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Monophyly and Phylogenetic Relationships of the Pelecaniformes: a Numerical Cladistic Analysis

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Monophyly and Phylogenetic Relationships of the Pelecaniformes: a Numerical Cladistic Analysis MONOPHYLY AND PHYLOGENETIC RELATIONSHIPS OF THE PELECANIFORMES: A NUMERICAL CLADISTIC ANALYSIS JOELCRACRAFT Departmentof Anatomy,University of Illinois,P.O. Box6998, Chicago, Illinois 60680 USA, and Divisionof Birds,Field Museum of NaturalHistory, Chicago, Illinois 60605 USA ABSTRACT.--Aphylogenetic analysis was undertakento evaluatethe monophylyof the Pelecaniformesand to determineinterfamilial relationships within the order.A total of 52 characterswas subjectedto a numericalcladistic analysis. Pelecaniform monophyly was highly corroborated,with 12 postulatedsynapomorphies supporting that hypothesis.Within the pelecaniforms,the phaethontidsare the sister-groupof the remainingfamilies, which are divided into two lineages,the fregatidson the one hand and the pelecanids,sulids, and phalacrocoracids(including anhingids) on the other. Within the latter clade, sulids and phalacrocoracidsare eachother's closest relatives. This pattern of interfamilialrelationships was stronglycorroborated by the data,and alternativehypotheses, especially those postu- lating a closerelationship between phaethontids and fregatids,are muchless parsimonious. The study also presentscorroborating evidence that pelecaniformsand procellariiforms are sister-taxa,although this hypothesisrequires further analysis.Evidence supporting a relationship between pelecaniformsand ciconiiformsis evaluated and consideredinsuffi- cient to warrant acceptanceof that hypothesisat this time. The hypothesisthat the Whale-headedStork (Balaeniceps rex) hasa relationshipto one or more pelecaniformtaxa was investigatedand rejected.The characterssaid to indicatea close relationshipare interpretedhere to be convergencesthat aroseas mechanicalresponses to similaritiesin feedingbehavior. Received 10 January 1985, accepted 16 May 1985. THE phylogenetic relationshipsof the Pele- within the order, the majority of previous caniformes have interested systematic orni- workers have concluded that pelecanids, su- thologists for many decades[see the detailed lids, phalacrocoracids,and anhingids comprise review by Sibley and Ahlquist (1972: 65-70)]. a natural group. Opinion has been divided, The most influential paper in this century however, regarding the phylogenetic place- probablyhas been that of Lanham(1947), whose ment of the fregatids and phaethontids, and phylogeneticconclusions were basedprimarily although some workers have placed them to- on an analysis of skeletal similarities and dif- gether, most have relegatedeach to a separate ferences.He supportedthe hypothesisof pele- suborderwithin the Pelecaniformes(e.g. Wet- caniform monophyly, with the frigatebirds more 1960,Storer 1971; see Sibley and Ahlquist (Fregatidae) and tropicbirds (Phaethontidae) 1972 for a summary). being sister-groupsand related to a lineage With respect to interordinal affinities, most composedof all the remaining families. Rela- systematistshave identified either the Procel- tionshipswithin the latter clade were postulat- lariiformes or Ciconiiformes as possible close ed to be subdivided into the pelicans (Pelecan- relatives of the pelecaniforms,but in no case idae)on the one hand, and gannetsand boobies has adequate documentation been provided. (Sulidae), cormorants (Phalacrocoracidae), and Some have noted a close anatomical resem- anhingas(Anhingidae) on the other; the sulids blancebetween fregatids and diomedeids(Bed- were consideredto be the sister-groupof the dard 1898, Lanham 1947, Simonetta 1963), and cormorantsand anhingas. virtually all who have studied the problem Although some systematists have raised agree that phaethontids are the most aberrant doubts about the monophyly of the Pelecani- family of the order. formes, the families usually have been placed I presentevidence supporting the hypothe- together. This decision has been based almost sis that the Pelecaniformes constitute a mono- entirely on a single shared character,namely phyletic group. Most of the phylogeneticrela- the totipalmatefoot. In assessingrelationships tionships postulated by Lanham (1947) are 834 The Auk 102: 834-853. October 1985 October1985] PelecaniformPhylogeny 835 confirmed,but the placementof fregatidswith ing prior(working) hypotheses about monophyly and phaethontids is not supported by evidence characterpolarity (Farris 1982,Maddison et al. 1984, based on an analysis of shared derived char- Swofford 1984): for a given character-taxonmatrix, acters. Furthermore, using cladistic analysis, the most parsimoniousnetwork is computedover all taxa using unordered characters(that is, character- much stronger evidence can now be provided statesare not polarized and evolutionary transitions to corroboratenot only the monophyly of the are allowedto occurfrom any one stateto any other). order but also the interrelationships of the in- Oncean undirectedtree is generated,it canbe rooted cluded families. In addition, evidence is pre- using one of several possible conventions, for ex- sentedthat suggestsa sister-grouprelationship ample by specifyingone or more outgroupsor by between the Pelecaniformes and Procellari- designatinga hypotheticalancestral taxon. If the in- iformes. group taxonemerges as a discretegroup within the undirectedcladogram, its monophyly can be accept- SYSTEMATIC METHODS AND MATERIALS ed and further testedby succeedinganalyses that in- corporatehypotheses of characterpolarity. Method- Using cladisticmethodology, hypotheses of mon- ologically,the useof unorderedcharacters is strictly ophyly are testedby the parsimoniousdistribution cladisticbecause congruence in character-statetran- of postulatedshared derived characters,or synapo- sitionsis being optimized on the tree. morphies (Hennig 1966, Eldredge and Cracraft 1980, In this analysisundirected trees or networkswere Nelson and Platnick 1981, Wiley 1981).Hypotheses computedusing the PhylogeneticAnalysis Using of synapomorphyusually are erectedusing the well- Parsimony(PAUP) program (version 2.2) of DavidL. known comparativetechnique of outgroup analysis Swofford(University of Illinois, Illinois Natural His- (Wiley 1975, 1981;Gaffhey 1979;Eldredge and Cra- tory Survey).As I will discussbelow, the Pelecani- craft 1980; Watrous and Wheeler 1981; Farris 1982; formesemerged as a naturalgroup in the analysisof Maddison et al. 1984).The argumentationscheme of unorderedcharacters. Accordingly, subsequent anal- outgroupanalysis takes the following form:If a char- yseson ordered(polarized) characters were under- acter(or characterstate) is sharedamong two or more taken using pelecaniformsas the ingroup. Relation- ingroup taxa and also is found within postulated shipswithin the pelecaniformswere investigatedby closelyrelated outgroups,the characteris primitive comparisonsto outgroup taxa that are frequently within the ingroup; if the characteris restricted to consideredto be closelyrelated, including the Pro- ingrouptaxa, it canbe consideredto be derivedwith- cellariiformes,Gaviiformes (including grebes,Podi- in that ingroup. Various workers have drawn atten- cipedidae),and Sphenisciformes(Sibley and Ahl- tion to someof the underlying(and often unstated) quist 1972;Cracraft 1981, 1982). Ordered trees were assumptionsof this method, namely, that the in- rootedby two procedures:first, by specifyingsphen- group is monophyleticand that the outgroupsare iscidsand/or gaviidsas the root, and second,by con- actuallyclosely related to the ingroup (seeMaddison structinga hypotheticalcommon ancestor (Kluge and et aL 1984 for a detailed discussion, and Raikow 1982 Farris 1969,Farris 1970,Lundberg 1972,Maddison et for an example using birds). Thus, the question of al. 1984)based on comparisonswith these outgroups ingroup monophylymay be an integral part of the and with many nonpasserinebirds aswell. study itself, and relationshipsamong the outgroups Parsimonyis the singlemost important philosoph- and their affinities to the putative ingroup may be ical principle underlying phylogeneticanalysis (Far- uncertain. ris 1982, 1983). Accordingto this principle, ad hoc One approachto this problem is to begin with a hypothesesof homoplasy(parallelism, reversal) higher-level phylogenetichypothesis (containing are minimized, and congruenceof characterdistri- both ingroup and outgrouptaxa), whose monophyly butions is maximized. The principle doesnot assume, is well established, and then document the mono- or assert,that the processof charactertransformation phyly of the ingroup by searchingfor characters is itself "parsimonious"(whatever that term might unique to those taxa. Raikow (1982), for example, mean in this context),only that our choice among adoptedthe classAves as his higher-leveltaxon, and competingscientific hypotheses must be basedon an then assessedthe monophylyof the Passeriformesby objectiveand rationalcriterion. Parsimony is that cri- searching for uniquely derived characters.Mono- terion. Thus, in the analysesdiscussed here, the phyly alsocan be corroboratedif charactersdefining number of character transformationsis the entity the group are derived but not unique, i.e. if they being minimized, with optimization proceduresas- have been acquired independently in the ingroup signing internode characterstates so as to minimize and one or more outgroups.Such a conclusion,how- the f-values (Farris 1972). Naturally, if the data set ever, is predicatedon acceptanceof a phylogenetic were changed,either by adding, subtracting,or re- hypothesisthat is globally parsimoniousover both coding charactersor taxa, tree lengths could be al- the ingroupand the outgroups(Maddison et al. 1984).
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