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ZOOLOGICAL SCIENCE 19: 1363-1375 (2002) @ 2002 Zoologica lSociet oyt Japan
NewSpecies of Metacaligus(CaligidCaoep,epoda)Parasitic
on the Cutlassfis(hTrichiuireupsturuS)of Taiwan,
CladistiAcnalysis the FamilyCaligidae
with a of
Ju-she yHoi" and Ching-Lon gLin2
" Department of BiotQgia aSlciences ,Calliomi aState Uhiversity;
Long Beach, Caliiomi a9,0840-37C1 2U;SA
2Dopartment BiosciencesN,ationaCJhiayUinivetsity;
ofAquatic
Chiayi6,0083,faivv:an
ABSTRACT-Metacaligus latu sn. sp. (Copepoda C,aligidae i)s describe dfrom specimens found p'alraaiswiatnic
in the oral and gil clavities of the cutlassfish, 77ichiuru slopturu sLinnaeus ,caught from the Strai otf
and landed at Dong-Shih Fishin gPort in Chiay iCounty l,aiwan ,l tis close to M, uruguayensis <Thomsen,
1949) ,but can be distinguish efdrom the latte bry the possessi oofn shorter caudal rami in both sexes and
wider cephalothorax and genit aclomplex in the male. Cladist i(cphylogen eatnailycsi)s of the Family
Caligida eb,ased on 23 selected morphological characters, revealed tha tMetacaligus Thomsen, 1949 is
distant lryelated to Caligus M"11er, 1758. I tis a valid genus and occurs in sister-group relationship with
Apogonia Cressey and Cressey ,1990.
Key words:parasitic Copepoda, Metacatigus latus c,utlassfish, caligi dcladistics
zuela, Ho and Bashirull a(h1977 r)eevaluated Thomsen's
INTRODUCTION (1949 c)oncept of MetacaLigu sand elevated i tto the generic
Two specjes of Metacaligus Thomsen, 1949 are known rank, Moreover, after examining the type-mater iofa lthe
parasit ionc the marinefishes of East Asla :M. hisae (Shen, three specles of Catigu sinclude din the Metacaligu sby
1957) from China and M. uruguayensis Thomsen, 1949 trom Thom$en (1949 )Ho, and Bashirull a(h1977 e)xcluded C.
Taiwan {Li annd Ho, 2000) ,Whil ethe tormer is confined to afurcatus and C, enomiis, and include dCaLigu hsilsa eShen,
China (Shen 1,957) the latt ehras been reported from Uru- 1957, in the genus Metacaligus.
guay (Thomse n1,949) ,lndi a(Pil l1a96i3,) ,Sr iLanka (Kirtis- The rationale for Ho and BashiruHa h{197 7t)o elevate
inghe ,1964) ,Venezuela (Ho and Bashlrulla h1,977) and Metaaaligu sto the generi rcank was the possessio nin com-
Braz i{lBoxsh aanldl Monta, 1997) ,ln thi spaper we report a mon of a combination of three character state$ by M, uru-
new species of this genus recovered from the cutlassfish, guayensis ,M. rufus, and M. hisae. Those three common
77iChiur ulsepturus Linnaeus ,caught in the Straigh tof Tai- morphological properti easre (1 )absence of sternal furca,
wan, (2 )absence of accessory processes on two middle terminal
ln his work on the copepod parasite sof marine fishes spines of second exopodal segment ot leg 1, and (3 )pos-
of Uruguay, Thomsen (194 9p)roposed the subgenus Meta- session of only one outer spine (inste aofd two) on the ter-
caiigus to accommodate a new species of Caligus found minal exopodal segment of leg 2. Since none ot the three
parasit ionc the cutlassfish from Uruguay I. twas called "Cafi- states of character are unique to MetacaLigu sand each of
gus (Metacali gutuuqgu)ayensis n. sp." Furthermore ,he them is respectivel yfound in some species of Caigus, Dojiri
include din this subgenus three other species of Catigus, (198 3wa)s skeptical about the recognition of Metacaligu sas
namely, C, afurcatus Wilson ,1913; C. enormis Wilson, a genus distin cftrom Caiigus .Therefore ,in thi sreport, in
1913; and C, rufus Wilson ,1908. Twenty-eigh tyears later, addition to providin gthe descripti oonf a new species of
on the occasion of the discover yof C, (Metacali guruu-q) MetaaaLigu sa, cladistic analysis based on the morphological
guayensi fsrom the same species of host caught off Vene- characters of the Famil yCaligid aies attempted, l tjs
believe dthis analysis shall provid emeans to explore phylo-
+ Correspondi anugthor:Tel.FAX . ++11--55626-29-895-8458-182;87B. gesetnaebtlii csrhemleatnito nsohf ipM$e twiatchainl itghue sfasa mai l gyaenndu st o seexpaamriantee it frtohme
E-mail:jsho@csulb,edu Caligu iss justifiable,
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MATERIALS AND METHODS Abdomen stocky and 1-segmented ,O.56 (O,40-O .m8m2)
long and O,23 (O, 81-O,38 )mm wide. Caudal ramus {Fi g3C.)
The cuUassfish (7?ichi utreputsuru sLinnaeus) caught in the 1.63 <O.13xO ,mm0>8 times longe rthan wide; with 3 short,
SCaytoiru aUnin ttioy vfw eaTsrasi ipwt uawyrhnec rheaands te hdl eaa nlndad betodr raaant stfoDero reynrxagemd-i nSiahnt iiaonhn Fiic seohbfio tnx htg eoP coNorapett pioidon nCah lCpihaairy--i sau bsteetrumlien-ablear isnegta ep aanpdi l3 l oanlo ndgo ,rtsear lmsuirnfaa clsee.ta eE gig n asdadci tiloonn gteor
asites was carried out. The copepod parasite sremoved from the than body, 3.9 mm and containing 35 eggs.
fis hwere preserve idn 70% ethanol. In studying the parasit etsh,e Antennule (Fi gI,B) 2-segmented; proxim aslegment
specimens were cleared in 85% lacti accid fo r1 to 2 hours before with 27 setae on anterodistal surface, dista slegment with 1
mdtahekev iindsgies d sdbeiycs tsHeeu dcmpteaisro ti ns an naadn dd rG ooappo pdoefnid nalggae sc(t1i ac9c i6du.n4 d weT)arshe ethmhapen logcyoiemndpgo-u dntdroo epm xeamtmihiconrdeo- sauebsttehremtiansacls soenta doins tpa olmsatrgeinr.i mo arrAginnte nannad (1F1 i sgett.aCe) p3-l$ues g2-
scope. Al ldrawings were made with the aid of a camera lucida, mented; proximal segment smallest and unarmed; second
Measurements of the body and body parts were taken from tO ran- segment rectangular and also unarmed; dista lsegment
domly selected specjmens, The measurements given in the stocky claw bearin g1 seta in proxima lregion and another
dth eas t cf1roni ltphltiois w opa.nraep etrh eon alvye qraugaeli wtiatth i icvthe asrraancgteer ssh oawrne usien dth ei n ptahre ecnlatdhisetsices osmnael lc laon$de tbo lmuidndtle lyp oriegnitone,d ,Pcaorsrytianngt e2n nba alpsrao lcpeasps i(llFaie wgIi,Cth)
analysis. Also ,autapemorphic characters are excluded because each bearin g4 setules; another similar papit llaocated
they do not aid in a search tor the wanted phylogeneti crelation- nearby on sternum. Mandible (Fi gI,D) 4-segmented; with
ships, except for help in boostin gconsistency index of the gener- 12 teeth on medial margin of dista bllade .Maxillu l(eFi g1,C)
asBaetsaleiesdtceti ed ccdol noan dtot hgoB reee axvnmterz{rats'ce)tsb .r( a tt1Tehsh9e,e w9 atn3 tcyEe)lpldaedi-orgs ryphaepmochmioeorsl uf o sMoogifro tluehsane c eEhcsdhi apwvrhaaaolcnritodd escsrta so,1lm i8afgt4oio0r di t dgahsneed nO uTphsUe a.i-rs- cbmeoanmstpeardl;i sipn agppr ioslx hliowrmiatta,hl s3set gonmuaektne ,tdde ns(tetliafea.ocr emMprar xtoilcualesrl sga)s (e aFanndid g I p.Eu)rn aor2mm-eisdn;eegn-t
bius Kreyer ,1838 were utilized as the outgroup to polari ztehe slender, dista slegment (brachi ucamrr}ying a iarge ,subter-
transformation of the identifi ehdomologous character states of the minal, hyalin emembrane on outer edge; both subterminal
Cftnaaorllra imnrgasooitftdioinaorg enm" s.aMe rtopiierrso oenwp o)acovsosne eter dani i,cbnn Qiycun ongOdt i'enarG gersd aim,ndulgy tlw ieesa tnba dtafe soDal ellc heboatiwrastec udtt{erhr1ces,a 9mte8it h7ow {ohnd)Iednn. it h coiafhs t"ocaritsaneo,ntsme-or-uscliaalnrnlagiep se a(tdnbdFuit t gue2n,rAarm)m ie3nd;-a slcae tgmammiudesdnlte ef dr( ;ispnrghoex a dwiifmatnahtdl )f sdieign msesetpnait nl{ulc esl{.ac woMsa)ergxp--us>
the state of character that received a code of "O" is not plesiomor- ments fused to form subchelum, with small, blun tknob in
phic. basal region. Sterna lfuFca absent.
The computer program HENNIG86 Versio n1.5 (Farr i19s8,8} Armature on rami of legs 1-4 as follow $(Roman
was used to analyze the phylogeneti crelationships between the numeral indicati nspgines and Arabi cnumeral, setae):
gener aof the Caligidae T.he commands `[mhennig'" and "bb'" were
Exopod Endopod
employed to produce multiple, shortest trees through performance
Leg l 1-O;111,1,3 (vestigial)
of extended branch swapping.
Leg 2 1-1 ;1-1 ;1,l,5 O-1; O-2i 6
Leg 3 1-O ;1-1 ;11L4 O-1; 6(absent)
DESCRIPTION Leg4 1-O:IV
Metacaiigu slatus sp, nov, L eg 1 ( F i g2 ,B ) protopo dwith long ,plumose outer seta and
(Fig s1-.4) an o t h e r s m a l l , p l u m o s e i n n e r s et a , in addition to a papilla
Materia lexamined: 261 g 9 , 105 88 , and 20 chaiimus b e ar i n g 2 l o n g se t u le s near outer margin of coxa; vestigial
larvae attached to adults of both sexes were recovered from en d op o d s m al l c on i ca l p r o t r u s i o nf;ir sstegment of exopod
oral and gi lclavities of cutlassfish, 77ichiur ulsepturus Lin- wi t h r o w o f s p in u le s o n posteri oerdge and $hort spiniform
naeus, caught in Strai otf Taiwan and landed at Dong-Shih se t a a t ou t er d i s t a cl o rn e r; middle 2 of 4 termina lelements
Fishin gPort on followi ndgates in 2001i 1 \ and 1 8 from on l a s t s eg m en t o f e x o po d w it h ou t accessory process but
1 fis hon 3t March; 59 ? ? , 19 r 8 and 3 larva terom 27 p i nn a t e b i l a te r a l l yl; a r ge p e c t e n a t base of each dista i3
fishe son 27 April a;nd 201 \ 9 , 85 (I" cl" , and 17 larvae sp i ne s ; 3 p l u m o s e se t ae o n posterior margin about half
trom 62 fishe son 22 June. One holotyp efemale ,1 ailotype l e ng t ho f t e r m i n a le l e me n t 4 , L e g 2 (Fi g2,C) coxa small,
male, and 60 paratype s(30 \\ and 30 8 '8) were with iarge ,plumose, jnne rseta on posteri oerdge and small
selected from the last collection and deposite din the papill taippe dwith a short seta on ventral surtace; basi swith
Nationa lMuseum of Natural History ,Smithsonia nlnstitution short ,naked outer seta; posterio mrargin ot basis with long,
in Washington ,D.C. narrow membrane and long marginal setule near midway of
Female: Body (Fi gI.A) 3.29 {2.42-3. m7m2 >long ,exclud- thi smembrane; outer spines on exopod {Fig 2.D) differen-
ing setae on caudal rami. Cephalothorac ischieid subcircu- tial lcoynstructed, with larges otne on proxima lsegment and
lar ,1.13 <O.82-1 .mm30 )jong and 1.01 (O.78-1. 32mm) smallest one on middie segment. Leg 3 (Fi g3,A) protopod
wide, excluding marginal hyalin emembranes. Fourth pedi- (apro nwi)th sma", plumose, outer seta and long ,plumose
ger 2.21 times wider than long .Genit aclomplex globular, inner seta in addition to an outer and a posterio mrarginal
1,13 (O,86-1 ,m4m0 )long and 1,11 (O.82-1, m4m2 >wide. membrane; one short and one long setule with each at tip
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Metacalig uansd Caligi dCladistics 1365
A
D
Fig, 1, Metacatigus latus sp, nov., female. A, habitus ,dorsal .B, antennule, dorsal ,C, antenna, postantenna prryocess and maxillule, ventral.
D, mandible. E, maxilla, Scale bars :O.8 mm in A; O.06 mm jn B; O.1 mm in C, E; O.98 mm i nD,
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Fig. 2. Metacaligus latu ssp, nov., female, A, maxiliiped, B, leg 1. C, leg 2, D, outer spines of le g2 exopod. E, le g4, F, termina lpart of leg 4
exopod. Scale bars: O.2 mm in A, E; O.1 mm in B. F; O.15 mm in C; O.04 mm in D.
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Metacaligu sand Caligi dCladistjcs 1367
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Fig, 3. Metacaigus latu ssp. nov, Female: A, leg 3. B, posterolat ecorranelr of genit acolmplex, ventral. C, caudal maus, dorsal .Male: D, habi-
tus dorsa lE., maxilliped. Scale bars: O.15 mm in A, E: 025 mm in B; O,05 mm in C; O.5 mm in D.
,
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1368 J.-S .Ho andC,-L Lin
of a papil ljaus itnner to baseline of posterior marginal mem- later amlargin of genit acolmplex, one tipped with a pinnate
brane ;2 patches of denticl oens ventral surface; velum well seta and another one, 2 pinnat seetae.
developed and fring ewidth marginal setules, Leg 4 (Fi g2E.) Male: Body (Fi g3D,) short and broad ,2.31 (1,72-2. 7m0m)
protopod with smatl, pinnat eseta at outerdistal corner; long ,excluding setae on caudal rami. Cehalothoraci schield
pecten at base of innermos touter spine on exopod (Fig. subcircular ,1,27 {O,96-1. m4m8 )long and 1.22 (1.00-t.56)
2F), Leg 5 (Fi g3,B) represented by 2 papilla eon postero- mm wide, excluding hyalin emarginal membranes. Fourth
C
l
x
Fig. 4, Metacaigus latus sp. nov., male, A, antennule, dorsal .B, antenna, postantennar yprocess and maxillule, ventral. C, dentifor mprocess
of maxillule, ventral. D, genita clomplex and abdomen, ventral. Scale bars: O,05 mm in A, C; O,15 mm in B; O,3 mm in D.
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MOtacafigu sand Caligi dCladistics 1369
pedige r4,44 times wider than long .Genita lcomplex shaped the followi nfgour characters: (1 )broader cephalothoracic
as a ja rO,.51 (O.36-O .mm58 }Iong and O,55 (O.44-O .m6m 0) shield and genita clomplex in both sexes, (2 )shorter caudal
wide. Abdomen 2-segmented ;proxima lsegment short, O.09 ramus in bot hsexes, (3 )papilla oen the postantennar ypro-
(O.06-O ,m1m2 )long and O.30 <O.22-O. m3m8) wide and cess tippe dwith 4 setules in both sexes, and (4 )the anal
anai segment about as long as wide, O,26 (O,20-O. ×34) somite in male distinc tlloynger than the proxima labdominal
O.24{O.18-02m8m). Caudalramus 1.29(O,09xO,mm0)7 somite.
times longe rthan wide and armed as in temale .Antennule Both M. uruguayensis and M. Iatus are found in the gill
(Fi g4.A) generall yas in female ,but proxim aslegment cavities of the cutlassfish taken from the Strai otf Taiwan.
armed with 29 setae, Antenna (Fi g4.B) with corrugated, However, in our past two years of examination of the cut-
adhesion pad on mediodistal surface of middle segment; lassfishes landed at Dong-Shih Fishin gMarket ,these two
terminal segment with lamella tecuticular tip and 2 basal congeners have never been found co-existing on a fis hhost,
setae with one on each side. Postantennary process (Fig, Another intrigu iponign tto mention ls about the occurrence
4B) stronger than in female, with prominent terminal hook. of the chalimus larva eof M. latu sO.n 27 Apri land 22 June
Dentifor mprocess of maxillule (Fi g4C.) bearin gsubterminal 2001, chalimus larva ein various stages of development
tine .MaxiHiped (Fi g3,E) with $mall medial seta in middle (fro cmhalimus 1 1to chalimus IV) were recovered attached
region of subchelum. Leg 5 as in female but leg 6 repre- randomly to the cephalothorax, genit acolmplex or abdomen
sented by a papill baearin g2 setae Iocate din dista rlegion ef the adult of both sexes. This is unusual in that the chali-
ot ventral ridge on genita clomplex (Fi g4.D). mus larva sltages are always found attaching directl yto the
Etymology: The species name latu sis a Lati nfor "broad" fis hhost .Of the 20 chalimus larvae ,8 were found attached
or "wide". I talludes to the possessio nof broad cephaiotho- to the female and 12, to the male,
racic shield and genit acolmplex in both sexes of thi sspe-
cies.Remarks: VALID CALIGID GENERA
Three species of MetacaLigu sare currently
known, they are: M. rufus (Wils o1n9,08) ,M. uiuguayensis ln their work on the parasit iccopepods of Brazilian
(Thomse n1,949) and M. hilsa e(She n1,957) .The last spe- fishes B,oxshal l& Monta "997 )provide da key to 29 genera
cies diffe rfsrom all other species of Metacaligu si,ncluding of the famil yCaligidae I. tis the lates ptublishe dwork show-
the presen tnew species, in the possessio nof a long ,4-seg- ing the recognized genera of the Caligidae H.owever, i tis
mented abdomen, which is longe rthan it sgenita lcomplex worthy to poin tout that while Doji r(i198 3c)onsidered Cafi-
(She n19,57) .The firs stpecies diffe rfsrom the presen tspe- gopsis and Diphyllogasterinsuffic ikennotwlny and relegated
cies in having the middle segment of the exopod ot leg 2 Heniochophilus to a synonym of Anuretes ,these three gen-
equipped with a larg eouter spine, which is as larg eas the era were recognized valid by Boxshall & MontU 0997) F,ur-
outer spine on the basal segment of the same ramus, and ther differen cbeetween those two works is the treatment ot
the outermost spine at the tip of leg 1 exopod longer than Mldias Wilson ,1911. This genus was not found in the key
the segment carrying i t(Wils o1n9,08; Ho and Bashirullah, give nby Boxshal l& MuntU (199 7b)ut considefed valid by
1977), Thus, M. utuguayensis bears the closest resem- Dojir i(1983),
blance to the present new species, Nevertheles sM,. latu sis According to Dojiri '($198 3r)edescription of Mldias
distinguisha bflreom M. uruguayensis in the possessio nof lobocte sb,ased on Wilson' s(191 1t)ype material kept at the
Table1. Thevalidcaligidgenera.
AbasiaWilson1,908 MappatesRangnekar,1958
AnchicatigusStebbing,1900 MetacaiigusThomsen, 1949
AnuretesHeller,1865 MidiasWilson,1911
ApogoniaCressey& Cressey,1990 RaralebionWilson1,911
Auama Dojir &i Cressey, 1991 Pa rapetalus Steenstru p& LUtken, 1861
BeliziaCressey1,990 RarechetusPillai1,961
CafigodesHeller1,865 PseuaCanuretYaemsaguti,1936
ClaipusMUIIer,1759 PseudechetusPrabha& Pillai,1979
CarituCsressey1,967 PseuclocaliguSscott,1901
Darteve"iaBrian,1939 Pseuclolepeophtheitus Markewitsch,1940
Echetu sKreyer ,1863 Pupulina van Beneden, 1892
Hermiliu sHeller ,1B68 Sciaenophilu svan Beneden ,1852
KlabataellParabha& Piila1i,988 SinocaligusShen,1957
LepeophtheirusNordmann, 1832 6ynestiSutseenstru&mLUtken,1861
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Smithsonia nlnstituti tohni, sparasit oef the grea tbarracuda, were coded in an ordered fashio nthat reflects a predeter-
Sphyrae nbaarTacuda (Walbaum) i,s not attributable to Cali- mined directi oont character evolution (se eAppendi xA),
gus as considered by Kabata (197 9a>nd followe dby Box- Inasmuch as an analysis with orderedcharacter coding has
shaN and Monta (1997 )Th.us, in followi ngDojiri ((s1983) been criticized as running the risk of predetermin itnhge
opinion, Midias is treate din this paper as a valid genus of topology of the resultant cladogram (O'Gra danyd Deets,
the Caligidae, 1987; Dojir iand Deets ,1988; Ho, 1991; Ho, 1994) ,both
ln their prelimina rryeview of the 19 species of Anuretes ordered and unordered analysis were pertormed to search
Ho and Lin (200 0c)oncurred with Dojiri '(s198 3t)reatment for the best resolved cladogram(s) of the Caligidae.
to relegate Heniochophitu tso a juni osyrnonym of Anuretes.
Our examination of the original works of Cafigopsi sby Analysis with orderedcoding
Markewit$ch (1940 )and Diphyllogast ebry Brian (1899) ln employjng the computer program HENNIG86 to per-
showed that both gener ahad not been full ycharacterized form the cladistic analysis, the algorithm "mhennig"' u$es
for making an evaluation of their taxonomic status. Thus, the available data ,such as the matrix in Appendix B, to con-
both Caligopsi asnd Diphytlogastera rcoensidered incettae struct initi calladogram(s) and saves all shortest trees (cla-
sedis and, therefor e2,8 genera of catigid copepods are rec- dograms} after running a Iimite dbranch-swapping .Five
ognized in this paper ,They are liste din Table 1. such trees were produced in this analysis. Then, with the
application of "bb'" command the computer uses those five
CLADISTIC ANALYSIS initi tarlees to perfor mextended branch-swappin gto pro-
duce multiple shortest trees. As a result of thi$ combined
lnasmuch as all caligids are known of their females and algorithms, a tota lof 668 equally parsimonio utrsees were
the male is unknown in seven genera, the homologous char- generate dwi,th a Iengt hof 96 $teps, a consistency index
acters utilized in the followi nagnalysis are confined to the (Cl )of 40 and a retention index (Rl o)f 55. Ctose examina-
adult females ,Furthermere i,n search for homologous char- tio nof these 668 trees revealed tha tthere are seven groups
acters in a genus comprising several species with different of cladograms distinguish ebdy the number of nodes that
states of character, the ones possessed by the type-species they carry.
are selected disregarding their status in the order of the As shown in Table 2, the number of nodes in those
character transformation .For instance, in the genus Caligus seven groups ef trees runs from 20 to 26, with 113 (226i668
where there are more than 200 species with many of them trees, 33.83 %) of the obtained cladograms showing 23
bearing 3-segmented exopod on leg 4, the state of 2-seg- nodes on the tree. The highest number of nodes on a tree
mented ramus is selected for this character even though 3- is 26 and only three of the 668 trees show the best resolu-
segmented state is more primltiv eTh,is is simply because tio nlike this .I ti scalled the best resoived because only one
the type-spe coif ethsi$ genus, C. curtus MllIle r1,785, has place (node }on these trees shows a polytomy where there
a 2-segmented exopod on it sleg 4. are three branches (clade Tsh)e.re are ten least re$olved
The characters setected for periormin gthe analysis of tree swith 20 nodes on the cladogram, which disptay sfive
phylogeneti crelationships between the 28 caligid genera places (node swi)th t7 clades (branch eocsc)urring in poly-
(Tabl 1e) are liste din Appendix A with the code for each tomy, three nodes with three clades and two nodes with four
state indicated in the parenthes ethsat follo wA,l ltogether 36 clades (se eTable 2).
apomorphic states were recognized and the coding of those In the output fil oef the analysis performed by employing
36 apemorphic states are given in Appendix B, l tis from this the algorithms "mhennigt" and "bb'" with the orderedcoding,
lis atn imput til ewas created to conduct the cladistic analy- the three best-resolv etdrees were designate das Trees 27,
sis. 50 and 280, They are reproduced in Fig ,5, with Tree 50
Ten of the 23 selected characters exhibit multiple state$ shown in it sentirety. As to Trees 27 and 280 only the parts
of character transformatio nT.he series of transformation differe nftrom Tree 50 are shown. The differenc beetween
Table 2, Number of trees in each of the seven groups of cladograms (identi fwiitehd the number of nodes tha tthey carry) obtained
in the analysis with orderedcoding.
Number of treeNumber of nodeNumber of branches on polytomou nsodes
1O trees 20 nodes21 17 branches (3 nodes each wi±h 3 branche s2, nodes each with 4 branches)
72 tree$191 nodes22 14 branche s(2 nodes each with 3 branches, 2 nodes each with 4 branches)
trees nodes23 11 branches <1 node with 3 branches, 2 nodes each with 4 branches)
226trees127 nodes24 1O branches {2 node each with 3 branches, 1 node with 4 branches)
trees nodes25 7 branches <1 node with 3 branches, 1 node with 4 branches>
39 trees nodes26 4 branche s(1 node with 4 branches)
3 trees nodes 3 branches (1 node with 3 branches)
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nttetacaligus and Caligi dCIadistics 1371
TREE 50
iil/:;,c.gg{,,g:,:ggheg?::"s
Pupulina
e54[
i
s3c:E,ligi!li
i
`O[3g[!:[i:Xi!
: '
'" '
Hermilius
p,),,
,iic,ic,lzlit Abasia
,,, ,lliiiiE 3i t1,f,[f Dartefveltligi:::
32 t1,f,[i Anuret:esgg:3:
;:;:g::::gtus
TREE 280
,,[
43[ Midias
46[ Parapetalus
47[ Sinocaligus
i:::gg:::r.g::theirus
U]49[
4,[
48[ Belizia
" fa'2g221:gus
42[
ChiCCaligus
41[3::/
Fi g . 5. enCtaligi dphylogeny produced through analysis of orderedcoding. Tree 50 shows entirety but forTree s27 and 280 only the part sdiffer-
from Tree 50 are shown. Note: the names of DarteveMa and kabataelL aon Node 31 and Mappates and Pseucianurete son Node 32 are
t22r13u((nOOc)) --at. e22 d1o3{n(1 1)T,) ra2et2 e(N 5Oo0)d ,-e T 4r28a2;n( s11fO) oa(rt1 m) Na-otd ie1o0 n5 (so32f;) c9aht(a tNr)a oc.dtee r9 4( 7sOt;) aa1ttOe Ns< o1ad}t e -e. a 5ct2hO; (7nO(o)d 1ea>t - No on7d (eTOr ) 4ea6et ; 5Nt0o2 d(ae1r5)e 1: -;7- ( 11O2() (O-O) ) - 7at( 1 1N{>1o, )d1 ea2t( 4ON5)o ;-d 2e 3 15(201<;) 1-1) Oa (t2 O3N) {o-3d.) e a1 t5O N4(o;1 d3) ae(tO N4) 4o-;d 1e33 (4<19O);),
-. 13{1) at Node 43; 6(O) - 6(1) ,16(O) -+ 16(1) at Node 42; 16(O) - 16{1) at Node 41; 14(O) - 14(1) at Node 40; 9(1) - 9(O) at Node 39;
2131<<1O)} -- . 2"3((D2) aatt NNooddee 3383;; 51(4O() 1-) - 5 (141<)O ,)15 a(tO )N o-d. e1 53(71;) ,9 2(1O() .1) -9.( 12)1 a (t 2N )aotd eN o3d6e; 13(21; ) 2-2{ 11) (-O) a2t2 N(o2d) eat 3N5o; d2e( O3>1 -; 4 {2O(}1 -) a t4 (N1o)d, e9 <314); .20 (9O<)2 }. at 2 N0o(d1e},
30 ;and 4(O} - 4(1) ,16(O) - 16(1), 17(O) . 17(1) ,19(O) . 19(1) at Node 29.
Tree 50 and Tree 27 is the displacemen tof Lepeophtheirus, Analysis with unorderedcoding
moving one step (nod ec)loser to the stem, and between Unlik ethe previo uasnalysis, application of the algo-
Tree 50 and Tree 280, the displacemen tof Sinocaligus,rithm "mhennig'" produced four initi aslho,rtest trees (cla-
jumpin fgrom Node 48 to Node 47. I tshould be emphasized dograms> ,and, with the execution of "bb'" i tgenerated
tha tthe topolog yof the 18 genera (Rseudolfrpeophthei3t,1u6s5 ,equally parsimoniou stree$ .The lengt hot these trees
Rseu(tocaligu sB,elizia ,Apogonia, Metacaligu s,Anchicati- is 87, with a consistency inde x(Cl o)f 44 and a retention
gus, SciaenQphitu sE,tuetus Caligocte sC,antus ,HerTniiius, index (Rl )of 55. Although the tree lengt hand Cl are
Abasia ,Dartevelt iaA,rtama, kabataeha ,Anuretes ,Atlap- improved over the previou sanalysis with ordered coding,
pates ,and Pseudanuretes ton CIade (Node 4)6 of Tree 50 the value of Rl remained unchanged and the overall resolu-
is exactly the same between these three best-resolved tion of the obtained trees is not as high.
trees, Nine groups ot cladograms are recognized with the
number of nodes on a givin gtree varying from 16 to 25. In
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TREE 496
F53[Pu p u lLienpeaophtheirus
[[[[[
I,,,
,
TREE 3061
g"s
r3sE3ftf,Ai.i･iiiii･i
L40L,,E figkg7:geS
[ 3
7Gi
Fig. 6, Caligi dphylogeny produce dthrough analysis of unordered coding. Tree 496 shows entirety but for Tree 3061 only the parts different
from Tree 496 are shown,
other words, the best resolved tree (wit 2h5 nodes) i$ not as any ot these three trees are next best to the completely
good as those produced in the previou sanalysis and the resolved tree where every node has djchotomy, Since Tree
leas tresolved tree (wi t1h6 nodes) is worse than those 50 occupies the central role with only one differen cferom
found in the previo uasnalysis, While there are 11 best each of the two other 26-node trees, i tis adopted as a work-
resolved trees (Tre e4s39, 454, 493, 496, 498, 3044, 3046, ing hypothesi sfor the phylogeny of the Caligidae.
3054, 3056, 3059 and 3061) there is but one Ieast-resolved Examinatio nof the transformati oofn each character on
tree (Tre e1180) .Those ld best-resolv etdrees can be Tree 50 {se eAppendix A) shows that Characte riO (out-
dMded int otwo groups with one carrying a polytom ycom- growth on the abdomen) changed at Node 49 from the ple-
prisin fgour branches (Tree 4s93, 496, 3056 and 3059) and siomorphy {"O "s,mooth and cylindrjcal) to an apomorphy
the other, two polytomie swith each carrying 3 branches ("1 "be,aring iatera laliform structures}, and at Node 46 it
(Tre e4s39, 454, 498, 3044, 3046, 3054 and 3061) ,One reverted to the origina lplesiomorphy .I tis this character
tree from each of these two groups was reproduced in Fjg. reversal (homopla soyf )the abdominal outgrowth that
6. Tree 496 is shown with it sentirety and Tree 3061, only serves as a synapomorphy to unite those 18 gener aon
the part sthat are differe nTth.e leas tresolved tree, Tree Clade 46 to a monophyietic clade,
1180, has five polytomie sw,ith two nodes containing three The absence of sternal furca (apomorp hoty Character
branches and another three nodes, fiv ebranches, 13) is a synapomorphy at Node 43 (Tab l3e) and that places
the afore-mentioned 18 genera in a monophyly excluding
Caligi dphylogeny Pseudocaligu sand Pseudolepeophtheirus. I tis interesting
l tis apparent from the above two sections that analysis to pojn tout tha tthe sternal furca is presen tin the species
of caligjd phylogeny with ordered coding generat ebdetter- of both Anuretes and Hennilius ,but these two genera are
resolved trees, producin gthree cladograms with each carry- shown on the thre etrees in Fig ,5 as the descendents of an
ing 26 nodes, Having only one node with three branches, ancestor lacking the sternaHurca.
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