the membranes can be altered by the movement
of the bronchial senirings which are, in turn, moved by the syringeal
muscles. The fourth membrane of the syringeal region is the bronch-
iodesmus which connects the two bronchial tubes with a thin sheet of
connective tissue.

The syrinx is supnlied by the long branch of the hypoglossal
nerve (Newton, 1899) and also by branches of the vagus ( Stresemann,
1927-193h). The innervation of the syrinx has likewise been stunied

by auppeii (19a3).
HISTORICAL REVIEW

The historical aspects of syrinqeal musculature have been well
sunmarized by Wunderlich (1886) and Kbditz (1925). It appears that
the first important recorded syringeal investication was made by F.
Vicq D'Azar in 1779. He gave a zeneral description of the vocal
organs in birds and came to the interesting conclusion that in son'
birds the syrinx is of simplest proportions. Later Cuvier (1802)

undertook an investigation of the EurOpean starlinz and came to a

-L’.
/

quite different conclusion. as maintained that the vocal apparatus
was, indeed, very complicated and that there are five muscles on
each side of the lower larynx.

Perhaps the ?irst complete and correct description of the svrinx
shomld be attributed to Savart (132 ). He noted a maximum of six
pairs of muscles and described their differences. He also noticed

an internal membrane and called it the nembrana semilunaris. Seven

 

years then passed before Yarrell (1833) completed a review of the
skeleton and misculature, includin* the svrinx, of the raven.

The most important fizure, perhaps, in the early nineteenth
century literature was 3 Berlin worker, Johannes Huller, who in
19h; published his survey of 125 genera and subgenera ( reclassification
since his time has changed the statis of some genera, elevating them
to the family level). Of special interest for this paper is that
he had access to an example in the genus Timalia fron Tenassarim
in northern Rurma. Muller noted that this Timalia had a characteristic
songbird syrinx. It is unfortunate that extensive as his work was,
he did not record his‘observations in detail and so a great part
of the value of his work is lost.

During the nineteenth century Herre (1859) described the lower
larynx of the Oscines and of the "Aves passerinae anomalae." Seven
years later Owen (1886) reported findin§:on the syrinx of tie
raven.

an English
Garrod,in 1878,9ublished untranslation of Muller's 18h5 work and

appended information pertaining to Meiura in which he described

-6-

three syrinqeal iuscles. Earlier (1876) he hai shed important limht
on the r.lationshins of Pitt .In this paper he also preposed the
term acromvodi Which referred to muscles of the svrinx inserting on
the extremities of the bronchial semirinzs. This is the most comnon
condition in the Oscines. Acromyodi along with mesmyodi (with the
insertion on the medial portion of t.e bronchial senirings) are
apprOpriate terms for use today.

Towards the end of the century Wunderlich (1886) gave a
description of syrinaeal historv up to his tine. Furbringer (1386)
also contributed to svringeal literature. doth authors found a
maximum of seven pairs of syringeal muscles. During this period
Gadowwgs actively compiling aspects of birds in dictionary and reference
form. He collaborated with Newton (1896) for a volume in English
after previously working with Selenka (1891) to do one in German.

Forbes (1880, a & h) discussed the postion of Philepitta and

 

the Eurylaimidae. In his 1882 paper he investigated the timaliid
genus firthgnyz.

At the turn of the century Hacker (190i) dealt with the variations
in the anatomy of the svrinx. Until this time the Terman and Englisi
workers were the prime syrinzeal investigators. A change occurred,
however, when Setterwall (1898) studied the nature of the syrinx in
many of his native Swedish birds. fie noted a maximim of nine pairs
of syringeal muscles, mentioning that on the ventral end of the third
semirinq he felt that two muscles attached - a lonier and a shorter

one. These two muscles were equivalent to Furhringer's (1889) long

‘H

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musculo tracheo brenchialis ohliquus.

It shoxld also be nentioned that durinf this tine 3eddard was
actively investigating the syrinx in cucloosg Cuculidae (1885, 1902)
and ofiier birds (1393) while rlso conniling a reference wirk (1°98).

There has not heen a qreat deal of work on svrineeal musculature
during the twentieth centurv althou~h several innortant papers have
appeared.

K6ditz(1925), worann under Prof. ”acker, presented a brief
and accurate acco;nt of five Old World passerine families, the

l
9

Pvcnonotidae, 7osteroridae, Mel nhagidae, teeteriniidae, and Dicaeidie.

We observed that Pvcnonotis his seven pairs of intrinsic svrinzeal

 

muscles - three lon: musculi larvnzosvrindei and four short musculi

 

’

svringei. One of his most imnortant contributions was a chart of
syrinqeal nuscles showinz the synonymy between the names used by eight
previous workers. He made a modiTied chart after an earlier work by
Setterwa1l (1901).

Mayr (1931) published an account of the Cinclosomatinae - an
interesting subfamily of the ”inaliidae found in the New Guinea

and the Australian region. The Cinclosomatinae include Helamnitta,

Orthon , and vanorhina (see page 37 - hOT. Resides describing

 

the syrinx of these birds he noticed a short dorso-lateral muscle

lying next to but ventral to the E. svr’ngeus dorsalis. This he called

 

the M. interbronchialis_laterali§. Incliding this muscle with the

 

nine others described by Setterwall (1901) there appear to be ten

“.

pairs of intrinsic syrinneal muscles (excluding the extrinsic a,

.43-

sternotrachealis and the M. laryniotrachealis) in the Cinclosomitinae.

 

 

Straseminn (192”-193h),in his monumental work on birds, devoted
several pares to descrihinz the svrinx.

'3 O . w \ +‘ O ' ‘T011

New Americans have done any morx on one syrinx, but miller

3% 35 V7

(1927, 1931, 193Q) published several interesttnj papers on owls.
He showed that there is a sinrle intrinsic muscle in owls and that
there is a slight sexual size dimorphism in the owls with which he
worked.

The question of sounc proiuetion is considered purther

hy Tole (lQLS) who published a general niner describing *he

I;

N

0 CeJ I
trachva and svrinx as a while, an! explaining in what ways thry

function. He attempted +o show some of the variations and

modipications of the svrinx and the trachea aid discussed the somewhat

linusual svrinx of a parrot.

(3

Miskinen (1951), usini ”resh material, ~xperimenta11y demonstrated
tliat the tympaniiorm membranes and not the semilunar membrane were
tdnea chief sound-producinr errans. She also dissect d some thirty
Siv201os of North lmerican birds, locating as a maximum numher five
P‘tirs of intrinsic muscles.

Supplementing investigations on wild birds are several studies

of f}allus domesticus (fiyers, 1917; Appel, 19?9)- The mOSt COWPTG'

D

‘enfifiive of these (Myers, 1917) included a discussion of the micro-
SCOFXiC anatomy and the embryology of the syrinx. Myers not d that
+":" ' ‘V‘ . ' . ' ' n a .

~aer¥3 ale no intrinSic muscles in the fowl syrinx, there 15 a Size

(1.?“ r‘ o 1 ’
al-1erxaite between the sexes in 'nllus, and that muscular tissue

J.
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. f 1 a, .
n toe 1'7-o—rm 1r e**:'.'r;.'o.

idlo

arentiafes
Apnel (17??) noticinz “Vers' run r un’vrtook to determine w:at

L

sexual diflmnurvisn mi'ht occur in the rzxmnx lcéhorn nreei of .he
domestic Chlfih°n. {e concluded th-' in this Free? tiere 15 no

dif”erence but noted that this mav not be the case in other hreeis and

in the wild Gallus gallus (nankiva of Ansel). He also presents a

 

review of the previous literature relatira to Callus svrinveal stu ies.
. . . bfi -, . ,
The syrinx in ducks has been studied humrhrey'(1955). He noted tnat
I“

the Anatidae svrinx possesses only a sinnle pair of extrinsic mus*1es and

that +he tracheal bulla is often enlaraed, esoeciallv on the left side.

\/

Thi host recent paper dealing with the svrinx is Rerser's (1968
review of the cuculine svrinx.

”RVSTNTATTUH 91 UiTi

Materials and Methods

 

The specimens used in this studv were collected with a 12—:uafe
shotqun (.blO aux. barrels, #12 shot) on an exnedition to 11am dist ict,
if. Nepal. Collecting altitudes ranged from ?,000 to 10,300 ffEt.

\fter the birds hid been skinned the thoracio and neck regions were
jxnmediatelv injected with and placed in five percent fornalin solution.
Fkor shipment to this country by air they were carefully wrapped in
fWDrmalin-scaked cloth and then pliced in plastic bass. Once in t'is

ccnuntry the specimens were kept in bottles contuininfi five percent formalin.

When dissections had been maie a variable rower binocular
micxrosc0pe was used as an aid in identifyine the tuscles. T‘e wost
usEdflll miqnifications were “onid to be between twentv ani fiftv

Dower. Drawinrgs from V"I‘Vln\"f .an‘rles were ma‘ie.

Forty snecimens representin' l’7 sbecies, 9 genera and 3
subfamilies were examined. A list of these follews. The names
Eire taken from Ripley (1961).

List of Species Dissected

 

Pellorneum rificens Swains n. Shetted quhler
———---—-. —. _J——. ‘

 

Pomstorhinus ruficollis FOJ”SOH. T‘ufnusnec‘md Joinitar Babbler

Pomatorbinus ervthroienvs "izers. Rustycheeked Scimitar fiabbler

A

Xiphirhyn nus surrrcilieris dlyth. Slenderbilled Scimitar Babbler

 

Hicroura albiventer (Wed sen). Slatvbelliei Hren-Bxhbler

 

Garrulax striatls Visors. Striated Leughing Thrush

 

Garrulax lineatus (Visors). Streakud Laughinz ”hrush

iarrulax ocellatus (Vigors). Spotted Laughing Thrish

 

Garrulax affinis Blvth. Blackfaced Lauénin* Thrush

 

Tarrulax erythrocephalus (Viwors). Yedheaded Laughing Thrush

Garrulax subunicolor (leth). Plaincelored “auqhing Thrush

 

larrulax leucolonhus (Hardwicke). White reisted Leaching Thrush

 

ilcipne castaneceps (Woieson). Thvstnutheaded Tit-Babhlyr

 

Alcigpe nipalensis (Vodrsen). Stripethrwqted Siva
Actinoiura nipalensis (led son). Henry Rarwing

HeterOphasia canistrata (Visors). Ylackcapped Sihia

Mu80112Ternhrflegv

 

'The terminology used in describing syrinjeal musculature is

 

 

 

 

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sciewhat~cnnfisinq. Several different canes have b2an five: to the
sane muscle and likewise the same 131e, at tines, has be,n need for
more thin one muscle. Also there has been 5016 disajreement as to
what constitutes an entire nuscle. This is evifent when we note
that the maximum number f intrinsic nuscles in sontbirds has been
variouslv civen as anywhere from five to ten pairs. Re*ent trends
tend toward the hifher number of muscles,except for Hiskimen (1351)
ho recorded f‘ve rairs. Yfldits (19?S), following Setterwall (1901),
reports4 nine and Vavr (1031) incluied ten in the Cinclosomatinae.

From my investinatiwns * have come to the conclusion that there nr.

six very distinct svriuqeal miscles. a seventh, W. laryngosyringeus

 

ventrilateralis, is quite distinct in west of my snacimens, while an

eizzth, H. laryngpsyrineeus dorsolateralis, can be distinguished in

__,2

 

:3 few speci~s. The ninth, T. syringeus ventriliteralis internus,

eand the tenth, H. inter-bronchialis latcralis are deep miscles

 

vehich I could not senarate from the overlying muscles in my specimens.

r.

#4

LlSCle fibers certainly ewist in the region described for these two
Wnlscles but in my spacinens they were inseparable as distinct nuscle..
In svrinfeal terninolO“y a well develoyed, distinct muscle

113 «considered "strong." If a muscle consists of only a few fibers
it ‘is considered ”very weak." Somewhere between and bordering on
919 "strong" is the "normal" situation.
To help clarify the confusing syrinqeal muscle terminclory
Setimarwwll (1901) prepared a chart showinz muscle synonymv. from

thlfii Kbditz (1925) prepared a revised chart which in turn I have

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modified and nresented below with the additions needed to bring it

up to date.

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Muscle Synonymy Thart - (showing intrinsic muscles)
ey t the Authors:

Cuvier - a; Savart - b: Herre - 0; Owen - d;
Wunderlich - c; Furorinrer (also “ficker) — f;
Setterwall (also Kofiitz and Nayr) - 3; Solo - h;
Hiskinen - 1.

Muscle Synongns Author

‘ I

1. Le releveur lonqitldinsl fies ueniunnesux 3
Le releveur antérieur flu deuxieme b
Levator anterior aT“JS secundi c
Qroncho-trachealis anticus d
Levator lonaus anterior arcus seoundi e
Tracher-hronchialis ventralis T
Larynnosvrinseus ventralis sive ventralis lonqus g
Dorsalis ventralis h
3roncho-trachealis snticus i

2. Le releveur lonn‘tddinsl Hes denianneux o.p. a
Le releveur anterieur in troisiene osselet p.p. b

- c

- d

lotqtor arcus tertii n.p. e
“racheobronchiilis ohliquus D.p. T
Larvnqos*rin:eus ventrileteralis lonrus w
- i

3. Le petit releveur ? 3
Le releveur posterieur nu prenier osselet b
Levator posterior rrimi arcus c
Broncho-trachealis brevis d
Tensor no branae tympanifornis internee e

Tracheo-bronchialis dorsalis brevis
Laryngosyringeus icrsalis sive dorsalis lonzus g

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Le relevenr rostertet

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Levator nosterior or

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3roncno-trfi.“hei‘is W
Le"1+or 10W 13 r“ *‘
Tracheo-oroncziqlis
Laryn"os*rintei5 ”“T
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Uorsalis orovis
Wronono-trnoneilis

Le releveur transver
Le reTeveur "if-3:3 deu‘r
Levator anterior arc
Tronchiul's anticus

Lavator brevis inter.

Syringeus ventralis
jvrinreus ventralis
3 ri_njeus ventra‘is
'3r oneliaiis notions

Le rele Jrur anteriF u
Leva tor anterior ere

Rotator arous tertii
Tracheo-bronchii‘is
Syringeus ventril

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soléterq‘is sive fiorso-
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sol

iEne et nrenier osselets
is sectuiiiuet firim
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give ventralis brevis

r iu troisiene osselet

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p.p.
obliquus p.n.
erm‘ is ever-nts sive ventri-

“alis orevis internus

Svrinqeis ventroii+er17is

Le relevnur antérieu
Levator anterior or

r du troisiene osselet p.p.

C18 tartii n.n.

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Laxator nenbrana, tjrmgiaaniornis e'rternae e

Syrin~eus ventriliterqlis‘ f

Syringeus v:ntri-11tera1is internus sive ventri- g
‘lztsraiis brewis internus

- h

- i

8. -

OO‘Q’

.. r3

Levator brevis rosterior arcus seeindi p.p.
Syrinzeis iorsalis D.n.

Syrinqeus norsalis sive dorsnlis brevis
Surin'eus dorsalis

H- 13‘th H: (D

9. Le releveur oblique 3
Le rotafeur rosterieir du dalxieme osselet ‘
Rotator oosterior secuudi arcis

Eronehinlis ‘OStiCLS

Levator brevis "osterior ircus seoiudi p.p.

Svringeus dorsalis

Syriuzeis dorso-literelis sive Horse-literilis

brevis
Jorsalis obliquus ii
Rronoriaiis nostieus i

C)

('1) (D Q,

Li

lCL 'Enter-bronohiwlis literalis ’ anr

The notation " n.?.“ in this CW7FL re“ers to additions

/‘ . J n
‘ufiffiitz(l995) made to JGLthW311'S (1931) work.

.~eSCr'}_7+.°on f' the Syringeal 3"useulature

 

The nomenclature and distributional ietw ls used in tois paper

n:

were tiken, Tor the nest "art, from 'inlev (1961).
The hronehiil senirinfs a"? nunberei from the most Cephqlsi
(as number one) to tNe vore cai‘il.

In the followin: lescription, ”i” reyreseris insertion, "o"

7-. a J- ‘ A . an . - . 1 'l " ‘ 'II' 3 v ’V‘ ' ‘
re,resenus :rr in 111 C ‘eters to contents.

This is a suhfanily of sone thirty species of smallish +"orest
birds. Thev are nost often foino in hilly re'ions at low iltitudes,

but have been found up to 7,000 feet in the himalavas.

Genus FEILOWNECW Swainson

 

Pellornaum ru5icegs Swainson

Distribitel +hrhzjh Innis, Yunnan, in} the Inloehinesa suorevion.
Phallorneun r1?ioe"s nqniollii qun?or4. Spotted lab lzr (91. I, 1138.
A, 7 31d C)
M. laryngosyrinteis ventralis
o. from the extr'nsio ”. lurfntotraehealis
i. on the ventral one third 0? the second semirinr,
coverin' the ventral end of the first seuirinj
c. sli “tly seeker than n~rmal

fl. leggnjosvrinjeus ventrilateralis

This muscle can be distinguishil from the rreoefiinq
in this sonoi 3 1nd lies life?“ll” to it.

-16-

T. lgryngosyrinreus Jorsalis

o. from the M. lsrynneotrqoh-slis

i. On t‘e iorsal tin o“ the first senirinv
C. normal

W. lirynlosgrin'e_§ w;rsolsteralis

lnsmoorahle frdm W. l. dorsslis

L, svringeus norSWlis

0. on the caidal mic?in of the third t"aoheal rinq
shove the tracheal drum

i. on the iorsal causal nirain o? the ’run

0. this muscle is GSD€CllllV stronr in this srecies

N. syrinqeus ventrilis
h. on the cethlic ventral margin of the ‘rum

1. on the TLULMTL ends of the first ”n4 second,
nostlv on soc>nd scmirinv.

.3: szringeus ventrilaterilis

 

 

0. On the lateral meflisn nariin of the lrum
i. on the ventril third of the thiri semiring
C. % cooperatively weak mujcle

 

H. svringeus dorsolateralig-

 

0. on the lateral margin of the drum
i. on the tip of the first semiring, slightly

ventral to w. 1. dorselis
c. a weak muscle in this species

”. sternotracheulis

 

i. on the literql mefiifln redion of the orum, near
the cenhnlic edge
0. Somewhat weak

The whole svrinx op this sneciss has a strikingly ”lit look

fin
‘he 'nuscles are n“t as hoovy as in the c‘hnr sp”cies oissected.

-l7-
Suhtanilv Ponatorhirinae
This suhfanilv comerises 33 species of tires which inhihit seni-
nrwn and scruh jungle oreis. T.ey are oriharilv t rrestrial or see
found in low bushes.
erlue F(lITVCR?Iiޣ3 H'nnsfielo
Ponstorhinus ruficollis Venison
The oistri“wtion of this sneci.s is from Lu non east through
fissan to northeri Turna, Yunwnn, ans northern Iniochina, Weinon, and
Formosa.
P natorhinus ruficollis vodwiui Kinnefir. Vufiusnedkrd lciniter

labhler (ll. I, fies. ; " ')

' r v’ . W 4 1n": ,'.;w~,+ “W '
_E,lerNIZSJrfifiluus *7.cr1_1s
o. “rom the lsrvnvotrschenlis
i. on the ventral fourth o? the seconi S ‘
co"erir1 the and of the third senirinq and
also coverin; the “. syrinqous iorsqlis
us ven‘rilsternlis

_. lorvntosrr'ngs

~ 4‘-

Kg)

It is riifpicult to senirate this muscle from the
preceding one; toqether thev form a strons

muscle.

M. larvniosyringels dorsslis

 

o. fron the lsrynrotrechealis
i. half on the dorsal tin of the first semirinq and

the other half on the dorsal caudal marqin o? the irln
c. mhis is a stony musclf in hhis soecies

W. Larynqosyringexs florsoleteralis

 

This nuscle is inseuarshle From the h. l. donsnlis
i. syringeus dorsslis

This muscle annears to be ntsent in this soecies

 

[.1;

-19-

~-

a. srrinreus ventreli

J __,
——

0. on the cephalic ventral rih of the tracheal drum

i. on the ventral thirfl o? the Second senirinq with
a iew ?ibe s to the first semirinq

c. although entirelv covered this is still a Strong
muscle in this so cies ‘

'. svrinvegs ventrilateralis

0. on the literql medial area of the drnr, cephaled
to h. s. dorsolwtsrslis

i. ow the ventral one-third of the third senirinz

0. strong

M. svrineeus florsolaterslis

 

0. From the lateral median area of the drum below
1. sternetrachealis
i. on the *orsal tin of of the first seniring; ventral
to} . l. derselis
H. sternotrachealis

 

i. on the literal edge of the drum just caudal to the
ceohalic rim
0. slithtly weak

Ponetorhinug ervthroqenvs Viqors

 

This species is distributed from the northwest Hi alayas, east
‘to western China and south to 7urma and northwest Thailand.

P%flnatorhinus ervthrogenvs ferrugilutas hodqson. Wustycheeked

 

Scimitar 3ab'ler (P1. II, firs. 4,1, 3 c)

H; larynvosvrihgefls ventreli§

 

o. from the H. liryfiqotracheal s

1. on the veetral end of the seconi Simiring with
some fib rs to t‘e first semirinq

c. stronq - completely overlaos the I. s. ventralis

i2 larghgosyringeus ventrileteralis

This is a weak wuscle and is fused with V. l. v ntralis

-1};-

. lirVfi‘OSVTln’th icrs.lis

 

o.from the T. larvnqotracheilis
i. on the iorssl ceuiel rim of the ”rum
c. serial

0 _ ‘ ‘ .
lwrfict'OSVdnri'gzs nirsolrfiz-ri‘is

 

 

_ L.’ ‘
fiuscle fihers occurinv in this rezion may he seoirehlc
tron ”, 1. dorsslis as tic? rin Tron the rim 0‘ +he
drum cenhqlefl elonr the lnt rel edze of the trachea
Y. svriifieus dorswlis

 

o. Seems to oe assent iL this speCies

W. svrinceus ventralis

 

o. fron the cephalic rim of the firum on tag ventral sise

i. on the ventral erds of the second awd first son riscs,
mostly on t e secOid

c. a weak musclein.this sh7ciss

N. sgrinveus v ntrilwtarzlis

 

o. from the 11*eral marqii o“ the drum below (caused to)
i. sternotracheilis

i. on the ventral t ird of the third semirinz

C. HOTTfll

a. svrinqeis dorsolaterelis

....--” ~— .—

 

 

0. Trom about the midflle one third of the lateral margin
on the drum

i. on the dorsal emi of the first semirine

c. This is a stran muscle in this species

h. s,,rootreche3lis

 

i. on the lateral edge 09 the drum just caudal to the
cephalic rim

Genus (IPhT937377U3 ‘lvth

Xiphfi—rhs'uchus supercil iiiris ‘lfrth

*3

1”is Spec » is distributed from the Himilsyas throufh eastern

U)

‘7
:9 T? r -_ _‘ fl . '
p 1- arr} assim to Burma and the Ihdochinese SdhréflOn,

-20..

‘Xinhirhvnchns sunerciliqris su erciliaris llvth. Slehdertilled

 

fl
.

4 l T
Lats. L g .4, ' L ll

Scinitnr “ahbler (Fl. II,

M. lirvnrosvringeis ventrslis

 

0. Fr‘n the H. larvnxotrachealis
i. on the ventral fourth of tie second semiriic
c. This seens to have a vernal develohmeut with
a li‘tle of the H. s. ventralis “is Eflx? from abcvs.

‘. larvnsosyringcus ventrilnteralis

—._....__ _

 

 

”his nuscle 18 difficult to seearate in tiis snecias

3: lazynflqsvrjnVeus dorsslis

 

o. pmm the M. larvnqntrachealis

i. on the dorsal caudal end of tie drum; no fibers
evtendine to the First semirinq

c. norhal to strong

n. larvngpsvrinfeas dorsnliteralis

 

 

This muscle is inseparable from ”. l. iorsqlis
fl, svrinceus dorsalis
This muscle seems to be absent

4 svrinceus ventrilis
0. from the caudal margin 0” the first tracheal riiq
aid the cephalic nnrrin (ventral) of the drun
1. on the ventral efid of the second semirin? with scie
fibers inserted on the first semirinq
c. "his is a relativel_ strcnz muscle
:1

:3 syringeus ventrilat.rali§

 

‘9

o. frcm he lateral narzid of the drum caudal to A.
_ sternotrachealis

i. on the ventral half of the third seniring

c. this muscle is strcnz and rounded

fl: svrinveus dorsolateralis

 

 

o. from the lateral margin of the drum
i. on the dorsal tin of the first senirinq with no fibers
extending on to the second semirinq

-21-

fl: sternotrachialis
i. on the lateral median margin of the cephalic drum
0. normal
Genus MICROURA Gould
Eicroura alhiventer (Hodqson)

The range of this species is from the northwestern Himalayas east
tJirough Assam to west China in Vunnan, Smechuan, 31d Formosa.
ELigroura alhiventer alhiventer (Hodqson).Scalvbreasted wren-qabbler

(Pl III, figs A a B)

M. larvnqosyringeus ventralis

 

o. from M. larynqotrachealis
i. on the ventral tip of the second semiring
c. extremely thin and weak

M. laryngosyrinqeus ventrilateralis

Indistinzuishahle from the preceding

H. larngoslringgus dorsalis

 

0. ?rom 1. laryngotrachealis
i. on the dorsal end of the first semirina
C. a weak muscle

F. larynzosvringeus dorsolateralis
Inseparahle from the M. l. ‘orsalis
.3: syringeus dorsalis

0. From the third tracheal ring (dorsal surface) with
a few fibers from v*e fourth and fifth tracheal
rinas

i. on the dorsal caudal margin of the drum

0. extremely weak with a large margin between the pair.
The right muscle is slightly more developed than
the left

M. syringeus_ventralis

.—

0. on the ventral cephalic rim of the drum

i. on *he ventral end of tie seconi sem’rini
c. hroad but 3!". t": little fienfih so it is f‘airlj' flat and weak

 

‘ l

”7 f‘!“1~‘. 0'31‘ ‘7' ,\

;. .3 l I. , >4 ~\-. lo-
...—I .....— ____... - ...—.-

.“ro'n the m ‘ian ‘orso-lateral refion of the irim

. on ‘he ventral caudal marvin o? the thir.’ a=miring
. this is a Sli'htlv wefik n scle - but ccrtzinly one
o” the strcn-est in this svrinx

o
i
c

“ol'1*¢% 9‘ '

  

I, . I ‘
:_-_- 53:23.12? 1: do

   

o. from the median lateril part 0? the drlm
i. on the dorsal tip of tie second semiring
0. weak

W. sternotracheglis
i. on the late“?l and medial pcavtion of the drum,
sonewtat.crv1ial to tiw3<xfliialic eive cf‘fiua drlw

C. hermil to week

The svrinx 1'n ”icréi“? alhivent r is the weakest of any

l

 

.3i_sscct d. The misclos wee not lzfiy and the “hole impre: ion 15

CH1e of ‘lxtness.

‘

‘ubfamily Turdoidioae

(J

Within this subfamilv are th-ood soecies of hiris iicludinq

tile larze lauqhiné fhrushes ani the small tit-haéhlers 13 well 13 other

l

t roe. There seems to be no one characteristic aspect abozt the

a

\

*1afiits or habitat of this wroup. dome soeCIe

(f.

, like the sihi's, are
esnixirely firhjrcql, while the laufihin: thrushes ire often laréelv
't€”‘TV?strial. "“13 lauvhirr*ifinfilshes are,zuy1+llv qroqarirxms, whereis
the? Eflirike-babhlers acre oFten are SOlltlTY. Bone speci s are met
“fiVth .at low al itudes, but the :olden filvetta is T913“h seen below
SEV73F1 thousand feet. Reqardins the voice, which *~ o? Stecial ’n‘zrc=t
héI"?, there is little correlation octween members of the sihiavilv.

Tlrrne ‘is so each '1riatioo in the cwll notes and soars t;it it is

imno 'L . . .
531+»le to say that any type of son: 15 tvpical of the firoup.

Genus GARRULAK Lesson

f

Garrulax striitus iiqors

 

This species ranges from the Kulu vallev east along the dimilayas
‘to Assam north and east of t e Drahmaputra, and northern Burma.

Elarrulax striatns sikkimensis (Ticehurstl Striated hauéhing Thrush

 

 

(Pl. III, ”iqs. 3 a 7)

larvrgggjringens ventralis

... -~._

h.

 

o. frim the ". larvnzotrachealis
i. on the ventral third of the second senirinq
C. stronfi

. larvn ‘osvrinrgezxs ventrilateralis

 

 

V’

This muscle is distinru‘shable. Lt lies lateral
to the preceding muscle and inserts on t*e second seniring

M. larynfiosvrinqeus dorsalis

o. 'ron the d. larvnfeotrachealis

i. on the dorsal tip of the first seniring with about
half of the fibers inserting on the end of the
second semiring

c. normal

gt larznggsyringeus iorsolateralis

This muscle is distinguishable, inserting on the
dorsal end of die second semiring

M. syringeus dorsalis

 

0. some fibers from the dorsal region of the lower trachea
i. a few fibers insert on the caudal edge of the drum
0. This is an evtreqelv weak muscle

’. svringeis :enjrbalis

 

o. from the ventral surface of the drum a half tracheal
rinq caudad from the cephalic rim of the drum

1. on the ventral third of the second semirinc

c. normal

-Qh-

M. svrinveus Ventrilateralis

 

“a

o. from the edge e the drum near W. sternotrachealis
i. on the ventral fourth of the third semiring
0. normal to weak

V"!
r‘

' .

svringeus dorsolateralis

 

o. from the lateral mardin of the drum caudad to M.
sternotrachealis
i. on the dorsal end of the first semirinz
c. normal
'3. sternétrachealis
i. on the lateral margin of the drum somewhat near the

caudal edge of the Tirst fused tracheal ring
0. normal

fiamlax iflW (Hardwicke)
This colorful species inhabits the northwest Himalayas east
tJirough the lndochinese subregion.
C311rrulax leucolonhus leucolgphus (Hardwicke).flhitecrested Laughing

Thrush (P1. IV, figs. A, B h C)

l_ryngosyringeus ventralis

I?

o. from the M. larynqotrachealis

i. on the ventral third of the second semiring (fibers
.do not seem to insert on the first semiring)

c. strong

M, arvnnos inreus ventrilater '

This muscle is distinnuishahle here and inserts on the
second semiring

fl: largngosyringeus dorsalis

 

o. from the H. laryngotrachealis

i. half on the caudal margin (dorsal) of the drum and
half on the end of semirinz number one

c. normal

M. larynggsxrin;ejs dorsolsteralis

 

This muscle issaiistinju‘shalfllexit is larvelj ihufimlzrith
M. l. lorsalis

H. svringeus dorsalis

 

o. finers seem to originate fro: the sixth tracheal ring
i. on the dorsal caudal rim of the cenhalic drum
c. a weak muscle

X1 szrjnceus ventralis
0. on the ventral cephalic rim of the drum
1. on the ventral end of the second semiring

c. strnnq

'3. svrirgeus ventri la teralis

 

 

o. from the lateral edqe of the irih
i. on the ventral third 0” the third 3;
c. nornal

0

hixfixié

\

I: . .‘-I ., a 1 - -. ‘
1.. syrin13.s fikn5$JLaterglfb

 

0. on the lateral marfin of the trzcheal irim
i. partly on the dorsal and of the first sehirin; and
partly near the dnrsal end of the second semirina

’; ‘ I"; . t“
m. sternntraciealis

 

 

i. on the lateral nargin of the drum just next to the
cephalic margin
0. strong

Garrulax ocellatus {Vigors)

 

This is a hirh altitude species ranging from Garhwal east

ifllrcnagh “anal and Sikkim to southeast Tibet and soathwest China

0 v ,‘
1n -unnan and :nechuan.

f‘ o ‘ n 0 ”'1
‘fl1233£lgi£ ocellatus ocellatus Vigorsh.ypotted Laughing .hrush

, . . . \
kPl. TV, figs. ), ‘/

L‘J

M. laryngasyringeas_zentnalii

o. from the H. larvngotrachealis

1. on the ventral third of the second semiring
C . normal

3

lgrgn’ouvrinveas ventrilateralis

This muscle is separadle, inserting more laterally
than the nrecedin: {an the second semirin;

1. lirngcsyringeas ;orsalis

 

o. from Phe larynmotrichealis

i. mostly on the iirst semiring with a few fibers inserting
on the caudal rim of the drum

c. normal

dorsolateralis

 

This muscle is idseparaole
. syrinjeus dorsalis
o. ”ron the fourth tracheal ring (dorsal sarface)

i. 01 the caudal rim of the tracheal drum
c. rather weak

 

M. svringeus ventralis
0. on the ventral side of the drum, Oie mm. from the
cephalic edge
i. on the ventral half 0' the second semiring
c. normal
M. svrinfieus ventriliteralis

 

 

c. on the lateral surface of the drlm
i. on the ventral and cephalad half of the third semiring
c. somewhat weak

3: syringe; dorsolateralis

0. on thr lateral surface of the drum, caudal to W. s.
ventrilateralis

i. mostly on the First sen rinw but also partly on the
cephalic and dorsal edge of the second semiring

c. normal

w.

e27-

sternotrachealis
i. on the cephalic lateral edge of the drum (actually

on the third Fused tracheal ring)
c. somewhat weak

garrulax lineatus (Vigors)

This bird prefers the neighhorhood of cultivation from Tad-

'zhikistan, Afghanistan, Vest Pakistan, India and southeast Tibet

ill the mountainous areas.

ngrznlaz_lineatus setafer (Hoigson) Streaked Laughing Thrush

1.3

n.

4
.1.
o

(P1.V , figs. A " 9)

_. largngosyringous ventralis

 

o. from the M. laryngotrachealis

1. on the ventral tip of the second semiring, inserting
on about one fourth of its length
c. slightly weak
larypaosyringeus ventrilateralis
This muscle is difficult to separate in this species
larvnaoslringeus dorsalis

0. From H.1arynqotrachealis

1. on the dorsal tip of U16 first semirint
c. normal

laryncosyringeus dorsolateralis
This muscle is inseparable in this species
syringeus dorsalis
0. on the dorsal surface 0” the fourth tracheal ring
1. on the caudal rim of the tracheal drum and a few
fibers to the tip of the first semiring

sgringeus ventralis

0. on the ventral cephalic edge of the syringeal drum

-28-

M. svringeus ventrilateralis

 

0. on the lateral margin of the drum
i. on the ventral half of the third seniring
c. so ewhat weak

r at
....3

. syrinzeus dorsolateralis

0. on the cephalic lateral portion of the drum
1. on the dorsal tip of the second semiring

fl; sternotrachealis
i. on the caudal lateral edge of the first ring (tracheal)
c, weak
Garrulax subunicolor (leth)

This bird is usually found in the semi-open and forested areas
fhccn Nepal east to the Assam hills north of the Brahmaputra, northern
{iirma, Yunnan and Tonkin.

CPIrrulax Sibunicolor subunigolor (Glyth) Plaincolored Laughing
flirlsh (Pl. V, figs. C & D)

.3: laryneosyringeus ventralis

 

o. from the d. laryngotrachealis
i. on the ventral end of the second semiring, covering
the ventral end of the first semiring
c. normal
M. larrn;osvringeus vent ilateralis

This muscle is difficult to distinguish in this species
M.larvn'oszrin:eus dorsalis

o. from the M. larynzotrachealis

i. on the dorsal caudal margin of the drum

c. nornal

_fl: laryngosyrinqeus dorsolatertlis

 

This muscle is difficult to separate from the H. l.
dorsalis in this species

svrinqeus dorsalis

.31

 

o. a few fibers spring from the fifth and sixth tracheal
ring

i. the fibers run beneath the M. l. dorsalis and attach
to the caudal surface of the drum

0. a v-rv weak muscle here

fl, vri e~ ventralis

0. just caudad to the cephalic margin of the drum

i. runninz beneath M.1. ventralis it inserts mostly on
the second semiring with perhaps a fiber or two
inserting on the first semiring

c. normal

He syringeus ventrilateralis

0. 6n the lateral margin of the drum, caudal to the
M. sternotrachealis

i. mostly on the ventral t ird of the third semiring. A
few fine fibers may run to the medial part of the
second rihg

c. normal

fl

.3, sternotrachealis

 

i. on the lateral margin of the drum just caudal to the
cephalic rim
Qazrula§_affinis Elyth
This high altitude bird ranges up to lh,OOO feet in forested
areas and is found from Nepal east along the Himalayas to Bhutan,
southeast Tibet, northwest Burma, Tonkin and western China.
EEEClflélgg§.affiinis hethelae Rand and Fleming Blackfaced Laughing Thrush
(Pl. VI, figs. A & 9)

EL laryndpsyringeus ventralis

 

o. from the M. laryngotrachealis
i. on the ventral third of the second semiring
c.nmmd

.-30-

M. larvngosvrinveus ventrilateralis
This muscle is distinguishable in this species and
inserts on the second semiring

11- 1._e_1‘;m'~:.0_8.:e:iasea§ dorsalis

o. from the l. larvngotrachaalis
i. on the dorsal end of the first semirinv with a few
fibers to the caudal rim of the drum
0. normal
L: larynaosvringeus dorsolateralis

 

In this species this muscle is fused with the M.l. dorsalis

M. svringeus dorsalis

 

o. a few fibers spring from the fifth tracheal ring
1. on file dorsal caudal rim of the tracheal drum
0. a weak muscle

M. syringeis ventralis

 

0. on the ventral cephalic rim of the drum
i. on the ventral end of the second semiring
C. normal

M. svringeus ventrilateralis

 

 

o. from the lateral edge of the drum
1. on the ventral third of the third semiring
C. normal

M. syringeus dorsolateralis

 

0. on the lateral margin of the tracheal drum

i. a few fibers on the dorsal end of the third semirinq
but mostly along the dorsal fourth of the second
scmiring

c. a weak muscle here

M. sternotrachealis

 

i. on the lateral cephalic edge of the tracheal drum
0. normal

Garrulax erythrocephalus (Vigors)

 

m o a Y '
efllJS colorful speCies ranges from the western Himalayas in

-21-

(Shamba and Murree east throush a pal to Assam, Burma, Vunnan,
the Indochinese subrezion a1d malava.

Clarrulax e_vthnoeephalus QiZZLELQtQN. 'Gutes)- Tedheaded lairhin

0Q

Thrush (Pl VI, figs. 3 & U)

34 . l»_rj.tn £9,331“ ‘7 n 3:: .1 s V'r.‘ n tr :1 l i S_

 

o. fron the E. larvngotrachealis
i. on the v~ntral fourtr of the second samirinf
c . nor'qal

H. laryngosgringels ventrilateralis

 

 

This nuscle is cenarahle, rltdnith weak

L; laryngosyringeas dorsalis

o. from the W. lirvnvotrachealis
i. on the dorsal end of the first seiirinm
c. normal

V. largnggsvringeus iorsoliferalis

 

3 “ll*% Senarfi’l: - more so than is
O I a ‘
rOSSihle in othe sneCies exanined

‘3. S‘.Ti-fl"’:Q-IS doqulis

o. fron the ‘ourth tracheal ring
i. on the caudal rin of the tracheal drlm
c. a stront nascle

syringe s ventralis
o. freq a sliqht distante cawdal from the cephalic rin
, _. _. . \
of tne drum (ventral s; e)
i. on the ventrll and of the second
C. normal

semirine

u. svringels ventrilateralis

 

0. on the lateral sarface of the tracheal lrum
i. on the ventral fourth of t*e third seniring
c. nnrmal

Ea Svrinneig jorsoliteralié

 

o. from the lateral Sirface of the srrinqenl drum

i. on the dorsal end of the first semiring; covered
completely hv 1.1. dorsalis

C. somewhat weak

:1. at ermuachea 113'

i. 01 the lateral cephalic edqe of the tracheal dram
0. normal

Genus ACTINfilWTA Gould
Act nodura higjlensis (Hodgson)
This arhoreal species is distributed froi Nepal to Assam
and northern Burma.
Actinodura njyal,ensi_s 1mm Riplev, Hoarv liming (Pl. VII, figs. A .9: 3)

;; laryn;g§;§;ngels ventralis
o. from the M. laryngotrachealis
i. nostly on the ventral end of the second seniring
with a few fibers attaching at the hiddle of the
second semirina
0. normal

74'
4..

larvngosvringeus ventrilateralis

 

This muscle is fused to the M. l. ventralis in this species

M. largngosyringeus dorsalis

 

from the M. larvnqotrachealis

. on the first semiring - not quite at the dorsal end;
a few fibers attaching an the caudal edge of the
tracheal drum

C. This muscle is extremely massive just above the

tracheal drum; normal otherwise

H‘O
O

‘l

A. larvngosvringeus dorsoliteralis

This muscle is fused to K. 1. dorsalis in this srecies

 

 

. syringeus dwells

0. “rom the fourth aid fifth tracheal rings
1. on the dorso-lateral and caudal edge of the drun,
leaving a separation of 1.5 mm.

3: sgzinaegs ventralis

0. from the ventral cephalic edge of the drum and the
ventral caudal edge of the first tracheal ring

i. on the ventral end of the second semiring; a few
fibers to the first semiring

...~\

_, EXEE?IGB§ ventrilateralis

 

o. from the lateral cephalic margin of the drum
1. on the Ventral end of the third semiring plus
attaching along the cephalic margin of the
third semiring for a third of its length
c. normal

in

-. syrinteus dorsolateralis

 

0. on the lateral region of the tracheal drum (actually
on the second fused tracheal ring)
1. on the dorsal end of the third semiring

N. sternotrachealis

.—

 

i. on the lateral cephalic edge of the tracheal drum
‘0. normal
Genus MIELA Hodgson
Eisla airfizsla Uodgson
'The minlas are found in the Himalayas from the northwest through
Asseun ;and into the hills of Runna, western and southern China and Tonkin.

filfllfi: :stripnla §§quu1§m (Hodgson) Stripethroated Minla

(Pl. VII, figs. C, D a E)
.M; larynzoayrinseus ventralis

o. from the 3. larvngotrachealis
i. on the ventral end of the second seniring

I

———

V
\

 

:.‘
O

u

 

,-__

c. tier“nal

1.7-}: '23-;9‘a::r.‘i.n:;:11_5 mllfihs
.v ~-.j ' Lu'c- an; '.;q x'+1 l'; 1 7+ NH
hufDQ‘ 1" '/.1.Q Qt .01 r» WI]. '1 A. .-.0 VP .;r-i..-S

larvnrnsvringens dorsalis

irthotrachealls
P

o. fren the ‘
(T. nordal

larvn305”riw{exs dorsol teralis

..__- ..

 

. l ‘ . C ,
lnscoaranle in this SIQClRS

v ' ‘n ’1 y“ ' o ‘
sirzn'e.s .orsllis

-..—.-._ .. _

3. ”red the ceuhalic dorsal Odie i: the first

1. on the CEHTQI Tlrflw 0: mid fr10“3“i firlm
c. strvnf

f" - . .‘ ' A 1 0
913133910 vc trails

...-...

0. fron the vent“al cephalic rim of the dram
i. on the ventral half of the second Sinirin‘
c. nornal

3"rinqeu3 ventrilateralis

 

o. from the lateral medial margin of the trac

i. on the veltrll third of the third seniring
0. normal

syrigggus dorsolateralis

 

o'— )q
. A

i on the iorsal cerhilic eige oi the first senirinv

v: r31§ [n I}

.931 drzh

rini

o. from the lateral surface 0‘ the draw vewtral (candid)

to '. sternotrachealis

i. on the dorsal cephalic edge of the second semirinq with

I."

a third inscrtinq on the end oi the ji_

7. sternotrachealis

--_.—-_,___._,-__-‘ ...---

i. on the lateral surface 0? the drum near the cerhalic

elqe o? the second Fised tracheal ring
0. weak

5‘;

C
.;A‘ ..1 l \
g;~_,.. , - ..

.‘t r

Genus ALTTPVW llvth
ilcinne cggtgneceps (Woiqson)
The tit-habhlers ire ioznd in the ”i alavas from ”e‘al east
throagh 91:35am to huria ani Vunnan.

Alciojg gastanece s chunbiensis (Kinneir), Chestnutheafied Tit-

?asnler (P1. VIII, £123. A, 3 a)

‘.‘.2

E: lerrnsosrrifi":ae tajlaalis

0. iron tie larvnrotrachealis

i. on the ventral tiird of the second semirinv with a few
fihers to the third semiring

c. weak, esoeciallv at the too where it fuses into the
V. larvniotrachealis

h- larynzosxtiaasus ventriivternlis

...—..-. .7-

 

“-—

This muscle is finite distinct and also llSBTtS on
the second senirinq

M. larvnrosvringeis dorsalis

 

o
q

o. from the H. larvngotrachealis
? rst semirina

i. on the dorsal and o” the -I _
c. weak
1. la‘”yfinfo s‘fri n-“eus riorsol iteralis

 

Fused in this species

I

[L_ Sfrin 8!S dorfi

'?
u - J

alis

O
U

0. on the caudal marjin of the ’orsal sarface of the
first tracheal ring

1. on the dorsal caudal margin of the irum

c. sonewhat short

.3: svrinqeus ventralis

o. fron the ventral cephalic edne of the drim

i. on the ventral medial tin of the second senirinq
comoletelv covering the first seniring (ventral and)

0. rather nronounced

M. svringcus vontriliteralis

-m ”H-

 

n. fr‘n the literal cenhalal portion of the drum just
caudal to V. stornotriohealis

i. on the ventral tir of the third senirinq

0. normal

js firsolateralis

 

 

o. frriije rwdial lateral margin of the drum
i. on the ventral end of the second semirinq
c. weak
,2. sternotriciaalis
i. on the lateral cephalic edge of the drum
0. nornal
flair .e {lijsljfllsié (Hodgson)
This unobtrusive species is found from central Nopal east along

the Himalayas through Assam, Burma in the hills to Yunnan, the

-£ndochinesc snbrcgion and Malaya.

.3

.Alcipn nigalensis inalgnsis (Hodgson) Nepal Babbler

.—

 

. -..—......—

1

(Pl. VIII, figs. C h D)

I. larynqosyringeus ventralis

 

l

o. from the L. laryngotrachealis
i. on the ventral and of the second seniring
c. somewhat narrow

M. laryngosyringeus ventrilateralis

 

 

 

This muscle, although thin, is distinguishable here and
inserts on the second semiring

y. laryngogyrinqeus dorsalis

o. from the H. larynqotrachsalis

i. on the dorsal caudal edge of the drum with a few
fibers continuing to the dorsal end of the first
sgniring

c. broad

:v
r‘
O

laryagos"rinvais tonsolateralis

mhis nuscle nopenrs to be fzsed in this species

'1’
0

syringeus dorsalis

 

o. tron the ‘irst tracheal ring
i. on the ventral caudal edge of the drlm
c. extrenely weak

"I"
§

;. svringeus ventralis

1

o. fron the ventral cephalic edge of the drum
1. on tie ventral end of the second seniring

c. strong

m. Svrznveus ventrilateralis

0. on the lateral aargin of the drum below 1. sternotrachealis
i. on the ventral third of the third semiring
C. strong

w. svrinqeus dorsolateralis

 

 

0. on the lateral nedial side of the drum below M.
sternotrachealis and dorsal to M. s. ventrilateralis

i. on the first senirinq with a few fibers running to the
dorsal end of the second seniring

c. strong to normal

9:

e. sternotrachealis

i. 01 the cephalic literal edge of the drum
0. normal

Goi1s WETZRCFHASIA llyth
Ustsrpyhasia caviataata (Visors)

This is an arboreal snecies ranging in the Himalayas from

‘

:hxrree east to Rhutan and southeast Tibet.

‘igpgrgphasia gapistratg bayleyi (Kinnear) Blackcanoed Sibia (Pl.IX,
“' figs. A & 3)
g; larvnzosvrindeus ventralis

o. from the V. l rvnaotrachealis
i. on the ventral end of the second senirina
c. normal

 

-.acr-
_ o n

. lnflr ......- ,4

 

‘0

p-o
«i

I 5

l1.

0
r:\

 

— 3(2-
qv "' v ' 4 ‘j
lar n19.5r1.3o.s fiJril 1121a 1 s
Th‘s a1 cle ‘Ls h tin 'ii"“ 01' - 'Jtsertfrir on fir:
.nco d 5‘..r11'
larvc'co‘rln"c:s 1ors ‘ l5

0. fret the I. larvn‘ztrachcalis
i. on the dorsal end of the first 53miring
c. lior"'al

lorvn'ocvrinW-Is

-w—a~__.——_—~"__————._— -

Isolateralis

This muscle is i senarahle in this snecies

.sgv'rin 333-: (baselie

o. frcv the ”irst tracheal ring
i. on the caudnd margin 0? the

C. weak

svrinreos 1e.. lis

o. Tron the cebhalic ventral nargin of the drtm

1. on the ventral thirfl o? the
c. normal to otron

s7r1n1ezs ventrjlaferelis

_—-—- a-.---

0. on the lateral margin of the

1. on the ventral third 0? the
0. strong

gyri nccus d_ors lolateralis

second senirinz

irnn

third sehirinq

0. on the lateral margin of the drum

i. on the dorsal end of the se

;§Q?39tr39h931i3

i. on the lateral elze of the
cephalic rim
c. weak

fond and third semir

ch

ll

74
(.0

”run just caudad +0 the

-31..

Suhfahil* Cincloswnatinae

This is a ironp of forest-iwollinz birds onlv rarely met with
in the New Gainea and Australian hills. The Sdbfahily is included

n
Q

hgre for nirnoses oi com arisen? althonvh I have not iissvctei any
members of the so group.

The first svrinqeil investidation in the ?imaliidae was nade
hv Muller (1955). He placed his Timelia with the other "norhal"
songbirds. Forbes (1882) resorted his findings on Lrthonvx. He
said that the svrinx in this genus was typical of songbiris and had
the normal conplenont of foar pairs of muscles.' He mentioned that the

"short anterior" huscle rins to the anterior end of the third

bronchial semiriif in O. soinicaid but that in O. ochrocgghala

F‘J

this semiring receives its muscular supply from a faociCilus o the
"lone anterior" nnscle. The "short anterior" nuscle is host likely

7‘. laryneosvringens ventracis combined with h. larvngosgrinwens ventri-

L—

 

'lateralis ani V. syrindens ventilateralis,
wifty vears later Mayr (1931) investigated the confignaration

«of the svrinx in four flew Guinoa species of three genera - Gymnorhina,

 

’“elampitta and Orthonyx. He located ten intrinsic syrinteal muscles
'inclndinq an extrenely short 2, intarhronchidlis lateralis which
runs from the iirst to the second bronchial somirinfis and does not
reach the tracheal drum. Mayr described many details of his work

n these in mind I am oresentina excerpts from his

arui so as to kee

lflnvcr on the following Wages.

.-h0-

Key:
fiymnorhina - Gy; Orthonvx - Or; Melanoitta luoubris - 31

U.

 

 

v

Melamgitta qigantica — Hg.

 

M. lfiyngosyrinfeus ventralis

 

dy. vigorozs: from the tranhea to the_second semirinz

Or. ins rts on the second semirine; weak

Fl. well develonei.; inserted on the secon? semirin: literal
and caldal to I. svringeus ventralis

”2. relatively thick aid not vsrv lonr; ?ron t‘e sevezth
and eifhth tracheal rini to the second sehirin:

M. larvnqosvrjnqeus ventrilateralis

 

 

 

3y. clearly strong; "ovariaz V.s. ”entralis

Or. 5 eninzly wile and strong;to the third seniring

Tl. lateral to ”.s. vn tralis

Ego relatively vigora.s; difficult to tell fron M. s.
veitralis

F. lgjynrosvrinceus dorsalis

c—_. v.» - _ ......

 

3y. caudal heads are divided into two - one to the
drum, the oth r to the first semiring

Or. weakly developed; inseparable from M. l. dorso-lateralis
El. Fused with H. l. iorsolateralis
fig. 71sed with H.l. dorsolateralis

§L_larvngosyrinqens dorsolateralis

 

 

Gy. thin; onlv on the second seniring
Or. weak; on the second senirin:

H1. fused with T. l. dorsalis

“r. "ased with W. l. dorsalis

*‘y

U

Eb swringeas dorsalis

fiy. stronq

Or. not verv stronj
Fl. “ery weak

Tfig. ”weak

5- Siriosee insatra'iis

Gy. snall, entirely covered; on the eniring

3
Or. wile, vigorous; medially pashel togetfi

-L1-

"

m1. large, almost uncovered
HS. Completely covered by g, 1. ventralis

M. svrinjeus ventrilateralis

3y. not stronf; to the second 3 hiring

Or. weak; scarcely senarahle from H. l. ventrolateralis

Ml. short and stronz

M2. 10»: and stroni, partially to the €irst senirin”
also "-rtlv covered \y M. l. ventralis

M. syringeus dorsclateralis
1v. stron9° ventral to it is the V. interbronchialis
n, 3
laterilis
Cr. clear, strong
”1. strona - to the second senirind
“T. very strong - to the first and secind senirinas

M. sternotrachealis

all normal

'7)

The foregoin: data indicate that there is not a great degrie of
‘Variation in syringeal musculwture thriuthout the wh~le family.

1';
lfiiskimen (1951), however, noted a great deal of variation withfithe
P‘r’mg‘llidae and other families with which she worked.

Host of the variation described here is of a minor nature. These
V«‘ar‘iations allow ior some flexibility depending unon where the muscles
iJisert on the semirina and to which rings the muscles attach. If the
nInsole inserts on the ventral third of the senirinq instead 0? the
Verflxral fourth, then one would consider the ?ormer less ?lerible.
ThEBVnost advanced species, in svrinqeal developnenf, are {hose that

‘have the greatest nimber of nuscles and have then attached to the

be
- L“

extreme ends of most or all the senirinzs. A less advanced or poorly
zievelonei svrinx mav, however, not he an indication of primitiveness
VJ en other str;ctures are taken into consideration. The less fletible
syrinx is merely an indication that the bird is less developed and

his should he considered in conjunction with all other pertinent
=anatonical, morphological and behavioral features before de’icite ,
rfiivlozen,tic relationships can be dete mined.

From the data nresented in this saber it apnears that the [

:subfanily Pomatorhininae has the weakest and most noorlv developed

yrinx. The M. svrinceus dorsalis is ahsent in the true scimitar
laabhlers and is verv weak in the wren-babhlers. In ”act the one
ivren—babhler (Microura) had the weakest syrinx of any srecimens
dissected. Only the V. svrinzeus ventriliteralis was as large as
one would exnect. Another point here, recardin: the Pomstorfiininae,
is that the long ventral 'uscles (the M. laryncosyringeus ventralis
eand the M. larynqosvrinqeus ventrilateralis) are broader than normal,
curvering the deener muscles completely. This indicates that even
ttuiugh the wiole svrinx is somewhat weak, at places it is stronger
than normal.

In the Pellorneinae the svrinx is also quite weak. The ventral

rnusculature is esnecially flat and weak but the dorsal muscul tire
iS rwuwuflu Another characteristic that should be noted is that there
is rue muscle attachment on the dorsal end of either the third or the
SGCCHnd semiring. This is also the case in the Pomatorhininae tnd

Cerfluainly renders the syrinx less flexible. The Pellorneinae syrinx

-L3-

is more ievelooed thin those in the “omatorhininae but less developed
than all examples studied within the Turioidinae.

It is imnortant, also, to see how much vari«tion occurs within
a subtaiilv. From dissections of twelve see:ies within tee Tard-
oidinae it ancears that there is not a great deal oi variation in

‘

this subfamily. Onlv in darrulat subunicolor is the dorsal end 0:

 

the second senirine left without mus ular attichment. The svrinx of

ilcipre castan,ceos differs from others in the subfamilv. Yer: tie

 

lone ventral muscles, the H. lirvnxosyrin'els ventralis and the i.
laryngosyringeus ventrilateralis as well 38 M. svrinteis dorso-
lateralis are rather weak. however, the M. syrinreis ventralis

is sonewhit cronoanced -nd this in cart strengthens the musculatire
on the ventral and of the second semirina. It might also be noted

that the small Vinla strigula svrinx is remarkahlv similar to the

 

larger Girrulax svrinces.

Another interestin7 diestiin revolves around whether or not
there is much variatioc in the svrinx within one genus. Many veneric
studies will hvve to be m-de beiore this can be conclusivelv deternin d.
From the data oresentei here it is tossihle to examine one jenus.
Seven snecies o? garrulax were dissected which is about a sixth of tie

snecics in the genus. Comnarinr these syrin es shows a remarkable

'Jo

stability in the configuration. All eieht muscles are present n
all the snecies examined and wit} arrroxinately the same developt nt

in each. There is a slirht difference in Garrilax subunicolor in

 

imich the SVrinx is so ewhat elcnva+e. The differences that do

41);-

appear in these snecies ire nainly in the insertions o? the nascles.
The M.svrinreus v ntrilxteralis in,erts on the ventral third 07 the
third sehiring in four sneoies and on the ventral half in three soecies.
The M.svringeus iorsoliteral‘s inserts on the dorsal ends of both the
second ani first senirinfis (with sliqht VlrlltiOnS) in six species

of Garrulax while in larrulax suhunicolor it inserts only on the first

 

semiring. These are merely minor dif’erences which indicate that
some svri-qes are slifhtlv more Flevihle in certain snecies than in
others in the same genus.

From Tayr's (1931) work it annears that there is more variation

within the Cinclosonwtin 9 than in the Turdoidinae. Within the

genus Telajgitta he noticed several dif”eren,es in the develonnxnt

 

0? various nuscles between two snecies. His work indicwtes thit
there mav he considerahle dif”erence between snecies in sone genera
but other investigations need to he hide. The svrinx in the
Cinclosom.tinae is as well developed as it is in the Turdoidinae
and in host cases is ah01t as Fletihle.

Two neglected areas of syrinceal investigation are the histology
of the syrinx and its enhryolovical develohment. Comntritive
histology of the svrinx may not he too siqniTicant but at leasF we
need to know the histology ‘ld how the nicroscocic anatomy varies
“rem group to qroup. r‘rnbrvoloqical studies would he usetul in trving
to deternine the homology 0“ certain muscles in the less developed
svrinx. heretoéore the only paper coverinz these asnects arre2rs to

have bean done on the domestic chicken (Tvers, 19175.

'T‘Vfil't' fVfi‘f

.. J“.

This naner rerorts on syrinacal inves+iqations in three subfanilies
within the Avian ’amily Tinaliidie. The purroee of this study was
to detorhine whether or not the svrinx is chiracteristically uniform
throughout a family mid whether or :ot it is a useful tool in
determining the phylo: netic relationship of birds. The history of
svrihgeal investi*atfons was traced from F. ”icq D' Azar in l7“?
up to the nresent century. V

The specim3ns used in this study were collected in Ilam distict,
E. Neral. Forty snecioens renresentihg l7 snecies, 9 tenera and
3 subtanilies were examinei. The venue Warrulax was the host
thoroughly studied.

The syrinx is the "voice hox” o? birds. It is an enlwrged reqion
locwted at the hifurcation of the trachea +nd the bronchi and is
usually associated with one or nore pairs of extrinsic and intrinsic muscles.

...

The syrinx in the Timaliid'e is remark hly regular.

“3 1

L .

A

git
intrinsic muscles are present in the Turdoidinne ani the Pellorneinae
whereas in the Pomatorhininae the true sc'nitar hadhlers have only

seven. The syringes in the Turdoidinae and the Qinclosomatinae are

similar and are equally well developed. The variations that are

present Are minor ani orobablv af7ect only slightly the flexio‘lity

of the svrinx. The degree o? develohment of the syrinx in the Tellorueinae
is intennediite between the Fonatorhininae and t‘e Turioiiinae.'£ithin

a narticular genus, the Clarrilax, the syrinx is also quite uni"orm.

The only Viriation noted was in ‘arrulax suhunicolor in which *he

 

dorsal eud of the second senirin; is left without easellar attaehnont.

r» 4
"3
L '1

LITEQI“WQE 1

Appel, F. H. 1929. Sexual dihorphisn in the syrinx of the fowl,
Jour. Morph., h7:h97-Sl7.

Reddard, Frank E. qu5. On the itlrcturql Uhqfiaqte‘s and slag; _
iiioution of he Cuckoos, Tree. 7001. Soc.

London, 1995:168-l7h.

 

1383. On CGFtBll points in the visceral anatony
(sic) of inlaenceps rex, beiring upon its

affinities, Proc. Zool. “oc. London, lghfitfh3-288.

 

 

1898. The 3truct re an? Classificatiin of Wirds,
pp. 60-71; 177-193, Lonrnans, Green and 00.,
London.

 

1902. On the syrinx and other points in the
structure of hierococgyr and sons allied tenera
cuckoos, Ibis, 1932:59V-508.

 

 

1erger, odrew J. 1960. Some anatomical characters of the Juculidae
and Vusoohazidae, flilson Bulletin, 72:60-1Ob.

Cole, Arch E. 19b5. Song-production mechanisms of dirds, Kentucky
Warbler, 21:17-27.

 

Cuvier, G. 1802. Uber den untern Larvnx der Yerl, nuts. Arch.
5. d. Physiologic, 1d. V, 33113.

 

Delacour, J., k C. Vaurie. 1957. A Classification of the Oscinos
(Aves), Los Anneles County Mus. Contrih. S ience,

No. 16.

F orhes, W} A. 1880a. Contributions to the Anatomy of Passerine Bird: -
part II, Free. 7eo1. Qoc. London, 1380:383-385.

 

1380b. Contribitions to the Anatongr of Passerine

Birds - part III - on some points in the structure
of Philegitta and its postion anonqst the Passeres,
Proc. 3001. Soc. London, 1880:339-91.

 

 

1882. Contributions to the Anatomy of Passerine
lirds - part V, Proc. 2001. 000. London ,
1882:5Lb.

 

Far43rineer, M. 1988. "Untersuchungen sur Morphologie und Systematik
der V82e1 zugleich ein Beitrac zur Hnatomie der
Statz - und Bewegungsorgane," 2 vols., Van
”olkena, kneterdam.

 

‘ 7v

Gadow, Hons., & E. selenka. 1391. :63e1. In Bronn's Klassen 1nd
Uninungen des Thier-ReiCis ljltoqlnfijon (5311,

 

 

 

Earrod, A. U. 1873. On sohe points in the anitonv of Stestornis,
Free. 5061. 30c. Lonion, 1973:531-533.

 

 

 

lfl76. 7n inutonicel chiracter which hear upon
the naior divisions of Passerine girds, Proc.
7001. Soc. London, 1876:305-516.

 

 

 

 

Wicker, Valentin . 1900. ler lesnni der o3e 1z sine aiatohisehen uii
hiotogischen Wrundlizei, 3. Fisch)r, Jeia.

 

 

Herre, Richard. leQ. We eviun passérinsrum larvnze hronchiuli,
Dissertatio anatomica, lrvphiae

 

himohrev, P. 3. 1758. Classi fies ion and S"3tematic position of the
Eiders, ,onior, 60:129-133.

Koditz, J. 1925. err die Svrinx einifler Clanstores uhd auslhndischer
Oscines, Zeits. f. wiss. 7oo1., 126:70-1hh.

 

avr, E. 1931. Die Svrinx einieer Jinqv63el aus SeuGuinea,z Jour.
fur Ornith., 79:333v737.
Miller, A.H. 193h. The Vocal lpparatus o? sohe North American
Owls, Gondor, 36:20h-211.

35. The Vocql ipparatus of the Elf L)wl and
Spotted Screech Owl, Condor, 37:239.

 

19h7. The struct tural base; of the voice of the
F1 anmulited Owl, Alk, 6h:133-l35.

 

Miskimen, M. 1951. Iound Production in Passerine Birds, Auk,

1:13, 68 : 13.19 3' 5011 o

1957. Absence of syrinx in the Turkev Vulture
Cathartes aura), Auk, 7h:1Ob-S.

Muller, Johannes. lBhS. Uber die hisher unhekannten tvpischen
Verschiedenheiten 1er Stinrorzane der Passerinen,
Abhandl. k. Akad. Berlinz 18h5.

Myers, J. A. 1917. Studies on the syrinx of Gillus

 

Newton, A. t H. @nlow. 1896.
Adam and

Dictionlry oi' Birds, pp. 937-9h2,
Irles Black, Lond .on.

 

 

‘
1
jh

Owen, R. 1835. Article "Aves" in Todd's Cyclopaeiia of Anatomy
and Physioloi Vol. I London.
I 3 9

1866. 0n the Anatomy of Vertebrates, Vol.11, London

 

 

 

Ripley, 3. D. 1961. .A.Svno sis of the Birds of India ani Pakistan,
pp. 389-518, 90m. Nat. Hist. Soc., Gombay.

thpell N. 1933. Physiologie und Akustik der Vogelstimme Jonr
, ’
g. Ornith., 81:h33-5h2.

Savart, M. Felix. 1826. Memorie sur la voix des oiseaux, Ann. 23
Chimie 3} Ph si.ue, T XXXIII, Paris. Ubersetzt
in Frorieps Notizen, V01. XVI, 1826.

 

Setterwall, Carl G. 1901. Studier ggver sgrinx hos oolvmxoda
Passers, Lund.

 

Stresemann, Erwin. 1927-193h. Aves in Handhuch der Zoologie, Vol. 7,
pt. 2, pp. 186-190, Kukenthal‘Walter de Gruyter,
Berlin.

 

Vicq D' Anyr, F. 1779. De las structure des organes qui servent
a' la formition de la voix consideres dans
- 1
l'iomne et dans les differente classes d'Animaux, et
compares ent' reux, Hem. de 1'acad. d. sci., Paris

 

Wunderlich, L. 1886. Beitrage zur vergleichenden Anatomie und
Entwicklungsgeschichte des unteren Kehl-Kopfes der
Vdgel, £913 Asia d. Kais. Leflp.- arQl. gggtggh.
gggggg}e_g, flgtggf., h8(1):1-80, Halle.

Yarrell, W. 1833. On the organs 0? voices in dirds, Trans. Linn.
Soc. London, 16:305-321.

 

 

  

LLTSCLf 7fi§T7I'Tfllfi?Y
. p
’ ‘ ............. trachea
// ...y..;........... larynqosyrinqeus ventralis

3...;)ad............................ sternotraéhealis

7;" ..o' \
If . .i' \\ ooooooooooooooooooooooooo llrynflosyringeus Vintrilateralis
r .0 / K
5 f I K
, r ..q......................... syrlngeus ventrllaznralls

...~......fl....................a.. svringeus dorsolaterqlis
I
L x

I ~ ...... syringeus ventralis
.... ....................... .......... .. larynjosyringeus

’ ‘ :orsolateralis

' \ M . . .- . .
.............?$;<1¢...... thlrd bronchlaL semlrlnq

    

 

 

 

 

svrinzeus dorswlis ..................... .~;,--

trAChea 0-00000000000000000000o... 0000.00.00.

/ .“h -.w \

SternOtFECheflliS 000000000319. '1 \\

"1'21;

:Laryngosyringeus
dOFSOliteraliS 0000000000000.

ilaryngeosyringeus dorsalis ...........

    
 
 

/' V
, n . , . I \
Eifrlngeus d0r5013teralls 0000090 I f{” \\ i
. ‘ I ‘ ' I ‘ l ‘
\~ . If [I I ' . I .‘

Iflassulus ......................Tk,g' .. ......f \;.' a?
X W 4 s A
,‘~‘. fl , "u” I

r 3? 9' 4 «9;

Fig. B - dorsal view

 

 

PLATE I

Pellorneum ruficqps Swainson

 

Fig. A.... lateral
Fig. Bo... dorsal
Fig. 0.... ventral

Pomatorhinus ruficollia Hodgson

Fig. D.... lateril
Fig. E.... ventral

I

PL HE

avg .1L

 

A .

‘u’.’
Lav. \ \1- /.
;,
.\
x w
* I
y
.
_
a
b!” Inf-cl Y
z
.‘ J

 

PLATE II

Pomatorhinua erythrogeqys Vigors

 

Fig. A. ..... lateral
Fig. B ...... ventral
Fig. C ..... derail

Xiphirhynchus superciliaril Blyth
Fig. D ...... latennl

Fig. E .0000. Ventrd
Fig. F ...... dorsal

 

 

,. .
I.
I ,
V C
1‘ l .
/
TL.
I
“0 rl
H.) :4.
. f -

L.
w...

s v

:40 ¢
I 0 .
x. \
§ \
I I In .... 1‘!
\“\,\\ I . I P .
’.
x o
1 RH

 

93-;
\‘r- »
IV”
,2
1'":
,x /

PLATE III

Microura albiventer (Hodgson)

 

Fig. L........latera1
Fig. B 000.00 ventral

Garrulax striatus Vigor:

 

Fig. c 0000 laterd
Fig. D .0000. ventr‘l

PT
l...’

.fi"
1...)

III

-. \.-;
a M” ' I
c‘w— -.—
A‘ - \ ‘
f \ I /
‘._ _ 'w-o‘
«a; ’ ‘
/\‘ i'j ‘ 3A
'. - I
/" \K ‘ I".
5“; .
I . . ' -’
ow

 

 

 

PLATE IV
Garrulqg leucolophue (Hardwicke)
Fig. A ..... ventral
Fig. 8 coo-e lateral
Fig. 0...... dorsal

Garrulnx ocellatue (Vigors)

Fig. D coco. Ventral
Fig. E ...... lateral

" 1/11-

e..u\\

 

q,
no
..
L
N
M
....
[A
a

I

....- '

i\_

\J

/

‘l

7/

I

r

M
‘I

‘n.
x

,)

1

o‘ f]

. 2 i 3'.
_‘,__A__';C:1‘)
__‘

.-.. .---.l-t~-r --»- ‘

.

.

.V, a.

.. . .-

. ‘ .

o a I
.

. .

f
5:. _.,
w4~- - ---

4 - —-—-—r -‘-o

6- \. ‘-~~-
\
D

\E: “"
Cf."

PLATE V

Garrulax lineatus (Vigors)

 

Fig- A 0000.0 lateral
Fig. B ...... ventrel

Garrulax subunicolor (Blyth)

 

Fig. C .......1ateral
Fig. D 00000 ventral

V
-93
A ‘

‘T._

 

 

PLATE VI

Garrulax affinis Blyth

Fig. A. .... lateral
Fig. 8. a... dorsal

Garrulax erythroceppalgg (Vigors)

‘Fig. 0...... lateral
Fig. D ..... dorsal

a
I
W
l
A
I
. .
I '1
__¢
\‘l I I ..-"
.I"
, \
- J"
."
a

‘a‘

N‘ ..
\\ -- a-
‘
. _ H- .. .-. -
l'fi
.' o

O
l
A_-‘ I
‘7 ,
S..—

 

 

PLATE VII

Actinodura nipalensie (Hodgaon)

 

Fig. A....... lateral

Minla strigula Hodgean

 

Fig. B0000... laterd
Fig. C....... ventral
Fig. D....... dorsal

 

 

 

II

 

'7'"? ‘7
J

L

PL A

A.

 

 

 

 

PLATE VIII
Alcipge castanecepe (Hodgson)

Fig. A..... lateral
Fig. B ...... ventral

Alcinpe‘gipalensis (Hodgaon)

Fig. C000... lateral
Fig.0....... dorsal

VIII

n
d
u

"L n

 

'*.—. "

..-._ I ,

 

 

 

PLATE IX

Heterophasia capistrata (Vigors)

 

Fig. A ...... lateral
Fig. B....... ventral

IX

NE

T
.1

P;

 

 

 

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