Monophyly and phylogeny of cuckoos (Aves, Cuculidae) inferred from osteological characters

Zoological Journal of the Linnean Society (2000), 130: 263–307. With 15 figures doi:10.1006/zjls 1999.0216, available online at http://www.idealibrary.com on

Monophyly and phylogeny of cuckoos (Aves, Cuculidae) inferred from osteological characters JANICE M. HUGHES∗ Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada M5S 2C6 Received October 1997; accepted for publication August 1999

A reanalysis of 32 characters from the literature previously deemed diagnostic of the Cuculidae revealed only five to be synapomorphic. I subsequently examined skeletons from 54 avian families and identified nine additional synapomorphies that supported cuckoo monophyly. My cladistic analysis of 33 cuculid genera using 135 skeletal characters differs markedly from currently accepted taxonomies. The most striking deviation is the placement of both New and Old World parasitic cuckoos in the Cuculinae, supporting the evolution of brood parasitism in a single event rather than three times as previously proposed. Unlike earlier classifications, the Cuculinae also includes the facultative parasites Coccyzus. This suggests that the ancestral Coccyzus was an obligate parasite, and is consistent with the many behavioral adaptations to parasitism exhibited by this genus. Other changes include the placement of three subfamilies, comprising non-parasitic, terrestrial cuckoos of Old World (Centropodinae and Carpococcystinae) and New World (Neomorphinae) distribution, in basal positions on the tree. Nineteen characters support a sister relationship between the Hoatzin (Opisthocomus hoatzin Mu¨ller) and turacos (Musophagidae), and not cuckoos. Three synapomorphies of the os carpi ulnare were found to unite the Cuculidae, turacos, and the Hoatzin, suggesting that these three diverse taxa may constitute a monophyletic group.  2000 The Linnean Society of London

ADDITIONAL KEY WORDS:—systematics – birds – Hoatzin – Musophagidae – Cuculiformes – brood parasitism – Opisthocomus – Coccyzus. CONTENTS

Introduction . . . . Material and methods Monophyly . . Phylogeny . . . Results . . . . . Monophyly . . Phylogeny . . . Discussion . . . . Monophyly . . Phylogeny . . .

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∗ Present address: Dept. Biology, Lakehead University, 955 Oliver Road, Thunder Bay, Ontario, Canada P7B 5E1. E-mail: [email protected] 0024–4082/00/100263+45 $35.00/0

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Acknowledgements References . . Appendix 1 . . Appendix 2 . . Appendix 3 . .

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INTRODUCTION

The Cuculidae is an ancient and diverse avian family comprising 129 species in 38 genera (Morony, Bock & Farrand, 1975). Most currently accepted classifications include cuckoos and their putative sister group, Musophagidae, in the Cuculiformes (Peters, 1940; Wetmore, 1960; Morony et al., 1975; Howard & Moore, 1991). The musophagids, or turacos, are an enigmatic family (20 species in 6 genera) of frugivorous, arboreal birds that are restricted to tropical Africa (Howard & Moore, 1991). However, some taxonomists have suggested that these taxa are too divergent to be in the same order (e.g. Bannerman, 1933; Lowe, 1943; Berger, 1960; Sibley & Ahlquist, 1990). More recently, the Hoatzin (Opisthocomus hoazin Mu¨ller) has been considered the sister taxon to cuckoos (Hedges et al., 1995; Mindell et al., 1997), or is in fact a cuckoo (Sibley & Ahlquist, 1972, 1973, 1990). The Hoatzin is a peculiar and anatomically divergent bird (Seibel, 1988) that inhabits the tropical forests of South America (Strahl, 1987). It is the only avian folivore that ferments leaves in a modified crop within a skeleton that is highly adapted to accommodate this unique digestive system (Grajal et al., 1989). In addition, the chick possesses wing claws that are used to climb among the vegetation of the nesting tree (Grimmer, 1962). Although cuckoo monophyly has been accepted for over 100 years, many characters used commonly to distinguish the Cuculidae from other taxa are either plesiomorphic or homoplasious among avian families (e.g. zygodactyl feet, holorhinal nares, desmognathous palate). Furthermore, Sibley & Ahlquist (1972, 1973, 1990) questioned cuculid monophyly when they classified the Hoatzin among cuckoos. Consequently, it is important that traditionally accepted diagnostic characters be reviewed to determine their validity (Appendix 1). In addition, I examine characters offered by Seibel (1988) as being synapomorphic of cuckoos. Finally, I present a list of synapomorphies compiled following my examination of skeletons from 54 avian families. This list is sufficient to establish cuckoo monophyly for the following purposes: (1) phylogenetic reconstruction, (2) evaluation of Hoatzin taxonomy, and (3) provision of basic guidelines for inclusion of all extinct and fossil taxa currently attributed to the Cuculidae. The classification of cuckoos has had a long and enigmatic history. Systematists have attempted to interpret relationships within the family by studying morphology (e.g. Beddard, 1885; Shufeldt, 1901; Pycraft, 1903; Verheyen, 1956a; Berger, 1960; Seibel, 1988), genetics (Sibley & Ahlquist, 1990; Avise, Nelson & Sibley, 1994), and behaviour (Hughes, 1996b). However substantial, and often ‘idiosyncratic’, differences observed among cuckoos have not yielded a consensus among several investigators. Although problematic, the widely accepted classification of Peters (1940) remains the basis for most contemporary cuculid taxonomies. Most contentious arrangements of taxa occur in two of six subfamilies: Neomorphinae and Phaenicophaeinae. The Neomorphinae comprise 13 species of parasitic and non-parasitic birds. Although they differ substantially in anatomy and life history, Peters based these species’

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inclusion in the subfamily on their New World distribution (except Carpococcyx) and their predilection for terrestrial foraging (Berger, 1960). Before Peters, most systematists had placed the neomorphine obligate parasites Tapera and Dromococcyx in a separate subfamily (Diplopterinae; e.g. Sclater & Salvin, 1873; Shelley, 1891) or with Old World parasites (Cuculinae; e.g. Beddard, 1885; Gadow & Selenka, 1891). Peters admitted dissatisfaction with the Phaenicophaeinae, referring (Peters, in litt. to Berger, 1952: 567) to the subfamily as “a general ‘catch-all’ group for genera that cannot be satisfactorily allocated in the other subfamilies”. This unnatural, albeit convenient, assemblage of 31 New and Old World species includes at least seven monotypic genera, as well as the facultative parasites Coccyzus. Peters’ justification for this grouping remains unknown. He may have assigned arbitrarily all arboreal, non-obligate parasitic cuckoos to the Phaenicophaeinae. In the decades since Peters’ volume, several morphological, molecular, and behavioral systematists have undertaken limited assessments of cuckoo relationships (Berger, 1952, 1955, 1960; Verheyen, 1956a; Sibley & Ahlquist, 1972; Brush & Witt, 1983; Avise et al., 1994; Hughes, 1996b). Although these workers did not produce definitive classifications, they nonetheless revealed inconsistencies in traditional classifications. Seibel’s (1988) phylogenetic reconstruction of the Cuculidae based on 48 postcranial osteological characters remains unpublished. In addition, a new cuculid classification was proposed by Sibley & Monroe (1990) based on a DNADNA hybridization study of Class Aves (Sibley & Ahlquist, 1990). In the present study, I reconstruct the phylogeny of cuckoos using both cranial and postcranial osteological characters. The dichotomy of generalized locomotory habits (arboreal vs terrestrial) among cuckoos justifies the inclusion of cranial characters, which are independent of the appendicular skeleton. In addition, postcranial characters are largely independent of feeding methods and, hence, may be useful in resolving relationships in an ancient and diverse group that occupies many habitats and uses a variety of foods. I also re-evaluate and subsequently revise the list of postcranial characters used by Seibel (1988). My study contrasts with previous analyses by inclusion of: (1) most cuculid genera and (1a) a character set sufficiently large to resolve all relationships among the taxa, and (2) the use of cladistic methodology. I then propose a new phylogenetic classification of the Cuculiformes, and follow by discussing it in light of the evolution of brood parasitism, the informed relationship of locomotory strategies to habitat use, the logical interpretation of biogeography, and the rationale of cuculid origins.

MATERIAL AND METHODS

Monophyly Many phylogenetic studies in ornithology merely assume ingroup monophyly. However, some workers have demonstrated that several well-accepted groups may not be monophyletic; e.g. Hawaiian honeycreepers (Drepanididae; Raikow, 1978), manakins and cotingas (Pipridae, Cotingidae; Prum, 1990), and blackbirds (Agelaius; Lanyon, 1994). Thus many widely accepted taxa may be paraphyletic or polyphyletic, and taxonomists have consequently assumed monophyly due to their familiarity with established classification and lack of empirical evidence to think otherwise.

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Now, as systematists re-evaluate avian classification using robust methodologies unavailable to earlier taxonomists, it is imperative that contemporary workers first demonstrate monophyly so that subsequent analyses can be used effectively to evaluate traditional sequences. I used outgroup comparison method of Raikow (1982) to establish monophyly of the Cuculidae. This procedure dictates that monophyly of a study group is demonstrated through character state comparisons within a more inclusive group that itself can be shown to be monophyletic (Gaffney, 1979). Traditionally, turacos (Musophagidae) were considered the sister group to cuckoos and both families were included in the Cuculiformes. However, this association was questioned by Sibley & Ahlquist (1990) following their DNA-DNA hybridization analyses. Consequently, cuculiform monophyly could not be assumed and it was imperative that I select a higher-level taxon to serve as the ingroup. Class Aves is monophyletic (Raikow, 1982). Furthermore, primary divisions of Class Aves have been demonstrated to be monophyletic (Sheldon & Bledsoe, 1993). Therefore, I used Parvclass ‘other neognaths’ (fig. 6b in Sheldon & Bledsoe, 1993) as ingroup. The outgroup was Parvclass Galloanserae, which includes Galliformes and Anseriformes. Character states that were present in some ‘other neognaths’ taxa, but not in Galloanserae, were considered to be derived. Support for cuckoo monophyly was given by character states that were derived within ‘other neognaths’ and unique to the Cuculidae. Derived character states that were not unique to the Cuculidae could be either plesiomorphic at a higher taxonomic rank (symplesiomorphic), or homoplasiously derived. Because neither condition corroborates a hypothesis of monophyly, such character states were considered non-diagnostic. I examined 306 disarticulated skeletons from 54 families in 25 avian orders. This included 209 specimens from 68 cuckoo species representing 33 of 41 genera. The three-species genus Cercococcyx and monotypic genera Rhamphomantis, Caliechthrus, Microdynamis, Scythrops, Taccocua, Zanclostomus, and Phaenicophaeus were excluded because no specimens were available for study. In the initial analysis, I reviewed 32 osteological characters from the literature that have been previously considered diagnostic of the Cuculidae, including eight potential synapomorphies proposed by Seibel (1988; Appendix 1). In a second analysis, I surveyed all specimens for previously undescribed osteological synapomorphies of cuckoos.

Phylogeny Taxa and specimens The genera- and species-level taxonomy of cuckoos has changed many times in past decades. In recent classifications, the Cuculidae has comprised at least 129 species (Morony et al., 1975) arranged in 29 (Sibley & Monroe, 1990) to 40 (Howard & Moore, 1991) genera. This wide discrepancy results primarily from varied treatment of approximately 20 monotypic taxa within the family. I followed Howard & Moore (1991) because their arrangement of genera divides the family into the largest number of operational units. In addition, I include the genus Coccycua, usually placed in Piaya (e.g. Peters, 1940; Sibley & Monroe, 1990), but viewed as a monotypic genus in a few older classifications (e.g. Ridgway, 1916; Cory, 1919). My decision to partition accepted generic groupings is not an assertion that other classifications that merge them are invalid. This approach merely simplified the identification of

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intrageneric variability when constructing characters, and also allowed for heuristic examination of other classifications once a tree was produced. In total, 33 cuculid genera were included in my study. Eight genera—Cercococcyx (3 spp.), Rhamphomantis (1 sp.), Caliechthrus (1 sp.), Microdynamis (1 sp.), Scythrops (1 sp.), Taccocua (1 sp.), Zanclostomus (1 sp.), and Phaenicophaeus (1 sp.)—were not used because skeletal material was limited or unavailable. Two or more specimens were examined for all genera except Pachycoccyx, Urodynamis, and Rhinortha. Full skeletons were available for all taxa but Pachycoccyx. A list of specimens used for this study is available from the author upon request. Outgroup selection Outgroup selection was problematic. Sibley & Ahlquist (1990: 370) suggested that cuckoos “have no close living relatives, but are the sister group of a large assemblage that includes more than half of the groups of living birds”. By convention, Seibel (1988) used turacos (Musophagidae) as outgroup because this family is most commonly classified in Cuculiformes (e.g. Peters, 1940; Wetmore, 1960; Howard & Moore, 1991). However, Seibel identified only three synapomorphies of the os carpi ulnare to support this relationship implying that, if cuckoos and turacos are sister taxa, they diverged early from one another. Other authors (De Queiroz & Good, 1988; Hedges et al., 1995; Mindell et al., 1997) submit that the Hoatzin is the sister taxon to the Cuculidae. Furthermore, Sibley & Ahlquist (1972, 1973, 1990) suggested that the Hoatzin is a cuckoo. They placed the Musophagidae (=Musophagiformes, their study) in a different superorder (Strigimorphae) from the Cuculidae. In contrast, Hughes & Baker (1999) demonstrated that the hoatzin is sister to turacos, not cuckoos. The uncertainty of appropriate outgroup selection led me to examine skeletons from 54 avian families. Rather than make an a priori decision regarding cuckoos’ closest relatives, I considered all plausibly related groups that possessed at least superficial similarities to cuckoos in major skeletal elements. In addition, I included two non-passerine families of unknown affinities, mousebirds (Coliidae) and trogons (Trogonidae), as well as individuals from three families of Galliformes. In total, 22 taxa from 11 avian families were selected as preliminary outgroups and coded for the same characters used to reconstruct the cuckoo phylogeny: Alectura (Megapodidiae); Ortalis, Crax (Cracidae); Phasianus, Tympanuchus, Tragopan (Phasianidae); Crinifer, Corythaixoides, Musophaga, Tauraco (Musophagidae); Opisthocomus (Opisthocomidae); Colius, Urocolius (Coliidae); Trogon, Apaloderma (Trogonidae); Merops (Meropidae); Coracias (Coraciidae); Tockus, Bycanistes (Bucerotidae); Bucco, Monasa, Malacoptila (Bucconidae). Additional states were added to characters to accommodate morphological divergence observed among some more distantly related taxa. An initial phylogenetic analysis included all outgroups. Character analysis A total of 135 skeletal characters, 56 cranial and 79 postcranial, were used to reconstruct a phylogeny of cuckoos (Appendix 2). Osteological nomenclature follows Baumel & Witmer (1993). Sixteen postcranial characters were taken directly from Seibel (1988). An additional 37 postcranial characters were based on 29 of Seibel’s 48 characters, but were altered from his original analysis in one or more ways: (1) multistate characters were divided into two or more binary characters, (2) informative intermediate conditions were assigned individual character states, (3) character states

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not previously recognized were added, and (4) ordering hypotheses were modified. In several cases, all character conditions were reanalysed. Characters were polarized into a plesiomorphic state and one or more apomorphic states through outgroup comparison with turacos and the Hoatzin. More distant outgroups were considered when necessary. Characters for which information was unavailable or irretrievable for particular taxa were coded as missing (?). Multistate characters were considered unordered unless a reasonable hypothesis of character evolution could be formulated. A 35×135 data matrix depicting all assigned character state codes can be found in Appendix 3. To ensure that the resulting topology was not influenced by differences in locomotory function among cuckoo species, a supplementary analysis was performed using characters independent of the appendicular skeleton. Fifty-eight characters from the tibiotarsus, tarsometatarsus, femur, pelvis, sternum, humerus, radius, ulna, and manus were deleted. Tree derivation and analysis Trees were constructed using PAUP 4.0b2a (Swofford, 1998). Supplementary tree and character analyses were performed using MacClade 3.01 (Maddison & Maddison, 1992). Optimal trees were found using the branch and bound algorithm. MULPARS option was in effect. MAXTREES was set at 1000 trees with automatic increase. Zero-length branches were collapsed to yield polytomies. Separate analyses were performed using all character state optimization schemes (ACCTRAN, DELTRAN, MIN-F), and addition sequences (furthest, as is, simple). Implementing these different options did not alter the outcome of the analysis. The consistency index (CI), retention index (RI), and rescaled consistency index (RC) were calculated. All characters were given equal weight. Character support for the optimal tree was assessed by a full heuristic bootstrap analysis of 1000 replicates.

RESULTS

Monophyly My reanalysis of 32 characters deemed diagnostic of cuculid affinities by other workers found 27 uninformative. Of these, 20 (Appendix 1: characters 2–9, 12–14, 16–21, and 23–32) could not establish monophyly because they were either symplesiomorphic, or convergent derived conditions observable in other families. Seven additional characters (9, 12, 14, 16, 17, 24, and 25) were uninformative at the familial level as they were not found in all cuculid taxa studied. Representation of these seven characters as diagnostic was likely attributable to previous investigators examining a single individual from an incomplete series of taxa. Only five potentially diagnostic characters were identified as being synapomorphic (characters 1, 10, 11, 15, and 22), including only two of eight proposed by Seibel (characters 10 and 15). Seibel’s other purported synapomorphies separated cuckoos from turacos in his phylogenetic analysis, but did not uphold monophyly when examined in other families. These synapomorphies are described below in extended form and supplemented with nine additional cuculid synapomorphies resulting from my examination of the skeletal material. This list, by no means exhaustive, is nonetheless

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Figures 1 & 2. Greater Roadrunner (Geococcyx californianus). Skull in (1) right lateral and (2) right caudolateral aspect. Os jugale not shown. OE=os ectethmoidale; OQ=os quadratum; OP=os pterygoideum.

sufficient to establish cuckoo monophyly for purposes of my phylogenetic reconstruction that follows. (1) Os quadratum. Condylus medialis, caudalis, and lateralis prominent, well rounded, and separated by a broad notch; condylus caudalis tapered to a rounded point, deflected and extending well beyond caudal margins of quadrate in lateral aspect; processus orbitalis quadrati truncated and somewhat anvil-shaped (Fig. 1). (2) Os pterygoideum. Mesially deflecting spur-like processus dorsalis located caudally on facies dorsalis near, but not articulating with, os quadratum (Fig. 1). (3) Os ectethmoidale. Large, quadrilateral in form, marginis lateralis and ventralis slightly to moderately excised (Fig. 2). (4) Os palatinum. Marginis medialis meeting in midline below, fully or nearly concealing, rostrum parasphenoidale; lamella caudolateralis being widest midway between processus pterygoideus and margo distalis of pars choanalis, tapering towards rostrum; processus interpalatinus extending nearly to margo caudalis of processus maxillopalatinus; in lateral aspect, cristae lateralis are deflected ventrally (Fig. 3). (5) Os maxillare. In ventral aspect, processus palatinus bilobed, extending caudally nearly to marginis distalis of crista ventralis; margo medialis of processus palatinus not fused caudally to margo distalis of os maxillare (Fig. 3). (6) Os articulare, crista intercotylaris. Prominent, half-disk shaped with radius aligned approximately 45° to planum medianum (Fig. 4). (7) Tibiotarsus, incisura intercondylaris. Deep, rounded pit extending caudally at least

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Figures 3 & 4. Greater Roadrunner. Fig. 3. Skull in ventral aspect. OPL=os palatine; OM=os maxillare. Fig. 4. Os articulare in dorsal aspect.

half the length of extremitas distalis (Gilbert, Martin & Savage, 1981); trochlea cartilaginis tibialis poorly developed (Fig. 5). (8) Tarsometatarsus, hypotarsi. Two oblong canales hypotarsi, completely enclosed in bone, positioned side by side in planum medianum both almost entirely caudal to cotylae medialis and lateralis, and corpus tarsometatarsi (Seibel, 1988: TM13; Fig. 6). (9) Tarsometatarsus, trochlea metatarsi quarti. Margo distalis proximal to incisura intertrochlearis medialis (Seibel, 1988: TM21); caudal trochlea accessoria prominent and strongly inflected medially (Gilbert, Martin & Savage, 1981; Fig. 7). (10) Coracoideum, facies articularis sternalis. Crista ventralis prominent, centred, rounded, and facing sternally (Gilbert, Martin & Savage, 1981; Fig. 8). (11) Scapula. Facies articularis humeralis directed dorsally; facies articularis clavicularis prominent, knob-like, and directed dorsally; extremitas cranialis scapulae, delineated by acromion and tuberculum coracoideum, truncated and flattened cranially (Fig. 9). (12) Humerus, condylus ventralis. Rounded, not flattened or oblong, in axis proximodistalis and both plana transversalia and dorsalia (Fig. 10). (13) Synsacrum, extremitas caudalis synsacri. Crista dorsalis reduced and fused to form

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Figures 5 & 6. Greater Roadrunner. Fig. 5. Right tibiotarsus in caudal aspect. Fig. 6. Right tarsometatarsus in cranial aspect.

Figure 7. Greater Roadrunner. Right tarsometatarsus in (A) dorsal and (B) caudal aspect.

a single low ridge; marginis lateralis of processes transversus widened in axis rostrocaudalis, facies lateralis projecting cranially (Fig. 11). (14) Vertebrae caudales, primus. Processes transversus tapered, not widened, paddleshaped in ventral aspect; projecting strongly caudally (Fig. 12). Phylogeny Cuckoos The preliminary analysis using all outgroups indicated that the most likely sister taxa to cuckoos were turacos and the Hoatzin, which were then used as outgroup in subsequent analyses. This resulted in one fully-resolved shortest-length tree of

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Figures 8 & 9. Greater Roadrunner. Fig. 8. Right coracoideum in dorsal aspect. Fig. 9. Right scapula in dorsal aspect.

Figure 10. Greater Roadrunner. Right humerus in (A) cranial and (B) distal aspect.

238 steps with CI=0.798, RI=0.941, and RC=0.751 (Fig. 13; with cuckoos only: length=191 steps, CI=0.948, RI=0.939, RC=0.890) that differs significantly from the currently accepted classification of cuckoos based primarily on Peters (1940). Perhaps the most striking deviation is the placement of all obligately parasitic cuckoos in a single monophyletic subfamily (Cuculinae; Fig. 13, node 66). Within the Cuculinae are the (1) ‘higher parasites’ (node 43) comprising Cuculus, Cacomantis, Penthoceryx, Surniculus, Chrysococcyx, Chalcites, Misocalius, and Pachycoccyx; (2) terrestrial, New World obligate parasites Tapera and Dromococcyx, formerly in the Neomorphinae; and (3) facultatively parasitic Coccyzus, previously in the Phaenicophaeinae, now sister taxon to the Clamator-Oxylophus complex. This contrasts with Peters (1940), which suggested that brood parasitism arose independently at least three times in cuckoos. In addition, my results indicate that terrestrial habits of Tapera and Dromococcyx have evolved secondarily. Arboreal, non-parasitic cuckoos of the Phaenicophaeinae (except Coccyzus) form a monophyletic group that is sister to parasitic cuckoos (Fig. 13, node 53). The New World genera—Saurothera, Hyetornis, Piaya, and Coccycua—form a clade (node 47). An

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Figures 11 & 12. Cuban Lizard-cuckoo (Saurothera merlini). Fig. 11. Pelvis in dorsal aspect. Fig. 12. Pelvis in ventral aspect.

additional clade (node 50) includes four genera of malkohas, Rhopodytes, Rhamphococcyx, Lepidogrammus, and Dasylophus, which have been merged into Phaenicophaeus by some authors (e.g. Delacour & Mayr, 1945; Sibley & Monroe, 1990). Ceuthmochares and Rhinortha occupy basal positions in the Phaenicophaeinae. The communally breeding cuckoos of the Crotophaginae, Crotophaga and Guira, are monophyletic (node 54) and are the sister group to the Cuculinae and Phaenicophaeninae. In basal positions are three paraphyletic taxa of terrestrial cuckoos. Non-parasitic New World neomorphine cuckoos, Morococcyx, Geococcyx, and Neomorphus, form a clade (Fig. 13, node 56). Old World terrestrial cuckoos, Centropus and Coua (Centropodinae; node 60), are sister to the Neomorphinae. The monotypic subfamily Couinae has been dissolved with the transfer of Coua to the Centropodinae. Carpococcyx, an Old World form classified in the Neomorphinae (Peters, 1940, and most subsequent classifications) or Cuculidae (=Cuculinae) by Sibley & Monroe (1990), occupies a monotypic subfamily (Carpococcystinae) at the base of the tree (node 59). The basal position of these taxa suggests that the ancestral cuckoo was at least in part adapted to a terrestrial existence. One measure of topological stability is the number of synapomorphies defining the basal stem of each clade. My reconstruction of cuculid phylogeny is well supported at most subfamily and tribe levels. The Cuculinae is supported by nine character state changes, eight of which are synapomorphic. Within the Cuculinae, ‘higher parasites’ are united by 14 character state changes (nine synapomorphies). The Crotophaginae are united by four synapomorphies, and the Phaenicophaeinae by two synapomorphies in four character state changes. The Neomorphinae is supported by nine character state changes, six being synapomorphic. Less well defined are the Centropodinae and Phaenicophaeinae, being supported by two and four character state changes, respectively, of which only two are synapomorphic in the latter subfamily. The position of Carpococcyx as basal taxon is well supported, with 10 character state changes and four synapomorphies defining the Cuculidae without this genus (node 61). Overall, the Cuculidae is strongly supported by 28

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Figure 13. Optimal hypothesis of phylogeny for 33 cuckoo genera based on 135 osteological characters. Musophagidae and Opisthocomus are outgroups.

synapomorphies. Bootstrap support values for nodes defining cuculid subfamilies are Cuculinae 98% (node 66), Neomorphinae 100% (node 56), Centropodinae 80% (node 60), Crotophaginae 100% (node 54), and Phaenicophaeinae 54% (node 53). All bootstrap values are indicated on Figure 13. The supplemental analysis using only non-appendicular characters resulted in nine trees (L=129 steps; CI=0.798; RI=0.926; RC=0.739) that differs from the full-character optimal tree only by minimal resolution loss among some higher parasites. My analysis supports, in part, merging the following genera: Oxylophus and Clamator (
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The status of the malkohas is unclear. Although this taxon is monophyletic, within malkohas there are two clades separated by nine character state changes—two uniting Rhopodytes and Rhamphococcyx, seven joining Lepidogrammus and Dasylophus— three of which are synapomorphic. This degree of divergence may controvert the suppression of these four genera into Phaenicophaeus. Outgroups The preliminary analysis suggested that the Hoatzin and turacos form a sister clade to cuckoos (Fig. 13; node 58). Three synapomorphies of the os carpi ulnae (this study; Seibel, 1988) corroborate the results of Hughes & Baker (1999) and imply that cuckoos, turacos, and the Hoatzin constitute a monophyletic group. Hence, the support for the Hoatzin-turaco clade can be tested using Ortalis (Cracidae) as outgroup. The Hoatzin (Opisthocomidae) has been considered the sister group to cracids by some authors (Sibley & Ahlquist, 1990). The Hoatzin-turaco clade was supported by 19 character state changes (eight synapomorphies) with 100% bootstrap support. Moving the Hoatzin to sister taxon of cuckoos, as suggested by Hedges et al. (1995) and Mindell et al. (1997) would require 10 additional steps. Furthermore, placing the Hoatzin basal to the Crotophaginae, as proposed by Sibley & Ahlquist (1972, 1973, 1990) would require an additional 56 steps. Character consistency Cranial characters had slightly lower mean CIs than did postcranial characters, 0.843 and 0.900, respectively. Although most characters had CIs of 1.00 (37 cranial; 57 postcranial), certain skeletal elements showed more homoplasy than others: os articulare (mean CI=0.600, n=5), os basioccipitale (0.611, 3), ulna (0.746, 5), and os exoccipitale (0.750, 4). In addition, many potential characters of the os palantinum and os ectethmoidale were examined and subsequently discarded because they were phylogenetically uninformative (CI<0.250). The most informative skeletal elements, in terms of useful characters extracted, were the tarsometatarsus (19 characters), pelvis (14), and os quadratum (9). When analysed separately, cranial characters resulted in 95 (L=97 steps, CI= 0.742, RI=0.904, RC=0.671), and postcranial characters in 40 (L=144 steps, CI=0.819, RI=0.953, RC=0.781) equal-length trees that were similar to the allcharacter optimal tree. Majority-rule (50%) consensus trees indicated that both cranial and postcranial characters successfully resolved all taxa, except the New World phaenicophaeine genera, at the tribe level and above.

DISCUSSION

Monophyly Many early systematists attempted to define a taxon using a single character and, as a result, classifications often included species of questionable affiliations. The Cuculinae of Nitzsch (1840), based on the superficial structure of the oil gland, included cuckoos, honeyguides (Indicator spp., Indicatoridae, Piciformes), trogons (Trogon spp., Trogoniformes), and the Cuckoo-roller (Leptosomus discolor Hermann, Leptosomatidae, Coraciiformes). Likewise, Lilljeborg (1886) placed honeyguides in

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the Cuculidae after observing similarities in their zygodactyl feet. In contrast, Goodchild (1891) concluded that cuckoos were polyphyletic based on the arrangement of secondary wing coverts. He suggested that ‘normal cuckoos’ (i.e. arboreal) were most closely associated with picarian birds (=Piciformes) and pigeons (=Columbidae), and ground cuckoos were allied with gallinaceous birds, in particular guans (Cracidae) and megapodes (Megapodiidae). However, by the late 19th century, most systematists considered cuckoos, as currently defined, to be a natural group that is anatomically divergent from other avian families. Although cuckoo monophyly was not questioned seriously for more than 100 years (but see below regarding the inclusion of the Hoatzin), previous workers failed to define and describe accurately those traits that they considered diagnostic. Most traditional characters were either plesiomorphic or homoplasious, and taxonomists generally described a taxon using a list of monothetic characters, all of which had to be satisfied for inclusion. In contrast, Seibel (1988) offered eight potential synapomorphies of the Cuculidae. However, my examination indicated that only two characters were synapomorphic, being found only in cuckoos and not among other avian families. My presentation of 14 cuculid synapomorphies sufficiently defines the Cuculidae to the exclusion of other purportedly related taxa, for purposes of phylogenetic reconstruction, and as a guideline for inclusion of extinct and fossil taxa. Hoatzin Sibley & Ahlquist (1972, 1973, 1990) concluded that the Hoatzin is a cuckoo most closely allied with anis (Crotophaga spp. and Guira guira Gmelin) or roadrunners (Geococcyx spp.) on the basis of protein electrophoresis and DNA-DNA hybridization data. However, aspects of Sibley & Ahlquist’s methodology have been criticized (e.g. Brush, 1979; Mayr & Bock, 1994). Their results were also questioned by Bock (1992) who suggested that the Hoatzin’s anisodactyl foot structure was sufficient to exclude it from zygodactylous cuckoos. In addition, many morphological characters cited by Sibley & Ahlquist as support for the Hoatzin’s inclusion in the Cuculidae show general similarities to anis (e.g. communal nesting, pelvic muscle formula, and arrangement of secondary wing coverts; see Sibley & Ahlquist, 1972, 1973, 1990 for a complete list), but have not been demonstrated to be homologous. Other characters, such as short processes mandibularis, large fossae temporalis, and absence of processes basipterygoideus, are symplesiomorphic and cannot be considered evidence for inclusion in the Cuculidae. Based on my results, I conclude that cuckoos are monophyletic without the Hoatzin. The taxonomic relationship between the Hoatzin and the Cuculidae is discussed below.

Phylogeny Taxonomy of cuckoos has remained virtually unchanged for many decades, with Peters (1940) still being the most widely accepted. Several subsequent studies (Verheyen, 1956a; Berger, 1960; Seibel, 1988) provided compelling evidence favoring the revision of cuculid classification, but failed to gain support from the ornithological community. Both Verheyen and Berger included too few taxa in their analyses to render any comprehensive conclusions, and Seibel’s work has remained in dissertation

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form only. Another cuckoo phylogeny (Hughes, 1996b) has been recently published. My analysis of cuculid relationships based on osteological data addresses most genera of cuckoos using a large character set and robust methodology of phylogenetic systematics unavailable to earlier taxonomists. Therefore, the results should be used not only to re-evaluate previous classifications, but also to propose a new hypothesis of evolutionary relationships within the group (Brooks & McLennan, 1991). A new phylogenetic classification of cuckoos based on the optimal tree (Fig. 13) is shown in Table 1. Single evolution of brood parasitism The Cuculidae are best known for the 55 species of brood parasites that lay their eggs in other birds’ nests. Of these, 51 species are obligate parasites (Wyllie, 1981), and at least four Coccyzus species are facultative parasites, which may lay their eggs both intra- and interspecifically when certain environmental conditions prevail (Nolan & Thompson, 1975; Ralph, 1975; Sick, 1993). One could infer from many earlier classifications that the evolution of brood parasitism in cuckoos resulted from at least three independent events: two origins of obligate parasitism in the Cuculinae and Neomorphinae, and at least one origin of facultative parasitism in the Phaenicophaeinae, sensu Peters (1940). However, it seems unlikely that so rare a behaviour in birds (about 1% of all species; Payne, 1977) would have arisen so many times in a single family. In contrast, my analysis supports a single evolution of brood parasitism in cuckoos. Behaviour can be used to subdivide the clade of parasitic genera into three clades of ‘lower parasites’—(1) Tapera and Dromococcyx; (2) Eudynamys, Urodynamis, and probably Scythrops not included in this study; and (3) Clamator, Oxylophus, and Coccyzus—in basal positions, and one clade of ‘higher parasites’— Pachycoccyx, Surniculus, Cacomantis, Cuculus, Chrysococcyx, Misocalius, and Chalcites—in an apomorphic position, consistent with the evolution of anti-host adaptations typically associated with parasitic cuckoos (Fig. 14). Based on my hypothesis of cuculid evolution, the first anti-host behaviours to arise among cuckoos were: (1) egg mimicry, whereby a species or gente has evolved an egg with base colour and/or maculations that resemble the eggs of frequently used host(s); and (2) host egg removal, in which a female cuckoo removes a single egg from a host nest prior to depositing her own egg. These behaviours are found among all Old World parasitic cuckoos, but are not exhibited by New World genera; suggesting that they arose early in the evolution of brood parasitism, but at some point after the divergence of Tapera and Dromococcyx. Most workers agree that egg mimicry in brood parasites has arisen in response to the selective pressure of hosts able to discriminate between the cuckoo’s and their own eggs. Mimetic eggs are the cuckoo’s evolutionary response to an egg-rejecting host because they are less likely to be discovered (Rothstein, 1990). However, the implied concurrent origin of egg mimicry and host egg removal lends support to an alternate hypothesis that suggests that egg mimicry may have evolved as a result of interspecific and/or intraspecific competition between parasitic females that are removing an egg from the host nest before depositing their own egg (Davies & Brooke, 1988; Brooker & Brooker, 1989, 1990). A mimetic cuckoo egg would have a selective advantage over a non-mimetic egg because it is less likely to be replaced by subsequent parasitic female cuckoos removing the most conspicuous egg in the clutch. Parasitic cuckoos are well known for host-offspring ejection behaviour of their nestlings, whereby a young cuckoo,

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T 1. Phylogenetic classification of cuckoos and sister taxa derived from optimal tree (Fig. 13). Taxonomic ranking after the methods in Wiley (1981) Order Opisthocomiformes (L’Herminer, 1837) Family Opisthocomidae (Fu¨rbringer, 1888) Family Musophagidae (Bonaparte, 1831) Order Cuculiformes (Wagler, 1830) Family Cuculidae (Vigors, 1825) Subfamily Carpococcystinae (Verheyen, 1956a) Genus Carpococcyx Subfamily Centropodinae (Gray, 1840) Genus Centropus Genus Coua Subfamily Neomorphinae (Shelley, 1891) Genus Morococcyx Genus Geococcyx Genus Neomorphus Subfamily Crotophaginae (Swainson, 1837) Genus Crotophaga Genus Guira Subfamily Phaenicophaeinae (Gray, 1840) Tribe Rhinorthini (Seibel, 1988) Genus Rhinortha Tribe Ceuthmocharini (Hughes, new tribe) Genus Ceuthmochares Tribe Phaenicophini (Salvin, 1882) Subtribe Rhopodytina (Hughes, new subtribe) Genus Rhopodytes Genus Rhamphococcyx Subtribe Phaenicophina (Hughes, new subtribe) Genus Lepidogrammus Genus Dasylophus Phaenicophaeini incertae sedis Genus Taccocua Genus Zanclostomus Genus Phaenicophaeus Tribe Saurotherini (Gray, 1840) Genus Saurothera Supergenus Piaya Genus Hyetornis Genus Piaya Genus Coccycua Subfamily Cuculinae (Vigors, 1825) Tribe Taperini (Verheyen, 1956a) Genus Tapera Genus Dromococcyx Tribe Eudynamini (Baker, 1927) Supergenus Eudynamys Genus Eudynamys Genus Urodynamis Eudynamini incertae sedis Genus Microdynamis Genus Scythrops Tribe Coccygini (Swainson, 1837) Genus Coccyzus Supergenus Clamator Genus Clamator Genus Oxylophus Tribe Cuculini (Vigors, 1825) Subtribe Pachycoccystina (Hughes, new subtribe) Genus Pachycoccyx Subtribe Chrysococcystina (Hughes, new subtribe) continued

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T 1—continued Supergenus Chrysococcyx (Berger, 1955) Genus Chrysococcyx Genus Misocalius Genus Chalcites Subtribe Cuculina (Vigors, 1825) Genus Surniculus Genus Penthoceryx Genus Cuculus Genus Cacomantis Cuculini incertae sedis Genus Cercococcyx Genus Rhamphomantis Genus Caliechthrus

Figure 14. Hypothesis of phylogeny for the Cuculinae based on osteological characters indicating the origin of brood parasitism and subsequent evolution of anti-host adaptations to parasitism.

within 3 hours to 4 days following hatching, uses the shallow depression in its scapular region to push host eggs or hatchlings over the nest rim. This behaviour ensures that the cuckoo is the sole occupant of the nest, and significantly improves its chance of fledging ( Jourdain, 1925; Payne, 1977). In my analysis, host-offspring ejection behaviour is synapomorphic among higher parasites. In contrast, lower parasite nestlings do not exhibit ejection behaviour, and are raised with one or more host chicks in the nest. Like other brood parasites, such as cowbirds (Icterinae;

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Figure 15. Pelvis of Cuban Lizard-cuckoo indicating sections A–C of crista dorsolateralis ilii.

Friedmann, 1963), whydahs and indigobirds (Viduinae; Payne, 1973), and the Cuckoo Weaver (Anomalospiza imberbis Cabanis, Ploceidae; Maclean, 1985), these cuckoos outcomplete host young by using their larger size, aggressive begging behaviour, and hyperstimulating mouth coloration (Hamilton & Orians, 1965). One notable exception is the Striped Cuckoo (Tapera naevia Linnaeus), a New World obligate parasite that uses mandibular hooks to kill host chicks in the same manner as parasitic honeyguides (Indicator, Piciformes; Morton & Farabaugh, 1979). It seems logical to separate the Cuculinae into two grades of parasites based on host-ejection because its presence or absence is often accompanied by a suite of other behaviours, such as host-chick mimicry ( Jourdain, 1925; Friedmann, 1964), begging-call mimicry (Courtney, 1967; Mundy, 1973; Morton & Farabaugh, 1979), and mouth-pattern mimicry (Payne, 1977) that are dependent on whether a cuckoo chick is raised alone or with nestmates. Inclusion of Coccyzus in the Cuculinae My classification differs markedly from Peters (1940) in the transfer of facultatively parasitic Coccyzus from the non-parasitic Phaenicophaeinae sensu Peters (1940) to the parasitic Cuculinae. It is likely that Peters grouped Coccyzus with New World phaenicophaeine cuckoos, such as Piaya and Saurothera, based on similarities in behaviour and vocalizations, New World distribution, and non-obligately parasitic breeding strategies (Berger, 1960). However, behavioral and vocal similarities between Coccyzus and New World phaenicophaeine cuckoos have not been shown to be homologous. In fact, many aspects of Coccyzus behaviour and ecology strongly link this genus to the Cuculinae (Hughes, 1996b). Earlier this century, most systematists classified Coccyzus with the Cuculinae based on internal anatomy, osteology, myology, and pterylosis (Beddard, 1885, 1901; Shufeldt, 1886, 1901, 1909; Shelley, 1891; Sharpe, 1900; Pycraft, 1903; Chandler, 1916). Coues (1897) suggested that the subfamily Coccyzinae—erected by Ridgway

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et al. (1899) to distinguish between New and Old World ‘tree cuckoos’—be abandoned based on the work of Beddard and Shelley. More recent examinations of Coccyzus osteology (Verheyen, 1956a, 1961; Seibel, 1988), myology (Berger, 1952, 1960), and behaviour and ecology (Hughes, 1996b) continue to support its placement in the Cuculinae. Sibley & Ahlquist (1990) addressed avian classification using DNA-DNA hybridization and produced a phenogram that clearly positions Coccyzus in the Coccyzinae, a ‘clade’ of New World phaenicophaeine cuckoos. However, it is unclear how these conclusions were formed because Coccyzus did not cluster consistently with that subfamily (see figs 56, 76, 80, and 84 in Sibley & Ahlquist, 1990). Likewise, a limited molecular analysis of cuculiform relationships (Avise et al., 1994) using cytochrome b sequences was inconclusive. Coccyzus associated with both phaenicophaeine and cuculine species using different character weighting schemes and substitution models. My analysis of cuckoo osteology strongly supports the inclusion of Coccyzus in the Cuculinae. Moving the genus to the base of the Saurotherini clade would require 29 additional steps, and would precipitate a reduction in the optimal tree CI from 0.798 to 0.700. In addition, Coccyzus differs from the Saurotherini in 33 characters (mean CI=0.822). That Coccyzus is a New World genus among an Old World subfamily is inconsequential. There are many reports of vagrant parasitic cuckoos (e.g. Oriental Cuckoo Cuculus saturatus Blyth and Common Cuckoo C. canorus Linnaeus) in western North America (Roberson, 1980; Cramp, 1985). Therefore, it is plausible that an ancestor could colonize the New World from Eurasian stock. Seibel (1988) was unable to resolve the position of Coccyzus within the Cuculinae. His preferred tree placed it in a tricotomy with Clamator and higher parasites (my term). Seibel’s (1988) difficulties stemmed from his failure to consider seriously the ‘puzzlingly intermediate’ (p. 82) conditions shared by Coccyzus, Clamator, and Oxylophus (Oxylophus), Coccyzus, and Pachycoccyx. However, I find that Pachycoccyx is paraphyletic to the Coccygini. Placing Coccyzus in the Cuculinae suggests that its ancestor was an obligate brood parasite. This implies that facultative behaviour exhibited by Coccyzus represents loss of obligate parasitism, rather than de novo development of parasitism from a nonparasitic ancestor. Traditional views suggest that Coccyzus represents an intermediate in a lineage that is becoming parasitic and that cuckoos possess some inherent predispositions that are fuelling the repeated evolution of parasitism (Miller, 1946; Berger, 1960; Hamilton & Orians, 1965). Although parasitism reversal seems unlikely, it is, ostensibly, the return to an ancestral state, and requires no ad hoc hypothesis of ‘preadaptation’ towards brood parasitism. Atavistic behaviour has been described among obligately parasitic cuckoos. There are numerous well-documented observations of adult cuckoos of the genera Chrysococcyx, Cuculus, Eudynamys, Scythrops (Friedmann, 1968), and Cacomantis (Ambrose, 1987) feeding fledglings of their own species. Fledgling feeding by Clamator has been observed, but reports have not been substantiated (Friedmann, 1964). Fleischer, Murphy & Hunt (1985: 125) commented that both intraspecific and interspecific parasitism were “regular aspects of Coccyzus breeding biology”. Although

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Yellow-billed (C. americanus Linnaeus) and Black-billed (C. erythropthalmus Wilson) cuckoos most frequently use each other as hosts where their ranges overlap, their eggs have also been found in nests of 15 other species (Herrick, 1910; Bent, 1940; Hughes, 1997), and at least seven host species have successfully hatched or fledged cuckoo young (Darwin, 1859; McIlwraith, 1894; Macon & Macon, 1909; Nickell, 1954; Nolan & Thompson, 1975; Wolfe, 1994). Unlike non-parasitic cuckoos, Coccyzus shares several life-history traits with the Cuculinae that seem adaptive to a parasitic life style. These include: (1) disassociation of egg-laying from the ‘normal’ nesting sequence (Kendeigh, 1952); (2) short incubation period that allows a parasitic chick to hatch first; (3) short nestling period that reduces the dependency period; (4) nestlings with an omnivorous diet (Hamilton & Orians, 1965); (5) delayed breeding season that allows the host to raise a brood unaffected by parasitism (May & Robinson, 1985); and (6) readiness to breed in response to exogenous stimuli (Hamilton & Hamilton, 1965; Ralph, 1975). In addition, Yellow-billed and Blackbilled cuckoos may lay mimetic eggs (Hughes, 1997). Although some traits may have adaptive value in a non-parasite that faces predation pressure or fluctuations in food availability, it is unlikely that their manifestation in both Coccyzus and the Cuculinae is entirely convergent. Ecological factors responsible for the reversal of parasitism in Coccyzus may have also contributed to the overall paucity of obligately parasitic cuckoos in the New World, in spite of the large number of potential hosts available. Unlike other parasitic taxa worldwide, New World cuckoos face significant distribution and host usage overlap with the parasitic cowbirds (Molothrus spp.). Cowbirds could be formidable competitors. They are more common than cuckoos, less specialized in host usage, and are capable of laying many more eggs per season. Local parasitism rates by cowbirds may approach 100% in some host species. In contrast, cuckoos usually occur at low density and are frequently specialized in host usage. Local parasitism rates among cuckoos’ hosts may approach 20%, but are usually below 5% (Hughes, 1997). If these general patterns of host usage existed in the early radiation of New World parasitic cuckoos, competition for host nests may have played a role in limiting their success. Hughes (1996b, 1997) provides additional discussion of evolutionary implications of Coccyzus behaviour. Facultative brood parasitism has also been recorded in Dark-billed (Coccyzus melacoryphus Vieillot; Sick, 1993) and Dwarf (C. pumilis Strickland; Ralph, 1975) cuckoos of South America. The remaining five Coccyzus species are poorly known and, consequently, parasitism has not been documented. Many aspects of Coccyzus life history are virtually unknown, and to date, there has been no large-scale examination of brood parasitism frequency in any Coccyzus species. Future work may show that brood parasitism occurs with greater frequency and success than has been considered to date. Re-evolution of terrestriality About 45 cuculid species, including the familiar Greater Roadrunner (Neomorphinae: Geococcyx californianus Lesson) of the American Southwest, are primarily terrestrial birds, which forage and nest on or near the ground. Although capable of flying, most prefer to flee from a disturbance on foot (Wyllie, 1981). Peters (1940) placed the obligate parasites Tapera and Dromococcyx in the Neomorphinae based on

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their New World distribution and terrestrial habits. He also may have been influenced by Friedmann (1933) who suggested that the Striped Cuckoo (T. naevia) developed brood parasitism independently in the New World. More recently, Berger (1960) proposed that ancestral cuculine stock brought parasitism to the Americas from the Old World. In most classifications, Tapera and Dromococcyx have been set apart from the non-parasitic, terrestrial cuckoos based on their osteology (Shufeldt, 1901; Verheyen, 1956a; Seibel, 1988), myology, syringeal morphology, and pterylosis (Beddard, 1885; Shelley, 1891; Berger, 1960), and behaviour and ecology (Hughes, 1996b). Some systematists have put Tapera and Dromococcyx within the Cuculinae, while others have placed these genera in their own subfamily (Diplopterinae: Sclater & Salvin, 1873; Shelley, 1891; or Taperinae: Verheyen, 1956a; Seibel, 1988) occupying some intermediate position between the Cuculinae and terrestrial, nonparasitic cuckoos of the Centropodinae and Neomorphinae. Friedmann, himself, commented (in litt. to K.C. Parkes, 7 August 1969) that he “would not object to the Neomorphinae being split into two—the Neomorphinae with Neomorphus, Geococcyx, and Morococcyx, and the Taperinae with Tapera and Dromococcyx” (K.C. Parkes, pers. comm.). Sibley & Ahlquist (1990) did not include either Tapera or Dromococcyx in their analysis, but merely maintained the taxonomic arrangement of Peters (1940). My study clearly supports the removal of Tapera and Dromococcyx from the Neomophinae. Reconstructing Peters’ (1940) Neomorphinae requires 82 additional steps, and reduces the optimal tree CI from 0.798 to 0.584. The Taperini differ from the Neomorphinae by 45 highly informative characters (mean CI=0.857), many of which are multistate characters that differ by more than one state. This is consistent with both the inclusion of these taxa in the Cuculinae, or the erection of a separate subfamily positioned as sister taxon to the Cuculinae to contain them. However, nine synapomorphies unite the Taperini and the other obligate parasites; therefore, I see no reason to exclude Tapera and Dromococcyx from the Cuculinae. I described only three characters, most probably linked to locomotion and foraging behaviour, which are markedly similar in the Taperini and Neomorphinae. Two convergent characters are found on the caudal end of the mandibular apparatus and are associated with M. depressor mandibulae attachment sites: (1) the U-shaped hiatus subtympanicus (character 40), and (2) the shape of the caudal surface of os articulare (character 50). The M. depressor mandibulae serves to raise and lower the mandible (George & Berger, 1966). Lowe (1938) and Berger (1957) suggested that the relative development of this muscle was strongly indicative of foraging strategies and, therefore, could not be considered a valid taxonomic character in some species. In cuckoos, this specific M. depressor mandibulae structure and its affiliated skeletal elements may be associated with ground foraging. The Neomorphinae and Taperini are among those that use a characteristic ‘chase and capture’ mode of foraging. While pursuing prey, the neck is outstretched and head and tail held horizontally to the body (Slud, 1964; Ridgely & Gwynne, 1989; Sieving, 1990; Hughes, 1996a). The third convergent character, found on the tarsometatarsus, is an enlarged tuberositas m. tibialis cranialis that is positioned immediately distal to the mesial foramina vascularia proximalia (character 69). This corresponds to the relative insertion point of the muscle tendon and is an adaptation to terrestrial locomotion (Berger, 1952). If Tapera and Dromococcyx were truly cursorial—like Geococcyx, Neomorphus, and Morococcyx—one might expect a greater similarity in skeletal elements. However, the Taperini have been poorly studied and, as such, their proclivities for terrestrial

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behaviour may be overestimated. Although they spend some time on the ground and are capable of running well, Tapera and Dromococcyx both perch, display, and vocalize at some height in the vegetation (Slud, 1964; Hilty & Brown, 1986; Stiles & Skutch, 1989; Sieving, 1990). Sternal keel depth is often used to predict flying abilities because the relative development of the keel is directly related to the development of the two major flight muscles, Mm. pectoralis and supracoracoideus, that have their origins on the keel (George & Berger, 1966). Keel depth-sternum length ratios, where a higher number characterizes a deeper sternum, indicates that Tapera (mean=0.39, n=11) and Dromococcyx (mean=0.41, n=3) have significantly deeper keels than other terrestrial cuckoos (Geococcyx, mean=0.31, n=6, P<0.003; Neomorphus, mean=0.30, n=3, P<0.002; Morococcyx, mean=0.32, n=4, P<0.005; Coua, mean=0.32, n=5, P<0.025; Centropus, mean=0.31, n=7, P<0.0004 for Tapera, P<0.001 for Dromococcyx; adult birds, sexes pooled; ANOVA). However, keel depthsternum length ratios of the Taperini are similar to the sedentary, arboreal cuckoos of the Saurotherini that fly moderately well (Saurothera, mean=0.37, n=4; Piaya, mean=0.41, n=6; Hyetornis, mean=0.39, n=4). Migratory cuculine cuckoos, such as Cuculus, Chalcites, and Coccyzus, have keel depth-sternum length ratios in excess of 0.48. Hence, Tapera and Dromococcyx may not be fully terrestrial. Slud (1964) observed Tapera flying with steady wing beats for a considerable distance at a height of about 15 metres. Wetmore (1968) noted markedly underdeveloped leg muscles in Dromococcyx. These birds have been most frequently observed while foraging on the ground and this behaviour may represent their predominant terrestrial activity. Phaenicophaeinae Peters (1940) had difficulty classifying this diverse subfamily of arboreal, nonparasitic cuckoos. However, my analysis supports its logical partitioning into four tribes that include all taxa assigned to this taxon by Shufeldt (1901), Verheyen (1956a), and Seibel (1988). Rhinortha and Ceuthmochares are anatomically distinct (Berger, 1960; Seibel, 1988). Thus, their placement in two paraphyletic taxa— Rhinothini and Ceuthmocharini—is well justified. The remaining two tribes, Old World Phaenicophini and New World Saurotherini, are associations that have not been disputed historically. My classification supports an Old World origin for the Phaenicophaeinae with a single, subsequent colonization of the New World by the ancestral saurotherine cuckoo. Some taxonomists have suggested that the malkohas—eight Old World phaenicophaeine genera, not including Ceuthmochares—should be merged into Phaenicophaeus (Delacour & Mayr, 1945; Delacour, 1946, 1947; Sibley & Monroe, 1990). The group is represented in this study by Rhinortha, Rhopodytes, Rhamphococcyx, Lepidogrammus, and Dasylophus. My results indicate that malkohas are paraphyletic, with the divergent Rhinortha positioned outside the clade containing the other four genera. In addition, within the Phaenicophini two subtribes are separated by three synapomorphies. However, it is possible that at least a few members (e.g. Rhopodytes and Rhamphococcyx) could be merged into a single genus, but this recommendation cannot be made without examining multiple specimens from all genera in question. However, due to the general lack of malkoha skeletal material, in particular Phaenicophaeus (one complete specimen) and Taccocua (none available; Wood & Schnell, 1986; and various pers. comm.), the question of appropriate allocation to genus may remain unanswered for some time.

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Centropodinae Peters (1940) likely placed the 10-species Coua in a monotypic subfamily, Couinae, because it is restricted to Madagascar. However, he recognized its association with Centropus and placed it next to the monotypic Centropodinae. Many other systematists have suggested that Coua and Centropus are close relatives, based on similarities in osteology, pterylosis, behaviour, protein assay and DNA. Consequently, they have been placed together repeatedly in higher-ranking taxa, often with other terrestrial cuckoos, such as Carpococcyx and Geococcyx (Beddard, 1885, 1898; Verheyen, 1956a; Sibley & Ahlquist, 1972; Seibel, 1988; Sibley & Monroe, 1990; Hughes, 1996b). My analysis indicates that Coua and Centropus are sister taxa that together comprise the Centropodinae. Berger (1960) thought that the anatomical divergence observable between them did not justify individual subfamily status. My examination of several Coua and Centropus skeletons supports this view. Centropus comprises 25 species of terrestrial cuckoos, known as coucals, that are distributed widely throughout Asia and Africa. Originating in Asia, and being of forest origin, individuals probably dispersed over land when a forest connection existed between the two continents (Irwin, 1985). Coua may represent a divergent lineage from ancestral African Centropus stock. Berlioz (1948) suggested that Coua, like many other endemic Madagascar birds, was originally a forest dweller that moved secondarily into more open, arid habitat types after its ancestor colonized the island from mainland Africa. Madagascar has been colonized by other African terrestrial birds that are incapable of long flights. For example, guinea-fowl (Numididae) are present on both sides of the Mozambique Channel (Dorst, 1972). Another centropodine cuckoo, the forest-dwelling Malagasy Coucal (Centropus toulou Mu¨ller), is also found on Madagascar and the Seychelles (Sibley & Monroe, 1990). Terrestriality in a plesiomorphic position Carpococcyx is a poorly known genus of two species that occurs in humid forests of southeast Asia and Indochina. Shelley (1891) placed it in the Neomorphinae due to its terrestrial habits and generalized morphological similarities to New World roadrunners (Geococcyx) and ground-cuckoos (Neomorphus). Coues (1897) argued that Carpococcyx could not be included logically in the Neomorphinae because of its Asian distribution, but its position was retained by Peters (1940). Verheyen (1956a) transferred Carpococcyx from the Neomorphinae to the Centropodidae. Seibel’s (1988) taxonomic position for Carpococcyx is less clear. Although he included it in the Geococcyginae (Neomorphinae sensu Peters, excluding Tapera and Dromococcyx) by convention, Carpococcyx also could be interpreted as sister taxon to the Neomorphinae, which is paraphyletic to Coua and Centropus. More recently, Sibley & Monroe (1990) transferred Carpococcyx to the Cuculinae based on DNA-DNA hybridization evidence of Sibley & Ahlquist (1990). However, Carpococcyx tissue was not included in their biochemical analyses; therefore, there was no data to support this change. Despite these different opinions, Carpococcyx continues to be included in the Neomorphinae sensu Peters (1940) in all other currently accepted classifications. My analysis indicates that Carpococcyx is the most plesiomorphic genus, suggesting that the ancestral cuckoo was at least partially adapted for terrestrial locomotion. My analysis of non-appendicular characters produced the same topology; hence, the basal position of terrestrial cuckoos is not a function of using the poorly flying turacos and Hoatzin to polarize characters. This placement of terrestrial cuckoos

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was corroborated by Verheyen (1956a, 1961) and Seibel (1988), and in part by Hughes (1996b). In addition, Shufeldt (1909: 356) commented that many typical cuculine characters were expressed more strongly in terrestrial types and less so in the arboreal Cuculinae. He proposed that the ancestral form was terrestrial, and suggested that Geococcyx may be a “modern, highly specialized representative of the ancestral stock”. Based on metacarpus structure, Stegmann (1978: 50–51) concluded that the “round-winged, forest-dwelling” cuculid form is plesiomorphic and shows greater affinities to turacos. Van Tyne & Berger (1959) argued that terrestriality could not be plesiomorphic among cuckoos because the zygodactyl foot is considered an adaptation to perching and climbing, not cursorial locomotion. Many other zygodactyl groups, such as woodpeckers (Picidae), parrots (Psittaciformes), and toucans (Ramphastidae), are predominantly arboreal. However, this statement disregards the role of phylogenetic constraint in explaining zygodactyly in cuckoos. Although the ordinal relationships of birds are poorly known, cuckoos have been commonly associated with other zygodactylous non-passerine taxa, such as turacos (Musophagidae; semi-zygodactyl), mousebirds (Coliiformes), and parrots (Psittaciformes). Therefore, it is reasonable to assume that cuckoos inherited zygodactyly from an ancestor. Turacos and the Hoatzin—putative sister taxa to cuckoos—exhibit an intermediate condition between true arboreal and terrestrial locomotion. Although both are primarily arboreal, they have a slow and laborious flight (Stegmann, 1978). Their locomotory habits have been compared to those of some terrestrial-type cuckoos, such as the neomorphine Morococcyx and several species of Coua, that walk in a dovelike fashion along branches (Berger, 1960). Other cuckoos, such as the malkohas, Saurotherini, and Crotophaginae, also exhibit intermediate locomotory habits (Berger, 1960). Strong flying ability is a derived characteristic found predominantly among the Cuculinae. Cuckoo origins The Old World origin of the Cuculidae has been generally accepted because the Musophagidae occur in Africa, and most cuckoo species are found in Old World tropical and subtropical regions (Berger, 1960; Cracraft, 1973). The Asian distribution of Carpococcyx is consistent with the subsequent divergence of early, forest-dwelling centropodine cuckoos that eventually gave rise to Centropus and Coua. My phylogeny implies that the New World was colonized by an ancestral terrestrial cuckoo derived from Old World stock, establishing a lineage that eventually gave rise to Morococcyx, Geococcyx, and Neomorphus. Cuckoos are an ancient family with a scanty fossil record dating from the Upper Eocene of France (Dynamopterus velox; Milne-Edwards, 1892). Two early New World cuckoo fossils are known from the Lower Oligocene of Saskatchewan (Neococcyx mccorquodalei; Weigel, 1963) and Lower Miocene of Colorado (Cursoricoccyx geraldinae; Martin & Mengel, 1984). The latter was assigned to the Neomorphinae, indicating that terrestrial cuckoos have been in North America for at least 20 Myr. Cursoricoccyx was found near the northern range limit of the most northerly neomorphine cuckoo, the Greater Roadrunner (Hughes, 1996a). This implies that Cursoricoccyx may have had a more northerly distribution than its extant relatives. Other neomorphine species range from Mexico to central South America (Sibley & Monroe, 1990). The existence of terrestrial cuckoos in the New World 20 Myr ago suggests that Old

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World terrestrial stock could have dispersed to North America by the Bering land bridge, which connected Asia and North America prior to the Miocene (McKenna, 1983). Neomorphine cuckoos may have been broadly distributed at northern latitudes prior to the late Tertiary climatic deterioration (Cracraft, 1973), with the northern boundary of their range being continually pushed southward with the onset of cooler temperatures. The distribution of extant roadrunners is limited by persistent snow cover as it impairs their ability to forage effectively for reptiles and arthropods (Norris & Elder, 1982). More recent events in the biogeographic history of cuckoos are difficult to diagnose. Cracraft (1973) suggested that the near global distribution of cuckoos could only be explained by several Northern and Southern Hemisphere dispersal events. This statement is supported by my phylogeny that requires at least five major intercontinental movements to explain the current distribution of the family. However, given the age of the Cuculidae and the propensity of some members for long-distance flight, this many dispersal events does not seem unreasonable. Hoatzin Perhaps no species has proven to be a greater systematic enigma than the Hoatzin. Since first described as Phasianus hoazin by Mu¨ller (1776), this species has been allied with Galliformes in 17, turacos in four, and cuckoos in eight classifications. In addition, it has been placed in the monotypic order, Opisthocomiformes, 12 times (see Sibley & Ahlquist, 1973, 1990 for a review). Peters (1934) considered the Hoatzin to be an aberrant galliform and placed it in the monotypic family Opisthocomidae (Galliformes). Despite much evidence to the contrary, this position has persisted in many current taxonomies. Since Peters’ (1934) work, several studies have supported the alliance of the Hoatzin with cuculiforms (=Cuculidae and Musophagidae) based on similarities in osteology (Verheyen, 1956b; De Queiroz & Good, 1988), and myology (Stegmann, 1978). Other workers used mitochondrial (Mindell et al., 1997) and nuclear gene sequences (Hedges et al., 1995) to suggest more specifically its association with cuckoos. Furthermore, protein electrophoresis and DNA-DNA hybridization evidence of Sibley & Ahlquist (1972, 1973, 1990) placed the Hoatzin within the Cuculidae, most closely allied with roadrunners (Geococcyx) and anis (Crotophaginae). However, Hughes & Baker (1999) demonstrated that the Hoatzin is sister to turacos, not cuckoos. The present study corroborates Hughes & Baker (1999) in the suggestion that turacos and the Hoatzin are sister taxa. Similarities in pterylography and internal anatomy between the Hoatzin and turacos were first described by Nitzsch (1829, 1840). Verheyen (1956b) listed 50 osteological characters that linked these taxa. In addition, Stegmann (1978) noted that both young turacos and hoatzins use their wings and claws of digits II and III for climbing among branches of the nesting tree long before their flight feathers have fully developed. Both taxa share a characteristic growth retardation of the outer primaries that facilitates this form of locomotion. He added that, although wing and associated structures of the Hoatzin most resemble those of cuckoos, if the taxonomic importance of these characters and the peculiarity of their ontogenetic development are considered, the Hoatzin should be allied with the Musophagidae. Interestingly, elements of both taxa combine in the Lower Eocene fossil Foro

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panarium of Wyoming. This species has a skull and mandible most like the Hoatzin, but shows some similarities to turacos in postcranial skeletal elements. Its elongated hind limb elements, particularly the tarsometatarsus, “suggest a more terrestrial mode of life than the modern species of Musophagidae or Opisthocomidae, perhaps not unlike some of the terrestrial Cuculidae” (Olson, 1992: 127). This does not imply that Foro panarium represents the ancestor of a Hoatzin-turaco-cuckoo radiation, because there are other Eocene fossils more closely associated with extant taxa (Milne-Edwards, 1892; Cracraft, 1971). Rather, it indicates the existence of a lineage of primitive, generalized, terrestrial non-passerines with which these taxa may have shared an ancestor. Cuculiformes The relationship between cuckoos and turacos was first recognized by Linnaeus (1758), but it was not until Fu¨rbringer (1888) and Shufeldt (1904) combined them in Coccygiformes that the association became widely accepted. Peters (1940) continued the tradition but renamed the families Cuculidae and Musophagidae, and the order Cuculiformes. This nomenclature remains currently in use. However, many systematists have noted significant anatomical and biochemical differences between these two avian families. Lowe (1943, after Bannerman, 1933) and Berger (1960) concluded that the Cuculidae and Musophagidae could not be placed in the same order based on their osteology and myology. Lowe also noted that feather tracts, which are generally consistent and characteristic within orders including Passeriformes, differ in cuckoos and turacos. Furthermore, Van Tuinen & Valentine (1986) suggested that differences in chromosome banding patterns set turacos apart from all other avian orders. Sibley & Ahlquist (1990) placed cuckoos (Cuculiformes) and turacos (Musophagiformes) in separate, non-sister orders in Parvclass Passerae with several other non-passerine orders and Passeriformes. In contrast, Stegmann (1978) noted different carpometacarpus structure in cuckoos and turacos, but added that similarities in wing myology suggested that they descended from a common ancestor after evolving independently over a long time. He concluded that they should be placed in the same order to recognize their “mutual relationships” (p. 51). Seibel (1988) tentatively upheld the inclusion of the Musophagidae in the Cuculiformes based on synapomorphies of the os carpi ulnare. My study corroborates Seibel (1988) in uncovering synapomorphies of the os carpi ulnare (characters 121, 122, and 123) that unite cuckoos, turacos, and the Hoatzin. Monophyly was also implied by Hughes & Baker (1999). Nevertheless, these three taxa are otherwise too diverse anatomically to be accommodated in the same order. Therefore, by convention, cuckoos should be classified in the monotypic order Cuculiformes, and turacos and the Hoatzin placed together in the order Opisthocomiformes—which has nomenclatural priority over Musophagiformes— adjacent to the Cuculiformes.

ACKNOWLEDGEMENTS

I thank G. F. Barrowclough, C. Blake, and P. Sweet (American Museum of Natural History), R. M. Zink and J. T. Klicka (Bell Museum of Natural History), B. Livezey and R. Panza (Carnegie Museum of Natural History), D. E. Willard and

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J. Bates (Field Museum of Natural History), T. Webber and D. Steadman (Florida Museum of Natural History), J. V. Remsen and D. Dittman (Louisiana State University Museum of Zoology), J. Dick and G. Murphy (Royal Ontario Museum), R. C. Banks and R. Browning (United States National Museum of Natural History), C. Cicero and N. K. Johnson (University of California Museum of Vertebrate Zoology), and R. B. Payne, L. Payne, and J. Hinshaw (University of Michigan Museum of Zoology) for their invaluable assistance in providing the many specimens required for this study. I also thank J. C. Barlow for his help. Finally, my sincere thanks to my brother, M. E. Hughes, for providing the illustrations. This work was supported by a Natural Sciences and Engineering Research Council fellowship. In addition, aspects of this work were funded by a NSERC grant (A3472) to J. C. Barlow.

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Sibley CG, Ahlquist JE. 1972. A comparative study of the egg white proteins of non-passerine birds. Peabody Museum of Natural History Bulletin 39. Sibley CG, Ahlquist JE. 1973. The relationships of the Hoatzin. Auk 90: 1–13. Sibley CG, Ahlquist JE. 1990. Phylogeny and classification of birds: a study in molecular evolution. New Haven: Yale University Press. Sibley CG, Monroe BL Jr. 1990. Distribution and taxonomy of birds of the world. New Haven: Yale University Press. Sick H. 1993. Birds in Brazil. Princeton: Princeton University Press. Sieving KE. 1990. Pheasant Cuckoo foraging behavior, with notes on habits and possible social organization in Panama. Journal of Field Ornithology 61: 41–46. Slud P. 1964. The birds of Costa Rica. Bulletin of the American Museum of Natural History 128. Stegmann BC. 1978. Relationships of the superorders Alectoromorphae and Charadriomorphae (Aves): a comparative study of the avian hand. In: Paynter JRA, ed. Nuttall Ornithological Club Publication 17. Stiles FG, Skutch AF. 1989. A guide to the birds of Costa Rica. Ithaca: Cornell University Press. Strahl SD. 1987. The social organization and behavior of the Hoatzin Opisthocomus hoazin in central Venezuela. Ibis 130: 483–502. Stresemann E. 1934. Aves. In: Ku¨kenthal W, Krumbach T, eds. Handbuch der Zoologie. Volume 7. Part 2. Berlin: Walter de Gruyter. Stresemann E. 1959. The status of avian systematics and its unsolved problems. Auk 76: 269–280. Swainson W. 1837. On the natural history and classification of birds. Volume 2. London: Longman, Rees, Orme, Brown, Green, and Longman. Swofford DL. 1998. PAUP 4.0: Phylogenetic analysis using parsimony and other methods. Sunderland: Sinauer. Van Tuinen P, Valentine M. 1986. Phylogenetic relationships of turacos (Musophagidae; Cuculiformes) based on comparative chromosome banding analysis. Ibis 128: 364–381. Van Tyne J, Berger AJ. 1959. Fundamentals of ornithology. New York: J. Wiley. Verheyen R. 1956a. Contribution a` l’anatomie et a` la syste´matique des touracos (Musophagi) et des coucous (Cuculiformes). Bulletin de Institut Royal de Sciences Naturelles de Belgique 32 (23): 1–28. Verheyen R. 1956b. Note syste´matique sur Opisthocomus hoazin (St. Mu¨ller). Bulletin de Institut Royal de Sciences Naturelles de Belgique 32 (32): 1–8. Verheyen R. 1961. A new classification for the non-passerine birds of the world. Bulletin de Institut Royal de Sciences Naturelles de Belgique 37 (32): 1–8. Vigors NA. 1825. Observations on the natural affinities that connect the orders and families of birds. Transactions of the Linnean Society of London 14: 395–562. Wagler JG. 1830. Natu¨rliches System der Amphibien mit vorangehender Classification der Sa¨ugethiere und Vo¨gel. Munich: J. G. Cotta. Weigel RD. 1963. Oligocene birds from Saskatchewan. Quarterly Journal of the Florida Academy of Sciences 26: 257–262. Wetmore A. 1960. A classification for the birds of the world. Smithsonian Miscellaneous Collections 139 (11): 1–37. Wetmore A. 1968. The birds of the Republic of Panama. Part 2, Columbidae (pigeons) to Picidae (woodpeckers). Smithsonian Miscellaneous Collections, No. 150. Wiley EO. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. New York: J. Wiley & Sons. Witherby HF. 1938. The handbook of British birds. Vol. 2, warblers to owls. London: H.F. & G. Witherby, Ltd. Wolfe DH. 1994. Yellow-billed Cuckoo hatched in Mourning Dove nest. Bulletin of the Oklahoma Ornithological Society 27: 29–30. Wood DS, Schnell GD. 1986. Revised world inventory of avian skeletal specimens. Norman: American Ornithologists’ Union and Oklahoma Biological Survey. Wyllie I. 1981. The cuckoo. New York: Universe Books.

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Characters purportedly diagnostic of the Cuculidae This list is not exhaustive, but includes most characters used previously to define the Cuculidae. However, many represent questionably homologous conditions that are not synapomorphic at the family level and cannot be used to support monophyly. From Beddard (1885, 1898), Shelley (1891), Gadow (1892), Pycraft (1903), Witherby (1938), Sibley (1955), Van Tyne & Berger (1959), Berger (1960), Sibley & Ahlquist (1972, 1990), Stegmann (1978), Gilbert, Martin & Savage (1981), and Seibel (1988). (1) Os lacrimale cuculine; (2) Os uncinatum absent; (3) Holorhinal; (4) Nares more or less impervious; (5) Tomia smooth; (6) Processes basipterygoideus absent or rudimentary; (7) Desmognathous palate; (8) Vomer small; (9) Margo medialis of os palantinum meeting in plana medianus below, and concealing, rostrum sphenoidale; (10) Incisura intercondylaris of tibiotarsus a rounded pit extending caudally at least half of length of extremitas distalis; (11) Two bony canales hypotarsi, subequal in size, with proximal openings side by side in planum medianum; (12) Tuberositas m. tibialis anticus positioned immediately distal to foramina vascularia proximalia; (13) Mesial ‘wing’ on trochlea metatarsi secondii not larger than lateral ‘wing’; (14) Presence of protuberance on facies lateralis of trochlea metatarsi tertii; (15) Trochlea metatarsi quartii of tarsometatarsus with mesially inflected caudal trochlea accessoria, and with margo distalis proximal to incisura intertrochlearis medialis; (16) Margo medialis of ala preacetabularis ilii not meeting above crista iliaca dorsalis in planum medianus; (17) Margo cranialis of apex carinae not continued cranially as far as free end of spina externa; (18) Sternum usually with two notches on each side, one sometimes lost or closed to form fenestra; (19) Four ribs reaching sternum; (20) Coracoideum not fused or ankylosed; (1) Processus procoracoideus not fused with processus acrocoracoideus; (22) Facies articularis sternalis ventralis of coracoideum centred, rounded, and facing sternally; (23) Facies articularis sternalis ventralis of coracoideum large, facies articularis sternalis dorsalis small to large; (24) Clavicula without distinct facies articularis acrocoracoideus; (25) Clavicula with distinct ‘procoracoidal facet’ sensu Seibel; (26) Apophysis furculae present; (27) Os metacarpale minus almost as broad in planum transversus as os metacarpale majus; (28) Atlas perforated; (29) 13 to 14 cervical vertebrae; (30) Four vertebrae thoracica; (31) 17 to 18 vertebrae; (32) Margo lateralis of vertebrae thoracica not produced into spikes.

APPENDIX 2

Osteological character descriptions Zero (0)=ancestral condition. Multistate characters are unordered unless marked as ordered. Consistency indices (CI) are calculated for both cuckoos only, and cuckoos and outgroup (Musophagidae), respectively. NC=no change in ingroup. Abbreviations: m., musculus; n., nervus, nervi; proc., processus, processes; sync., synchondrosis, synchondroses.

Cranial characters 1. Os cranii, size of fossa temporalis: (0) small to moderate; (1) large, meeting, or nearly meeting sagitally. CI=0.50; 0.50. 2. Os cranii, fissura cranio-facialis: (0) well defined; (1) moderately to poorly defined. CI=0.50; 0.50. 3. Os frontale, size of proc. orbitalis (ordered): (0) large; (1) small to moderate; (2) vestigial. CI= 1.00; 1.00. 4. Os frontale, shape of margo supraorbitalis (ordered): (0) not flared; (1) slightly flared; (2) strongly flared. CI=0.50; 0.50. 5. Os squamosum, size of proc. postorbitalis: (0) large; (1) moderate; (2) small. CI=0.67; 0.50. 6. Os squamosum, size of proc. suprameaticus relative to proc. postorbitalis: (0) small; (1) large. CI=1.00; 1.00. 7. Os squamosum, shape of proc. suprameaticus: (0) triangular, flattened in planum medianum; (1) bulbous in dorsal aspect. CI=1.00; 1.00.

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8. Os squamosum, length of proc. suprameaticus: (0) short; does not obscure head of proc. oticus quadrati; (1) long; obscures head of proc. oticus quadrati. CI=1.00; 1.00. 9. Os lacrimale, extent of articulation with os frontale: (0) large; (1) small to moderate. CI=NC; 1.00. 10. Os lacrimale, width of distal end of proc. orbitalis: (0) wide to moderate; (1) narrow. CI=0.50; 0.50. 11. Os lacrimale, shape of proc. supraorbitalis: (0) ovoid, triangular, or rod-shaped; (1) semi-lunar, margo dorsalis well caudal of fissura cranio-dorsalis and sutured to lateral extensions of os frontale. CI=1.00; 1.00. 12. Os lacrimale, shape of proc. supraorbitalis: (0) not as in state 1; (1) triangular, directed laterally, margo dorsalis horizontal, not curved. CI=1.00; 1.00. 13. Os lacrimale, shape of proc. supraorbitalis: (0) not as in state 1; (1) triangular, directed caudolaterally, dorsal margin curved, not horizontal. CI=1.00; 1.00. 14. Os lacrimale, shape of proc. supraorbitalis: (0) ovoid, triangular, or semi-lunar; (1) rod-shaped, wide in axis rostrocaudalis, positioned in V-shaped notch formed by os frontale and os nasale. CI=1.00; 1.00. 15. Os ectethmoidale, as in monophyly character 3 (see Results of above monophyly analysis): (0) no; (1) yes. CI=NC; 1.00. 16. Os ectethmoidale, generalized form: (0) triangular, margo ventralis horizontal; (1) not as in state 0. CI=NC; 1.00. 17. Os ectethmoidale, margo lateralis: (0) straight or concave; (1) convex. CI=1.00; 0.50. 18. Os ectethmoidale, margo lateralis: (0) not as in state 1; (1) prominent, bulbous protrusion directed ventrally. CI=1.00; 1.00. 19. Os ectethmoidale, margo lateralis: (0) not as in state 1; (1) straight distally, strongly excised proximally. CI=1.00; 1.00. 20. Os jugale, point of articulation of distal end: (0) juncture of os premaxilla and os nasale; (1) dorsal to juncture. CI=NC; 1.00. 21. Os quadratum, as in monophyly character 1: (0) no; (1) yes. CI=NC; 1.00. 22. Os quadratum, shape of proc. orbitalis: (0) blunt; (1) strongly pointed. CI=1.00; 1.00. 23. Os quadratum, shape of proc. orbitalis: (0) not as in state 1; (1) narrow dorso-ventrally, elongate. CI=1.00; 1.00. 24. Os quadratum, angle of proc. oticus and proc. orbitalis in lateral aspect (ordered): (0) large; (1) moderate; (2) small. CI=0.67; 0.67. 25. Os quadratum, mesial inflection of proc. orbitalis (ordered): (0) none or little; (1) moderate; (2) strong. CI=0.67; 0.67. 26. Os quadratum, proc. mandibularis, relative sizes of condylus caudalis and condylus pterygoidus: (0) condylus medialis slightly larger than condylus pterygoidus; (1) condylus medialis much larger than condylus pterygoidus. CI=1.00; 1.00. 27. Os quadratum, inflection of distal end of condylus caudalis in lateral aspect: (0) not as in state 1; (1) pointed dorsally. CI=1.00; 0.50. 28. Os quadratum, inflection of distal end of condylus medialis in lateral aspect: (0) not as in state 1; (1) pointed strongly ventrally. CI=1.00; 1.00. 29. Os quadratum, expansion of distal end of cotyla quadrojugalis (ordered): (0) more than two times wider than neck of proc. quadratojugalis; (1) approximately two times wider; (2) less than two times wider. CI=1.00; 1.00. 30. Cavitas tympanica, position of canalis nervus facialis: (0) at same level caudo-rostrally as meatus acousticus externas; (1) caudal to meatus acousticus externas. CI=NC; 1.00. 31. Cavitas tympanica, shape of otic region defined by ala tympanica and proc. oticus quadrati in lateral aspect: (0) short oval; (1) long oval. CI=1.00; 1.00. 32. Cavitas tympanica, otic region as above, angle of longest chord: (0) 15 to 45°; (1) approximately 90°; (2) 45 to 90°. CI=0.50; 0.50. 33. Os basioccipitale, form of crista basilaris transversa: (0) poorly defined; (1) well defined. CI= 0.50; 0.50. 34. Os basioccipitale, form of crista basilaris transversa: (0) separated at midpoint rostral to condylus occipitalis; (1) entire. CI=0.67; 0.67. 35. Os basioccipitale, position of crista basilaris transversa: (0) caudal or at same level caudo-rostrally as ostium canalis carotici; (1) rostral to ostium canalis carotici. CI=0.67; 0.67. 36. Os basisphenoidale, bar-like process between ostium canalis carotici and lamina basiparasphenoidalis: (0) absent; (1) present. CI=1.00; 1.00.

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37. Os exoccipitale, shape (ordered); (0) flat or only slightly raised; (1) produced into bulbous expansion raised above contour of os occipitale; (2) as in state 1, but greatly expanded. CI= 1.00; 1.00. 38. Os exoccipitale, position of foramen n. vagi relative to ostium canalis ophthalmic externi: (0) caudal or at same level in axis rostrocaudalis as ostium canalis ophthalmic externi; (1) rostral to ostium canalis opthalmic externi. CI=1.00; 1.00. 39. Os exoccipitale, size of hiatus subtympanicus: (0) large; (1) small. CI=1.00; 0.50. 40. Os exoccipitale, shape of hiatus subtympanicus; (0) V-shaped; (1) widened U-shape. CI=0.50; 0.50. 41. Os parasphenoidale, rostrum parasphenoidale: (0) exposed; (1) obscured. CI=1.00; 1.00. 42. Os parasphenoidale, proc. basipterygoideus: (0) absent; (1) present. CI=1.00; 1.00. 43. Os parasphenoidale, size of proc. medialis parasphenoidalis: (0) vestigial; (1) absent. CI=NC; 1.00. 44. Os pterygoideum, as in monophyly character 2: (0) no; (1) yes. CI=NC; 1.00. 45. Os pterygoideum, facies ventralis of rostral end: (0) flanged; (1) not flanged. CI=1.00; 1.00. 46. Os pterygoideum, shape of facies articularis sphenoidalis in ventral aspect: (0) not as in state 1; (1) flared with rostrally projecting spurs. CI=1.00; 1.00. 47. Os palatinum, as in monophyly character 4: (0) no; (1) yes. CI=NC; 1.00. 48. Os maxillare, as in monophyly character 5: (0) no; (1) yes. CI=NC; 1.00. 49. Os maxillare, caudal extremes of proc. palatinus in ventral aspect: (0) enclosed within medial margins of os palatine, visible; (1) displaced laterally, obscured by os palatine. CI=1.00; 1.00. 50. Os articulare, shape of cotylae lateralis in dorsal aspect: (0) not as in state 1; (1) pointed with appearance of two cotylae, upper rounded, lower pointed and deflected latero-caudally. CI= 0.50; 0.50. 51. Os articulare, shape of caudal surface: (0) semi-circular; (1) U-shaped. CI=0.50; 0.50. 52. Os articulare, size of caudal surface in plana dorsalis (ordered): (0) deep; (1) moderate; (2) shallow. CI=0.67; 0.50. 53. Os articulare, tuberculum pseudotemporale: (0) small or vestigial; (1) prominent. CI=0.50; 0.50. 54. Os articulare, crista intercotylaris, as in monophyly character 6: (0) no; (1) yes. CI=NC; 1.00. 55. Apparatus hyobranchialis, shape of os basibranchiale rostrale in lateral aspect: (0) margo ventralis straight; (1) prominent protrusion near midpoint of margo ventralis. CI=1.00; 1.00. 56. Apparatus hyobranchialis, relative width of distal and proximal end of os basibranchiale in lateral aspect: (0) similar; (1) distal end wider than proximal. CI=1.00; 1.00.

Postcranial characters Numbers of characters based on Seibel (1988) are given in parentheses. Revised characters are based partially on those of Seibel (1988) but have been recoded, split into several characters, extended to include more character states, or otherwise modified from the original character description. 57. Tibiotarsus, form of incisura intercondylaris, as in monophyly character 7: (0) no; (1) yes. CI= NC; 1.00. 58. Tarsometatarsus, position of canales hypotarsi, as in monophyly character 8: (0) no; (1) yes. CI= NC; 1.00. 59. Tarsometatarsus, number of canales hypotarsi: (0) one; (1) two (Seibel, 1988: character TM 13, revised). CI=NC; 1.00. 60. Tarsometatarsus, relative size of canales hypotarsi: (0) only one canal; (1) two canales equal, or almost equal in size; (2) canales medialis much larger than canales lateralis (Seibel, 1988: TM 13, revised). CI=1.00; 1.00. 61. Tarsometatarsus, size of cristae hypotarsi: (0) moderate to large; (1) small (Seibel, 1988: TM 13, revised). CI=1.00; 1.00. 62. Tarsometatarsus, direction of deflection of crista medialis hypotarsi: (0) caudally; (1) laterally (Seibel, 1988: TM 13, revised). CI=1.00; 1.00. 63. Tarsometatarsus, shape of crista hypotarsi in lateral view: (0) broadly curved; (1) flattened. CI= NC; 1.00. 64. Tarsometatarsus, shape of lateral surface: (0) broad in planum medianum, facies lateralis flattened; (1) flat, narrow throughout length (Seibel, 1988: TM 16). CI=1.00; 1.00.

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65. Tarsometatarsus, degree of torsion of shaft relative to cotylae (ordered): (0) none; (1) moderate; (2) high torsion (Seibel, 1988: TM 17, revised). CI=1.00; 1.00. 66. Tarsometatarsus, relative position of foramina vascularia proximalia in axis proximodistalis: (0) foramina lateralis proximal to foramina medialis; (1) not as in state 0 (Seibel, 1988: TM 26, revised). CI=1.00; 0.50. 67. Tarsometatarsus, relative position of foramina vascularia proximalia in axis proximodistalis (ordered): (0) foramina medialis not proximal to foramina lateralis; (1) slightly proximal; (2) highly proximal (Seibel, 1988: TM 26, revised). CI=0.67; 0.67. 68. Tarsometatarsus, position of tuberositas m. tibialis cranialis: (0) distal to foramina vascularia proximalia medialis only; (1) borders margo distalis of both foramina (Seibel, 1988: TM 27, revised). CI=0.50; 0.50. 69. Tarsometatarsus, position of tuberositas m. tibialis cranialis: (0) tuberositas immediately distal to foramina; (1) distinct gap between tuberositas and foramina (Seibel, 1988: TM 28, revised). CI= 0.50; 0.50. 70. Tarsometatarsus, form of sulcus between tuberositas m. tibialis cranialis and foramina vascularia proximalia medialis (ordered): (0) no sulcus; (1) shallow; (2) distinct; (3) deep (Seibel, 1988: TM 29, revised). CI=1.00; 1.00. 71. Tarsometatarsus, protuberance on facies lateralis of trochlea metatarsi tertii: (0) absent; (1) present (Seibel, 1988: TM 20, revised). CI=NC; 1.00. 72. Tarsometatarsus, depression on facies lateralis of trochlea metatarsi tertii: (0) absent; (1) present (Seibel, 1988: TM 20, revised). CI=1.00; 1.00. 73. Tarsometatarsus, size of ‘wings’ of trochlea metatarsi secundii: (0) mesial wing much larger than lateral wing; (1) wings equal in size or indiscernible (Seibel, 1988: TM 24). CI=NC; 1.00. 74. Tarsometatarsus, position of facies distalis of trochlea metatarsi quarti (ordered): (0) distal to incisura intertrochlearis medialis; (1) proximal to incisura; (2) highly proximal to incisura (Seibel, 1988: TM 21, revised). CI=1.00; 1.00. 75. Tarsometatarsus, inflection of trochlea metatarsi quarti, as in monophyly character 9: (0) no; (1) yes. CI=NC; 1.00. 76. Tarsometatarsus, contour of facies distalis of trochlea metatarsi quarti (ordered): (0) straight transverse; (1) curved proximally; (2) notched (Seibel, 1988: TM 25, revised). CI=1.00; 1.00. 77. Femur, extremitas proximalis femoris, pneumatic foramina caudalis: (0) absent or small; (1) large (Seibel, 1988: FE 2, revised). CI=1.00; 1.00. 78. Femur orientation of fovea ligamentum capitis: (0) faces mesially or proximo-mesially; (1) faces proximally (Seibel, 1988: FE 3, revised). CI=1.00; 1.00. 79. Femur, contour of margo lateralis of trochanter femoris: (0) distinct proximal expansion; (1) no distinct proximal expansion (Seibel, 1988: FE 4, revised). CI=NC; 1.00. 80. Femur, size of crista trochanteris (ordered): (0) large, square; (1) moderate, flap-like; (2) small (Seibel, 1988: FE 4, revised). CI=0.67; 0.67. 81. Pelvis, relative widths of facies dorsalis (ordered): (0) narrowest point less than half the width between processes antitrochantericus; (1) approximately half; (2) more than half (Seibel, 1988: PE 15, revised). CI=0.67; 0.67. 82. Ilium, form and degree of fusion of crista iliaca dorsalis and margo medialis of ala preacetabularis ilii (ordered): (0) crista and margo medialis fused; (1) crista contact margo medialis without fusing, or have bridges of bone that approach or anklylose; (2) no contact, ridges straight, tall, and close to margo medialis; (3) as in 2, but ridges lower and widely spaced (Seibel, 1988: PE 18, revised). CI=0.50; 0.50. 83. Ilium, form of anterior bridge connecting crista iliaca dorsalis and margo medialis of ala preacetabularis ilii: (0) not as in state 1; (1) broad, ankylosed to median ridge; fossa iliaca dorsalis diamond shaped in cranial aspect (Seibel, 1988: PE 22). CI=1.00; 1.00 84. Ilium, contour of crista iliaca dorsalis in dorsal aspect (ordered): (0) not as in state 1 or 2; (1) straight or curving slightly mesially at margo caudalis; (2) crista remains free farther caudally, curving laterally at margo caudalis (Seibel, 1988: PE 23). CI=1.00; 1.00. 85. Ilium, contour and orientation of margo cranialis of crista dorsolateralis ilii immediately lateral to tip of proc. antitrochantericus: (0) broadly curved, directed obliquely cranio-caudally; (1) straight, directed obliquely meso-laterally (Seibel, 1988: PE 21). CI=1.00; 1.00. 86. Ilium, size of tuberculum preacetabulare (ordered): (0) moderate to large; (1) small; (2) vestigial (Seibel, 1988: PE 5, revised). CI=0.60; 0.60. 87. Ilium, form of tuberculum preacetabulare: (0) not as in state 1; (1) short, blunt, broad in axis dorsoventralis (Seibel, 1988: PE 19). CI=1.00; 1.00.

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J. M. HUGHES

88. Ilium, cross-section shape of margo medialis (Fig. 15, section A) of crista iliaca dorsolateralis in axis dorsoventralis: (0) broad, forming smooth arc; (1) compressed into thin, blade-like shelf (Seibel, 1988: PE 10). CI=1.00; 1.00. 89. Ilium, posterior extent of spina dorsolateralis ilii (ordered): (0) very long; (1) moderate; (2) short (Seibel, 1988: PE 12, revised). CI=0.75; 0.75. 90. Ilium, relative lengths of sections B and C (Fig. 15) of crista dorsolateralis ilii (ordered): (0) B longer than section C; (1) broadly rounded, no clear distinction; (2) B and C same length; (3) B shorter than C (Seibel, 1988: PE 13, revised). CI=1.00; 1.00. 91. Ilium, contour of margo lateralis of crista dorsolateralis ilii, relative to tip of proc. antitrochantericus: (0) not as in state 1; (1) margo lateralis and antitrochantericus meeting, margo lateralis straight to slightly convex (Seibel, 1988: PE 17, revised). CI=1.00; 1.00. 92. Ilium, contour and cranio-caudal position of crista dorsolateralis ilii: (0) margo lateralis of crista displaced caudally relative to proc. antitrochantericus forming a gap, margo lateralis straight to concave; (1) intermediate between state 0 and 2; (2) as in state 0, but margo lateralis of crista immediately caudal to the proc. antitrochantericus convex; (3) not as in state 0, 1, or 2 (Seibel, 1988: PE 24, revised). CI=1.00; 0.75. 93. Ilium, relative length of sync. ilioischiadica from crista caudalis fossa renalis to proc. marginis caudalis in axis rostrocaudalis, with reference to depth of recessus caudalis fossae (ordered): (0) sync. ilioischiadica long, recessus caudalis fossae deep; (1) intermediate between state 0 and 2; (2) sync. ilioischiadica short, recessus caudalis fossae shallow (Seibel, 1988: PE 20, revised). CI= 0.67; 0.67. 94. Os coxae, foramen obturatum: (0) margo ventralis closed by bone, ischium and pubis ankylosed; (1) closed, but ischium and pubis not ankylosed; (2) margo ventralis not closed by bone (Seibel, 1988: PE 8, revised). CI=1.00; 0.50. 95. Sternum, cranial extent of apex carinae (ordered): (0) cranial tip of apex caudal to labrum externum of sulcus articularis coracoideus; (1) apex at same cranio-caudal level as sulcus; (2) apex cranial to sulcus (Seibel, 1988: ST 5, revised). CI=0.50; 0.60. 96. Sternum, form and fusion of rostrum sterni: (0) spina externa only; (1) spina interna and externa present, separate; (2) both present, fused, spina interna protrudes; (3) both present, fused, spina externa protrudes; (4) both present, fused, and widened in axis dorso-ventralis (Seibel, 1988: ST 2, revised). CI=0.80; 0.80. 97. Sternum, size of spina externa: (0) prominent; (1) greatly reduced. CI=1.00; 1.00. 98. Sternum, shape of spina interna: (0) not as in state 1; (1) wide, flattened in axis dorso-ventralis. CI=1.00; 1.00. 99. Coracoideum, form and inflection of facies articularis sternalis, as in monophyly character 10: (0) no; (1) yes. CI=NC; 1.00. 100. Coracoideum, relative size of facies articularis sternalis: (0) dorsal large, ventral small; (1) dorsal small, ventral large; (2) both small, equal in size; (3) both small, but dorsal larger; (4) both large (Seibel, 1988: CO 19, revised). CI=1.00; 1.00. 101. Coracoideum, size of proc. procoracoideus: (0) large; (1) small. CI=NC; 1.00. 102. Coracoideum, direction of inflection of proximal tip of proc. acrocoracoideus: (0) dorso-laterally; (1) dorso-mesially (Seibel, 1988: CO 20, revised). CI=1.00; 1.00. 103. Coracoideum, size and form of ‘brachial tuberosity’ sensu Howard (1929) of proc. acrocoracoideus (ordered): (0) small or absent; (1) moderate, inflected meso-ventrally; (2) large, strongly inflected (Seibel, 1988: CO 20, revised). CI=1.00; 1.00. 104. Coracoideum, shape of distal extreme of cotyla scapularis: (0) blunt or rounded; (1) sharply pointed, elongate. CI=1.00; 1.00. 105. Clavicula, facies articularis procoracoideus: (0) absent; (1) present, distinct; (2) present, indistinct (Seibel, 1988: FU 1, revised). CI=1.00; 1.00. 106. Clavicula, orientation of facies articularis acrocoracoideus (ordered): (0) dorso-laterally to dorsally; (1) latero-dorsally to laterally; (2) ventro-laterally (Seibel, 1988: FU 2, revised). CI=1.00; 1.00. 107. Scapula, as in monophyly character 11: (0) no; (1) yes. CI=NC; 1.00. 108. Scapula, extremitas cranialis scapulae, contour of margo dorsalis and ventralis: (0) obscured by fusion, or not as in other states; (1) margo dorsalis straight from mesial of facies articularis humeralis to acromion, margo ventralis with abrupt angle between facies articularis humeralis and acromion; (2) intermediate between state 1 and 3; (3) both margo dorsalis and ventralis almost straight from facies articularis humeralis to acromion where both bend abruptly dorsally (Seibel, 1988: SC 15, revised). CI=1.00; 1.00.

CUCKOO MONOPHYLY AND PHYLOGENY BASED ON OSTEOLOGY

299

109. Humerus, protuberance near attachment of m. supraspinatus: (0) absent; (1) present (Seibel, 1988: HU 9). CI=1.00; 1.00. 110. Humerus, extremitas proximalis humeri, ‘bulbous convexity’ on facies cranialis just lateral to midpoint: (0) absent; (1) present (Seibel, 1988: HU 16). CI=1.00; 1.00. 111. Humerus, size of foramen pneumaticum: (0) large; (1) small to moderate. CI=1.00; 1.00. 112. Humerus, position of fossa m. brachialis in cranial aspect: (0) mesial to centre of corpus humeri; (1) far mesial; (2) at or lateral to meso-lateral centre of corpus humeri (Seibel, 1988: HU 47, revised). CI=1.00; 0.67. 113. Humerus, form and position of proc. flexorius in axis proximodistalis (ordered): (0) square shape in distal aspect, finger-like in caudal aspect, tip distinctly distal to condylus ventralis; (1) shape as in state 0, but tip not distal to condylus ventralis; (2) tip of proc. flexorius ‘cut off’, condylus ventralis protrudes distally to tip of processus (Seibel, 1988: HU 14, revised). CI=0.67; 0.67. 114. Humerus, form of condylus ventralis, as in monophyly character 12: (0) no; (1) yes. CI=NC; 1.00. 115. Ulna, margo distalis of proc. cotyla dorsalis relative to proc. cotyla ventralis: (0) same level in axis proximodistalis; (1) markedly proximal; (2) markedly distal (Seibel, 1988: UL 1, revised). CI=0.40; 0.40. 116. Ulna, wide fossa between proc. cotylae dorsalis and ventralis: (0) absent; (1) present (Seibel, 1988: UL 1, revised). CI=1.00; 1.00. 117. Ulna, shape of proc. cotyla dorsalis: (0) not elongate; (1) elongate. CI=0.33; 0.33. 118. Ulna, prominent spur on margo distalis of proc. cotyla dorsalis: (0) absent; (1) present. CI=1.00; 1.00. 119. Ulna, incline of longest chord passing through face of proc. cotyla ventralis: (0) meso-distally to latero-proximally; (1) not as in state 0 (Seibel, 1988: UL 4). CI=NC; 1.00. 120. Radius, extremitas distalis radii, facies articularis radiocarpalis: (0) concave or convex; (1) straight. CI=1.00; 1.00. 121. Os carpi ulnare, angle of juncture of crus breve and crus longum: (0) not approximately 90°; (1) approximately 90° (Seibel, 1988: CU 1). CI=NC; NC. 122. Os carpi ulnare, shape of crus longum: (0) curved or bent; (1) nearly straight (Seibel, 1988: CU 2). CI=NC; NC. 123. Os carpi ulnare, relative thickness of crus breve and crus longum: (0) facies articularis ulnaris thicker than crus longum; (1) approximately equal (Seibel, 1988: CU 3). CI=NC; NC. 124. Os carpi ulnare, shape of distal end: (0) blunt, expanded, or bulbous; (1) pointed, margo distalis 45° to crus longum; (2) strongly pointed, approximately 30°. CI=0.67; 0.67. 125. Phalanx proximalis digiti majoris, shape in planum transversalia: (0) triangular, concave sides; (1) rectangular, flat sides (Seibel, 1988: PH 1). CI=NC; 1.00. 126. Atlas, form: (0) notched; (1) perforated. CI=NC; 1.00. 127. Axis, form of proc. articularis caudalis in lateral aspect: (0) elongate with rounded tip; (1) anvilshaped. CI=NC; 1.00. 128. Vertebrae cervicales, number: (0) 14 or more; (1) 13. CI=1.00; 1.00. 129. Vertebrae thoracica, number: (0) 5; (1) 4. CI=1.00; 1.00. 130. Synsacrum, extremitas caudalis synsacri, as in monophyly character 13: (0) no; (1) yes. CI=NC; 1.00. 131. Vertebrae caudales, primus, as in monophyly character 14: (0) no; (1) yes. CI=NC; 1.00. 132. Pygostylus, cranial extent of margo dorsalis in lateral aspect: (0) not as in state 1 or 2; (1) caudal of facies articularis; (2) cranial of facies articularis. CI=1.00; 1.00. 133. Pygostylus, approximate angle of juncture of margo dorsalis and caudalis in lateral aspect: (0) 45°; (1) 90°. CI=1.00; 1.00. 134. Pygostylus, angle of juncture of basis pygostyli and margo caudalis in lateral aspect: (0) straight, 180°; (1) obviously angled, about 135°. CI=1.00; 1.00. 135. Pygostylus, shape of caudal region between basis pygostyli and canalis vascularis in lateral aspect: (0) tapered to point, projecting cranially; (1) blunt, squared-off. CI=1.00; 1.00.

300

J. M. HUGHES APPENDIX 3

Osteological character data matrix Character data matrix for 33 genera of cuckoos, turacos (Musophagidae) and the Hoatzin (Opisthocomus hoazin). See Appendix 2 for character descriptions. Characters Taxa

1

2

3

4

5

6

7

8

9

1 0

1

2

3

4

5

6

7

8

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

0 0 ? 0 0 0 0 0 0 0 1 1 0 1 1 0 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 01 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 0 1 1 0 0

1 1 ? 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1

2 2 ? 2 1 ? 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 0 ? 0 0 0 0 0 1 1

0 0 ? 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 1 1 0 0 0 0 0 0 0 0 0 0

0 0 ? 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

continued

CUCKOO MONOPHYLY AND PHYLOGENY BASED ON OSTEOLOGY

301

APPENDIX 3—continued

Characters Taxa

1 9

2 0

1

2

3

4

5

6

7

8

9

3 0

1

2

3

5

6

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 2 0 0 0 0 1 0 0 1 0 0 1 1 2 1 2 1 1 0 0 0 0 1

1 1 ? 2 2 ? 2 2 2 2 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0

1 1 ? 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1

0 0 ? 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 2 2 ? 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 0 0 1 0 1 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 2 2 2 0 0 0 0 0 0 0 0 0 0 ? 1 1 1 1 0 1 0 1 1 0 0 0 0 0

1 0 0 1 0 0 ? ? ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 01 0 0 1 0 0 1 01 01 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 1 ? ? ? 1 1 1 1 1 1 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 1 1 0 0 0 0 0

4

continued

302

J. M. HUGHES APPENDIX 3—continued

Characters Taxa

3 7

8

9

4 0

1

2

3

4

5

6

7

8

9

5 0

1

2

3

4

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

1 1 ? 2 1 1 2 1 1 1 1 1 1 0 0 0 0 0 0 0 ? 0 0 0 0 1 0 1 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1 1 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 0 0 1 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 1 0 1 1 1 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 01 1 0 1 0 0 0 0 01 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 01 1 01

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 2 1 2 2 1 1 1 1 1 1 1 1 1 1 0 1 0 0 1 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

continued

CUCKOO MONOPHYLY AND PHYLOGENY BASED ON OSTEOLOGY

303

APPENDIX 3—continued

Characters Taxa

5 5

6

7

8

9

6 0

1

2

3

4

5

6

7

8

9

7 0

1

2

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0

1 1 ? 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

0 0 ? 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 2 2 2 2 2 2 2 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 1 1 0 0

1 1 ? 2 1 1 2 1 1 1 1 1 1 0 0 0 1 1 1 1 1 1 1 0 0 1 0 1 0 0 0 0 0 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 3 2 2 3 2 2 2 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0

continued

304

J. M. HUGHES APPENDIX 3—continued

Characters Taxa

7 3

4

5

6

7

8

9

8 0

1

2

3

4

5

6

7

8

9

9 0

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0

0 0 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 2 2 2 0 2 0 0 0 0 0 0 0

1 1 2 2 2 2 2 2 2 2 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 1 0

2 2 3 3 3 3 3 3 3 3 2 2 2 2 2 2 1 1 2 2 2 1 1 1 1 2 1 2 1 1 0 1 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0

1 1 1 2 2 2 2 2 2 2 1 1 1 0 0 1 0 1 0 0 1 0 0 1 1 1 0 12 0 0 0 0 0 2 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0

1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 1 1 1 2 0 12 0 0 0 0 0 0 0

2 2 3 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 1 1 1 1 0 0

continued

CUCKOO MONOPHYLY AND PHYLOGENY BASED ON OSTEOLOGY

305

APPENDIX 3—continued

Characters

Taxa

9 1

2

3

4

5

6

7

8

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 3 3 3 3 3 3 3 3 3 3 0 0 0 2 0 2 2 1 1 1 0 0

2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 0 0 0 0 2 0 2 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 1 0 1 1 1 1 1 2 0

2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 0 0 0 0 0 0 1 1 2 0 2 0 0 0 0 0 0 01

2 2 0 1 1 1 1 1 1 1 0 0 2 3 3 3 3 4 4 4 0 4 4 0 0 2 0 2 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 01 0 0

0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

9

1 0 0

0 1

0 2

0 3

0 4

0 5

0 6

0 7

0 8

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

3 3 3 3 3 ? 3 3 3 3 2 2 3 2 2 2 2 2 2 2 2 2 2 1 1 2 1 2 1 1 0 0 0 4 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0

2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0

1 1 0 0 0 ? 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 ? 2 1 2 1 1 1 1 1 0 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 3 1 3 1 1 2 2 2 0 2

continued

306

J. M. HUGHES APPENDIX 3—continued

Characters

Taxa

1 0 9

1 0

1 1

1 2

1 3

1 4

1 5

1 6

1 7

1 8

1 9

1 2 0

2 1

2 2

2 3

2 4

2 5

2 6

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 2 0 0 0 0 0 0 0 0 0 0 0 0 2 1 2 1 1 1 1 1 0 0

1 1 ? 2 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 ? 2 2 2 2 2 2 2 2 2 1 1 1 1 1 2 2 2 2 2 2 1 1 1 1 1 1 1 0 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

2 2 ? 2 2 2 2 2 2 2 2 ? 2 1 1 1 1 1 1 1 1 1 1 1 1 2 0 2 0 0 0 1 1 1 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 0 0

continued

CUCKOO MONOPHYLY AND PHYLOGENY BASED ON OSTEOLOGY APPENDIX 3—continued

Characters

Taxa

1 2 7

2 8

2 9

1 3 0

3 1

3 2

3 3

3 4

3 5

Clamator Oxylophus Pachycoccyx Cuculus Surniculus Penthoceryx Cacomantis Chrysococcyx Misocalius Chalcites Eudynamys Urodynamis Coccyzus Hyetornis Piaya Coccycua Saurothera Ceuthmochares Rhopodytes Rhamphococcyx Rhinortha Lepidogrammus Dasylophus Crotophaga Guira Tapera Morococcyx Dromococcyx Geococcyx Neomorphus Carpococcyx Coua Centropus Opisthocomus Musophagidae

1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 ? 0 0 0 0 0 0 0 0 ? 1 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 ? 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 1 0 1 1 0 0

1 1 ? 1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

2 2 ? 2 2 ? 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 1 0 0 0 ? 0

1 1 ? 1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0

0 0 ? 0 0 ? 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 1 0

0 0 ? 1 1 ? 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0

307