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Monophyly of the Piciformes: a Reply to Olson

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Monophyly of the Piciformes: a Reply to Olson MONOPHYLY OF THE PICIFORMES: A REPLY TO OLSON ROBERTJ. RAIKOW1 AND JOELCRACRAFT 2 •Departmentof BiologicalSciences, University of Pittsburgh,Pittsburgh, Pennsylvania 15260 USA and Carnegie Museumof Natural History, Pittsburgh,Pennsylvania 15213 USA, and 2Departmentof Anatomy,University of Illinoisat the MedicalCenter, Chicago, illinois 60680 USA and FieldMuseum of Natural History,Chicago, illinois 60605 USA ABSTP•CT.--Olson(1983) questioned the hypothesisthat the order Piciformesis mono- phyletic and suggestedinstead that each piciform suborder is allied with a nonpiciform group. His attempt to discreditthe synapomorphiesjoining the Galbulae and Pici is refuted by correctionsof his interpretationsof previous work. The Piciformes share a complex derived hindlimb morphologyinvolving zygodactyly,a tripartite flexorhallucis longus, and Type VI flexortendons. Olson's argument for polyphylycombines inadequate data with the inappropriatetechnique of equatinggeneral overall similarity with affinity.His hypothesis is thereforerejected. Problems concerning fossil taxa are also discussed.Based on current information,we believethat a monophyleticorigin of the Piciformesremains the hypothesis of choice.Received 18 August1982, accepted18 October1982. SWIERCZEWSKIand Raikow (1981)and Simp- more primitive than that of the Pici, which is son and Cracraft (1981) studied the phyloge- more "specialized," and thereforethat mono- netic relationshipsof the Piciformes.Both con- phyly requiresthe conditionin the Pici to have cluded that the order is monophyleticand that evolved from that in the Galbulae. Here, he it contains two monophyletic suborders, the confusessister-group relationship with ances- Galbulae (Bucconidae and Galbulidae) and Pici tor-descendentrelationship. We did not pro- (Capitonidae, Ramphastidae, Indicatoridae, pose that the Pici evolved from the Galbulae and Picidae). Olson (1983) criticizesthe hy- (higher taxa cannotbe ancestors).Olson asks pothesisof monophyly. First, he questionsthe "... why would such a transformationtake argumentssupporting this view. He then pre- place?Once a group of birds has becomeper- sents an alternative hypothesis that the Gal- manently zygodactyl,is it possibleto become bulae are most closely related to the Coracii more zygodactyl?"This question manifestsa and the Pici to the Passeriformes.Finally, he confusion. One group is not "more zygodac- criticizes Simpson and Cracraft'sdiscussion of tyl" than the other;both are describedby this certain fossil birds. After carefullyexamining term. It may be expected,however, that dif- Olson's critique, we believe that monophyly ferenceswill accumulatein separatelyevolving remains the more strongly supportedhypoth- lineagesafter they have split; it is this process esis. The possiblevalue of these discussions of evolutionary divergence or charactertrans- goesbeyond the questionof a singlebranching formation that gives rise to the hierarchical point in a phylogeny,as it providesan oppor- structureof organicdiversity. We suggestthat tunity to compareand contrasttwo profoundly the zygodactylconditions of the Galbulae and different approachesto systematicanalysis. Pici are homologousbecause other characters (see below) corroboratethe unity of the Pici- formes. PICIFORM MONOPHYLY M. flexorhallucis longus.--Swierczewski and Z!/godact!/l!/.--Olsonsuggests that zygodac- Raikow (1981)reported that the flexor hallucis tyly aroseindependently in the Pici and Gal- longusmuscle (FHL) shows a derived condi- bulae becauseSteinbacher (1935) found differ- tion in its origin by a certainpattern of three ences in their tarsometatarsi. He does not heads. In most birds there are one or some- consider the possibility that the differences times two femoral heads, but the Piciformes aroseafter the two groupsdiverged from a zy- have as well an extensiveorigin from the fib- godactylouscommon ancestor. Olson consid- ula. Olson dissectedone capitonid (Trachy- ers that the tarsometatarsus of the Galbulae is phonus;Pici) and one bucconid(Hypnelus; Gai- 134 The Auk I00: 134-138.January 1983 January1983] Monophylyof thePiciformes 135 FIlL MED each of two species.In contrast,Swierczewski INT LAT dissectedbetween one and four specimenseach of 45 species, and concluded unequivocally (1977: 58) that "Three heads of origin were foundin allspecies studied herein." Nevertheless, one of us (R.J.R.) has, in re- sponseto Olson'scomments, dissected the limb of Colaptes auratus (Picidae), Bucco tamiata (Bucconidae), and Galbula dea (Galbulidae). Like Olson, Raikow confirmed in Colaptesthe three heads previously described(Fig. 1). In Hypnelus, Olson "... could not detect any separateheads of origin . ." and questioned whether "... the nature of the origin of this muscle is homologousbetween the Galbulae and the Pici or even that it can really be said to have three heads in the Galbulae." The dis- section of Galbula and Bucco has fully con- firmedour previousreport that the threeheads, including the large fibular head, are presentin the pattern described (Fig. 1). This corrobo- ratesthe hypothesisof piciformmonophyly and refutes Olson's claim that the condition is not found in both piciform suborders. Type VI flexor tendons.--Did this condition Fig. 1. Lateralviews of the regionof the knee and upper crus in three speciesof piciform birds to show arise oncein the Piciformesor separatelyin the Galbulae and Pici? Olson states that it is the modeof origin of M. flexorhallucis longus (FHL). In all Piciformesthe musclearises by three separate "probably a convergentsimilarity," but offers heads, medial (MED), intermediate (INT), and lateral no evidence. He suggeststhat the Galbulae (LAT), all of which lie medial to the tendon of M. would be difficult to identify with their feet cut iliofibularis (IL. FIB.). The representativeforms il- off, but fortunately all of our specimenspos- lustrated are Colaptesauratus (Picidae; Pici), Bucco sessedfeet. (Actually, the relevant structures tamiata (Bucconidae; Galbulae), and Galbula dea are legs, not feet, as the FHL belly lies in the (Galbulidae;Galbulae). The drawingswere madewith crus.) Olson feels that such characters do not a camera lucida and dissectingmicroscope and are not to scale. justify ordinalrank, but we do not agree that taxonomic rank is related to the corporeal lo- cation of the relevant characters. Here, Olson bulae). In Trachyphonushe confirmedthe report confusesphylogeny with taxonomy;the ques- of Swierczewski and Raikow (1981), but in tion at issue is whether the group is mono- Hypnelus he did not. Olson quotes Swier- phyletic, not what rank it shouldhave in a clas- czewski (1977: 57) as saying that in the Gal- sification. Perhaps the FHL should rank the bulae the headsare "somewhatdifficult to sep- group at the generic level, becauseits origin arate from each other." Swierczewski's full lies at the knee joint (Juncturagenus). statement, however, conveysa different mean- ing: "In the Galbulidae and Bucconidae, the ANALYSIS OF OLSON'S HYPOTHESIS medial head arises semitendinous from the in- tercondyloidregion of the femur, and the in- Inadequatedata.--Olson comparescharacters termedlate head arises fleshy from the proxi- in only some of the relevant groups, so that mocaudal surface of the external femoral comprehensivecomparisons cannot be made. condyle;the headsbeing somewhatdifficult to For example, he notes that "in plumage pat- separatefrom each other." This statement re- tern, the ground roller Brachypteraciaslepto- fers to only two of the three heads;it doesnot somusis quite similar to certain of the Buccon- refer to the fibular head, which is the signifi- idae such as Malacoptila." But what are the cant structure. Olson dissected one specimen patterns in the other Coracii, the other Buc- 136 RAIKOWA•r) CRACRAFX [Auk, Vol. 100 conidae, and the Galbulidae?Why are they not ic similarity. He claimsthat we acceptthe idea discussed?Convergence in plumagepatterns is that "... differences between taxa [are] evi- common, which is why comprehensivecom- dence of nonrelationship .... "but this is in- parisons are needed to distinguish between correct.We mean only that the absenceof sim- synapomorphiesand spuriousresemblances. ilarities fails to corroboratea hypothesis of Again, in discussingthe skull Olson argues monophyly, not that the presenceof differ- for similarities between the Galbulae and Cor- encesrefutes such a hypothesis."Nonrelation- acii, but restricts his comparison almost en- ship" has no intrinsic meaning;we are search- tirely to the Bucconidaeand Coraciaswhen list- ing for patternsof commonancestry identified ing severalfeatures. "In all of thesecharacters," by synapomorphy. he notes, "the Bucconidaeare consistentlydif- The problem with phenetic comparisonsis ferent from the Pici." But what about the Gal- that charactersare not analyzed so as to iden- bulidae and other Coracii? tify the level within the phylogenetichierarchy This patternof selective,almost casual choice at which they define taxa. To illustrate this of data characterizes Olson's entire presenta- point, we will consider one example from a tion. It is also difficult to assess his data, be- recentstudy of the relationshipsof Pedionomus cause they are presented in a narrative form torquatus.This speciesis placed in the mono- within the text but are not tabulated in a way typic family Pedionomidae, and the problem that would allow one to determine the state of is to find its closest relatives. Olson and Stead- each character in all the relevant taxa. A tab- man (1981) rejected
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