Myology of the Pectorla Appendage in Kingbirds (Tyannus) and Their Allies

The Condor 87:402-417 0 The Cooper Ornithological Society 1985

MYOLOGY OF THE PECTORAL APPENDAGE IN KINGBIRDS (TYZWWUS) AND THEIR ALLIES

MARY C. McKITRICK

ABSTRACT. -The forelimb musclesof several speciesof the kingbird group are here described and illustrated, with the goal of discovering character statesof use in delineating cladeswithin the Tyrannidae. These musclesconform in all species to the oscine pattern as described by other authors, with the exception of a few features. Special attention is given to M. latissimus dorsi caudalis, the only muscle that varies widely among, as well as within, the speciesexamined. Absence of this* musclein the speciesofMyiozetetes could mean that this group is a clade. Variation in other tyrannids, however, indicates that closely related speciesmay be losing M. latissimus dorsi caudalis independently.

The present study of wing muscles in the Flycatcher (Legatus leucophaius USNM “kingbird assemblage”(Traylor 1977) was un- 5 10867), and Variegated Flycatcher (Empi- dertaken as part of a larger study, to determine donomusvarius FMNH 288494), all members whether the appendicular musculature of New of the kingbird group. One muscle, M. latis- World flycatchers (Tyrannidae) can provide simus dorsi caudalis, was intraspecifically information of use in constructing phyloge- variable, and I examined this muscle in 39 netic hypotheses for this group. Anatomical other speciesof Tyrannidae to assessthe extent data from the work of Garrod (1876), Ames of variation. Dissections were done under a (197 l), Midtgard (1982), and W. E. Lanyon stereomicroscopeat 6 x , 12 x and 25 x , with (pers. comm.) have recently been used in a the use of iodine stain (Bock and Shear 1972) cladistic analysis of relationships within the to enhancevisibility of muscle fibers. Anatom- Tyrannoidea (McKitrick 1985). Other ana- ical nomenclature is from Baumel et al. (1979) tomical data for this group are scarce and in- and abbreviations are, for the most part, from complete, and much additional information is Zusi and Bentz (1984). Descriptions of mus- needed from a variety of anatomical (and mo- cles refer to Tyrannusmelancholicus, and are lecular) systemsto clarify relationships within the same in the other speciesunless otherwise this group. stated. Comparisons with other passerine A few published studiesof the forelimb mus- speciesare noted where differences occur. cles of oscinesexist, e.g., Swinebroad (1954) Abbreviations for museums from which Hudson and Lanzillotti (1955), George and specimens were borrowed are as follows: Berger (1966), Borecky (1977), and Raikow AMNH, American Museum of Natural His- (1977, 1978), but I have found virtually no tory; CM, Carnegie Museum of Natural His- such information for suboscines.I undertook tory; FMNH, Field Museum of Natural His- to describe and illustrate the forelimb muscles tory; KU, University of Kansas; LSU, of several speciesof the kingbird group, and Louisiana State University; USNM, National compare them with what is known of these Museum of Natural History; YPM, Peabody musclesin other passerinebirds. My purpose Museum, Yale University. Collection data were was to determine polarities of character states, reported in McKitrick (1984). and thereby assessthe potential of this anatomical system for phylogenetic analysis.Sim- ilar data for the hindlimb musclesare provided RESULTS AND COMPARISONS in McKitrick (in press). My working hypoth- M. rhomboideussuperjcialis (Fig. l:RS) is a esis was that the Tyrannidae is monophyletic thin, flat, essentially parallel-fibered muscle, (M&it&k 1985) and, further, that the king- lying deep to M. latissimusdorsi pars cranialis bird group is also a clade (Lanyon 1984). and superficial to M. rhomboideus profundus. It arises by an aponeurosis from the neural MATERIALS AND METHODS spines of the last two cervical vertebrae and I dissected fluid-preserved specimens of the the neural spines of the first dorsal vertebrae. following species:Tropical Kingbird (Tyrun- It passescraniolaterally to insert by a narrow nus melancholicus USNM 504538 and aponeurosison the dorsomedial surface of the 504539), Rusty-margined Flycatcher (Myio- cranial two-thirds of the scapula.The cranial- zetetescayanensis USNM 504542), Piratic most fibers insert at the very cranial tip of the

[4021 FORELIMB MYOLOGY OF KINGBIRDS 403

I SSM ’ SHCA

FIGURE 1. Superficialdorsal musclesof the forelimb (all figuresillustrate ~jwunnusmelancholicus). Abbreviations for Figs. l-l 1: ABA, M. abductor alulae;ADA, M. adductor alulae;ADM, M. abductordigiti majoris; B, M. brachialis; BB, M. biceps brachii; CCA, M. coracobrachialiscaudalis; DMACA, M. deltoideus major caudalis; DMACR, M. deltoideus major cranialis; DMI, M. deltoideus minor; EC, M. ectepicondylo-ulnaris;EDC, M. extensor digitorum communis: ELA. M. extensor longus alulae: ELDM. M. extensor longus digiti maioris: EMR, M. extensor metacarpi radialis; EMU, M. extensor metac&pi ulna& ES, M. extensor secundario&m; FCU, M. flexor carpi ulnaris; FCU a, M. flexor carpi ulnaris, accessorybelly; FDM, M. flexor digiti minoris; FDP, M. flexor digitorum profundus;FDS, M. flexor digitorum superficialis; HUP a, humero-ulnar pulley, pars accessoria;HUP ha, humero-ulnar pulley, pars humeralis accessoria;HUP u, humero-ulnar pulley pars ulnaris; ICR, M. iliotibialis cranialis; ID, M. interosseus dorsalis;IV, M. interosseusventralis; LDCA, M. latissimusdorsi caudalis;LDCR, M. latissimusdorsi cranialis; P, M. pectoralis;PPB, M. pectoralispars propatagialisbrevis; PPL, M. pectoralispars propatagialislongus; PRP, M. pronator profundus: PRS, M. pronator superficialis;RP, M. rhomboideus profundus; RS, M. rhomboideus superficialis;SBC h, M. subcoracoideusidorsal head; SBC v, M. subcoracoideus,ventral head; SBS LA, M. subscapulariscaput laterale; SBS M. M. subscauulariscauut mediale: SC. M. sternocoracoideus:SHCA, M. scapulohumeraliscaudalis; SHCR, M. scapuldhumeraliscranialis; SP, M. serratusprofundus; SP ca, M. serratusprofundus, caudal head, SP cr, M. serratus profimdus, cranial head; SSCA, M. serratussuperficialis pars caudalis;SSCR, M. serratussuperficialis pars cranialis; SSM, M. serratussuperficialis pars metapatagialis; SU, M. supinator;SUP, M. supracoracoideus;TH, M. humerotriceps; TPB, M. tensor propatagialispars brevis; TPL, M. tensor propatagialispars longa; TS, scapulotriceps;UD, M. ulnometacarpalisdorsalis; UV, M. ulnometacarpalisventralis. 404 MARY C. McKITRICK

DMI

SUP

SHCR

-TH

SSCA

FIGURE 2. Second layer of dorsal musclesof the forelimb.

scapulaand are slightly separatedfrom the rest tendinous sheet on the ventromedial surface of the belly. of the distal end of the scapula. In Myiozetetesand Legatus,this muscle in- Pars metapatagialis(SSM) is a narrow, strap- serts on the cranial three-fourths of the scap- shapedbelly, arising fleshily on the lateral sur- ula. face of the true ribs ventral to the distal end M. rhomboideusprofundus (Figs. 1,2:RP) is of the scapulaand inserting under the patagial a thin, flat, triangular muscle, lying deep to M. skin near the elbow. rhomboideus superficialis. It arises by a nar- Pars cranialis (SSCR) has lateral and medial row aponeurosisfrom the neural spinesof the heads of origin. The medial head ariseson the dorsal vertebrae, caudal to the proximal origin cervical rib, from just distal to the uncinate of M. rhomboideus superficialis; it inserts processto nearly the distal end of the rib. The fleshily on the caudal two-thirds of the scapula. lateral head arisesfrom the first true rib, from In the other species that I examined, this its uncinate processall the way to the articu- muscle is as described above, except for the lation with the sternal rib. Both bellies are es- relative length of the insertion (see McKitrick sentially strap-shaped, the lateral one being 1984). wider than the medial one. They are separated M. serratussuperficialis (Figs. 1, 2:SS) con- by a slight gap on the dorsal surface of the sistsof a caudal, a metapatagial, and a cranial origin. The two bellies extend cranidorsally be- part. Pars caudalis (SSCA) is a fan-shaped bel- tween the two heads of M. subscapularis,and ly, arisingby an aponeurosisfrom the true ribs. insert together by a tendon on the ventral mar- The caudal fibers insert fleshily on the apex of gin of the cranial end of the scapula. the scapula and the cranial fibers insert by a Whereas in Tyrannusmelancholicus the lat- FORELIMB MYOLOGY OF KINGBIRDS 405

SBS M

FIGURE 3. Left: muscleson the lateral surfaceof the scapulaand coracoid. Right: muscleson the medial surfaceof the scapulaand coracoid.

era1belly of pars cranialis completely overlies M. scapulohumeraliscaudalis (Fig. 1:SHCA) the medial belly, in Myiozetetesand Legatus, is a bipennate, fan-shaped muscle arising on the two bellies are farther apart because the the caudal two-thirds of the scapula. The su- origin of the lateral belly is limited to the region perficial fibers arise fleshily on the dorsal sur- of the uncinate process.In Tyrannus,the two face and medial border of the scapular blade, bellies fuse at the insertion and insert by a while the deeper ones arise by an aponeurosis singletendon, but, in Myiozetetesand Legatus, on the caudolateral margin of the blade. The the two bellies remain separated.The insertion muscle narrows considerably as it passescra- is, therefore, slightly broader in Myiozetetes nially; it inserts by a broad, dense tendon on and broader still in Legatus.In Empidonomus, the dorsal surface of the bicipital crest of the the lateral belly is as in Tyrannus,extending humerus, on the medial border of the pneu- to the articulation with the sternal rib. The matic fossa. bellies are nevertheless separated becausethe M. subscapularis(Fig. 3:SBS) arises by two cervical rib protrudes slightly. unipennate heads. The lateral head (SBS LA) In Empidonomusvarius the two bellies of arises fleshily on the dorsolateral, lateral, and pars cranialis are separated by a gap of about ventrolateral surfacesof the proximal end of 2 mm at the origin, but on the ventral surface the scapularblade (not near the glenoid lip as rather than the dorsal surface as in Tyrannus in Loxops[Raikow 19781). The medial head and Myiozetetes.The two bellies fuse two- (SBS M) arisesfleshily deep to this on the me- thirds of the way distally to form a common dial surface of the scapula,beginning near the belly, unlike Tyrannus,in which the two bellies acromion. The two heads fuse to insert by a do not fuse until their insertion. short tendon on the ventral tuberculum of the M. serratusprofundus (Fig. 2:SP) possesses humerus. The heads are separated by the in- three more-or-less strap-shapedheads, some- sertion of M. serratussuperficialis cranialis. times poorly defined. The caudal and middle M. subcoracoideus(Fig. 3:SBC) arisesby two ones arise from the dorsal surface of the true heads.The ventral head (SBC v) is narrow and ribs, ventromedial to the scapula,and the cra- bipennate, arising fleshily on the ventral sur- nial one arisesfrom the cervical vertebrae, lat- face of the caudal half of the coracoid. The eral to the scapula.All three insert on the me- smaller, dorsal head (SBC d) ariseson the ven- dial surface of the scapula, cranial to the tromedial surface of the head of the scapula, insertion of M. rhomboideus profundus. just ventral to its articulation with the clavicle. M. scapulohumeraliscranialis (Fig. 2:SHCR) The fibers of the two headsconverge on a stout is an essentiallyparallel-fibered muscle, slight- tendon, which is closely associated with the ly wider at its origin than at its insertion. It tendon of insertion of M. subscapularis,with arisesfleshily on the dorsolateral surfaceof the which it passesover the ventral surface of the proximal end of the scapula, and passescra- head of the coracoid to insert on the ventral nially between the ventral and dorsal heads of tuberosity of the humerus. M. humerotriceps to insert fleshily on the cra- M. coracobrachialiscranialis (CCR) is ves- nial margin of the pneumatic fossa of the hu- tigial, being reduced to ligamentous tissue. merus. M. coracobrachialis cranialis is vestigial in 406 MARY C. McKITRICK

the “corvid assemblage”(Borecky 1977). It is examined by Hudson and Lanzillotti (1955) absent in most New World nine-primaried os- and the “corvid assemblage” (Borecky 1977); tines, but well developed in Vireoand Vireo- it is fleshy (but minute) in Dendroicakirtlandii lunius(Raikow 1978). It was present in Fre- (Berger 1968) and fleshy in Fregilupusvarius gilupusvarius (Sturnidae; Berger 1957). It is (Berger 1957). reported to be “weakly developed” in Procnias M. supracorucoideus(Fig. 8:SUP) is a large, nudicollis(Cotingidae; George and Berger 1966: asymmetrically bipennate muscle, arising 315). fleshily on the dorsal fourth of the entire length M. coracobrachialiscaudalis (Fig. 8:CCA) is of the sternal carina, the craniomedial two- a bipennate muscle, lying lateral to M. supra- thirds of the sternal plate, and the dorsolateral coracoideus and arising fleshily on the lateral surfaceof the coracoclavicularmembrane. The surface of the distal half of the coracoid. The muscle narrows to a stout tendon that passes fibers insert on the dorsal surface of an apo- through the triossealcanal to insert on the dor- neurosis that arises on the ventral surface of sal surface of the deltoid crest of the humerus. the belly and gives rise to the tendon of inser- The dorsal fibers occupy about one-fourth of tion. The latter insertson the Tuberculum ven- the surface of the belly. trale of the humerus, dorsal to the insertion of M. latissimusdorsi (Fig. 1:LD) consistsof a M. subscapularis. cranial and a caudal belly in Tyrannusmel- M. sternocoracoideus(Fig. 8:SC) is a short, ancholicus.Pars cranialis (LDCR) is a narrow, broad, parallel-fibered muscle that arisesflesh- strap-shaped belly, lying superficial to M. ily from the distal ends of the sternal ribs and rhomboideus superficialis. It arises by an the sternocoracoidalprocess of the sternum. It aponeurosisfrom the neural spinesof the last inserts fleshily, and by an aponeurosisarising cervical and first dorsal vertebrae. It passes on its ventral surface, on the sternocoracoidal craniad between Mm. scapulotricepsand hu- impression of the coracoid. merotriceps, and superficial to M. latissimus M. pectoralis(Fig. 7) comprises three parts. dorsi pars caudalis, to insert fleshily on the Pars thoracicus (P) is a large, fan-shaped mus- dorsal surface of the proximal end of the hu- cle, arisingfleshily from the sternalcarina, from meral shaft. the caudolateral third of the sternal plate, the Pars caudalis (LDCA) is a strap-shapedbel- lateral trabecula of the sternum, the membrane ly, about equal in width to pars cranialis. It overlying the sternal ribs, the lateral surfaceof lies superficial to M. rhomboideus profundus the clavicle, and the coracoclavicular mem- and arises by an aponeurosisfrom the neural brane. It inserts on the Crista pectoralis (del- spinesof the dorsal vertebrae. It passescraniad toid crest) of the humerus by fleshy fibers and between Mm. scapulotricepsand humerotri- by a broad, dense, tendinous sheet arising on ceps, narrows to a tendon about 2 mm long, the dorsal surface of the belly. and inserts on the dorsal surface of the prox- Pars propatagialis longus (PPL) is a strap- imal end of the humeral shaft just cranial to shaped belly (large compared to its condition the insertion of pars cranialis. in drepanidids [Raikow 1977]- 13 mm in Ty- I examined M. latissimusdorsi for the pres- rannusmelancholicus USNM 504538), arising ence of pars caudalisin an additional 22 species fleshily on the ventral surface of pars thora- of kingbirds, as well as in 17 speciesof Tyran- cicusnear the clavicle. A narrow tendon arises nidae outside the kingbird group. I found no from its cranioventral surface and joins the instancesof asymmetry in the presenceor ab- tendon of M. tensor propatagialis longus at a senceof pars caudalis in any of the specimens right angle. dissected, although Raikow (1978) did find Pars propatagialisbrevis (PPB) is a tiny, flat asymmetry in M. latissimusdorsi in some nine- tendon, arising on the cranioventral surface of primaried oscines. pars thoracicus near the Crista pectoralis of Pars caudalis is present in the following the humerus. The tendinous origin of pars pro- tyrannid species (sample size in parentheses patagialisbrevis is about four times the width where greater than one): Pitanguslictor, Mega- of the tendon itself. Pars propatagialis brevis rynchuspitangua (2), Conopiasinornatus, C. passesdistally to form a common tendon with parva,Empidonomus varius (3) E. aurantioa- M. tensor propatagialisbrevis, at the proximal trocristatus,Tyrannus niveigularis, T. melan- end of the tendon of insertion of the latter (Fig. cholicus(7), T. voczjerans(2), T. verticalis(14), 1). T. forficatus(3), T. savana(3) T. tyrannus(3), Pars propatagialis longis is aponeurotic or T. dominicensis,T. caud&sciatus,Pyroceph- tendinous, rather than fleshy in the following: alus rubinus,Savornis phoebe, Agriornis livi- Agelaius phoeniceusand Spizella arborea da, Fluvicolapica, and Hirundineaferruginea. (George and Berger 1966) Paradisaearubra Pars caudalis is absent in the following tyr- (Berger 1956), and the 19 speciesof Corvidae annid species:Myiozetetes luteiventris, M. ca- FORELIMB MYOLOGY OF KINGBIRDS 407 yanensis(4), M. similis(15) M. granadensis, sorius of the carpometacarpus dorsal to M. Conopiastrivirgata (2) Myiodynastesluteiven- abductor allulae and just distal to the insertion tris (4) M. bairdii, Legatus leucophaius, of M. extensor metacarpi radialis. The other Camptostomaobsoletum, Elaenia flavogaster, main branch attachesbroadly on the distal end Mionectesoleagineus, Phaeomyias murina, of the radius, and on the joint capsule, insert- Lophotriccusgaleatus, Tolmomyiassulphu- ing on the OS radiale. rescens,Platyrinchus coronatus, Contopus vi- Pars brevis (TPB) is a spindle-shapedbelly, rens,Empidonax virescens, Machetornis rix- lying superficial to Mm. deltoideus major osus,Attila spadiceus,Myiarchus crinitus (2). cranialis and deltoideus major caudalis from Pars caudalisis variable in the following tyr- the dorsal surface of the upper arm. It arises annid species(+ denotes presence, - denotes by a short tendon on the dorsal surface of the absence): Pitangussulphuratus (6 +, 1 -), apex of the clavicle, passesdistally about half- Myiodynastesmaculatus (3 +, 2 -). way down the humerus, and gives rise to a Pars caudalis is present in Paradisaearubra tendon. This tendon is joined by the tendon (Paradisaeidae;Berger 1956); Sturnusvulgaris of insertion of M. pectoralis pars propatagialis and Aplonistabuensis (Sturnidae; Berger 1957); longus. It anchors on the proximal end of M. 19 speciesin 14 genera of Corvidae (Hudson extensor metacarpi radialis about 6 mm distal and Lanzillotti 195 5); most of the “corvid as- to the humeral origin of the latter. The tendon semblage” (Borecky 1977); ApalisJlavida,Or- then passesalong the surface of M. extensor thotomusatrogularis (Sylviinae), Turdusmi- metacarpi radialis and inserts on the dorsal gratorius, Catharus ustulatus (Turdinae), epicondyle of the humerus. It has its own, dis- Dumetellacarolinensis (Mimidae), Nectarinia tinct insertion, separate from that of M. ex- sperata(Nectarinidae), four genera of Vireo- tensor metacarpi radialis, as do all other pas- nidae, Peucedramustaeniatus (incertae sedis), serines studied (Raikow 1982). Thraupisepiscopus (Thraupinae), and Cacicus There is a scapular tendon as described for cela (Icterinae; Raikow 1978); Vidua and all drepanidids by Raikow (1977). Ploceinae, Bubalornithinae, Lonchurae, and M. deltoideus major cranialis (Fig. 1: Poephilae examined by Bentz (1979). The DMACR) is a large, fan-shaped muscle, situ- muscle is absent in Streperagraculina (Crac- ated cranial to M. deltoideus major caudalis. ticidae; Leach 19 14, confirmed by Hudson and It arises fleshily on the OS humeroscapulare Lanzillotti 1955); Fregilupusvarius (Sturni- and surroundingligaments. The muscle inserts dae) and Artamella (Vangidae; Berger 1957); by fleshy fibers on the caudal half of the dorsal Agelaiusphoeniceus (Icterinae; George and surface of the Crista pectoralis, and by tendi- Berger 1966); Dendroicakirtlandii (Parulinae; nous fibers on the proximal one-half of the Berger 1968); Lamprotornis,Cinnyricinclus, humeral shaft (i.e., the entire insertion extends Speculipastor,Neocichla and Saroglossa(Stur- two-thirds the length of the humerus). nidae), Dendrocittaand Pseudopodoces(incer- In Legatusleucophaius the entire belly ex- tae sedis),and the Cracticidae (Borecky 1977); tends lessthan half the length of the humerus, the majority of the New World nine-primaried or about one-third of the length of the shaft. oscines(Raikow 1978); numerous Estrildidae In the other speciesthat I examined, this mus- and three speciesof Passerinae(Bentz 1979). cle is asdescribed for Tyrannusmelancholicus. M. latissimus dorsi caudalis is widespread George and Berger (1966) reported that M. in non-passerines (George and Berger 1966: deltoideus major cranialis is composed of two 288-294). parts in Agelaiusphoeniceus, one arising from M. tensorpropatagialis (Figs. 1, 7) consists the OS humeroscapulare as in the tyrannids, of two separate bellies. Pars longa (TPL) is a the other from the scapula. This is also the small, triangular belly that arises by a short casein corvids (Hudson and Lanzillotti 1955) tendon from the tip of the clavicle. It gives rise Fregilupusvarius, Sturnus vulgaris,Aplonis to a long tendon (tendo longa) which forms the tabuensisand Artamellaviridis, (Berger 1957); cranial border of the patagium. The tendon is Dendroicakirtlandii (Berger 1968) and the joined at its proximal end by the tendons of “corvid assemblage”(Borecky 1977). The condition in Procniasnudicollis is apparently like Mm. cucullaris capitis pars propatagialis and that in tyrannids (George and Berger 1966) as pectoralis pars propatagialis longus. is that in most passerinesfor which data exist Tendo longa divides into two main branch- (seeRaikow 1978). In Pityriasisgymnocephala es. One branch splits into three parts, of which (Laniidae), Mm. deltoideusmajor cranialis and one inserts on the craniodistal surface of digit deltoideus major caudalis are fused for most I, one at the base of the follicle for the remex of their length (Raikow et al. 1980). on the dorsal surface of digit I, and one on the M. deltoideusmajor caudalis(Fig. l:DMA- craniodorsal surface of the Processusexten- CA) is a long, strap-shapedmuscle that arises 408 MARY C. McKITRICK

EDC

ELDM

EMU

FIGURE 4. Third layer of dorsal musclesof the forelimb.

fleshily from the medial surface of the proxi- kow 1978). Bentz (1979) reported a coracoidal mal end of the clavicle. It extends down the head for 13 speciesof Estrildidae. dorsal surface of the upper arm, superficial to M. triceps brachii comprises Mm. scapulo- M. scapulotricepsand caudal to M. deltoideus triceps and humerotriceps. M. scapulotriceps major cranialis. It becomes tendinous in its (Fig. l:TS) has one head of origin in Tyrannus distal fourth, and the narrow tendon insertson melancholicus,arising by a short tendon from the dorsal surface of the distal end of the hu- the dorsocaudalborder of the Faciesarticularis merus, just proximal to the Processussupra- humeralis of the scapulaand on the surround- condylaris dorsalis. A narrow scapularanchor ing ligaments. The strap-shapedbelly proceeds arisesfrom the deep portion of the medial mar- distally to become tendinousin its distal fourth; gin and inserts on the dorsomedial surface of the tendon contains a sesamoid between the the scapula cranial to the insertion of M. la- humerus and ulna. Here, a short tendon (hu- tissimus dorsi. meral anchor) branches off the main tendon I did not detect a scapular anchor in Myio- and passesto the medial surfaceof the humeral zetetescayanensis. shaft just proximal to the dorsal condyle. The M. deltoideusminor (Fig. 8:DMI) is a small, main tendon inserts on the caudolateral sur- strap-shapedmuscle, lying deep to M. tensor face of the Processuscotylaris dorsalis of the propatagialis brevis on the dorsal surface of ulna. the shoulder. It arises fleshily from the lateral M. scapulotriceps arises by two scapular surface of the acromion of the scapula and heads in Fregilupus varius (Berger 1957), Age- inserts by semitendinous fibers on the head of laius phoeniceus (George and Berger 1966), the humerus, just cranial to the insertion of Dendroica kirtlandii (Berger 1968) and nine M. supracoracoideus. speciesof Drepanididae (Raikow 1977). Only In many passerines,M. deltoideusminor has one head is present in corvids (Hudson and only one, scapularhead of origin. George and Lanzillottii 1955) and Paradisaea rubra (Ber- Berger (1966) noted an additional, coracoidal ger 1956). head in Agelaius phoeniceus.This additional The humeral anchor apparently hasnot been head occurs in the Ptilonorhynchidae, Par- reported for other passerines, although it is adisaea (Paradisaeidae), and Corcorax (Gral- present in a number of non-passerines(Bed- linidae; Borecky 1977) as well as “universal- dard 1898, George and Berger 1966). ly” in the Carduelinae and Drepanididae; in A4. humerotriceps(Figs. 1, 2, 4, 5, 8, 9:TH) most Emberizinae, Icteria and Seiurus (Paru- consists of a ventral and a dorsal head, that linae) and Rhodinocichla (Thraupinae; Rai- arise fleshily from the ventral and dorsal bor- FORELIMB MYOLOGY OF KINGBIRDS 409

ELDM

FIGURE 5. Fourth layer of dorsal musclesof the forelimb. ders, respectively, of the pneumatic fossa of M. expansor secundariorum (Figs. 10, 11: the humerus. The dorsal fibers of the ventral ES) is a small, triangular muscle that arises belly arise from within the pneumatic canal. fleshily from the ventral surfaceof the humero- The two fuse to form a single, bipennate belly ulnar pulley. The belly passessuperficial to all about half-way down the humeral shaft, and the ventral muscles of the forearm and fans insert by tendinous fibers on the olecranon of out to insert at the base of the proximal three the ulna. Some fibers insert on the tendon of secondary remiges. insertion of M. scapulotriceps,proximal to the M. brachialis (Fig. 10:B) is a short, parallel- sesamoid. The two heads are separated at the fibered, rectangular muscle, arising fleshily on origin by the insertion on the humerus of M. the ventral surface of the Fossa M. brachialis scapulohumeraliscranialis. of the distal end of the humerus, and passing M. bicepsbrachii (Fig. 8:BB) is a large, spin- acrossthe joint to insert fleshily on the prox- dle-shaped muscle, arising by two heads. One, imoventral surfaceof the ulna between the two caput coracoideum, arises by a stout tendon heads of origin of M. flexor digitorum profun- on the ventromedial surface of the proximal dus. end of the head of the coracoid and by elastic M. pronator superficialis (Fig. 9:PRS) is a fibers from the humerocoracoidalligament; the spindle-shapedmuscle that arisesby a tendon other, caput humerale, arisesby a short tendon on the ventral surface of the distal end of the from the ventral border of the peneumatic fos- humerus, distal and ventral to the origin of M. sa of the humerus. It inserts by three tendons: brachialis. The tendon of origin passesdeep one on the proximocranial end of the radius within the belly, and the ventral fibers of the (Tuberculum bicipitale of the radius), and two belly originate along it, about 3 mm distal to side by side on the Tuberculum bicipitale of the origin of the dorsal fibers. The insertion is the ulna. on the ventral surfaceof the shaft of the radius, The insertion of M. biceps brachii is by two by a tendinous sheet arising along the distal tendons in most birds, one on the radius and two-thirds of the length of the belly. a larger one on the ulna, rather than by three. M. pronator profundus (Fig. 9:PRP) is a Swinebroad (1954) reported only one tendon spindle-shapedmuscle on the ventral surface of insertion, on the ulna, in Passerdomesticus, of the forearm between Mm. extensor meta- Cardinalis cardinalis, Zonotrichia albicollis carpi radialis and flexor digitorum superfici- and Melospiza melodia; however Bentz (1979) alis, deep to M. pronator superficialis.It arises who also dissected Passer domesticus,made by a tendon from the ventral epicondyle of the no reference to such a condition. humerus between the origins of Mm. pronator 410 MARY C. McKITRICK

FIGURE 6. Deep dorsal musclesof the forelimb. superficialisand flexor digitorum superficialis. humero-ulnar pulley deep to M. expansor se- It inserts by fleshy fibers, and, at its proximal cundariorum and givesrise to a spindle-shaped and distal ends, by tendinous ones, on the me- belly. This belly extends half-way down the dial surfaceof the radius, immediately deep to forearm, and inserts by a stout tendon on the the insertion of M. pronator superficialis. The OS ulnare. The humerocarpal band, which in- belly has the appearance of being bipennate, vests the belly of M. flexor carpi ulnaris and with two sets of fibers converging on an apo- the tendon of insertion of Mm. flexor carpi neurosisthat covers part of the ventral surface ulnaris and flexor digitorum superficialis,sends of the belly. tendinous branchesto the base of the six outer On the right side of Tyrannus melancholicus secondaries(see description for M. flexor dig- CM 345, M. pronator superficialis was at- itorum superficialis). tached, at the distal end of its belly, to the distal A smaller, accessory belly (FCU a) arises end of the belly of M. pronator profundus. In from the caudal surface of the proximal end addition, it had a long tendon of insertion that of the main belly, and passesone-third of the attached near the distal end of the forearm to way down the forearm, giving rise to a slender the tendon of M. flexor digitorum profundus. accessorytendon. This tendon is attached to The distal end of M. pronator profundus was the base of the six outer secondaries,not by also attached to this evidently extra tendon. simple, slender, tendinous bands as in Loxops This condition seemsanomalous, as I did not (Raikow 1977), but broadly by a fibrous tissue, observe anything like it on the left side or in either muscular or elastic in composition. other specimens. The humero-ulnar pulley (trochlea humer- In many passerines,M. pronator profundus oulnaris of Baumel et al. [ 19791)(Fig. 11) arises originatesfrom the humero-ulnar pulley aswell by a narrow strap from the caudal surface of as from the humerus. This is true of corvids the ventral epicondyle of the humerus (pars (Hudson and Lanzillotti 195 5), Agelaiusphoe- humeralis accessoria).Two narrow strapsarise niceus (George and Berger 1966) Dendroica on the flexor processof the humerus, one (not kirtlandii (Berger 1968) the “corvid assem- describedby Bentz and Zusi 1982) passingdeep blage” (Borecky 1977) most New World nine- to M. flexor carpi ulnar-isand the other (pars primaried oscines (Raikow 1978), and 13 humeralis), more delicate strap passingsuper- speciesof Estrildidae (Bentz 1979). ficial to it. Both attach on pars humeralis near M. jlexor carpi ulnaris (Fig. 9:FCU) is the its junction with pars ulnaris. Pars ulnaris is a most caudal muscle on the ventral surface of broad band that arisesfrom the ventral surface the forearm. It arises by a tendon from the of the proximal end of the ulna and attaches cranioventral surface of the flexor process of on the dorsal surface of the pulley. the humerus. The tendon passesthrough the A slender tendon, arising on the ulna prox- FORELIMB MYOLOGY OF KINGBIRDS 411

TPL

FIGURE 7. Superficial ventral musclesof the forelimb. imal to the ulnar anchor of M. extensor me- envelops the belly and tendon of insertion of tacarpi radialis, inserts on the deep surface of M. flexor digitorum superficialisand the main the humero-ulnar pulley (ligamentous sling of belly of M. flexor carpi ulnar-is.With the ten- Bentz and Zusi [1982]). dons of those muscles,it passesacross the OS M. j7exordigitorum superficialis (Fig. 9:FDS) ulnare, whence it sendstendinous branchesto is a spindle-shapedmuscle that arisesby a stout the base of the alula, to the bases of the first tendon from the caudoventral surface of the three primary remiges, to the distal end of the flexor process of the humerus, distal to the radius, and to ensheathe the tendon of inser- origin of M. pronator profundus.It passesabout tion of Mm. flexor digitorum profundus and one-third of the way down the forearm before ulnometacarpalis ventralis at the point where giving rise to the tendon of insertion. This ten- these enter the manus. don is closely associatedproximally with that This muscle is essentially the same in the of M. flexor carpi ulnaris via the humerocarpal other speciesthat I examined, except that in band (described below). The tendon passes Legatusand Empidonomus,the belly extends around the OS ulnare and acrossthe manus to only about one-fourth of the way down the insert on the ventrodistal surface of the first forearm. phalanx of digit II. M. flexor digitorumprofindus (Fig. 9:FDP) The humerocarpal band ariseson the medial is a spindle-shaped muscle along the ventral epicondyle of the humerus as part of the origin surface of the ulna caudal to M. pronator pro- of M. flexor digitorum superficialis.It partially fundus and superficial to M. ulnometacarpalis 412 MARY C. McKITRICK

DMI

FIGURE 8. Second layer of ventral musclesof the forelimb. ventralis. It arises fleshily by two heads from belly arisesby two distinct tendons on the Pro- the proximoventral surface of the ulna, the cessussupracondylaris dorsalis, one (Tendon dorsal head arising just distal to the ulnar in- 1) from the dorsal surface and one (Tendon 2) sertion of M. biceps brachii. M. brachialis in- from the caudal surface. The two tendons fuse sertsbetween the two heads, and distal to this a few millimeters distad to form a common point, the two heads fuse. The single belly belly. Pars intermedius arises by a tendon on passeshalf-way down the forearm and gives the dorsocaudal surface of the Processussu- rise to a tendon. The tendon passesaround the pracondylaris dorsalis, distal to the origin of Processusintermetacarpalis, deep to a liga- Tendon 1 (see above), and by fleshy fibers in ment connecting the distal end of the radius the intercondylar region between pars ancon- and the Processes intermetacarpalis, then alis and pars palmaris (i.e., just ventral to the acrossthe manus to insert on the proximal end Processussupracondylaris dorsalis). Pars pal- of the second phalanx of digit II. mark arises by fleshy and tendinous fibers in M. extensormetacarpi radialis (Fig. 9:EMR) the intercondylar region just ventral to pars is the most superficial muscle on the dorsal intermedius. surface of the forearm. It is a large, bipennate Pars intermedius fuses with pars palmaris muscle with four points of origin. The anconal about one-fourth of the way down the forearm, FORELIMB MYOLOGY OF KINGBIRDS 413

PRP

FIGURE 9. Third layer of ventral musclesof the forelimb. but the point of attachment is narrow, and pars and into the manus, bifurcating at digit I. The palmaris continues distad as a unipennate bel- smaller branch inserts on the caudodorsalsur- ly. Pars palmaris and anconalis fuse three- face of the base of digit I; the main branch fourths of the way down the forearm and give continues distad within the Sulcus tendineus rise to the common tendon of insertion. This of the carpometacarpus,passes over the distal tendon inserts on the Processusextensorius of end of the carpometacarpuswith a sesamoid, the carpometacarpus. and inserts on the dorsal surface of phalanx 1 This condition is similar to that in Loxops of digit II. virens(although not noted as such by Raikow The aponeurosis antebrachialis dorsalis [ 1977]), with some exceptions. The interme- ariseson the dorsal epicondyle of the humerus, diate head is considerably smaller in Loxops in common with M. extensor digitorum com- (pers. observ.). In Loxops,pars anconalis and munis. It attaches along the distal two-thirds pars palmaris are about equal in size at their of the shaft of the radius, and ensheathesthe origin, with pars intermedius barely visible in bellies of Mm. extensor digitorum communis between. In Tyrannusmelancholicus the fleshy and extensor metacarpi ulnaris. The aponeu- origins of pars intermedius and of pars pal- rosis attaches to the bases of the follicles for maris are about equally large and pars ancon- the first eight secondary remiges, and then to alis is relatively slender. Also, in that species, the dorsal surface of the distal end of the ra- pars anconalis has no fleshy origin on the Pro- dius. cessussupracondylaris dorsalis, only a dorsal M. extensormetacarpi ulnaris (Figs. 4, 5: tendinous one, while in Loxops,there are both EMU) is a bipennate muscle that arises by a a fleshy and a dorsal tendinous origin. tendon from the dorsal surface of the distal AL extensordigitorum communis (Figs. 4, end of the humerus just cranial to the dorsal 5:EDC) is a spindle-shapedmuscle, caudal to epicondyle, and by a broader tendon (the ulnar M. extensor metacarpi radialis on the dorsal anchor) from the craniodorsal surface of the surface of the forearm. It arises by a tendon proximal end of the ulnar shaft. Arising just from the dorsal epicondyle of the humerus and proximal to the ulnar anchor is a slender ten- the belly extends nearly half-way down the don that passesdown to insert on the deep forearm before giving rise to the tendon of surface of the humero-ulnar pulley, the “lig- insertion. This tendon passesaround the ulna amentous sling” of Bentz and Zusi (1982). It 414 MARY C. McKITRICK

ELA

FIGURE 10. Deep layer of ventral musclesof the forelimb.

doesnot contain a sesamoidin thesetyrannids, the manus to insert on the craniodorsal border as it does in hummingbirds. of the base of phalanx 2, digit II. The single belly narrows to a tendon about M. supinator(Fig. 10:SU) is a narrow, fan- two-thirds of the way down the shaft, and shapedmuscle, arising by a slendertendon from passesaround the Tuberculum ventrale to in- the dorsal epicondyle of the humerus. It passes sert on the carpometacarpusat the base of the about one-fourth of the way down the radius minor metacarpal. and inserts fleshily on the craniodorsal surface M. extensor longusalulae (Figs. 6, 10:ELA) of that bone. is a tiny muscle, arising fleshily on the dor- M. ectepicondylo-ulnaris(Figs. 6, 10:EC) is solateral surface of the ulna medial to M. ex- a fan-shaped muscle that arises by a tendon tensor longus digiti majoris. Half-way down from the craniodorsal surface of the dorsal the forearm, it gives rise to a hair-like tendon condyle of the humerus. It passesacross the that passesdistally along the ulna deep to the elbow joint, giving rise to a long, narrow belly tendon of insertion of M. extensor metacarpi that extends about half-way down the cra- radialis, and inserts lateral to that tendon on nioventral surface of the ulna and inserts the Processusextensor% of the carpometa- fleshily on that bone. carpus. M. ulnometacarpalisdorsalis (Fig. 4:UD) is The inserting tendon of this muscle fuses a small, fan-shaped muscle, arising by a nar- with that of M. extensor metacarpi radialis for row tendon on the ligament ensheathing the a common insertion on the carpometacarpus distodorsal end of the ulna. It inserts fleshily in Fregilupusvarius and Sturnusvulgaris (Ber- on the proximocaudal surface of the minor ger 1957) and Agelaiusphoeniceus(George and metacarpal. Berger 1966). In other passerinesreported in The muscle is absent in Spizella arborea the literature, the two tendons are separatebut (George and Berger 1966) and Dendroicakirt- closely associated,as in the tyrannids. landii(Berger 1968). George and Berger (1966) M. extensorIongus digiti majoris(Figs. 4, 5, reported it to be present as a ligament in Age- 9, 10:ELDM) is a narrow, bipennate muscle laius phoeniceus,but Berger later (1969) that arises fleshily on the caudodorsal surface claimed it to be absent in Agelaius.He may of the proximal two-thirds of the shaft of the have been referring to a functional rather than radius. It narrows to a tendon about three- an actual absence. He did not mention the fourths of the way down the shaft. The tendon muscle for Paradisaearubra (Berger 1956) or passesaround the distal end of the ulna into Fregilupusvarius (Berger 1957); in other lit- FORELIMB MYOLOGY OF KINGBIRDS 415

position, as described for Loxopsby Raikow 1977). M. abductoralulae (Figs. 9, 10:ABA) is a small, spindle-shapedmuscle, arising fleshily on the ventral surface of the distal end of the inserting tendon of M. extensor metacarpi radialis. It passesventral to the Processusexten- sorius of the carpometacarpus,giving rise to a fine tendon that inserts on the caudoventral surface of digit I. M. adductoralulae (Figs. 4, 5, 9, 10:ADA) is a narrow, spindle-shapedmuscle that arises fleshily at the baseof the Processusextensorius of the carpometacarpus. It inserts fleshily on ES the caudal surfaceof digit I and on the inserting FIGURE 11. M. expansorsecundariorum and humero- tendon of M. extensor digitorum communis. ulnar pulley. M. abductordigiti majoris (Figs. 9, 1O:ADM) is a tiny, fan-shaped muscle, arising fleshily on erature on passerines,the muscle is reported the cranioventral surface of the alular meta- as present, but always small. carpal. It extends the length of the cranial sur- M. ulnometacarpalisventralis (Figs. 9, 10: face of the carpometacarpus,and inserts by a UV) is an elongate, fan-shaped muscle along fine tendon on the base of the first phalanx of the ventral surfaceof the ulna deep to M. flexor digit II. digitorum profundus. It arises fleshily on the M. abductor digiti majoris was reported to cranioventral surface of the proximal end of be “vestigial or absentin many passerinebirds ” the ulna, and on the caudoventral surface of (George and Berger 1966). It is absent in Age- the ulna about one-third of the way down the laius phoeniceus(ibid), Passer domesticus, shaft. This “double” origin gives the muscle Cardinalis cardinalis, Zonotrichia albicollis the appearanceof having two heads. The com- and Melospizamelodia (Swinebroad 1954) and mon belly extends to two-thirds of the way Dendroicakirtlandii (Berger 1968). It is “small down the ulnar shaft, and its tendon of inser- and inconspicuous” in corvids (Hudson and tion arises as an aponeurosis on the ventral Lanzillotti 1955) and rudimentary in Para- surface of the belly. The tendon passesover disaea rubra (Berger 1956). Borecky (1977) the ventral condyle of the ulna and with a ses- found the muscle in the “corvid assemblage,” amoid over the OS radiale. It inserts near the and Raikow (1977, 1978) found it in many base of the Processusextensorius of the car- oscines. Its presence may have been over- pometacarpus. looked in some speciesbecause it is difficult M. ulnometacarpalis ventralis has a proxi- to see without staining. mal and a ventral head of origin in Dendroica M. flexor digiti minoris(Figs. 9, 10:FDM) kirtlandii (Berger 1968) which is presumably is a long, narrow, bipennate muscle that arises an exaggeration of the condition described fleshily on the caudal surface of the minor above. metacarpal, and by a tendon on the distal end M. interosseusdorsalis (Figs. 4, 9:ID) is a of the ulna. It extends the length of the car- tiny, bipennate muscle, arising fleshily on the pometacarpus and inserts by a fine tendon on proximal margin of the interossealspace of the the caudal surface of digit III. carpometacarpusand the surface of the alular In Agelaiusphoeniceus, this muscle has a and minor metacarpal. It is about one-third of tiny, deep head as well (George and Berger the length of the adjacent M. interosseusven- 1966). tralis. It narrows to a fine tendon that inserts on the dorsoproximal surface of the second DISCUSSION phalanx of digit II. My investigation confirms, for the group that M. interosseusventralis (Figs. 4, 5, 9, 10:IV) I studied, that the suboscine forelimb mus- is a tiny, bipennate muscle that arises fleshily culature conforms to the passeriform pattern on the proximal margin of the interossealspace reported by other authors for the oscines.Dif- and the margins of the alular, major, and mi- ferencesamong speciesin the forelimb muscles nor metacarpals. It narrows to a fine tendon appear to occur primarily in M. latissimusdor- that passesover the dorsal surface of phalanx si caudalis. My survey of this muscle in pas- 1 of digit II, and inserts half-way down the serines and nonpasserines suggeststhat its caudodorsal margin of the second phalanx of presence is the primitive condition in birds, digit II (it has the anchor of unknown com- and that it has been lost numerous times, es- 416 MARY C. McKITRICK pecially within the Passeriformes.It has been (National Museum of Natural History). Illustrations were lost repeatedly within the Tyrannidae as well. preparedby J. Anderton. The manuscript was greatly im- proved by the comments of W. E. Lanyon and R. L. Zusi. In the kingbird group, most species possess This material is based upon work supported by the Na- pars caudalis, but all members of Myiozetetes tional Science Foundation under Grants DEB-8010898 that I examined invariably lacked the muscle and BSR-83 14729 to R. J. Raikow. (15 specimensof similis,four cayanensis,one granadensis,one luteiventris),as did Legatus LITERATURE CITED leucophaius(one specimen dissected), Conop- AMES, P. L. 1971. The morphology of the syrinx in pas- ias trivirgata(two dissected),and Myiodynas- serine birds. Bull. Peabodv Mus. Nat. Hist. No. 37. tes luteiventris(four dissected). Of seven Pi- BAUMEL,J. J., A. S. KING, A. M. LUCAS,J. E. BREAZILE, ANDH. E. EVANS.[EDS.]. 1979. Nomina anatomica tangussulphuratus, one specimen lacked it, as avium. Academic Press, London. did two out of five specimensof Myiodynastes BEDDARD,F. E. 1898. The structure and classification maculatus.Many tyrannids outside of the of birds. Longmans, Green and Co., New York. kingbird group lack the muscle. The speciesof BENTZ,G. D. 1979. The appendicularmyology and phylogenetic relationshipsof the Ploceidae and Estrildi- Myiozetetesmay indeed constitute a clade; dae (Aves: Passeriformes).Bull. Carnegie Mus. Nat. however, it would be unsatisfying to cluster Hist. No. 15. them with the other speciesthat lack M. latis- BENTZ,G. D., AND R. L. Zusr. 1982. The humeroulnar simus dorsi caudalis. This muscle is clearly pulley and its evolution in hummingbirds. Wilson prone to frequent, and probably independent, Bull. 94~71-73. BERGER,A. J. 1956. On the anatomy of the Red Bird of lossand, therefore, an unreliable basisfor phy- Paradise,with comparative remarks on the Corvidae. logenetic hypotheses.The individual variation Auk 73~427-446. in the occurrence of M. latissimus dorsi cau- BERGER,A. J. 1957. On the anatomy and relationships dalis may representa polymorphism in species of Fregilupus varius, an extinct starlingfrom the Mas- carene Islands. Bull. Am. Mus. Nat. Hist. 113:225- undergoing loss of the muscle, an idea that I 272. have explored elsewhere (McKitrick 1984). BERGER,A. J. 1968. Appendicular myology of Kirtland’s Several features of the forelimb musculature Warbler. Auk 85:594-616. of the kingbird group differ from those in other BERGER,A. J. 1969. Appendicular myology of passerine birds; these are the occurrence of a humeral birds. Wilson Bull.81:220-223. _ -_ _ BOCK.W. J.. ANDC. R. SHEAR. 1972. A staining method anchor in M. scapulotriceps,three (rather than for grossdissection of vertebrate muscle. ALat. Anz. two) tendons of insertion of M. biceps brachii, 130:222-227. and the condition of the humero-ulnar pulley BORECKY,S. R. 1977. The appendicular myology and and of M. extensor metacarpi radialis. The hu- phylogenetic relationships of the avian “corvid assemblage.” Unpubl. Ph.D. diss., University of Pitts- meral anchor occursin non-passerinesand may burgh, Pittsburgh, PA. be primitive for tyrannids, but the other fea- GARROD,A. H. 1876. On some anatomical characters tures are probably derived at some level. Fur- which bear upon the major divisions of the passerine thermore, the occurrenceof a ligamentoussling birds. Part I. Proc. Zool. Sot. Lond. 1876:506-519. between the ulnar shaft and the humero-ulnar GEORGE,J. C., AND A. J. BERGER.1966. Avian myology. Academic Press, New York. pulley is apparently limited in its distribution, HUDSON,G. E., AND P. J. LANZILLOTTI. 1955. Gross having been reported so far only in suboscines, anatomy of the wing musclesin the family Corvidae. Apodidae, and Trochilidae (Bentz and Zusi Am. Midl. Nat. 53:i-44. 1982). Future studies of tyrannids and other LANYON.W. E. 1984. A vhvloeenv of the kinebirds and their allies. Am. Mus.~Nbvi?.No. 2797. - suboscinesshould pay particular attention to LEACH,J. A. 1914. The myology of the Bell Magpie these character statesto determine their utility (Strepera) and its position in classification.Emu 14: for delineating clades within the suboscines. 2-38. As I found virtually no variation in most of MCIOTRICK,M. C. 1984. The appendicular myology of the forelimb musclesamong the speciesthat I the kingbirds and their allies (Tyrannidae) and its evolutionary significance. Ph.D. dissertation, Uni- examined, however, these muscles appear to versity of Pittsburgh, Pittsburgh, PA. be of little use for clarifying relationshipswith- MCI(ITRICK,M. C. 1985. Monophyly of the Tyrannidae in the kingbird assemblage. (Aves): comoarison of moroholoav__ and DNA. Svst. Zool. 34:35-45. ACKNOWLEDGMENTS MCKITRICK.M. C. In Dress. The velvic mvoloav of the kingbirds and their allies (A&: Tyrannidae). Ann. For continual advice and support,I thank M. Kreithen, Carnegie Mus. 54. K. C. Parkes,J. H. Schwartz,W. A. Searcy,and, especially, MIDTGARD,U. 1982. Patterns of the blood vascularsys- R. J. Raikow, who criticized an earlier draft of this paper. tem in the pelvic limb of birds. J. Zool. (Land.) 196: The following people generouslylent specimens for his- 545-567. - section:G. F. Barrowcloughand W. E. Lanyon (American RAIKOW,R. J. 1977. Pectoral appendagemyology of the Museum of Natural History); K. C. Parkes, D. S. Wood, Hawaiian honeycreepers(Drepanididae). Auk 94:33l- and J. M. Loughlin (CarnegieMuseum of Natural History); 342. J. W. Fitzpatrick (Field Museum of Natural History); M. RAIKOW,R. J. 1978. Appendicular myology and rela- A. Jenkinson (University of Kansas); J. V. Remsen, Jr. tionships of the New World nine-primaried oscines (Louisiana State University); C. G. Sibley and E. Stickney (Aves: Passeriformes).Bull. CarnegieMus. Nat. Hist. (Peabody Museum of Natural History); and R. L. Zusi No. 7. FORELIMB MYOLOGY OF KINGBIRDS 417

RAIKOW, R. J. 1982. Monophyly of the Passeriformes: ZUSI, R. L., AND G. D. BENTZ. 1984. Myology of the test of a phylogenetichypothesis. Auk 99:43 l-445. Purple-throatedCarib (Eulampisjugularis) and other RAIKOW,R. J., P. J. POLUMBO,AND S. R. BORECKY.1980. hummingbirds (Aves: Trochilidae). Smithson. Con- Appendicularmyology and relationshipsof the shrikes trib. Zool. No. 385. (Aves: Passeriformes:Laniidae). Ann. CarnegieMus. 49:131-152. Department of Biological Sciences, University of Pitts- SWINEBROAD,J. 1954. A comparative study of the wing burgh, Pittsburgh,Pennsylvania 15260. Present address: myology of certain passerines.Am. Midl. Nat. 51: Departmentof Ornithology,American Museum of Natural 488-514. History, Central Park Westat 79th Street,New York, New TRAYLOR,M. A., JR. 1977. A classificationof the tyrant York 10024. Received 22 October 1984. Final acceptance flycatchers(Tyrannidae). Bull. Mus. Comp. Zool. 148: 27 March 1985. 129-184.

The-Condor 87:417 0 The Cooper Ornithological Society 1985

RECENT PUBLICATIONS

International registry of poultry genetic stocks/A direc- colonies in the north Atlantic and western Arctic waters. tory of specializedlines and strains, mutations,breeds and Commendably, the book managesto report a great deal varietiesof chickens,Japanese quail and turkeys.-Ralph of accurate and detailed information, yet in a readable, G. Somes, Jr. [ 19851Bulletin 469, Storrs Agricultural Ex- non-technical style. Various additional data are given in periment Station, The University of Connecticut, Storrs. 20 tables at the back. The volume is well illustrated with 96 p. Paper cover. $3.00. Source:Storrs Agricultural Ex- maps, diagrams,monochrome photographs,and drawings periment Station, Storrs, CT. This is the fifth edition of a by Keith Brockie. List of references,index. Harris is cor- reference for poultry fanciers and researcherswho use rect in saying that his book complements rather than poultry in their work. It is organized like its predecessor supersedesLockley ’s. Read the new book for up-to-date (noted in Condor 84:202), but has again been enlargedso facts and analysis,and the old one for a vivid, admittedly as to include stocksfrom more suppliers.Incidentally, the personal, account of life in a puffin colony. registry demonstratesthe major roles played by domes- ticated birds in studies of avian genetics. Ecogeographicalvariation in size and proportionsof Song Bobwhite thesaurus.-Thomas G. Scott. 1985. Intema- Sparrows(Melospiza melodia).-John W. Aldrich. 1984. tional Quail Foundation. 306 p. $29.95. Source: I.Q.F., Ornithological Monographs No. 35, American Omithol- P.O. Box 550, 206 Buncombe St., Edgefield, SC 29824. ogists’union, Washington,DC. 134 p. Papercover. $10.50 This is an indexed bibliography of Colinus virginianus, ($8.50 to AOU members). Source:Frank R. Moore, De- including more than 3,000 publications that were issued partment of Biology, University of Southern Mississippi, before 1983. The index is unusually thorough for works Southern Station Box 5018, Hattiesburg, MS 39406; all of this kind, as it comprises over 100 principal subject- ordersmust be prepaidand includea $0.50 handlingcharge. categoriesand up to three levels of subordinatecategories. In this monograph, Aldrich examines patterns of mor- Even the .page - referencesfor each specific piece of infor- phological variation in a species with a wide breeding mation are given. The book is nice&made and has a color rangein order to learn their probableadaptive significance. frontispiece by wildlife painter Bob Carey. It will be a Measurements and proportions of the wing, bill, tarsus, useful tool for quail biologistsand managers. tail, and middle toe are correlated with certain ecological categories(Life Areas and Ecoregions)of North America. Multitudes of data are reported and analyzed.The oatterns The Puffin.-M. P. Harris. 1984. T&AD Poyser, Carlton, are, for the most part, clear and irrefutable, but the adap- England. 224 v. $32.50. Thirtv vears after Locklev’s Pufi tive reasonsbehind them, while plausible, remain to be fins, here is a new book aboui &ratercula arctica,.one of established.Aldrich concludesthat “the ecologicalforces the most generally familiar and widely appealing of sea- selectingadaptive genetic differenceshave a greater effect birds. It offers comprehensive treatment of the species’ on morphological changeor microevolution than do geo- distribution, behavior, breeding and feeding biology, nat- graphicalseparation or isolation.” His rich findings should ural and man-caused threats to survival, and population stimulate further researchinto the ways by which envi- dynamics. Material has been drawn from the author’s own ronmental factors influence the evolution of avian taxa. studiesand those of other ornithologistsat several island Maps, graphs, list of references,appendices.