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The Relationships of the Hoatzin

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The Relationships of the Hoatzin THE AUK A QUARTERLY JOURNAL OF ORNITHOLOGY Vo•,. 90 J^NU^R¾1973 NO. 1 THE RELATIONSHIPS OF THE HOATZIN CH^R•,ESG. Sm•,E¾^N•) JON E. AHLQmST THE taxonomicposition of the Hoatzin (Opisthocomushoazin) of South Americahas long been one of the mostdebated problems in avian system- atics. It hasusually been placed in the Galliformes,but someauthors have allied it with the African turacos(Cuculiformes: Musophagidae), the pigeons(Columbidae), or the rails (Rallidae), and othershave placedit in its own monotypicorder. The Hoatzin occursin riparianvegetation alongthe streamsof the Orinocoand Amazonriver drainagesin northern SouthAmerica. It is a slenderbird, about 25 inchesin lengthand generally brownishin color, somewhatresembling a chachalaca(Cracidae). The smallhead bears a ragged,bristly, reddish-browncrest and the bare facial skin is bright blue. As is apparentfrom the Frontispiece,the Hoatzin is actuallymost like the smallerGuira Cuckoo(G. guira) in coloration;in fact, the two speciesare remarkablysimilar, except in size. This paperreviews the taxonomichistory of the Hoatzinand presents newevidence from a studyof the egg-whiteproteins indicating that the resemblanceto Guira is due to a closerelationship, not merelyto con- vergenceor coincidence.The correctposition of the Hoatzinis as a genus withinthe neotropicalsubfamily Crotophaginae (Cuculidae) and its dosest relativesare Guira and Crotophaga.The longassociation with the Gal- liformes,we believe,was based upon little morethan the prejudiceestab- lishedby the originaldescription as Phasianushoazin (Mi•ller 1776) and the inabilityof subsequentworkers to interprettheir anatomicaldata. The link to the Musophagidaeseems to be basedupon the commonpossession of a numberof anatomicalcharacters (see Verheyen 1956), the vegetarian diets, poor flying ability, generalproportions, and similar size. In the turacosand in the Hoatzin the younghave wing clawsand clamberabout near the nest before they can fly. THE TAXONOMIC HISTORY OF THE HOATZIN In generalappearance a museumspecimen of the Hoatzin resemblesa galliformbird andit is not surprisingthat M[iller (1776) originallynamed 1 The Auk 90: 1-13. January 1973 The tlOATZIN(lower tigure) and the GUIBACUCKOo dr•twn to the samesize. Insert shows relative sizes of the sanac birds. From an original acrylic painting by Jon Ahlquist. SIBLEY AND AllLQUIST [Auk, Vol. 90 TABLE 1 SOME OPINIONS ON THE SYSTEMATIC POSITION 01* OPISTHO½OMUS Evidence of relation- Placed in Galliformes Placed in separateorder Placed near Musophagidae ship to Cuculidae Miiller, 1776 Linnaeus, 1788 Illiger, 1811 Cuvier, 1817 L'Herminier, 18372 Nitzsch, 1829, 1840 Carus, 1868 Huxley, 18682 Gray, 1844-49, Garrod, 1879• 1869-71 Reichenow, 1882 Sclater, 1880• Cabanis, 1847ø Newton, 18842 Fiirbringer, 18884 Elliot, 1885a Goodchild, 1886 Sharpe, 18915 (secondary coverts) Gadow, 1892, 18934 Beddard, 1898• Pycraft, 1895 (pterylography) Shufeldt, 1904 Stresemann, Beebe, 1909 1927-34, 1959ø (behavior) Lemmrich, 1931 (sclerotic ossicles) Wetmore, 1930, 1934, 1940, 1951, 1960 Peters, 1934 Berlioz, 1950 Mayr and Amadon, Barnikol, 1953 Verheyen, 1956 1951 Storer, 1960 Verheyen, 1961 Sibley and Ahlquist, 1972 • Between Galliformes and Cuculiformes. a Between Galliformes and Columbiformes. a Next to Ga]]iformes. a Noted ties to Musophagidae. • Between Co]umbae and Ralli, following Seebohm (1888, 1890, 1895) who believed that Opistlto- comushad gruiform affinities. o Cabanis was apparently the first to note the similarity in co]or and pattern between Opistltocomus and Guira. it Phasianushoazin. Apparently this was enoughto establisha bias in favor of galliformaffinities, and in most classificationsto the presenttime, the Hoatzin has beenplaced in or near the Galliformes. Someof the opinions concerningthe classificationof the Hoatzin are summarizedin Table 1. The first suggestionthat the Hoatzin might be related to the Muso- phagidaewas that of Nitzsch who found similaritiesin the carotid arteries (1829) and pterylography(1840). In both papersNitzsch placedOpistho- comuswith Colius and the Musophagidaein the "family" Amphibolae. January 1973] Hoatzin Relationships 3 Gray (1844-49, 1869-71) followeda similarscheme, placing these three groups,along with the Bucerotidaeand severalpasserine families, in the Conirostres. Huxley (1867) first examinedonly an incompleteskull and the feet of a Hoatzin. He found that the slendervomer bifurcates anteriorly in an unusualway but he consideredthe palate to be schizognathus,as in the Galliformes,rather than desmognathous,as are the cuckoos.Huxley con- sideredthe tarsometatarsusto be galliform and assignedOpisthocomus to a "specialsubdivision," Heteromorphae, in his large "suborder"Schizog- nathae. He placed the Heteromorphaebetween the sandgrouse("Ptero- clomorphae")and the pigeons("Peristeromorphae"), not far from the galliforms ("Alectoromorphae").Huxley soonobtained a completeskele- ton and (1868) decidedthat in the majority of its osteologicalcharacters the Hoatzin resembles the Galliformes and the Columbiformes and that where it differs from these it is either unique or like the Musophagidae. He concludedthat Opisthocomusis a highly modified form derived from "Gallo-columbine"stock, worthy of distinctionin his monotypic"Hetero- morphae." Huxley's prestigeand influencewere great and these pronouncements must have had a strong effect upon subsequentworkers. The galliform affinities of the Hoatzin seemedconfirmed, but an alliance to the cuckoos, via the turacos,had beenleft open. This basicconcept has prevailedto the presentday, in spite of evidenceto the contrary. Garrod (1879) was amongthe first to follow Huxley's lead by placing the galliform and cuculiformbirds togetherin his Galliformes(which also included the ratites, tinamous, rails, seriemas,and bustards). Garrod expressedhis beliefthat Opisthocomusis an intermediateform, linking the Cuculidaeand Musophagidaeto the typical galliforms.He thus emphasized that Opisthocomushas a relationshipto the cuckoos,although he retained the supposedlink to the Galliformesas well. Otherswho adoptedOpisthocomus as the evidencefor a galliform-cuculi- form allianceincluded Ftirbringer (1888) who placedthe family Opistho- comidaenext to the Gallidae in the "gens" Galli, but also noted a dozen characterslinking Opisthocomuswith the Musophagidae. Gadow (1892: 229) also consideredOpisthocomus to be a connection betweenthe cuckoosand the galliformsbut (1893) he placed the Opistho- comiin the Galliformes.Like Ftirbringer,Gadow consistently emphasized the allianceto the cuckoos.Gadow (1893: 177) noted the followingchar- acters in comparisonsamong the three groups: I. Opisthocomusagrees with the Galliformesand differs from the Cucu- liformes in: 1) downy precocialyoung, not blind at hatching, 2) 4 SlBLEYAND AI-ILQUIST [Auk, Vol. 90 schizognathouspalate, 3) fusion of many dorsalvertebrae, 4) syrinx, 5) foot structure, 6) large crop. II. Opisthocomusagrees with the Cuculiformesand differs from the Galliformesin: 1) deep temporal fossae,2) short mandibularproc- esses,3) lack of basipterygoidprocesses, 4) lack of spina interna, 5) metasternalconfiguration, 6) large procoracoid,7) spottedeggs, 8) 10 rectrices. III. Opisthocomusdiffers from both Galliformes and Cuculiformesin: 1) lacking apteria on the sidesof the neck, 2) arrangementof the adult downs,3) numberof cervicalvertebrae, 4) small thoracichae- mapophyses,5) shapeof the liver, 6) other specializedcharacteristics, for example,sternum, stomach, crop. Becausevirtually all subsequentclassifications, including those of Strese- mann (1927-34), Wetmore(1930, 1934, 1940, 1951, 1960), Peters(1934) and Mayr and Areadon (1951) are basedupon Gadow and FiJrbringer,it is easy to understandwhy the Hoatzin has been placed so consistentlyin the Galliformes. The anatomical evidenceindicating a relationship to the Cuculidae includesa few additionalpoints. Pycraft (1895: 362) found a general resemblancein the pterylographyof the spinal tract amongOpisthocomus, Tauraco, and Centropusbut declinedto offer an opinion on their relation- ships. Lemmrich (1931) studied the bony platelets of the scleroticring in severalgroups of birds. He found that all of the 10 speciesof galliforms he examinedhad 13 to 15 plates but that Opisthocomushas only 12. Lemmrich noted (p. 534) that such a differenceis "very remarkable" becausethere tends to be little variation within a group. Among the few other birds with 12 plates is Cuculus. The skull musculatureand its innervationin Opisthocomuswere studied by Barnikol (1953). He comparedthe Hoatzin with representativespecies of galliforms,turacos, and pigeonsand summarizedthe study in a table (p. 520) listing 40 anatomicalcharacters. Of these Opisthocomusshared 8 with the Galliformes,9 with the Columbidaeand 13 with the Muso- phagidae. Barnikol concludedthat Opisthocomusis an isolatedspecies deservingrecognition as the soleoccupant of the order Opisthocomae. Verheyen (1956) defined 66 osteologicalcharacters of the Hoatzin and found that 50 of them are shared with the Musophagidae. This led him to combinethe two groupsas the orderMusophagiformes, placed near the Galliformesand the "Anhimiformes."However, in his later (1961) clas- sification Verheyen included Opisthocomusnext to the Cracidae in his Galliformes,"owing to new information" (p. 21). January 1973] Hoatzin Relationships 5 Some especiallyconvincing sets of data arguing against a galliform alliancefor Opisthocomushave appearedin recentyears. In 1959 Hudson, Lanzillotti, and Edwardsfound 13 aspectsof the
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