The Auk 116(1):116-130, 1999
PHYLOGENY
OF THE FALCONIDAE
INFERRED
FROM
MOLECULAR
AND MORPHOLOGICAL
DATA
CAROLE S. GRIFFITFIS •
Department
of Ornithology,
American
MuseumofNaturalHistory,CentralParkWestat 79thStreet,
New York, New York 10024, USA
ABSTR^CT.--Molecular
data and variationin syringealmorphologywere usedto infer a
phylogenyfor 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
signalcaused
by rapidly changingsitesin the moleculardata were not solvedby combiningdata sets.
Differentiallyweightingsaturatedpartitionsof the sequencedata, prior to phylogenetic
analysis,provideda phylogenycongruentwith morphological
analysis.Moleculardataprovide substantially
moreinformativecharacters
thanmorphological
data.However,morphologicaldataprovidea higherproportionof unreversedsynapomorphies.
A reclassification
of 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•ephylogenyimpliesthattheoriginandearlydiversification
of thefamilyoccurredin SouthAmerica. Received
12 November
1997,accepted
18 June1998.
THE FAI•C¸NIDAE,one of three families in the
orderFalconiformes
(thediurnalbirdsof prey),
includessomeof the fastestand mostspectacular birds in the world (Brown and Amadon
1968).Althoughthefamilyis renownedfor the
huntingabilityof its species
(e.g.thePeregrine
Falcon[Falco
peregrinus]),
foraginghabitswithin thefamilyarediverse.Thus,theNeotropical
caracaras(Daptrius,Milvago, Polyborus,and
Phalcoboenus)
are scavengers
that subsistmainly on carrionor invertebrates.
In additionto adaptationsrelatedto foraging
habits,the four caracaragenerashareother
morphological
traits(Friedmann1950)andhistoricallyhavebeenconsidered
to be closelyrelated(Sharpe1874,Gurney1894,Swann1922,
Peters1931).Classification
of the othergenera
in thefamilyhasbeenlessstable(Sharpe1874,
genera:thefourcaracaragenera,Micrastur(forest-falcons),Herpetotheres
(Laughing Falcon)
and Spiziapteryx
(Spot-wingedFalconet).The
relationships
of thelatterthreegenerahistoricallyhavebeenproblematic
(Sharpe1874,Gurney 1894,Swann1922,Peters1931).Theplacementof thesegenerawith the caracaras(Amadon and Bull 1988) is in conflictwith results
from four cladisticanalyses:(1) an osteological
study (Becker1987;Fig. 1A), (2) a preliminary
morphologicalanalysis (Kemp and Crowe
1990;Fig. 1A), (3) an analysisbasedon syringealmorphology
(Griffiths1994a;Fig. lB), and
(4) an analysisbasedoncytochrome-b
sequences (Griffiths1997;Fig. 1C). Basedon Griffiths
(1994a),theAOU (1998)reclassified
thefamily,
placing Herpetotheres
and Spiziapteryx
within
the Falconinae and Micrastur
in its own sub-
Gurney 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
continuum
of ideas. At one end is the idea that
data setsshouldbe analyzedseparately.The
E-mail:[email protected]
116
January1999]
A
t
B
c
Falconidae
Phylogeny
•
117
nentsof totalevidenceproposegivingall charactersequal weight initially in phylogenetic
analysis.Assessments
of 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,
Brewer
and
DeSalle
1994,
Chippindale
Polihierax
insignis
and Wiens 1994).
Horpotothoros
As part of the empiricalexplorationof the
Caracara
species
analysis
of multipledatasets,I examinethe efMicrastur
species
Outgroups
fectsof differentialweightingon phylogenetic
inference.I then comparethe morphological
and moleculardata. The phylogenyinferred
Falco
from thesedatais usedto reclassifytheFalconidaeandtwo generawithin thatfamily.Finally,
Microhiorax
the phylogenyprovidesa frameworkfor disPolihioraxsomitorquatus
cussingthebiogeography
of thefamily.
Spiziapteryx
Falco
Microhierax and Polihierax
Spiziaptoryx
•
•
Caracaraspecies
Horpotothoros
Micrastur
species
MATERIALS
AND METHODS
Morphological
data.--Characters
werederivedfrom
variation in syringeal supportingelements,memFIC. 1. Phylogenetic
hypotheses
inferredfrom branes,andmuscles(Griffiths1994a).Supportingelcladisticanalysesof the Falconidae.
(A) Phylogeny ementsincluderinglikeelementson the tracheaand
inferred from osteologicaldata (Becker1987) and a bronchi(A andBelements;Ames1971), thepessulus,
preliminaryanalysisof skinsandcharacters
fromlit- and accessory
cartilaginousstructures.Hemologyof
erature(Kempand Crowe1990).(B) Phylogenyin- the ringlikeelementswastraditionallybasedon the
Accipiter
ferredfromsyringealmorphology
(Griffiths1994a). relativepositionof thesestructuresto the tracheo(C) Phylogeny
inferredfromdifferentially
weighted bronchial
junction(King1989).Thisanalysis
postucytochrome-b
sequences
(Griffiths1997).
lateshomologous
elementsfollowingAmes' (1971)
definitionsof syringealstructures(Griffiths1994b).
Dataon variationin syringealmorphology
fortwo
congruence
of 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 B1
element,changed
fromoneto zero.Character
3, dorsal fusionof the tympanum,changedfrom one to
i.e.combination
onlyif thedatasetsarenotsignificantlyheterogeneous
(Bullet al. 1993,Huel- two. In addition,threecharacters(characters
5, 6 and
senbecket 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
*
*
*
no
*
*
*
*
*
no
no
*
*
no
no
E vespertinus
E femoralis
Accipiterstriatus
Otus asio
Pelecanus onocrotalus
matrixfor the reanalysisconsisted
of 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.
missingcharacters
in themorphological
datasetreMoleculardata.--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-membrane
the
Falconidae(Griffiths 1994b),was used to root the
regionsof 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 rootedthe
fiths 1997). Osteologicalcharacters(Becker1987) cladogramin the sameplaceand producedthe idenwere not included because the data matrix for that
tical phylogenyas the analysiswith multiple outanalysisand details of the analysiswere not avail- groups.The additionaloutgroupspecies
weredelet-
January
1999]
Falconidae
Phylogeny
ed to facilitatethe detailedexaminationof changein
molecularcharacters
within the family.
To test the differentmethodsproposedfor combining data for phylogeneticinference,two setsof
phylogeneticanalyseswere performed.In the first
set (i.e. the total evidencetest), all characterswere
weightedequallyfor an initial phylogenetic
analysis.
Data were thenweightedagainto assess
the effects
of a posteriori
differentialweighting.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 the
first method, the maximum value of the rescaledcon-
sistencyindexwasusedto reweightcharacters
(suc-
Phalcoboenus
2 23
16
cessiveapproximations;Farris 1969, 1989;Carpenter
1988). In the secondmethod, the characterswere
PolihieraXinsignis
3
Spiziapteryx
3
Microhierax
Polihieraxsernitorquatus
weightedconcurrentlywith 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 femoralis
of characters
to accommodate
heterogeneity
of data
Falcosparverius
sets (unconditionalcombination;Chippindale and
Falco 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 the
syringeal
morphology;
F
=
Falconinae,
C = Polyboradditionof the morphologicaldata on supportfor
inae.
Cladogram
represents
the
strict
consensus
tree
nodes.Bootstrapanalyseswere performedon the
of 54most-parsimonious
treesof length50;CI = 0.59
molecular data and then on the combined data to
provide 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 morphological
characters
wasexamined.
Ordering effectivelyweights character-state
transformationsby 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,morphological
characters
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 the
ordering morphologicalchangeson support for
osteologicaldata (Becker1987),the molecular
nodes, bootstrap analyseson the combineddata
data
were 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 of
heuristic algorithm, which included 100 replicates coninae clade rather than with the caracaras.
that randomlyvaried the order in which taxa were The positionof Herpetotheres
in thistree differs
added.
from the originalsyringealanalysis(Griffiths
1994a;Fig. lB).
RESULTS
As 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. GRIFFITHS
A
i_•j.• caracara
species
I
•
Spiziapteryx
I
[•---Falcospecies
I
--•Mlcrohierax
[Auk,Vol. 116
erain the family (BrownandAmadon1968).In
addition,the node supportingmonophylyof
theMicrasturspecies
is oneof themoststrongly
supportedin the morphologicalphylogeny
(Fig. 2), with four synapomorphies,
three of
these with a CI of 1.0. The alternative result, in-
L........
Z;;[;;;Td.
TvZ;
u'us
.............
Accipitor
ferredusingdataweighteddifferentiallyprior
to phylogenetic
analysis,is congruent
with the
morphologicalhypotheses;
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 only
moleculardata(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
Accipiter
thesecladesnorof the relationships
of thetwo
FIG. 3. (A) Phylogenyinferred from equally
weightedmoleculardata(Griffiths1997).(B)Phylogeny inferredfrom combineddata setswith equally
weightedmoleculardata. Strict consensus
tree of
five most-parsimonious
cladograms
(length= 1,089,
CI = 0.46,RI = 0.45),morphological
characters
unordered.The samecladogramsare inferred if morphologicalcharacters
are treatedasordered.Dashed
linesconnectspeciesin the genusMicrastur.
Micrasturspeciesand Herpetotheres.
A posteriori
differentialweighting,usingassessments
of characterreliabilityderivedfrom
the phylogeneticanalysis(i.e. successive
approximations),
or weightsderivedduringphylogenetic analysis, produced one cladogram
similarto that inferredfrom equallyweighted
moleculardata(Fig.3A). Resultsare the same
with the morphological
characters
orderedor
unordered.
Spiziapteryx
is
sister
taxon
to thecanusin eitherdifferentsubgenera
or generahas
racaras,
and
the
genus
Micrastur
is
not
monobeen advocated(Brown and Amadon 1968).
phyletic.
Analysis
of molecular
data.--Results
fromthe
Combined data: Unconditional combination.analysisof variationin cytochrome-b
sequences
Combining
data setswith a prioridifferential
are describedin detail in Griffiths (1997). A
weighting
of
moleculardata(i.e.unconditional
summaryis presentedto serveas background
combination)
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,this
is 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 molecular
more anomalousresultis that the genusMi- data [Fig. 1C] and syringealmorphological
crasturis notmonophyletic.
Contraryto there- data [Fig. 2]). Differencesbetweenthe comsuitsfor Daptriusand Polihierax,
polyphylyof bined and separateanalysesappearin two arof
the Micrasturspecieshasnot beensuggested easof thetree:(1) thesistertaxarelationship
previously.The specieswithin this cladeare MicrasturandHerpetotheres
is unresolved
in the
similar 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
reviews
of
this issuehavebeenpresentedpreviously(Hillis 1987,de Queiroz et al. 1995,Miyamoto and
Fitch 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 relevant evidence;Kluge 1989). Analyzing data
setsseparatelyis disputedbecause
thecreation
of separatedatasetsassumes
thattherearenatural classesof data, an assumptionthat is not
justifiedscientifically
(Kluge1989).This argumentcarriesoverinto the appropriatemethod
of weighting characters.A priori character
weighting requires partitioning data and
would thereforebe unjustifiedby thosewho
Daptrius
americanu
Z
Micrasturgilvicollis
M. semitorquatus
(•.•oo)
HerpetoEleres
FIC. 4. Phylogenyinferred from combineddata
setswith a prioridifferentialweightingof molecular
data, and morphologicalcharacters
orderedor unordered. F = Falconinae,C Polyborinae(caracaras).
Strictconsensus
tree of two most-parsimonious
clad-
ograms.Bootstrapvaluesat nodesareshownfor the
molecular data set and for the combined data sets;
numbers
above the branches are for the molecular
data only,numbersbelow the branchesare for the
combineddatasets.Rangeof numbersfor the combined data sets are for ordered
and unordered
mor-
phologicalcharacters
(left = unordered,right = ordered).
canusare sistertaxa, a result congruentwith
the morphologicalanalysis.In the secondtree,
Polyborus
is basalto the othercaracaras.
Bootstrapanalysisof the combineddataindicatesadditionalsupportfor key nodescomparedwith the analysisof moleculardataalone
(Fig.4). Thepositionof Spiziapteryx
in theFalco
cladeis not supportedin the majorityof bootstrap replicateswith molecular data alone.
When morphologicaldata are added, that relationshipis supportedin 61 to 71%of the replicates.The supportfor monophylyof Micrastur
increasesfrom 72% to 94 to 97%, and the support for the sister-taxarelationshipof the caracara
and Falco clade increases
from
65 to 78
believe
that
data
should
be combined
before
analysis.A posteriori
weighting,determinedeither iterativelyor directly,is appropriatebecauseweightsare inferred throughphylogenetic analysis.
Separateanalysisof multipledatasetsis supported by the argument that congruence
amonghypotheses
from differentdatasetsis a
powerfultool for assessing
accuracyof the hypotheses(Miyamoto and Fitch 1995).An assumptionof this methodologyis that dividing
data into classescan be justifiedscientifically
(e.g. different codonpositions).Advocatesof
the initial separateanalysisof data setsdiffer
on the final treatment of multiple data sets.
Opinionsrangefrom alwayskeepingdata sets
apart (Miyamotoand Fitch 1995) to always
combiningdata sets,after any conflictsare accommodatedthrough differential character
weighting(Chippindaleand Wiens 1994).An
intermediatepositionis to combineonly congruent data sets,using either biological(e.g.
genetreesfor a species;de Queirozet al. 1995)
or statistical (significant differencesamong
to 82%. Orderingthe four multistatemorphologicalcharacters
doesnot changethe topolo- data sets;Huelsenbecket al. 1996) criteria for
gy, but it provides strongersupport for six of
determiningincongruence.
One problemwith
sevennodeson the cladogram(Fig.4).
this approachis that the two criteria are not
equivalent;datasetsrespondingto differentbiDISCUSSION
ologicalprocesses
may or may not differ sigAnalysis
ofmultipledatasets.--Oneof themost nificantly.In addition,althoughvarioussignifcontentiousissuesin systematicsis the treat- icancetestshavebeenproposed(bootstrap[de
122
CAROLE
S. GRIFFITHS
Queiroz1993],tree lengths[Farriset al. 1995],
and likelihood ratios [Huelsenbeckand Bull
1996]), the power and sensitivityto the assumptionsof thesetestsare unknown.
If the goalof phylogenetic
analysisis to provide the hypothesis
with the greatestexplanatory power,then comparingthe variousmethods is unnecessarybecauseonly a combined
analysismeetsthatgoal.If thegoalis accuracy
of phylogenetic
inference,thenempiricaltests
comparingthesemethodsare useful(de Queiroz et al. 1995).Oneimplicitassumption
of the
total evidenceapproachis that the sameunderlyingsignalis containedin eachdata set
(Hillis et al. 1996).Thus,any potentialconflict
amongthe hypotheses
inferredfrom individ-
[Auk,Vol. 116
stateat nodes(Kim 1996).Thesekinds of characters occur in the molecular data in this re-
search.For thesedata, ratesof changediffer
among partitions of sequences,
with several
partitions containingcharacterswith high
ratesof change(Griffiths1997).Substitutions
are saturatedin thesepartitions,and recovering phylogeneticsignalis problematic(Griffiths1997).Whena prioridifferentialweighting
is usedto correctfor the high ratesof change
in these partitions, the phylogenyindicates
monophylyof Micrastur,a result congruent
with the morphological
analyses.
Theseresultssupportthe intermediateposition,the initial separateanalysisof multiple
data sets,and differentialweightingto accomual data sets will be overcome when data sets
modateheterogeneity
in ratesof change.After
are 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
was
tent methodology
will resultin convergence
to accompaniedwith assertionsthat molecular
the correcttopologyasdataare added(Felsen- evidencewasmorereliablethanmorphological
stein 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 cladisticmorphological
analyses
andwith 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 assessments
of characterreliability to molecularcharacterswere an order of magnireweightthe characters
(aposteriori
differential tude greater in number than morphological
weighting)also resultedin polyphyly of Mi- characters(328 vs. 23). The relativepaucityof
crastur.
Thisis not surprising;searches
usinga characters
is 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 assumptions
of the methodare
Theadvantage
of thesyringealdatasetisthe
violated (e.g. Debry 1992, Huelsenbeck1995, proportionof unreversedsynapomorphies;
15
Swofford et al. 1996, Sullivan and Swofford
of 31 charactersstateshave a C! of 1.0, and 40%
1997). Theseviolationscan be accommodated of the characters
changeonlyoncein 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 obscures
the 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
0
0-0.24
0.25-0.49
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 characters
with a given
numbersof steps.(A) Morphologicaldata, (B) mo-
10.
lecular
data.
5'
0
0-0.24
0.25-0,49
0.50-0.74
0.75-0.99
1.00
Consistency
Index
FIG.5. The proportionof charactersfor eachvalue of the consistency
index.(A) Morphological
data,
(B) molecular data.
nodes.Alternatively,additionalsequencedata
may not resolvethesebranchesif the additional
datahavethe samepatternsof noiseandsignal
asthe presentdata (Brunset al. 1992)or if the
data do not fit the assumptionsof the model
used (Sullivan and Swofford 1997). Because
these genera are monotypic,increasedsampling of taxawithin the familywouldprobably
not resolvethe problem.
The resultsof my studysupportthe general
conclusionthat both typesof data are valuable
for phylogenetic
inference(DonoghueandSan-
derson 1992, de Queiroz et al. 1995). Differen-
tially weightedmoleculardata stronglysupport the sisterrelationshipof Herpetotheres
and
Micrastur,which is unresolvedin the morphological tree. The positionof Spiziapteryx
has
weak supportin the moleculartree.The inclusionof Polihierax
insignisin the morphological
analysis(Fig. 2) strengthensthe supportfor
Spiziapteryx
within the Falcoclade.
In this study,the moleculardata were more
homoplastic,thus disputingthe assertionthat
moleculardata are more reliablethan morphologicaldata for phylogeneticinference(Sibley
and Ahlquist 1987, Hedgesand Sibley1994).
The occurrenceof saturateddata partitionsin
the falconidsequences
follows a pattern illustratedin studiesof cytochromeb and othermitochondrialprotein-codinggenes(e.g. Kornegay et al. 1993,Hackett1996,Yoderet al. 1996).
The valueof differentialweightingfor rapidly
124
CAROLE
S.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 in
mutabilis)
partitionsof data (e.g.in cranialversuspostGENUSPo[yborus
cranial osteologyin birds; Livezey 1986).FiGENUSIbycter
nally,homoplasymay not be as prevalentin
GENUSMilvago
GENUS Phalcoboenus
somemorphologicaldata setsbecausecharactersin thesedataare generallyfilteredbefore
GENUSDaptrius
phylogenetic
analysis(Livezey1986,Novacek
To minimize the numberof highertaxa in
and Wheeler1992,Griffiths1994b).
taxaof equalrank within a
Phylogenetic
relationships
in the Falconidae.-- this classification,
monophyletic
higher
categoryare 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 the
gestionthat thesetwo be split hasbeenmade placementof a taxonas sedismutabilis(order
and 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 thehighercategory
This
classification
divides
thefamilyintotwo
Falcoamericanus
Boddaert 1783) is resurrected
subfamilies.
Herpetotheres
is
removedfrom the
for 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 over
I alsorecommend
that thetwo species
of PoMicrasturinae
and is resurrected for that clade.
lihieraxbe splitbasedon polyphylyof thisgeis retainedand
nusrevealedby 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,therelationship
of
previously,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 morphometric
hieraxinsignis.
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 thistribeare
quatus.
Althoughthereis supportfor the
Therelationships
revealedin theanalyses
of unresolved.
and
syringealmorphology
(Fig.2), DNA sequence cladecontainingDaptriusater,Phalcoboenus
Milvago,
further
resolution
of
the
relationships
data(Fig. 1C), and osteology(Fig. 1A; Becker
1987)are reflectedin thephylogenetic
classifi- amongthesethreegenera,and of this cladeto
cationpresentedbelow,which is a revisionof the othertwo genera,awaita detailedphylogeneticanalysisof speciesand subspecies
in
the classification
of AOU (1998):
that tribe.
FAMILY Falconidae
SubfamilyHerpetotherinae
GENUSHerpetotheres
GENUS Micrastur
SubfamilyFalconinae
Tribe Falconini
GENUSNeohierax
(incertae
sedis)
GENUSSpiziapteryx
Biogeography.--I
usedthe phylogenetic
classification
asa frameworkforexaminingthebiogeographyof the Falconidae.
Two approaches
basedon cladisticmethodology
canbe usedto
infer biogeographic
history.Cladisticbiogeography(i.e. vicariancebiogeography),
the
firstapproach,
wasdeveloped
asan alternative
to "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 congruence
of multiplecladistichypotheses.
Forthismethod, an area cladogramrepresentingthe distribution of one group is interestingonly to the
World-wide
India to the
Microhiorw(
Philippines
ß
Polihiorax
Aftica
Spiziapteryx•
Daptriu$
Phalcoboenu$
•
extentthat it confirmsor conflictswith a general pattern.The alternativeapproachconsiders the historicalbiogeography
of 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-vicariance
analysis(Ronquist1997),havebeendeveloped
thatusecharacter
optimizationto interpretthe
biogeographic
patternof an area cladogram.
The ancestral-areamethodologyestimates
theoriginaldistributionof a groupby comparing the numberof gainsversuslossesof areas
under two models,forward (all gains) or reverse(all losses)Camin-Sokalparsimony(Bremer 1992).Forexample,if thereare moregains
than losses,the most-parsimonious
interpretationis that the areawaspart of the ancestral
F•G.7. Area cladogramof the Falconidae.
Distributionof falconidgeneramappedontothe phylogeny of the Falconidae.
distribution.
J. Bates,K. Burns, N. Caithness,T. Chesser,J. Groth,
ly diversificationof the Falconidaeoccurredin
the Neotropics.
ACKNOWLEDGMENTS
I thank G. Barrowclough
and R. Rockwellfor adviceandsupportthroughoutthecourseofthisstudy.
Dispersal-vicariance
analysisusesFitchop- 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 combinations
of Barrowclough(AMNH), E Gill (ANSP), E Sheldon
ancestraland descendantdistributions(the difference between the distribution
of 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).
I usedthe basicprinciplesof the secondapproachto inferthebiogeography
of theFalconidae.The areacladogram,illustratingthe distributionsof the falconidgeneramappedonto
the phylogeny,
is shownin Figure7. The pattern of distributionof thesegeneraenablesa
simpleinterpretationof biogeographic
history
for the family.Distributionof speciesin two of
the threelineages,and the basalbranchin the
9321486,and City University of New York Dissertation ImprovementGrants. The researchreported
hereis a contribution
fromtheLewisB. andDorothy
CullmanResearchFacilityat the AmericanMuseum
of Natural History and has receivedgeneroussupport from the Lewis B. and DorothyCullmanProgram for MolecularSystematics
Studies,a joint initiative of the New York Botanical Garden and the
AmericanMuseum of Natural History.
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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 unordered
multistate.In all cases,the hypothesized
plesiomorphic
stateis state0.
Tympanum:FusedA elementsnearthe tracheo-bronchial
junction
(1) Presence
of 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 elementsthroughossification
of 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 parenthetical
shape;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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CAROLE
S. GRIFFITHS
[Auk, Vol. 116