The Auk 116(1):116-130, 1999

PHYLOGENY
OF THE FALCONIDAE
INFERREDFROM
MOLECULAR
AND MORPHOLOGICALDATACAROLE S. GRIFFITFIS •Departmentof Ornithology,
AmericanMuseumofNaturalHistory,CentralParkWestat 79thStreet,
New York, New York 10024, USA

ABSTR^CT.--Molecular
data and variationin syringealmorphologywere usedto infer aphylogenyfor the family Falconidaeand to addressthreeissuescurrentlyof interestin systematics:(1) the treatmentof multipledatasetsin phylogeneticanalysis,(2) a priorianalysis
and differentialweightingof moleculardata,and(3) thereliabilityof molecularversusmorphologicaldatain phylogenetic
analysis.Problemsin recoveringphylogenetic
signalcausedby rapidly changingsitesin the moleculardata were not solvedby combiningdata sets.
Differentiallyweightingsaturatedpartitionsof the sequencedata, prior to phylogenetic
analysis,provideda phylogenycongruentwith morphologicalanalysis.Moleculardataprovide substantiallymoreinformativecharacters
thanmorphological
data.However,morphologicaldataprovidea higherproportionof unreversedsynapomorphies.
A reclassificationof the family basedon the phylogenyresultsin two subfamilies:(1) the Herpetotherinae,(forest-falcons[Micrastur]and LaughingFalcon[Herpetotheres
cachinnans]);
and (2) the Falconinae,whichincludesthe tribesFalconini(Spot-wingedFalconet[Spiziapteryx
circumcinc-tus],pygmy-falcons[Polihierax],
falconets[Microhierax],
andthegenusFalco)andCaracarini(caracaras).
Thephylogenyalsoindicatesthattwo genera,Daptrius
andPolihierax,
arepolyphyletic,and thesetwo are split.Finally,a biogeographic
hypothesisderivedfrom tl•ephylogenyimpliesthattheoriginandearlydiversificationof thefamilyoccurredin SouthAmerica. Received
12 November
1997,accepted
18 June1998.

THE FAI•C¸NIDAE,one of three families in the genera:
thefourcaracaragenera,Micrastur(fororderFalconiformes(thediurnalbirdsof prey), est-falcons),Herpetotheres(Laughing Falcon)
includessomeof the fastestand mostspectac- and Spiziapteryx(Spot-wingedFalconet).The
ular birds in the world (Brown and Amadon relationshipsof thelatterthreegenerahistori1968).Althoughthefamilyis renownedfor the callyhavebeenproblematic(Sharpe1874,Gurhuntingabilityof its species(e.g.thePeregrine ney 1894,Swann1922,Peters1931).TheplaceFalcon[Falco
peregrinus]),foraginghabitswith- mentof thesegenerawith the caracaras(Amain thefamilyarediverse.Thus,theNeotropical don and Bull 1988) is in conflictwith results
caracaras(Daptrius,Milvago, Polyborus,and from four cladisticanalyses:(1) an osteological
Phalcoboenus)
are scavengers
that subsistmain- study (Becker1987;Fig. 1A), (2) a preliminary
ly on carrionor invertebrates.
morphologicalanalysis (Kemp and Crowe
In additionto adaptationsrelatedto foraging 1990;Fig. 1A), (3) an analysisbasedon syrinhabits,the four caracaragenerashareother gealmorphology(Griffiths1994a;Fig. lB), andmorphologicaltraits(Friedmann1950)andhis- (4) an analysisbasedoncytochrome-b
sequenctoricallyhavebeenconsidered
to be closelyre- es (Griffiths1997;Fig. 1C). Basedon Griffiths
lated(Sharpe1874,Gurney1894,Swann1922, (1994a),theAOU (1998)reclassified
thefamily,Peters1931).Classificationof the othergenera placing Herpetotheres
and Spiziapteryx
withinin thefamilyhasbeenlessstable(Sharpe1874, the Falconinae and Micrastur in its own subGurney 1894, Swann 1922, Peters1931). Ama- family.
don and Bull (1988)allocatedthe 10 currently
In thispaper,I reanalyzethesyringealdata,
recognizedgenerainto two subfamilies.The incorporatingnew information.Having collectFalconinaeincludedthe speciosegenusFalco ed both morphologicaland moleculardata, I
and the smallestdiurnal raptors, Polihierax then examinea controversialissuein phylo(pygmy-falcons)and Microhierax(falconets). geneticinference,the appropriateanalysisof
The PolyborinaeincludedsevenNeotropical multiple data sets.This controversyspansa
continuumof ideas. At one end is the idea thatdata setsshouldbe analyzedseparately.The

E-mail:csg@amnh.org
116January1999]
A

t
B

c

Falconidae
Phylogeny•

117

nentsof totalevidenceproposegivingall charactersequal weight initially in phylogenetic
analysis.Assessmentsof characterreliability
derivedfrom phylogenetic
inference(i.e. sucCaracaraspecies
cessiveapproximations;
Farris1969,1989;CarMicrastur
penter1988)canthenbe usedto weightcharHorpotothoros
actersdifferentially(a posteriori
weighting).A
prioridifferentialweightingis criticizedfor inFalco
curring unwarrantedassumptionsabout proMicrohiorax
cessor requiringpartitionsof datafor weightPolihiorax
somitorquatus
ing that maybe arbitrary(Eernisse
and Kluge
Spiziaptoryx
1993,
Brewerand
DeSalle1994,
Chippindale
Polihierax
insignisand Wiens 1994).
Horpotothoros
As part of the empiricalexplorationof the
Caracara
speciesanalysisof multipledatasets,I examinethe efMicrastur
species
Outgroups
fectsof differentialweightingon phylogenetic
inference.I then comparethe morphologicaland moleculardata. The phylogenyinferred
Falcofrom thesedatais usedto reclassifytheFalconidaeandtwo generawithin thatfamily.Finally,
Microhiorax
the phylogenyprovidesa frameworkfor disPolihioraxsomitorquatus
cussingthebiogeographyof thefamily.
Spiziapteryx

Falco
Microhierax and Polihierax

Spiziaptoryx••

Caracaraspecies

Horpotothoros
Micrasturspecies

MATERIALS

AND METHODSMorphological
data.--Characters
werederivedfrom
variation in syringeal supportingelements,memFIC. 1. Phylogenetic
hypotheses
inferredfrom branes,andmuscles(Griffiths1994a).Supportingelcladisticanalysesof the Falconidae.
(A) Phylogeny ementsincluderinglikeelementson the tracheaandinferred from osteologicaldata (Becker1987) and a bronchi(A andBelements;Ames1971), thepessulus,
preliminaryanalysisof skinsandcharactersfromlit- and accessory
cartilaginousstructures.Hemologyof
erature(Kempand Crowe1990).(B) Phylogenyin- the ringlikeelementswastraditionallybasedon the
Accipiter

ferredfromsyringealmorphology(Griffiths1994a). relativepositionof thesestructuresto the tracheo(C) Phylogeny
inferredfromdifferentiallyweighted bronchialjunction(King1989).Thisanalysis
postucytochrome-b
sequences(Griffiths1997).
lateshomologous
elementsfollowingAmes' (1971)
definitionsof syringealstructures(Griffiths1994b).
Dataon variationin syringealmorphology
fortwocongruenceof phylogenies
inferredfrom each specieswereaddedto the originaldataof Griffiths
dataset(taxonomic
congruence)
thenprovides (1994a).The first, Accipiterstriatus(Sharp-shinned

a measureof the reliabilityof the phylogenies Hawk; AMNH 18761), was substituted for Gampso(Lanyon1993,Miyamoto and Fitch 1995).At nyx swainsonii
to be consistent
with the molecular
the oppositeendis the ideathat multipledata data.The second,Herpetotheres
cachinnans(LSUMNS
setsshouldbe combined
forphylogenetic
anal- 123200),recentlybecameavailable.Becausethe Hersyrinxusedto codecharacters
fortheorigysis (total evidenceor charactercongruence; petotheres
inal analysis(Griffiths1994a)had been damaged,
Kluge 1989).
Between these two extremes, proposed thisnew specimenwasexaminedandthe resulting
information used to recede two of the characters.
methodsfor analyzingmultipledata setsinclude: (1) conditional combinationof data sets,

Character16, the modificationof the endsof the B1element,changed
fromoneto zero.Character
3, dorsal fusionof the tympanum,changedfrom one toi.e.combination
onlyif thedatasetsarenotsignificantlyheterogeneous(Bullet al. 1993,Huel- two. In addition,threecharacters(characters5, 6 andsenbecket al. 1996);or (2) unconditionalcom- 7) that wereoriginallycodedasbinaryrepresentabinationof data setswith differentialweight- tions of alternative character states were receded as
ing of charactersto accommodate
heterogene- one unordered multistate character.
ity (Chippindaleand Wiens1994).Differential Syringealcharacter
descriptions
with thesemodarepresented
in Appendix1.Thefinaldata
weightingis, in itself, controversial.
Propo- ifications

118

CAROLES. GRIFFITHS[Auk, Vol. 116

TABLE
1. Species
includedin thephylogenetic
analyses.
Species

Combinedanalysis

Morphological

Molecular

Milvagochimachima
M. chimango
Polyborus
plancus

*
no
*

*
*
*

*
no
*

Phalcoboenus australis

*

*

*

Daptriusater

*

*

*

D. americanus

*

*

*

Spiziapteryx
circumcinctus
Micrasturgilvicollis
M. semitorquatus
Polihieraxinsignis
P.semitorquatus
Microhieraxerythrogonys
Herpetotheres
cachinnans
Falcoberigora
E sparverius

*
*
*
no
*
*
*
no
*

*
*
*
*
*
*
*
*
*

*
*
*
no
*
*
*
no
*

E mexicanus

no

*

no

E peregrinus
E rufigularis

*
no

*
*

*
no

E biarmicus
E columbarius
E cenchroides

no
no
no

*
*
*

no
no
no

E vespertinus
E femoralis
Accipiterstriatus

*
*
*

no
*
*

*
*
*

Otus asio
Pelecanus onocrotalus

no
no

*
*

no
no

matrixfor the reanalysisconsistedof 25 taxaand 23 able.All 10 currentlyrecognizedgenerawere samcharacters,
six of which were multistate(Appendix pledfor bothmolecularandsyringealcharacters(Ta2). Transformation series for multistate characters ble 1).However,taxonsamplingwasnot identicalbewere proposedif adjacentderivedstateswere simi- tweenthe two datasets.Somespecies
sampledwithlar, and eachsucceedingstatewas a modificationof in the genusFalcovaried betweenthe two, and no
the previousstate;that is, the derivedstatesformed tissuesamplesof Polihieraxinsignis(White-rumped
a nestedset of synapomorphies.
Multistatecharac- Pygmy-Falcon)were available. An initial analysis
ters were coded as unordered if states were alternawas run that includedall species.The numberof
tive variations of a character.
missingcharactersin themorphologicaldatasetreMoleculardata.--The completesequencesfor cyto- suited in conflictin relationshipsamongthe Falco
chromeb werecollectedfor speciesfromeachof the speciesand, therefore,thousandsof most-parsimo10 generain the family (Griffiths1997;Genbankac- nioustrees.Thetopologyof thestrictconsensus
tree
cessionnumbers U83305 to U83320). Details of DNA

extraction,primersused,PCR and sequencing
pro-

fromthatanalysiswasthesameasthetopologypro-

ducedwhen specieswith missingdatawere deleted
tocols,and data analysisare describedin Griffiths
from the analysis.Because
this studywasnot an at(1997).In that study,moleculardata were differentemptto resolverelationships
withinthegenusFalco,
tially weightedprior to phylogeneticanalysisby filspeciesnot includedin the molecularanalysiswere
tering saturatedsubsetsof data. These partitions
pruned from the combineddata matrix for the phywere identified in an assessment of saturation of substitutions within domains of cytochromeb (the logenetictests.
Accipiterstriatus(Accipitridae),the sistertaxonto
transmembrane,
561basepairs;two extra-membranetheFalconidae(Griffiths 1994b),was used to root theregionsof 240and342basepairs;Griffiths1997).The
filteredpartitionswerefirst-andthird-positiontran- tree. The rationalefor using one outgroupwas desitions,and first-positiontransversionsand second- tailed by Griffiths (1997).Basically,severalspecies
positiontransitionsin theextra-membrane
domains. within theAccipitridaewereincludedinitially to deCombined
data.--Thesyringealmorphological
data terminethe root (Smith 1994).In the final analysis,
were combinedwith cytochrome-b
sequences(Grif- only oneoutgroupspecieswasused.This rootedthefiths 1997). Osteologicalcharacters(Becker1987) cladogramin the sameplaceand producedthe idenwere not included because the data matrix for thattical phylogenyas the analysiswith multiple outanalysisand details of the analysiswere not avail- groups.The additionaloutgroupspecies
weredelet-

January
1999]

Falconidae
Phylogeny

ed to facilitatethe detailedexaminationof changeinmolecularcharacterswithin the family.
To test the differentmethodsproposedfor combining data for phylogeneticinference,two setsof
phylogeneticanalyseswere performed.In the firstset (i.e. the total evidencetest), all characterswere
weightedequallyfor an initial phylogenetic
analysis.
Data were thenweightedagainto assess
the effects
of a posterioridifferentialweighting.Weightswere
derivedfrom the fit of the characters
to a phyloge-

119
Accipiter

Herpetotheres
23
2

9*

E icrastur
gilvicollis
M, semitorquatus

5'12'

Daptriusamericanus
6

Polyborus

4* 8•

Daptriusater

Milvagochirnachirna--

Milvagochimango
22

netichypothesis,
usingtwodifferentmethods.
In thePhalcoboenus

first method, the maximum value of the rescaledcon-

2 23sistencyindexwasusedto reweightcharacters
(suc-

16

cessiveapproximations;Farris 1969, 1989;Carpenter1988). In the secondmethod, the characterswere

PolihieraXinsignis
3

Spiziapteryx
3

Microhierax

Polihieraxsernitorquatusweightedconcurrentlywith the analysisbasedon
7•
Falco cenchroides
the homoplasyimpliedby eachtree (Goloboff1993).
•
Falcobedgora
The weightingfunctionusedto evaluatetreeswasa
F•
Falco colurnbarius
modificationof the consistency
index.
The secondsetof analysesexaminedthe effectof
Falcorufigularis
combiningdatausinga prioridifferentialweighting
Falco femoralisof charactersto accommodate
heterogeneityof data
Falcosparverius
sets (unconditionalcombination;Chippindale andFalco biarrnicus
Wiens 1994). Molecular data were differentially•8'
Falco mexicanus
weightedas in Griffiths(1997).Becausethe molecFalcoperegdnus_ __ular phylogeny
inferredin thatstudywascongruent
with the morphologicalphylogeny,the combined
FIG.2. Phylogenyof theFalconidae
inferredfrom
analysiswasperformedto determinetheeffectof thesyringeal
morphology;
F=
Falconinae,
C = Polyboradditionof the morphologicaldata on supportfor
inae.
Cladogram
represents
thestrictconsensus
tree
nodes.Bootstrapanalyseswere performedon theof 54most-parsimonious
treesof length50;CI = 0.59
molecular data and then on the combined data toprovide heuristic measuresfor comparingnodal and RI = 0.81. Characters(Appendix2) supporting
each node are labeled. * = character with CI = 1.0.

support.

Characters3, 4, 5, 6, 16, and 19 are multistate.

Withinthesesetsof analyses,
theeffectof ordering
four of the morphologicalcharacterswasexamined.
Ordering effectivelyweights character-statetransformationsby forcingintermediatestepsin thetran- (1) caracaras;and (2) Falcospecies,including
sition from the first to the last character state. Thus, the two small falconets (Microhieraxand Poliin eachset,morphologicalcharacters
werefirsttreat- hierax)and Spiziapteryx.
Herpetotheres
and Mied as unordered, and then four of the six multistate

crasturform a dichotomybasal to thesetwo
clades.This phylogenyis congruentwith theordering morphologicalchangeson support forosteologicaldata (Becker1987),the molecular
nodes, bootstrap analyseson the combineddata
datawere performedwith morphologicalcharactersor- data (Griffiths1997),the morphological(Kemp and Crowe 1990), and the original sydered and then unordered.
All phylogeneticanalyseswere conductedusing ringeal data in: (1) establishingthese two
the testversion4.0d54of PAUP*,writtenby David L. clades, (2) placing Micrastur basal to these
Swofford.Eachphylogeneticanalysisemployedthe clades,and (3) placingSpiziapteryx
in the Falcharacters were ordered. To determine

the effect ofheuristic algorithm, which included 100 replicates coninae clade rather than with the caracaras.
that randomlyvaried the order in which taxa were The positionof Herpetotheresin thistree differs
added.

from the originalsyringealanalysis(Griffiths
1994a;Fig. lB).
RESULTSAs with Griffiths(1994a),thisphylogenyindicatesthat two genera,Polihierax
and Daptrius,Morphological
data.--Phylogenetic
analysisof are polyphyletic.Differencesin morphology,
thesedata resultedin 54 most-parsimoniousbehavior,andhabitatusebetweenthe two spetrees, which are summarized in the strict con- ciesin eachof thesegenerahavebeendescribed
sensustree (Fig.2). Therearetwo majorclades: previously,and placingthe speciesin eachge-

120

CAROLE
S. GRIFFITHSA[Auk,Vol. 116

in the family (BrownandAmadon1968).In
i_•j.• caracara
species era
addition,the node supportingmonophylyof

I•

Spiziapteryx

theMicrasturspecies
is oneof themoststronglyI[•---Falcospecies supportedin the morphologicalphylogeny

I--•Mlcrohierax(Fig. 2), with four synapomorphies,
three ofthese with a CI of 1.0. The alternative result, in-

L........
Z;;[;;;Td.
TvZ;
u'us

.............
Accipitor

ferredusingdataweighteddifferentiallyprior
to phylogenetic
analysis,is congruent
with themorphologicalhypotheses;
i.e. the Micrastur
speciesare monophyletic.
Combineddata: Total evidence.--Combining

B•J...................
...................

equallyweightedmoleculardataandmorphoCaracaraspecies
logical characters(either ordered or unordered)producedfive most-parsimonious
trees.
Spiziapteryx
The strictconsensus
of thesefive (Fig. 3B) is
Falco species
lessresolvedthan the tree inferredfrom onlymoleculardata(Fig.3A), reflectingthe conflict
Microhierax
Polihierax
semitorquatus betweenthe moleculardata and the morphoMicrasturgilvicollis
logicaldata.Supportexistsfor threeclades:(1)the Falcospecies,(2) the caracaraspeciesand
Herpetotheres
and (3) Microhierax
and Polihierax.
Micrastur
semitorquatus Spiziapteryx,
Thereis no resolutionof relationships
among
Accipiterthesecladesnorof the relationshipsof thetwo

FIG. 3. (A) Phylogenyinferred from equallyweightedmoleculardata(Griffiths1997).(B)Phylogeny inferredfrom combineddata setswith equallyweightedmoleculardata. Strict consensus
tree offive most-parsimonious
cladograms(length= 1,089,
CI = 0.46,RI = 0.45),morphologicalcharactersunordered.The samecladogramsare inferred if morphologicalcharacters
are treatedasordered.Dashedlinesconnectspeciesin the genusMicrastur.

Micrasturspeciesand Herpetotheres.

A posterioridifferentialweighting,usingassessmentsof characterreliabilityderivedfrom
the phylogeneticanalysis(i.e. successive
approximations),
or weightsderivedduringphylogenetic analysis, produced one cladogramsimilarto that inferredfrom equallyweighted
moleculardata(Fig.3A). Resultsare the samewith the morphologicalcharactersorderedor
unordered.
Spiziapteryx
issistertaxon
to thecanusin eitherdifferentsubgenera
or generahas
racaras,
and
the
genus
Micrasturisnotmonobeen advocated(Brown and Amadon 1968).
phyletic.
Analysisof molecular
data.--Results
fromtheCombined data: Unconditional combination.analysisof variationin cytochrome-b
sequences
Combiningdata setswith a prioridifferential
are describedin detail in Griffiths (1997). A
weightingofmoleculardata(i.e.unconditional
summaryis presentedto serveas backgroundcombination)
produced two most-parsimonifor the combinedanalysis.Equal weighting
Thesedifferedonly in reso(Fig.3A) anddifferentialweighting(Fig.1C)of ous cladograms.
sequencedata provided two different results. lution within the caracaraclade (Fig. 4). BeIn the phylogenyinferred from the equally causestepmatriceswereusedfor the molecusummarystatistics
werenotproweightedmoleculardata(Fig.3A), Spiziapteryx lar characters,
As expected,thisis the sister taxon to the caracaras,a result in- ducedfor thesecladograms.
with thesepcongruentwith that inferredfrom osteological phylogenyis basicallycongruent
and morphologicaldata (Figs. 1A, lB, 2). A arateanalyses(a prioriweightingof molecularmore anomalousresultis that the genusMi- data [Fig. 1C] and syringealmorphologicalcrasturis notmonophyletic.
Contraryto there- data [Fig. 2]). Differencesbetweenthe comsuitsfor Daptriusand Polihierax,
polyphylyof bined and separateanalysesappearin two arof
the Micrasturspecieshasnot beensuggested easof thetree:(1) thesistertaxarelationshippreviously.The specieswithin this cladeare MicrasturandHerpetotheresis unresolvedin thesimilar in morphologyand habitat use and morphologicalanalysis(Fig. 2); and (2) in one
havebeenconsidered
distinctfrom othergen- of the two trees,Polyborus
and Daptriusameri-

January
1999]

Falconidae
Phylogeny
Falco sparveriu$

ß
--

-- --

Falcovespergnus
Falco femoralis

Falco
peregrinusF
Polihierax
sernitorquatu$
Microhierax

Spiziapteryx

121

ment of different

data sets. Detailed

reviewsofthis issuehavebeenpresentedpreviously(Hillis 1987,de Queiroz et al. 1995,Miyamoto andFitch 1995,Huelsenbecket al. 1996),and only
main pointsof the argumentwill be discussed.
The two main questionsto be answeredare:(1)Shoulddata setsbe combinedprior to anyphylogeneticanalysis?and (2) If not combinedinitially, shoulddatasetsbe combinedafterinitial
analyses?
The initial combinationof data sets__ Phalcob•enus•C

hasbeenjustifiedphilosophically(i.e. the hypothesiswith the greatestexplanatorypoweris
providedby the concurrentanalysisof all relMicrasturgilvicollis
evant evidence;Kluge 1989). Analyzing data
M. semitorquatussetsseparatelyis disputedbecause
thecreation(•.•oo)HerpetoEleresof separatedatasetsassumes
thattherearenatural classesof data, an assumptionthat is not(Kluge1989).This arguFIC. 4. Phylogenyinferred from combineddata justifiedscientificallysetswith a prioridifferentialweightingof molecular mentcarriesoverinto the appropriatemethod
data, and morphologicalcharactersorderedor un- of weighting characters.A priori character
ordered. F = Falconinae,C Polyborinae(caracaras). weighting requires partitioning data andStrictconsensus
tree of two most-parsimonious
clad- would thereforebe unjustifiedby thosewho

Daptrius
americanu
Z

ograms.Bootstrapvaluesat nodesareshownfor themolecular data set and for the combined data sets;

believe

that

data

shouldbe combined

beforeanalysis.A posteriori
weighting,determinedeither
iteratively
ordirectly,
is appropriatebedata only,numbersbelow the branchesare for thecauseweightsareinferredthroughphylogecombineddatasets.Rangeof numbersfor the comnetic analysis.
bined data sets are for ordered and unordered morSeparateanalysisof multipledatasetsis supphologicalcharacters(left = unordered,right = ordered).
ported by the argument that congruenceamonghypotheses
from differentdatasetsis apowerful
tool
forassessingaccuracyof the hycanusare sistertaxa, a result congruentwith
potheses(MiyamotoandFitch1995).An asthe morphologicalanalysis.In the secondtree,
sumptionof this methodologyis that dividing
Polyborus
is basalto the othercaracaras.
Bootstrapanalysisof the combineddatain- data into classescan be justifiedscientificallydicatesadditionalsupportfor key nodescom- (e.g. different codonpositions).Advocatesof
paredwith the analysisof moleculardataalone the initial separateanalysisof data setsdiffer(Fig.4). Thepositionof Spiziapteryx
in theFalco on the final treatment of multiple data sets.
cladeis not supportedin the majorityof boot- Opinionsrangefrom alwayskeepingdata sets
strap replicateswith molecular data alone. apart (Miyamotoand Fitch 1995) to always
When morphologicaldata are added, that re- combiningdata sets,after any conflictsare aclationshipis supportedin 61 to 71%of the rep- commodatedthrough differential character
licates.The supportfor monophylyof Micrastur weighting(Chippindaleand Wiens 1994).An
increasesfrom 72% to 94 to 97%, and the sup- intermediatepositionis to combineonly conport for the sister-taxarelationshipof the ca- gruent data sets,using either biological(e.g.
racara and Falco clade increases from 65 to 78
genetreesfor a species;de Queirozet al. 1995)
to 82%. Orderingthe four multistatemorpho- or statistical (significant differencesamonglogicalcharactersdoesnot changethe topolo- data sets;Huelsenbecket al. 1996) criteria forgy, but it provides strongersupport for six ofdeterminingincongruence.
One problemwith
sevennodeson the cladogram(Fig.4).
this approachis that the two criteria are notequivalent;datasetsrespondingto differentbiDISCUSSION
ologicalprocesses
may or may not differ sigAnalysis
ofmultipledatasets.--Oneof themost nificantly.In addition,althoughvarioussignifcontentiousissuesin systematicsis the treat- icancetestshavebeenproposed(bootstrap[denumbers

above the branches are for the molecular

122

CAROLE
S. GRIFFITHS[Auk,Vol. 116

Queiroz1993],tree lengths[Farriset al. 1995], stateat nodes(Kim 1996).Thesekinds of charand likelihood ratios [Huelsenbeckand Bull acters occur in the molecular data in this re1996]), the power and sensitivityto the as- search.For thesedata, ratesof changediffer
sumptionsof thesetestsare unknown.
among partitions of sequences,with several
If the goalof phylogenetic
analysisis to pro- partitions containingcharacterswith highvide the hypothesis
with the greatestexplana- ratesof change(Griffiths1997).Substitutions
tory power,then comparingthe variousmeth- are saturatedin thesepartitions,and recoverods is unnecessarybecauseonly a combined ing phylogeneticsignalis problematic(Grifanalysismeetsthatgoal.If thegoalis accuracy fiths1997).Whena prioridifferentialweightingof phylogenetic
inference,thenempiricaltests is usedto correctfor the high ratesof change
comparingthesemethodsare useful(de Quei- in these partitions, the phylogenyindicatesroz et al. 1995).Oneimplicitassumptionof the monophylyof Micrastur,a result congruent
total evidenceapproachis that the sameun- with the morphologicalanalyses.
Theseresultssupportthe intermediatepoderlyingsignalis containedin eachdata set(Hillis et al. 1996).Thus,any potentialconflict sition,the initial separateanalysisof multiple
amongthe hypotheses
inferredfrom individ- data sets,and differentialweightingto accomual data sets will be overcome when data setsmodateheterogeneity
in ratesof change.Afterare combined,circumventingthe need to ad- these processeswere performed,combining
dress conflict among data sets (Brower and the congruentdata setsincreasedthe support
DeSalle 1994).
for nodesin thephylogeny(Fig.4).
This is a testablehypothesisand is directly
Molecules
andmorphology.--Although
the inirelatedto the ideaof consistency;
i.e. a consis- tial useof biochemical
datain systematics
wastent methodology
will resultin convergenceto accompaniedwith assertionsthat molecular
the correcttopologyasdataare added(Felsen- evidencewasmorereliablethanmorphologicalstein 1978, Pennyet al. 1992). In this study, evidence(e.g. Sibleyand Ahlquist1987),the
combiningmorphologicaland moleculardata value of both morphologicaland molecular
produceda phylogenyindicatingpolyphylyof charactersfor systematics
is now generallyacthe genusMicrastur.This resultis inconsistent knowledged(Donoghueand Sanderson1992,
with cladisticmorphologicalanalysesandwith de Queirozet al. 1995;but seeHedgesand Sib~
generallyacceptedconclusions
basedon mor- ley 1994).For this analysis,eachmethodhas
phological,behavioral,and geographicdata. advantages and disadvantages.Informative
Using assessmentsof characterreliability to molecularcharacterswere an order of magnireweightthe characters(aposterioridifferential tude greater in number than morphologicalweighting)also resultedin polyphyly of Mi- characters(328 vs. 23). The relativepaucityof
crastur.
Thisis not surprising;searches
usinga charactersis themostsubstantial
disadvantage
posterJori
weightingare dependenton the val- of the syringealdata.For this analysis,there
uesobtainedfromthestartingtreeandmaybe- are 31 character states for 23 characters. There
cometrapped in local optima (Swoffordet al. are few characters
supportingmostnodes,and
1996).
the relationshipof Herpetotheres
and Micrastur
All phylogeneticmethodscan be inconsis- is unresolved.
tent when the assumptionsof the methodare
Theadvantageof thesyringealdatasetistheviolated (e.g. Debry 1992, Huelsenbeck1995, proportionof unreversedsynapomorphies;
15
Swofford et al. 1996, Sullivan and Swoffordof 31 charactersstateshave a C! of 1.0, and 40%1997). Theseviolationscan be accommodated of the characterschangeonlyoncein the phythroughthe useof a moreappropriatemodel logeny(Figs.5A, 6A). Homoplasyis moreprevor by differentiallyweightingdata(e.g.Steelet alent in the molecular data. Of the 328 characal. 1993, Sullivan and Swofford 1997). The ters, only 10% (33) have a C! of 1.0, and only
problem,then, lies in recognizingthe condi- 5% changeonetime (Figs.5B,6B).Homoplasy

tionsin a datasetunderwhicha phylogenetic obscuresthe signalin the morebasalbranches
method is inconsistent.
connectingthe monotypicgeneraHerpetotheres
Inconsistencycan be causedby characters and Spiziapteryx;
differentialweightingcorwith high ratesof changeand,subsequently,
a rectstheseproblems.
It ispossible
thatmoreselow probabilityof recovering
the homologousquencedatawouldincreasethe signalat theseJanuary1999]

Falconidae
Phylogeny

123

50

45

A

40'

35'

30'

25'

15

20'

1

15'

10.

O' 1

'

2 '

5'

3 '

4 ' 5

'

6

Numberol• steps

00-0.240.25-0.49

0.50-0.74

0.75-0.99

1.00

Consistency
Index

30

B
25
50'

B

45,

.• 20

40'

(D

10

35'

o• 5
o

1

2

3

4

5

6

7

Numberol• steps

20,
15'

FIG.6. The proportionof characterswith a given
numbersof steps.(A) Morphologicaldata, (B) mo-

10.

lecular

data.

5'

0

0-0.240.25-0,49

0.50-0.74

0.75-0.99

1.00

derson 1992, de Queiroz et al. 1995). Differen-

tially weightedmoleculardata stronglysupConsistency
Index
port the sisterrelationshipof Herpetotheres
andFIG.5. The proportionof charactersfor eachval- Micrastur,which is unresolvedin the morphohas
ue of the consistency
index.(A) Morphological
data, logical tree. The positionof Spiziapteryx(B) molecular data.
weak supportin the moleculartree.The inclusionof Polihierax
insignisin the morphologicalanalysis(Fig. 2) strengthensthe supportfor
nodes.Alternatively,additionalsequencedata Spiziapteryx
within the Falcoclade.
may not resolvethesebranchesif the additional
In this study,the moleculardata were moredatahavethe samepatternsof noiseandsignal homoplastic,thus disputingthe assertionthat
asthe presentdata (Brunset al. 1992)or if the moleculardata are morereliablethan morphodata do not fit the assumptionsof the model logicaldata for phylogeneticinference(Sibley
used (Sullivan and Swofford 1997). Because and Ahlquist 1987, Hedgesand Sibley1994).
these genera are monotypic,increasedsam- The occurrenceof saturateddata partitionsinpling of taxawithin the familywouldprobably the falconidsequences
follows a pattern illusnot resolvethe problem.
tratedin studiesof cytochromeb and othermiThe resultsof my studysupportthe general tochondrialprotein-codinggenes(e.g. Korneconclusionthat both typesof data are valuable gay et al. 1993,Hackett1996,Yoderet al. 1996).
for phylogenetic
inference(DonoghueandSan- The valueof differentialweightingfor rapidly

124

CAROLES.GRIFFITHS

evolvingpartitionsof molecularcharacters,as

GENUS Falco

shown in this research,has also been demon-

GENUS Microhierax[Auk,Vol.116

GENUS Polihierax
stratedpreviously(e.g.Simonet al. 1994,Sullivan and Swofford1997). For morphological
Tribe Caracarini(all generasedis
data,the levelof homoplasy
mayalsovary inmutabilis)
partitionsof data (e.g.in cranialversuspostGENUSPo[yborus
cranial osteologyin birds; Livezey 1986).FiGENUSIbycternally,homoplasymay not be as prevalentin
GENUSMilvago
GENUS Phalcoboenus
somemorphologicaldata setsbecausecharactersin thesedataare generallyfilteredbefore
GENUSDaptrius
phylogenetic
analysis(Livezey1986,Novacek
To minimize the numberof highertaxa inand Wheeler1992,Griffiths1994b).
taxaof equalrank within a
Phylogenetic
relationships
in the Falconidae.-- this classification,
monophyletic
highercategoryare listedin orCladogramsinferredfrom variationin syrinder;
each
taxon
is
the
sistergroupto the regealmorphology
and cytochrome-b
sequences
bothprovideevidence
thatthe two species
in mainingtaxaat thatlevel(Raikow1985).ProbthegenusDaptrius
arenotsistertaxa.Thesug- lematic relationshipsare indicatedby thegestionthat thesetwo be split hasbeenmade placementof a taxonas sedismutabilis(orderand interchangepreviouslybased on differencesin habitat use within groupis ambiguous
sedis(uncertainposition)withand foraging(Brown and Amadon 1968).The able)or incertae(Raikow1985).
nameIbycterVieillot 1816(type,by monotypy, in thehighercategoryThis
classification
divides
thefamilyintotwo
Falcoamericanus
Boddaert 1783) is resurrectedsubfamilies.
Herpetotheresisremovedfrom thefor Daptriusamericanus.
The name Daptrius
and placedin a subfamilywith MiVieillot1816(type,by monotypy,Daptrius
ater Falconinae
crastur.Herpetotherinae(Lesson,1843, type
Vieillot1816)is retainedfor Daptriusater.
Herpetotheres
Vieillot,1816)
has priority overI alsorecommendthat thetwo speciesof PoMicrasturinae
and is resurrected for that clade.
lihieraxbe splitbasedon polyphylyof thisgeis retainedandnusrevealedby syringealcharacters.
The sub- ThesubfamilynameFalconinae
includes the tribes Falconini and Caracarini.
stantial differencesof these speciesin size,
plumagecolor,and tail shapehavebeennoted The compositionof the Falconini is not
Withinthattribe,therelationshipofpreviously,with the suggestionthat they be changed.
insignisis unresolved.Polihierax
semplaced in different subgenera(Brown and Neohierax
Amadon 1968). The name NeohieraxSwann itorquatusand Microhieraxare sister taxa, a re1922 (type, by original designation,Polihierax lationshipsupportedby molecularsequences,insignisWalden 1872) is resurrectedfor Poli- syringealmorphology,and a morphometrichieraxinsignis.
The namePolihierax
Kaup1847 analysis(Kempand Crowe 1993).The Caracarini is a new tribe created to include the five ca(type, by monotypy,Falco semitorquatus
A.
Smith 1836) is retained for Polihieraxsemitor- racarageneraformerlyplacedin the Caracarinae.Relationships
ofthegenerain thistribearequatus.
Althoughthereis supportfor the
Therelationships
revealedin theanalysesof unresolved.
andsyringealmorphology(Fig.2), DNA sequence cladecontainingDaptriusater,Phalcoboenus
Milvago,furtherresolutionoftherelationships
data(Fig. 1C), and osteology(Fig. 1A; Becker1987)are reflectedin thephylogeneticclassifi- amongthesethreegenera,and of this cladetocationpresentedbelow,which is a revisionof the othertwo genera,awaita detailedphylogeneticanalysisof speciesand subspeciesinthe classificationof AOU (1998):

that tribe.

FAMILY Falconidae

SubfamilyHerpetotherinae
GENUSHerpetotheres
GENUS Micrastur

SubfamilyFalconinae
Tribe Falconini

GENUSNeohierax(incertae
sedis)
GENUSSpiziapteryx

Biogeography.--I
usedthe phylogeneticclassificationasa frameworkforexaminingthebiogeographyof the Falconidae.
Two approaches
basedon cladisticmethodology
canbe usedto
infer biogeographic
history.Cladisticbiogeography(i.e. vicariancebiogeography),thefirstapproach,wasdeveloped
asan alternativeto "center of origin" and "improbable"dis-January1999]

Falconidae
Phylogeny

125
Falco

persalscenariosprevalentin the 1970s(Nelson
1978,Bremer1992).Vicariancebiogeographyis
an attemptto recoverthe biogeographicpattern of an entirebiotathroughthe congruenceof multiplecladistichypotheses.
Forthismethod, an area cladogramrepresentingthe distribution of one group is interestingonly to theWorld-wide
India to theMicrohiorw(
PhilippinesßPolihiorax

AfticaSpiziapteryx•

Daptriu$
Phalcoboenu$•

extentthat it confirmsor conflictswith a general pattern.The alternativeapproachconsiders the historicalbiogeographyof an individual groupto be an importantpart of its natural
history(Bremer1992).Twomethods,ancestral-

Miivago South
Ibycter America
Polyborus•

Micrastur

Horpotothoro$

area (Bremer 1992) and dispersal-vicarianceanalysis(Ronquist1997),havebeendeveloped F•G.7. Area cladogramof the Falconidae.Distrithatusecharacter
optimizationto interpretthe butionof falconidgeneramappedontothe phylogbiogeographic
patternof an area cladogram. eny of the Falconidae.
The ancestral-areamethodologyestimatestheoriginaldistributionof a groupby compar- ly diversificationof the Falconidaeoccurredin
ing the numberof gainsversuslossesof areas the Neotropics.
under two models,forward (all gains) or reverse(all losses)Camin-Sokalparsimony(BreACKNOWLEDGMENTS
mer 1992).Forexample,if thereare moregains
than losses,the most-parsimonious
interpre- I thank G. Barrowcloughand R. Rockwellfor adtationis that the areawaspart of the ancestral viceandsupportthroughoutthecourseofthisstudy.
J. Bates,K. Burns, N. Caithness,T. Chesser,J. Groth,

distribution.

Dispersal-vicarianceanalysisusesFitchop- S. Hackett,and N. Klein provideduseful conversation and thoughtfulcriticisms.B. Livezey and C.
timizationto reconstructthe historyof bioge- Crajewskiprovided valuablecommentson the manographyby minimizingthe dispersal-extinc-uscript.I thank the followingcuratorsand institutioneventson the areacladogram.
A stepma- tionsfor providingtissuesused in this research:G.
trix, whichspecifies
the costof combinationsof Barrowclough(AMNH), E Gill (ANSP), E Sheldon
ancestraland descendantdistributions(the difference between the distributionof the ancestor(LSUMNS), and M. Braun (USNM). This work was

supportedby grantsfrom the Frank M. Chapman
Memorial

Fund of the American Museum of Natural

andeachdaughterarea),isusedin a three-pass
History, a Frank M. ChapmanPredoctoralFellowalgorithm to find the optimal reconstruction ship, NSE DissertationImprovementGrant BSR(Ronquist1997).

9321486,and City University of New York Disserta-

I usedthe basicprinciplesof the secondap- tion ImprovementGrants. The researchreported
proachto inferthebiogeographyof theFalcon- hereis a contributionfromtheLewisB. andDorothyidae.The areacladogram,illustratingthe dis- CullmanResearchFacilityat the AmericanMuseum
tributionsof the falconidgeneramappedonto of Natural History and has receivedgeneroussupthe phylogeny,
is shownin Figure7. The pat- port from the LewisB. and DorothyCullmanProgram for MolecularSystematics
Studies,a joint initern of distributionof thesegeneraenablesa tiative of the New York Botanical Garden and thesimpleinterpretationof biogeographic
history AmericanMuseum of Natural History.
for the family.Distributionof speciesin two of
the threelineages,and the basalbranchin the
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Associate Editor: R. M. Zink

128

CAROLE
S. GRIFFITHS[Auk, Vol. 116

APPENDIX
1. Descriptionsof syringealanatomyand the 23 syringealcharacters;17 charactersare binary
and 6 are multistate. Characters 3, 4, 6, and 16 are ordered multistate; characters5 and 19 are unorderedmultistate.In all cases,the hypothesized
plesiomorphic
stateis state0.
Tympanum:FusedA elementsnearthe tracheo-bronchialjunction(1) Presenceof a tympanumor trachealdrum[0] No fusion of A elements[1] Ossification and fusion of first A elements(2) Dorsal fusionof tympanumby medial bar[0] Medial bar absent[1] First A elementsfused medially by ossifiedbar(3) Fusionalongedgesof A elementsthroughossificationof cartilaginous
bordersseparatingelements[0] Ossifiedbordersnot apparent[1] Threeor four A elementsfusedalongmarginsof elements[2] Heavierfusion,marginssomewhatobliterated,light suturesapparent(4) At least five A fused entirely along edges[0] Light fusionor fusionabsent[1] First five elementsheavilyfused,somesuturesapparent[2] Suturesobliterated(5) Degree of ventral fusionof tympanum[0] No ventral fusion of A elements[1] First threeor four A elementsfusedpartially alongmargins[2] First threeor four A fusedentirelyalongmargins[3] At leastfive A elementsfusedentirelyalongmargins,somesuturesapparent(6) Shapeof fusedand ossifiedtympanum(partiallyorderedcharacter;state3 is a modificationof state2)[0] Tympanum absentor not ossified[1] Tympanum graduated,widens caudally[2] Tympanum cylindrical[3] Tympanumcylindrical;A1 laterallyflattenedcausing"pinchingin" of most-caudalelement
A Elements:Occuron the tracheaassingleringsbut may extendontothe bronchias paireddoublerings;
they are ossifiedand flattenedin cross-section(7) Ridge coveringdorsalA1 ends[0] Ridge absent[1] Ridgeof ossifiedtissueforms medial ridge over A1 ends,coversdorsalascentof pessulus(8) Separationof dorsalA1 ends[0] A1 endsseparated[1] Endsclosemedially,fusedby ossifiedtissue(9) Size of A1 ends[0] A1 singleor width of endssimilarto width medial[1] Dorsal endsvery enlarged(10) Flatteningof paired A1 elements[0] A1 singleelement,or roundedhalf rings[1] A1 flatteneddorsoventrally
into parentheticalshape;laterally,A1 dorsalandventralendsprotrude(11) Appearanceof A1 on lateral view[0] A1 concaveup medially[1] A1 flattenedmedially
B Elements:Occuras paired rings on the bronchiand are generallycartilaginousand D-shapedin crosssection(12) Abrupt changein shapeand orientationof B1 ends[0] B1 endschangegraduallyor are unchanged[1] B1 endsthickand wide,ascendabruptlyin L-shapeto fusewith A1(13) Changein orientationof B1 ends[0] B1 endschangeshapeor endsmaintainorientation[1] B1 endsthin, ascendgraduallyto fusewith A1(14) Knobbingof B1 ends[0] No knobbing[1] Craniadedgesknobbed,fusedwith A1 ends
(15) Gradualchangein shapeand orientationof B1 ends[0] B1 endsthin or L-shaped[1] B1 endsthicken;endsare roundedand ascendgraduallyto fusewith A1(16) Fusionof B element ends[0] No fusion of B elements[1] B2 ends fused to B1/A1[2] B3 endsfusedformingridge borderingmedial (internal)membraneJanuary 1999]
APPENDIX 1.
Muscles

FalconidaePhylogeny
Continued.

and Membranes(17) Existenceof lateral (external)membraneon bronchi '[0] Membrane absent[1] Membranepresentand locatedbetweenA1 and B1(18) Presenceof cartilaginousbar on lateral membrane[0] Bar is absent[1] Bar is presentM. tracheolateralis
insertson bar(19) Presenceof membranousextensionof lateralmembraneontowhich M. tracheolateralis
inserts[0] Membranous extension is absent[1] Thick, bulbous membrane on dorsal half of membrane[2] Thick, bulbousmembrane coversentire width of membrane(20) Insertionof M. tracheolateralis
onto lateralmembrane[0] Membrane absent or modified[1] M. tracheolateralis
insertsdirectlyontodorsalhalf of lateralmembrane(21) Cartilaginousborder on (medial) membrane[0] Border is absent[1] Internal (medial)membranehassomecartilageon caudalborder(22) Completeborder on internal membrane from A1 dorsal to A1 ventral ends[0] Borderabsentor incomplete[1] Thick, even, concaveborder on internal membrane from A1 dorsal to A1 ventral ends

Accessory
CartilaginousStructures:
Occuron internalbronchialmembranes(23) Presenceof thin, amorphousaccessory
structureextendingfrom border[0] Accessorystructureabsent[1] Cartilageformsstraightcaudaledgefrom dorsalto ventralends

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S. GRIFFITHS[Auk, Vol. 116