OCCASIONAL PAPERS

OF THE

California Academy of Sciences

No. 121^ 23 pages, 2 figures, 1 table

THE PROCURRENT SPUR:
AN UNDESCRIBED PERCIFORM CAUDAL CHARACTER
AND ITS PHYLOGENETIC IMPLICATIONS

By

G. David Johnson

SAN FRANCISCO
PUBLISHED BY THE ACADEMY
September 11, 1975

COMMITTEE ON PUBLICATION
George E. Lindsay, Chairman
Diana R. Young, Editor

OCCASIONAL PAPERS
OF THE

California Academy of Sciences

No. 121, 23 pages, 2 figures, 1 table

THE PROCURRENT SPUR:
AN UNDESCRIBED PERCIFORM CAUDAL CHARACTER
AND ITS PHYLOGENETIC IMPLICATIONS

By

6. David Johnson

Scripps Institution of Oceanography
La Jolla, California 92037

Introduction

The morphology of the supporting elements of the caudal
skeleton is considered by most fish systematists to provide
important taxonomic and phylogenetic information (Gosline,
1961; Nybelin, 1963; Patterson, 1968). The principal and
procurrent rays associated with these elements however, are
often not illustrated or described in studies of caudal
osteology, and where they are, details of their morphology
are usually not considered. In consequence, a phylo-
genetically significant character within the Perciformes is
to date undescribed. This character, involving a unique
configuration of the two posteriormost ventral procurrent

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

rays (fig. 2) , was first observed during the course of an
investigation into the interrelationships of several percoid
families. It was originally taken as a condition unique to
the Haemulidae, but further exploration has shown it to be
present with surprising consistency in other percoid
families. The purpose of this paper is to describe the
character, present a synoptic survey of its distribution
within the Perciformes, and discuss some of its phylogenetic
implications.

Material and Methods

Both cleared and stained and alcoholic specimens were
examined. In the latter, the morphology of the caudal ray
bases could be clearly determined by examination with
transmitted light under a dissecting microscope after minor
dissection and folding back of the skin and flesh in this
region.

The majority of the specimens examined are in the
collections of the Scripps Institution of Oceanography (SIO)
and the California Academy of Sciences (CAS) . A few speci-
mens are part of a cleared and stained collection housed at
the Southwest Fisheries Center, National Marine Fisheries
Service (NMFS) . In order to shorten table I, catalogue
numbers are not included. However, a complete listing of
all specimens examined and their catalogue numbers is on
file with the California Academy of Sciences.

Description

The usual number of principal rays in the perciform
caudal is 17, consisting of 15 branched and 2 unbranched
rays (Regan, 1913). These 17 rays support the major surface
of the caudal fin. Anterior to these principal rays, dor-
sally and ventrally, are the procurrent rays, a series of
much shorter rays not typically included in the posterior
margin of the fin.

Figure 1 shows the usual arrangement of principal and
procurrent caudal rays. There is no modification of the
proximal ends of the procurrent rays , and the ventral series
is very nearly a mirror image of the dorsal one. This is
the situation in the majority of teleosts. However, in a
group of about 30 percoid families and the Polynemidae,
Sphyraenidae , and Stromateoidei , there is a unique speciali-
zation of the ventral series. The posteriormost procurrent
ray bears a spur that projects ventrally to overlap the
preceding ray (fig. 2). Ordinarily, in fishes with a spur
the preceding ray does not extend forward proximally to meet
the bases of the remaining procurrent rays, but is basally
shortened, often originating just anterior to the overlapping
spur. The presence or absence of the spur and the shortening
of the preceding ray are indicated for the species examined

No. 121] JOHNSON: PROCURRENT SPUR

(arranged by family) in table I. Not shown in table I are
those species observed in a cursory search for the spur or
shortening in various non-acanthopterygian orders; this
search failed to reveal any trace of either condition.

Shortening of the preceding ray does not always accompany
the development of the spur, but this is by far the more
frequent condition. In some genera, particularly within the
Sciaenidae and the Stromateoidei , the spur is well developed,
but the preceding ray extends forward normally. A condition
which may or may not be homologous with that shown in figure
2 is found in the Beryciformes and in the genus Symphysanodon;
the preceding ray is shortened with no trace of spur develop-
ment. This condition is also evident in two genera of Poly-
nemidae, but the presence of the spur in other members of
that family possibly indicates a secondary reduction. A few
species, as in the genera Siniperaa and Scombrolabrax , have
no typically developed spur but do have a small projection
interpretable as a remnant of the spur. This condition is
indicated by "r" in table I. Although usually the preceding
ray base is obviously either shortened or not, the situation
has proved to be difficult to interpret in a few species. A
very slight basal shortening is indicated by ^ in the table.
In the text it is assumed that the spur is accompanied by
shortening of the preceding ray except where otherwise
indicated.

The caudal musculature was compared in genera with and
genera without the spur. All muscle bundles described and
illustrated by Nursall (1963) occur in each group and ex-
hibit similar origins and insertions. The ventral bundle of
the main body of the deep ventral flexor inserts on the last
ventral procurrent ray, which is the one that bears the spur.
The spur, when present, forms the point of insertion. In
its absence, insertion is directly on the ray axis._ No
difference was noted in the insertion of the remaining ten-
dons of the ventral bundle.

Discussion

The presence of such a character as the procurrent spur
in a large number of fairly diverse families leads to two
possible conclusions. Either it has been evolved indepen-
dently in several separate lineages and fixed by strong
selection pressures resulting from its functional importance,
or it has been evolved only once and thus indicates relation-
ship by common ancestry of all those groups possessing it.

The association of the procurrent spur with the caudal
musculature suggests a possible function. Nursall (1963)
pointed out that, in Hoplopagrus , the chief action of the
deep ventral flexor "is flexion with abduction also being
applied owing to the form of the heads of the fin rays and
sites of attachment. " If any functional significance for
the procurrent spur is to be inferred, it must surely in-
volve an increased ability to abduct the last procurrent ray

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

(and through their connective association, all ventral rays
of the caudal) . A shift in insertion of the uppermost ten-
don of the ventral bundle from a point on the lateral sur-
face of the ray to a ventrolateral projection (procurrent
spur) could alter the forces acting on the ray when the
muscle contracts in such a way as to increase the abduction
vector. Increase in size of the spur would increase
abduction leverage. The associated shortening of the base
of the preceding ray might prevent its interference with
full abduction, allowing the base of the next preceding ray
to project up into the resulting space as its posterior part
is pushed downward. In essence, this may mean that those
fishes possessing a procurrent spur have a greater ability
to spread the ventral rays of the caudal fin or at least a
more efficient way of spreading them to the same degree.

The establishment of this rather speculative possible
function for the procurrent spur does not necessarily lead
to the acceptance of the alternative multiple-origin
hypothesis. It is not the existence of the function itself
but its adaptive value that is crucial to the concept of
convergence. Any functional advantage in this case would
surely be associated with the operation of the caudal fin
and locomotion. Yet a quick survey of table I shows that
there is no obvious correlation between the presence or
absence of the spur and caudal fin structure, swimming type,
or life style. The spur is present in such diverse swimming
and feeding types as Stereolepis and Emmeliohthys and yet
absent in E-pinephelus and Ftevooaesio , two forms representing
a similar ecological range. It is absent in the fast
swimming Carangidae and yet present in Pomatomus , also a
powerful swimmer. Its presence or absence within families
is very consistent, despite variation therein in swimming
and feeding types, and where a few members of a family have
lost the spur, no unifying pattern is apparent. The pro-
current spur appears to be associated in no obvious way with
adaptive radiation.

Ventral spurs are developed on one or more of the
principal caudal ray bases in certain groups with a reduced
number of rays, most notably in the Scorpaeni formes ,
Blennioidei , and Gobioidei. These spurs would probably
facilitate abduction of the rays as is hypothesized for the
procurrent spur. However, the procurrent spur has not been
found in any members of these groups. Its absence, in spite
of the apparent importance of abduction in these fishes as
evidenced by spurs on the principal rays, strengthens the
concept of the procurrent spur as a specialization of unique
origin.

Several facts point to the primitiveness of the pro-
current spur within the percoids. The Beryciformes exhibit
a basal shortening of the next to last procurrent ray which
might be interpreted as a foreshadowing of the percoid
condition. The spur and the associated shortening charac-
terize the Polynemidae and Sphyraenidae , two groups often
placed at the pre-perciform or basal perciform level.

No. 121] JOHNSON: PROCURRENT SPUR

Finally, the presence of the spur in the lower percoid
families Percichthyidae , Kuhliidae, Centropomidae , Scorpidi-
dae , Kyphosidae, and Monodactylidae directly demonstrates
its early occurrence within the percoids.

These facts, and the fact that the presence or absence
of the procurrent spur agrees for the most part with well
known and accepted taxonomic groupings , particularly
families, support the interpretation that the presence of
the spur is a primitive perciform character, uniquely
derived and thus indicative of phylogenetic relationship,
i.e. all groups possessing it were derived from a common
ancestor. The relationship of those groups lacking the spur
is less clear. It is evident that the spur and associated
shortening can be lost with no trace (essentially a reversion
to the generalized teleost condition) , as this has apparently
occurred independently within the Sciaenidae and the
Stromateoidei. It cannot be argued therefore, that other
families without a spur represent a single lineage within
the percoids. However, it does seem likely that many if not
all of these families were derived from ancestors possessing
the spur.

The value of the procurrent spur as a taxonomic and
phylogenetic indicator can perhaps best be illustrated by
considering its occurrence in relation to some current
concepts of familial relationships and generic placements
within the Perciformes:

Axillary process. The presence or absence of the scaly
process in the axil of the pelvic fin was used by Regan
(1913) and Gosline (1966) to suggest an arrangement of
percoid families, the general phylogenetic validity of which
is still uncertain. A comparison of the distribution of the
procurrent spur with that of the axillary process shows only
that the spur occurs more frequently in those families with
the axillary process than in those without it (of 35
families, excluding beryciforms, listed by Gosline as having
an axillary process, 15 have the spur; of 36 listed as not
having the axillary process, only 8 have the spur). This
higher frequency of coincidence of the two characters is
probably a reflection of the fact that they are both
primitive at the percoid level.

Ramus lateralis aaoessorius . Freihofer (1963) described
various patterns of the ramus lateralis accessorius nerve
(RLA) in the Perciformes and discussed their possible
phylogenetic importance. There is a striking correlation
between the occurrence of Freihofer 's pattern 10 and the
presence of the procurrent spur. The following are pattern-
10 families: Arripidae , Theraponidae , Pomatomidae,
Kuhliidae, Scorpididae, Kyphosidae, Girellidae, Nematistiidae ,
Centrolophidae, Nomeidae, Stromateidae (Freihofer, 1963),
and Oplegnathidae (Freihofer, pers. comm. ) . Of these, only
the Girellidae and Nematistiidae lack the spur. In the

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Girellidae, the probably primitive genus Graus^ retains a
remnant of the spur, indicating that the absence of the spur
in the other members of this family is secondary.
Nematistius was described by Freihofer as having a "reduced
pattern 10" which he considered homologous to the typical
pattern 10. The absence of the spur then could be explained
as a secondary loss, perhaps resulting from the modified
caudal fin structure (deeply cleft fin ray bases, overlapping
and closely applied to the hypurals) . The possibility also
exists that the "reduced pattern 10" is not homologous to
the typical pattern 10 and that the usual placement of
Nematistius in or near the Carangidae is correct. Inte-
restingly, of the eleven pattern-10 families listed in table
I of Haedrich (1967), only two, Girellidae and Nematistiidae ,
lack a bony bridge over the anterior vertical canal of the
inner ear (AVC bridge) , and these are the same two that lack
the procurrent spur.

Perciohthyidae . Gosline (1966) removed several genera
from the Serranidae and placed them in the somewhat hetero-
geneous family Perciohthyidae. The presence of the pro-
current spur in all these genera (except Niphon and
Maocullochella , of questionable pertinence to the group on
other grounds) supports their exclusion from the Serranidae,
in which the spur is consistently absent.

Symphysanodon. On the basis of an osteological study,
Katayama (1968) agreed with previous authors in placing
Symphysanodon in the Lutjanidae. Anderson (1970), while
presenting no new anatomical evidence, concurred with
Katayama. As part of an ongoing study on the limits and
relationships of the lutjanids and some associated groups,
I have recently examined the osteology and cheek myology of
Symphysanodon and am unable to find any sound evidence for
relating it to the Lutjanidae. It appears to be more
primitive than the lutjanids in several respects and
resembles most closely the Perciohthyidae (sensu Gosline)
except that it has an axillary process. The larva of
Symphysanodon is distinctive and resembles no known percoid
larva (E. H. Ahlstrom, pers. comm. ) . The presence of a
marked shortening of the next to last ventral procurrent ray
supports the removal of this genus from the Lutjanidae. The
absence of the spur can be interpreted as a secondary loss
or as a condition homologous to that seen in the beryciforms,
suggesting that Symphysanodon may occupy a very primitive
position within the percoids.

The monotypic Graus nigra {=Pinguilabrum punctatum) was
originally described as a labrid by R. A. Philippi in 1887.
However examination by the author and R. A. Fritzsche of
several specimens of this fish from Chile indicates that it
is a primitive girellid (to be reported on further) .

No. 121] JOHNSON: PROCURRENT SPUR

Verilus. Most previous authors have treated this genus
as a lutjanid closely allied to Etelis . However, on the
basis of both external and internal morphology, I find no
evidence of a lutjanid affinity, but instead a striking
resemblance to the genera Aovopoma, Doderleinia , and
Malakiohthys , which Gosline placed in the Percichthyidae.
The presence of the procurrent spur in Verilus and its total
absence in all lutjanid genera support the new placement of
this genus.

Aipogonidae . The procurrent spur is absent in all
apogonid genera examined. Eraser (1972) was uncertain as to
the placement of the genus Brinkmanella , and the presence of
the spur in this genus indicates that it is probably not
closely related, unless antecedently, to the Apogonidae.
The absence of the spur in Howella and Bathysphyraenops
provides no support for the placement of these genera in the
Percichthyidae as was done by Eraser.

Soombrolabrax. Gosline (1968) stated that "except in a
few characters, the genus Soombrolabrax could serve morpho-
logically as an ancestral form for the trichiuroids and, in
most respects, for the Scombridae as well." The presence of
a remnant of the spur and shortening of the preceding ray
suggest that if this genus does share any close relationship
with the Scombroidei , it must indeed be a primitive one. If
we accept Soombrolabrax as a basal scombroid, the presence
of the spur supports Gosline 's objection to the postulation
of Starks (1911) that scombroid origins probably lie in the
area of the carangids, and his suggestion, instead, of a
pomatomid origin.

The above discussion and specific examples demonstrate
the value of the procurrent spur in perciform systematics.
It appears to be a phylogenetically significant character
and one which should be considered in any systematic
investigation of the perciform fishes, particularly the
Percoidei. Because there are clear indications that the
spur has been lost independently several times, it is not
possible to draw any broad inferences concerning familial
relationships solely from the distribution of the spur.
Only a more comprehensive knowledge and understanding of all
aspects of perciform morphology will eventually allow us to
elucidate the complex evolutionary relationships of this
diverse group of fishes.

Acknowledgments

I would like to thank William Eschmeyer (CAS) and E. H.
Ahlstrom (NMFS) for allowing me to examine specimens in
their care. Thanks are also due Carl L. Hubbs for providing
editorial assistance with the manuscript; E. H. Ahlstrom,
W. A. Gosline, and Donn E. Rosen for reading and commenting

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

on the manuscript; Warren C. Freihofer for his assistance
during my work at the California Academy of Sciences and for
providing specific information on RLA patterns. In parti-
cular I thank Richard H, Rosenblatt for his guidance and
editorial assistance during the course of this investigation,

Literature Cited

ANDERSON, WILLIAM D., JR.

1970. Revision of the genus Symphysanodon (Pisces:
Lutjanidae) with descriptions of four new
species. Fishery Bulletin, vol. 68, no. 2,
pp. 325-346, 5 figs., 3 tables.
FRASER, THOMAS H.

19 72. Comparative osteology of the shallow water

cardinal fishes (Perciformes : Apogonidae) with
reference to the systematics and evolution of
the family. Ichthyological Bulletin of the
Rhodes University, no. 34, pp. 1-105, 44 pis.,
6 tables.

WARREN C.

Patterns of the ramus lateralis accessorius and
their systematic significance in teleostean
fishes. Stanford Ichthyological Bulletin,
vol. 8, no. 2, pp. 79-189, 29 figs., 2 tables.
W. A.

The perciform caudal skeleton. Copeia, no. 3,
pp. 265-270, 3 figs.

The limits of the fish family Serranidae, with
notes on other lower percoids. Proceedings of
the California Academy of Sciences, ser. 4,
vol. 33, no. 6, pp. 91-112, 10 figs., 1 table.

The suborders of perciform fishes. Proceedings
of the United States National Museum, vol. 124,
no. 3647, 78 pp., 12 figs., 3 tables.
RICHARD L.

The stromateoid fishes: systematics and a
classification. Bulletin of the Museum of
Comparative Zoology, vol. 135, no. 2, 56 figs.,
2 tables.
MASAO

Notes on the osteology and systematic position of
Symphysanodon typus Bleeker. Bulletin of the
Faculty of Education, Yamaguchi University,
vol. 17, pt. 2, pp. 105-111, 3 figs.
R.

The caudal musculature of Hoplopagvus guntheri
Gill (Perciformes: Lutjanidae). Canadian
Journal of Zoology, vol. 41, pp. 865-880,
6 figs.

FREIHOFER,
1963.

GOSLINE,
1961

1966,

1968,

HAEDRICH,
1967.

KATAYAMA ,
1968.

NURSALL, J,
1963.

No. 121]

JOHNSON: PROCURRENT SPUR

NYBELIN, 0
1963.

PATTERSON,
1968.

PHILIPPI
1887

REGAN, C,
1913.

STARKS, E
1911.

Zur Morphologie und Terminologie des Schwan-
skelettes der Actinopterygier. Arkiv for
Zoologie, ser. 2, vol. 15, pp. 485-516, 22 figs.

C.

The caudal skeleton in Mesozoic acanthopterygian
fishes. Bulletin of the British Museum
(Natural History), Geology, vol. 17, pp. 47-102,
28 figs.
R. A.

Sobre los tiburones y
Chile. Universidad
pp. 1-42, 8 pis.
T.

The classification of

and Magazine of Natural History,
12, pp. 111-145.

C.

Osteology of certain scombroid fishes. Leland
Stanford Junior University Publications,
University Series, no. 5, 49 pp., 2 pis.,
1 fig.

algunos otros peces de

de Chile, Anales , vol. 71,

the percoid fishes,
ser. 8,

Annals
vol.

10 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

TABLE 1. The list below follows the olassification of
Greenwood et al. (1966) with certain modifications. The
proouvrent spur is present (-fJ^ absent (-)j or a remnant (r) .
The preceding ray base is shortened (+), not shortened (-)i
or slightly shortened (J-).

Procurrent Preceding
Spur Ray Base
Shortened

BERYCIFORMES

Polymixoidei

Polymixiidae

Polymixia lowei - -

Berycoidei

Trachichthyidae

Gephyroberyx japonicus , - +

Hoplostethus japonicus ,
Paratrachichthys fernandezianus

Anomalopidae

Anomalops katoptron , - +

Photoblepharon palpebratus

Berycidae

Beryx affinis - +

Holocentridae

Myripristis leiognathus , - ■*■

Adioryx suborbitalis

Anoplogasteroidei

Anoplogasteridae

Anoplogaster cornuta - +

STEPHANOBERYCIFORMES

Stephanoberycoidei
Me 1 amp h ae i dae

Poromitra crassiceps ,
Poromitra oapito ,
Melamphaes spinifer,
Melamphaes macrocephalus

PERCIFORMES

Mugiloidei

Mugilidae

Mugil oephalus , Agonostomus
montioola

No. 121] JOHNSON: PROCURRENT SPUR 11

Sphyraenoidei

Sphyraenidae

Sphyraena barracuda,
Sphyraena luoasana

Polynemoidei

Polynemidae

Polydactylus approximans ,
Galeoides polydactylus

Pentanemus quinquarius ,
Eleutheronema tetradactylus

Percoidei

Percichthyidae

Verilus sordidus , Aoropoma
japonioum , Doderleinia
berycoides , Malakiohthys
wakiyae , Synagrops bella,
Stereolepis gigas , Morone
saxatilis , Lateolabrax
japonicus , Dicentrarchus
punctatus , Perciohthys
trucha, Macquaria australasica ,
Percilia gillissi

Siniperca chuatsi

Niphon spinosus , Maccullochslla
macquariensis

Incertae sedis

Symphy sanodon berryi

Kuhliidae

Kuhlia marginata , Kuhlia arge

Labracoglossidae

Labracoglossa argentiventris

Scorpididae

Scorpis aequipinnis ,
Medialuna calif avniensis ,
Mi or acanthus strigatus

Kyphosidae

Kyphosus elegans , Sectator
ocyurus , Hermosilla azurea

Girellidae

Graus nigra

Girella nigricans , Melambaphes
zebra, Doydixodon fremenvillii

12 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Monodactylidae

Monodaotylus argenteus +

Centropomidae

Centropomus nigresoens , Lates +

niloticus , Psammoperaa

waigiensis

Ambassidae

Ambassis interrupta , Ambassis
gymnoGephala , Priops lungi

Glaucosomidae

Glauoosoma burgeri +

Serranidae

Epinephelus labriformis , -

Paralabrax maculatofasoiatus ,
Centropristes philadelphicus ,
EZlevkeldia macoulloohi ,
Cratinus agassizii , Hypopleotrus
lamprurus , Variola louti ,
Hapalogenys mucvonatus , Serranus
phoebe , Diplectrum paoifioum ,
Sohultzea beta, Liopvopoma sp . ,
Pikea aurora, Anthias anthias ,
Callanthias platei , Pseudanthias
thompsoni

Grammistidae

Rypticus bicolor -

Pseudochromidae

Pseudochromis fusous -

Plesiopidae

Plesiops nigrioans ~

Acanthoclinidae

BeZonepterygion fascioZatus -

Theraponidae

Therapon theraps , Pelates +

quadriZineatus , Batnia pZumbea

Nannopercidae

Nannoperca australis -

I

1

\

No. 121] JOHNSON: PROCURRENT SPUR 13

Centrarchidae

Miaropterus salmoides , Lepomis
maohrochirus , Enneacanthus
obesus , Elassoma zonatum ,
Chaenobvyttus gutasus , Fomoxis
annularis , Centrarahus
macroptevus I Aoanthavchus
pomotis , Arohoplites interruptus

Percidae

Perca flavesoens , Percina
oaprodes f Gymnocephalus cernua ,
Gymnooephalus sohraetzer ,
Etheostoma nigrum

Apogonidae

Apogon retrosella , Fowleria
aurita, Archamia bleekeri ,
Cheitodipterus macrodon ,
Sphaerarnia orbicularis ,
Rhabdamia gracilis , Pseudamia
gelatinosa , Gymnapogon
philippinus , Howella sp . ,
Bathysphyraenops simplex

Incertae sedis

Brinkmanella elongata

Pri acanthi dae

Priacanthus cruentatus ^
Cookeolus boops

Malacanthidae

Caulolatilus prinoeps ,
Malaoanthus parvipinnus

Sillaginidae

Sillago sp.

Mullidae

Pseudupeneus grandisquamis ,
Mullus auratus , Parupeneus
barberinus , Upeneus tragula ,
Mulloidichthys martinicus

Rachycentridae

Rachycentron canadum

Echeneidae

Remora remora

14 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Carangidae

Hemiaaranx zetotes ,
Traohinotus faloatus ,
Vomer deotivifrons ,
Nauorates duotor, Traahurus
symmetrious f Etegatis
bipinnulatus , Gnathanodon
speoiosus , Deoapterus punctatus ,
Chloroseombrus ovqueta , Seriola
doTsalis

Nematistiidae

Nematistius pectoralis

Coryphaenidae

Coryphaena equiselis

Apolectidae

Parastromateus niger

Menidae

Mene maoulata

Bramidae

Brama japoniea

Caristiidae

Cavistius maderensis

Pomatomidae

Pomatomus saltatrix ,
Scombrops boops

Lactariidae

Laotarius laotarius

Leptobramidae

Leptobvama mullevi

Arripidae

Avripis georgianus

Emmelichthyidae

Emmeliahthys oyanescens ,
Evythvocles sohZegeti ,
Plagogenion sp .

Banjosidae

Banjos banjos

No. 121] JOHNSON: PROCURRENT SPUR 15

Haemulidae

Pomadasys panamensis , +

Orthopristis reddingi ,
Haemulon sexfasoiatum ,
Lythrulon flaviguttatum ,
Ovthostoechus maculicauda ,
Anisotvemus davidsoni , Conodon
serrifev , Microlepidotus
inornatus , Xenichthys xanti ,
Xenistius oaliforniensis ,
Xenocys jessiae , Isaoia
conoeptionis , Genyatvemus
luteus , Brachydeuterus auritus ,
Gaterin ahvy sotaenia ,
Parapristipoma trilineatum ,
Diagvamma pictum

Sciaenidae

Soiaenops oaellata , Soiaena +

dussumieri , Otolithes dux,
Lavimus acclivus , Cynosoion
vivescens , Mioropogon
undulatus , Argyrosomus iharae

Cheilotrema satuvnum, +

Cynosoion regalis , Nebris

miorops

Baivdiella icistia, +

Bairdiella chrysura , Seriphus
politus , Pareques viola,
Plagioscion sp . , Stellifer sp . ,
Corvula maorops , Paralonchuvus
bvasiliensis , Lonohurus
lanoeolatus , Aplodinotus
grunniens , Paohypops furohaeus ,
Pogonias ohromis , Collichthys
fragi lis

Pachyurus bonaviensis , Umbrina r

ronoador

Collichthys niveatus -

Lobotidae

Lobotes surinamensis +

Datnioides polota -

Gerreidae

Gevres oineveus , Eugerres +

brasilianus , Diapterus
peruvianus , Eucinostomus
dovii , Sohizoptevus auveolus

16 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Leiognathidae

Leiognathus sTplendens , -

Secutor vuoonius

Enoplosidae

Enoplosus armatus + +

Pent ace rot idae

Histiopterus typus +

Pentaoeros japoniaus + ■•■

Oplegnathidae

Oplegnathus fasciatus + +

Toxotidae

Toxotes insidiatov + +

Bathyclupeidae

Bathyclupea argentea + +

Ephippidae

Chaetodipterus zonatus , + +

Platax orhioulavis , Dvepane

punctata

Parapsettus panamensis + ■**

Scatophagidae

Soatophagus sp . ~ ~

Lutjanidae

Lutjanus avgentiventris , - -

Rabirubia inermis , Ooyurus

chrysurus , Rhomboplites

aurorubens , Hoplopagrus

guntheri , Pinjalo pinjalo ,

Macolor niger, Symphoriohthys

spilurus , Aprion viresaens ,

Pristipomoides micvolepis ,

Etelis marshi , Aphareus

furoatus , Apsilus dentatus

Caesionidae

Caesio cuning , Pterocaesio ~ -

tile, Gymnocaesio argenteus ,
Dipterygonotus gruvelii

Nemipteridae

Nemipterus japoniaus , ~ ~

Pentapodus setosus , Scolopsis
bi lineatus

i

No. 121] JOHNSON: PROCURRENT SPUR 17

Lethrinidae

Lethrinus rhodopterus r -

Monotaxis grandooutis ,
Gnathodentex aurolineatus ,
Gymnooranius griseus

Sparidae

Acanthopagrus bevda,
Calamus brachysomus ,
Evynnis cardinalis , Lagodon
rhomhoides , Lithognathus
mormyrus , Taius tumifrons ,
Avohosavgus pouvtalesii ,
Boops hoops , Scatharus
graeous

Centracanthidae

Centracanthus cirrus, - -

Spioava smaris

Incertae sedis

Inermia vittata + +

Pempheridae

Pempheris sokomburghi , - -

Parap viae an thus dispar

Embiotocidae

Cymatogaster aggvegata ,
Amphistichus argenteus ,
Hyperprosopon argenteum ,
Ditrema temninaki

Cichlidae

Tilapia mossambica ,
Geophagus pappaterra ,
Cichlasoma meeki

Pomacentridae

Hypsypops Tuhicunda , - -

Miorospathodon dovsalis ,

Abudefduf tvosohelii ,

Chromis punatipinnis ,

Nexillarius declivifrons ,

Eupomaoentrus re otifvaenum ,

Dascyllus albisella

18 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Chaetodontidae

Forcipiger flavissimus ,
Prognathodes aouleatus ,
Heniochus nigrivostris ,
Pomaoanthus semicirculatus ,
Holaoanthus tvimaculatus ,
Centroipyge bispinosus

Nandidae

Nandus nandus , Monocirrhus
poly acanthus , Polycentvus sp

Badidae

Badis badis

Pristolepidae

Pristolepis sp .

Gadopsidae

Gadopsis mavmoratus

Cirrhitidae

Cirrhitus rivulatus ,
Pavacivvhites forstevi ,
Oxycirvhites typus

Aplodactylidae

Aplodaotylus ethevidgi

Cheilodactylidae

Cheilodaotylus variegatus ,
Aoantholatris gayi

Owstoniidae

Owstonia sp.

Cepolidae

Cepola sohlegeli

Stromateoidei

Centrolophidae

Sohedophilus maoulatus ,
Centrolophus japonious

Seriolella orassus

laichthy s lookingtoni

Stromateidae

Stromateus xanthurus ,
Peprilus similtimus ,
Peprilus palometa ,
Pampus avgenteus

No. 121] JOHNSON: PROCURRENT SPUR 19

Nomeidae

Norneus gronovii +

Psenes pacificus, +

Psenes sio

Cubiceps caevuleus , r

Cuhioeps pauoiradiatus

Tetragonuridae

Tetragonurus cuvieri -

Ariommidae

Aviomma melanum,
Aviomma regulus

Amarsipidae

Amarsipus oarlshergi +

Labroidei

Labridae

Bodianus diplotaenia,
Halichoeves dispilus ,
Tautoglabrus adspersus

Scaridae

Soavus ghobban, Nicholsina
denticulatus

Odacidae

Neodax balteatus , -

Siphonognathus avgyrophanes

Trachinoidei

Trichodontidae

Triohodon trichodon

Opistognathidae

Opistognathus punatatus ,
Lonahopisthus sp .

Ba t hy mas te r i dae

Bathymastev caeruleofasoiatus

Mugiloididae

Parapercis cephalopunotata

Trachinidae

Traohinus draco -

Percophididae

Bembvops gobioides -

20 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Dactyloscopidae

Daotylosaopus tridigitatus

Uranoscopidae

Uranos copus japonicus

Champ sodontidae

Champsodon sp .

Chiasmodontidae
Chiasmodon niger

Scombroidei

Scombrolabracidae

Soombrolabrax hetevolepis

Gempylidae

Gempylus serpens,
Nealotus tripes

Trichiuridae

Lepidopus xantusi

Scombridae

Scomber japonicus , Thunnus
atlanticus , Scomberomorus
conco tor

Istiophoridae

Istiophorus platyptera

Notothenoidei

Nototheneidae

Trematomus nicolai ,
Notothenia Zongipes

Bovichthyidae

Cottoperca gobio

Gobeoidei

Gobiidae

Eleotris sandwicensis ,
Bathygobius ramosus ,
Coryphopterus nicholsi

Mi erode smidae

Microdesmus floridanus ,
Cerdale ionthas

Aitvmody toi de i

Ammodytidae

Ammodytes tobianus

No. 121] JOHNSON: PROCURRENT SPUR 21

Blenniodei

Blenniidae

Hypsoblennius gilberti

Clinidae

Labr-isomus multiporosus ,
Neoolinus stephensae

Chaenopsidae

Chaenopsis alepidota ,
Covalliozetus angelica

Tripterygiidae

Axoclinus oavminalis

Acanthuroidei

Acanthuridae

Aaanthurus xanthopterus ,
Zanclus covnuta

Siganidae

Siganus doliatus

Anabantoidei

Anabantidae

Betta sptendens

Osphronemidae

Trichogaster leeri

SCORPAENIFORMES

Scorpaenoidei

Scorpaenidae

Sebastes serrioeps ,
Soovpaena guttata , Pontinus
furairhinus

Triglidae

Pvionotus stephanophvys

Hexagrammoidei

Hexagranunidae

Oxylebius pictus

Anoplopomatidae

Anoptopoma fimbria

Zaniolepidae

Zaniolepis frenata

22

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Platycephaloidei

Platycephalidae

Platy oeiphalus malayanus

Cottoidei

Cottidae

Orthonopias triads ,
Hemilepidotus jordani

. Agonidae

Asterotheoa pentaoantha

A,

01

P(
si

FIGURE 1. Caudal skeleton of Lutjanus argentiventris .
A. Full view, principal rays shaded, area enlarged in B
outlined. B. Enlargement showing typical ventral procurrent
ray series, arrow marks posteriormost (last) procurrent ray.

No. 121]

JOHNSON: PROCURRENT SPUR

23

FIGURE 2.. Caudal skeleton of Ly thrulon flaviguttatum.
A. Full view, principal rays shaded, area enlarged in B
outlined. B. Enlargement showing procurrent spur on
posteriormost ventral procurrent ray (marked by arrow) and
shortening of preceding ray base.

iiniiiuniiH

mill iiiii"ii>' ""'■;:•-; v