Table Of ContentBull. not. Hist. Mus. Lond. (Zool.)61(1): 1-10 Issued29June 1995
A
revised familial classification for certain
THE NATURAL
cirrhitoid genera (Teleostei, Percoidei HISTORY MUSEUM
Cirrhitoidea), with comments on the groifp'ss jul 1995
monophyly and taxonomic ranking PRESENTED
GENERAL LIBRARY
P.H. GREENWOOD1" g/2J l^lf.01
,
Honorary Research Associate, J.L.B. Smith InstituteofIchthyology, Private Bag1015, Grahamstown
6140, SouthAfrica and Visiting Research Fellow, The NaturalHistory Museum, Cromwell Road,
London SW75BD
CONTENTS
Introduction 1
Methodsand materials 2
Urohyal morphologyand thesupragenericclassificationofthecirrhitoidfishes, particularlythe genus
Acantholatrisandspeciescurrentlyreferred tothe genusNemadactylus 2
The familialclassificationofAcantholatris Gill, 1862, and Nemadactylus Richardson, 1839 5
Possible lineageswithin the Latridae asnowexpandedtoinclude thegeneraAcantholatrisand
Nemadactylus 5
Taxonomicand phylogeneticconclusions 6
Taxonomy 6
Thecirrhitoidsasamonophyleticlineage 6
Intragrouprelationshipswithin thecirrhitoids 8
Acknowledgements 9
References 10
Synopsis. Previous suprageneric classifications of cirrhitoid fishes were based mainly on superficial characters.
Recentanatomicalstudiesshowthatfivemorphologicallydistincttypesofurohyalboneoccurwithinthegroup,and
that on this basis certain taxa have been misplaced at both the generic and familial levels. The monophyly ofthe
cirrhitoidfishes, tacitlyassumed bypreviousauthors,isreviewed, andanhypothesisoftheirmonophylyerectedon
the basis of several synapomorphic features. It is also proposed, on the basis of those synapomorphies, that the
lineagebegivensubordinal rankwithin the Perciformes.
INTRODUCTION features. Apart from some comments by Regan on cranial
features, vertebral numbers and vertebral morphology, nei-
ther paper included any other anatomical information on
Duringaninvestigationintothecranial andbranchial muscles these fishes.
associated with feedingin certain cirrhitoid fishesfrom South Although Regan (1911) expressed some doubts about the
African waters, it became apparentthatfive distinct morpho- familialstatusgivenbyhimtothefivegenericassemblageshe
types of urohyal occur within the group. Two very distinct recognised (suggesting that subfamilial rank could be more
types are found in genera currently classified as members of appropriate) he did not comment on Gill's (1862) earlier
the Cheilodactylidae; of these two forms, one also occurs in classficationwhichrecognisedfoursubfamilieswithinasingle
genera referred to the family Latridae by Regan. Despite the family, the Cirrhitoidae; indeed, and inexplicably, Regan
passage of over 80 years since Regan's (1911) paper was {op. cit.) makes no reference at all to Gill'spaper.
published, it remains the most recent suprageneric classifica- Gill's {op.cit) foursubfamiliescorrespond, almostentirely,
tion of these fishes, which he arranged in five families, viz to Regan's families, except that Gill's Latridinae included
Cirrhitidae, Cheilodactylidae,Aplodactylidae, Chironemidae two genera, Nemadactylus Richardson, 1839, and Cheilodac-
and Latridae, grouping them informally as the 'cirrhitiform tylusLacepede, 1803, which Reganincorporatedinhisfamily
percoids'. Cheilodactylidae; Gill's list ofincluded taxa in his concept of
Like its predecessor, namely Gill's more detailed paper of the Latridinae also contained two genera not mentioned by
1862, Regan's later analysis was based mainly on external Regan, despite their being recognised at that time and still
beingrecognisedtoday {viz Latridopsis Gill, 1862, andAcan-
tDrGreenwooddied3March 1995. tholatris Gill, 1862).
©TheNaturalHistoryMuseum, 1995
PH. GREENWOOD
Thepresentcontribution, itishoped, willplaysomepartin Chirodactylus brachydactylus: 3 specimens, DIFS
rewakening an interest in a phylogenetically based taxonomy unregistered (D).
of Regan's 'cirrhitiform percoids', and of their interrelation- 2specimens, DIFS unregistered (S).
shipswithin, orperhapsoutside, the Percoidei. Regan (1911) 2 'paperfish' larvae, 33 & 38 mm standard length.
gave no formal suprafamilial ranking to his 'cirrhitiforms', RUSI, unregistered (D).
but in some recent classifications (Nelson, 1994) the group C. grandis: 1 'paperfish' larva, 57 mm standard
hasbeen raisedto asuperfamily, the Cirrhitoidea. The use of length. RUSI 18404 (D).
BMNH
the terms 'cirrhitoid' and 'cirrhitoids' in this paper is thus to Dactylophora nigricans*: 1869.2.24:8 (S).
beinterpretedasareferencetothesuperfamilyandnottothe Chironemidae: Chironemus marmoratus*: BMNH
subordersuggested forthese fishes on p. 9. 1871.3.29: 28 (S).
Chironemusmarmoratus: BMNH 1861.11.7: 6 (D).
Cirrhitidae: Amblycirrhitus pinos: 2 alizarin preparations
BMNH
METHODS AND MATERIALS ex 1976.7.14: 453-7 (S).
BMNH
Amblycirrhituspinos: 1984.7.16: 96 (D).
BMNH
Cirrhitusmaculatus*: unregistered (S).
Species of the percoid families Serranidae and Centropomi- Cirrhitichthys oxycephalus: RUSI 11658; BMNH
dae served as outgroup representatives of basal members of 1929.9.20: 8 (D).
BMNH
the Percoidei. The condition and composition of the sterno- Cirrhitichthysoxycephalus: 1908.3.23: 77-79
hyoideusmuscles, andthemorphologyoftheurohyalinthese (X).
taxa were taken to represent the plesiomorphic percoid Cyprinocirrhitespolyactis: RUSI 12339 (D).
conditions. Gymnocirrhites arcatus (type species of the
Wherever possible, dry skeletal preparations and alizarin genus):BMNH 1965.12.20: 10(D).
BMNH
transparencies were examined, as were radiographs and Gymnocirrhites arcatus*: unregistered, col-
dissections ofpreservedspecimens. lected by Richardson (S).
BMNH
Studymaterial. Thesymbolsusedinthefollowinglistare: Gymnocirrhitesarcatus: 1865.3.2:82-83(X).
Oxycirrhites typus (type species of the genus):
D. Sternohyoid muscle dissected and the urohyal removedor BMNH
1929.6.12: 2(D).
examined in situ; X. Radiographed; S. Dry skeletal material BMNH
examined; an asterisk following a binomen indicates that this Paracirrhitesforsteri: BMNH 1852.9.13: 119 (S)
name appears on the box housing the skeleton, and that its Paracirrhitesforsteri: 1969.7.16: 28-32; one
specimen(D).
specific identity could not be checked on the basis of the BMNH
Latridae: Acantholatris bergi 1936.18.26: 439 (D)
characters preserved.
&(X).
The familial grouping used in the list is that resulting from BMNH
Acantholatris gayi: 1879.5.14: 278;
theconclusionsreachedinthispaper, andthusdiffersinsome 1890.2.26: 49 (D) & (X).
respects from the classifications of Gill (1862) and Regan
Acantholatris monodactylus (type species of the
(1911), and in some cases that of later authors (see p. 5 BMNH
relating to the species 'bergf and 'gayi' previously placed in genus): 1960.1.8: 6-13B; MonNeH(D), all (X).
Acantholatris monodactylus: unregistered,
thegenus Cheilodactylus andin the family Cheilodactylidae).
Institutional abbreviations are: BMNH, The Natural His- AecxaGntohuoglahtrIisls.;m2osnpoedcaicmteynlsu,sb:otRhU(SS)I. 33484; 33485;
tory Museum, London; DIFS, The Department of Ichthyol-
33624 (D).
ogy and Fisheries Science, Rhodes University, South Africa;
RUSI, J.L.B. Smith Institute of Ichthyology, South Africa; BLaMtrNidHopsis ciliaris (type species of the genus):
SAM, South African Museum, CapeTown. Latridops1i8s72.ci7l.i1a:ris3:2B(SM).NH 1873.12.13: 55 (D) &
Studymaterial. (X).
(i) Outgroups. Latris lineata: BMNH 1855.9.19: 124 (D).
SAM
Centropomidae; Glaucosomatidae; Ambassidae: The osteo- Latrislineata: 22623 (D).
logical and othermaterial listed in Greenwood (1976). Mendosoma lineatum (type species of the genus):
BMNH
Serranidae: as above, together with Epinephalus alexandri- 1960.1.8: 14-21; one (D), all (X).
nus BMNH 1867.2.1: 69-72; (D). Mendosoma lineatum: RUSI33613; 33625 (D).
BMNH
Nemadactylus macropterus: 1855.9.19: 314
(ii) Cirrhitoid fishes. (X).
BMNH
Aplodactylidae: Aplodactyluspunctatus * (type species of Nemadactylus macropterus: 1872.7.1: 21
thegenus): BMNH 1873.4.3: 157 (S). (D) & (X).
BMNH
A. lophodon: 1914.8.20: 214 (D).
Cheilodactylidae: Cheilodactylusfasciatus (type species of
thegenus): 3 specimens, DIFS unregistered (D).
C.fuscus: 1 specimen, DIFS unregistered (D). Urohyal morphology and the suprageneric
C. pixi: 6 specimens, DIFS unregistered (D).
2specimens, DIFS unregistered (S). classification ofcirrhitoid fishes, particularly the
4 'paperfish' larvae, 44-50 mm standard length, genusAcantholatris and species currently referred
RUSI 19842 (D). to the genusNemadactylus
1 'paperfish' larva, 44 mm standard length, RUSI
19842 (S). Within the cirrhitoid fishes, five morphologically distinct
C. zonatus: BMNH 1907.12.23: 238 (S). types of urohyal can be recognised (Figs 1 & 2). Since each
REVISED CLASSIFICATION FOR CERTAIN CIRRHITOID GENERA
Fig. 1 The urohyalof:A. Paracirrhitesforsteri; Cirrhitidae; left lateral view (BMNH 1852.9.13: 119). B. Chironemusmarmoratus;
Chironemidae; left lateral andventralviews. (BMNH 1871.3.29:28). C. Aplodactyluspunctatus; Aplodactylidae; left lateral and ventral
views. (BMNH 1873.4.3: 157). D. Dactylophoranigricans; Cheilodactylidae; left lateralview. (BMNH 1869.2.24:8). Relative toother
figures, thisbone hasbeen rotatedthrough90°tothe left; arrowindicatesdorsal prominence. Scale inmillimetres. DrawnbyGordon
Howes.
type can be correlated with one of the five family groups the bone'suppermargin issharporsomewhatflattened. Like
recognised by Regan (1911), these will be referred to as the the cirrhitid type, the latrid urohyal isalso ofapleisomorphic
cirrhitid, latrid, chironemid, aplodactylid and cheilodactylid form, one occurring in such basal percoids as the Serranidae
typesrespectively. and Centropomidae (Kusaka, 1974; Greenwood, 1976: 39,
The cirrhitid type (Fig. 1A) appears to be a plesiomorphic fig. 21, and other personal observations).
form, onefundamentallysimilartothatfoundinseveralbasal Departure from the basal percoid form of urohyal is most
percoids (see Kusaka, 1974; also personal observations). pronounced in the chironemid, aplodactylid and cheilodac-
The latrid urohyal (Fig. 2) differs markedly from the tylid types of bone. In chironemids (Fig. IB) the bone is
cirrhitid type, and also shows slightly more intrafamilial shallow, the ventral margin greatly flattened and expanded
variation, particularly with regard to its posterior margin's bilaterally to form a broad shelf, while the dorsal margin is
degree ofindentation, the extent to which the ventral margin alsonoticeablyflattenedandbilaterallyproducedintoashelf,
is produced bilaterally into a narrow or broader shelf, the albeit one relatively narrower than that on the ventral aspect
extent to which the bone is produced ventrally, and whether of the bone; when compared with the urohyal in cirrhitids,
GREENWOOD
P.H.
Fig. 2 Theurohyaloffourlatridspecies.A. Acantholatrismonodactylus (BMNHunregistered;exGoughIsland);leftlateralandventral
views. B. Latrislineata (BMNH 1855.9.19:194);leftlateralview. C. Mendosomalineatum (BMNH 1960.1.8:14-21);leftlateralandventral
views. D. Latridopsisciliaris (BMNH 1872.7.1:31); left lateral andventralviews. Scaleinmillimetres. DrawnbyGordon Howes.
the chironemid type is relativelyforeshortened (cfFigs 1A & from a latrid type by a marked anterior extension and
IB). deepening of the latter's posterior indentation, coupled with
A pronounced ventral shelf and overall foreshortening of an increase in the angle subtended by the two arms so
the bone is also characteristic of the aplodactylid urohyal formed, the two morphotypes are readily distinguishable.
(Fig. 1C), butinthistype theboneis relativelydeeperthanis Interestingly, the urohyal in the so-called 'paperfish' juvenile
mm
the chironemid urohyal, and the dorsal surface is produced stage (see p. 7) of a 44 standard length Cheilodactylus
into only a narrow shelf. pixi Smith, 1980, resembles that of the latrid type more
When compared with all other types, the cheilodactylid closelythan does this bone in larger specimens; nevertheless,
urohyal (Fig. ID) is very distinctive. In lateral view it has the upper and lower arms of the urohyal in the 'paperfish'
virtually the shape of an arrow-head with its tip directed stage are more widely separated, the division between them
anteriorly, and with the two arms meeting at the somewhat extends further anteriorly, and the anterior body ofthe bone
thickened apex from which a dorsally directedprocess arises. islesscompressedandmorebarrel-likethanthatinanyofthe
The anterior edges of both arms are slightly broadened to adult latrid urohyals I have examined.
form a very narrow bilateral shelfthat does not quite extend In his monograph on urohyal bones Kusaka (1974)
to the posterior tip of either arm. Although, morphogeneti- described and illustrates this bone in Goniistius zonatus
cally, the cheilodactylid type of urohyal could be derived (Cuv., 1839), a taxon now, and previously (Gill, 1862),
,
REVISED CLASSIFICATION FORCERTAIN CIRRHITOID GENERA
referred to the family Cheilodactylidae (see Allen & Heem- single subfamily, as did Gill (1862).
stra, 1976, forcomments on the status ofthisgenus; alsop. 6 Thus, in my view, based essentially on their urohyal
below). Kusaka, however, lists the species underthe heading morphology and not negated by other characters (see, how-
Aplodactylidae. The bone depicted is certainly ofthe cheilo- ever, the pectoral fin character discussed below), Acanthola-
dactylid type and not ofthe aplodactylid type, and I presume tris and Nemadactylus should be included in the family
Kusaka'splacingthespeciesinthe Aplodactylidaeisalapsus. Latridae, currently comprising species of the genera Latris,
This author (op. cit: ) also figured and described the urohyal Latridopsis and Mendosoma, the latter recently shown to be
from a specimen supposedly of Cirrhitichthys aureus Temm. monotypic by Gon & Heemstra (1987). In addition to the
& Schl., 1843 (Cirrhitidae). Unfortunately I have not been urohyal characters, the genera listed above lack a suborbital
able to examine a specimen of this species, but the bone shelf, which in cheilodactylids is a prominent feature formed
illustrated (and described as 'shaped like a standing rat') is from the posterior upper margin of the lachrymal bone and
unlike that in any cirrhitoid taxon I have examined, particu- the entire upper margins ofthe second and third suborbitals.
larly amongst membersofthe Cirrhitidae andeven in a taxon Also, unlike cheilodactylids, these genera have the basal
such as Oxychirrhites typus Bleeker, 1857, whose elongate scaly sheath to the soft dorsal fin somewhat higher and thus
and tubular snout is an unusual morphotype within both the more prominent than that at the base of the spinous part of
Cirrhitidae and the cirrhitoids as a whole. If Kusaka's figure the fin.
and description are accurate and the specimen was correctly As in cheilodactylids, the latrids (here taken to include
identified, then asixth and highly distinctive form ofurohyal, Acantholatris and Nemadactylus) have 35 vertebrae (14
one far removed from that of other cirrhitids must be abdominal + 21 caudal elements including the urostylar
recognised, andthe highertaxonomicpositionofitspossessor element; data from radiographs and dry skeletons listed on
or posessord be reconsidered (assuming, that is, the bone p. 2). To judge from the dry skeletal and dissected material
Kusaka examined was not teratological or damaged during available to me, parapophyses are present on all precaudal
preparation). centrain both families, and no ribs are sessile.
A typical cheilodactylid urohyal (Fig. ID) occurs in all
members of the family (sensu Regan [1911] and subsequent Possible lineages within the Latridae as now
authors) I have examined apart from Nemadactylus and
members of the genus Acantholatris, viz the type species A. expanded to include the generaAcantholatris and
monodactylus (Carmichael, 1818), and the species A. gayi Nemadactylus
(Kner, 1869) andA. bergi (Norman, 1937).
Parentheticallyit should be noted thatA. gayiandA. bergi AcantholatrisandNemadactylusdiffernoticeablyfrom Latris,
were both placed in the genus Cheilodactylus, and the family Latridopsis and Mendosoma in having one of the lower,
Cheilodactylidae, by Norman (1937). The formerspecieswas unbranched pectoral rays (i.e. the fifth, sixth or seventh ray
latertransferred by Fowler (1945) to the genusAcantholatris from the bottom of the series) greatly elongated, its tip,
with noexplanation given forthe change, but was retained in which extends beyond the fin's margin, reaching to at least
the family Cheilodactylidae. Neither author appears to have the level ofthe anus and sometimes as far as the midpoint of
been aware, however, that Gill (1862) had includedAcantho- the anal fin.
latris in his subfamily Latridinae. Mann (1954: 266) followed There are also differences in the followingfeatures:
Fowler'sgenericandfamilial placingofA. gayi, and listedthe (i) In scale size, as shown by lateral-line scale counts. In
speciesbergiunderAcantholatris in the index to that publica- Latris, Latridopsis and Mendosoma these range from 112 to
tion. The reader is there referred to page 266 ofthe text. No 120 in the two former taxa, and from 68-78 in Mendosoma
mentionismadeofA. bergion thatpage, buton page 85 (op. (data from Last et. ai, 1983; Gon & Heemstra, 1987;
cit.) Acantholatris bergi Norman (the author's name not pers.obs.). In Nemadactylus macropterus the count is 59 or
eAthtnilcsalnotpsiaecrdot'i.ciunlMabarrnacncokem(tbosip)naciittsi.)olniisstoetdfhusanmatomhneegssfitrfsottrhaeutthhTeonrvsaptesocoireeesms.pldAoesyl 6ar0an,ndgaeinndisAcifanrnootmhtoh5le0art-rs6ip0esc(imNeoosnro4md7aa-nc6,t8y1l(9up3es7r;,sAp.oebrssb..e;rogLbisa.sa)tn.det.A.aig,ay1i98t3h)e
noted earlier (p. 1) Regan did not include Acantholatris in (ii) Anal fin length. In Latris, Latridopsis and Mendosoma,
any ofhiscirrhitiform families. the number of branched anal rays ranges from 17-35 (the
lowest counts occurring in Mendosoma, viz. 17-21, whereas
The familial classification ofAcantholatris Gill, in Acantholatris species and Nemadactylus macropterus the
range is from 12-15, and other species of the genus, 16-19
1982 and Nemadactylus
(sources as above).
The urohyal in all three Acantholatris species examined, and Pending a detailed generic revision of the various taxa
inNemadactylusmacropterus, isvirtuallyidentical and differs involved, especially the several Australian and New Zealand
markedlyfrom thatinthe cheilodactylids, cirrhitids, aplodac- speciescurrentlyreferredtothe genus Nemadactylusitwould
tylids and chironemids (see pp. 2-5 and cf Fig. 1 with Fig. be premature to formally recognise the two groups as, for
2A). Instead, it resembles the latrid type, both in detail and instance, tribes or subfamilies of the Latridae, although
in its gross morphology (c/Figs. 2A, B and C), especially in phylogeneticallysomesplitseemstohave occurredwithin the
its fan-like outline. This marked difference would suggest lineage.
thatthelatridgenera(aslistedin Regan, 1911), togetherwith The condition ofthe pectoral fin in the Latris-Latridopsis-
Acantholatris and Nemadactylus shared a recent common Mendosoma group of latrids provides something of a puzzle
ancestry distinct from that of the cheilodactylids. It also since these taxa are the only cirrhitiforms not showing any
suggests that the phyletic relationships ofthe two groups are marked elongation of the uppermost unbranched ray in the
obscured by uniting the cheilodactylids with the latrids in a lower section of the pectoral fin, nor, as in most other
PH. GREENWOOD
cirrhitoids, do any ofthese raysclearly extend beyondthe fin ThegenusAcantholatrisGill (1862, typespecies Chaetodon
membrane, and none is markedly thickened. In having the monodactylus Carmichael, 1818), was overlooked by Regan
lowermost 5-9 rays unbranched, these fishes are, however, (1911) in his synoptic review of cirrhitoid families, but is
typically cirrhitoid. This latter condition can be considered currently placed in the family Cheilodactylidae (see p. 5).
one ofthe synapomorphies unitingcirrhitoid fishes. However, on the basis of its urohyal morphology, and its
A typically derived pectoral fin configuration occurs in lacking a suborbital shelf (see p. 5) the genus should be
the Cirrhitidae, yet the family would appear to be the least classified in the Latridae. Regan (1911) differentiated the
derived of all cirrhitoid taxa (see p. 8). In contrast, except Latridae from the Cheilodactylidae on the basis ofthe latrids
for the condition of the pectoral fin, members of the having feeble, unbranched pectoral rays that are not pro-
Latris-Latridopsis-Mendosoma assemblage within the duced beyond the fin's margin, and by their lacking a
Latridae share with Acantholatris and Nemadactylus, and suborbital shelf; in other feaures he noted that the two
with the cheilodactylids, aplodactylids and chironemids, families are similar. With the inclusion ofAcantholatris and
the derived condition for all the osteological and myologi- NemadactylusintheLatridaethenatureofthepectoralfin no
cal features discussed on page 5. That being so, it is longer serves as a differentiating feature (see p. 5), the
unlikely that the pectoral fin form in Latris, Latridopsis principal diagnosticcharactersforthe family now lying in the
and Mendosoma can be interpreted as a true retention of form of the urohyal bone, the absence of a suborbital shelf,
the plesiomorph condition. If that was the case, then the and in the more prominent arrangement ofthe basal sheath-
derived condition must have evolved more than once ingscales ofthe soft dorsal fin (see p. 5).
within the cirrhitoids. A more parsimonious solution to the Gill's (1862) suprageneric classification included Nemadac-
problem therefore, would, be to interpret the pectoral fin tylus as a division-Nematodactyli-ofhis subfamily Latridi-
form in Latris, Latridopsis and Mendosoma as a secondary nae, in which subfamily but as another division to which he
reversal to a seemingly more plesiomorphic condition than gave the name Latrides he also included Latris, Latridopsis,
is found in any other cirrhitiforms, including the family Mendosoma, Acantholatris, Chirodactylus, Cheilodactylus
with the greatest number of plesiomorphic features, the and Goniistius. Regan (1911) on the other hand, but without
Cirrhitidae (see p. 8). reference to Gill's paper, treated the latter author's four
The geographical distribution of the two groups within subfamilies asfamilies, and recognised afifth, the Cheilodac-
the Latridae has an interesting pattern. Of the taxa in the tylidae, fortwogenera, viz CheilodactylusandNemadactylus;
long-finned assemblage, Nemadactylus (see p. 5) occurs no mention is made in Regan's paper of the other taxa in
only in Australia, Tasmania and New Zealand, thus over- Gill's Latridinae except for Latris and Mendosoma, which
lapping the entire range of Latridopsis a member of the Regan retained in hisfamily Latridae.
short-finned group and one restricted to that region; it The evidence presented here (pp. 2-5), especially that
overlaps in part (New Zealand and Tasmania) that of the basedon urohyal morphology, wouldsupport Regan's (1911)
widely distributed Mendosoma lineatum, also a member of classification with regard to the separation of Cheilodactylus
the short-finned group, and in part, that of Latris, another (and, although not mentioned by Regan, Chirodactylus and
member of the short-finned group (Australia; New Goniistius) from the other taxa included in Gill's Latridinae,
Zealand; Gough and Tristan da Cunha islands; Vema and would support the inclusion of all three taxa in one
Seamount; St Paul and Amsterdam islands). The other family, the Cheilodactylidae. The same evidence would also
long-finned taxon, Acantholatris does not occur in Aus- support Gill's inclusion of Latris, Latridopsis, Acantholatris,
,
tralasia, but has a wide western distribution, including St Mendosoma and Nemadactylus in a single suprageneric
Paul, Amsterdam, and Gough Islands, Tristan da Cunha, taxon. Since Regan's (1911) familial ranking has been
Vema Seamount Chile, Juan Fernandez and the western accepted and used since that time, and until contraindicative
coast of South America from Rio de Janeiro southwards. evidence is available to suggest otherwise, that ranking (i.e.
This distribution thus widely overlaps that of the short- Latridae) is retained.
finned, monotypicgenus Mendosoma lineatum, viz St Paul, The anatomical and other features used in this paper
Amsterdam and Gough islands, the coast of Chile and, as (pp. 2-5) would support the recognition of Gill's (1862) and
noted above, New Zealand and Tasmania (the latter being Regan's (1911) other suprageneric lineages, again, for the
areas where Acantholatris does not occur); data from reasons given above, as families and not subfamilies, viz. the
Norman, 1937; Fowler, 1945; Mann, 1954; Smith 1984; Cirrhitidae, Aplodactylidae (Gill'sHaplodactylinae) andChi-
Last et. al., 1983; Gon & Heemstra, 1987; Paulin et. al. ronemidae.
1989; Andrew & Hecht, 1992; Andrew, pers.comm., 1993). At an intrafamilial taxonomic level, Allen & Heemstra
(1976) note that 'The currently accepted classification of the
Cheilodactylidae ... is most unsatisfactory' a sentiment I
would not only endorse, but would extend to other cirrhiti-
TAXONOMIC AND PHYLOGENETIC form families as well. In part this situation has resulted from
CONCLUSIONS the use of mainly superficial characters, with little or no
attention paid to anatomical features, especially myological
and osteological ones. Thus on those grounds I cannot agree
Taxonomy with Allen & Heemstra's (op.cit.) treatingAcantholatris as a
subjective synonym of Cheilodactylus and its consequent
The material studied indicates, on osteological and myologi- placement in the Cheilodactylidae (see above, p. 5). How-
cal grounds (p. 5), that the species currently named Nema- ever, at least on the characters and specimens I have exam-
dactylus macropterus (the type species of Gill's (1862) genus ined, I would endorse their synonymy of Whitley's (1957)
Dactylopagrus; see Wheeler, 1986) should be classified in the genus Morwong (type species Cheilodactylus fuscus Castel-
Latridae and not the Cheilodactylidae as it is at present. nau, 1879) with Cheilodactylus.
REVISED CLASSIFICATION FOR CERTAIN CIRRHITOID GENERA
The cirrhitoids as a monophyletic lineage of all but two of the cirrhitoid families I have dissected (see
p. 2). The exceptions are a chironemid, Chironemus marmora-
On the basis ofseveral apparently synapomorphic character- tus Giinther, 1860 (160 mm standard length) and an aplodac-
istics (see below) the cirrhitoids would seem to be a mono- tylid, Aplodactylus lophodon Giinther, 1859 (180 mm S.L.).
phyletic lineage, aconclusion implied by both Gill (1862) and Since, however, the sac is a juvenile {i.e. sub-adult) feature in
Regan (1911) who described the group as a 'natural' one but the other taxa, and the exceptional specimens were, to judge
gave no reasons for that conclusion. The derived characters fromtheirgonadialdevelopment, youngadults, Isuspectthatit
on which I would base an hypothesised monophyly of the would also be present in smaller specimens ofthese species. In
cirrhitoids are, taken in conjunction, a reduced number (15) Cheilodactyluspixi, for example, the sac is well-developed in a
mm
ofprincipalcaudal fin rays, the unbranched lowermostfive to specimenof44 standardlength, buthasdisappearedinone
nine rays in the pectoral fin (usually with their tips produced of 46 mm. Likewise, in Chirodactylus brachydactylus (Cuv.,
beyond the fin membrane), the lower part ofeach cleithrum 1830),itispresentinafishof38mmstandardlength,butabsent
greatly expanded anteroposteriorly and meeting its antimere inoneof42mm.Thepresenceofalipoidsacinspecimensfrom
in a deep, carinate symphysis, an increased number of 128to 243 mm standard length ofotherspecies (referred in the
vertebrae relative to other percoids (26-35, comprising 10-16 current literature to the Latridae and Cheilodactylidae) whose
abdominal and 15-21 caudal elements), and the presence, maximum adult lengths are from 50 cm to one metre, suggests
ventrally, in subadults of a peculiar, lipoid-filled sac (Fig. 3), that the size at which the lipoid sac disappears is positively
free from the overlyinghypaxial muscles, andextendingfrom correlated with that at which members of a species become
the urohyal, to which it is attached, to the anus, with the adult.Thissuppositionisborneoutbythepresenceofthesacin
mm
lipoidal material apparently contained in hexagonal compart- a juvenile Chirodactylus grandis (Giinther, 1860) of 57
ments. standard length, a species whose adults reach a length of one
To the best of my knowledge, this lipoid sac has not previ- metre, whereas it has disappeared, at a length of 42 mm, in
ously been noted as a feature ofsubadult cirrhitoid fishes, nor young Chirodactylus brachydactylus, whose adults reach a
indeed ofany other perciforms except the stichaeid Lumpenus lengthof40cm. Again, itispresentinaspecimenofAcanthola-
mm
maculatus (see Falk-Petersen etal., 1984). I first observed it in tris monodactylus 243 S.L.; adults of this species attain a
smallspecimens(the so-called 'paperfish' stage) ofCheilodacty- standard length of at least 65 cm. Thus, the sac's apparent
lus pixi ca 43 to 44 mm standard length (Fig. 3), where its absence in chironemids and aplodactylids could be artefactual,
presenceresultsinthe 'pouter-pigeon'-like ventralprofileofthe and related to the size-range of the specimens I was able to
paperfish stage in this and other cirrhitoid species (see photo- examine.
graphsin Whitley, 1957; Allen& Heemstra, 1976, andNielsen, It is hoped to carry out a more detailed examination ofthe
1963). Subsequent dissections revealed a lipoid sacin members lipoid sac when specimens suitably fixed for detailed histo-
Fig.3 Chirodactyluspixi,49mmstandardlength (RUSI 19842) in rightlateralview;partiallydissected, andwiththegreaterpartofthe
pectoralandtheentire pelvicfinremoved. The large anteriorportionofthe lipoidsac(LS)isclearlyvisible; partofitsposteriorportionis
alsovisible (x). Throughoutitslength, thewallofthesac,unlikethe musclesabove it, isheavilypepperedwith melanophores.
PH. GREENWOOD
logical and histochemical study are available. and right cleithra, its origin thus being immediately before
that ofthe anterior infracarinalis muscle insertion.
Intragroup relationships within the cirrhitoids, In both the Cirrhitidae and in the other families, the
and the ranking ofthe group sternohyoideus myotomes, except the horizontal first myo-
tome in the latter group, are chevron-shaped with the apex
Withinthecirrhitoids,theCirrhitidaeshouldbe rankedasthe directed anteriorly. However, in the latrids, cheilodactylids,
most plesiomorphic taxon, a view seemingly implicit in chironemids and aplodactylids, the angle between the upper
Regan's (1911) diagram of relationships. My reasons for and lower arms of the chevron is more acute, and the lower
giving the family this ranking are based on urohyal form, the arm is relatively longer than in cirrhitids.
low vertebral count (26-28) relative to that in other families, Judgingfrom the ratherscant literature on the sternohyoid
the absence of parapophyses on the first three abdominal muscle in teleosts (see Winterbottom, 1974), and from a
vertebrae, sessile ribs associated with these vertebrae, the personalknowledgeofthesituationinpercoids, thecondition
presence of 3 predorsal bones (2 in the other cirrhitoids of the muscle in the Cirrhitidae should be ranked as plesio-
radiographed or dissected) and the presence of a suborbital morphic, that in the other cirrhitoid families as derived and
shelf(which, however, is also developed in the Cheilodactyl- possibly a unifying synapomorphy for the Aplodactylidae,
idae [but see below]). Furthermore, the Cirrhitidae are the Chironemidae, Latridae and Cheilodactylidae within the lin-
only cirrhitoids with a basal percoid type of myotome eage.
arrangement in the sternohyoideus muscle, a feature not In Regan's (1911) figure of cirrhitoid intrarelationships
previously noted. That is, one in which the three pairs of referred to above, the Cirrhitidae occupy a basal (i.e. stem
hypaxialmyomeresformingthesternohyoideusmuscleare all position) and are linked, on the one hand to the Chironemi-
arranged in a vertical series, with the first block covering the dae and Aplodactylidae, and on the other to the Cheilodac-
anterior part of the urohyal (Fig. 4A). Members of all other tylidae and Latridae (the generic composition ofthe families
cirrhitoid families, in contrast, have the first (i.e. anterior) being those given by Regan). No reasons were provided by
hypaxial myotome of each side displaced ventrally so that it Regan for these supposed relationships, which presumably
now lies medially and horizontally (not, as in cirrhitids, were based mainly on superficial characters as well as a few
laterally and vertically) to form, with its antimere, a ventral anatomicalones. With the anatomical information now avail-
muscle, paired in most species but in some with the left and able a different scheme of intragroup relationships at the
right parts fused over most of their lengths to form a single family level can be proposed (see Fig. 5, and Table 1).
median muscle (Fig. 4B). Single orpaired, thisventro-medial In this scheme, apomorphic features (see Table 1) are
component of the sternohyoid runs from the urohyal tip to taken to be: (i) The derived form of urohyal, ofwhich there
the prominent ventral projection at the symphysis ofthe left are three distinct types (see p. 3). (ii) The presence of a
Fig.4 A. Cirrhitidtype organizationofsternohyoid
myotomes. Drawnfrom Paracirrhitesforsteri;
semi-schematic. In thisarrangement, the
sternohyoidmusclescompletelyenvelopthe urohyal
bone. B. Basicorganizationofthesternohyoid
myotomesinallotherfamilies. Drawnfrom
ce Nemadactylusmonopterus;semi-schematic. Intaxa
ofthesefamilies,varyinganteriorextentsofthe
urohyalare notcoveredbythesternohyoidmuscle.
1-4: 1sttofourthmyotomesofthesternohyoid
muscle;c: cleithrum;ce: cutedgesofthesterno-
hyoidmyotomes;uh: urohyal. DrawnbyAnthea
Ribbink
ce
REVISED CLASSIFICATION FORCERTAIN CIRRHITOID GENERA
.v6*
•
\0
~N?
o*
4 ,15 .15*1 3*.15*|
|
7* # 3*
1 1 ,1
1 or 2,6,8,10,12,14
Fig. 5 Tentativeschemeofphyletic relationshipswithin the Cirrhitoidei. Asterisksindicate apomorphiccharacters(see alsoTable 1). Since
both thecirrhilid (1) and latrid (2) conditionofthe urohyal are rated aspleisomorphic, that bone in thecommon ancestorofall lineagesis
taken tobe 1 or2.
derived myotomal arrangement of the sternohyoideus synapomorphic features discussed (above pp. 6-7) it seems
muscles (p. 8). (iii) More than 28 vertebrae, (iv) Parapophy- reasonable to hypothesize that the five families comprising
ses developed on the first three abdominal vertebrae, with the lineage, given informal ranking as the 'cirrhitiform per-
the first pleural rib associated with the second or third coids' by Regan (1911), and suprafamilial status by recent
vertebrae, (v) Predorsal bones reduced to 2. The three authors (Nelson, 1994), should be elevated to subordinal
differenttypesofderivedurohyalmorphology (pp. 2-5) (iein status (Cirrhitoidei) within the Perciformes.
chironemids, aplodactylids and cheilodactylids) are each
taken to be independently evolved apomorphies. Relation-
ships (Fig. ) suggested by these data are: (i) That the Acknowledgements. I amparticularlyindebtedtoDrColinBuxton
cirrhitids are the plesiomorph sister group to the other four of Rhodes University's Department of Ichthyology and Fisheries
families combined, (ii) The chironemids are the sister taxon Science who first aroused my interest in the anatomy of cirrhitoid
of the aplodactylids, latrids and cheilodactylids combined, fishes, and who provided many specimens for dissection. Professor
andthatforthe moment this assemblage should be treated as Tom Hecht of that department is to be thanked for personally
an unresolved trichotomy since no two lineages uniquely obtainingaspecimenofCheilodactylusfuscusfromSalamanderBay,
share a recognisable synapomorphic feature. For example, NewSouth Wales, inwhichoperation he wasaided by BillTalbot of
tahlethouurgohhytahlatinbotnheeinlatthriedschiesiloofdacatyblaisdaslapnedrcaopildodatcytpyel,idasnids gtrhaeTtoBitruaDdcrekifsoPhrhitWlhaetHmeeraenmFsyitsdhriasCcuuolsftsuirtoehnesRweJse.eLha.arBvc.ehShSmtaaidttiahobnoIutnthsetcriietru.rtheitogiodess,amnyd
highly derived, each is unique to the families respectively. forhis adviceon the relevant literature, and toTony BoothofDIFS
The cheilodactylids it will be noted, retain the plesiomorphic myappreciation forhispainstakingpreparationofseveralskeletons.
suborbital shelf, whereas it is lost in the latrids and aplodac- Once again it is a great pleasure to thank my old colleague Gordon
tylids (and in the chironemid lineage as well). The value of Howes for his elegant and accurate draftmanship, and also to thank
this feature as an indicator of relationship, however, is another formercolleague, OliverCrimmen ofthe Fish Section, The
problematical because it involves a loss (and not an acquisi- Natural History Museum, London for his untiring helpwith matters
tion) in the lineages concerned. Clearly, a greater number radiographicandbibliographical. Yet again, myspecial thanksgo to
and variety ofcharacters must be sampled and their polarity Huibre Tomlinson, for her patience and skills in producing the
determined before this hypothesis of cirrhitoid intrarelation- typescripts, and to Robin Stobbs for producing certain of the
shipscan be improved and the trichotomy resolved. radiographs, andthephotograph, usedinthispaper.
The same reservation would apply before any sister-group
hypothesis can be erected regarding the relationships of the
cirrhitoids within the Percomorpha. However, based on the
10 PH. GREENWOOD
Table 1 Datamatrixandcharacters.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Cirrhtidae
Amblycirrhituspinos + + + + + +
Cirrhitusmaculatus + + + + + +
Cirrhitichthysoxycephalic + + + + + +
Cyprinocirrhitespolyactus + + + + + +
Gymnocirrhitesarcatus + + + + + +
Oxycirrhitestypus + + + + + +
Paracirrhitesforsteri + + + + + +
Chironemidae
Chironemusmarmoratus
Latridae
Acantholatrisbergi + + + + + +
A. gayi + + + + + +
A. monodactylus + + + + + +
Latridopsisciliaris + + + + + +
Latrislineata + + + + + +
Mendosomalineatum + + + + + +
Nemadactylusmacropterus + + + + 6- +
Aplodactylidae
Aplodactyluspunctatus + + + + + +
A. lophodon + + + + + +
Cheilodactylidae
Cheilodactylusfasciatus + + + + + +
C.fuscus + + + + + +
C. pixi + + + + + +
C. zonatus + + + + + +
Chirodactylusbrachydactylus + + + + + +
C. grandis + + + + + +
Dactylophoranigricans + + + + + +
Characters: 1. Cirrhitid-type urohyal (i.e. ofabasalpercoidtype, butone differingfrom thelatridcondition);2. Latrid-type urohyal {i.e. of
basalpercoidtype; see 1); *3. Aplodactylid-typeurohyal; *4. Chironemid-type urohyal; *5. Cheilodactylid-type urohyal;6. Lessthan28
vertebrae; *7. More than 28vertebrae;8. Parapophysesnotdevelopedonthefirstthree (sometimes4) abdominalvertebrae,butsessile
pleuralribsonone ortwoofthesecentra; *9. Parapophysesdevelopedonthe firstthree abdominalvertebrae. Nosessilepleuralribs; 1st
pleural ribarticulatingwith the parapophysesofthethirdabdominalvertebrae; 10. Three predorsalbones; *11. Twopredorsalbones; 12.
Basalpercoid-typeofsternohyoid muscle; *13. Derivedconditionofthe sternohyoidmuscle; 14. Suborbitalshelfpresent; *15. Suborbital
shelfabsent. Asteriskindicatesan apomorphicfeature.
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