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Library of the 

Museum of 

Comparative Zoology 

aulletin of the 

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Volume 135 







No. 1. The Postcranial Skeleton of the Giant Permian Pelycosaur CotijJo- 
rhynchus lomeii B> J. Wilhs Stovall, Llewellyn I. Price, and 
Alfred Sherwood Romer. September, 1966 1 

No. 2. The Stromateoid Fishes: Systematics and a Classification. By 

Richard L. Haedrich. January, 1967 31 

No. 3. Morphology and Relationships of the Holocephali with Special 
Reference to the Venous System. By Barbara J. Stahl. January, 
1967 - ' 141 

No. 4. A Review of the Mesochrysinae and Nothochrysinae (Neuroptera: 

Chrysopidae). By Phillip A. Adams. February, 1967 215 

No. 5. Marine Nematodes of the East Coast of North America. I. Florida. 

By Wolfgang Wieser and Bruce Hopper. April, 1967 239 

No. 6. The Ameiva (Lacertilia, Teiidae) of Hispaniola. III. Ameiva 

taeniura Cope. By Albert Schwartz. April, 1967 345 

No. 7. New Cyclopoid Copepods Associated with Polychaete Annelids in 

Madagascar. By Arthur G. Humes and Ju-Shey Ho. April, 1967 — 377 

No. 8. Proterochompso banionuevoi and the Early Evolution of the 

Crocodilia. By William D. Sill. April, 1967 415 

No. 9. Sihcified Silurian Trilobites from Maine. By H. B. Whittington 

and K. S. W. Campbell. June, 1967 447 


uUetin OF THE 

The Postcranial Skeleton of the Giant Permian 
Pelycosaur Cotylorhynchus romeri 


Museum of Comparafive Zoology, Harvard University 



VOLUME 135, NO. 1 
SEPTEMBER 22, 1966 





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Bigelow, H. B. and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
Reprint, $6.50 cloth. 

Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In- 
sects. $9.00 cloth. 

Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth. 

Lyman, C. P. and A. R. Davve (eds.), 1960. Symposium on Natural Mam- 
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Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 15. (Price list on 
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Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae 
(Mollusca: Bivalvia). $8.00 cloth. 

Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution 
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Proceedings of the New England Zoological Club 1899-1948. ( Complete sets 
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© The President ond Fellows of Harvard College 1966. 




Introductory note. In 1937 fragmentary 
remains of a gigantic new pelycosaur from 
the Oklahoma Pennian were brought to the 
attention of Dr. Stovall of the University of 
Oklahoma, who shortly after published a 
preliminary account of the animal, as Cotij- 
Jorhijnchus romeri, aided by notes and draw- 
ings b\' Llewellyn I. Price and myself ( Sto- 
vall, 1937 ) . Subsequently, many additional 
specimens of this form were found; these 
were prepared under Dr. Stovall's direc- 
tion, and studies and drawings were made 
of much of the material by Mr. Price. It 
was planned that the material be described 
in a joint paper by Stovall and Price, but 
owing to a variety of circumstances, this 
was never written. Nearly thirty years have 
passed, and except for a brief notice and 
figures of the skull in the "Review of the 
Pelycosauria" ( Romer and Price, 1940: 
419-421), no further account of Coty- 
lorhynchus has appeared. Meanwhile Dr. 
Stoxall has died, and Mr. Price is now 
resident in Brasil. Adding to the need for 
description is the fact that Dr. E. C. Olson 
has found additional remains of Coty- 
lorhynclms, and a number of related fomis 
have been discovered in a variety of locali- 
ties and horizons in Texas and even in 
Russia (Olson. 1962: 24-47). In this situa- 
tion, it has been agreed that I should pub- 
lish a general account of the postcranial 
anatomy of Cotylorhynchus. But although 
I alone should be held responsible for any 
inaccuracies or misinterpretations that this 

paper may contain, I have felt that the 
paper should be considered as a joint effort, 
to give credit to Dr. Stovall for his work 
in the collection and supervision of prep- 
aration of the material, and to Mr. Price for 
his invaluable notes and drawings of the 
Oklahoma materials. 

— Alfred Sherwood Romer 


Although a broad band of continental 
Permian beds extends across Oklahoma 
from the Kansas border to the Red River, 
relatively few vertebrates have been dis- 
covered in the Lower Permian of the state 
— quite in contrast to the situation to the 
south, in northern Texas. This relative pau- 
cit>' of finds appears to be due to the gen- 
erally higher rainfall and consequent better 
vegetative covering in Oklahoma, and to 
the more level topography of Oklahoma, in 
contrast to the Texas area where the pres- 
ence of a number of limestones and heavy 
sandstones makes for a more rugged to- 
pography and the development of poten- 
tially fossiliferous "breaks." 

The discovery of abundant remains of a 
large new type of caseid pelycosaur, Coty- 
lorhynchus romeri, was in consequence a 
major event in the history of paleontological 
work in Oklahoma. As noted in Stovall's 
paper (1937: 308), the first find of Coty- 
lorhynchus material was made by Dr. W. S. 
Strain (then a graduate assistant at the 

Bull. Mus. Comp. Zool., 135(1): 1-30, September, 1966 1 

2 Bulletin Miisriiw of Comparative Zoology, Vol. 135, No. 1 

University ot Oklahoma). The site lay in 
the Hennessey shales, 4% miles west of 
Navina, Logan County. The type specimen 
consisted of an incomplete skull and jaw, a 
front foot, and intercla\icle. Sliortly after, 
a nearly complete postcranial skeleton was 
found at about the same horizon, in the 
Hennessey Formation, but close to the city 
of Norman and hence some 50 miles or so 
from the region of the first discovery, and 
during the years 1937-1939 a very con- 
siderable number of further Cotijlorhijnchus 
specimens were collected in the Hennessey 
shales of the Norman district and prepared 
by laborers under Works Progress Admini- 
stration funds. Most of this material is now 
in the Stovall Museum at Norman, luit 
specimens were also obtained for the mu- 
seums in Chicago, New York, Washington, 
and Cambridge. Preparation of the mate- 
rials at Norman was carried out under the 
general direction of Dr. Stovall, and notes 
and figures on much of the material were 
made by Mr. Price. In the preparation of 
the present description, Romer has seen 
all tliis material, and has utilized Price's 
notes and figures in addition to his own ob- 
servations. A word of caution must be 
given, however. Although prc>paration was 
carried out under Dr. Stovall's direction, 
it is not improbable that, with a consider- 
able number of workers and with a number 
of specimens undergoing simultaneous prep- 
aration, a certain amount of inaccuracv 
may have occurred in the restoration and 
as.sociatioii of materials. Further, in the 
time that has since elapsed, some of the 
specimens have been moved several times 
for storage and re-cataloging and further 
chances of error have crept in. 

The reptilian remains from the Hennessey 
.shales of the Norman region appear, curi- 
ously, to pertain almost entirely to Cotij- 
lorlnjncluis, with very few remains of other 
reptiles and amphibians — a situation in 
strong contrast to the usual mixed assem- 
blages generally present in typical Texas 
redbeds deposits. As far as can be seen, 
the Cotylorhynchus materials from the Nor- 

Table 1 

Specimen No. 









4-0-2 (1249) 








CiN'HM 272 





AMNH 7517 


1250 (?) 










251 + 


MCZ 3416 










man area can be assigned to a single species 
as far as morphological characters are con- 
cerned.^ There is however, considerable 
variation in size. In reptiles generally, early 
growth is rapid, followed by a slower if 
steady growth; the natural result is that the 
greater part of specimens recovered in the 
case of fossil forms should represent rela- 
tively young mature animals, with a small 
percentage of overly large specimens repre- 
senting indi\'iduals which were exception- 
ally long-lived and consequently exception- 
ally large, and a small percentage of young 
and immature individuals. The present ma- 
terials tend to show a rather greater size 
range than is common. As a rough index to 
size, I list the length (in mm) of major limb 
elements in a number of the better speci- 
mens (Table 1). 

These specimens are deposited in the 
following institutions: Chicago Natural 
History Museum ( CNHM ) ; American Mu- 
seum of Natural History (AMNH); United 
States National Museum (USNM); Museum 
of Comparati\'e Zoology, Harvard ( MCZ ) . 
Other specimens (numbered) are in the 
University of Oklahoma collection. 

As can be seen from this table, the first 
five specimens listcxl appear to be of rela- 
tively uniform large size. Below this there 
is a sharp drop to three specimens which 
are 20 per cent or so smaller than the first 

^ Dr. Olson informs mo that there is a possible 
specific tUfferencc Ix-lwecn the Norman material 
and the type from Navina. 

COTYLORHYNCHUS SKELETON • StovaU, Price, and Rotner 

group but appear to be essentially mature; 
below, with the MCZ specimen as a tran- 
sition, there is a drop to small and seem- 
ingly immature specimens such as 4-0-3. 
Possibly there is a specific difference be- 
tween the first and second groups. Equally 
possible, however, the contrast is a sexual 
one; in Dimetrodon limhatiis, for example, 
there is a seemingly clear-cut size difference 
of 10 per cent between the sexes (Romer 
and Price, 1940: 341-342). 

In typical Texas redbeds localities, fossil 
reptile remains are most commonly found 
in disarticulated condition, and when found 
articulated appear to have undergone de- 
position in a variety of poses. One gets the 
impression that this is the result of stream 
action, and that many of the "bone pockets" 
represent back eddies in streams in which 
cadavers brought downstream tended to 
collect (and decompose). But in the Hen- 
nessey shales of the Nomian region, many 
of the specimens show clearly that the 
CotylorhyncJnis individuals were generally 
buried in articulated and undisturbed 
fashion, right side up, with the limbs spread 
outward at the sides. This strongly suggests 
that we are here dealing with entombment 
of quite another sort. South African Per- 
mian pareiasaurs, equally large and clumsy 
herbivores, are typically preserved in simi- 
lar fashion (Watson, 1913). There are two 
possible interpretations. Watson, in the 
case of the pareiasaurs, implies death from 
starvation, and suggests a covering of the 
cadavers in situ by loess-like materials. I 
would favor an alternative interpretation in 
both cases — that the animals were bogged 
down and entombed in swamps. 

Study has been greatly handicapped by 
the nature of the specimens. Due in part, 
perhaps, to a rather spongy structure of the 
skeletal elements, and to the nature of the 
sediments in which they were embedded, 
much of the material has been subjected to 
crushing and distortion, with a consequent 
limitation of the accuracy with which res- 
toration and illustration can be made. In 
great measure description and illustration 

of structures given here are based on a 
synthesis of a number of specimens. Wher- 
ever possible the illustrations are based on 
specific examples, although frequently with 
the addition from other individuals of de- 
tails missing or obscured in the specimen 
primarily utilized. 

It became apparent, even from the ma- 
terial described in Stovall's preliminary 
paper, that CotijJorhijnchus was a giant rel- 
ative of Casea, a small Clear Fork Texas 
pelycosaur described by Williston (Willis- 
ton, 1910: 590-592; 1911: 111-131, etc.; 
Romer and Price, 1940: 412-419). Despite 
cranial differences, Casea and a number of 
other Texas pelycosaurs show such a large 
number of diagnostic postcranial similari- 
ties to Edaphosaurus that Romer (Romer 
and Price, 1940: 21, 366-378) felt justified 
in including these forms with Edaphosaurus 
in a common suborder Edaphosauria. As in 
Casea, the CotylorJiynchus postcranial skele- 
ton agrees in almost every particular with 
the characteristics assigned to the Edapho- 
sauria as a whole. In consequence, in the 
description which follows, comparisons are, 
in general, with Casea or other edapho- 
saurian types. 


The general characteristics of the Coty- 
lorhynchus vertebral column are almost ex- 
actly those cited in the "Review of the 
Pelycosauria" for the suborder Edaphosauria 
as a whole: 

"The number of presacral vertebrae is sometimes 
reduced. The dorsal vertebrae are moderately to 
greatly elongated; the cervicals are small in all 
dimensions. The dorsal centra are spool-shaped, 
with rounded bottoms, and lack any trace of ven- 
tral keel or lip. Intercentra are small. The dorsal 
transverse processes are moderately elongated and 
rise from a more antero-dorsal position on the arch 
than in other pelycosaurs. The zygapophysial 
surfaces are large, moderately tilted, and extend 
farther laterally than in sphenacodontids. The 
neural arches are not excavated above the dia- 
pophyses. . . . The atlas centrum reaches the ven- 
tral surface of the column." 

Bulletin Museum of Comparative Zoology, Vol. 135. No. 1 

Onl>- a very few qualifications on this tenninal, are present in seemingly articu- 
definition need be made to fit Cotylorlujn- lated fashion. However, there is a suspi- 
chiis. (1) It was stated that the presacral cionsly sharp drop in the size of centra 
count is "sometimes" reduced. At the time following element 27, and a photograph of 
this was written, the figure was known the specimen made during preparation 
only in Casca where apparently 24 or 25 shows that the last 16 vertebrae were not 
presacrals were present ( Romer and Price, part of the block containing the main part 
1940: 417). It is now known that Edapho- of the material, although they may well 
saiirus boanerges has likewise reduced the pertain to the same individual. Measure- 
primitive pelycosaur number from 27 to 23 ments suggest that there is a gap here of 
presacral vertebrae (Shuler and Witter, about 10 vertebrae, to raise the probable 
1942). It is thus a reasonable inference that total count to about 53. In the CNHM 
presacral reduction was general in the sub- specimen, 37 caudals are present in articula- 
order, and reduction is present in Coty- tion with the trunk; following a gap of 
lorhynchus. (2) "Intercentra are small." about 23 cm in the slab there are eight 
Few are present in any Edaphosaurus ma- further, much smaller, vertebrae, the last 
terial; only a single intercentrum was found apparently terminal or sub-terminal. The 
in the Casea material (in the sacral region); gap is of such length as to suggest that 10 
none are known in Cotylorhynchus except vertebrae are missing here, giving a prob- 
for the atlas-axis. Small gaps between the able total length of 55 vertebrae. Tlris is a 
ventral edges of the centra in all three reasonable pelycosaurian number, 
genera suggest that tiny intercentra were Dorsal veriehrae. Such a dorsal vertebra 
present in cartilaginous fonn. (3) "The as, for example, vertebra 12 of 4-0-6 (Figs. 
atlas centrum reaches the ventral surface of IC, 2C ) , exhibits a typical pelycosaurian 
the column." This appears to be true of and, further, edaphosaurian character. The 
Edaphosaurus, and WilUston (1911: pi. centra are edaphosauroid — spool-shaped 
XIV ) restores this centrum as reaching the structures without any of the tendency seen 
ventral surface in Casea. As described be- in ophiacodontoids and sphenacodontoids 
low, however, it does not reach this surface for development of a ventral keel. The 
in Cotylorhynchus, and Williston's material centrum, as in Casea, is relatively short and 
of Casca was obviously imperfect and open stout, its length, averaging about 60 mm in 
to misinterpretation. large individuals, being but about IV2 times 

The Cotylorhynchus romcri column in- the end height of 38-40 mm. The ventral 

eludes definitely 25 and probably 26 pre- surface is broad, essentially flattened, but 

sacral vertebrae, 3 sacrals, and about 55 slightly convex; at the margins of this area 

caudals. It has proved difficult to deter- the sides curve upward sharply and, above 

mine the presacral count in Co^y/or/if/nc/H/.s. this point, are slightly in-pinched. There 

There are several specimens in which dor- is no visible suture between arch and cen- 

sals, lumbars, and sacrals are present in trum. On the sides of the centrum, at about 

well-connected series; generally, however, the level where the base of the neural arch 

the cervicals are poorly preserved or absent, pedicel would be expected, there is a low 

In 4-0-6, however, the cervicals are present, longitudinal ridge, rounded at its summit, 

There are definitely 25 well-preserved pre- running lengthwise from a point well below 

sacral vertebrae, and apparently one further the upper margin of the centrum anteriorly 

poorly preserved one. Several specimens back to a somewhat higher position pos- 

show a sacrum of three vertebrae. The teriorly. Above this point the upper part of 

tail is preserved in but few cases. In the the surface of the centrum and the lower 

mounted slab of 4-0-2 (1249), a hind leg lateral surface of the neural arch form a 

and tail, 43 caudals, the last apparently longitudinal depression extending length- 

CoTYLORHYNCHUS SKELETON • StovdU, Piicc, and Romcr 5 

Fig. 1. Lateral views of representative vertebrae, in side view. A, atlas-axis, X %; 8/ vertebra 6, X Vi; C, vertebra 12, 
X Vv D, vertebra 21, X Vv f' proximal caudal, X 'A- 

wise beneath the base of the transverse 

The ends of the centrum are of a typical 
pelycosaurian nature — essentially circular in 
outline, with thickened edges. At either 
end this circle contains a cone-shaped de- 
pression for the notochord, the two cones 
connected by a small foramen. Both an- 
teriorly and posteriorly the rim of the cen- 
trum is somewhat thickened laterally for 
the articulation of the capitulum, which 
was thus inter-central in position. 

The transverse processes are highly de- 
veloped in this region, extending outward 
on either side to a distance from the mid- 
line approximately equal to the vertebral 

length. In edaphosauroid fashion the trans- 
verse processes arise from a far anterior 
position on the neural arch, and their front 
margins extend almost directly outward from 
the level of the prezygapophyses. There is 
in this region no ventral expansion of the 
process, which is thin dorsoventrally but 
broad anteroposteriorly; there is thus no 
continuum in articular areas between ca- 
pitulum and tuberculum. The transverse 
process here is directed almost straight lat- 
erally, with, however, a slight upward and 
forward tilt, rather than the downward slant 
of most pelycosaurs ( Cosea excepted ) . The 
base of the process is very broad, extending 
nearly the whole length of the vertebra. 

6 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 2. Posterior views of the same vertebrae as those of Figure 1. 

The broad articular surface for the tuber- 
cukxm faces ventrally and only slightly lat- 
erally on the under surface of the process. 
Since the tuberculum is little raised, the 
proximal portion of the rib continues out- 
ward in the line of the transverse process. 
The zygapophyses are normally constructed, 
with a very slight median tilt of the articular 
surfaces. The neural arches, as in edapho- 
saurs generally, lack the lateral excavation 
seen in sphenacodontoids above the trans- 
verse processes. The neural spines, as in 
Casea, are slender transversely but long 
anteroposteriorly, and are low (although 
not as low, relatively, as in the last-named 
genus). In such mid-dorsals as are com- 

pletely preserved the rugose end of the 
spine is expanded laterally on either side 
and indented medially, to give a somewhat 
bifurcate appearance, much as in, for ex- 
ample, some parts of the En/ops column. 

CcrvicaJs. Progressing forward through 
the anterior dorsals into the cervical series, 
the vertebrae are increasingly lightly built, 
as tends to be true of pelycosaurs generally, 
but more especially of edaphosauroids, such 
as CotylorJiyncJws; in which the head is 
relatively small. The centra decrease steadily 
in diameter and length anteriorly, and the 
width tends to equal the length. In the 
6th vertebra of 4-0-4, for example (Figs. 
IB, 2B), the length is about 50 mm, the 

CoTYLORHYNCHUS SKELETON • StovciU, Price, and Romer 7 

width about 45 mm. The flattened ventral 
surface of the centra persists into the cer- 
vical region. In the cervicals the transverse 
processes are shorter and lightly built, in 
correlation with the decreased size of the 
ribs. Anteriorly, the cleft in the summit of 
the neural spines disappears, although some 
distal expansion may persist; the spines be- 
come shorter and somewhat more rounded 
— ovoid — in section. Proceeding forward, 
the direction of the transverse processes 
gradually shifts. In a typical dorsal it points 
some\\'hat forward and upward, but in the 
cervicals the processes become somewhat 
ventrally and posteriorly directed, in cor- 
relation with the direction of the ribs in the 
relatively slender neck region. Even as far 
forward as vertebra 3, the capitular facet is 
high up on the rim of the centrum, showing 
little of the tendency to descend anteriorly, 
seen in most pelycosaurs. The cervical 
zygapophyses are, like the dorsals, widely 
separated and with essentially horizontal 
articular surfaces. 

Atlas-axis. The atlas-axis complex is pres- 
ent and well preserved in 4-0-6, and is 
present also in 4-0-4 (Figs. lA, 2A). A 
facet on the atlas neural arch indicates the 
presence in life of a proatlas. The atlas 
arch is bipartite and typically pelycosaurian, 
with a long posterior process articulating 
with the prezygapophysis of the axis and 
with a spur running farther back along the 
side of the axis neural arch. There is a 
short but distinct transverse process for rib 
attachment, turned strongly downward and 
backward. The atlantal intercentrum is 
highly developed for cranial articulation; 
laterally there is a distinct facet for the 
rib capitulum. As is well known, the atlas 
centrum is variable in development in pely- 
cosaurs, reaching the ventral surface in 
Dimetrodon, for example, but excluded from 
this surface bv the second intercentrum in 
Ophiacodon. The situation in Cotyloihyn- 
chiis is not too clear. In both specimens 
in which the atlas-axis is preserved, there 
is a distinct ventral gap between inter- 
centra 1 and 2, but the atlas centrum does 

not reach the surface here; possibly this gap 
may have been filled by a ventral car- 
tilaginous continuation of the bone. 

Intercentrum 2 is very well developed 
and bears a distinct rib facet. The axis cen- 
trum and arch are built in typically pely- 
cosaurian — and especially edaphosaurian — 
fashion. The transverse process slants 
strongly downward and to a slight degree 
backward. The neural spine is, for an axis, 
rather short, but is stout, long anteropos- 
teriorly, and expanded in a 3-pronged ar- 
rangement posteriorly toward the summit. 

Posterior dorsals. Passing backward along 
the dorsal series, the vertebral centra in- 
crease somewhat in length to about verte- 
brae 17-19, beyond which a progressive 
reduction takes place so that beyond this 
point (as is also the case anteriorly) width 
exceeds length. In 4-0-4, for example, the 
length of vertebra 21 (Figs. ID, 2D) is 51 
mm, the breadth about 55 mm. An increase 
in central width, however, persists, width 
reaching its maximum in the sacral region, 
and the posterior dorsals are the most mas- 
sive vertebrae in the column. In the more 
posterior presacrals the neural spines are 
increasingly long fore-and-aft and increas- 
ingly narrow in transverse diameter, al- 
though the arches are broadly expanded at 
the spine bases. In the posterior dorsal 
series the two members of each pair of 
prezygapophyses are joined by a horizontal 
ridge of bone bridging the space between 
them and forming a strengthening trans- 
verse element wliich crosses the front end 
of the vertebra between the ribs of either 
side. Progressing backward, the transverse 
processes are increasingly stout but decrease 
in length. In the last members of the series 
the processes are fused with the ribs. In 
the posterior vertebrae the area of attach- 
ment of the rib capitulum moves upward 
and backward to an oval area on the an- 
terodorsal surface of the side of the cen- 
trum below the transverse process. Here, 
as in the transverse process, fusion with the 
rib takes place in the last few members of 
the dorsal series. 

8 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 3. Left, ventral view of last presacral, the three sacrals, and first two caudal vertebrae, X 'A- '?ighf, obove, lateral 
view of mid-caudal vertebrae, X Vi. Right, below, posterior and ventral views of a mid-caudal vertebra and a mid- 
caudal chevron, X 'A- 

Sacrals (Fig. 3). Three sacral vertebrae 
are present, as noted in the description of 
the ribs. The centra are broad but appear 
to be shallow dorsoventrally ( an effect per- 
haps due to crushing). In one specimen, at 
least, the centra of the first and second 
sacrals appear to be fused; whether the last 
two were fused is uncertain. On the first 
sacral the zygapophyses are still well sep- 
arated, but on the third vertebra the two 
members of each pair are much closer to- 
gether, a situation transitional to that in 
the tail. There appears to be no ossification 
of apposed zygapophyses. The transverse 
processes — firmly fused to the ribs — are 
short, but very stout, and are continuous 
with a large area on the centrum represent- 
ing the capitular attachment. A depression 
which represents the point of junction of 
tubercular and capitular areas is seen in 
some specimens; it cannot be detennined 
whether an arterial foramen perforates the 

structure. The neural spines of the first two 
sacrals are similar to those of the "lumbars"; 
that of sacral 3 is shorter fore-and-aft and 
less compressed from side to side. 

Caudah (Figs. IE, 2E, 3). The most 
anterior caudals resemble the last sacral 
closely. The breadth of the centrum is 
greater than the length or height; the rib 
attachment areas are short but greatly ex- 
panded, with a groove, presumably for an 
intersegmental artery, marking the line of 
distinction between tubercular and capitu- 
lar components. More posteriorly, the trans- 
verse processes become greatly reduced in 
size (with a concomitant reduction in rib 
size), and process and rib have essentially 
disappeared by vertebra 12, although a 
slightly projecting ridge is present as far as 
vertebra 20. Meanwhile, the centra have 
begun a change in their proportions, so that 
by mid-length of the tail the length is con- 
siderably greater than the width and the 

CoTYLORHYXCHUS SKELETON • StovciU, Piicc, and Roiuer 9 


5 4 3 2 


Fig. 4. Above, right ribs 1-8, seen from the posterior aspect, X Vi- Below, the heads of right and left ribs from the 
same mid-dorsal segment of a single individual, to show contrasts in post-mortem distortion, X 'A- 

height also exceeds the \\'idth. The zygapo- 
ph)'sial pairs have come to He close to each 
other near the mid-Hne, and their articular 
surfaces have become sharply tilted to a 
plane close to the vertical. There is, further, 
a gradual reduction in height and stoutness 
of the neural spines, so that by the time 
the mid-caudals are reached, the spine is 
a low nubbin projecting a short distance 
above tlie postzygapophyses. The neural 
arch as a whole is much reduced, and in the 
most posterior part of the tail the material 
shows httle evidence of any structure above 
the centra. 

The first of the chevrons appears, in the 
CNHM specimen (No. 272), between the 
3rd and 4th caudals; they continue back 
as far as vertebra 36, at least in one speci- 
men. The first elements are stoutly built, 
with a length equal to two centra; as is 

normal, the length decreases posteriorly, 
although but slowly. In one specimen, as in 
pelycosaurs generally ( and in Casea ) , there 
is, in the proximal elements, a basal inter- 
central component connecting the two arms 
of the chevron; but in the CNHM specimen 
not even the most proximal chevrons have 
a proper intercentrum, there being merely 
an enlargement of each of the two proximal 
ends of the chevron structure. Although 
both of the adjacent centra are bevelled 
for reception of a chevron, the centrum 
anterior to the chevron has the most ob- 
vious articulations; it bears ventrally a pair 
of well developed facets, each supported by 
a longitudinal ridge on the centrum. In the 
tail the bottom of the centrum shows a 
longitudinal depression, bounded on either 
side by these ridges. 

10 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 5. The proximal part of right ribs 9-21, seen from the posterior aspect, X 'A- 

CoTYLORHYNCHUS SKELETON • Stovdl, Piicc, and Romer 11 


As in all known pelycosaurs, ribs were 
present on every vertebra from the atlas to 
and including the proximal caudals. For 
most parts of the series, specimens 4-0-6 
and CNHM 272 furnish the best material. 

Dorsal ribs (Fig. 5). In a typical dorsal 
rib the shaft extends outward (and in life 
somewhat upward) from the capitulum, 
the articular surface of which is an oval, 
narrow anteroposteriorly, occupying the en- 
tire head of the rib. The tuberculum in the 
mid-dorsals does not project to any marked 
degree from the shaft; it apposes to the 
transverse process a concave articular sur- 
face with an oval shape, broader distally, 
the length about twice the width. 

The proximal end of the rib appears to 
run distally slightly upward to and past the 
tubercular region. Shortly beyond this point 
the rib curves markedly outward and down- 
ward to encase the flank. The degree of 
curvature in life is difficult to detennine, 
due to variable post-mortem crushing and 
compression; see, for example, the marked 
contrast between the two members of a rib- 
pair from the same segment of a single in- 
dividual shown in Figure 4. In most in- 
stances there is a considerable segment of 
the rib, lying beyond the point of proximal 
cunature, which is nearly straight; this in- 
cludes about half the total rib length. Tliis 
segment appears to have been directed di- 
agonally outsvard and downward. Distally, 
beyond this straight section, the rib curves 
gradually inward toward the mid-line. Al- 
though seldom is a complete rib length 
preserved, it would appear that the greatest 
length of a rib (measured from the tuber- 
culum) is about the length of 13 dorsal 
centra; two mid-dorsals of CNHM 272 mea- 
sure 721 and 718 mm. In such dorsals as are 
completely preserved the distal end of the 
rib exhibits a cupped tenuination with 
which the cartilaginous section of the rib 
presumably articulated. 

Despite the handicap noted as due to 
post-mortem distortion, articulation of ribs 

and \ ertebrae gives, on the average, a pic- 
ture closely comparable to that gi\en by 
Williston (1911: pi. XVII, fig. 1) for a 
Casea mid-dorsal, and shows the presence 
of a very broad, barrel-like trunk, with a 
probable transverse diameter of about 60 
cm — i.e., about two feet. 

Except for the rather flattened capitular 
region, typical dorsal ribs are essentially 
oval in section, and so tilted in life that the 
long axis of the oval slants downward and 
forward well over 45°, with the two major 
surfaces essentially anterior and posterior 
in position. The anterior margin is in all 
typical dorsals smoothly rounded. On the 
posterior surface a ridge develops at a point 
somewhat distal to the tuberculum and 
rather toward the ventral margin. This ridge 
extends outward along the straight lateral 
segment of the rib, gradually approaching 
the ventral margin of the rib and, decreas- 
ing in prominence, disappears at about the 
point where the median curvature begins. 

There is little expansion of typical dorsal 
ribs at any region. The diameter of the rib 
is greatest a short distance distal to the 
tuberculum; beyond this region there is 
generally a very gradual decrease to a mini- 
mum of about two-thirds of the maximum. 

Progressing anteriorly, from such a typi- 
cal dorsal as that described, to rib 9, a 
series of gradual changes may be observed. 
The length decreases, rib 9 in 4-0-6, for 
example, being but 430 mm in length, i.e. 
about 60 per cent as long as the longest rib, 
and the ribs are more lightly built. In an- 
terior dorsals, in contrast with mid-dorsals, 
the tuberculum projects markedl\- above 
the general outline of the dorsal rib margin, 
and, in contrast, the capitulum is sharply 
turned downward medially from the line of 
the shaft. There is here only a slight curva- 
ture beyond the tubercular region; when 
the rib is articulated with the vertebra, it is 
obvious that its shaft runs more directly 
downward than in the typical dorsals, in- 
dicating a narrowing of the bod>' in the 
"chest" region, as the level of the shoulder 
girdle is approached. Distal to the straight 


Bulletin Museum of Comparative Zoology, Vol. 135, Xo. 1 

descending portion of the shaft, there is a 
final segment curving somewhat inward. 
The total distance from the tuberculum to 
the beginning of the median curvature is 
little less than in rib 14, indicating that the 
"chest" is nearly as deep as the belly, al- 
though narrower. 

The longitudinal ridge found on the pos- 
terior rib surface changes in position as one 
progresses fonvard. It has shifted upward, 
so that its proximal portion lies along the 
dorsal margin of the rib, and its distal end 
lies at about the middle of the posterior 
surface, rather than toward its ventral mar- 
gin. It has, further, moved proximally, so 
that its proximal end has attained the distal 
margin of the tubercular projection, and its 
distal end extends only part-way down the 
vertical segment of the rib. Distally the 
anterior dorsal ribs remain sub-circular in 
section. Proximally, however, the posterior 
surface becomes much hollowed out, with 
the proximal part of the longitudinal ridge 
forming a very prominent projection dor- 
sally. As far as can be determined in the 
usually crushed condition of the specimens, 
the most anterior dorsal ribs are somewhat 
expanded distally — notably rib 9 in 4-0-6. 

Cervical ribs (Fig. 4). In the few pre- 
viously known pelycosaurs in which the 
distal ends of the ribs have been well pre- 
served, it appeared that rib 8 was stout 
distally and presumably had a sternal con- 
nection, establishing 7 as the probable num- 
ber of non-sternal, i.e., cervical, ribs. In 
4-0-6, however, rib 8, although elongate 
(with a length of about 360 mm) and 
resembling the dorsal rib following it in 
most regards, tapers to a point distally. It 
thus failed, obviously, to reach the sternum 
and must be considered technically to be a 

In 4-0-6, remains of all the cervical ribs 
are present (but the third is poorly pre- 
served). Although the rib-tips are incom- 
plete, it is clear that all were slender and 
pointed distally. From rib 8 forward to rib 
4 there is a sharp and steady diminution in 
length, and tlic preserved portions of the 

most anterior ribs indicate that these ribs 
were shorter still. In correlation with the 
wide separation of the two points of attach- 
ment to the vertebrae, tuberculum and 
capitulum are strongly divergent, giving a 
V-shape to the proximal part of the rib. 
The slender distal portions of the ribs ap- 
pear to be somewhat compressed antero- 

In the cervicals, as in the most anterior 
dorsals, the rib shaft runs straight distally 
from the tuberculum, indicating a narrow 
neck region; the tuberculum, as far as pre- 
served in these ribs, retains the somewhat 
distinct character seen in the most anterior 
dorsals. On rib 8 the longitudinal ridge 
retains the character seen in the rib follow- 
ing for much of the proximal half of the 
shaft, fonning a sharp dorsal margin of the 
rather thin rib, but it fails to reach the 
tuberculum. On rib 7, the ridge is present 
on the middle third of the shaft; on the 
more anterior ribs it has disappeared. It 
is in a sense replaced by a thin dorsal flange 
extending distally on rib 7 a short distance 
outward from the tuberculum, but not con- 
tinuous distally with the typical dorsal ridge. 
This flange is present in reduced form on 
rib 6; more anteriorly, as far as can be seen 
from the material, no noticeable structures 
are present on the rib shafts. Ribs 6-8 
appear to have lain beneath the scapula; 
these reduced flanges, presumably func- 
tioning for the origin of serratus muscula- 
ture, contrast with the much greater flange 
development seen in many early tetrapods. 

Posterior dorsal ribs (Fig. 6). Proceed- 
ing backward along the dorsal series, the 
posterior dorsal ribs as far as about rib 20 
appear to be essentially similar to more 
anterior dorsals in character, and show little 
decrease in length; however, the proximal 
end of the longitudinal ridge described for 
the anterior dorsals retreats distally to a 
small extent in this region. Rib 20 becomes 
broader proximally; in the posterior ribs 
there is a gradual approximation of tuber- 
cular and capitular heads so that, from 
about rib 22 back, capitular and tubercular 

COTYLORHYNCHUS SKELETON • StovaU, Pitcc, and Roiyier 13 






Fig. 6. Left, posterior presacral ribs of the right side, X 'A- Rig^^i, left clavicle, external and internal views, X Vi- 

areas are essentially fused into a single 
articular surface. A decrease in length of 
ribs is not marked until approximately rib 
23, which is notably shorter than that pre- 
ceding it (with a length in CNHM 272, 
as preserved, of 413 mm) and ribs 24-26 
are increasingly short, the last being about 
158 mm long in CNHM 272. As in pely- 
cosaurs generally, the freedom of rib artic- 
ulation with the vertebrae decreases pos- 
teriorly; the last 3 short ribs appear to be 
well fused and immovable and hence are to 
be considered as lumbars, and the next pre- 
ceding may also have been immovable in 
life, although in 4-0-4 a suture between rib 
and vertebra was seen during preparation 
for the fourth presacral. 

All the more posterior ribs tend to have 
shafts which are relatively broad and thin 
for most of their length; the last 5, however, 
definitely taper to a point distally, and thus 
lack a sternal connection, and the same 
may be true of rib 21. As far as segment 
23 the ribs continue to be curved, turning 
downward and backward in life. Ribs 24- 
26, however, are nearly straight and di- 
rected laterally, their length being little 
more than that of the proximal nearly 
straight segment of rib 23. The breadth of 
the proximal part of the ribs increases pos- 

teriorly to a maximum in rib 24, which is 
very broad in proportion to its length. 

Sacral ribs (Fig. 3). In correlation with 
the width of the trunk and pelvic region, 
the sacral ribs are longer than in sphena- 
codonts and ophiacodonts. Three sacral 
ribs are present. These are tightly fused 
to their vertebrae over a broad area; this 
includes the short transverse processes which 
arise from much of the lateral surface of the 
centra. No clear sutures are seen, but dor- 
sally a rugose anteroposterior ridge marks 
the line of fusion of rib and transverse proc- 
ess. The rib head is deep dorsoventrally, 
as well as anteroposteriorly. Distinction be- 
tween capitular and tubercular attachment 
is indicated by depressions on both anterior 
and posterior surfaces about half way down; 
these depressions have not been excava- 
ted, but may have been connected by a 
canal for the intervertebral artery. 

The first sacral rib is short but stout. 
Beyond the head it contracts somewhat in 
width and extends outward horizontally and 
slightly posteriorly to terminate in a rela- 
tively thin expanded blade extending down- 
ward and apposed laterally to the inner 
face of the ilium. The second rib extends 
directly laterally; it is similar to the first 
but slightly less developed. The third rib 


Bulletin Muscitm of Comparative Zoology, Vol. 135, No. 1 

Fig. 7. Left, external, and right, internal views of the left scapulocoracoid. Broken lines on the internal view outline the 
portion preserved in No. 4-0-6. Center, dorsal, right lateral, and anterior views of the interclavicle. X \/b- 

is essentially accessory in nature; it is more 
slenderly built and curves forward to but- 
tress the second rib as well as apposing the 
ilium with an only slightly expanded tip. 

The first two sacral ribs are fairly com- 
parable to those of Casca. In that genus, 
however, sacral rib 3 is well developed, and 
Cotylorhyndnis here exhibits a condition 
which is less advanced. Edapliomiini.s 
shows an intermediate condition in the de- 
velopment of sacral rib 3. 

Caudal ribs. As in pelycosaurs gener- 
ally, ribs are present in the proximal tail 
region. Tlie first five show a fused attach- 
ment to transverse process and centrum with 
a pattern similar to that of the sacrals, al- 
though \\'ith a steadily diminishing attach- 
ment area; the heads are pierced antero- 
posteriorly by canals for the intervertebral 
artery. The first caudal is comparable to 
the third sacral in most regards, but it ex- 
tends directly laterally, with a slight distal 
posterior curvature, to terminate in a pointed 
tip. Posterior to this, all the caudal ribs, as 
in pelycosaurs generally, continue to extend 
outward in a horizontal plane and to taper 
to distal extremities. In caudals 2-5 a back- 

ward curvature of the distal end is pro- 
nounced. These ribs show a steady decrease 
in length and stoubiess. Rib 6 is notably 
shorter, with little development of a distal 
curved segment, and from this point back 
the caudal ribs, fused to the transverse proc- 
esses, are laterally projecting, tapering, 
and pointed structures which decrease to 
small nubbins and disappear, except for 
low rugosities, beyond vertebra 11. In gen- 
eral the caudal ribs are comparable to those 
of Casea. 


Shoulder girdle. A nearly complete scap- 
ulocoracoid, as seen from the inner side, 
is present in 4-0-6, and the figure is based 
primarily on this specimen. The outer sur- 
face is not available in this specimen, but 
several others show this aspect. The scapu- 
lar blade, incomplete in 4-0-6, is better 
preserved in other specimens; its exact 
height, however, is difficult to determine, 
due to the lack of distinctive features in this 
area. The blade in large specimens may 
have been somewhat more developed. In 


all available specimens the dorsal margin of 
the blade shows an unfinished surface, so 
that there may have been a considerable 
cartilaginous suprascapula. No sutures be- 
tween the presumed three elements could 
be determined. 

The scapulocoracoid (Fig. 7) is edapho- 
saurian in general character. It is short dor- 
soventrally, broad anteroposteriorly, with 
much the proportions of Lupeosaurus 
( Ccisca, presumably in relation to its smaller 
size, has a girdle of more slender build). 
The scapular blade is very short and broad, 
its breadth distally due to a backward flare 
of the posterior margin greater than is seen 
in any other pelycosaur. Below the point 
of cla\'icular attachment the margin of the 
scapula curves out widely anteriorly, to 
give great breadth to the lower part of the 
blade, as in Edaphosaurus and Lupeosaurus. 
In contrast to Edaphosaurus and Lupeosau- 
rus, but in agreement with Casea, Nito- 
satirus, and Mycterosaurus, there is no 
supraglenoid foramen. There is, as in all 
pelycosaurs, a screw-shaped glenoid cavity; 
it is here, as in other edaphosaurians, re- 
markably deep anteriorly. Below its anterior 
end is the external opening of the supra- 
coracoid foramen. Posteriorly, the coracoid 
region shows little development of a tu- 
bercle for origin of the coracoid head of the 
triceps. On the inner surface, the upper 
portion of the subcoracoscapular fossa is 
shallow, in correlation with the absence of 
a supraglenoid foramen. 

No cleithrum is present in the available 
material. Much of the clavicles and the 
interclavicle are present in 4-0-6 and in the 
type (Figs. 6, 7). In contrast to Edapho- 
saurus and sphenacodonts, there is little 
ventral expansion of the clavicle (the clavicle 
of Casea is unknown ) . Its upper portion is 
well grooved posteriorly to clasp the anterior 
edge of the scapula. Tlie blade of the in- 
terclavicle is unusuallv broad and short, 
and there is no development of the longi- 
tudinal ridge seen on the ventral surface of 
the shaft in most pelycosaurs. As preserved, 
the head turns sharply upward on the shaft. 

in contrast to the gentler curvature found 
in pelycosaurs generally, so that the sur- 
faces apposed to the clavicles are essen- 
tially in a vertical plane. The head of the 
interclavicle is short and T-shaped, in con- 
trast to the diamond-shaped head of many 
pelycosaurs, and is nearly completely oc- 
cupied by the pair of crescent-shaped sur- 
faces for the clavicles. These surfaces, which 
extend far laterally, are covered by rugose 
striations; the areas on the clavicles which 
meet them are similarly rugose, and set 
in distinctly below the general level of the 
"inner" surfaces of that bone. Obviously 
the contact bet\veen clavicles and inter- 
clavicle was an intimate one. 

Pelvic girdle. Pelvic girdle material 
(Fig. 8) is relatively poorly represented. 
The Cotylorhynchus skeletons were nearly 
all buried with the dorsal surface upward; 
the ilia in consequence have been generally 
subject either to damage by crushing dur- 
ing entombment or to erosion prior to dis- 
covery, and this element is well preserved 
in only a few cases. The iHac blade is 
moderately high, as in sphenacodonts and 
other edaphosaurians, and in contrast to 
ophiacodonts, and agrees with other eda- 
phosaurians in that ( in contrast with sphen- 
acodonts) there is little posterior elonga- 
tion. There is a narrow but well-defined 
longitudinal area for muscle attachment 
at the top of the inner surface, above the 
areas for the sacral ribs. The first sacral 
appears to have been in contact with a de- 
pressed area at the anterior margin of the 
inner surface of the blade, the second pre- 
sumably apposed to a flat area posterior to 
this; still farther posteriorly, a well-marked 
internal depression received the small distal 
end of the third sacral. 

The acetabular region is of a typical pely- 
cosaurian nature, with the usual primitive 
dorsal buttress. In most specimens sutures 
between the three peKic elements are not 
clearly seen; in one specimen part of the 
sutures could be made out on the external 
surface, and in several instances lines of 
striae on the inner surface indicated the 

16 BuUf'lin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 8. Right pelvic girdle in dorsal and lateral views, X '/3- 

areas of fusion of ilium with pubis and 

The puboischiadic plate is of very large 
size, with an anteroposterior length in 4-0-4, 
for example, of 380 mm, a depth below and 
internal to the base of the ilium of 112 mm 
and a breadth of the pubis, measured at 

right angles from the front end of the pubic 
symphysis, of 173 mm. There is but a slight 
indication in the material of the develop- 
ment of a pubic tubercle, such as is found 
in Edaphosaurus, Nitosatinis, and Casea. 
The great development of the puboischiadic 
plate is comparable to the type of structure 

CoTYLORHYNCHUS SKELETON • StovciU, Piicc, and Roiuer 17 

seen in Edaphosaiirus and Casea, and is and well preserved in a single specimen, 
associated with bodily breadth, present in The pose was obviously that common to all 
the pelvic region as well as farther forward, pelycosaurs, with humerus and femur pro- 
The plate was strongly tilted outward at jecting nearly straight outward horizontally, 
somewhat more than a 45° angle, so that it and with the lower limb segment essentially 
is not seen to any great degree in side view, vertical in position. Front and hind legs 
Internally, the puboischiadic plate ex- appear to be nearly equal in length, but 
hibits, as in pelycosaurs generally, a gently with the humerus slightly shorter in over- 
hollowed out area, in which the bone is all length than the femur. In all pely- 
relatively thin, along the middle third of its cosaurs the lower limb is much shorter than 
length. Anterior and posterior to this, on the proximal segment, but this relative 
pubis and ischium respectively, thickened shortness is very marked in Cofylorhyn- 
bony areas— essentially supporting struts— chits. Here radius and tibia are only ap- 
descend from the iliac region to the sym- proximately 60 per cent as long as humerus 
physis. The presence of these thickened and femur, respectively. In Ophiacodon the 
areas is reflected in the symphysis, which radius is about 77 per cent of the humeral 
is thickened in both pubic and ischiadic length, the tibia about 83 per cent of the 
regions, with a relatively thin intermediate length of the femur; in Dimetrodon the 
zone. Anterior to the ridge descending the comparable figures are 82 and 83 per cent, 
pubis is a very broad area of origin for The C otijlorhynchm proportions, however, 
puboischiofemoralis internus, which is but are comparable to those in other edapho- 
slightly turned outward from the general sauroids; in Edaphosaiirus hoanerges we 
plane of the internal surface of the plate, find figures of 62 and 57 per cent. Like 
Within this area is the usual internal open- other edaphosauroids, Cotijlorhynchus was 
ing of the obturator foramen which opens "low-slung." 

externally below the pubic portion of the Humerus (Fig. 9). The humerus is of 

acetabulum. There is a slight indication the primitive tetrahedral type, although the 

in certain specimens of a small area along shaft is stouter than in typical (and smaller) 

the dorsal margin of the ischium for an pelycosaurs. As figured, the "twist" of the 

ischiotrochantericus origin. ends on one another is extreme; this, how- 
ever, appears to be due to crushing of the 

LIMB ELEMENTS (none too well preserved) specimens upon 

In almost all instances the limb elements ^T^^^^'^' *!^^ t^^^'"?," '' l^^'^^^; specimen 4-0-3 

1 1 1 J J ri j.i. 11 snows the twist to nave been one or ap- 

nave been crushed and flattened dorsoven- . ^ , ^.^o * • i i . i 

.11 i.1 ,. ^1 ..11 1. proximatelv 90 . As is sphenacodonts and 

trally, so that they are essentially two-di- i i 'a n ^.u • i 

. , . . , ^ edaphosauroids generally, the proximal ar- 

mensional-a situation making correct re- ^^^^^i^^. ^^^^.^^^^ ^^^^^^^ posteriorly well 

construction difficult. ^^^^^^ ^^^^^ ^^^ ^^^^^^^j ^^^^f^^^ ^f ^j^^ ^^^^ 

The major elements differ markedly from j^ ^^^ ^erv considerable proximo-distal 

those of httle Casea, and to a lesser extent breadth the entepicondyle is edaphosauroid. 

from those of Edaphosaiirus and Lupeosau- The entepicondylar foramen appears to 

rus, in the fact that they are very stoutly have been unusually large, 

built. This is, of course, a feature associated The region of the ectepicondyle and the 

with the large size and great weight of supinator process is imperfect in most speci- 

Cotylorhynchus. In most regards, however, mens. In small specimens, such as University 

they are not only typically pelycosaurian but of Oklahoma specimens bearing the num- 

show a number of distinctive edaphosauroid bers 4-0-3 and 2.3-38, in which the distal end 

features. In few instances do we find all, or is well preserved, the supinator process is 

most, of the major Hmb elements present broad and close to the ectepicondyle but 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 9. Right humerus, viewed in dorsal and ventral aspects in the plane of the distal end, X 'A- 

separated from it by the noteh typical of 
pelycosaurs. Tliis separation, however, is 
obviously a growth stage only, for in 4-0-16 
and a further University of Oklahoma speci- 
men of uncertain number the notch is closed 
and an ectepicondylar foramen present. Tlie 
foramen is otherwise found in pelycosaurs 
only in Edaphosaurus, and it is stated to be 
absent in Casca. In Caseci, however, the 
gap between the tip of the supinator proc- 
ess and the ectepicondyle is small, and 
may well have been bridged in cartilage. 
It is obvious that the ectepicondylar fora- 
men has developed more than once, in 
parallel fashion, in early reptiles; its pres- 
ence here may perhaps be correlated with 
changes in limb mechanics and muscula- 
ture, due to increased size, rather than attrib- 
uted to inheritance from a basal edapho- 
sauroid ancestor. 

Radiu.s (Fig. 10). The radius is pre- 
served (although not too well preserved) 
in several instances. As noted above, it is 

short, with a length rather less than three- 
fifths that of the humerus. Although the 
effect is in all specimens accentuated by 
crushing, the bone was obviously relatively 
thin dorsoventrally, as in pelycosaurs gener- 
ally. As in the case of other limb bones of 
Cotylorliynchus, the radius is broad in pro- 
portion to its length. 

Tlie proximal articular surface, where 
preserved, has the appearance of an oval, 
thin dorsoventrally; presumably it was sub- 
circular in life. The dorsal (extensor) sur- 
face of the shaft is convex in section; the 
ventral surface apparently was flattened. A 
rugose area for ligament or muscle attach- 
ment is visible on the lateral edge of the 
dorsal surface just below the head of the 
bone. From the head the bone (when un- 
crushed) constricts to a somewhat thinner 
shaft. A short distance below the head, 
however, a ridge arises on the medial sur- 
face of the bone. Proximally it begins some- 
what toward the ventral surface; it ex- 

COTYLORHYNCHUS SKELETON • Stovdll, Price, and Romer 19 

Fig. 10. Left, left ulna in extensor and flexor aspects. Right, comparable views of tfie radius. Right, below, proximal 
and distal surfaces of radius (dorsal aspect above). X '/s- 

panels, however, to attain the lateral margin 
and nuis do\\ai\vard mueh of the length 
of the bone. Distally, the ridge is absorbed 
in a distal expansion of the bone leading 
toward the temiinal articulation. As in 
pelycosaurs generally, the distal end of the 
bone is somewhat curved ventrally, so that 
the oval distal articular surface (for the 
radiale) faces somewhat ventrally as well 
as distally. The lateral margin of the shaft 
is also rather thin, but there is no develop- 
ment of a projecting ridge. Medioventrally 
there is an abiaipt out-turning of the lateral 
margin above the articular surface. 

Ulna (Fig. 10). The olecranon appears 
to have become well developed at a rela- 
tively early stage of growth, for it is nearly 
complete ( although ^^'ith a small unfinished 
terminal surface) in University of Okla- 
homa specimen N-7-37, a small specimen. 
As preserved, the head of the ulna is thin 
where seen, but this is presumably an effect 

due to crushing; very probably the head 
in life was as thick as in Edaphosaurus. As 
in the case of the radius, the bone is very 
short compared with the humerus. It is, 
further, exceedingly broad; the breadth of 
the distal end in one complete specimen 
measures about 40 per cent of the length of 
the bone as measured from the lower mar- 
gin of the sigmoid notch; and the proximal 
width, across the notch, is about 50 per cent 
of the length. These figures are far in ex- 
cess of those of other groups, in which the 
highest figures available to me are 29 per 
cent and 39 per cent for a specimen of 

Femur (Fig. 11). The femur is typically 
edaphosaurian in nature, closely compar- 
able in every major way to a well-preser\'ed 
Lupcosaurus femur in the Harvard collec- 
tion and likewise comparable, except for its 
stouter build, to the femora of Casea and 
Edaphosaurus. As in other edaphosaurs. 


Bulletin Museum of Comparative Zoology, Vol. 135, Xo. 1 


Right femur, In ventral and dorsa! views; at right, proximal and distal viev/s {dorsal aspect above). X /3- 

the curvature of the shaft characteristic of 
sphenacodonts is absent. Particularly char- 
acteristic is the ventral trochanteric system. 
There is a well-developed internal tro- 
chanter from which, in contrast to sphena- 
codonts and ophiacodonts, a ridge descends 
the under side of the shaft diagonally to- 
ward the external condyle. There is little 
indication of a distinct fourth trochanter 
along this ridge, and likewise little develop- 
ment of the posterior proximal branch of 
the Y-shaped ridge system, the intertro- 
chanteric fossa thus being shallow posteri- 
orly. In sphenacodonts, and to a lesser 
degree in some ophiacodonts, the proximal 
articular surface extends along the proximal 
portion of the posterior margin of the shaft; 
here, as in other edaphosaurs, this surface 
is confined to the proximal end of the bone. 
Proximally, on the dorsal surface of the 

shaft, there is sometimes seen, toward the 
posterior margin, a rugose area for muscle 
attachment. Unique is the presence at the 
anterior margin of a very distinct rugose 
ridge, about 25 mm long, likewise presum- 
ably for muscle attachment; this is clearly 
seen in two specimens. The external condyle, 
as in other edaphosauroids, projects very 
markedly beyond the internal (medial) 
one, and in a well developed specimen the 
tip of this condyle markedly overhangs the 
articular surface for the tibia below it, as 
it does in other edaphosaiuoid femora in 
which ossification is well advanced. 

Tibia (Fig. 12). The tibia is, as noted 
earlier, relatively short, with a length only 
three-fifths or less that of the femur. As 
with other limb bones, the tibia is very 
broad, notably its head. The width of the 
head in one specimen is about 57 per cent 


Fig. 12. Lefl, extensor aspect of right tibio; center, extensor and flexor aspects of right fibula; right, above, proximal 
and distal surfaces of tibia, and below, distal surface of fibula, extensor surface above. X '/s- 

the length of the bone, and the distal width 
nearly 33 per cent of the length. The closest 
approach to these proportions is in Edapho- 
saunis, in which these two figures approxi- 
mate 50 per cent and 30 per cent. In all 
other kno\vn pelycosaurs the figures are 
much lower — Dimetrodon limbatiis, for ex- 
ample, gi\'ing figures of 38 per cent and 22 
per cent, Ophiocodon 37 per cent and 26 
per cent. The two articular surfaces of the 
head are distinctly separated and set off 
from one another at a considerable angle. 
As in other ophiacodonts, the lateral femoral 
articular area is relatively narrow dorso- 
ventrally. The cnemial crest is little de- 
veloped. As in Edaplwsaurus, and in con- 
trast with most other non-edaphosaurian 
pelycosaurs, the distal articular surface 
curves strongly toward the lateral side of 
the bone. 

Fibula (Fig. 12). As in the case of the 
tibia, the fibula is very short as compared 
with the femur, and is very broad distally. 
Tlie mean breadth here in three specimens 
is 38 per cent of the length. This figure is 
comparable in Edaphosoiirus, but pely- 
cosaurs generally have a much slimmer 

fibula, with distal widths in Dimetrodon 
and Ophiacodon, for example, of 20 per 
cent and 29 per cent of the length. 


Mantis (Figs. 13, 14). As noted above, 
the specimens of Cotylorhynchus are not 
infrequently found in articulated fashion, 
and the feet are sometimes well preserved. 
We may note, for example, well preserved 
front feet in 4-0-1, 4-0-6 and 4-1-S2, hind 
feet in 4-0-10, 4-0-2 (1249), and both front 
and hind feet in the Chicago skeleton. Be- 
tween the various specimens nearly all fea- 
tures of carpus, tarsus and digits are seen. 
As in the case of the major limb bones, the 
feet are broad and short, and thus differ 
considerably at first glance from those of 
most pelycosaurs. Study, however, shows 
that, apart from questions of proportions 
related to the size of the animals the feet 
are typically pelycosaurian and, despite the 
contrast in shape, resemble closely those 
of Casea. 

The manus has the usual pelycosaur ele- 
ments, including a pisiforme, two centralia 
and a series of five distal carpals. Tlie 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

Fig. 13. Left, left manus of No. 4-0-6; right, restored left menus. X 'A- Abbreviations for Figs. 13-16: o, astragalus; c, 
centralia; col, calccneum; F, fibula; i, intermedium; p, pisiforme; R, radius; r, radiale; T, tibia; U, ulna; u, ulnare; 1-5, 
distal carpals or tarsals; /-V, digits. 

radiale has a very deep, essentially square, 
proximal articular surface for the foot of 
the radius. The intermedium is short and 
broad, with well developed processes on 
both radial and ulnar sides at mid-height, 
and a broad proximal articular surface for 
the ulna. The ulnare is far shorter than in 
most pelycosaurs, but comparable in pro- 
portions to that of Casea. The proximal end 
is much less convex in outline than in most 
pelycosaurs; it forms a nearly continuous 
articular surface for the very broad ulna; 
this surface extends medially from a con- 
tact with the intermedium to a lateral facet 
for the pisiforme. Tliis last element, as 
usual, is a thin plate. In 4-0-6 the bone is 
curved sharply toward the ventral surface 
at its outer margin; this may, however, be 
an effect of crushing. The medial or proxi- 
mal centrale is again relatively short. The 
usual arterial gap is present between proxi- 
mal centrale, intermedium and ulnare. The 
lateral centrale is not too well preserved; it 
is, as in pelycosaurs generally, a small ele- 
ment when viewed from the exterior sur- 

face, short proximodistally, broad mediola- 
terally. The articulated feet suggest that it 
was placed well in toward the center of the 
manus, with an unossified gap between 
radiale and distal carpal 1. 

Distal cai-pal 1 appears to be essentially 
a simple rectangle in dorsal outline, short 
proximodistally but broad mediolaterally, 
covering the entire width of the head of 
metacarpal I. Element 2 is longer but less 
broad, its width less than the overall width 
of its metacarpal; its lateral border is 
straight, its proximal and medial borders 
a continuous curve. Element 3 is about 
as broad as 2, but longer proximodistally. 
As in pelycosaurs generally, 4 is by far the 
largest of the distal series, with a width 
double that of element 3 and a somewhat 
greater length proximodistally. Its proxi- 
mal end has, as in pelycosaurs generally, 
two articular faces at somewhat of an angle 
to each other, a laterally tilted surface for 
apposition to the ulnare, a shorter medially 
tilted face for the proximal centrale. Distally 
the bone articulates broadlv with the ex- 

COTYLORHYNCHUS SKELETON • StovdU, Piicc, and Romer 23 

Fig. 14. leh, left manus of No. 4-1-S2; nghl, left pes of No. 4-0-10. X '/j. Abbreviations as in Fig. 13. 

panded head of metacarpal IV. Element 5 
is narrow proximodistally but is expanded 
mediolaterally to meet the entire breadth of 
metacarpal V. Proximomedial and proxi- 
molateral surfaces meet element 4 and the 
ulnare, respectively. 

In the metapodials (and in the phalanges) 
the shortness and breadth of the elements 
and the almost complete absence of a dis- 
tinct shaft region give the foot a clumsy ap- 
pearance. There is a steady increase in 
length from metacarpal I to metacarpal IV, 
and metacarpal V is, exceptionally, some- 
what longer still. Metacarpal I appears to 
have a broad, flat head; in metacarpals II 
and III the proximal articular surface is a 
concave area not occupying the full width 
of the bone. In metacarpal IV the proximal 
articulation is a concavity, but a very broad 
one. In metacarpals II and III there is a 
pronounced expansion of the head toward 
the lateral side, and in IV this lateral ex- 
tension is very pronounced. In metapodial 
V the proximal articulation does not appear 
to be cupped; it is tilted so that the outer 

margin is much more proximal than the 

The phalanges, hke the metapodials, are 
short and massive in appearance. The na- 
ture and structure of the articulations of the 
elements is in general of a typically pely- 
cosaurian type (Romer and Price, 1940: 
167-169). From the massive build of the 
feet and the presumed herbivorous habits 
of the animal, one would expect the toes to 
terminate, like those of diadectids or parei- 
asaurs, in hoof-like structures. Instead, 
however, there are (as in Casea) long and 
powerful bony claw supports, suggesting 
that in life the animal did considerable dig- 
ging for its food supply. 

Pelycosaurs, in which good articulated 
feet are known, typically have a phalangeal 
fomiula in the manus of 2-3-4-5-3.^ Coty- 
lorhynchus, as is definitely proven by the 
material, has the surprisingly low formula 
of 2-2-3-3-2 — even lower than in typical 
therapsids and rivalled for reduction among 

^ Edaphosatinis, formerly in doubt, is now known 
to have this formula also. 

24 Bulletin Museum of Comparative Zoology, Vol 135, No. 1 

Fig. 15. Dorsal and ventral views of right pes of CNHM 272; tfie clav/ed toes are strongly flexed and bent under the 
tarsus. X 'A- Abbreviations as in Fig. 13. 

Paleozoic reptiles only by the pareiasaurs. 
In Williston's material of Casca, the manus 
was nearly complete bvit for the most part 
disarticulated. Williston, not expecting re- 
duction, utilized the material available to 
give a manus with the typical reptilian for- 
mula (1911: fig. 13). To do this he was forced 
to assume that a number of elements were 
missing from the toes. However, new ma- 
terial described by Olson (1954) shows that 
the formula of the manus was 2-3-3-4-3 — a 
definite reduction, although not as marked 
as in its giant relative. The series of ele- 
ments present in Williston's specimen was, 
thus, actuallv nearly complete. 

Pes (Figs. 14, 15, 16). Although the 
material of the hind foot is not as good as 
that of the manus, nearly all the structure 
can be clearly made out. There is a series 
of typical tarsal elements — astragalus and 
calcaneum proximally, and five distal tar- 
sals; there is a lateral centrale, but whether 
a small medial centrale was present is un- 
certain. The proximal tarsal elements are 
relatively short, as compared with those 
of most other pelycosaurs except for the 
ophiacodonts; this presumably in relation to 
ponderous build. The astragalus appears 
to have had a relatively flat facet for the 
tibia. There is a typical arterial notch be- 
tween artragalus and calcaneum. 

As in the manus, the reduced phalangeal 
formula of 2-2-3-3-2 was present. Williston 

attempted to restore the foot of Casea with 
the primitive formula. But, as his descrip- 
tion suggests, little of the material was actu- 
ally articulated and it seems reasonable to 
believe that Casea had a reduced pha- 
langeal foiTuula in the pes similar to that in 
the manus; if so, Williston's foot material 
was nearly complete. 


In agreement, it would seem, with the 
fact that in the Edaphosauria generally the 
gastralia are little developed, no trace of 
abdominal ribs was discovered during prep- 
aration of the materials except in one in- 
stance. Here there were found numerous 
slender elements with tapering ends. The 
maximum length as preserved was 65 mm; 
the widths 3 to 5 mm. Tlie material was 
not sufficient to determine their arrange- 
ment, although they were presumably ar- 
rayed in the usual series of V-shaped seg- 
ments along the course of the abdomen. 


A lateral view of a restoration is shown 
in Figure 17, based on the series of larger 
specimens. The general appearance is com- 
parable to that sho\Mi in restorations of its 
smaller relative, Caseo (Williston, 1911: 
frontispiece; Romer and Price, 1940: fig. 
71), except for the somewhat more massive 
build of Cotylorhynchus associated with its 

CoTYLORHYNCHUS SKELETON • Stovdll, Price, and Ronier 25 

Fig. 16. Restored left pes, X Vj- Abbreviations as in Fig. 

larger size, and the absurdly small size of 
the head. The lateral view does not, of 
course, give proper emphasis to the great 
breadth of the barrel-like trunk, commented 
on previously, and well shown in Wil- 
liston's photograph of the mounted Casea. 
The build of CotyloihyncJuis is in agree- 
ment with the portrait of a generalized 
edaphosaurian given by Romer and Price 
(1940: 377): 

"We find a tiny head armed \\ith a powerful 
battery of blunt teeth, the trunk a large, broadly 
rounded barrel, the legs spread out broadly, but 
the lower segments so short that the belly cannot 
have been far clear of the ground. Such an ani- 
mal was obviously not a carnivore, and, in the 
discussion of habits, we have already cited data 
supporting Williston's belief that these reptiles 
were herbivores. The enormous storage capacity of 
the abdomen further suggests that the food was 
probably of a bulky, watery nature, low in nutri- 
tive value, so that it was necessary for the animal to 
ingest large quantities. The curiously small head of 
the advanced edaphosaurs is matched among later 
plant-eating reptiles by the sauropods, whose food 
may have been of a comparable type." 

Cotijlorhyncluis romeri exceeds in bulk 
any of the known pelycosaurs from the 
typical Texas Wichita and Clear Fork red- 
beds deposits, and is exceeded only by its 
presumed descendant, C. hancocki from the 









26 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

San Angelo (Olson and Beerbower, 1953; 
Olson, 1962: 28-45). In the stndy of pely- 
cosauis generally, Romer ( Romer and Price, 
1940; Romer, 1948) used as an index to 
relative size — and \\eight — an "orthometric 
linear unit" based on the dimensions of 
dorsal vertebrae. At the time of publication 
of the "Review of the Pelycosauria" such 
infonnation as was then available concern- 
ing C. romer i suggested that this unit 
was approximately 8.32, and this appears 
still to be a reasonable figure. Closest to 
Cotylorliynchu.s romeri among Wichita and 
Clear Fork pelycosaurs were the large ter- 
minal member of the Ophiacodon phylum, 
O. major., with an orthometric linear unit of 
7.37, and tiie large tenninal Clear Fork 
Dimcirodon grandis, at 7.61. Since the 
Casea unit is but 3.30, the average linear 
measurements of elements of Cotylorhyn- 
chus should be approximately two and one- 
half times that of Casea, and the weight 
more than 15 times as great. With an esti- 
mated weight of about 331 kg — roughly 
about one-third of a ton — Cotylorhynehu.s 
was the giant of its times. 


When Casea was first described it oc- 
cupied an isolated position among pely- 
cosaurs — so isolated, indeed, that Watson 
(1917: 173) suggested that it was not a 
pelycosaur at all. Possibly Tricha.murus (cf. 
Romer and Price, 1940: 422-423) is a rela- 
tive, but until the discovery of Cotylo- 
rhynchus no further members of the family 
Caseidae were recognized. In recent years, 
however, Olson (1962: 24-47; and earHer 
papers ) has added a number of new fonns 
to the group from the middle and upper 
Clear Fork fonnations and the lower part 
of the Pease River group, including further 
species of Casea and Cotylorhynchiis, and 
the new genera Caseoides, Ca.^eo))sis, and 
An<ie}osaurus; further, the caseids are now 
known to have ranged widely, for Ennato- 
sauriis of the Russian Kazanian is clearly a 
caseid (as is possibly the poorly known 
Phreatophasma) . 

The caseids are the last major group of 
pelycosaurs to appear in the geological 
record. Casea broilii, the earliest acknowl- 
edged member of tlie family, only appears 
at about the Arroyo-Vale boundary in the 
Clear Fork group; other forms occur in the 
later Clear Fork formations and the roughly 
equivalent Hennessey of Oklahoma, or the 
still later Pease River group of Texas and 
the Russian Kazanian. Although some mil- 
lions of years must be allowed for the de- 
velopment of caseid specializations, it is 
quite possible for the family to have evolved 
from some more generalized group during 
Wichita and early Clear Fork times. Until 
recently clues as to caseid ancestry were 
few. Tricha.satirus of the Arroyo Forma- 
tion and Glaucosaiirus of the Clyde Forma- 
tion of Texas were suggested by Romer and 
Price (1940: 421-423) as presumed eda- 
phosauroids possibly related to the caseids. 
In the first-named genus the skull is un- 
known.^ The second is represented only 
by a single small skull which is extremely 
short-faced and with an isodont dentition; 
this suggests possibilities of relationship to 
the caseid pedigree. As to a more remote 
ancestry, it was suggested by Romer 
(Romer, 1937; Romer and Price, 1940: 405- 
412) that Mycterosaurus and Nito.saiiriis, 
small early Permian pelycosaurs which 
seemed to be primitive in many ways but 
showed definite edaphosaurian characters 
in the postcranial skeleton, might represent 
the ancestral stock of the caseids and per- 
haps of the edaphosaurians as a whole. 

In recent years several new finds have 
added somewhat to the picture. Vaughn 
(1958) has described as Colobomycter an 
imperfect skull from the Fort Sill quarry 
(an Arroyo equivalent in Oklahoma), and 
considers, reasonably, that its characters in- 
dicate that "it provides a good structural if 
not an actual ancestor for the family Casei- 
dae." Fox (1962) has described as Delo- 

1 A toothplate provisionally referred to this genus 
(Romer and Price 1940: 423) is now known to per- 
tain to the cotylosaur Labidosaurikos. 

COTYLORHYNCHUS SKELETON • Stovull, Piicc, and RoTTier 27 

rhynchus three isolated maxillae from this saurus had more teeth than Oedaleops. In 

same quarry; the bone itself is caseid-like, the latter genus the dentary is not known 

but the dentition is primiti\e and hence from associated material. In the two max- 

Fox classes it amongst the presumably an- illae of Oedaleops found by Langston, the 

cestral Nitosauridae rather than placing it tooth counts appear to be IS andl6; in the 

in the Caseidae. As noted by Langston incomplete Nifosaunis maxilla 15 teeth and 

(1965) there is little to distinguish Delo- alveoli are present, and the total count was 

rhynchus from its quarr>'-mate Colobomyc- probably about 18. (4) There appears to 

ter. Langston (1965) has recently described be no significant difference in the height of 

as Oedaleops a small pelycosaur from the the maxilla bet^^'een Oedaleops and Nito- 

New Mexican Permian represented by two .sa»;f/.s— particularly if the ob\'ious crushing 

skulls and other fragmentary material. As undergone by the Oedaleops skull be taken 

Langston points out, the Oedaleops skull into account. 

is of a type quite surely expected in a caseid The one possibly valid generic distinction 

ancestor, but differs from proper members lies in the greater isodontv of the maxillary 

of that family in that, for example, the face dentition in Oedaleops. In the type there is 

is not as abbreviated and, most especially, a modest development of a "canine" pair 

the dentition is primitive; as proper for a at maxillary positions 2 and 3; in a second 

primitive pelycosaur of any sort, the teeth specimen the third tooth is large; in the 

are sharp-pointed and somewhat recurved A' /fo.sfl«/f/5 maxilla a maximum is not gained 

and \\ith a modest development of a maxi- until we reach teeth 4 and 5. Considering 

mum tooth size in the canine region, in one the constant tooth replacement characteristic 

specimen, at least. As possibly attributable of reptiles and the consequent continual 

to Oedaleops, Langston describes a number changes in the aspect of a dentition, this one 

of small postcranial elements found in the feature seems hardly safe ground for generic 

same quarry. Of these, the ilium, as Langs- distinction. ^ 

ton notes, is of a very primitive type, cer- Although the material of the genera dis- 
tainly not expected in a pre-caseid. Other cussed above is quite incomplete, in most 
elements, most notably the scapulocoracoid instances, it appears that in this we have 
(lacking, significantly, the supraglenoid at least the beginnings of a phyletic series 
foramen), are comparable to those of Nito- leading from such a primitive but edapho- 
saiinis and caseids. They may well pertain sauroid pelycosaur as Myctewsauriis up- 
to A itosaurus. ward toward the caseid condition. A com- 
Moreover, may not Oedaleops and Nito- plicating and confusing factor in the sit- 
saitnis, contemporaries from the same region nation, however, was introduced by Watson 
and horizon, be identical? Of Oedaleops (1954: 356) with his suggestion that £of/?{/m 
we ha\'e no certain knowledge of postcranial might be related to caseid ancestry. Both 
material; of N itosaurus we have no skull Vaughn and Langston have adopted this 
material except of maxilla. Langston briefly point of view, placing such forms as Col- 
mentions this possibility but says that Nito- obomycter and Oedaleops in the Eothv- 
saurus had longer jaws, a more slender j-ididae, and Langston goes to the extreme 
dentary, considerably more teeth (sub- ^f excluding the Nitosauridae from anv re- 
isodont m fonii) and a higher maxillary lationship to the Caseidae, despite the num- 
bone. But 1 ) we do not have a complete ^ i i • •£• ' ^ . , 
• „.  v/ /T\ .1 11 erous and surelv significant postcranial re- 
aw in A /fo.rai/n/.s; (2) the seeming slender- i, u ^ u i_ 

r .1 TVT-x 1 . , 1 , semblances between the two. 

ness of the N itosaurus dentary is probablv t^i i . r j • r- ^7 • 

1^.1 r i.u ^i- • 1  / r ^ he basic reason tor considering Eothiiris 

due to loss of the thm lower margin (cf. 

Romer and Price: fig. 70, and Langston: 1 a \. a- ^ a,.- v .. n • t 

r. ^., ° ' ° '■A. subordinal distinction, actually, in Langston s 

tig. 2a ) ; ( 3 ) there is no evidence that Nito- chart, page 43. 

28 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

as a possible relative of the caseids is, of 
course, the fact that Eothyris, like the 
caseids, is remarkably short-faced. This in 
itself is no more a valid reason for associat- 
ing them than would be the association of 
the sphenacodont Secodontosaiirns with the 
ophiacodont Varanosaurus because they 
are both extremely long-snouted. If the 
Eothyris-cAseid relationship is to be sub- 
stantiated, more positive reasons must be 

Langston (1965: 21) cites fourteen points 
in which Oedaleops and Eothyris are in 
partial or complete agreement. As he says, 
this seems to be, at first sight, an imposing 
list. Included, of course, is the fact that the 
face is short, not necessarily meaningful, 
and the fact, of no systematic value, that 
both are small. Correlated with small size, 
and hence likewise without other necessary 
significance, is the relatively large size of 
the orbits and of the pineal foramen. A 
number of other common features are such 
as are liable to be present in any relatively 
primitive pelycosaur, including: (1) rela- 
tively flat skull; (2) jaw articulation on a 
level with the toothrow, as in the Ophia- 
codontia, the primitive sphenacodont Var- 
anops, and Mycterosoiirus; (3) outward 
slope of cheek plates (contrasting with 
sphenacodonts ) ; ( 4 ) a primitive long lacri- 
mal; (5) nonnal relationship of roofing 
bones; (6) an unusually large supratemporal; 
(7) a long, tapering postorbital (as, for ex- 
ample, in the ophiacodont Varanosaurus, 
and the sphenacodont Varanops); (8) 
some indications in the tabular-supratem- 
poral region of the otic notch of ancestral 
types; (9) a differentiated dentition, as in 
most pelycosaurs except Edaphosaurus and 

The two genera, thus, are short-faced, 
small in size, and have both retained various 
primitive characters. Little remains of the 
original fourteen points which can be con- 
strued as positive indication of relationship. 
Langston cites "relative position of orbits 
and pineal opening," but there does not 
appear to be any unusual condition fiere in 

either case. With regard to "enlarged nares 
and obtuse rostrum," the nares in Eothyris 
do not appear to be any larger, propor- 
tionately, than in many other pelycosaurs, 
and the rostrum does not appear to be any 
more obtuse than in pelycosaurs generally. 
The Eothyris prefrontals are said to be "in- 
flated" (i.e., somewhat expanded), but this 
does not seem significant. 

There is thus little positive reason to 
associate Eothyris with caseid ancestry, and 
one very strong objection — the dentition. 
In all early pelycosaurs — indeed, in nearly 
all primitive tetrapods generally — there is a 
trend for the development of somewhat 
enlarged teeth near the front of the maxilla 
as an incipient "canine" region. In sphena- 
codonts and their therapsid descendants 
this trend is accentuated; in edaphosaurians 
— both Edaphosaurus and the caseids — 
there is an opposite trend toward isodonty. 
In such forms as Oedaleops there is but a 
mild, essentially primitive, development of 
a canine "maximum" such as might be ex- 
pected in any relatively primitive pely- 
cosaurs. In Eothyris, on the other hand, we 
find the greatest exaggeration of canine tusks 
to be found in any pelycosaur. In their 
dentition, Eothyris and the caseids have 
evolved in such diametrically opposite 
directions that it is difficult to believe that 
they are at all closely related. 

But even if (as is not too probably the 
case) Eotliyris should prove to be related to 
the caseids, it is dangerous to base hypoth- 
eses of broader relationships on the "family 
Eothyrididae," as has been done by Langs- 
ton and, to a lesser degree, by Vaughn. 
As I have pointed out (Romer and Price, 
1940: 247; Romer, 1956: 676), this family 
is a purely provisional one, set up to receive 
fonus, presumably highly predaceous, which 
have marked canine development but are 
not members of the Sphenacodontia. Any 
unity the group might have is based on 
this dental feature — which is, of course, 
the one point in which all of them notably 
differ from the trend toward isodonty ex- 
pected in caseid ancestors. There is no 

COTYLORHYNCHUS SKELETON • Stovoll Pricc, and Romer 29 

evidence that any form assigned to this 
family, other than Eofhyiis, was short-faced. 
Such postcranial material as can be as- 
sociated with any of the genera included 
in this provisional family lacks any features 
indicative of caseid relationships, and such 
indications of systematic position as are 
shown by Stcrcorhachis, StcrcophaUodon, 
and Bcddwinonus suggest the Ophiacodon- 
tia. With regard to Eothyris, the lack of 
postcranial data is a stumbling block. 

On the assumption that the Ophiacodontia 
represent the basal stock of the Pelyco- 
sauria, it is a reasonable assumption that 
the caseids are of ultimate ophiacodontian 
derivation; but that any of the "eothyrids" 
are connecting links is very dubious. 

Romer and Price (1940: 366-376) pointed 
out numerous skeletal features which Eda- 
phosaiiriis and Casea, as two extremes, have 
in common, and advocated their being 
placed in a common suborder. There is, 
however, a seeming difficulty in assuming a 
common ancestry, because of chronological 
factors. The caseids seem quite surely to 
have specialized from primitive ancestors 
at a late date, for no form attributable to 
this stock is known earlier than fairly early 
Wichita times; on the other hand, Edapho- 
saunis had already evolved by the late 
Pennsylvanian. If both Edaphosourm and 
the caseids evolved from essentially primi- 
tive edaphosaurians, such as the nitosaurs, 
this must have been, as Langston (1965: 58) 
notes, a very bradytelic group, and nitosaurs 
should have been in existence in the Penn- 

This appears to have been the case. It 
seems highly probable that Pctrohcosaums, 
from the Pennsylvanian Garnett shales of 
Kansas (Lane, 1945; Peabody, 1949, 1952), 
is a primitive edaphosauroid. 

Lane, and Peabody at first, concluded 
that this small reptile was a pelycosaur. The 
latter author, however, impressed by cranial 
resemblances to Prolaceita (with which he 
was famihar), argued that Petrolacosaunis 
was an eosuchian — a primitive diapsid. 
There is no morphological proof, one way 

or another, of this suggestion. There was a 
lateral temporal opening, but the cheek is 
poorly preserved, and whether an addi- 
tional upper opening was present cannot be 
determined. Apart from the possible l)ut 
unproven diapsid nature of the temporal 
region, there is no reason to assign Petroh- 
cosaurus to the Eosuchia. Peabody points 
out a number of common features of the 
skulls in Petrolacosaunis and Prohceria, 
but these are essentially primitive characters 
which could have been inherited by both 
from captorhinomorph cotylosaurian an- 
cestors. The time of appearance of Petrola- 
cosmirus is one at which an eosuchian is 
hardly to be expected. Tliere are no traces 
of any diapsid in the Lower Permian, and 
no sure evidence in the Middle Permian; the 
first certain diapsid is Upper Permian in 
age — a full period after Petrolacosaunis — 
and Watson ( 1957 ) has argued that diapsids 
were only then evolving from millerettid 
cotylosaur derivatives. 

If, then, we abandon the possible but 
improbable suggestion that Petrolacosaurus 
was a precocious diapsid, all the features 
of this little reptile agree with the assump- 
tion that we are dealing with a pelycosaur, 
and several characters point strongly to the 
suggestion that it is a primitive edapho- 
saurian. Tlie skull is primitive and gen- 
erahzed, as it is in such archaic ophiacodonts 
as Clepsijdrops and Varanosaurus, such a 
sphenacodontian as Varanops, and such a 
nitosaurian as Mycterosaurus. Diagnostic 
features, however, can be found in the 
postcranial skeleton. (1) The postcervical 
vertebral centra are rounded ventrally as 
in ophiacodonts and edaphosaurs, in con- 
trast to the keeled sphenacodonts. (2) In 
the carpus the ulnare is short, in contrast 
to advanced sphenacodonts and as in ophia- 
codonts and edaphosaurs (however, Var- 
anops among the sphenacodonts also has a 
short ulnare). (3) In the tarsus there is a 
broad, higlily developed, medial centrale 
fonning the sole connection between astrag- 
alus and distal tarsals 1 and 2; edaphosaurs 
and sphenacodonts are similar, but ophia- 

30 Bulletin Museum of Comparative Zoology, Vol. 135, No. 1 

codonts have two small centralia of sub- 
equal size in this position. (4) On the 
femur, the ventral system of trochanters 
and ridges is one found among pelycosaurs 
in edaphosaurs only (cf. Romer and Price, 
1940: fig. 37). Distal to the internal tro- 
chanter an adductor crest slants diagonally 
across the bone towards the external tro- 
chanter. In sphenacodonts there is a pro- 
nounced fourth trochanter but no adductor 
crest; in opliiacodonts the crest descends 
the external margin of the bone. ( 5 ) Both 
Lane and Peabody ascribe to Petrolaco- 
■sauni.s a pelvis (University of Kansas no. 
1425) which is very distinctively edapho- 
saurian. It is of a type strongly contrasting 
wdth that of any other reptilian group — 
most notabh' in the shovel-shaped iliac 
blade, tall but without a marked posterior 
projection. Were this pelvis definitely as- 
sociated, Petwhicosxninis- could be assigned 
to the Edaphosauria without hesitation. It 
was, however, found isolated, and since a 
specimen of Edaphosainus has been found 
in the quarry, this pelvis may pertain to 
that genus. Several pelves with low, long 
iha of ophiacodont type are also present in 
the material; but these, too, lack association, 
and may belong to an ophiacodont, also 
present in the Garnett material. 

There are, thus, in the postcranial skele- 
ton of Petwlacosaiinis a number of features 
which strongly indicate that this genus be- 
longs to a group of archaic edaphosaurians 
from which both Edaphosaurus and, at a 
much later time, the caseids may have 

Preparation of material for this paper 
was aided by a grant (No. GB 500) from 
the National Science Foundation. 


Fox, R. C. 1962. Two new pelycosaurs from the 
Lower Permian of Oklahoma. Univ. Kansas 
Publ. Mus. Nat. Hist., 12: 297-307. 

Lane, H. H. 1945. New mid-Pennsylvanian rep- 
tiles from Kansas. Trans. Kansas Acad. Sci., 
47: 381-396. 

Langston, W. 1965. Oeduleops campi (Rep- 
tilia: Pelycosauria ) . A new genus and species 

from the Lower Permian of New Mexico, and 
the family Eothyrididae. Bull. Texas Mem. 
Mus., 9: 1-46. 

Olson, E. C. 1954. Fauna of the Vale and 
Choza. 7. Pelvcosauria: family Caseidae. 
Fieldiana, Geology, 10: 193-204. 

. 1962. Late Permian terrestrial verte- 
brates, U.S.A. and U.S.S.R. Trans. Amer. 
Philos. Soc, (N.S.) 52 (part 2): 1-196. 

Olson, E. C. and J. R. Beerbower, Jr. 1953. 
The San Angelo Formation, Permian of Texas, 
and its vertebrates. Jour. Geol., 61 : 381-423. 

Peabody, F. E. 1949. Mid-Pennsylvanian pely- 
cosaurs from Kansas. Bull. Geol. Soc. Amer., 
60: 1913. 

. 1952. Pctrolaco.miints kansensis Lane, a 

Pennsylvanian reptile from Kansas. Univ. 
Kansas, Palcont. Contrib. Vertebrata, 1: 1-41. 

Romer, A. S. 19.37. New genera and species of 
pelvcosaurian reptiles. Proc. New England 
Zool. Club, 16: 89-96. 

. 1948. Relative growth in pelvcosaurian 

reptiles. Robert Broom Commemorative Vol- 
ume. Pp. 45-55. 

1956. Osteology of the reptiles. Chi- 

cago, Univ. Chicago Press, xxi -f- 772 pp. 
Romer, A. S. and L. I. Price. 1940. Review of 

the Pelycosauria. Geol. Soc. Amer., Spec. Pap. 

No. 28: 1-538. 
Shuler, E. W. and R. V. Witter. 1942. The 

mounted skeleton of Edaphosaurus boanerges 

Romer at Southern Methodist University. 

Field and Laboratory, 10 (2): 140-144. 
Stovall, J. W. 1937. Cotijlorhynchus romeri, 

a new genus and species of pelvcosaurian 

reptile from Oklahoma. Amer. Jour. Sci., (5) 

34: 308-313. 
Vaughn, P. P. 1958. On a new pelycosaur from 

the Lower Permian of Oklahoma, and on the 

origin of tlie family Caseidae. Jour. Paleont., 

32: 981-991. 
Watson, D. M. S. 1913. The Beaufort Beds of 

the Karroo System of South Africa. Geol. 

Mag., (5) 10: 388-393. 
. 1917. A sketch classification of the 

pre-Jurassic tetrapod \'ertebrates. Proc. Zool. 

Soc. London, 1917: 167-186. 
. 1954. On Bolosaurus and the origin 

and classification of reptiles. Bull. Mus. 

Comp. Zool, 111: 295-449. 

1957. On Millerosaunis and the early 

historv of the sauropsid reptiles. Philos. Trans. 

Roy. Soc. London, (B) 240: 325-400. 
WiLLisTON, S. W. 1910. New Permian reptiles: 

rhachitomous vertebrae. Jour. Geol., 18: 

. 1911. American Permian vertebrates. 

Chicago, Univ. Chicago Press, 145 pp. 
{Received 27 September 1965.) 

SulLetln OF THE 

Museum of 



The Stromateoid Fishes: Systematics and 

a Classification 


M««exmi 'Of" ConipernBtlve Zook>gy, Cambridge, Massachusetts 
- l5 t i tut oi^j_JipiP9io Generole, Universita di Pisa, Italy - . 



JANUARY 27, 1967 





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JoHNSONiA, Department of MoUusks, 1941- 

OcxziAsiONAL Papers on Mollusks, 1945- 

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Bigelow, H. B. and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
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Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In- 
sects. $9.00 cloth. 

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Lyman, C. P. and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- 
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Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 15. (Price list on 
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Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae 
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Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution 
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Proceedings of the New England Zoological Club 1899-1948. ( Complete sets 
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Publications of the Boston Society of Natural History. 

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@ The President and Fellows of Harvard College 1967. 




Historical introduction 31 

Methods 35 

Material 38 

Systematic section 43 

Suborder Stromateoidei 44 

Key to stroniateoid families 52 

Family Centrolophidae 53 

Key to centrolophid genera 54 

Genus Hypcro>iIijphe 54 

Genus Schedoplulus 58 

Genus Centrolopfnis 62 

Genus IcichtJiys 65 

Genus Seriolella 69 

Genus Psenopsis 72 

Family Nomeidae 76 

Key to nomeid genera 77 

Genus Cttbiceps 78 

Genus Nomeus 81 

Genus Psenes 84 

Family Ariommidae 88 

Genus Ariomma 90 

Family Tetragonuridae 94 

Genus Tetragonurus 96 

Family Stromateidae 98 

Key to stromateid genera 99 

Genus Stronuiteus 99 

Genus Peprilus 103 

Genus Pampus 108 

Evolutionary trends in the Stromateoidei 113 

Distribution of the Stromateoidei 122 

Acknowledgments 127 

Summary 128 

Literature cited 129 


The Stromateoidei are a small suborder 
of the perciform fishes, characterized pri- 
marily by toothed saccular outgrowths in 
the gullet immediately behind the last gill 
arch. The stromateoids are all marine, pe- 
lagic, and widely distributed in the tem- 
perate and tropical oceans of the world. 
Most species are rare and infrequently seen, 
but a few form the basis of fisheries. Adult 
stromateoids range from less than a foot to 
over four feet in length. 

^ This paper is based on a thesis presented to 
Harxard University in partial fulfillment of the re- 
quirements for the Ph.D. in Biology. Contribution 
No. 1685 from the Woods Hole Oceanographic 

- Woods Hole Oceanographic Institution, and 
Museum of Comparative Zoology, Harvard Univer- 

Certain stromateoids were recognized in 
classical times. Stromateiis was the name 
applied by the Greeks of Egypt to a fish 
probably from the Red Sea. The name, de- 
rived from the word for a brightly colored 
rug, may have referred to the fish's shape 
and coloration. Later, however, Rondelet 
( 1554 ) used the name for a similar Medi- 
terranean fish known in the contemporary 
Roman vernacular as fiatola. Linnaeus 
(1758) described the same fish as Stro- 
mateus fiatola. 

The oceanic fish pompilus was sacred to 
the Greeks. As pompilm accompanied ships, 
it brought a calm sea (Gesner, 1560). Pom- 
pilus has been equated with Centrolophiis 
(Gunther, 1860), and Gesner's figure 
(1560:113) certainly is of this fish. Thomp- 
son (1947), however, presents evidence that 
pompilus is the pilot fish Naucrates. Cuvier 

Bull. Mus. Comp. Zool., 135(2) : 31-139, January, 1967 31 

32 Bulletin Museum of Cotnpamtive Zoology, Vol. 135, No. 2 

and V^ilenciennes (1833), citing Cetti's 
"Historia Naturale di Sardagni" ( 1777 ) , 
suggested that }wmpilus might be a tuna. 
And while the subject of Ovid's heroic Hnes 

"Tuque comes ratiuin, tractique per aequora 


Qui semper spumas sequeris, pompile, nitentes" 

could well ha\'e been the centrolophid 
Schedophihis ovalis, it seems more likely 
that the poet is referring to Coryphaena, 
the dolphin. The classical name has been 
used in Coryphaena pompilus Linnaeus, 
1758, an unrecognizable fish; in Pompiltis 
Lowe, 1839, a synonym of Ceiitrolophus 
Lacepede, 1803; and in Pompilus Minding, 
1832, a synonym of Naucmtes Rafinesque, 

Identical figures of stromateoids were 
published by the Renaissance compilers 
Belon (1553), Rondelet (1554), Gesner 
(1560), and Aldrovandi (1613). Illustra- 
tions of clearly recognizable species appear 
in these works on the facing page with im- 
possible monsters. Nonetheless, the infor- 
mation gathered by these men was to prove 
very useful to later authors, and was con- 
sidered authoritative by many. Some of 
their work, for example, can be found al- 
most word for word in Cuvier and Valen- 
ciennes ( 1833 ) . 

John Ray's studies (Willughby, 1686) 
are marked by their care and attention to 
detail. His anatomical work revealed for 
the first time the structure most character- 
istic of the stromateoid fishes, the peculiar 
pharyngeal sacs. Ray mistakenly believed 
that the sacs constituted a second stomach. 
Nonetheless, care that was not to reappear 
for several centuries is apparent in his de- 
scription of the sacs of Stromateiis (p. 156) : 

"In palato duo oblonga ossicula aspera . . . 
Nam prime duos habet ventriculos; primum 
retro cor, prope ipsiun os situm, quem echinum 
nun ab re dixeris: carnosus enim est, & apopli- 
ysibus longis, asperis, crebris, pellis erinacei 
fere in modum intus consitur. ( Appendices 
hae in sex radios divaricantur cylindriae supra 
centrum stellae erectae. D. Will.)" 

During the first half of the nineteenth 
century a majority of the stromateoids were 
described and characterized reasonably 
well. The "Regne Animal" (Cuvier, 1817) 
and "Histoire Naturelle des Poissons" (Cu- 
vier and Valenciennes, 1833) were especially 
valuable. Cuvier (1817) provided the basic 
arrangement which was expanded upon in 
the later "Histoire Naturelle." 

Cuvier and Valenciennes (1833:381) 
added to Ray's description of the pharyn- 
geal sacs of Stromateus fiatola: 

"A I'exterieur, cette partie presente la forme 
d'une bourse; les epines dont elle est armee 
sont de different grosseur; les plus grandes 
sont un pen en forme fuseau; les petites garnis- 
sent les intervalles des grandes. Chacune de 
ces epines s'attache a la veloutee par sept ou 
huit ravines ou fibres disposees en etoile." 

They continue later with their own obser- 
vations on the sacs of Stromateus condidu.s 
{— Pampus argenteu.s) (p. 392): 

"Immediatement apres les os pharyngiens 
vient un oesophage en forme de sac ranfle et 
chamu, rond, un peu bilobe . . . garnies . . . 
de grosses epines osseuses . . ." 

and of Rhombus xanthurus ( = Peprilus 
pom) (p. 406): 

". . . un oesophage charnu, arme interieure- 
ment de dents osseuses coniques, les unes plus 
grandes, les autres plus petites . . ." 

Cuvier and Valenciennes not only recog- 
nized this unique structure in "les Stro- 
matees" but also described a similar struc- 
ture in "les Centrolophes." Concerning 
Ccntrolophus pom})ilus (=C. niger) they 
wrote ( p. 339 ) : 

"Le pharynx du centrolophe presente une 
particularite remarquable, qui donne au com- 
mencement de leur oesophage un armure 
puissante. Entre les os pharyngiens . . . I'os 
superieur du quatrieme arceau porte plusiers 
appendices alonges et garnis de dents semb- 
lables ... la partie laterale du pharynx a de 
profondes cannelures osseuses et dentees . . ." 

They continue, observing that this seems to 

". . . quelque analogic avec les epines dont 
le meme cavite est armee dans les stromatees." 

Stromateoid Fishes • Haedrich 33 

Here, for the first time, a relationship be- 
tween the centrolophids and the stromateids 
was indicated. 

Gi.inther recognized a unifying character 
here and, in his Catalogue (1(S60:355), 
noted that in the Stromateina "tooth-hke 
processes extend into the oesophagus." The 
"Catalogue" provided keys to the scombrid 
group Stromateina and to the two genera 
Giinther included in it, Stiomatcus ("ven- 
trals none in an adult state") and Ccntio- 
Jophiis ("ventrals well developed"). The 
other groups in his family Scombridae were 
the Scombrina, Cyttina, Coryphaenina, and 
Nomeina, the last composed largely of 
stromateoids. The diagnostic pharyngeal 
sacs of the Nomeina remained to be dis- 
covered, for they \\ere not mentioned in 
Giinther's account. 

Discussing the limits and arrangement of 
the scombroids. Gill ( 1S62 ) corrected some 
of Giinther's omissions. In doing so, how- 
ever, he l)roke up the convenient group 
Nomeina, and added little to the classifica- 
tion. The Stromateina, though mentioned, 
were not defined. 

"An Introduction to the Study of Fishes" 
(Giinther, 1880) was essentially the same 
as the earlier "Catalogue" in its treatment 
of the stromateoids. Here, however, each 
group was given full family status as Stro- 
mateidae and Nomeidae. The close relation- 
ship of the two was still not indicated, and 
Giinther continued to allocate a number of 
stromateoid genera to other families. 

Although Giinther's (1880) publication 
added almost nothing to the classification, 
it had an important incidental effect. 
Whether the cause was Giinther's failure 
to have noticed Gill's earlier ( 1862 ) paper 
or whether it was his casual accounting 
cannot be said. At any rate, Giinther's treat- 
ment of the stromateoids soon occasioned a 
vitriolic blast from Gill. In his "Notes on 
the Stromateidae," Gill (1884) united the 
fonns scattered by Giinther under the sin- 
gle family Stromateidae, still omitting 
Nomeiis but including, albeit reservedly, 
Psenes and Cuhiceps. The definition of the 

family noted "the gill-rakers of the upper 
segment of the last branchial arch enlarged 
and dentigerous or sacciform, and project- 
ing back\\'ards into the oesophagus" (p. 
665). Gill furthermore recognized a basic 
dichotomy in the group by dividing the 
family into two subfamilies, the Stro- 
mateinae and the Centrolophinae: 

"These are distinguished by differences in 
the development of the vertebrae, the former 
[Stromateinae] having 14-15 abdominal and 
17-21 caudal vertebrae, and the latter [Centro- 
lophinae] 11 alxloniinal and 14 caudal verte- 
lirae; these differences are supplemented by 
variations in the degree of complexity of the 
peculiar appendages representing and homol- 
ogous with the gill-rakers of ordinary fishes, 
developed from the last branchial arch, and 
extending into the oesophagus (p. 654)." 

He also observed that the Centrolophinae 
have normally persistent pelvic fins, while 
those of the Stromateinae are lost with 
growth. Gill considered the Centrolophinae 
to be the most generalized type; the Stro- 
mateinae he thought more specialized. 

"Spolia Atlantica" of Liitken (1880) con- 
tained accounts of the genera Psenes. Cuhi- 
ceps, Stwmoteus, and Schedophihis. The 
discussion of relationships was carefully 
done, and the listing of included species 
was especially good. Unfortunately, the 
work was in Danish, and has apparently 
been little used by subsequent investiga- 

Fordice ( 1884 ) reviewed the American 
species of the Stromateidae. No mention 
was made of the pharyngeal sacs. Only two 
genera, Strotiiateus and Leirus ( = Sclwdo- 
pJiilus) were mentioned, and, again, the 
division was based on the presence or ab- 
sence of pelvic fins. Fordice provided keys 
and neat synonymies of most American 
stromateids. His jiaper was essentially an 
extension of the foundation laid do\\'n by 
Jordan and Gilbert's ( 1882 ) "Synopsis of 
the Fishes of North America," a work which 
erroneously reported (p. 448) for the 
Nomeidae, "No tooth-like processes in the 

Relying heavily on the work of Gill, Jor- 

34 Bulletin Museum of Comparative Zoologij, Vol. 135, No. 2 

clan and (Tilbcrt, and Fordice, Jordan and 
EveiTnann's ( 1896 ) "Fishes of North and 
Middle America" provided a syndiesis of 
current thoughts on stromateoid classifica- 
tion. The Centrolophidae were considered 
a family apart from the Stromateidae, "dif- 
fering in appearance and in the smaller 
number of \'erte]:)rae, although agreeing in 
the possession of teeth in the oesophagus" 
(p. 964). Nonwu.s and Pscnc.s, in the family 
Nomeidae, remained distinct, and no men- 
tion of a relationship \\ ith the stromateids 
was made. 

"Oceanic Ichthyology" ( Goode and Bean, 
1896) drew on Jordan and Gilbert, Gill, 
and Giinther. for much of its information. 
An unexplained but correct innovation was 
the inclusion of Icichthijs among the stro- 
mateoids. The treatment of the group was 
extremely casual; genera were shuffled into 
families more or less randomly without 
checking familial characters. The Nome- 
idae constituted almost the same unnatural 
group as set up by Giinther (1860), with 
still no realization of its relationships. 
Goode and Bean's account confused, rather 
than improved, the stromateoid classifica- 
tion. Fortunately, it has been disregarded 
by most subsequent workers. 

The first, and the only, world-wide re- 
vision of the stromateoids was that of 
Regan (1902). Regan gave the group its 
modern dimensions by adding the genera 
^'Nomcus, Cuhiceps, Pscncs, Bathy.scriola, 
and SeriolcUo, all of which have a toothed 
oesophagus exactly similar to that of a 
Centrolophus" (p. 117). His definition of 
the family was based largely on osteology, 
and made important contributions. His 
warnings of the pitfalls of allometry and 
of the unusual ubiquity of certain char- 
acters recognized a recurrent problem. Re- 
gan treated the group as one family, the 
Stromateidae, but disregarded the conve- 
nient subfamilial distinction made earlier by 
(;ill (1884). Norman's much later "Draft 
Synopsis" ( 1957 ) differs from Regan only 
in this one respect, for Norman recognized 
two families based on the first couplet of 

Regan's key to genera, "ventral fins present" 
[Centrolophidae], or "ventral fins absent" 
[Stromateidae]. Citing correspondence with 
Boulenger, Regan suggested, for the first 
time, the affiliation of Tetrcifionunis to the 

Boulenger was much impressed by one 
of Regan's diagnostic characters, the loose 
attachment of the pelvic bones to the pec- 
toral arch in all stromateids. In his system- 
atic account of the fishes for the "Cam- 
bridge Natural History" (1904), he re- 
moved the stromateids from the scom- 
broids, where all previous workers had 
placed them, and ranked them among the 
Percesoces. Holt and Byrne (1903), using 
the same argument, also considered the 
stromateids to be allied with the Percesoces. 
Although in error with respect to the rela- 
tionships of the group, their account of local 
British and Irish species was otherwise 
carefully done, and showed broad under- 

Although the presence of teeth in sac- 
cular outgrowths in the gullet had long 
been used as a diagnostic character for the 
stromateoids, no one since John Ray had 
investigated the structure of this peculiar 
feature. Gilchrist (1922) examined teeth 
from the sacs of several South African 
stromateoids. He noted differences be- 
tween species, but, although he recognized 
the value of the teeth in taxonomy, he did 
not indulge in systematic speculations. The 
sacs had previously been referred to as 
"oesophageal"; Gilchrist pointed out that 
they were "not strictly oesophageal, but 
. . . derived from . . . pharyngeal epithe- 
lium . . ." (p. 254). Later, in an incisive 
review, Barnard (1948) corrected some of 
Gilchrist's errors, and extended his work 
by examining more species and publishing 
more illustrations. 

Biihler's (1930) monograph on the diges- 
tive system of the stromateoids pointed out, 
independently from Ciilchrist, the pharyn- 
geal origin of the toothed sacs. Biihler 
proposed the terai "Rachensiiche" [= pha- 
ryngeal sacs] to replace the misleading 

Stromateoid Fishes • Haedrich 35 

on any 




of the 




"oesophageal sacs" commonly in use. His 
work was done primarily with serial micro- 
scopic sections, allowing examination and 
description of great detail. It was a sub- 
stantial contribution to understanding the 
origin, nature, and probable function of the 
teeth in the pharyngeal sacs. For details 
morjohological aspect of the pha- 
sacs, Biihler's work, or the recent 
extension of this by Isokawa et al. 
should be consulted. Other parts 
digestive system were noted to 
in rough correspondence with 
in the Rachensiiche, and within 
( 1902 ) framework Biihler pro- 
posed two subfamilies, the Lirinae, corre- 
sponding to Norman's ( vide supra ) Cen- 
trolophidae, and the Stromateinae, corre- 
sponding to NoiTnan's Stromateidae. 

The work of Gilchrist, Barnard, and 
Biihler offered sound characters for the 
stromateoid classification. But, because 
each study dealt with only a limited array 
of characters, the observations could not 
be properly or safely interpreted. 

My work has dealt primarily with skeletal 
characters. I have looked at the soft anat- 
omy only cursorily, and have found little 
of use except in a most general way. The 
study has involved only Recent fishes. 

My conclusions are largely based on the 
presence or absence of pelvic fins, whether 
the dorsal fin is separated or continuous, 
the presence or absence of certain teeth, 
the number of vertebrae, the number of 
branchiostegal rays, and, in particular, the 
structure of the caudal region and the de- 
velopment of the papillae^ in the pha- 
ryngeal sacs. The comparative morphology 
of these characters not only provides a 
reasonable separation of the suborder into 
five families, but also, because the char- 
acters change in a correlated fashion, it 
suggests the course of evolution in the 
stromateoids. In the trunk and caudal 

^ This temi implies the unit composed of a bony 
base with teeth seated upon it. It is adopted here 
in conformance with past usage (Biihler, 1930; 
Barnard, 1948). 

region the number of vertebrae increases, 
while the elements in the tail become fused 
and reduced, and the pelvic fins are lost. 
In the branchial region, the number of 
branchiostegals decreases, while the papil- 
lae of the pharyngeal sacs become in- 
creasingly more complex. The present 
geographical distributions of the different 
taxa support the conclusions based on 
anatomical evidence. 

I propose for the stromateoids a hierarchy 
of five families and fourteen genera, as 
follows : 

Order Percifomies 
Suborder Stromateoidei 
Family Centrolophidae 






Family Nomeidae 



Family Ariommidae 

Family Tetragonuridae 

Family Stromateidae 





Measurements were made point-to-point 
with a pair of fine-point dial calipers. A 
dissecting microscope with an eyepiece 
dial micrometer was used for a few very 
small specimens. Measurements routinely 
made were: 

Total length (TL), from the tip of the 
snout to the farthest tip of the caudal fin. 

Standard length (SL), from the tip of 
the snout to the caudal fin base. 

Length of head, from the tip of the 
snout to the hindmost point on the oper- 
cular membrane, usually immediately above 
the pectoral fin. 

Length of pectoral fin, from the base 
of the uppermost ray to the farthest tip 
of the fin. 


Bulletin Museum of Comparative Zoologij. Vol. 135, No. 2 

Length of pel\'ic fin, from the base of 
the most anterior ray to the farthest tip 
of the fin. 

Longest Di spine, from the base of the 
spine to its extremity. 

Predorsal distance, from the tip of the 
snout to the base of the first element of 
the dorsal fin. 

Preanal distance, from the tip of the 
snout to the base of the first element of 
the anal fin. 

Maximum depth, the greatest depth of 
the body, exclusive of fleshy or scaly fin 

Depth of peduncle, the least depth of 
the caudal peduncle. 

Snout, from the tip of the snout to the 
anterior margin of the orbit. 

Eye diameter, the greatest distance be- 
tween the fleshy margins of the eye. 

Length upper jaw, from the symphysis 
of the premaxillaries to the hindmost point, 
often covered by the lacrimal bone, on the 

Interorbital width, the least distance 
between the bony rims over the eyes. 

The measurements used for showing allo- 
metry were length of head, length of pec- 
toral fin, length of pelvic fin, predorsal 
distance, preanal distance, and maximum 
depth. These were expressed as a percent- 
age of standard length. This percentage 
was plotted against standard length follow- 
ing the method advocated by Parr ( 1956 ) . 
The measurements of snout, eye diameter, 
length of upper jaw, and interorbital width 
were expressed as a percentage of length 
of head. 

Counts were made \\'ith a fine needle, 
usually under low magnification on a dis- 
secting microscope. For extremely small 
specimens, median finray counts were 
made more easily using transmitted, polar- 
ized light. Counts routinely made were: 

D, total dorsal fin elements, spines indi- 
cated by Roman numerals, rays by Arabic. 
In some cases it was impossible to distin- 
guish between spines and rays; these counts 

are followed by the expression "total ele- 
ments. " The last, double ray of both dorsal 
and anal fins was counted as one element. 

A, total anal fin elements. 

P, total pectoral fin elements, one side, 
spine not distinguished from rays. 

Gill rakers, the total number of rakers 
on the first arch, one side. Expressed as 
number on upper limb plus one, if at angle, 
plus number on lower limb (e.g. 8 + 1-1- 17). 

Lateral line scales, the number of 
scales along the position normally occupied 
by the lateral line, one side, terminating at 
the caudal fin base. In many stromateoids 
the tubed scales end on the peduncle, but 
the count was nonetheless continued to the 
caudal base. Often the deciduous scales are 
lost and scale pockets must be counted. 
Lateral line scales is a difficult count and 
cannot be made on most specimens. 

Counts less routinely made were: 

Branchiostegal rays (BR), left side. 

Vertebrae, number of precaudal verte- 
brae plus the number of caudal vertebrae, 
including the hypural plate ( = 1 ) . Almost 
all vertebral counts were made from radio- 
graphs. Determination of the first caudal 
vertebra is hence somewhat subjective. 
Where skeletal material has been used, the 
count is followed by the expression "skel." 

All measurements and counts confomi 
with the standards of Hubbs and Lagler 

The osteology was studied primarily with 
radiographs and cleared-and-stained prep- 
arations. With the use of soft X-rays, speci- 
mens as small as 20 mm SL could be suc- 
cessfully radiographed. 

Small fishes, usually no longer than 65 
mm TL, were cleared and stained. Gill 
arches and the associated pharyngeal sacs 
were dissected from larger fish, usually 
around 190 mm TL, and were also cleared 
and stained. To remove the arches and 
sacs, cuts were made between the hyal and 
opercular series, the tongue and dentaries, 
and the last gill arch and pectoral girdle. 
The unit so freed was carefully discon- 
nected from the base of the neurocranium. 

Stromateoid Fishes • llacdrich 37 












Figure 1. The bones of the caudal skeleton, schematic drawing of the basic centrolophid type. 

and the pharyngeal sacs pulled forward 
from between the cleithra. The esophagus 
was cut behind the sacs, and the complete 
branchial apparatus removed. This was 
divided sagitally, and one half, usually the 
right, was cleared and stained. A more 
legible preparation \\ as obtained if the gill 
filaments were stripped off prior to treat- 
ment. Teeth from the pharyngeal sacs were 
examined by macerating the sac in potas- 
sium hydroxide until the muscle was trans- 
lucent, staining, and allowing the sac to 
disintegrate. Individual teeth could then 
be picked out and examined. 

Excellent results were obtained follow- 
ing the clearing-and-staining method of 
Clothier (1950) modified from Hollister 
( 1934 ) . The entire clearing-and-staining 
procedure takes about two months. 

A few complete skeletons were prepared, 
mostly from fresh specimens, by picking the 
meat carefully from the bones. Skulls were 

prepared by boiling the specimen until it 
fell apart. Only partial dissections were 
made in some cases. Whenever possible, 
skeletons in museum collections were ex- 
amined. Scales were stained in alizarin, 
blotted dry, and mounted in balsam on 

Drawings of cleared-and-stained prepara- 
tions were made through a Wild Dissecting 
Microscope with a camera lucida attach- 
ment. All caudal skeletons were drawn at 
25 power, using transmitted light. Draw- 
ings of the branchial arches were made at 
6 power, using transmitted light to draw 
the orientation of the bones, and reflected 
light to draw the arrangement and structure 
of the bases of the papillae. These draw- 
ings were redrafted on tracing paper and 
the final drawing was made with reference 
to the specimen through the scope without 
camera lucida. All anatomical drawings. 

38 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 










Figure 2. Elements of the branchial region, schematic drawing of the basic stromateoid type. 

though made from specific preparations, 
are semi-diagrammatic. 

The various elements in the drawings of 
the caudal skeleton are identified in Figure 
1. Those of the branchial region are shown 
in Figure 2. Nomenclature of the caudal 
skeleton follows Gosline (1960, 1961); that 
of the head and branchial region follows 
Mead and Bradbury (1963). 


The specimens examined are arranged 
below under the classificatory scheme here- 
in proposed. Museum and institutional 
names, in alphabetical order under each 
species, are abbreviated as follows: 

ABE —Collection of Dr. T. Abe, I'okyo 
AMS — Australian Museum, Sydney 

ANSP — Academy of Natural Sciences of Phila- 

BC — University of British Columbia, Van- 

BCF — Bureau of Commercial Fisheries, Bio- 
logical Laboratory, Washington, D. C. 

BMNH — British Museum (Natural History), 

BOC — Bingham Oceanographic Collection, 
Yale University 

CF — Danish Carlsberg Foundation, Char- 
lottenlund Slot 

CNHM— Chicago Natural History Museum 

CTS — Guinean Trawling Survey 

nOE' — International Indian Ocean Expedition 

MCZ — xMuseum of Comparative Zoology, Har- 
vard University 

MNHN — Museum National d'Histoire Naturclle, 

^ These specimens will ultimately be catalogued 
in the MCZ. 

Stromateoid Fishes • Uacdrich 


NRF — Nankai Regional Fisheries Research 

Laboratory, Kochi 
NTU — Department of Zoology, Faculty of 

Science, Tokyo University 
SAM — South African Museum 
SIO — Scripps Institution of Oceanography 
SU — Natural History Museum, Stanford 

USNM — United States National Museum, 

Washington, D. C. 
WHOF — Woods Hole Oceanographic Institution 
ZMC — Zoological Museum, Copenhagen 

The number of specimens, the range of 
standard lengths, and the locaHty appear 
in that order within the parentheses. Types 
are marked with an asterisk (*), the spe- 
cies name follo\\'ing within the parentheses. 
Specimens radiographed are marked with 
a dagger ( t ) ; specimens that have been 
cleared and stained are marked CS. 


H. antarctica ( Carmichael ) : tAMS IB. 3825 
(I, 170 mm, near Sydney, N.S.W.). *t BMNH 
1855. 9. 19. 2 ( 1, 108 mm, coasts of Australia, 
Diagravima porosa Richardson). tSAM 23592 
(1, 105 mm. Cape Point, South Africa). 

H. hythitcs (Ginsburg): CNHM 46408 (1, 12 
mm. Ocean Spring, Miss.). *tUSNM 157776 
( 1, 197 mm, off Pensacola, Fla., Palimiricli- 
thijs hijthitcs Gimhing) ; *tUSNM 157778 ( 1, 
187 mm, Mississippi Delta, paratype Pali- 
nurichthiis hifthites Ginsburg). WHOI (1, 204 
umi, OREGON sta. 3762); WHOI (1, 203 
mm, OREGON sta. 4011); WHOI (7, 188- 
213 mm, OREGON sta. 4030). 

H. iaponica (Doderlein): ABE 58-258, 60-107 
to 111, 60-113 to 116. 60-139, 60-225, 60-370, 
60-709, 60-744, 60-775, 61-452, 61-458, 61- 
461 to 464, 61-500 to 511, 62-302, 62-303, 
62-336 to .368, 63-435, 63-476, 63-447, 63- 
480, 63-541 to 544, 63-547, 63-668, 63-708, 
6.3-728, 63-872 (35, 35-233 mm, Manazuru, 
Japan); ABE 64-2201 and 2202 (2, 400, 415 
mm, Tokyo market); ABE 2236 (1, 695 mm, 
Tokyo market); ABE plankton collection (1, 20 
mm, East China Sea). CNHM 59428 (1, 420 
mm, Tokyo). 

H. percifonna (Mitchill): tMCZ 36624 (2, 159. 
209 mm. Woods Hole, Mass.). WHOI (47, 
101-198 mm, 40°10'N 69°30'W, gill arch CS); 
WHOI (1, 50 mm, 38°37'N 69°24'W, CS). 


S. griscolineaius (Norman): tBMNH 1936.8. 
26. 1068-9 (2, 258, 258 mm, South Atlantic); 

*tBMNH 1936.8.26.1070-1 (2, 193, 196 
mm, South Atlantic, Palinurichthys grheo- 
Uneatus Norman ) . 

S. macukitus Giinther: *tBMNH 48. 3. 16. 150 
(1, 37 mm, China Seas, S. Duiculatus Gimther). 

S. huttoni (Waite): tZMC (1, ca. 40 mm, 
34°24'S 94M5'W). 

S. maniioratus Kner and Steindachner: *tHam- 
burg Museum H464 (1, 37 mm, Siidsee, S. 
luaiDiorafus Kner and Steindachner). tZMC 
(2, 28 and 46 mm, GALATHEA sta. 176). 

S. medusophagus Cocco: *BMNH 
(I, ca. 405 mm, stuffed, Cornwall, Centro- 
lopJius britannicits Giinther). tCF (7, 59- 
188 mm, North Atlantic). tUSNM 163880 
(1, 333 mm, Kitty Hawk). WHOI (19, 5.5- 
435 mm. North Atlantic, one CS). 

S. ovalis (Cuvier and Valenciennes): *tBMNH 
1860.7.17.2-3 (2, 171, 186 mm, Madeira, 
Leiws bennettii Lowe). *MNHN 264.4. 1.2 
( 1, 299 mm, Nice, Centrolophus avails Cuvier 
and Valenciennes); *MNHN (1, 
144 mm, Canary Islands, Crius hcrthclotti 
Valenciennes ) . 

S. pcmarco (Poll): BCF 928 (1, 95 mm, 4°31'S 
10°53'E). tGTS (11, 103-217 mm, Gulf of 
Guinea, gill arch CS ) . 


C. niger Lacepede: *tBMNH 1862.6.14.16 
( 1, 290 mm, Madeira, Schedophilus elongatus 
Johnson); BMNH (1, .385 mm, no 
locality); BMNH 55.9. 19. 1461 (1, 124 mm, 
Hasler Coll.); BMNH (1, 183 
mm, Falmouth); BMNH 1934.8.8.67 (1, 
362 mm, SW Ireland); BMNH 1934.8.8. 
68-9 (2, 498, 560 mm, SW Ireland). tCF 
(2, 147, 204 mm, DANA sta. 4205, gill arch 
CS). MCZ .34246 (1, 443 mm, 42°10'N 
66°45'W); MCZ 37983 (1, 284 mm, Province- 
town); MCZ 37984 (1, 490 mm. Sable Island 
Bank). *tMNHN (1, 287 mm, 
Fecamp, Centrolophus niger Lacepede); 
*MNHN, (3, 169-207 
mm, Nice, Centrolophus morio Cu\'ier and 
Valenciennes); MNHN (1, 180 
mm, Naples); *MNHN 264. 4. 2. 1 (1, 121 mm, 
Marseille, Centrolophus valenciennesi Moreau). 
tUSNM 44440 (1, 189 mm, Dennis, Mass.); 
USNM 48367 (1, 39 mm, Naples); tUSNM 
48906 (1, 265 mm. North Truro, Mass.); 
USNM 49335 (1, 220 mm, Genoa). WHOI 
(1, 1065 mm, 40°13'N 65°45'W). 


I. lockingtoni lordan and Gilliert: ABE 63-526, 
63-527, 63-529, 63-530, 63-548, 63-549, 63- 
555 ( 12, 72-173 mm, Manazuru, gill arch 
CS). BC 53-99A (1, 164 mm, Vancouver 

40 Bulletin Museum of Comparative Zoology, Vol. 135. No. 2 

Island); BC 59-652 (1, 132 mm, Vancouver 
Island). *SU 7442 (1, 65 mm, Monterey Bav, 
Schedophilu.s hcathi Gilbert); SU 17346 (1, 
42 mm, Seal Beach); SU 22955 (1, 41 mm, 
Catalina Island); SU 22971 (1, 26 mm, 
Monterey Bay); SU 41028 (8, 15-78 mm, 
Monterey Bay, CS). *tUSNM 27397 (1, 159 
mm, Point Reyes, Icichthij.s lockingtoni); 
*tUSNM 89398 (3. 28-65 mm, Monterey 
Bay, Centrolophu.s califomicus Hobbs). WHOI 
(2, 17, 83 mm, Monterey Bay, received 
through Giles Mead). 


S. }>r(nii(i Giinther: tAMS lA. 10170 (1, 233 
mm, Sydney Harbour, New South Wales ) ; 
tAMS 1.10333 (1, 152 mm, 40 miles W 
Kingston, South Australia). *BMNH (1, 298 
mm. New Zealand, stuffed, Neptomenu.s 
hrama Giinther). 

S. punctata (Bloch and Schneider): tAMS I. 
10840 (1, 192 mm. Oyster Bay, Tasmania); 
tAMS 1.14747 (1. 220 mm, Portobello). 
*tBMNH 1869. 2. 24. 42^44 (3, 234-243 mm, 
Tasmania, Neptoiuenus doJntIa Giinther). 
USNM 176915 (2, 190, 195 mm, Queensland, 
gill arch CS); tUSNM 176968 (1, 149 mm. 
New South Wales); tUSNM 177109 (2, 197, 
203 mm. New South Wales). 

S. pnrosa Guichenot: tUSNM 176478 (1, 218 
mm, Tictoc Bay, Chile); tUSNM 176535 (1, 
198 mm, Puerto Auchemo, Chile); tUSNM 
176593 (3, 197-203 mm, Auellon, Chile). 

S. viohcea Guichenot: tMCZ 17239 (2, 430, 
445 mm, Callao, Peru). *SU 9590 (1, 262 
mm, Callao, Peru, paratype Ncptomenus cras- 

sti-s Starks ' 

rUSNM 53465 (1, 265 mm. 

Callao, Peru, Neptoiuenus cra.s-s-us Starks); 
tUSNM 77513 (1, 130 mm, Mellendo, Peru); 
tUSNM 77593 (1, 150 mm, Mellendo, Peru, 
gill arch CS); tUSNM 77611 (1, 173 mm, 
Callao, Peru); tUSNM 77625 (1, 213 mm, 
Callao, Peru). tZMC (1, 87 mm, 14°S 77°W). 


P. anoniala ( Temminck and Schlegel); ABE 60- 
1232, 63-752, 63-1141 (3, 93-119 mm, Mana- 
zuru market); ABE 62-656 (1, 40 mm, Japan, 
CS); ABE 61-590 (1, 160 mm, Tsubaki); 
ABE 64-1223 to 1225 (25, 43-84 mm, 32^09' 
N 123°15'E); ABE 64-1972 to 1959 (8, 17- 
52 mm, Kozu); ABE 64-2014 to 2017, 64- 
2142 to 2144, 64-2148 to 2150 (14, 7.5-93 
mm, off Misaki); ABE 64-2037 to 2039 (3, 
30-42 mm, Amakusa Island); ABE, plankton 
collection (9, 7-28 mm. East China Sea); 
ABE (1, 150 mm, Tokyo market, gill arch 
CS). BC 56-29 (1, 136 mm, Tokyo market); 

BC 59-555 (1, 138 mm, Aberdeen market). 
CNHM 57288 (3, 91-124 mm, Kobe). tMCZ 
1186 (2, 122, 143 mm, Kanagawa); MCZ 
31150 (1, 125 mm, Yenosima). tUSNM 
6424 (1, 149 mm. Hong Kong); tUSNM 
49465, 71131, 151829 (3, 146-156 mm, 
Tokyo market); tUSNM 59618 (1, 141 mm, 
Matsushima Bay); tUSNM 177426 (2, 132, 
142 mm, Taipei market). 

P. ctjanea Alcock: *tBMNH 1890.11.28.9 ( 1, 
120 mm, Ganjam Coast, India, Bathuseriola 
cijanea Alcock); tBMNH 1937.6.28.1-8 (9, 
103-110 mm, off Cananore). 

Psenopsis sp.: tUSNM 98818 (1, 132 mm. 
Mare Island, Dutch East Indies). 


C. athenae Hacdrich: *tMCZ 42974 ( 1, 68 mm, 
38°36'N 71°24'W, Cubiceps athenae Hae- 
drich). *tUSNM 198058 (1, 81 mm, 28°54'N 
88°18'W, paratype Cubiceps athenae Hac- 
drich). WHOI (1, 33 mm, 24"2rN 8I°15'W, 

C. caeruleus Regan: *tBMNH 1913.12.4.28- 
29 (2, 86, 95 mm. Three King's Isl., New 
Zealand, Cubiceps caeruleus Regan); BMNH 
1926.6.30.50 (1, 282 mm. Lord Howe Isl., 
Tasman Sea ) . 

C. capensis (Smith): *BMNH (1, 905 mm. 
South Africa, Atimnstoma capensi.^ Smith, 
stuffed); tBMNH 1925.10.14.1-4 (3, all 
ca. 160 mm, 70 mi. WNW Saldanha Bay, 
South Africa). 

C. carinatus Nichols and Muri^hy: CNHM 
61958 (5, 93-99 mm, 83°15'W 5n5'N); 
tCNHM 61939 (2, both 110 mm, 125 mi. SW 
Cape Mala, Panama). SIO 63-538 (1, 101 
mm, 2I°35'N 107°00'W); SIO 63-882, 63- 
888, 63-892 (4, 65-101 mm, Golfo de Te- 
huantepec); SIO 63-1027 (1, 91 mm, 13°33' 
N 95°59'W); SIO H 49-77 (1, 87 mm, off 
El Salvador); SIO H 52-351 (2, 90, 94 mm, 
13°45'N 99°22'W). ZMC (1, 57 mm, 11°52' 
N 97°19'W); ZMC (2, 64, 71 mm, 12n4'N 
97°46'W); ZMC (1, 70 mm, 12°38'N 98°14' 
W); ZMC (1, 72 mm, 13°00'N 98MrW); 
ZMC (1, 71 mm, 13°4rN 97°34'W). 

C. gracilis Lowe: tBMNH (2, 
143, 150 mm, Madeira); BMNH 1960.12. 
19.8 (1, 165 mm, 5r51'N 13°43'W). CF 
(5, 40-57 mm, DANA sta. 855); CF (2, 50, 
59 mm, DANA sta. 856); CF (26, 5-35 mm, 
DANA sta. 939, one CS); CF (1, 69 mm, 
DANA sta. 1372); CF (1, ca. 70 mm, 
DANA sta. 1378); CF (1, 77 mm, DANA 
sta. 1380); CF (1, 43 mm, DANA .sta. 
4017, CS); CF (4, 43-76 mm, DANA sta. 
4185); CF (18, 16-80 mm, DANA sta. 
4192); CF (37, 11-61 mm, DANA sta. 4195); 

Strom ATEOiD Fishes • Hacdrich 41 

CF (50, 10-58 mm, DANA sta. 4197). 
*MNHN 42-29 (1, 195 mm, Sete, Trachelo- 
cirrhus meditcrraneiis Doumet). WHOI (8, 
20-51 mm, DELAWARE 63-4 sta. 3); WHOI 
12, 2.5-51 mm, 39°27'N 27°35'W); tWHOI 
(1, 92 mm, DELAWARE 63-4 sta. 13); 
tWHOI (2, 75, 81 mm, DELAWARE 63-4 
sta. 16). 

C. longimanus Fowler: *ANSP 55058 (1, 
42 mm, Durban, Natal, Culnceps lonfiitnamis 
Fowler). IIOE (1, 29 mm, ANTON BRUUN 
6 sta. 338). 

C. pauchmliatus Gimther: ABE 57-347 (1, 120 
mm, Manazuru, Japan); ABE 10832 (1, 80 
mm, Kochi, Japan). *ANSP 68380 (1, 124 
mm, 40 mi. S Christmas Isl., Line Islands, 
Cuhiceps nesiotes Fowler). *tBMNH 1870. 
8.31.124 (1, 113 mm, Misol, Molucca Isl., 
Cuhiceps pauciradiatus Giinther). SIO 60- 
216 (1, 93 mm, 10°26'N 128°22'W). 

C. squamiceps (Lloyd): ABE 57-348, 59-37, 
61-843, 62-13, 62-106 (5, 84-154 mm, Mana- 
zuru, Japan); ABE 64-1348 (1, 264 mm, 
6°15'N 164°10'W); ABE 10833 (1, 159 mm, 
Kochi, Japan). 


N. gronovii (Gmelin): ABE 64-1280 (1, 68 
mm, Kushimoto, Japan); ABE plankton col- 
lections (5, 8-38 mm, Japan). BCF 1002 
( 1, 61 mm, 3°41'N 0"05'E ) ; BCF 1003 ( 1, 40 
mm, GERONIMO 4-155). BOG 602 (6, 17- 
101 mm. Key West Harbor); BOG 3361 (21, 
10-118 mm, Atlantic Ocean); BOG 3515 (22, 
13-149 mm, Gulf of Mexico); tBOG 3516 
(8, 31-153 mm, Gulf of Mexico); BOC 3517 
(14, 16-133 mm, 28"07'N 89°53'W); BOC 
3518 (11, 13-114 mm, Atlantic Ocean). 
tCNHM (1, 225 mm, OREGON sta. 1178). 
IIOE (4, 20-45 mm, 2^20'N 65°54'E); IIOE 
(9, 12-41 mm, 8°00'S 65"00'E). tMCZ 35327 
(2, 144, 154 mm, 39°27'N 70°38'W gill arch 
GS). *MNHN (3, largest ca. 
42 mm, seas of Java, Nomeus peronii Guvier 
and Valenciennes). SIO 60-263 (4, 22-40 
mm, 5°18'N 160°05'W); SIO 61-84 (2, 20, 
25 mm, 5°58'S 149°31'W); SIO 61-87 (1, 
33 mm, 1°32'S 148°39'W); SIO 61-89 (3, 
32-41 mm, 5°32'N 146°09'W). WHOI (2, 
45, 66 mm, 17°00'N 65 05'W, CS). 

P series 

P. arafurensis Giinther: ABE 60-101 (1, 72 
mm, Manazuru, Japan); ABE 62-651 (1, 35 
mm, Japan, CS)"; ABE 64-212 (1, 150 mm, 
Nagasaki); ABE 64-1767 (1, 92 mm, Komat- 
subara, Japan); ABE, plankton collection (2, 
19, 37 mm. East China Seas). *tBMNH 

1889. 7. 20. 55 ( 1, 30 mm, China Seas, Fsenes 
arafurcmus Giinther). IIOE ( 1, 25 mm, 2°20' 
S 64°54'E); IIOE (1, 20 mm, 4°0rS 65°02' 
E). MCZ 41550 (5, 14-18 mm, 10°52'N 
29°26'W). WHOI (1, 20 mm, 41°33'N 54° 
P. hemmli Rossignol and Blache: *MNHN 264. 

9.1.1 (1, 63 mm, 3°38'S 9°22'E, Psenes 
hcmirdi Rossignol and Blache); *MNHN 264. 

9.1.2 (2, 63, 65 mm, 1°55'S 8°30'E, para- 
types, Psenes henardi Rossignol and Blache ) . 

P. cijanophrifs Guvier and Valenciennes: *t 
BMNH 1871. 7. 20. 156 ( 1, 111 mm, Manado, 
Philippines, Cuhiceps mtdtiradiatus Ciiinther). 
CF ( 1, 28 mm, Dansk Vestindien sta. 
132, CS). *tMNHN (1, 117 
mm. New Ireland, Bismarck Archipelago, 
Psenes cijanophnjs Guvier and Valenciennes). 
CNHM "46409 (1, 80 mm, 25"N 89°W). 
WHOI (1, 38 mm, CRAWFORD 62, CS). 
tWHOI (2, 119, 128 mm, OREGON sta. 
3715, gill arch CS). 

P. maculatus Liitken: ABE 64-1226 (1, 20 mm, 
Japan): ABE plankton collection (1, 20 mm, 
"East China Sea). CF (1, 48 mm, 15°31'N 
18°05'W). tMCZ 41122 (1, 59 mm, 40M9'N 
64°57'W). WHOI (1, 27 mm, 38°26'N 68° 
15'W); WHOI (1, 35 mm, 41°36'N 60^30' 
W). *tZMC (2, 57 and 72 mm, 39^00'N 
34°10'W, P. maculatus Liitken). 

P. pcllucidus Liitken: ABE 59-172, 60-102, 60- 
106, gill arch CS. 60-140 and 141, 61-21, 
61-82 and 83, 61-459 (13, 72-147 mm, Mana- 
zuru, Japan); ABE 63-1064 (1, 89 mm, 
Tokyo market). tBGF 957 (1, 130 imn, 
4°07'S 10°23'E). CF (1, 60 mm, 15°31'N 
18°05'W); CF (1, 39 mm, AGENT PETER- 
SEN sta. 769, CS). CNHM 5285 (1, 54 mm. 
Bermuda); tCNHM 49189 (1, 131 mm, Ber- 
muda); nCNHM 57097 (1, 193 mm, Oki- 
nawa, Icticus ischanus Jordan and Thomp- 
son). *tUSNM 49745 (1, 67 mm, Newport, 
Rhode Island, Psenes eduardsii Eigenmann). 
SU 43310 (1, 93 mm, Bermuda). WHOI 
(1, 25 mm, 38°38'N 68°50'W). *tZMC (1, 
38 mm, Surabaya, P. peUucidus Liitken). 


A. africana (Gilchrist and von Bonde): *t 
BMNH 1927. 12. 6. 45 (1, 166 mm, Agulhas 
Bank, South Africa, co-t\pe, Psenes africanus 
Gilchrist and von Bonde ) . 

A. hondi Fowler: *ANSP 52528 (1, 79 mm, 
Grenada, British West Indies, Arioninia hondi 
Fowler ) . 

A. doUfusi (Ghabanaud): *tBMNH 1931. 4. 16. 
1 (1, 112 mm. Gulf of Suez, co-type, Cuhi- 
ceps doUfusi Ghabanaud ) . 

A. evermanni Jordan and Snyder: *tUSNM 

42 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

57783 ( 1, 156 mm, Honolulu, Hawaii, Arioiii- 
ma evcrmanni Jordan and Snyder). 

A. irulica (Day)": BC 59-555 (1, 151 mm, 
Aberdeen market, Hong Kong). *tBMNH 
1889.2.1.3255-6 (2, 74, 90 mm, Madras. 
India, Psene.s- imlicm Day). NTU 51941 to 
51946 (6, 123-181 mm, near Hainan Island, 
sill arch CS). 

A. Ittrida Jordan and Snyder: *SU 8441 (1, ca. 
190 mm, Honolulu, Hawaii, paratype Ariomma 
Itirida Jordan and Snyder). *tUSNM 51400 
(1, 166 mm, Honolulu, Hawaii, Ariomma hirida 
Jordan and Snyder); tUSNM 109418 (1, 193 
mm, Honolulu). 

A. mdimu (Ginsburg): *tUSNM 157779 ( 1, 
154 mm, Mississippi Delta, Cubiceps melanus 

A. multisquumis (Marchal): *MNHN 264.7.2. 
1 (type), (paratype) (2, 159, 
147 mm, Ivory Coast, Parcictihiccps- miilti- 
sqiiamis Marchal). 

A. nigriargenteus (Ginsburg): *MCZ .37183 (1, 
113 mm. Sandwich, Massachusetts, paratype 
Cuhiceps nigriurgenieus Ginsburg). *tUSNM 
151954 (1, 190 mm, off Cape Romain, South 
Carolina, Cuhiceps nigriargcnteus Ginsburg). 

A. regidm (Poey): tUSNM 197110 (2, 142, 
150 mm, British Guiana). tWHOI (1, 136 
mm, 29°59'N 87°06'W). 

Ariomma sp. Western North Atlantic: MCZ 
40259 (1, 116 mm, Provincetown, Massachu- 
.setts); MCZ 40498 (1, 115 mm. Province- 
town, Massachusetts). SU 57297 (1, 86 mm, 
Bermuda). WHO! (3, 87-121 mm, OREGON 
sta. 3725); WHOI (2, 103, 109 mm, ORE- 
GON sta. 3733); WHOI (3, 122-134 mm, 
OREGON sta. 4014); WHOI (1, 143 mm, 
9°03'N 81^22^); WHOI (2, 141, 149 mm, 
9°13'N 80°44'W); WHOI (3, 139-140 mm, 
16°45'N 8r27'W); WHOI (3, 131-140 mm, 
28°57'N 88°4rW); WHOI (2, 125, 133 mm, 
29°07'N 88°34'W); WHOI (3, 1,30-140 mm, 
28°54'N 88°51'W, gill arch CS); WHOI (1, 
23 mm, CRAWFORD 62 sta. 29, CS); WHOI 
(1, 28 mm, 24°N 81 °W, CS). 

Ariomma sp. Japan: ABE 59-404, 59-408, 60- 
124, 60-144, 60-224, 60-478 and 479, 60-1611, 
61-1188, 62-738, 62-955, 62-1387, 62-1628 
(13, 99-226 mm, Manazuru, Japan). NRF 
1441 (1, ca. 800 mm, Bonin Islands). 


T. atlanticm Lowe: *tBMNH ( 1, 197 mm, 
Madeira, Tctragonurus ailanticas Lowe). 
MCZ 41726 (1, 21 nun, 39°47'N, 70'32'W, 
CS); MCZ 41791 (1, 66 mm, 39°4rN 69° 
54'W, CS). WHOI (2, 15, 20 mm, tropical 

T. cuvicri Risso: CNHM 64218 (1, 340 mm, 
46°5rN, I55°00'W, gill arch CS). 


S. Inasilicnsis Fowler: *ANSP 11354 (1, 276 
mm, Rio Grande do Sul, Brazil, Stromatcus 
hra.silicnsi.s Fowler). tMCZ 4599 (3, 254- 
285 mm, Rio Grande do Sul, Brazil). 

S. ficitola Linnaeus: tBMNH 87. 3. 2. 30 ( 1, 76 
mm. Lower Congo). tGTS (6, 200-240 mm, 
7°20'N I2°40'W). tMCZ 16729 (1, 257 mm, 
Mediterranean?). tSU 1537 (1, 44 mm, 
Palermo, Italy). tUSNM (I, 177 mm. Fed. 
Fish. Serv. Nigeria, No. 4046). 

S. steUattis Cuvier: tUSNM Ace. No. 167496 (2, 
230, 237 mm, 42°29'S 72°46'W). *MNHN (2, 144, 164 mm, Valparaiso, 
Chile, Stromatetis maculatus Cuvier and Va- 
lenciennes). tUSNM 176474 (3, 178-250 mm, 
Calbucco, Chile); tUSNM 176494 (1, 193 
mm, 41°52'S 73°53'W). 


P. alcpidotus (Linnaeus): tUSNM 127352, 
127353 (2, 122, 141 mm, Grand Terre, La.); 
tUSNM 23215 (1, 130 mm. Bay Chaland, 
La.). WHOI (1, 39 mm, SILVER BAY sta. 
4331, CS). 

P. palometa ( lordan and Bollman): *tUSNM 
41136 (5, 53-61 mm, Perlas Isl; Panama, 
Stromatcus palometa Jordan and Bollman; 
tUSNM .50.337 (2, 136, 169 mm, Panama). 

P. paru (Linnaeus): tMCZ 4600 (1, 55 mm, 
Brazil); tMCZ 41064 (2, 48, 57 mm, Port-au- 
Prince, Haiti). *MNHN (1, 93 mm, Rio de 
Janeiro, Sescrinus xautJitirus Quoy and Gai- 
mard ) . 

P. simillimus (Ayres): tMCZ 26875 (1, 110 
mm, San Diego, Calif.). tSU 48000 (1, 82 
mm, Oceanside, California). 

P. smjdcri Gilbert and Starks: *tBMNH 1903. 
5. 15. 190 (1, 217 mm, Panama, co-type 
Peprilus sntjderi Gilbert and Starks). *tUSNM 
50448 ( 1, 189 mm, Panama, PcprUus smjdcri 
Gilbert and Starks). tUSNM 76796 (1," 178 
mm, Panama City). 

P. triacanthus (Peck): tABE 64-1920, 1924, 
1930 (3, 80-110 mm, Florida). tWIIOI (7, 
157-195 mm, SILVER BAY sta. 4104); 
tWHOI (4, 127-155 mm, 40°0rN 7r23'W); 
tWHOI (1, 120 mm. Woods Hole, gill arch 
CS); WHOI (I, 36 mm, BEAR 188 sta. 371, 


P. argenteus (Euphrascn): tABE 64-1231, 12.33 
(2, 10,5, 147 mm, Bangkok); tABE 64-1929 
(I, 1,33 mm. Hong Kong); tABE 19,36, 1937 

Stromateoid Fishes • Haedrich 43 

(2, 48, 72 mm, off mouth of Sliiota River, 
Ariake Sound, Kyushu, CS); tABE 64-1964 
(1, 156 mm, East China Sea); tABE P 4347 
(1, 92 mm, Sarawak, North Borneo). tllOE 
(6, 65-161 mm, Bombay). tUSNM 44900 
(1, 299 mm, Japan). 

P. chinemis ( Euphrasen ) : tABE 64-1229 (1, 
89 mm, Bangkok); tABE P 2211, P 4319 
(2, 77, 112 mm, Sarawak, North Borneo). 
tCNHM 15917 (1, 74 mm, Batavia, Java). 
tMCZ 16772 (1, 84 mm, Singapore). 

P. echinogaster ( Basilewsky ) : tABE 64-902, 
64-906, 64-909, 64-911, 64-926, 64-1192 (5, 
165-207 mm. East China Sea); tABE 64-1112 
(1, 221 mm, Tokyo market); tABE 1743 ( 1, ca. 
180 mm, Tokyo market, gill arch CS); tABE 
64-1934 (1, 158 mm, south of Poi Toi Isl., 
Hong Kong). tCNHM 55810 (3, 81-106 mm, 
Chinnampo, Korea). tSU 22292 (3, 77-87, 
Chinnampo, Korea). tUSNM 75941 (1, 236 
mm, Japan?). 

In addition to stromateoid fishes, the 
following non-stromateoids were examined: 


Apolectus [= Parastromateus] niger MCZ 

15912 (3, Singapore). 

Arripis trutia ZMC lA.VAl^ (3, New Zealand). 

Menidia sp. WHOI (2, Morehead City). 

Curanx hippos WHOI ( 1, Woods Hole), hkw- 

cratcs ductor WHOI (1, ATLANTIS sta. 219). 

Selar crutucnoptliahuus WHOI ( 1, Woods 

Hole). Trachinotus glaucus WHOI (1, Tru- 

jillo, Honduras ) . 

Conjphaena equiselis WHOI (1, OREGON 

sta. 1297). 

Pkitax occUaim MCZ 2748 (1, Manila). 

Icosfetis aenigmaticus BC 63-98 (1, Alaska); 

BC 64-12 (1, Spiller Channel). MCZ 34915 

(1, California). SU 1171 (1, Pacific Grove); 

SU 25640 ( 1, Monterey Bay). *USNM 27398 

(1, Point Reyes); USNM 75159 (1, Pacific 

Grove ) . 

Boops vulgaris MCZ 21706 (1. France). 

Girella nigricans MCZ 10775 (1, Cahfornia). 

KuJdia main MCZ 29367 (1, Papeete). 

Kyphosus sectatrix WHOI (2, 32°08'N 67° 

lO'W). Pimelepterm hoscii MCZ 2610 ( 1, 

Florida ) . 


MonodacUjlus argcntcus MCZ 34101 (1, Dar 
es Salaam). 


Nematistius pcctoralis BC 60-15 (1, Acapulco 
market ) . 


Pomatomus saltatrix MCZ 16941 (5, New Jer- 
sey). WHOI (5, Woods Hole). 


Scomberesox saurus WHOI (1, 40°12'N 62° 


Auxis thazard WHOI (1, 41°16'N .57°37' 
W). Scomber scomber WHOI (1, Quisset, 


Scorpls calif orniensis MCZ 4896 (2, Cali- 
fornia ) . 


Autlsthes puta WHOI (2, Australia, received 
through James Moulton). Pelates sp. WHOI 
(2, Australia, received through James Moul- 
ton). Therapon jarbita MCZ 24823 (2, Java). 


Zaprora silcnus BC 61-573 ( 1, Cross Sound, 


In the classification which follows, the 
suborder, the families, and the genera are 
characterized. Categories below the genus 
are not. In a number of cases, subgeneric 
division is certainly called for. This action 
is not taken here, but is reserved for critical 
treatment in a planned series of mono- 
graphs. Likewise, the proper sorting of spe- 
cies is left for the future. As a preliminary 
step in clarifying the confusion which sur- 
rounds stromateoid classification, however, 
lists of nominal species are included under 
each genus. Species synonymies, whenever 
given, are to be considered tentative. 

The synonymy of the suborder is fairly 
complete and is intended as a guide to most 
works, especially those of a faunistic nature, 
that include references to stromateoids. 
Distributional notes concerning single or 
very few species, however, have not been 
included. The synonymies of each family 
contain only major references. Family 
names have been used with such confusion 
in the past that complete synonymies would 
be essentially meaningless. 

44 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Included under each taxon are: synon- 
ymy, diagnosis or brief characterization, 
description, distribution (suborder, fami- 
hes), natural history (suborder, genera), 
relationships, and key to included taxa or 
list of nominal species. An asterisk (*) 
precedes the names of species of which I 
have seen the types. 


Stromatini. Rafinesque, 1810:39 {Stronwtcus, 

Stromateini. Bonaparte, 1846:76 {Siwmatcm, 
Peprilus, Luvarus, Kurtus). 

Stromatiniae. Swainson, 1839:177 (Seserinus, 
Stromateus, Peprilus, Kurtus, Kcris). 

Stromateina. Giinther, 1860:397 (+ Nomeina p. 
387, clef., Stromateus, Ccntrolophu.s, Gastero- 
chisma, Nomcus, Ctibiceps, Neptomenus, 
Phtijsieth us, Ditrema ) . 

Stromateidae. Gill, 1884:66.5 (def., gen. synopsis, 
Centrolophus, Schcdophilus, Lirus, Stromateus, 
Stromateoides, Psenopsis). Day, 1875:246 ( + 
Nomeidae, p. 237. descr.; India). Jordan and 
Gilbert, 1882:449 (key, descr.; North Amer- 
ica). Fordicc, 1884: 311 (key, synon.; Amer- 
ica). Collett, 1896:26 ( -f Nomeidae, p. 31. 
descr.; eastern Atlantic). Goode and Bean, 
1896:213 (descr.; oceanic spp.). Regan, 1902: 
117 (major revi.sion, Nomeus, Cuhiceps, 
Psenes, SerioleUa, Psenopsis, Centrolophus, 
Lirus, Stromateus, Peprilus, Stromateoides). 
Holt and Byrne, 1903:71 (key, descr.; British 
Isles). Boulenger, 1904:64.3 (popular ac- 
count). Smith, 1907:221 (key, descr.; North 
Carolina). Miranda-Ribeiro, 1915 (key, descr.; 
Brazil). Gilchrist, 1922:249 (papillae). Gil- 
christ and von Bonde, 1923:1 (descr.; South 
Africa). Meek and Hildebrand, 1925:407 
(key, descr.; Panama). Biihler, 19.30:62 (di- 
gestive system). Fowler, 1936:658 (key, 
descr.; West Africa). Fowler, 1941:152 ( + 
Nomeidae, list; Brazil). Fowler, 1944:78 ( + 
Centrolophidae, p. 79; Nomeidae, p. 80. descr.; 
Chile). Hildebrand, 1946:416 (descr.; Peru). 
Barnard, 1948:394 (descr., sacs and papillae; 
South Africa). Smith, 1949:302 (key, descr.; 
South Africa); 1949a: 8.39 (revision; South 
Africa). Lozano y Rey, 1952:648 (descr.; 
Iberia). Mori, 1952:138 (+ Nomeidae, hst; 
Korea). Bigelow and Schroeder, 1953:363 { + 
Centrolophidae, p. 369, descr.; Gulf of Maine). 
Herre, 1953:258 (+ Nomeidae. list, synon.; 
Philippines). Tchang et al, 19.55:195 (descr.; 
Gulf of Pechili, Yellow Sea). Poll, 19.59:125 
(descr.; West Africa). Blache, 1962:70 (hst; 
West Africa). Lowe, 1962:694 (list; British 

Guiana). Chu et al, 1963:407 (descr.; East 
China Sea). 

[Stromateidae.] Liitken, 1880:513, 521 (disc, gen., 
oceanic spp.). Giinther 1889:10 (disc, gen., 
descr.; CHALLENGER). Nobre, 1935:332 
(descr.; Portugal). Okada, Uchida, and 
Matsubara, 1935:123 (descr., ill; Japan). 
Kamohara, 1940:173 (descr.; Japan). 

Nomeifonnes. Gregory, 1907:. 502 (relationships). 

Stromateiformes. Jordan, 1923:182 (list, fam., 
gen.). Jordan, Evermann, and Clark, 1930: 
226 (list, synon.; North America). 

Centrolophidae.' Fowler, 1928:138 (descr.; Pa- 
cific); 1931:325 (descr.; add. Pacific); 1934: 
403 (descr.; add. Pacific); 1949:75 (descr.; 
add. Pacific). Norman, 1937:115 (+ Stro- 
mateidae, p. 118. descr.; Patagonia). Sanz- 
Echeverria, 1949:151 (otoliths). Tortonese, 
19,59:.57 (revision; Gulf of Genoa). 

Stromateoids. Gregory, 1933:306 (skull of Rhom- 
l)us [= Peprilus] ). 

Stromateoidei. Berg, 1940:.324 (def., fam. synop- 
sis); 19.55:247 (def., fam. synopsis). Bertin 
and Arambourg, 1958:2441 (fam., gen. synop- 
sis). Munro, 1958: 117 (descr.; Australia). 
Duarte-Bello, 1959:119 (list; Cuba). Gosline 
and Brock, 1960:281 (descr.; Hawaii). Chu 
et al, 1962:759 (descr.; South China Sea). 
Scott, 1962:142 (key, descr.; South Australia). 

Stromateoidea. Blegvad and L0ppenthin, 1944:178 
(descr.; Iranian Gulf). Beaufort and Chap- 
man, 1951:85 (descr.; Indo-Australian Archi- 
pelago). Norman, 1957:216 (fam., gen. 
syTiopsis, after Regan). Herald, 1961:243 
(popular account). Marshall, 1964:398 (key, 
descr.; Eastern Austraha). 

Stromateoidae. Clemens and Wilby, 1961:230 
(de.scr.; British Columbia). 

Distinctive characters. There is no mis- 
taking the "stromateoid look." Though the 
characters given in the diagnosis are the 
only ones that absolutely identify a member 
of the suborder, these fishes nonetheless 
have a physiognomy that nine times out of 
ten says "Stromateoid!" to an experienced 
ichthyologist. Once recognized, the stro- 
mateoid expression is not likely to be for- 
gotten. It is a fat-nosed, wide-eyed, stuf fed- 
up look, smug and at the same time appre- 
hensive. Some stromateoids might even be 
accused of a certain prissiness. 

The stromateoid look results from the ex- 
panded lacrimal bone all but covering the 
maxillary, the slightly underslung lower jaw 
shutting within the upper, the large cen- 

Stromateoid Fishes • Haedrich 45 

trally located eye rimmed with adipose tis- 
sue, and the protruding, inflated, naked, 
and pored snout and top of the head. 

Diagnosh. Perciform fishes with toothed 
saccular outgrowths in the gullet immedi- 
ately behind the last gill arch. Small teeth 
approximately uniserial in the jaws. 

Description. Body slender to deep, com- 
pressed or rounded. Dorsal fin single or 
double; dorsal spines present, very weak in 
some species. One to three anal spines, 
never separated from the rays. Dorsal and 
anal fins coterminal. Pelvic fins present or 
absent. Rays in pectoral fin 16 to 25. Body 
scaled, snout and top of head naked. Scales 
usually thin, cycloid, deciduous, but very 
weakly ctenoid in some nomeids and 
Schcdophilus medusophagtis, and heavy, 
keeled, and adherent in Tefragonurus. 
Scales usually covering bases of median 
fins. Lateral line present, the scales with 
simple tubes, except no tubed scales in 
Tetragonurus. No bony scutes or keels as- 
sociated with lateral line. Usually a well 
developed subdennal mucous canal system 
communicating to the surface through small 
pores liberally scattered over head and 
body. Eyes small to large, lateral, not enter- 
ing into profile of head. Nostrils double, 
the anterior round, the posterior usually a 
vertical slit. Jaw teeth small, simple or 
minutely cusped, arranged more or less in 
a single series, close-set or spaced. Teeth 
on vomer, palatines, and basibranchials 
present or absent. No teeth on entoptery- 
goid or metapterygoid. Small teeth usually 
present on inner edge of gill-rakers. Gills 
4, a slit behind the fourth. Gill-rakers 10 
to 20 on lower limb of first arch. Well de- 
veloped pseudobranch usually present, but 
absent in Pampus; rudimentary gill-rakers 
under pseudobranch commonly present. 
Gill-membranes usually free from isthmus, 
but united in Pampus. 

Skeleton never strongly ossified. Epiotic 
forked, orbitosphenoid absent, 15 principal 
branched rays in caudal fin. Pelvic fin, 
when present, with one spine, five rays. A 
bony bridge partially covering the anterior 

vertical canal of the ear. Opercular bones 
thin, denticulate or entire, never with strong 
spines, except moderate preopercular spines 
in Schedophilus. No bony stay for the pre- 
opercle. Five to seven branchiostegal rays. 
Lacrimal bone usually enlarged, often cov- 
ering maxillary almost completely. Lacri- 
mal absent in Pampus. Premaxillary scarcely 
if at all protractile. A slender supramaxil- 
lary present or absent. Pelvic bones not 
finnly attached to coracoid. Vertebrae 25, 
26, or 29 to 60, including hypural. Caudal 
skeleton with two to six hypural elements, 
two or three epural elements, two auto- 
genous haemal spines, except three in 
Icichthys, hypuropophysis present on first 
hypural. Sometimes two but usually three 
free interneurals ahead of dorsal fin, but 
six or more in Icichthys. Lower pharyngeal 
bones not united, partially supporting pha- 
ryngeal sacs. In the sacs, numerous simple 
teeth on irregularly-shaped or stellate bony 
bases [= papillae] seated in the muscular 
walls, arranged in longitudinal bands or 

Distribution. All stromateoid fishes are 
marine. They are found in the three major 
oceans of the world on the high seas, over 
the continental shelves, and in large bays. 
Most species live in tropical and temperate 
waters, but a few occur in colder areas. No 
stromateoids have been reported from the 
Arctic Ocean, the Bering Sea, the Okhotsk 
Sea, the Baltic Sea, the Black Sea, and the 
Antarctic Ocean. 

Natural history. The association between 
stromateoid fishes and medusae or siphono- 
phores is widely known and well docu- 
mented. Mansueti (1963) has reviewed the 
literature concerning this unusual associa- 
tion. His lists of all fish species involved 
is made up largely of stromateoids, and in- 
cludes centrolophids, nomeids, tetragonu- 
rids, and stromateids. Only one ariommid 
has been reported from under a jellyfish, a 
.36-mm Ariomma indica taken in a 305-mm 
diameter ctenophore in Durban Bay, Natal 
(Fowler, 1934a). 

46 BiiUctin Museum of Couipaidtive Zoology, Vol. 135, No. 2 

Plate 1. A young stromateoid, Psenops/s onomo/o, under a medusa. Iwago photo. 

Young stromateoids are pelagic, and it is living host, and descend to deeper layers, 

not surprising that it is the young stages the adult habitat. Stromateoids also hover 

that are found with jellyfishes. As they beneath flotsam and Sarf],os.s-iim weed 

grow older. th(> fishes desert their surface- ( Besednov, 1960). It is this charactf^istic 

Stromateoid Fishes • Hacdrich 


J 100 120 140 

200 220 








"T I 1 1 1 




1 I 

1 1 1 

I I i 1 1 1 


, * 


• . ' • ••*, 

' ' 

1 ' 

1 1 1 


1 1 1 1 ! 1 

1 I - , 

1 I ■-] 

1 1 



•• • 





— 1 1 1 1 

1_ .1 1 1 

1 1 

1 • 


Figure 3. Scatter diagram of the allometric growth in the 
nomeid Nomeus gronovn. 

habit which gives the barrelfish, Hypcro- 
gjyphe, its common name. Young stromat- 
eoids typically have a banded pattern, 
whereas adults tend to be plain. Undoubt- 
edly, the banding is protective coloration 
for the fishes during that period of their 
lives when they live in the shifting shadows 
under a jellyfish. 

Shelter is not all the jellyfish provide. 
Many stromateoids have been observed 
actively feeding on their hosts {Schedo- 
philus. Maul 1964; Nomem, Kato, 1933; 
Peprihis, Mansueti, 1963, photo p. 63). The 
diagnostic pharyngeal sacs of stromateoids 
may have been perfected partially in re- 
sponse to this sort of diet. Tetragonurus, 
one of the most highly specialized stromat- 
eoids with great slicing teeth and a long 
pharyngeal sac, may feed almost entirely 
on soft-bodied medusae, ctenophores, and 
salps (Grey, 1955). 


60 80 100 120 140 



S 35 




S 551- •'■ 

"1 I I 




Figure 4. Scatter diagram of the early growth in the soft- 
spined centrolophid Schedoph/lus medusophogus, a fish 
which grows to 500 mm SL. 

Stromateoids do show high resistance to 
the toxins of jellyfish (Lane, 1960; Maul, 
1964 ) but they are by no means immune to 
it (Garman, 1896; Totton, 1960). Besides 
the relatively high resistance to the toxins, 
simple avoidance of the tentacles and the 
characteristic hea\'y coating of slime prob- 
ably are important in allowing the fishes 
to swim with impunity under their hosts. 

Because of the efficient shredding of food 
by the papillae, stomach contents of stro- 
mateoids are largely unidentifiable. Shred- 
ded transparent tissues, probably from jelly- 
fish, ctenophores, and salps, predominated 
in stomachs examined. But I have also 

48 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 




60 80 

100 120 140 160 




1 1 

1 1 1 1 










.* '•'*•'•• 







1 1 

1 1 1 1 




1 r 

• • • ** 

"T 1 1 r 


J I I L- 

>u 55 


. , 


1 1 


1 1 1 1 


•' 1 


1 1 



1 1 1 1 

T 1 1 r 


■•• <: . 



1 1 

1 1 1 




1 1 

1 1 1 




1 1 

1 1 1 1 1 






1 1 



1 1 1 1 1 


40 60 80 100 120 140 160 180 


Figure 5. Scatter diagram of the growth in the hord-spined 
centrolophid Psenops/s onomala. 

found remains of fishes, large crustaceans, 
and, rarely, squids. 

With growth, marked changes occur. The 
fish are no longer in the immediate upper 
layers, but tend to live deeper in the water. 
The typically banded pattern of the young 
fish gives way to the plainer colors of the 
adult. The most pronounced changes are 
in relative proportions. Allometry is the 
rule among stromateoids. Generally, the 
relative length of the head, length of the 
pelvic fins, predorsal distance, and preanal 
distance increase rapidly, then decrease 
with growth. The relative length of the 
pectoral fin and the maximum depth may 
increase steadily, or increase and then de- 
crease. Allometry is especially marked in 
nomeids (Fig. 3), most of which have a 

similar pattern of growth, and centrolophids 
(Figs. 4, 5). The growth curves given for 
Tetraiionurm (Grey, 1955) are fairly regu- 
lar, and rarely show a range of variation ex- 
ceeding ten per cent. McKenney's ( 1961 ) 
curves for Ariomma, perhaps based on two 
species, likewise show regular and little 
variation. The pectoral fin of the stroma- 
teids may relatively lengthen or shorten 
with age and the fin lobes may be very 
long in the young, but the allometry is in 
general not remarkable in this family. 

The largest stromateoids are the centro- 
lophids, many of which reach three feet in 
length. A large CentroJophus or Hijpero- 
glyphc will exceed four feet. A few no- 
meids attain two feet, but most are smaller. 
There is a diminutive species group in the 
family; its members rarely exceed six inches. 
Tetragonurids are said to reach two feet in 
length (Fitch, 1951). The largest reported 
ariommid was almost three feet long 
(Klunzinger, 1884), but most seem to be 
much smaller. A large stromateid barely 
exceeds 18 inches. 

Stromateids, the subjects of substantial 
fisheries in the eastern United States, India, 
China, and Japan, and ariommids, with 
commercial potential but as yet unfished, 
are schooling fishes. The centrolophids 
Psenopsis and Hijperoiijijphe, commercial 
fishes in Japan, and Seriolella, fished to a 
lesser extent in Peru, Ghile, and Australia, 
are also found in shoals. Numerous young 
specimens of the more oceanic stromateoids 
are often taken by dipnet collecting, but 
data is too sparse, and large specimens too 
few, to know whether these fishes occur in 
schools or not. 

Little is known of the breeding of stro- 
mateoids. The eggs are pelagic. Those of 
Vcprihis triacanthiis are described as buoy- 
ant, transparent, and containing a large oil 
globule (Bigelow and Schroeder, 1953). 
All species seem dioecious. There is some 
evidence that the sexes may be slightly di- 
morphic, with respect to color and or rela- 
tive proportions. 

Rclatiunships. The stromateoid fishes are 

Stromateoid Fishes • Haedrich 49 

a well-defined unit. Pharyngeal sacs are 
the diagnostic character. These are present 
in all species, and are readily apparent on 
dissection. A somewhat similar organ is 
found in Dorosoma, a clupeoid (Miller, 
1964), and in Trisotrophis, a serranid (Kata- 
yama, 1959). 

Within the group, there is a broad spec- 
trum from primitive to highly advanced 
forms. Between existing families, there is 
evidence of phyletic relationships. The 
natural coherence of the group and the ten- 
dency toward direct internal lines from the 
generalized to the specialized condition, 
makes it unlikely that the stromateoids have 
given rise to other groups. Tetragominis, a 
highly specialized form, is sometimes placed 
in a suborder of its own, in which case it 
would be considered a derived group. 
Tetragomirus is certainly far removed from 
the mainstream of stromateoid evolution, 
but its degree of divergence is probably not 
great enough to warrant subordinal recog- 
nition. It retains the characteristic sacs, and 
is here considered the sole genus of the 
stromateoid family Tetragonuridae. 

Giinther (1880) considered the stromat- 
eoids a subdivision of the scombroids. 
Little subsequent attention has been paid 
to the relationships of the suborder, with 
the exception of Boulenger (1904) and Holt 
and Byrne ( 1903 ) who found cause for in- 
cluding them in the Percesoces. General 
classifications have continued to place them 
near the Scombroidei. There is a close re- 
semblance between some stromateoids and 
the carangids, a group standing near the 
base of the scombroid stem ( Suzuki, 1962 ) . 

Freihofer's ( 1963 ) survey of patterns of 
the ramus lateralis accessorhis (a facial 
nerve complex) in fishes indicated that the 
stromateoids might be related to entirely 
different groups. His pattern-10 teleosts 
are a novel assemblage, composed of stro- 
mateoids ( centrolophids, nomeids, stroma- 
teids), pomatomids, kyphosids, scorpidids 
(excluding Scorpis), arripidids, girellids, 
theraponids, and kuhliids. The nematistiids 
have a reduced pattern 10. 

The pattern of a nerve complex, because 
of its basically conservative nature, should 
be a strong taxonomic character in dealing 
with higher categories. The common pat- 
tern of the ramus lateralis aecessoriiis in the 
above families is probably good cause for 
considering them a phylogenetic unit. But 
is there additional evidence for lumping 
these families together? The stromateoids, 
almost without exception, have a bony 
bridge across the inside of the anterior ver- 
tical canal of the ear. Because of its wide- 
spread occurrence in the diverse forms of 
the suborder, I consider this a conservative 
character, useful at higher categories. This 
bridge is also present, at least in some spe- 
cies and at some stage of growth, in pat- 
tern-10 families. The bridge is absent, how- 
ever, in other perciform families examined 
(see Table 1 and Material examined, p. 43). 

Many pattern-10 families have character- 
istics common to basal perciforms. Many 
have 25 vertebrae, seven branchiostegal 
rays, a suborbital shelf, and a caudal skele- 
ton near to the basal perciform type with 
six hypurals and three epurals. In most, 
the shape of the body is of the most gen- 
eralized type. The general impression is of 
an older group of fishes which have man- 
aged to remain successful without di- 
verging too far from the basal stock. 

The common ramus lateralis accessorius 
pattern and the bridge over the anterior 
vertical canal are strong evidence for 
considering that the relatively specialized 
stromateoids arose from somewhere in 
this relatively undifferentiated assemblage. 
Nonetheless, it is unlikely that the stromat- 
eoids are the descendents of any living pat- 
tern- 10 family, all of which are specialized 
in some respect. With the present imper- 
fect knowledge of the comparative osteology 
of these groups, the best that can be said 
is that all share a common heritage. 

The fin spines of stromateoids are not 
remarkably developed, and the teeth are 
uniserial in the jaws. Pattern-10 families 
with moderate-to-weak fin spines and a 
major row of uniserial teeth ( some have 

50 Bulleiin Museum of Comparative Zoology, Vol. 135, No. 2 

Table 1. Selected characters of some teleostean families. + denotes presence; — absence. 

RLA PatternI 

AVC Bridge 

















































reduced 10 































reduced 9 





* not examined 

1 From Freihofer 1963 

- Largely from Regan 1913, and Berlin and Arambourg 1958 

" From Smith and Bailey 1962 

^ Largely from Giinther 1859, 1860, 1861 

very reduced rows behind the major row) 
are the pomatomids, kyphosids, and scorpi- 
dids. These could be the groups most 
closely associated with the line leading to 
the stromateoids. The arripidids, girellids, 
theraponids, and kuhliids have teeth in 
bands and heavy fin spines. 

There is a strong resemblance between 
the kyphosids and the centrolophids, the 
primitive stromateoids. In both, the general 
rule is 10 + 15 vertebrae, a caudal skeleton 
with six hypural and three epural elements, 
a perforate ceratohyal, an expanded lacri- 
mal, and scaled fin bases. Behind the major 
row of uniserial teeth in KypJio.'iUs, there 
are rows of small villiform teeth ( Moore, 
1962); there is only one row of teeth in 
centrolophids. The kyphosids, however. 

lack the subocular shelf found in some 

The kyphosids are today one of the most 
primitive perciform families. They retain 
the teeth on the ectopterygoid and endo- 
pterygoid and the two foramina in the fa- 
cial wall of the ])ars jufi,ularis, conditions 
associated with the bcrvciform level. Pat- 
terson (1964) offers evidence for the deriva- 
tion of the kyphosids, scoq^idids, and mono- 
dactylids from the Cretaceous polymixioid 
Benjcopsis. If this is the case, and if the 
stromateoids arose from near the kyphosid 
stem, a direct line can be drawn from the 
berycoid level to Pampiis, one of the most 
advanced perciforms. 

The carangids were probably derived 
from the Cretaceous dinopterygoid Aipich- 
thys (Patterson, 1964). Their development, 

Stromateoid Fishes • Haedrich 


Figure 6. Parastromateus niger, drawing of a 15-inch specimen, from Day, 1875. 

from ber\'ciform to perciform level, has 
been independent of the line which pro- 
duced the stromateoids. The resemblances 
between certain members of these two 
groups must therefore be considered an 
evolutionary convergence. 

A problem remains in the genus Para- 
stromateus {= Apolectus) (Fig. 6). Bloch 
( 1795 ) described the sole representative of 
the genus as a species of Stromateus. The 
shape of the body and the small pelvics 
which are lost with age were good cause 
for this action. But Parasfromatcus lacks 
pharyngeal sacs and, because of this, Regan 
( 1902 ) removed the genus from the stro- 
mateoids and placed it in the carangids. 
More extensive comparisons by Apsangikar 
( 1953 ) supported the separation, but diver- 
gence from the carangids was noted and 
the new subfamily Parastromateinae of the 
Carangidae proposed. Suzuki ( 1962 ) , in 
his great review of Japanese carangids, fol- 
lowed Jordan (1923) in considering Paro- 
stromateus the sole representative of a 
monotypic family. 

Parastromateus is a pattem-9 teleost, as 
are the carangids (Freihofer, 1963), but 
has a bony bridge over the anterior vertical 
canal of the ear, as do the stromateoids. 
The only reason, other than the bony 
bridge, for relating Parastromateus to the 
stromateids is the similarity in body form. 
Parastromateus, however, has 10 + 14 ver- 
tebrae, while all stromateids have at least 
13 + 17. Even if Parastromateus is related 
to the stromateids, the relationship is at 
most a very distant one. 

Within the suborder Stromateoidei, there 
are three distinct groups, the primitive 
centrolophids, the intermediate nomeids 
with their specialized off-shoots the tetra- 
gonurids and the ariommids, and the ad- 
vanced stromateids. The stromateids are an 
obvious derivative of the centrolophid line. 
The nomeids, an evolutionary grade above 
the centrolophids, have evolved parallel to 
the centrolophids, but from an earlier com- 
mon ancestor. The probable relationships 
between the stromateoid families, discussed 

52 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



Figure 7. Dendrogram showing probable relationships of 
the five stromateoid families. 

more fully in the family accounts, are ex- 
pressed diagrammatically in Figure 7. 

Fossils. The Cretaceous Omo.soma, usu- 
ally considered a stromateid (Arambourg, 
1954 ) has been shown by Patterson ( 1964 ) 
to be a polymixioid, standing, interestingly 
enough, near Bcnjcopsis. Camnp,odcs ceph- 
alus, from the Monte Bolca beds of Austria, 
is well described and figured by Heckel 
( 1856 ) . Though it looks somewhat like 
some stromateoids and was considered a 
nomeid by Jordan (1923), the diagnostic 
characters are missing and it cannot be af- 
filiated with this group with any certainty. 
AspidoJepis Geinitz 1868, based on a scale, 
was considered a stromateid by Jordan 
(1923). But the scales of the majority of 
stromateoids are in no way distinctive, and 
thus the possible relationships of this fossil 
genus cannot be determined. Two new 
fossil genera have been found by Bonde 
(1966) in the lower Eocene Mo-clay of 

Key tu Stronuiteoid Families 

1 (6j. Two dorsal fins, distinctly, though 
scarcely, separated, the first usually 
with ten to twenty spines; if there are 
fewer than ten spines, the longest 
spine is about the same length as the 
longest dorsal finray. Pelvic fins al- 
ways present. Vomer, palatines, and 
basibranchials toothed or not. 2 

2 (5). The first dorsal fin with about ten 

long, slender spines, often folded into 
a groove, the longest spine nearly as 
long as, or longer than, the longest 
finray in the second dorsal. Anal fin- 
rays 14 to 30. Scales cycloid, thin, 
deciduous. Fleshy lateral keels on 
peduncle near caudal fin base absent 
or only slightly de\eloped. Vertebrae 
29 to 42 -- 3 

3 (4). Vomer, palatines, and usually basi- 

branchials with small, often almost 
indistinguishable, teeth. Caudal pe- 
duncle compressed, its least depth 
greater than 5 per cent of the stan- 
dard length, without lateral keels. 
Usually more than fifteen rays in both 

the dorsal and anal fins. 

NOMEIDAE, p. 76 

4 (3). Vomer, palatines, and basibranchials 

toothless. Caudal peduncle square in 
cross-section, its least depth less than 
5 per cent of the standard length, 
with two low lateral keels on each 
side near caudal fin base. Fourteen or 
fifteen rays in both the dorsal and 
anal fins. ARIOMMIDAE, p. 88 

.5 ( 2 ) . The first dorsal fin with ten to twenty 
short spines, the longest only half the 
length of the longest finray in the 
second dorsal. Anal finrays 10 to 16. 
Scales keeled, heavy, very adherent. 
Modified scales form two well de- 
veloped lateral keels on each side of 
peduncle near caudal fin base. Ver- 
tebrae 43 to 58. 


6(1). A continuous dorsal fin, or two dor- 
sal fins scarcely separated, the first 
with less than ten spines; if spines are 
present, the longest spine is less than 
half the length of the longest dorsal 
finray. Pelvic fins present or absent. 
Vomer, palatines, and basibranchials 
toothless. 7 

7 (8). Pelvic fins always present. None or 

one to five weak spines, or five to 
nine stout spines precede dorsal fin- 
rays. Anal finrays 15 to 30. Median 
fins never falcate; their bases rarely 
the same length. Jaw teeth all conical, 
simple. Supramaxillary bone usually 
present, but hard to find in some. 
Seven branchiostegal rays. Vertebrae 

25 to 30 or 50 to 60. 


8 (7). Pelvic fins never present in adults, 

rarely present in the young. No stout 
spines precede dorsal finrays, but, in 
.some species, five to ten small blade- 

Stromateoid Fishes • Haedrich 


like spines resembling the ends of free 
interneurals protrude ahead of the fin. 
Anal finrays 30 to 50. Median fins 
often falcate; their bases about equal 
in length. Jaw teeth laterally com- 
pressed, with three cusps. No supra- 
maxillary bone. Five or si.x branchi- 
ostegal rays. Vertebrae 30 to 48. __.. 



Type genus: Cenfrolophus Lacepede 1803 

Centrolophes. Cuvier and Valenciennes, 1833:330 

( descr. ) . 
Centrolophinae. Gill, 1861:34 (list); 1862:127 
(genera listed); 1884:666-667 (def., gen.). 
Jordan and Gilbert, 1882:450 (name, descr.). 
Centrolophidae. Jordan and Evermann, 1896: 
962 (descr.; North America). Jordan, 1923: 
182 (in part, list). Nomian, 1937:115 (descr., 
relationships; Chile). Tortonese, 1959:57 (in 
part, revision; Gulf of Genoa). 
Lirinae. Biihler, 1930:62 (in part, morph., diges- 
tive system ) . 
Nomeidae. Berg, 1940:323 (in part, dist.); 1955: 
248 (part, dist.). Norman, 1957:503 (in part, 
def., genera listed). 
Diagnosis. Stromateoid fishes with pelvic 
fins present in adults, continuous dorsal fin, 
toothless palate, seven branchiostegal rays, 
and six hypural bones in the tail. The papil- 
lae in the pharyngeal sacs with irregularly 
shaped bases, arranged in ten to twentv' 
longitudinal bands. 

Description. Body slender to deep, usu- 
ally somewhat compressed. The rayed por- 
tion of the continuous dorsal fin preceded 
by six to eight short stout spines in Hijpcro- 
ghjphe, Seriolella, and Psenopsis; none or 
one to five thin weak spines in Cenfro- 
lophus, Schedophihis, and Icichtht/s. In the 
latter group and in Psenopsis the spines 
graduating to the rayed portion of the fin; 
in the others not. Three anal fin spines, 
not separated from the rays. Pelvic fins 
usually attached to the abdomen by a thin 
membrane and folding into a broad shallow 
groove. Head conspicuously naked, usually 
covered with small pores. Scales cycloid, 
but with minute cteni in SchedopJiiJiis 
medusophagus, and usually deciduous. 
Tubed scales of lateral line extending onto 
peduncle. Margin of preopercle usually 

moderately denticulate, but spinulose in 
most young stages and in SchedophiJus. 
Opercle thin, with two flat, weak spines; 
the margin denticulate. Seven branchios- 
tegal rays. Mouth large, maxillary extend- 
ing at least to below eye. A nearly uniserial 
row of small conical teeth in the jaws; 
vomer, palatines, and basibranchials tooth- 
less. Supramaxillary bone present in most 
but absent in Psenopsis. Adipose tissue 
around eye usually not conspicuously de- 
veloped. Vertebrae 25 or 26 in most spe- 
cies, except 50 to 60 in Icichthys. Caudal 
skeleton with six hypurals and usually three 
epurals, except two in Icichthys. Pharyn- 
geal sacs with irregularly shaped papillae 
in ten to twenty longitudinal bands. Teeth 
seated directly on top of the bony base. 
Adults one to four feet in length, usu- 
ally dark-colored and without conspicuous 

Distribution. Centrolophids are pelagic, 
usually on the high seas and over the edge 
of the continental shelves, although Psenop- 
sis and SerioJeUa occur in shallow water 
near the coast. Some are found in tropical 
waters, but the majority are fishes of tem- 
perate seas. The soft-spined centrolophids 
(Cenfrolophus, Icichthys, and SchedopJjihis) 
tend to be more oceanic than the hard- 
spined centrolophids (Hyperogh/phc, Seri- 
olella, and Psenopsis) . To some extent, the 
distributions of these two groups comple- 
ment each other (Figs. 52, 53). 

The distribution of the centrolophids is 
in part a relict distribution. Cenfrolophus 
is bipolar, found in the North Atlantic, 
South Africa, and Southern Australia and 
New Zealand. Icichthys, very similar in 
appearance to Cenfrolophus, is bipolar in 
the Pacific. In the waters from Australia to 
the coasts of Chile, the endemic genus 
Seriolella has evolved. 

No centrolophids occur across the broad 
tropical Pacific or Indian Oceans. 

Relationships. The Centrolophidae stand 
at the base of the line leading to the Stro- 
mateidae. Of all stromateoids, they are the 
least differentiated from the percifomi an- 

54 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

cestor. Most have 25 vertebrae, the well- 
known basic perciform number. Most have 
a snpramaxillary bone. There are seven 
branehiostegal rays, and the caudal skeleton 
(Figs. 10, 12, 23) is of the basic perciform 
type (Gosline, 1961a). In this sense, the 
centrolophids can be considered the most 
primitive stromateoids. 

The caudal skeleton of the nomeids, with 
hypurals 2 + 3 and 4 + 5 of the basic six 
fused, could easily have been derived from 
the centrolophids. The stellate papillae, 
also, and loss of one branehiostegal repre- 
sent a grade above the centrolophid condi- 
tion. But teeth are present on the vomer, 
palatines, and basibranchials of the no- 
meids, in general a primitive condition 
(Liem, 1963), and are absent in the cen- 
trolophids. This makes it unlikely that the 
former group has been derived from the 
latter. Rather, the two must represent ap- 
proximately parallel lines, derived from an 
earlier form which had palatal dentition. 

There is a fairly close and probably direct 
relationship between the advanced centro- 
lophid genera SeholcUa and P.scnopsis and 
the stromateids Stromoteus and Pcprilus. 
Fishes in both these groups have well- 
ossified sclerotic bones, minute body pores, 
slender tapering branchiostegals, and ex- 
tremely deciduous scales. 

Key to Centrolo))hid Genera 

1 (6). Spines of the dorsal fin weakly de- 

veloped and all graduating to the 
dorsal rays. 2 

2 (5). Weak dentieulations on preopercular 

margin. Origin of dorsal fin usually 
well behind insertion of pectoral fins, 
])ut over pectoral insertion in very 
small specimens. Body elongate, maxi- 
mum depth usually less than 30 per 
cent of the standard length. 3 

3 (4). Total elements in anal fin 23 to 27. 

Scales small, very deciduous, pre- 
opercle and cheek naked. Scales in 
lateral line 160 to 230. Vertebrae 
25 Centrolophu.s, p. 62. Fig. 13 

4 (3). Total elements in anal fin 27 to 31. 

Scales moderate in size, not especially 
deciduous, present on preopercle and 
cheek. Scales in lateral line 100 to 

130. Vertebrae 50 to 60. 

Icichfhtjs, p. 65. Fig. 15 

5 (2). Nine to fifteen small spines on pre- 

opercular margin. Origin of dorsal 
fin usually before insertion of pec- 
toral fins, but over pectoral insertion 
in very large specimens. Body deep, 
maximum depth usually greater than 
.35 per cent of the standard length. __._ 
_. Schedophihi.s, p. 58. Fig. 11 

6 ( 1 ) . Five to nine stout dorsal spines, 

shorter than and not graduating 
(graduating slightly in Psenopsis) to 
the dorsal rays. 7 

7 (8). Dorsal finrays 19 to 25; anal finrays 

14 to 21. Preopercular margin spi- 
nulose. Scales not especially decid- 
uous. Lateral line arched anteriorly, 
straightening out over the anal fin. 
Adipose tissue around eye not well de- 
veloped. Sclerotic bones not well 
ossified; golden iris appears as a com- 
plete ring. - Hyperoghjphe, p. 54. Fig. 8 

8 (7). Dorsal finrays 25 to 40; anal finrays 

18 to 30. Preopercular margin entire 
or finely denticulate. Scales very de- 
ciduous. Lateral line follows dorsal 
profile. Adipose tissue around eye 
well developed. Sclerotic bones usu- 
ally well ossified; golden iris appears 

divided by a vertical bar. 9 

9(10). Insertion of pelvic fins behind inser- 
tion of pectorals. Supramaxillary bone 
present. At least seven more dorsal 
finrays than anal finrays. Usually 
eight dorsal spines, the third, fourth, 

and fifth the longest. 

Seriolella, p. 69. Figs. 18, 19 

10 (9). Insertion of pelvic fins before or just 
under insertion of pectorals. Supra- 
maxillary bone absent. Number of 
dorsal finrays never exceeds number 
of anal finrays by more than five. 
Five to seven dorsal spines, increasing 

in length posteriorly. 

Psenopsis, p. 72. Fig. 21 

Genus HYPEROGLYPHE Gunther, 1859 
Figure 8 

Palinurus DeKay, 1842:118. (Type species: Cory- 
phcunia pcrciformis Mitchill, 1818:244, by 
monotypy. New York Harlior. Preoccupied 
by Pulinurns Fabricius, 1798, Crustacea.) 

Hypero^hjphc Ciinther, 1859 (June):337. (Type 
species: *Diagramnia porosa Richardson, 1845: 
26, l)y monotypy. Coasts of Australia. A syn- 
onym of Perca antarctica Carmichael, 1818: 

Stromateoid Fishes • Hacdrich 55 

Figure 8. Hyperoglyphe perciiorma, drawing of an approximately 200-mm specimen, courtesy of the Smitfisonian Institution. 

Palimirichthys Bleeker, 1859 (November): 22. (Sub- 
stitute name for Palinurus DeKay, and there- 
fore taking the same type species, Conjphacna 
perciformis Mitchill, 1818:244.) 

Palimirichthys Gill, 1860:20. (Substitute name, 
proposed independently from Bleeker, for 
Palinums DeKay, and therefore taking the 
same type species, Coryphacna perciformis 
Mitchill, 1818:244.) 

PammcJas (Uinther, 1860:485. (Substitute name 
for Palinurus DeKay, and therefore taking the 
same type species, Coryphacna perciformis 
Mitchill, 1818:244.) 

Eurumetopos Morton, 1888:77. (Type species: 
Eurumetopos johnstonii Morton, 1888:77, by 
monotypy. Tasmania. A synonym of Perca 
antarctica Camiichael, 1818:501.) 

Tolcdiu Miranda-Ribeiro, 1915:5. (Type species: 
Toledia macrophihalma Miranda-Ribeiro, 
1915:5, by monotypy. Macahe, Brazil.) 

Ocycrius Jordan and Hubbs, 1925:226. (Type spe- 
cies: Centrolophus joponicus Doderlein in 
Steindachner and Doderlein, 1885:183, by 
original designation. Tokyo, Japan. ) 

The combination of less than 25 dorsal 
finrays, about eight short spines not in- 
creasing in length to the rays in the dor- 
sal fin, toothless palate, pelvic insertion 
under pectoral fin base, supramaxillary 
bone present, and lateral line arched an- 
teriorly straightening out over the anal fin, 
distinguishes Hyperoglyphe from all other 

stromateoid genera. The name, a feminine 
noun, is from the Greek vvkfj, above, + 
y\v<f>yi, groove, in reference to the deep 
longitudinal groove in the roof of the 

Description. Body moderately deep, 
maximum depth around 30 to 35 per cent 
of the standard length; musculature firm. 
Caudal peduncle broad, of moderate length. 
Dorsal fin originating over or a little behind 
insertion of pectoral fins, continuous, six to 
eight short stout spines not graduating to 
the longer rays. The longest spine half the 
length of the longest ray. Anterionnost fin- 
rays the longest, those that follow shorter, 
19 to 25 finrays in all. Anus at mid-body, 
in a slit. Anal fin originating a little behind 
middle of body, three spines precede the 
15 to 20 rays. Pectoral fin rounded in the 
young, pointed in adult. Pelvic fins insert- 
ing under end of pectoral fin base, attached 
to abdomen by a small membrane and fold- 
ing into a shallow groove. Caudal fin broad, 
emarginate to moderately forked in adult. 
Scales cycloid, moderate in size, somewhat 
deciduous, covering bases of median fins. 
Lateral line arched anteriorly, straightening 
out over middle of anal fin and extending 


Bulletin Miisciini of Comparative Zoology, Vol. 135. No. 2 

onto peduncle. Skin moderately thick; ex- 
tensive subdermal canal system communi- 
cating to the surface through small pores. 
Head around 33 per cent of the standard 
length, broad. Top of head not scaled, 
pores prominent, naked skin projecting 
backward over nape. Eye moderate to 
large, no adipose tissue. Nostrils located 
near tip of obtuse snout, large, the anterior 
round, the posterior a slit. Angle of gape 
extending below eye. Premaxillary not pro- 
tractile. Lacrimal bone partially covering 
anterior portion of upper jaw when mouth 
is closed, end of maxillary remaining ex- 
posed. Supramaxillary present. Jaw teeth 
very small, pointed, uniserial, close-set; 
vomer, palatines, and basibranchials tooth- 
less. Opercle and preopercle thin; opercle 
with two weak flat spines, scaled, margin 
very finely denticulate or entire; preopercle 
not scaled, striated, margin \\'ith numerous 
very small spinules. Angle of preopercle 
rounded, bulging backward slightly. Gill- 
rakers heavy, slightly longer than the fila- 
ments, finely toothed on inner edge, spaced, 
about 16 on lower limb of first arch. Seven 
branchiostegal rays, five on the ceratohyal, 
two on the epihyal. Scapula visible. Verte- 
brae 10 + 15 = 25. Stomach a simple sac; 
intestine long. Pyloric caeca numerous, in 
a mass resembling a raspberry. 

Base color green-grey or blue-grey to 
reddish brown. Back dark, sides and be- 
low lighter, sometimes silvery. Head dark, 
iris a golden ring, opercle often silvery. 
Median fins usually darker than the body. 
Color pattern irregularly striped, mottled, 
or clear, changeable in life. Inside of mouth 
and gill cavity light. Peritoneum light with 
minute dark speckles. 

Natural history. Though Ilypcro^hjphc 
occurs throughout the world and is fished 
commercially in Japan, very little is known 
of its habits. The young commonly occur 
under flotsam, but usually not under jelly- 
fish, in surface waters near the edge of 
the continental shelf. The larger adults 
form shoals in deep water, perhaps fairly 
near the bottom. 

Figure 9. Branchial region of Hyperoglyphe percilorma, 
drawing of a cleared-and-stained preparation from a 173-mm 
SL specimen. Elements identified in Figure 2. 

Small H. ))crciforma two to four inches 
long occur off the New England coasts 
under floating objects in great numbers 
during the summer. By fall, these fish have 
doubled or even tripled their size. With 
the approach of cold weather they dis- 
appear. Only recently has it been found 
that adult H. pcrciforma attain three feet 
in length, and live in deep water off the 
coast of west Florida (Schwartz, 1963). This 
discovery bears out an earlier suggestion 
by Merriman ( 1945 ) that the fish observed 
off New England were the young of a much 
larger bathypelagic species. In Japan, the 
large adults had been marketed long before 
the young were first discovered (Abe, 1955). 

Bigelow and Schroeder ( 1953 ) reported 
small fishes and crustacean remains from 
stomachs of Hypcroiijiiplic pcrcifornia. The 
fish may also feed occasionally on barnacles 
(Cornish, 1874; Holt and Byrne, 1903). At 
times //. porosa feeds heavily on the tuni- 
cate Pyrosoma atlanticum (Cowper, 1960). 

Rclation.sJiips. Hyperoglyphe is the cen- 
tral genus of the Centrolophidae. The mem- 

Stromateoid Fishes • Hacdrich 57 



Figure 10. Caudal skeleton of Hyperoglyphe perciforma, drawing of a cleared-and-stained preparation from a 50-mm SL 
specimen. All elements identified in Figure 1. 

bers of this genus are the most generalized 
fishes in the entire suborder, and are prob- 
ably not unlike the ancestral form. The 
relatively low number of median finrays, 
the stout spines in the median fins, the 
seven blunt branchiostegal rays (Fig. 9), 
the 25 vertebrae, the spiny preopercle, and 
the large size attained, are all basal char- 
acters. The caudal skeleton (Fig. 10) is of 
the generalized perciform type. 

Hyperog,Iy])he has given rise, on the one 
hand, to the more oceanic soft-spined cen- 
trolophids, through Schedophilus to Cen- 
trolophus and Icichthys. The major change 

has been the softening of the fin spines and 
of the tissues in general. On the other hand, 
Hyperoglyphe has given rise to the more 
coastal, advanced, hard-spined genera Scri- 
olella and Psenopsis. The change in this 
direction has been one of slight refinement 
in the branchial region and a tendency 
toward fusion of elements in the caudal 

Species. Hyperoglyphe is a wide-ranging 
genus. The species are found in the slope 
water off the east coasts of the New World, 
in the Gulf of Mexico, near St. Helena and 
Tristan da Cunha, along the west coast of 

58 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Africa, in Australia-New Zealand, and in 

There is little problem of synonymy, since 
the species are fairh' distinct from one an- 
other. The changes that occur with growth, 
however, remain a stumbling block. The 
species in Hypew^h/phe are: 

Hyperoglyphc antarctica (Carmichael, 
1818) = Pcrco antarctica Carmichael. Tris- 
tan da Cunha, South Africa, southern Aus- 
tralia, and New Zealand, type locality 
Tristan da Cunha. D VIII, 19-21. A III 
15-16. P 18-20. Gill-rakers 5+1 + 14. 
Vertebrae 10 + 15. This is the most prim- 
itive species in Hypcro<jJyphc, and stands 
nearer to the base of the stromateoid stem 
than any other fish. It attains a very large 
size. McCulloch (1914) reports a specimen 
1072 mm long. The low median finray 
counts, large mouth, and a characteristic 
patch of scales on the otherwise naked 
occiput distinguish this species from all 
others. Synonyms are: '''Diaiiramma porosa 
Richardson, 1845, from Australia; Eiirume- 
topos johmtonii Morton, 1888, from Tas- 
mania; and Scriolella ampins Griffin, 1928, 
from Bay of Plenty, New Zealand. 

Uypcroiijyphc hythitcs (Ginsburg, 1954) 
= *PaJinurichfhys hythitcs Ginsburg. Gulf 
of Mexico, type locality off Pensacola, Flor- 
ida. D VII-VIII, 22-25. A III 16-17. P 
20-21. Gill-rakers 6-7 + 1 + 15-16. Verte- 
brae 10 +15. Possibly a synonym of //. 
macrophthalma (Miranda-Ribeiro, 1915). 
This species has more dorsal finrays and 
a larger eye than H. pcrciforma (Mitchill, 

Hypcro'j^lyphc pcrciforma (Mitchill, 1818) 
= Curyphaena pcrciformis Mitchill. East 
coast of North America, Florida to Nova 
Scotia, type locality New York Harbor. D 
VII-VIII, 19-21. A III 15-17. P 20-22. 
Gill-rakers 5-7 + 1 + 15-17. Vertebrae 10 + 
15 (skel.). This fish is the common "barrel- 
fish" of the offings of New England. Young 
specimens have followed floating logs 
across the Atlantic to the British Isles ( Holt 
and Byrne, 1903). Pimeleptenis cornu- 
hiensis Cornish, 1874, is a synonym based 

on a specimen which floated to Cornwall 
in a box. 

Hypcro'^lyphc japonica (Doderlein in 
Steindachner and Doderlein, 1885) = Ccn- 
troloplius japonicus Doderlein. Seas of Ja- 
pan, type locality Tokyo. D VIII, 22-24. 
A III 17-19. P 21-23. Gill-rakers 6-7 + 1 
+ 15-16. Vertebrae 10 + 15. Tliis fish is 
the "medai" of Japanese fisheries literature. 
It is the subject of a small, deep, hand-line 
fishery. A prol)able synonym is *Linis 
paiicidens Giinther, 1889, based on three 
small specimens captured by the CHAL- 
LENGER somewhere between New Guinea 
and Japan. 

Hyperoglyphc mosclii (Cunningham, 
1910) = Lcirus mosclii Cunningham. St. 
Helena, and coasts of Angola and South 
Africa, type locality St. Helena. D VI, 23- 
25. A II i 19-20. P 20-22. Gill-rakers about 
7 + 1 + 15. The type is described as having 
but one spine and 25 rays in the dorsal fin. 
The specimen is so large and heavy that I 
was unable to lift it and its container off 
the shelf in the British Museum, and hence 
did not get to examine it closely during 
my brief visit there. At such a large size, 
the first five spines in the dorsal may be 
buried in the skin, where Cunningham 
could have overlooked them. Probable 
synonyms are Palinurichthys pringlci Smith, 
1949, and PalinuriclitJu/s matthewsi Smith, 
1960, both from South Africa. 

Hyperoglyphc macrophthalma ( Miranda- 
Ribeiro, 1915) = Tolcdia macrophthalma 
Miranda-Ribeiro. Brazil, type locality Ma- 
cahe. D VII, 26. A 20 (from Miranda- 
Ribeiro, 1915). Known from a single speci- 
men 68 cm long. Possibly a synonym of 
//. mosclii (Cunningham, 1910). 

Genus SCHEDOPHILUS Cocco, 1839 

Fi.uure 11 

Lcirus Lowe, 1833:143. (Type species: * Lcirus 
hcnnettii Lowe, 1833:143, by monotypy. 
Madeira, Atlantic Ocean. Preoccupied by 
Leirus Dahl, 1823, Coleoptera. A junior syn- 
onym of *Ccnirolo])}ius ovalis Cuvier and 
Valenciennes, 1 833 : 346. ) 

Stromateoid Fishes • Haedrich 



_ii_ ^^'•. 

*V. ,.-••.-. ^V^Sig^ : l?:';? C- -4; ;v^ .-i^iv ■■;^- i'-iv^-C^S^-.SySfeB- 

Figure 11. Scfiedoph/lus pemarco, drawing of a 245-mm specimen, from Poll, 1959. 

Schedophihts Cocco, 1839^:57. (Type species: 
Schi'dophihis medusophagus Cocco, 1839:57, 
by monotypy. Messina. ) 

Mupiis Cocco, 1840': 237. (Type species: Mitpus 
imperialis Cocco, 1840:237, by monotypy. 
Messina. A synonym of *Centrolophus ovalis 
Cuvier and Valenciennes, 1833:346.) 

Lints Agassiz, 1846:213. (Emendation of Leirus 
Lowe, 1833:143, and therefore taking the 
same type species, *Leinis hennettii Lowe, 
1833:143, a junior synonym of *Ceniwlophus 
ovalis Cuvier and Valenciennes, 1833: 346.) 

Crhis Valenciennes, 1848:43. (Type species: *Crius 
bertheloti Valenciennes, 1848:45, l>y original 
designation. Canary Islands, Atlantic Ocean. 
A synonym of *Centrolophus ovalis Cuvier 
and Valenciennes, 1833:346.) 

Hoplocorijphis Gill, 1862:127. (Type species: 
*SchedophiIus maculatus Giinther, 1860:412, 
by original designation. Seas of China.) 

Eucrotiis T. H. Bean, 1912:123. (Type species: 
Eucrotus ventralis T. H. Bean, 1912:123, by 
monotypy. Bemiuda, Atlantic Ocean. ) 

Ti/ii^j/a Whitley, 1943:178. (Type species: Tuhbia 
tasmanica Whitley, 1943:179, by original 
designation. Eastern Tasmania.) 

The combination of deep body, broad 
deep head, large eye, continuous dorsal fin 
with weak spines graduating to the rays 

^ Tortonese (1959) has clarified the confusion 
surrounding the publication dates of Cocco's names. 

and originating before the pectoral inser- 
tion, toothless palate, and prominent spines 
on the preopercular margin distinguishes 
Schedophihis from all other stromateoid 
genera. The name, a masculine noun, is 
from the Greek axeSta, raft, + c^iAos, friend, 
in reference to the fish's common associa- 
tion with floating objects. 

Description. Body deep, maximum depth 
generally greater than 35 per cent of the 
standard length; musculature soft. Pedun- 
cle fairly broad, short. Dorsal fin originat- 
ing before (or over in very large specimens) 
insertion of pectoral fins, continuous, three 
to seven weak spines graduating to the 23 
to 50 rays. Anus and genital pore at mid- 
body, in a slit. Anal fin originating behind 
middle of body, three long weak spines pre- 
ceding the 16 to 30 rays. Median fins with 
compressed fleshy bases. Pectoral fin 
rounded in the young, pointed in adult, 
relative length decreasing with growth. 
Pelvic fins inserting under end of pectoral 
fin base, reaching to anus in young and 
juveniles, attached to abdomen by a mem- 
brane and folding into a shallow groove; 
relative length of fin decreasing markedly 

60 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

with growth. Caudal fin broad, forked, take small erustaceans. At a length of 

Scales small to moderate, cycloid, or with about 200 mm, S. meduso])Jia^us deserts 

one or two minute cteni in young Schedo- its coelenterate companion, and descends 

/)/n7n.9 mc'dusophuiius, deciduous, covering to deeper water. 

fleshy bases of the median fins. Lateral line Adult Scliedophilus appear very different 
arched anteriorly, straightening out about from the younger stages. The relative 
mid-body and extending onto peduncle, length of the paired fins is greatly de- 
Skin thin; extensive subdermal canal system creased, the body is much more elongate, 
communicating to the surface through small and the mottled or barred pattern, typical 
pores. Head soft, broad and deep, usually of juveniles, is gone. 

greater than 25 per cent of the standard Relatiun.ships. ScJicdophiJus provides the 

length, not scaled, naked skin projecting link between the soft-spined and the hard- 

slightly backward over the nape. Eye large, spined centrolophids. The range of varia- 

no adipose tissue. Nostrils located near tip tion in the genus is great, and the species 

of obtuse snout, anterior nostril round, the grade from the one condition to the other, 

posterior a slit. Angle of gape extending The caudal skeleton ( Fig. 12 ) is most like 

below eye. Premaxillary not protractile, that of Centrolophus and Icichthijs. The 

Lacrimal bone covering anterior portion of pharyngeal sacs and teeth are intermediate 

upper jaw when mouth is closed, end of between those of Centrolophus and Hy- 

maxillary remaining exposed. Slender supra- peroii.lyphe. Schedophilus ovalis has fairly 

maxillary present. Jaw teeth very small, stout spines ahead of the median fins; in 

pointed, uniserial, close-set; vomer, pala- S. mcdusophagus the spines are soft and 

tines, and basibranchials toothless. Opercle flexible. 

and preopercle thin; opercle with two weak Schedopluhi.'i is derived from the central 

flat spines, scaled, margin denticulate; pre- Jhjpero'^Jyphc stock. As it has moved into 

opercle not scaled, margin set with nine to a more oceanic environment, the spines on 

eighteen prominent spines, angle of pre- the preopercle have become more pro- 

opercle rounded, bulging back slightly. Gill- nounced, while the fin spines and the body 

rakers heavy, about half the length of the in general have become softer, 

filaments, toothed on inner edge, spaced; Species. The species in Schedophilus are 

10 to 16 on lower limb of first arch; a few in general well differentiated. Almost all 

rudimentary rakers present under large descriptions are based on young specimens, 

pseudobranch. Seven branchiostegal rays. The large adults differ greatly in appear- 

five on the ceratohyal, two on the epihyal. ance from the young. Adults are so very 

Scapula visible. Vertebrae 10 + 15, 16 or rarely seen that only three have entered 

20 = 25, 26 or 30, or 12 + 17 = 29. Stomach the literature, two of them assigned to other 

a simple sac; intestine long. Pyloric caeca genera. Unfortunately, the species from the 

numerous, dendritic. Australian region are very poorly known. 

Base color brown, bluish, or silvery. Me- Because of their isolated geographic dis- 

dian fins, pectorals, and pelvics usually tribution, critical examination of these spe- 

darker than the body. Color pattern irreg- cies will doubtless provide much insight 

ularly striped, mottled, or clear. Young of into the evolution of the soft-spined centro- 

some have dark vertical stripes. lophids. 

Natural history. Most species in ScJwdo- The species in the genus are: 

/j/nVn.s' are oceanic, rare, and, consequently, Schedophilus ovalis (Cuvier and Valen- 

little is known concerning their biology, ciennes, 1833) = ^Centrolophus ovalis Cu- 

The young of S. medusophw^us occurs vier and Valenciennes. Eastern Atlantic 

commonly with jellyfish. The fish may Ocean from Spain to South Africa and 

feed very largely on medusae, but will also Mediterranean Sea, type locality Nice. D 

Strom ATEOiD Fishes • HaedricJi 61 



Figure 12. Cauda! skeleton of Schedophilus medusophagus, drawing of a cleared-and-stained preparation from a 39-mm SL 
specimen. All elements identified in Figure 1. 

VI-VIII, 30-32. A III 20-24. P 21-22. Gill- 
rakers around 6+1 + 16. Vertebrae 10 + 
15. Silvery to greenish. Synonyms are: 
CentwJopluis crassus Cuvier and Valen- 
ciennes, 1833, from west of the Azores; 
*Leirus bennettii Lowe, 1833, from Ma- 
deira; Mupiis impcrkiUs Cocco, 1840, from 
the Mediterranean; *Crins- bertheloti Valen- 
ciennes, 1848, from the Canary Islands; 
Centrolophiis rotund icauda Costa, 1866, 
from Naples; Centrolophus porosissimus 
Canestrini, 1865, and Schedophilus bottcri 
Steindachner, 1868, from Barcelona. 

Schedophihis medusophagus Cocco, 1839. 
Atlantic Ocean and western Mediterranean 

Sea, type locality Messina. D 44-50 (total 
elements). A 28-31 (total elements). P 
18-21. Gill-rakers around 5 + 1 + 11. Ver- 
tebrae 10 + 15. Major preopercular spines 
usually about 12. Lateral line scales 160- 
230, increasing in number with growth. 
Brown, often mottled. The report of this 
species from the South Pacific (Giinther, 
1876), is undoubtedly that of a closely 
related form, Schedophilus huttoni (Waite, 
1910). The adult of S. medusophagus has 
long been known under the name ''Cen- 
trolophus britannicus Giinther, 1860a. 

"^Schedophilus maculatus Giinther, 1860. 
China Seas. D 36 (total elements). A 27 

62 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

(total elements). P 19. Gill-rakers 5 + 1 
+ 13. Vertebrae 10 + 15. Major preopercu- 
lar spines 13. This speeies is known only 
from the t\pe, a 37-mm SL specimen that is 
soft and in poor condition. 

*SchedophiIus marmomtiis Kner and 
Steindachner, 1866. "Siidsee," presumably 
near Australia. D 38 (total elements). A 
27 (total elements). Vertebrae 12+17. 
This species is usually treated as a synonym 
of S. maculatus Giinther, 1860. A probable 
synonym is HopJocoryphis pJii/.sdUanini 
Whitley, 1933, from New South Wales. 

Schcdophilii.s- huttoni (Waite, 1910) = 
Centrolopliu.s huttoni Waite. Seas of New 
Zealand, eastern Australia, and Tasmania, 
type locality Sumner, New Zealand. D 57 
(total elements). A 38 (total elements). 
Gill-rakers 5 + 12. Vertebrae 10 + 20. Lat- 
eral line scales in the 776-mm holotype near 
240. Brownish. As in S. mcdu.sophuiius, 
the number of lateral line scales probably 
increases with age. A probable synonym 
is Tuhhia tosmanica Whitley, 1943, from 
Tasmania, known only from a 10-cm speci- 
men reported as having 144 scales in the 
lateral line. 

Schedophilu.s ventmlis (Bean, 1912) = 
Eucrotus vcntmlis Bean. Bermuda. D IV- 
VII, 31-34. A III 20-23. P 22. Gill-rakers 
around 5 + 1 + 16. Vertebrae 10 + 15. Ma- 
jor preopercular spines about 9. The type 
is apparently lost. This nominal species has 
been synonymized with S. ovalis (Cuvier 
and Valenciennes, 18.33) by Fowler (1936). 

Schedo))hiJn.s iiriseoUneatus ( Norman, 
1937) = ^'PalinuriclitJuj.s griseolineatus Nor- 
man. Southern Atlantic Ocean, type local- 
ity 49'^00'S 6r58'W. D VII-VIII, 31-33. A 
III 20-21. P 19-21. Gill-rakers around 6 + 1 
+ 14. Vertebra(> 10 + 16. Lateral line 
scales about 120. Major preopercular 
spines around 14. Blue-brown, horizontally 
striped. This species can be distinguished 
at once by the increased number of caudal 
vertebrae. The large specimens which Nor- 
man ( 1937) doubtfully referred to "Palintir- 
icJithys caendeiis" belong to this species. 

Schcdopliilu.s pcmarco (Poll, 1959) = 
Palinurichthijs pemarco Poll. Gulf of 
Guinea, tropical Atlantic Ocean. D V-VII, 
23-26. A III 16-18. P 19-22. Gill-rakers 
around 5 + 1 + 16. Vertebrae 10 + 15. Lat- 
eral line scales about 95. Major preopercu- 
lar spines 15-19. Blue-brown, horizontally 
striped. The median finray counts in this 
species are lower than in any other. 

Genus CENTROLOPHUS Lacepede, 1803 
Figure 13 

Ccntrolophii.^ Lacepede, 1803:441. (Type species: 
Perca ni^ro Cinelin, 1788:132, by monotypy. 
"Rivers of Cornwall.') 

Acentrolophus Nardo, 1827:28. (Substitute name 
for CentroJophiis Lacepede, 1803, and there- 
fore taking the same type species, Perca nigra 
Gmelin, 1788:132. Centrolophiis deemed in- 
apphcable. ) 

Gijtnnucephahi.s (non Bloch, 1793:24) Cocco, 
1838:26. (Type species: Gymnocephalus 
mcs.sinen.sis Cocco, 1838:26, by monotypy. 
Messina. A synonym of Perca nigra Cmelin, 
1788:132.) (Vi^e Jordan, 1923.) 

Pompihis Lowe, 1839:81. (Type species: *Ccntro- 
lophu.'i morio Cuvier and Valenciennes, 1833: 
342, by absolute tautonymy, C. pompilus [ = 
P. pompihi.s] Cuvier and Valenciennes, 1833: 
334, considered a synonym. Madeira. A 
synonym of Perca nigra Cmelin, 1788:132. 
Preoccupied in Pompilus Schneider, 1784, 
Cephalopoda. ) 

Centrolophodes Gilchrist and von Bonde, 1923:2. 
(Type species: Centrolojyhode.s irwini Gil- 
christ and von Bonde, 1923:3, by monotypy. 
South Africa. A synonym of Perca nigra 
Gmelin, 1788:132.) 

The combination of elongate body, small 
head with prominent pores, continuous dor- 
sal fin with very weak spines graduating to 
the rays, toothless palate, very small scales, 
and 160 to 230 scales in the lateral line, 
distinguishes Centrolojdni.s from all other 
stromateoid genera. The name, a masculine 
noun, is from the Greek Kei'Tfjov, spine, 
+ Aoc/)os% crest of a helmet, probably in 
reference to the manner in which the dor- 
sal fin rises from the back. 

De.^cription. Body elongate, maximum 
depth rarely exceeding 30 per cent of the 
standard length except in very small sp(X'i- 
m(>ns; musculature firm. Peduncle broad, 

Stromateoid Fishes • Hacdrich 


y^ " 

'"•••I' ^z,-^?,,. .^«'55r^ 





Figure 13. Cenfro/ophus n/ger, drawing of a 223-mm specimen, USNM 44440, courtesy of tfie Smithsonian Institution. 

thick, long. Dorsal fin originating a little 
behind insertion of pectoral fins, contin- 
uons, about fi\'e very weak spines graduat- 
ing to the 32 to 37 rays. Anus and genital 
pore at mid-body, in a slit. Anal fin orig- 
inating a little behind middle of body, 
three weak spines precede the 20 to 23 rays. 
Pectoral fin rounded in the young, pointed 
in adult, relative length decreasing slightly 
with growth. Pelvic fins inserting under 
posterior portion of pectoral fin base, at- 
tached to the abdomen by a small mem- 
brane and folding into a shallow groove. 
Caudal fin broad, moderately forked. Very 
small cycloid scales, deciduous, covering 
fleshy bases of the median fins. Lateral 
line slightly arched anteriorly, straightening 
out about mid-body and extending onto pe- 
duncle; lateral line scales around 190. Skin 
fairly thick; extensive subdermal canal sys- 
tem communicating to the surface through 
small pores. Head usually less than 25 
per cent of the standard length, not scaled, 
pores very prominent, naked skin not pro- 
jecting backward over the nape. Eye of 
moderate size, no adipose tissue. Nostrils 
near tip of rounded snout, the anterior 
round, the posterior a slit. Angle of gape 
extending below eye. Premaxillary not pro- 
tractile. Upper jaw covered completely by 
lacrimal bone when mouth is closed. Slen- 
der supramaxillary present. Jaw teeth small, 
pointed, uniserial, spaced, increasing in 

number with growth; vomer, palatines, and 
basibranchials toothless. Opercle and pre- 
opercle thin, margins finely denticulate; 
opercle with two weak flat spines, scaled; 
angle of preopercle rounded, bulging back 
slightly; preopercle and cheek not scaled. 
Gill-rakers heavy, about half the length of 
the filaments, toothed on inner edge, 
spaced, about 13 on lower limb of first 
arch; rudimentary rakers present under 
large pseudobranch. Seven branchiostegal 
rays, five on the ceratohyal, two on the 
epihyal. Scapula prominent. Vertebrae 10 
+ 15 = 25. Caudal skeleton with six hy- 
purals and three epurals. Stomach a simple 
sac; intestine long. Pyloric caeca about 10, 

Base color brown. Range is from russet 
through chocolate to dark bluish. Median 
fins and peKics darker than the body. No 
pattern, hardK* any countershading in 
adults; young have three or four dark ver- 
tical stripes. 

Natural history. Young Centrolophus 
have been taken under jellyfish (Collett, 
1896) and swimming with Mola (Munro, 
1958). Some described as "small" were 
found in the stomachs of bottom-living hake 
trawled west of the British Isles (Blacker, 
1962 ) . Presumably these Centrolophiis had 
not been in association with pelagic medu- 
sae. While young fish are found near the 
surface, the large fish are taken at depth. 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 14. Branchial region of Cenfro/ophus n/ger, drawing o' a cleared-and-stained preparation from a 190-mm specimen. 
Elements identified in Figure 2, 

In the North Atlantic, the adults seem 
widespread, but the young have been 
found only in the eastern Atlantic and 
Mediterranean areas. A spawned-out fe- 
male, however, has been caught south of 
New England (Templeman and Haedrich, 

The young are at first vertically banded, 
but by the time they are about 100 milli- 
meters long they have become a uniform 
brown. Growth is very rapid; from De- 
cember to May, five months, a Mediterra- 
nean specimen grew from 20 to 170 milli- 
meters (Padoa, 1956). Growth is regular 
and the allometry is not marked. The num- 
ber of jaw teeth does increase, however, 
from about 17 in a 150-millimeter specimen 
to near 100 in one of 1,200 millimeters. 

Centrolophus is one of the largest stro- 
mateoids known. Specimens a meter or 

more in length have been taken in Australia 
(Mees, 1962), South Africa (Barnard, 1948), 
and the western North Atlantic. 

Autumn spawning, from October into 
winter, is indicated by the occurrence of 
eggs and very small fish at this time in the 
Mediterranean (Padoa, 1956) and by the 
capture of a large, recently spawned-out 
female in December 1963 in the western 
North Atlantic. Fraser-Brunner (1935) noted 
dimorphism in the coloring of the sexes, the 
females said to be lighter than the males. 
This difference, however, is not always ob- 

Lo Bianco (1909) observed young Centro- 
lophus feeding on medusae, but Chabanaud 
and Tregouboff (1930) found that their 
aquarium specimen preferred small fish and 
plankton. It never attempted to eat the 
medusae which were j^resent in the tank. 

Stromateoid Fishes • Hacdrich 65 

The large specimen from south of New 
England was taken on a long-line baited 
with squid. Fish and large crustacean re- 
mains occurred most often in stomachs ex- 
amined, and, on one occasion, bits of po- 
tato and an onion were found. 

As Nielsen ( 1963 ) has suggested, in re- 
porting the seining of five near Skagen, 
Centrolophus may school. Blacker (1962) 
reports several hundred\\'eight trawled off 
Ireland. Potentially a good fish with fine 
white meat, those offered experimentally 
in Milford Market found no sale (Blacker, 

Relationships. Centrolophus is one of the 
most primitive stromateoids. The small 
pharyngeal sac with few rows of large pa- 
pillae (Fig. 14), the heavy blunt-ended 
branchiostegal rays, and the large size at- 
tained, are all primitive characters. Centro- 
troIopJius shows much affinity of fonn 
towards Icichthijs, from which it differs 
mainly in having far fewer vertebrae. 

Centrolophus, Icichthijs, and Schedopliilus 
are the soft-spined centrolophids. This 
group is in general a little more primitive 
than the hard-spined centrolophids, Hijpero- 
ghjphe, Seriolella, and Psenopsis. The soft- 
spined centrolophids usually have smaller 
sacs with fewer papillae, coarser jaw teeth, 
and attain a larger size than the hard-spined 

Species. Centrolophus is known from the 
Australian region, from South Africa, and 
from the North Atlantic, where numerous 
species have been described. The counts of 
the Southern Hemisphere specimens, of 
which only a handful are known, o\erlap the 
range of those for the North Atlantic spe- 
cies. Some differences may exist in rela- 
tive proportions, but these are only at cer- 
tain stages of growth. Lacking comparative 
material, the safest course is to follow Waite 
(1910) and Mees (1962) in recognizing but 
one bipolar species: 

^Centrolophus niger (Gmelin, 1788) = 
Perca nigra Gmelin, 1788. North Atlantic, 
western Mediterranean Sea, Adriatic Sea, 
South Africa, southern Australia, and New 

Zealand, type localitv "Rivers of Cornwall." 
D 37-41 (total elements). A III 20-23. P 
19-22. Gill-rakers 5-6 + 1 + 12-15, usually 
19 total. Vertebrae 10 + 15. The name 
Centrolophus pompilus (Linnaeus, 1758) 
is often used for this fish. Linnaeus's 
Conjphaena pompilus, however, is too 
poorly characterized, and differs too much 
in certain respects to be considered the 
same species. Gmelin's (1788) Perca nigra 
is the first available name. Synonyms from 
the North Atlantic are: Centrolophus liparis 
Risso, 1826, from Nice; Acentrolophus 
maculosus Nardo, 1827, from the Adriatic 
Sea; ^Centrolophus pompilus Cuvier and 
Valenciennes, 1833, from Marseille; "^Cen- 
trolophus morio Cuvier and Valenciennes, 
1833 (ascribed to Lacepede), from the 
Mediterranean Sea; "^Schedopliilus elon- 
gatus Johnson, 1862, from Madeira; and 
"^Centrolophus calenciennesi Moreau, 1881, 
from Marseille. 

The two species described from the South- 
ern Hemisphere, here considered synonyms 
of niger, are: Centrolophus maoricus 
Ogilby, 1893, Australia and New Zealand. 
Counts made on two specimens of this 
nominal species fell at the high end of the 
range for C. niger, as do the counts for one 
small specimen reported by Regan ( 1914 ) . 

Centrolophus incini (Gilchrist and von 
Bonde, 1923) = Centrolophodes incini Gil- 
christ and von Bonde. South Africa. The 
counts reported for the holotype likewise 
fall at the high end of the range for C. 
niger. Mupus bifasciatus Smith, 1961, based 
on two small specimens, is almost certainly 
the same fish. There is little cause to doubt 
that the South African and Australian forms 
belong to the same population. 

Genus ICICHTHYS Jordan and Gilbert, 1 880 
Figure 15 

Icichthijs Jordan and Gilbert, 1880:305. (Type 
species: *Icichthys lockingtoni Jordan and 
Gilbert, 1880:305, by original designation. 
Point Reyes, California."! 

The combination of elongate soft body, 
continuous dorsal fin originating well be- 


BiiUctin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 15. Icichthys lockingtoni, drawing of a 390-mm specimen, from Parin, 1958. 

hind pectoral insertion, toothless palate, 
moderate scales covering opercles and 
cheek, 100 to 130 scales in the lateral line, 
and 50 to 60 vertebrae, distinguishes 
Icichthys from all other stromateoid genera. 
The name, a masculine noun, is from the 
Greek Hkoj, to yield, + Ixdv^i, fish, in refer- 
ence to the fish's flexible soft body. 

Description. Body elongate, maximum 
depth less than 25 per cent of the standard 
length except in small specimens; muscula- 
ture soft. Peduncle broad, compressed, of 
moderate length. Dorsal fin originating 
well behind insertion of pectoral fins, con- 
tinuous, a few very weak spines graduating 
to the rays, 39-43 elements in all. A mid- 
dorsal ridge preceding the fin. Anus at 
about mid-body. Anal fin originating slightly 
behind middle of body, three weak spines 
precede the rays, 27 to 32 elements in all. 
Median fins with compressed fleshy bases. 
Pectoral fin rounded, base fleshy. Pelvic 
fins small, inserting directly under insertion 
of pectoral fins, not attached to abdomen 
with a membrane, folding into an insignifi- 
cant groove. Caudal fin broad, slightly 
rounded or emarginate. Moderate cycloid 
scales with prominent circuli, not especially 
deciduous, covering bases of median fins. 
Lateral line slightly arched anteriorly, 
straightening out over anterior part of anal 
fin and extending onto peduncle; lateral 
line scales around 120. Skin fairly thick; 
subdermal canal system not well developed, 
pores very small. Head around 25 per cent 
of the standard length, its profile sloping 

and the pores not prominent. Top of head 
not scaled, naked skin not projecting back- 
ward over the nape. Eye of moderate size, 
no adipose tissue. Nostrils near tip of trun- 
cate snout, both round. Angle of gape ex- 
tending below eye. Premaxillary not pro- 
tractile. Only upper margin of upper jaw 
covered by lacrimal bone when mouth is 
closed. Very slender supramaxillary pres- 
ent. Jaw teeth minute, pointed, uniserial, 
close-set; vomer, palatines, and basibran- 
chials toothless. Opercle and preopercle 
thin, both well scaled, margins with very 
fine spinules; opercle with two weak flat 
spines; angle of preopercle rounded, bulg- 
ing backward. Cheek scaled. Gill-rakers 
heavy, a little shorter than the filaments, 
toothed on inner edge, spaced, about 10 on 
lower limb of first arch. Pseudobranch 
small. Seven branchiostegal rays, five on 
the ceratohyal, two on the epihyal. Scapula 
not prominent. Vertebrae 50 to 60. Caudal 
skeleton with three autogenous haemal 
spines, six hypurals, and two or three 
epurals. Stomach a simple sac; intestine 
long. Pyloric caeca about 10, digitiform, 

Color in preservative tan to dark brown, 
the median fins and pelvics darker than the 
body. No pattern, slight countershading. 

Natural Jiistory. Young Icichthys are 
commonly found swimming under or within 
medusae (Jordan, 1923a; Ilobbs, 1929; 
Fitch, 1949), and sometimes appear in fair 
number off the California coast. Large 
adults have been taken by drift-nets (Parin, 

Stromateoid Fishes • Hacdrich 


Figure 16. Branchial region of Icichthys lockingtoni, drawing of a cleared-ond-stained preparation from a 173-mm specimen. 
Elements identified in Figure 2. 

1958) and by deep trawl ( Ueno, 1954), but 
are very rare. All recorded captures are 
from deep water. Icichthys is certainly 
oceanic, and, judging from its soft tissues, 
somber color, and rare occurrence, it may 
well live as an adult in the bathypelagic 
realms. Ueno's (1954) 362-mm SL speci- 
men is the largest known. 

Icichthys is found in cool waters. The 
appearance of twelve small specimens off 
the Cape of Manazuru, Japan, in the spring 
of 1963 corresponded with an unusual in- 
flux of ca. 15°C water in this normally 
warmer area (Abe, 1963). 

Relationships. Externally, Icichthys very 
closely resembles Centwlophus, with which 
it has been synonymized by Parin ( 1958 ) . 
However, in several respects — the scalation 
on the cheeks, the caudal skeleton, and the 
greatly increased number of vertebrae — 

Icichthys differs from Ccnirolophus enough 
to warrant generic recognition. 

The structure of the pharyngeal sacs (Fig. 
16, cf. Fig. 14) and the general appearance 
of IcichtJiys suggest a very close relation- 
ship with Centrolophus. Icichthys has lost 
an epural in the caudal skeleton (Fig. 17), 
and is almost certainly the derived form. 
But Centrolophus, having lost the cheek 
scales retained in Icichthys, cannot be the 
direct ancestor. Both must have branched 
from a common stem. It is perhaps signifi- 
cant that the ranges of the two genera com- 
plement each other nicely (Fig. 52). 

Icichthys, a member of the most primi- 
tive group of stromateoids, has a very high 
number of vertebrae, an advanced condi- 
tion. The number, between 50 and 60, is 
slightly more than twice the basic perciform 
number, 25, found in other centrolophids. 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 






Figure 17. Caudal skeleton of Icichfbys lockingtoni, drawing of a cleared-and-stained preparation from a 43-mm specimen, 
SU 41028. All elements identified in Figure 1. 

The number of elements in the median fins 
is about the same as in Centrolophus, but 
there are more than twice the number of 
free intemeurals ahead of the dorsal fin. 
The evidence is at least suggestive that 
Icichthys may have arisen by polyploidy; 
chromosome counts would be most instruc- 
tive. The three autogenous haemal spines 
in the tail (Fig. 17), in contrast to the two 
of all other perciforms (Gosline, 1961a), 
are undoubtedly a by-product of the in- 
creased number of vertebrae. 

S})ccics. The genus is restricted to the 
cooler waters of the North Pacific and of 
New Zealand, from whence a new species is 
being described ( Haedrich, in press ) . Abe 
( 1963 ) reports more pyloric caeca and 
slightly fewer vertebrae for his Japanese 
specimens than are found in specimens 
from off California. Many more specimens 
will be needed to see whether these differ- 
ences are significant. From knowledge of 
Jcichthijs apparent bathypelagic habitat, it 

Stromateoid Fishes • Haedrich 


Figure 18. Senolella punctata, an elongate species, drawing of on approximately 250-mm specimen, from McCulloch, 191' 

seems best for the time being to recognize 
but one North Pacific species: 

*IcicJit]iys Jocking,toni Jordan and Gilbert, 
1880. Cahfornia to Japan, type locaHty 
Point Reyes, Cahfornia. D 39-43 ( total ele- 
ments). A 27-32 (total elements). P 18- 
21. Gill-rakers 4-6 + 1 + 11-13, usually 18 
total. Vertebrae 56-60. Synonyms, both 
based on small specimens from the coast 
of California, are *Schedo))JiiIus hcatJii Gil- 
bert, 1904, and "^Centrolophus californiciis 
Hobbs, 1929. 

Genus SERIOLELLA Guichenot, 1848 

Figures 18, 19 

Seriolella Guichenot, 1848:238. (Type species: 
Seriolella porosa Guichenot, 1848:239, by sub- 
sequent designation of Jordan, 1923:238. 
Chile. ) 

Neptomenus Giinther, 1860:389. (Type species: 
Neptomemis brama Giinther, 1860:340, by 
original designation. New Zealand. ) 

The combination of at least seven more 
dorsal than anal finrays, short stout spines 
not increasing in length to the rays in the 
dorsal fin, toothless palate, pelvic insertion 
behind the pectoral insertion, supramaxil- 
lary bone present, and lateral line following 
the dorsal profile, distinguishes Seriolella 
from all other stromateoid genera. The 
name, a feminine noun, is the diminutive 
of Seriola, a carangid genus. Ultimately 

from the Latin seria, an oblong earthen ves- 
sel, it doubtless refers to the shape of the 

Description. Body moderately deep to 
elongate, maximum depth 25 to 40 per cent 
of the standard length, compressed but 
fairly thick; musculature firm. Peduncle 
stout. Two dorsal fins, the first originating 
over or slightly behind insertion of pectoral 
fins, with seven to nine short spines. Usu- 
ally the third, fourth, and fifth spines are 
the longest, the longest spine less than half 
the length of the longest dorsal finray. 
Second dorsal with 25 to 40 finrays, the an- 
teriormost the longest. Anal and genital 
pore slightly before or behind mid-body, in 
a slit. Anal fin originating at or behind 
mid-body, three spines increase in length to 
the 18 to 25 ravs, the anteriormost finravs 
the longest. Number of dorsal finrays ex- 
ceeds number of anal finrays by more than 
seven. Pectoral fins rounded in the young, 
long and falcate in the adult. Pelvic fins 
inserting just under end of or behind pec- 
toral fin base, attached to the abdomen by 
a small membrane and folding into a shal- 
low groove. Caudal fin broad and forked. 
Large cycloid scales, very deciduous, cov- 
ering fleshy bases of the median fins. Lat- 
eral line moderately high, following dorsal 
profile and extending onto peduncle. Skin 
thin; main subdermal canal along inter- 

"() Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 19. Ser/o/e//a brama, a deep-bodied species, drawing of an approximately 250-mm specimen, from McCulloch, 191' 

muscular septum and side branches usually 
visible, pores small. Head about 30 to 35 
per cent of the standard length. Top of 
head naked, fine canal network and small 
pores usually visible, naked skin projecting 
backwards over the nape. Eye moderate 
to large. Adipose tissue around eye well 
developed and extending forward around 
the nostrils. Nostrils near tip of pointed or 
truncate snout, small, the anterior round, 
the posterior a vertical slit. Maxillary ex- 
tending below eye but angle of gape be- 
fore eye. Premaxillary not protractile. Lac- 
rimal bone partially covering upper jaw 
when mouth is closed, ventral border of 
premaxillary and end of maxillary remain- 
ing exposed. Supramaxillary present. Jaw 
teeth small to minute, pointed, uniserial, 
close-set or slightly spaced, covered later- 
ally by a membrane; vomer, palatines, and 
basibranchials toothless. Opercle and pre- 
opcrcle thin, margins entire or finely den- 
ticulate; opercle with two weak flat spines, 
scaled, the scales covered by skin; pre- 
opercle not scaled, angle rounded, bulging 
backward. Cheek scaled, the scales cov- 
ered by thick skin and not visible without 
dissection. Gill-rakers one-half to one-third 
the length of the filaments, toothed on inner 

edge, slightly spaced, 14 to 18 on lower 
limb of first arch; no rudimentary rakers 
under the small pseudobranch. Seven bran- 
chiostegal rays, five on the ceratohyal, two 
on the epihyal, the tips of the branchioste- 
gals pointed. Posterior border of scapula 
free from the body. Vertebrae 10 + 15, or 
11 + 14 = 25. In the adult, hypurals 2 -f 3 
and 4-1-5 closely conjoined or even par- 
tially fused, three epurals. Sclerotic bones 
well ossified, subocular shelf present on 
second suborbital. Stomach a simple sac; 
intestine long. Pyloric caeca numerous and 
fonning a dendritic mass. 

Color in preservative brown or bluish, 
darker above than below, the sides some- 
times with a silvery overlay. Usually a 
prominent dark blotch on the shoulder at 
the beginning of the lateral line; smaller 
spots often present on sides. Fins usually 
a little lighter than the body, but black- 
edged. Inside of mouth and gill cavity light. 

Natural history. In contrast to most other 
centrolophids, the species of ScrioIcUa are 
coastal fishes. Schools of them occur from 
150 fathoms in towards the coasts, and 
some species even enter estuaries ( Munro, 
1958). Others live in kelp beds, apparently 
not deeper than 40 fathoms (Scott, 1962). 

Strom ATEOiD Fishes • Hacdricli 71 

\ ■»^"^<..£^*- =^" - 1 err y^-  - s 

Figure 20. Branchial region of Seriolella vio/aceo, drawing of a cleared-and-stained preparation from a 188-mm specimen, 
USNM 77593. Elements identified in Figure 2. 

Nichols and Murphy ( 1922 ) report a 
young Peruvian specimen from under a 

Seriolella is the subject of a modest fish- 
ery in Chile (Mann, 1953). In Peru, nine- 
or ten-inch specimens are at times so com- 
mon that they are caught by jigging ( Nich- 
ols and Murphy, 1922). These fish are oc- 
casionally taken by fishennen in Australia 
and New Zealand, but apparently are not 
sought-after commercial species there. 

Relationships. Seriolella, with its prob- 
able off-shoot Psenopsis, represents the ad- 

vanced condition among centrolophids. 
Seriolella is derived from a Hypcroghjplie- 
like stock, with which it shares the short 
stout spines in the dorsal fin and the fluted 
first haemal spine curving backward to 
meet the first interhaemal. The slender 
pointed branchiostegal rays (Fig. 20), the 
numerous bands of small papillae in the 
pharyngeal sacs, the well ossified sclerotic 
bones, and the partial fusion of hypurals 
2 + 3 and 4 + 5 with growth are all ad- 
vanced characters, and approach the no- 
meid grade. S. violacea, from Peru, comes 


Bulletin Musctini of Comparative Zoologtj, Vol. 135, No. 2 

near to bridging the gap between Ilijpcro- 
glyphe and Seriolella. 

ScriolcIIa has given rise to Psenopsis. The 
pharyngeal sacs and caudal skeleton of both 
are very similar. Both genera have, in most 
species, well ossified sclerotic bones and a 
dark blotch on the shoulder. Seriolella, how- 
ever, is closer to Hypcroiijiiphe in the pos- 
session of a supramaxillary, which has been 
lost in Psenopsis. 

Species. Seriolella is restricted to the 
cool temperate waters of the Southern 
Hemisphere. About a dozen species have 
been described; the majority are known to 
me only from published descriptions. I 
have been able to examine only a few 
Seriolella, most of them from South Amer- 
ica. The nominal species in the genus are: 

Seriolella punctata (Bloch and Schneider, 
1801) = Scomber puncfafus Bloch and 
Schneider. Southern Australia, Tasmania, 
and New Zealand. Elongate. D VI-VII, 
34-39. A III 21-24. P 19-22. Gill-rakers 
usually 6 + 1 + 14-15. Vertebrae 10 + 15. 
Synonyms are "^'Neptomenus dohula Giin- 
ther, 1869, from Tasmania, and Neptotiienus 
hilincatus Hutton, 1872, from Wellington 
Harbor, New Zealand. 

Seriolella violacea Guichenot, 1848. Chile 
and Peru, type locality \^alparaiso. Mod- 
erately deep. D VII-VIII, 25-28. A III 
18-20. P 21-22. Gill-rakers 5-7 + 1 + 16- 
18. Vertebrae 11 + 14. This fish is the 
"cojinoba" of Chilean fisheries literature 
(Mann, 1953). Synonyms are Centrolophus 
peruanus Steindaehner, 1874, from Callao, 
Peru, and '"Neptomenus crassus Starks, 
1906, also from Callao. 

Seriolella porosa Guichenot, 1848. Chile 
and Peru, type locality Valparaiso. Elon- 
gate. D \T-VIII, 34-38. A III 22-23. P 
19-21. Gill-rakers usually 6 + 1 + 14-15. 
Vertebrae 10 + 15. This species has the 
same counts as S. punctata (Bloch and 
Schneider, 1801), and was synonymized 
with S. dohula (Giinther, 1869) [here con- 
sidered = S. punctata] by Regan (1902). 
It is unlikely that an essentially coastal fish 
such as Seriolella would regularly cross the 

broad expanse of ocean between South 
America and Australia. With closer study 
S, porosa, S. punctata, and possibly S. 
dohula will probably prove distinct. 

Seriolella hrama (Giinther, 1860) = 
Neptomenus hrama Giinther. Southern 
Australia and New Zealand, type locality 
New Zealand. Deep-bodied. D VI-VIII, 
26-33. A III 21-23. P 20-21. Gill-rakers 
7 + 1 + 16. Vertebrae 10 + 15. Neptomenus 
travale Castelnau, 1872, from New Zealand 
is a synonym. 

Seriolella velaini Sauvage, 1879. Island 
of St. Paul, Indian Ocean. Moderately deep. 
D VIII, 27. A III 20. ( From Regan, 1902. ) 

Seriolella christopherseni Sivertsen, 1945. 
Tristan da Cunha. Atlantic Ocean. D VI, 
28. A III 20. Moderately deep. (From 
Sivertsen, 1945. ) 

Seriolella noel Whitley, 1958, is based on 
one battered specimen from Sydney, Aus- 
tralia, standard length 331 mm. The counts 
given are D X + 31?; A 2-30; P 14; gill- 
rakers 8 + 16; lateral line scales 95 + 8. 
The description is inadequate to tell even 
to what genus this fish belongs, but it is 
decidedly not a Seriolella. The ten dorsal 
spines indicate it may belong in the family 

Genus PSENOPSIS Gill, 1862 

Figure 21 

Ps-cnopsis Gill, 1862:127. (Type species: Trachi- 
iwtus anoinalus Temminck and Schlegel, 1850: 
107, by monotypy. Japan. ) 

Bathyscriola Alcock, 1890:202. (Type species: 
*Bathi/seriola ajanca Alcock, 1890:202, by 
monotypy. Ganjam Coast, India. ) 

The combination of dorsal and anal fin- 
rays in almost equal numbers, spines grad- 
uating to the rays, toothless palate, pelvic 
insertion directly under the pectoral inser- 
tion, broad forward scoop in the opercle 
below the second opercular spine, and no 
supramaxillary distinguishes Psenopsis from 
all other stromateoid genera. The name, a 
feminine noun, is from the Greek i/zTJir/, 
Psenes + oxpis, appearance, drawing atten- 

Stromateoid Fishes • Hacdrich 


Figure 21. Psenops/s cyoneo, an elongate species, drawing of a 139-mm specimen, BMNH 1890. 11. 28. 9, from Alcock, 

tion to the superficial similarity between 
these two genera. 

Description. Body moderately deep to 
deep, maximum depth 30 to 45 j^er cent of 
the standard length, compressed but fairly 
thick; musculature soft. Peduncle short, 
deep, and compressed. Dorsal fin originat- 
ing over or slightly behind insertion of pec- 
toral fins, continuous, with five to seven 
short spines increasing in length to the 27 
to 32 rays. The last spine the longest, but 
less than half the length of the longest dor- 
sal finray. Anal and genital pore well 
before or at mid-body, in a slit. Anal fin 
originating well before or slightly behind 
mid-body, three spines increase in length to 
the 22 to 29 rays. Number of dorsal 
finrays never exceeds number of anal fin- 
rays by more than five. Pectoral fins 
rounded in the young, usually produced 
in the adult. Pelvic fins inserting directly 
under origin of the pectoral fin, attached 
to the abdomen by a small membrane and 
folding into a grooxe which reaches to the 
anus. Caudal fin broad, slightly forked. 
Small cycloid scales, very deciduous, cov- 
ering fleshy bases of the median fins. Lat- 
eral line moderately high, following dorsal 
profile and extending onto peduncle. Skin 
very thin; main subdennal canal along inter- 
muscular septum and side branches clearh' 
visible, canals particularly dense on back. 

pores very small. Head around 30 per cent 
of the standard length. Top of head naked, 
minute pores faintly visible, naked skin not 
projecting or projecting only slightly back- 
wards over the nape. Eye moderate to 
large. Adipose tissue around eye developed 
and extending forward around the nostrils. 
Nostrils near tip of truncate snout, mod- 
erate in size, the anterior round, the pos- 
terior a slit. Maxillary extending below eye, 
angle of gape at anterior border of eye. 
Premaxillary not protractile. Upper jaw 
covered completely by lacrimal bone when 
mouth is closed. Supramaxillary absent. 
Jaw teeth minute, pointed, uniserial, close- 
set, covered laterally by a membrane; vo- 
mer, palatines, and basibranchials toothless. 
Opercle and preopercle thin, not scaled, 
margins entire or finely denticulate; opercle 
with two weak flat spines; under the second 
spine the bone is 3-shaped, the upper in- 
dentation reaching almost to the preopercle 
and covered with uncalcified membrane; 
angle of preopercle rounded, bulging back- 
wards significantly, the margin scalloped 
in very small specimens. Gill-rakers about 
half the length of the filaments, toothed 
on inner edge, spaced, about 13 on lower 
limb of first arch; no rudimentary rakers 
under small pseudobranch. Seven bran- 
chiostegal rays, fi\e on the ceratohyal, t^^'o 
on the epihyal, the tips of the branchi- 

74 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



Figure 22. Caudal skeleton of Psenops/s anomala, drawing of a cleared-and-stained preparation from a 40-mm specimen, 
ABE 62-656. All elements identified in Figure 1. 

ostegals pointed. Scapula visible. Verte- 
brae 10 + 15 = 25. In the adult, hypurals 
2 + 3 and 4 + 5 closely conjoined, three 
epurals. Sclerotic bones well ossified, sub- 
ocular shelf present on second suborl^ital. 
Stomach a simple sac; intestine long. Py- 
loric caeca very numerous, in a mass 
resem])ling a raspberry. 

Color in preservative brown or bluish, 
deep-bodied form often with a silvery or 
whitish overlay. Deep-bodied form coun- 
tershaded, others uniform. Usually a prom- 
inent ]>lack spot on shoulder at beginning 
of lateral line. Fins a little lighter than the 
body. Opercles and peritoneum silvery or 
blackish. Inside of mouth light, gill cavity 

Natural histonj. Though fished commer- 

cially in Japan, very little is known of the 
habits of these fishes. Young Psenopsis 
have been reported in association with 
medusae (Shojima, 1961). The adults of 
P. anomala, at least, live nearer the coasts 
and in shallower water than most centro- 
lophids. Large schools are taken by near- 
shore trap nets in Japan. Adult specimens 
of P. cijanca were taken off Cananore in 
Wj. fathoms. 

Psenopsis is one of the smaller ccntro- 
lophids. Specimens of 180 mm SL are fully 
mature. Few exceed 200 mm. 

Relationships. Psenopsis, with Seriolella, 
is the most evolutionarily advanced cen- 
trolophid. The slender tapering branchio- 
stegal rays and the conjunction of hypurals 
2 + 4 and 4 + 5 ( Fig. 22 ) \\'ith growth 

Stromateoid Fishes • Hacdrich 


Figure 23. Branchial region of Pier\op%\% anomala, drawing of a cleared-and-stained preparation from a 150-mm speci- 
men. Elements identified in Figure 2. 

approach the nomeid grade. The pharyn- 
geal sacs (Fig. 23) are larger, and there 
are more numerous bands of small papillae 
than are found in Hijperoglijphe or Ccntro- 
lophits. The well ossified sclerotic bones, 
the absence of a supramaxillary bone, the 
smallish mouth, the deciduous scales, and 
the dorsal fin with only slightly more fin- 
rays than the anal suggest that Fscnopsis 
may be near the base of the line leading to 
the Stromateidae. ScrioJella, which retains 
the supramaxillary lost in Psenopsis, is its 
closest relative within the centrolophids. 

Species. Psenopsis is an Indo-Pacific 
genus, found in India, Japan, northwest 
Australia, and the East Indies. There are 
four allopatric species, one of them un- 

described. Little confusion has arisen re- 
garding the identification of these fishes, 
and there are no problems of synonymy. 
The species are: 

Psenopsis anomala (Temminck and Schle- 
gel, 1850) = TracJunotus anomahis Tem- 
minck and Schlegel. China and southern 
Japan, type locality Tokyo. Deep-bodied. 
D V-\TI, 27-32. A III 25-29. P 20-23. 
Gill-rakers usually 6 + 1 + 13, 12-15 on 
lower limb of first arch, 18-21 total. Verte- 
brae 10 + 15 ( skel. ) . This species is the 
"ibodai" of Japanese fisheries literature, 
and is common from Hong Kong to Tokyo 
and into the Sea of Japan as far north as 
Hokkaido. It fomis the basis of an impor- 
tant fishery. P. sliojimai Ochiai and Mori, 

76 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



HYPURAL 4+5 — 

HYPURAL 2 + 3 — 

Figure 24. Caudal skeleton of Nomeus gronovii, drawing of a cieored-and-stained preparation from an 87-mm specimen. 
All elements identified in Figure 1. 

1965, from the Sea of Japan is a probable 

Psenopsis Munro, 1958. Dam- 
pier Archipelago, N. W. Australia. Deep- 
bodied. D VII, 28. A III 25. P 22. Gill- 
rakers 12 on lower limb of first arch ( from 
Munro, 1958). Probably a good species, 
little differentiated from P. anumala. 

Fsenopslsciianea (Alcock, 1890) = *B(ithy- 
seriola cijanea Alcock, type locality, Ganjam 
Goast, India. Elongate.' D VI, 25-26. A III 
22-23. P 20. Gill-rakers 5+1 + 14. Verte- 
brae 10 + 15. 


Type genus: Nomeus Cuvier, 1817 

Fasteur.s. Cuvier and \'aleiicieiines, 1833:242 
( descT. ) . 

Nomeina. Giinther, 1860:387 (in part, def.). 

Nonieidae. Giinther, 1880:455 (in part, def.). 
Jordan and Gilbert, 1882:448 (descr.). Jor- 
dan and Evermann, 1896:948 (descr.. North 
America). Jordan, 1923:183 (in part, hst). 
BerK, 1940:323 (in part, dist.); 1955:249 (in 
part, dist.). Norman, 1957:503 (in part, def., 
genera hsted ) . 

Psenidae. Auctorinn. 

Diagnosis. Stromateoid fishes with pelvic 
fins present in adults, two dorsal fins, teeth 
on vomer and palatines, six branchiostegal 
rays, and four hypural and three epural 
bones in the tail. The papillae in the pharyn- 
geal sacs with stellate bases, arranged in 
about five broad longitudinal bands. 

Description. Body slender to deep, com- 
pressed. Two dorsal fins, the first with 
about ten slender spines folding into a 

Stromateoid Fishes • HaedricJi 

I I 

Figure 25. Branchial region of Nomeus gronovii, drawing of a cleared-and-stained preparation from a 187-mm specimen, 
MCZ 35327. Elements identified in Figure 2. 

groove, the longest spine at least as long 
as the longest ray of the second dorsal fin. 
One to three anal spines, not separated 
from the rays. Soft dorsal and anal fins 
approximately the same length. Bases of 
median fins sheathed by scales. Pelvic fins 
attached to the abdomen by a thin mem- 
brane, folding into a narrow groove, the 
fins greatly produced and expanded in 
young Nomeus and some Psenes. Scales 
small to very large, cycloid or with ver\' 
small weak cteni, thin, extremely deciduous. 
Lateral line high, following dorsal profile 
and often not extending onto peduncle. 
Skin thin; subdermal mucous canal system 
well developed and visible in most; the 
main canal down the side of the body may 
be mistaken for a lateral line. Opercular 
and preopercular margins entire or finely 
denticulate. Opercle very thin, with two 
flat, weak spines. Six branchiostegal rays. 

Mouth small, maxillary rarely extending to 
below eye. Teeth small, conical, or cusped 
in some Psenes, approximately uniserial in 
the jaws, present on vomer, palatines, and 
basibranchials. Supramaxillary absent. Adi- 
pose tissue around eye only moderately 
developed in most. Vertebrae 30 to 38, 41, or 
42. Caudal skeleton with four hypurals and 
three epurals. Pharyngeal sacs with papil- 
lae in upper and lower sections, papillae in 
five to seven broad longitudinal bands. 
Bases of the papillae stellate, teeth seated 
on top of a central stalk. Adults usually 
about a foot long, although a giant 
Cubiceps may exceed three feet. Silver\' 
to bluish-brown, some with conspicuous 
striped or blotched pattern. 

Distribution. Nomeids are oceanic fishes 
of tropical and subtropical waters. They 
occur in the Gulf of Mexico, the Caribbean 
Sea, the Atlantic Ocean, the western Medi- 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

terranean Sea, the Indian Ocean, and across 
the Pacific. Numerous in the waters of the 
Philippines and soutliern Japan, they do not 
seem to enter the shallow South China Sea 
(Fig. 54). 

Relationships. From an evolutionary 
standpoint, the nomeids are a grade above 
the centrolophids. There are more verte- 
brae, fusions have occurred in the hypural 
fan ( Fig. 24 ) , a branchiostegal ray has been 
lost, and the papillae in the pharyngeal sacs 
have stellate bases (Fig. 25). Nomeids 
have teeth on the palate and basibran- 
chials, however, which precludes their deri- 
vation from a centrolophid. Probably both 
families have a common ancestor, and de- 
velopment has been somewhat parallel. 
The palatal dentition, lost in the Centro- 
lophidae, remains in nomeids. The Nome- 
idae have passed through the centrolophid 
stage without leaving living representatives 
at that level. 

The Nomeidae have given rise to two 
other families, each with a single genus. 
The tetragonurids, a very highly specialized 
group, arose early, perhaps from the same 
line which produced Psencs. The similarity 
between the teeth of Tetragonurus and 
Psenes pcUucidus is striking, but need not 
imply too close a relationship. The re- 
appearance of characters in divergent lines 
of common ancestry is not an unusual phe- 
nomenon (Simpson, 1953), and seems wide- 
spread in stromateoids. 

The ariommids may have been derived 
more recently. Superficially, they resemble 
nomeids very much, but the teeth on the 
palate have been lost, further fusions have 
taken place in the caudal skeleton, and the 
pharyngeal sacs are strikingly divergent. 
The species of the Cuhiceps pauciradiatus 
group may share a common ancestor with 
the Ariommidae. These show a tendency 
toward the ariommid condition in the re- 
duced palatal dentition, and share with 
them the very large, deciduous scales and 
the extremely slender (sometimes even ab- 
sent ) bridge over the anterior vertical canal 
in the ear. 

Key to Nonwid Genera 

1 (4). Body elongate, maximum depth us- 

ually less than .35 per cent of the stan- 
dard length, greatest in small speci- 
mens. Origin of dorsal fin behind, or 
directly over in very small specimens, 
insertion of pectoral fins. 2 

2 (3). Anal count I-III 14-25. Insertion of 

pelvic fins under end or behind base 
of pectoral fin. An oval patch of 
knoblike teeth on the tongue. Ver- 
tebrae 30 to 33. Cuhiceps, p. 78. Fig. 26 

3 (2). Anal coimt I-II 24-29. Insertion of 

pelvic fins before or under insertion 
of pectoral fin, possibly behind in very 
large specimens. No patch of teeth 

on the tongue. Vertebrae 41. 

Nomeits, p. 81. Fig. 27 

4 ( 1 ) . Body deep, maximum depth usually 

greater than 40 per cent of the stan- 
dard length, but possibly less in very 
large specimens. Origin of dorsal fin 
before, or directly over in large speci- 
mens, insertion of pectoral fins. 

Psenes, p. 84. Fig. 28 

Genus CUBICEPS Lowe, 1843 

Figure 26 

Cuhieeps Lowe, 1843:82. (Type species: Seriola^ 
gracilis Lowe, 1843:82, by subsequent desig- 
nation of Jordan and Evermann, 1896:950. 
Madeira. ) 

Atiniostoma A. Smith, 1849, plate XXIV. (Type 
species: AtiniDstoina capensis Smith, 1849, 
plate XXIV, by monotypy. South Africa. ) 

Navarchiis Filippi and Verany, 1859:187. (Type 
species: Navarchtis suJcatus Filippi and Ver- 
any, 1859:187, by monotypy. Mediterranean. 
A synonym of Cuhiceps gracilis Lowe, 1843: 
82. ) 

TraclielocirrJnis Doumet, 1863:220. (Type species: 
Trachelocirrhus mediterraneus Doumet, 1863: 
222, by monotypy. Sete, France. A synonym 
of Cuhiceps gracilis Lowe, 1843:82.) 

Miilichthys Lloyd, 1909:1.56. (Type species: Muli- 
chtJiys s<piatuiceps Lloyd, 1909:158, by 
monotypy. Arabian Sea.) 

MamlclichtJujs Nichols and Murphy, 1944:247. 
(Subgenus. Type species: Cuhiceps cari- 
natus Nichols and Murphy, 1944:245, by 
monotypy. 180 miles SW of Cape Mala, 
Panama. ) 

^ I^owe described his fish as a species in the 
genus Seriola, but noted (p. 82), "Still it is not 
unlikely that a comparison of the two fishes 
[gracilis and S. hijiinnulata ( Quoy and Gaimard)] 
may warrant . . . their separation from Seriola 
into a genus, winch may be called Cuhiceps." 

Stromateoid Fishes • HaedricJi 


The combination of elongate body, long 
winglike pectoral fins, insertion of pelvics 
behind pectoral fin base, scales on top of 
head, cheeks, and opercles, and a patch of 
teeth on the tongue distinguishes Cubiceps 
from all other stromateoid genera. The 
name, a masculine noun, is from the Greek 
Kvf^o'i, cube, + K£<^oA?/, head, in reference to 
the square profile of the fish's head. 

Description. Body elongate, maximum 
depth 25 to 30 per cent of the standard 
length; musculature firm. Peduncle short, 
deep, and compressed. Two dorsal fins, 
scarcely divided. First dorsal originating 
behind insertion of pectoral fins, with about 
ten stiff spines folding into a groove, the 
longest spine longer than the longest ray of 
the second dorsal. Anterior rays of the 
second dorsal the longest, those that follow 
decreasing in length, 14 to 23 finrays in 
all. Anal and genital papillae behind mid- 
body, in a slit. Anal fin originating behind 
origin of second dorsal fin, one to three 
short spines preceding the rays. Anterior 
rays the longest, those that follow decreas- 
ing in length, 14 to 21 finrays in all. Pec- 
toral fin pointed, becoming very long and 
winglike, the relative length increasing 
markedly with growth; base of the fin 
inclined at an angle of 45°. Pelvic fins 
inserting just under end of or behind pec- 
toral fin base, attached to the abdomen by 
a small membrane and folding into a deep 
groove. Expanded coracoid often forming 
a conspicuous keel along mid-ventral line 
ahead of pelvics. Caudal fin forked, the 
lobes often folding over one another. Scales 
large, cycloid, very deciduous, covering 
bases of the median fins. Simple tubed 
scales of lateral line high, following dorsal 
profile and ending under last dorsal finray 
or extending onto peduncle. Skin thin; sub- 
dermal canals on flanks easily traced. Main 
canal may be confused with lateral line. 
Pores to surface small. Head around 32 
per cent of the standard length. Top of 
snout naked, minute pores in naked skin. 
Scales extending forward on top of head 

almost to level of the nostrils. Eye large, 
bony supraorbital ridge pronounced. Adi- 
pose tissue around eye well developed, 
extending forward around the nostrils. Nos- 
trils near tip of blunt snout, small, both 
round. Maxillary ending under anterior 
border of eye, angle of gape well before 
eye. Premaxillary not protractile. Lacrimal 
bone completely covering upper jaw when 
mouth is closed, ventral border of maxillary 
sometimes remaining exposed. Supramaxil- 
lary absent. Jaw teeth small, pointed, 
slightly recurved, usually spaced. Very 
small teeth usually present on vomer and 
in a single series on the palatines and basi- 
branchials. An oval patch of low knoblike 
teeth on the glossohyal. Opercle and pre- 
opercle thin, scaled, margins entire or finely 
denticulate; opercle with two weak flat 
spines; angle of preopercle slightly rounded, 
not bulging backward. Cheeks scaled. Cill- 
rakers slender, almost the length of the 
filaments, toothed on inner edge, fairly 
close set, 14 to 18 on lower limb of first 
arch; no rudimentary rakers under the large 
pseudobranch. Scapula prominent. Verte- 
brae 13 + 17 to 15 + 18 = 30 to 33. Scle- 
rotic bones usually well ossified in adults. 
Stomach a simple sac; intestine of moderate 
length. Pyloric caeca very numerous, in a 
large dendritic mass. 

Color in preservative either brownish, 
darker on the back than on the sides, or 
bluish above and silvery on the sides. First 
dorsal blackish, other fins usually the same 
color as the body, caudal dusky. Iris grey- 
ish, often with golden semicircles at anterior 
and posterior borders. Inside of mouth, gill 
cavity, and peritoneum dark. 

NofumI history. All the nomeids are 
oceanic. Most species are very rarely seen, 
and little is known of their habits. 

Small Cubiceps gracilis are very numer- 
ous near the Azores, where they are taken 
in surface nets and from under medusae. 
By the time these fishes reach about 200 
mm SL, they are mature. With the attain- 
ment of maturity growth does not stop, but 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



Figure 26. Cubiceps gracilis, drawing of a 164-mm specimen, from Gunther, 1889. 

continues significantly. Specimens near 800 
mm SL have been reported from the Medi- 
terranean (Ariola, 1912). With growth, the 
relative length of the pectoral fin increases 

In the Philippines, Cubiceps is caught by 
using night-lights and lift-nets ( Herre and 
Herald, 1950). Large specimens are taken 
occasionally by the near-shore winter long- 
line fishery for swordfish off southern 

Relationships. Cubiceps occupies the 
central position in nomeid evolution. The 
two other nomeid genera, Nomeus and 
Psenes, are certainly derived from Cubi- 
ceps. In both derived genera the patch of 
teeth on the tongue has been lost, and 
there has been a tendency towards an 
increase in the number of vertebrae and 
finrays. The papillae in the pharyngeal 
sacs of Cubiceps are very similar to those 
of Nomeus (Fig. 25), but those of Psenes 
are in general smaller (Figs. 29, 30). The 
caudal skeletons of all three genera are 
almost identical (Fig. 24). 

The Ariommidae have probably evolved 
from a Cubiceps, although the systematic 
position of the family is far from clear. 
The ariommids do share certain characters, 
however, with the fi.shes of the Cubiceps 
pauciradiatus group, as discussed on pages 
77 and 90. 

Cubiceps and the tetragonurids both have 
patches of teeth on the tongue. The jaw 
teeth, the pharyngeal sacs, and the scales, 

however, are very different in these two 
groups. It seems likely that the Tetrago- 
nuridae branched off very early from the 
nomeid stem, but their ancestor may have 
been a fish similar in many respects to the 
present-day Cubiceps. 

Species. Cubiceps is found in the tem- 
perate and tropical waters of the Atlantic, 
Pacific, and Indian oceans. It also occurs in 
the western Mediterranean Sea, and the 
Gulf of Mexico and Caribbean Sea. 

Much confusion surrounds the species of 
Cubiceps. The counts of all described are 
fairly close. Lacking sufficient comparative 
material, it is difficult to evaluate the small 
differences which do occur, for example in 
vertebral number. There seems to be a 
great differential in the size attained by the 
adult. The species of the Cubiceps paucira- 
diatus group may never exceed 200 mm SL 
(Haedrich, 1965), whereas fishes allied to 
Cubiceps gracilis are reported (Smith, 1849; 
Ariola, 1912; Abe, 1955a) to approach a 
meter in length. 

The nominal species in the genus Cubi- 
ceps are: 

Cidjiceps gracilis (Lowe, 1843) = Seriola 
gracilis Lowe. Atlantic Ocean and western 
Mediterranean, type locality Madeira, per- 
haps a world-wide species. D IX-XI, I-II 
20-22. A I I-II I 20-23. P 20-24. Gill-rakers 
8-9 + 1 + 14-17. Vertebrae 15 + 18. Syn- 
onyms are: Navarchus sulcatus Filippi and 
Verany, 1859, from the Mediterranean; 
Trachelocirrhus mediterraneus Doumet, 

Stromateoid Fishes • Haedrich 


1863, from Sete; Ciihiceps lowei Osorio, 
1909, from the Cape \'erde Islands; and 
Aphareus ohtusirostrls Borodin, 1930, from 
the Azores. 

Cuhiceps capensis (A. Smith, 1849) = 
*Atimostoma capensis Smith. Type locahtv' 
South Africa. D IX-X, I-III 24-26. A I'l 
22-23. P 16-18. Gill-rakers 8-9 + 1 + 16- 
17. Vertebrae 14 + 17. The stuffed type is 
about 900 mm SL. The margin of the oper- 
cle between the two flat opercular spines 
may be strongly serrate in this species. A 
probable synonym is Cubiceps niger Franca, 
1957, from Angola. 

*Cubiceps pauciradiatiis Giinther, 1872. 
Central and western Pacific Ocean, type 
locality Misol Island. D X-XII, I 16-18. A 
I-II 14-17. P 18-19. Gill-rakers 8-9 + 1 + 
16. This is a diminutive species, rarely 
exceeding 160 mm SL. Closely related 
forms are: *C. longimanus Fowler, 1934a, 
C. carinatus Nichols and Murj^jhy, 1944, and 
*C. athenae Haedrich, 1965. ' "^Cubiceps 
nesiotes Fowler, 1938, from Christmas Is- 
land, Central Pacific, is a probable synonym. 

Cubiceps sqiiomiceps (Lloyd, 1909) = 
Midichthys squamiceps Lloyd. South Africa 
to Japan, type locality Arabian Sea. D IX- 
XI, I-II 19-21. A II-III 18-21. P 18-20. 
Gill-rakers 8-9 + 1 + 16-17. This is a 
chunky-looking fish, with a very short 
peduncle and broad, winglike pectoral. 
*Cubiceps nataJensis Gilchrist and von 
Bonde, 1923, is probably a synonym. 

^Cubiceps caendeus Regan, 1914a. Tas- 
man Sea, tvpe locality Three Kings Island. 
D X-XI, I-II 21-24. A II-III 21-24. P 19- 
21. Gill-rakers 7 + 1 + 16-17. Vertebrae 13 
+ 18. A few of the jaw teeth on the types 
are long and project like fangs. Cubiceps 
baxteri McCulloch, 1923, based on a dam- 
aged specimen 371 mm long, may be the 

"^Cubiceps longimanus Fowler, 1934. 
V^estern Indian Ocean, type locality Dur- 
ban. D X-XI, I 15-16. a'I-I1 15. p' 18-20. 
Gill-rakers 9+1+14. All specimens known 
are less than 50 mm SL long. 

Cubiceps carinatus Nichols and Murphy, 

1944. Pacific Coast of Central America, 
type locality Gulf of Panama. D IX-X, I 
14-16. A II 14-15. P 17-19. Gill-rakers 
7-8 + 1 + 14-16. Vertebrae 13 + 17. This 
species has fewer median finrays than the 
closely related C. pauciradiatus Giinther, 
1872, from the Central and Western Pacific. 
*Cubiceps athenae Haedrich, 1965. East 
coast of North America. D X-XI, I 15-16. 
A II 14-15. P 18-19. Gill-rakers 8 + 1 + 
16-17. Vertebrae 13 + 18. 

Genus NOMEUS Cuvier, 1817 

Figure 27 

Nomeus Cuvier, 1817:315. (Type species: Gobius 
gronovii Gmelin, 1788:1205, by subsequent 
designation of Jordan and Gilbert, 1882: 449. ^ 
Atlantic Ocean. ) 

The combination of elongate body, black 
fanlike pelvic fins with the full length of 
the trailing edge attached to the abdomen, 
insertion of the pelvics (usually) ahead of 
the pectorals, blotched and spotted pattern, 
and 41 vertebrae distinguishes Nomeus 
from all other stromateoid genera. The 
name, a masculine noun, is from the Greek 
vofxeis, herdsman, a translation of the Dutch 
vernacular "Harder" (Marcgrave, 1648), 
probably in reference to the fishes' habit 
of following Fhijsalia. 

Description. Body elongate, maximum 
depth around 30 per cent of the standard 
length; musculature firm. Peduncle slightly 
tapered, compressed. Two dorsal fins, 
scarcely divided. First dorsal fin originat- 
ing over or a little behind insertion of the 
pectoral fin, with about ten soft spines 
folding into a deep groove, the longest 

1 Cuvier and Valenciennes (1833:242) desig- 
nated Nomeus nuniritii Cuvier ( 1817:.315) type 
for the genus. Cu\ier's species, howe\er, based 
on the "Harder" of Marcgrave (1648:153), ap- 
peared in name only, the description being later 
supplied by Cuvier and Valenciennes (1833:243). 
Under the International Code, a uotncn uiiduru 
is unavailable as a type, and Nomeus mauritii 
Cuvier, 1817, is thus rejected. Nomeus mauritii 
( non Cuvier, 1817 ) Cuvier and Valenciennes, 
1833, is a synonvm of Nomeus gronovii ( Gmelin, 

82 Bulletin Museum of Comparative Zoolo<:,ij, Vol. 135, No. 2 

Figure 27. Nomeus gronov/i, drawing of an approximately 40-mm specimen, courtesy of tfie Smitfisonian Institution. 

spine .slightly longer than the longest ray 
of the second dorsal. Anterior rays of the 
second dorsal fin the longest, those that 
follow shorter, subequal, 24 to 28 finrays 
in all. Anal and genital papilla at mid-body, 
in a deep slit. Anal fin originating under 
or slightly behind origin of second dorsal 
fin, one or two weak spines preceding the 
rays. Anterionnost rays the longest, those 
that follow shorter, subequal. Pectoral fin 
rounded in the young, pointed, winglike, in 
the adult; the relative length increases 
markedly with growth. Pelvic fins inserting 
before, or in large specimens under, pec- 
toral fin base, fan-shaped, innermost ray 
the longest, attached to the abdomen for its 
entire length by a strong membrane and 
folding into a deep groove which reaches 
to the anus. Relative length of the fin 
decreasing markedly with growth. Caudal 
fin deeply forked, lobes very long. Scales 
small, cycloid, very thin, deciduous, covering 
bases of median fins. Simple tubed scales 
of lateral line high, following dorsal profile 
and not appearing to extend onto peduncle. 
End of lateral line under last dorsal finray 
in most specimens (in the single large 
specimem known, the lateral line reaches 

the caudal base). Skin thin; subdermal 
canals on flanks easily traced. Main canal 
may be confused with lateral line. Pores to 
surface minute. Head around 30 per cent 
of the standard length. Top of snout naked, 
minute pores visible in naked skin. Scales 
extending forward over nape to level of an- 
terior border of the eye. Interorbital space 
and top of snout covered with adipose 
tissue. Eye of moderate size, bony supra- 
orbital ridge pronounced. Adipose tissue 
around eye very well developed, extending 
forward to cover the lacrimal and surround- 
ing the nostrils. Nostrils on tip of truncate 
snout, small, the anterior round, the pos- 
terior a slit. Maxillary ends under, or before 
in large specimens, anterior border of the 
eye, angle of gape well before eye. Pre- 
maxillary not protractile. Lacrimal bone 
almost completely covering upper jaw when 
mouth is closed, ventral border of maxillary 
remaining exposed. Supramaxillary absent. 
Jaw teeth small, pointed, slightly recurved, 
spaced. Small recurved teeth present on 
vomer and in a single series on the pala- 
tines and basibranchials. No teeth on the 
glossohyal. Opercle and preopercle thin, 
scaled, margins very finely denticulate or 

Stromateoid Fishes • Haedrich 


entire; opercle with two weak hardly de- 
fined flat spines; angle of preopercle 
rounded, bulging backward in large speci- 
mens. Cheeks scaled. Gill-rakers slender, 
half the length of the filaments, toothed on 
inner edge, fairly close-set, about 16 on 
lower limb of first arch; sometimes a few 
rudimentary rakers under the large pseudo- 
branch. Six branchiostegal rays, four on 
the ceratohyal, two on the epihyal. Scapula 
prominent. Vertebrae 15 + 26 = 41. In the 
single large specimen known, the sclerotic 
bones are well ossified. Stomach a simple 
sac; intestine of moderate length. Pyloric 
caeca numerous, in a small dendritic mass. 

Color in life bright blue above, blotched 
and spotted with blue on the brilliant 
silvery sides. In preservative, the base 
color is tan, the blotches and spots appear- 
ing dark brown. Median fins \\'ith about 
three dark stripes. First dorsal and pel vies 
black, pectorals light. The single large 
specimen known is uniform dark brown. 
Eye usually greyish; in the large specimen 
the iris is golden, divided by a dark ver- 
tical bar. Inside of mouth, gill cavity, and 
peritoneum light. 

Natural history. The association between 
Nomeus and PhysaJia is commonly cited 
as an example of commensalism, but ac- 
tually very little is known about the true 
nature of the association or about the life 
histories of the animals involved. No77ieus 
has been observed eating Physalia (Kato, 
1933) and vice verso (Carman, 1896). In 
comparison to other fishes, however, No- 
meus is relatively immune to the toxin of 
the siphonophore (Lane, 1960). 

The eggs and larvae of Nomeus are not 
known. Fishes of 10-mm SL have been 
taken from under Physalia, however, indi- 
cating that the association must fonn early 
in the fish's life. I have seen 150-mm SL 
specimens also taken with Physalia. These 
fishes appeared to be adults, but none were 
ripe. The largest known specimen of No- 
meus was taken with a bottom trawl in the 
Caribbean Sea. This 225-mm specimen was 
apparently not in association with Physalia, 

was living fairly deep in the water, and was 
a uniform dark brown instead of blotched 
like smaller specimens. This fish appeared 
to be a mature male. Many young stromat- 
eoids live with jellyfishes, have a blotched 
or mottled color pattern, and both desert 
their coelenterate host and become uni- 
formly colored with growth. The discovery 
of this large, dark Nomeus suggests that the 
familiar small, blotched Nomeus found un- 
der Physalia may only be the young form 
of a bigger fish which lives in the depths. 

Relationships. Nomeus is derived from 
the central Cubiceps stock. It is very simi- 
lar in appearance to Cubiceps. The differ- 
ences between the two are slight, but 
sufficient to consider Nomeus a genus in 
its own right. In Nomeus the pelvic bones 
have become much shortened, the patch of 
teeth on the glossohyal has disappeared, 
the number of vertebrae and finrays has 
increased, and there are only two spines 
preceding the anal finrays. 

Nomeus lives in a very specialized en- 
vironment, under Physalia. The features 
which distinguish it from Cubiceps doubt- 
less reflect the demands of this unusual 
habitat. Adapted to a particular way of 
life, Nomeus has given rise to no other 

Species. Nomeus is a wide-ranging genus, 
found in the temperate and tropical waters 
of all the major oceans. It does not occur, 
however, in the eastern Atlantic or the 
Mediterranean. I have examined specimens 
from the Atlantic Ocean, the Culf of Mexico 
and Caribbean Sea, the Indian Ocean, and 
the central and western Pacific Ocean. The 
counts of all these are essentially the same. 
I cannot but conclude that in the genus 
Nomeus there is but one species: 

Nomeus g,ronovii (Gmelin, 1788) = Gobius 
gronovii Gmelin. Temperate and tropical 
regions of the Atlantic, Pacific, and Indian 
Oceans, Gulf of Mexico and Caribbean Sea, 
type locality "American Ocean in the Tor- 
rid Zone." D IX-XII, 24-28. A I-II 24-29. 
P 21-23. Gill-rakers 8-9 + 1 + 15-18. Ver- 
tebrae 41. Synonyms are: Eleotris mauritii 


Bulletin Museum of Comparative Zoology , Vol. 135, No. 2 

Bloch and Schneider, 1801, from Mauritius; 
Nomeu.s maculosus Bennett, 1831, from the 
Atlantic coast of North Africa; Nomeus 
maculattis Valenciennes, 1836; Nomeu.s 
oxytini.s Poey, 1861, from Cuba; and No- 
meu.s dijscritus Whitley, 1931, from New 
South Wales. *Nomeus peronii Cuvier and 
Valenciennes, 1833, from the seas of Java 
is known only from three specimens in very 
poor condition. It is hard to be certain that 
these fish belong to Nomeus, though they 
are almost surely nomeids. I was able to 
make counts on only one. The counts were 
D IX, 30; A 31 (total elements); P 21; gill- 
rakers 8 + 1 + 16. No subsequent material 
has appeared, and I consider *iV. peronii a 
synonym of N. gronovii. 

Genus PSENES Cuvier and Valenciennes, 

Figure 28 

Psenes Cuvier and Valenciennes, 1833:259. (Type 
species: *P.senes cyanophrys Cuvier and Val- 
enciennes, 1833:260, by original designation. 
New Ireland, western Pacific Ocean. ) 

Icticus Jordan and Thompson, 1914:242. (Type 
species: * Icticus i.schaini.s- Jordan and Thomp- 
son, 1914:242, by original designation. Oki- 
nawa, western Pacific Ocean. A synonym of 
*Pscucs pcUiicidus Liitken, 1880:516.) 

Papyrichthys J. L. B. Smith, 1934:90. (Type .spe- 
cies: *Psenes pcUucidus Liitken, 1880:516, by 
original designation. Straits of Surabaja, 
Java. ) 

Thecopsenes Fowler, 1944a: 63. (Type species: 
*P.'ienes chapmani Fowler, 1906:119, by orig- 
inal designation. Cape Verde Islands, Atlantic 
Ocean. A synonym of *Psene.s cyanophrys 
Cuvier and Valenciennes, 1833:260.) 

Carlstioides Whitley, 1948:87. (Type species: 
Cari^ioides amplipinni.s- Whitley, 1948:88, by 
monotypy. Lord Howe Island, Tasman Sea. 
A svnonym of '*P.sencs pclhicidus Liitken, 

Parapsenes J. L. B. Smith, 1949a :847. (Type 
species: Psenes rotundus Smith, 1949:307, by 
original designation. Dassen Island, South 
Africa. ) 

The combination of two dorsal fins, the 
first dorsal fin originating before or over 
the pectoral insertion, pelvic fins present, 
deep body, teeth on the palatines and basi- 
branchials, and no teeth on the glossohyal 

distinguishes Psenes from all other stromat- 
eoid genera. The name, a feminine noun, 
is from the Greek i//?;!?/, the osprey Pandion, 
the allusion not evident. The authors of the 
name, Cuvier and Valenciennes (1833), 
may have been impressed by the resem- 
blance of the "sourcil bleu" on their little 
fish to the similar brow of the fish hawk. 

Description. Body deep, maximum depth 
usually greater than 40 per cent of the stan- 
dard length, but sometimes less in large 
specimens; musculature firm to soft and 
flabby. Regions at bases of median fins may 
be very compressed and translucent. Pe- 
duncle short, compressed, may be fairly 
slender. Two dorsal fins, scarcely divided. 
First dorsal fin originating before insertion 
of pectoral fins, with about ten soft spines 
folding into a deeper groove. Rays of the 
second dorsal fin nearly as long as the long- 
est Di spine, all approximately the same 
length or decreasing in length posteriorly, 
18 to 30 finrays in all. Anal papilla a little 
before mid-body, in a depression. Anal fin 
originating at mid-body, slightly behind 
origin of second dorsal fin, two or three 
weak spines preceding the 17 to 30 rays. 
Pectoral fin rounded or winglike; relative 
length of fin decreasing slightly or increas- 
ing markedly with growth. Pelvic fins in- 
serting under posterior portion of pectoral 
fin base, attached to the abdomen by a 
small membrane and folding into a groove. 
Pelvics very long in the young of some, the 
relative length decreasing markedly with 
growth. Caudal fin deeply forked. Scales 
small to minute, with a few weak cteni, 
very thin, deciduous, covering bases of me- 
dian fins. Simple tubed scales of lateral 
line high, following dorsal profile and end- 
ing under last dorsal finray or extending 
onto peduncle. Skin thin; main subdermal 
canals along intermuscular septum appar- 
ent, may be confused with lateral line, side 
branches not visible. Pores to surface mi- 
nute or absent. Head around 30 per cent 
of the standard length. Top of snout naked, 
minute pores in naked skin. Scales extend- 
ing forward on top of head almost to level 

Stromateoid Fishes • Haedrich 85 

20 mm 

Figure 28. Psenes pellucidus, drawing of a 1 30-mm specimen by Margaret Bradbury. 

of anterior border of the eye. Eye moderate 
to large. Adipose tissue around eye mod- 
erately developed. Nostrils near tip of trun- 
cate snout, small, the anterior one round, 
the posterior a slit. Maxillary ending under 
anterior portion of the eye. Premaxillary 
not protractile. Lacrimal bone almost com- 
pletely covering upper jaw when mouth is 
closed, ventral border of maxillary remain- 
ing exposed. Supramaxillary absent. Teeth 
in both jaws uniserial, pointed; teeth in 
upper jaw small, slightly recurved, spaced; 
teeth in lower jaw larger, may be long and 
bladelike with small cusps, close-set. A 
few small recurved teeth in a patch on the 
head of the vomer and in a single series on 
each palatine and on the basibranchials. 
Opercle and preopercle thin, scaled, mar- 
gins very finely denticulate or entire; oper- 
cle with two weak, hardh" defined, flat 
spines; angle of preopercle rounded, pro- 
jecting backward very slightly. Gill-rakers 
slender, a little shorter than the filaments, 
toothed on inner edge, moderately spaced, 
14 to 19 on lower limb of first arch; some- 
times a iew rudimentary rakers under the 
long pseudobranch. Six branchiostegal rays, 

four on the ceratohyal, two on the epihyal. 
Scapula not prominent. Vertebrae 13 to 15 
+ 18 to 23 = 31 to 38, or 15 + 26 or 27 = 41 
or 42. Stomach a simple sac; intestine very 
long. Pyloric caeca numerous, in a dendritic 

Color in preservative bro\\'n to yellowish, 
some species with a conspicuous, dark, 
blotched or longitudinally striped pattern. 
Median fins and pelvics often darker than 
the body. Region at bases of median fins 
translucent in P. pcUucidus. Inside of mouth 
light brown, gill cavity usually darker. 
Peritoneum dark or light. 

Natural history. The young of Psenes are 
fairly common in the surface layers on the 
high seas. They do not seem to associate 
with jellyfishes to any extent, but are \ery 
often dipnetted from under floating Sargas- 
sum. The larger adults, as with most other 
stromateoids, probably live deeper in the 
water. Most species in Psenes are strictly 
oceanic, and large specimens are rarely 
seen. Longley and Hildebrand ( 1941 ) re- 
port the remains of 120-mm P. cijanophnjs 
from bird rookeries in the Tortugas. Other 
species found there included Monacanthus 

86 BuUcfiu Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 29. Branchial region of Psenes cyanophrys, drawing of a cleared-and-stoined preparation from a 128-mm specimen. 
Elements identified in Figure 2. 

lii.spichis, Trachurops crumenoptJialmo, and 
Caranx ruber, all fishes commonly associ- 
ated with floating Gulf weed. 

In the Caribbean area, the monthly dis- 
tribution of post-larval and juvenile Psenes 
cyanophrys suggested that spawning oc- 
curred from March through October 
( Legaspi, 1956 ) . The young fish fed at 
first on copepods, but later a variety of 
foods — copepods, amphipods, chaetognaths, 
fish eggs and larvae — was taken (Lloyd, 
1909; Legaspi, 1956). In large P. pellucidus, 
a sombre-colored probably mesopelagic spe- 
cies, I have found gonostomatids of the 
genus Mauroliciis. 

Relationships. Psenes, like N omens, is a 
derivative of the central Cubiceps stock. In 
Psenes, the number of median finrays and 
vertebrae has tended to increase, the first 
interhaemal has moved forward fonning an 
abrupt angle with the haemal process of the 
first precaudal vertebra, the teeth on the 
glossohyal have disappeared, the tooth- 
bases in the pharyngeal sac have become 
smaller, and the body has become deeper. 
In the meso- or bathypelagic species the 
teeth are highly differentiated. Those in the 
lower jaw are long and knifelike, while 
those in the upper jaw are small and 
strongly recurved. In many species of 

Stromateoid Fishes • Hacdrich 87 


Figure 30. Branchial region of Psenes pellucidus, the type species for the nominal genus /cficus, drawing of a cleared-and- 
stained preparation from a 166-mm specimen, ABE 60-106. Elements identified in Figure 2. 

Psenes, the teeth in the lower jaw are 

Psenes has given rise to no other forms, 
although species such as Psenes pellucidiis, 
which have entered the bathypelagic realm, 
seem to be diverging rapidly from the cen- 
tral bauplan. This fish has been described 
as a separate genus, Icticus. Were it not 
for the great similarity between the struc- 
ture of the pharyngeal sacs of this nominal 
genus and Psenes (Figs. 29, 30), and for 
the presence of species intennediate in 
other characters, Icticus would stand as a 
genus derived from Psenes. 

Species. Psenes is widespread in the tem- 
perate and tropical parts of the Atlantic, 
Pacific, and Indian oceans. There are nu- 
merous records of the genus from the Gulf 

of Mexico and the Caribbean, but none 
from the Mediterranean Sea. 

As in Cubiceps, there is some confusion 
surrounding the species of Psenes. Most 
seem to be world-^\ide, with minor differ- 
ences from ocean to ocean. Larger collec- 
tions and more extensive series than those 
now available are necessary for proper com- 
ment on the significance of these differ- 

The nominal species in the genus are: 
*Psenes cyanophiys Cuvier and Valen- 
ciennes, 1833. Atlantic, Pacific, and Indian 
oceans, type locality New Ireland. D IX- 
XI, 24-28. A III 24-28. P 17-20. Gill-rakers 
8-9 + 1 + 19. Vertebrae 13 + 18. The counts 
recorded for specimens from the Gulf of 
Mexico ( Legaspi, 1956 ) are modally higher 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

than those of Pacific specimens. A very 
characteristic feature of this species is the 
numerous longitudinal streaks on the sides. 
According to Le Danois ( 1962 ) , the follow- 
ing are synonyms: Pscnes jovanicus Cuvier 
and Valenciennes, 1833, from Java, Psenes 
aurafus Cuvier and Valenciennes, 1833, 
from Guam, and Psenes ftiscus Guichenot, 
1866, from Madagascar. Psenes leucurus 
Cuvier and Valenciennes, the color of which 
is described by the authors (1833:265) as 
"jaunatre, marbre de noiratre et finement 
raye de traits longitudinaux noiratres," is 
also probably a synonym, but Le Danois 
(1962) does not agree. Other synonyms 
are: *Cubiceps niultimdiatus Gimiher, 1871, 
from Manado, Philippines; Psenes chapmani 
Fowler, 1906, from the Cape Verde Islands; 
Psenes pocifieus Meek and Hildebrand, 
1925, from the Bay of Panama; and Psenes 
kamoharai Abe, Kojima, and Kosakai, 1963, 
from Kyushu. 

*Psenes pelhieidtis Liitken, 1S80. Atlan- 
tic, Pacific, and Indian oceans, type locality 
Surabaja Strait, Java Sea. D X-XI, I-II 27- 
32. A III 26-31. P 18-20. Gill-rakers 8-9 
+ 1 + 14-16. Vertebrae 15 + 26-27. This 
meso- or bathypelagic species is immedi- 
ately recognizable by the soft, flabby mus- 
culature, the long knifelike teeth in the 
lower jaw, the sombre coloration, and the 
high vertebral and median finray counts. 
With growth, this species becomes quite 
slender. Synonyms are: *Psenes edtcardsii 
Eigenmann, 1902, from the Atlantic Ocean 
south of Rhode Island; '^Ictieus ischanus Jor- 
dan and Thompson, 1914, from Okinawa; 
Caiistioides ampUpinnis Whitley, 1948, 
from Lord Howe Island, and Cidneeps 
ismaelensis Dieuzeide and Roland, 1955, 
from the coast of Algeria. 

*Psenes macuhtns Liitken 1880. Atlan- 
tic, Pacific, and Indian oceans, type locality 
central Atlantic Ocean. D IX-XI, I 22-24. 
A III 21-23. P 20-21. Vertebrae 15 4- 18- 
20. Psenes ni^rescens Lloyd, 1909, from the 
Andaman Sea is a possible synonym. 

*Psenes arafurensis Giinther, 1889. Atlan- 
tic, Pacific, and Indian oceans, type locality 

Arafura Sea. D X-XI, I-II 19-21. A III 20- 
21. P 18-20. Gill-rakers 7-9 + 1 + 15-16. 
Vertebrae 13 + 18. This species is very 
similar to *P. moculatus Liitken, 1880, but 
the body is deeper at comparable sizes. The 
teeth in the lower jaws of both nominal 
species are like those of *P. pellueidus. P. 
rotundtis Smith, 1949, from South Africa is 
a probable synonym. 

Psenes wlutcle^gii Waite, 1894. Coasts of 
New South Wales, type locality Maroubra 
Bay. D XI, I 19. A III 18. P 18. (From 
Waite, 1894.) Psenes hiUii Ogilby, 1915, 
from Queensland is a proliable synonym. 

Psenes guttatns Fowler, 1934a. Coast of 
Natal. D XI, I 20. A III 18. Gill-rakers 7 
+ 18. (From Fowler, 1934a). This name 
appears as a nomen nudum in Fowler 
(1906). Psenes stigmapleuron Fowler, 1939, 
is a synonym. Perhaps a synonym of P. 
liliiteleggii Waite, 1894. 

*Psenes henardi Rossignol and Blache, 
1961. Gulf of Guinea. D XI, I 19-20. A III 
21-22. P 19. Gill-rakers 9 + 1 + 16-17. *P. 
arafurensis Giinther, 1889, has a much 
deeper body, and is striped and mottled on 
the sides. P. henardi is uniform light brown. 
It is perhaps a svnonym of P. guttatus Fow- 
ler, 1934a. 

Family ARIOMMIDAE, new family 

Type genus: Ariomma Jordan and Snyder, 


Diagnosis. Stromateoid fishes with pelvic 
fins present in adults, two dorsal fins, tooth- 
less palate, six branchiostegal rays, two hy- 
pural bones in the tail, and well ossified 
sclerotic bones. Bases of papillae in the 
pharyngeal sacs round; papillae not in bands 
and in upper halves of the sacs only. 

Deseription. Body slender or deep, 
rounded to compressed. Peduncle slender, 
with two low fleshy lateral keels on each 
side. Two dorsal fins. The first dorsal with 
about ten slender spines folding into a 
groove. The longest spine twice the length' 
of the longest ray of the second dorsal fin. 
Three anal spines, not separated from the 
rays. Soft dorsal and anal fins approxi- 

Stromateoid Fishes • Haedrich 



Figure 31. Bronchial region of Anomma cf. nigriorgenfeo, 
an elongate species, drawing of a cleared-and-sfained prep- 
aration from a 140-nrim specinnen. Elements identified in 
Figure 2. 

mately the same length, each with 14 or 15 
finrays, the large basals protruding into the 
body profile. Bases of median fins not 
sheathed by scales. Pelvic fins attached to 
the abdomen by a thin membrane and fold- 
ing into a groove. Scales large, cycloid, 
thin, extremely deciduous. Lateral line 
high, following dorsal profile and not ex- 
tending onto peduncle; tubes in the lateral 
line scales sometimes branched. A branch 
of the lateral line extending forward over 
the eye in a bony tract. Skin thin; subder- 
mal mucous canal system well developed. 
Opercular and preopercular margins entire 
or very finely denticulate. Opercle very thin, 
brittle, with two weak, ill-defined, flat 
spines. Six branchiostegal rays. Mouth 
small, maxillary barely extending to below 
eye. Teeth small, simple or three-cusped, 
uniserial in the jaws. Vomer, palatines, and 
basibranchials toothless. Supramaxillary 
bone absent. Eye large, adipose tissue well 
developed and covering the lacrimal bone. 
Sclerotic bones well ossified. Vertebrae 29 
to 32. Caudal skeleton with tvvo hypurals 
and three epurals. Pharyngeal sacs with 
papillae in the upper halves only. The pa- 
pillae not in bands, their bases rounded 
with a stalk with teeth seated all along it 
arising off-center. Adults usually about a 
foot long, but in some species exceeding 
two feet. Silvery to blue-brownish, some 


Figure 32. Brancfiial region of Anomma indica, a deep- 
bodied species, drawing of a cleared-and-stained prepara- 
tion from a 164-mm specimen, NTU 51942. Elements iden- 
tified in Figure 2. 

species with conspicuous spotted or counter- 
shaded pattern. 

Distri])ntion. Ariommids apparently live 
near bottom in deep water of the subtropics 
and tropics. They occur along the east 
coast of North America, in the Gulf of Mex- 
ico and Caribbean Sea, along the coasts of 
West and South Africa, along Asian coasts 
from the Red Sea to Japan, and off Hawaii 
(Fig. 55). 

Relationships. Ariomma, the single genus 
in the family, superficially appears to be a 
nomeid. The two dorsal fins, persistent pel- 
vies, and six branchiostegal rays have been 
the cause for placement in this group close 
to Ciibiceps (Psenidae of Jordan and 
Snyder, 1907; Nomeidae of Katayama, 1952). 
Some authors (Regan, 1914a; Jordan, 1923) 
have even considered Ariomma a synonym 
of Cubiceps. But the complete absence of 
teeth on the vomer, palatines, and basi- 
branchials in Ariomma contrasts with the 
situation in the Nomeidae. The structure 
of the caudal skeleton ( Fig. 33 ) and of the 
pharyngeal sacs (Figs. 31, 32) in Ariomma 
is unique among stromateoids, and diver- 
gent enough from any others to warrant 
separation at the family level. The unique 

90 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



HYPURAL 4 + 5 + 6 

HYPURAL 1 + 2 + 3- 

Figure 33. Caudal skeleton of Ariomma cf. n/griorgenfeo, drawing of a cleared-and-stained preparation from a 28-mm 
specimen. All elements identified in Figure 1. 

characters of the family, far from the con- 
dition of others in the suborder, are prob- 
ably due to the shift the ariommids have 
made to an adaptive zone atypical for stro- 
mateoids (see below: Natural history, p. 

Ariomma presents a confusing array of 
characters which could suggest affinities 
with the centrolophids, the nomeids, or the 
stromateids. The well ossified sclerotic 
bones, minute body pores, and extremely 
deciduous scales are characters held in 
common with the advanced Seriolella group 
of the centrolophids and the diminutive 
Cuhiccps pauciradiatus group of the no- 
meids and the stromateids. The teeth on 
the papillae occur all along the bony stalk, 
and the jaw teeth of Ariomma indica are 
cusped. Both characters are typical of stro- 
mateids. The general body shape is like 
Seriolella. The complete absence of palatal 

dentition suggests an affiliation with the 
line connecting the advanced centrolophids 
with the stromateids. 

However, Ariomma has two distinct dor- 
sal fins and very large scales, and the bony 
bridge over the anterior vertical canal of 
the ear is either very reduced or absent, 
all in marked contrast to the situation in 
centrolophids and stromateids. These con- 
ditions are found in the Cuhiceps pauci- 
radiatus group, nomeids which in addition 
have reduced palatal dentition. The Ariom- 
midae are probably derived from some- 
where in this line, and have lost the teeth 
on the palate and basibranchials. 

Genus ARlOtAhAA Jordan and Snyder, 1904 

Figures 34, 35 
Ariomma Jordan and Snyder, 1904:942. (Type 
.species: Ariomma hirida Jordan and Snyder, 
1904:943, by original designation. Honolulu, 
Hawaii. ) 

Stromateoid Fishes • Haedrich 


Figure 34. Anomma ledanoisi, an elongate species, drawing of a 189-mm specimen, from Poll, 1959. 

Paracithiceps Belloc, 1937:356. (Type .species: 
Paracuhiceps ledanoisi Belloc, 1937:356, by 
original designation. Coast of West Africa. ) 

The combination of slender caudal pe- 
duncle with lateral keels, deeply forked, 
stiff caudal fin, about fifteen dorsal and anal 
finrays with their basal elements pro- 
nounced and entering into the body profile, 
well-developed adipose tissue around the 
eye, two dorsal fins, and toothless palate 
distinguishes Ariomma from all other stro- 
mateoid genera. The name is a feminine 
noun of unknown derivation. 

Description. Body either elongate, maxi- 
mum depth about 25 per cent of the stan- 
dard length, rounded, or deep, maximum 
depth 40 per cent of the standard length, 
compressed; musculature firm, often oily. 
Peduncle short, slender, square in cross- 
section, with two ill-defined, low, fleshy 
keels on each side at base of caudal fin. 
Two dorsal fins, scarcely separated. First 
dorsal originating directly over, or a little 
before, insertion of pectoral fin, with about 
ten long, brittle spines, folding into a deep 
groove. Second dorsal fin \\'ith 14 or 15 
rays, each about half as long as the longest 

Di spine; the anterior rays a little longer 
than those which follow. Anal papilla a 
little behind mid-body, in a slit. Anal fin 
originating behind middle of body and be- 
hind origin of the second dorsal fin, two or 
three spines preceding the 14 to 15 rays; 
rays short, the anterior ones the longest. 
Rays of the median fins close-set anteriorly, 
becoming more widely spaced posteriorly. 
Basals of the finrays often projecting above 
the body profile. Pectoral fin rounded in 
the young, becoming pointed with growth; 
relative length decreasing slightly with 
growth of elongate form but increasing 
markedly with growth of deep-bodied form. 
Pelvic fins inserting under end or behind 
base of pectoral fin, attached to abdomen 
with a membrane and folding into a pro- 
nounced groove which reaches to the anus. 
Caudal fin stiff, deeply forked, rays on the 
leading edge stiff and spinelike. Scales 
large, cycloid, very thin, extremely decidu- 
ous, not covering bases of the median fins. 
Scales of the lateral line \\'ith branched 
tubes, located high on the body, following 
dorsal profile and not extending onto pe- 
duncle. A branch of the lateral line ext(Mid- 

92 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 35. Ariomma regu/us, a deep-bodied species, from McKenney, 1961. 

ing forward over the eye in a bony tract 
from the head of the hyomandibular. Skin 
thin; main subdennal canal along inter- 
muscular septum very apparent, may be 
confused with lateral line; side branches not 
as conspicuous, pores to surface seem to be 
wanting. Head around 30 per cent of the 
standard length. Top of snout naked, pores 
and subdermal canals barely visible. Scales 
extending forward over nape to level of an- 
terior border of the eye. Eye large, bony 
supraorbital ridge pronounced. Adipose tis- 
sue around eye well developed, extending 
forward over the lacrimal and around the 
nostrils. Nostrils near tip of the obtuse 
snout, small, the anterior round, the pos- 
terior a slit. Maxillary scarcely reaching to 
under eye, angle of gape well before eye 
and nearer to tip of snout. Premaxillary not 
protractile. Lacrimal bone transparent, al- 
most completely covering upper jaw when 
mouth is closed, the ventral border of the 
maxillary remaining exposed. Supramaxil- 
lary absent. Jaw teeth minute, covered 
basally with a membrane, usually pointed 
but three-cusped in a few deep-bodied 
forms, uniserial, close set or slightly spaced; 

vomer, palatines, and basibranchials tooth- 
less. Opercle and preopercle thin, brittle, 
margins very finely denticulate or entire; 
opercle rounded, with two weak, ill-defined, 
flat spines; angle of preopercle rounded 
and not bulging backward. Gill-rakers 
slender, half the length of the filaments, 
toothed on inner edge, fairly close set, 
about 19 on lower limb of first arch; no 
rudimentary rakers under the well-devel- 
oped pseudobranch. Six branchiostegal 
rays, four on the ceratohyal, two on the 
epihyal. Scapula prominent. Vertebrae 12 
to 14 + 17 or 18 = 29 to 32. Two hypurals 
and three epurals. Sclerotic bones well os- 
sified. Stomach large, a simple sac; intes- 
tine very long. Pyloric caeca numerous, in 
a dendritic mass. Air bladder present, ex- 
tending the length of the abdominal cavity. 
Color in preservative brown, bluish, or 
silvery. First dorsal blackish, pelvic fins 
dark or clear, other fins usually light. Color 
pattern may be blue above, silvery below, 
the shades not intergrading, uniformly dark, 
or light with dark spots. Young have three 
to five dark vertical stripes. Opercles sil- 
very or blackish. Iris usually golden, di- 

Stromateoid Fishes • Hacdrich 


vided by a dark vertical bar. Mouth and 
gill cavity light or dark. Peritoneum silvery. 

Natural history. Most stromateoids are 
pelagic, but Ariomma is a bottom or near- 
bottom fish of deep water. Very small 
Ariomma are pelagic, for they are taken in 
surface collections, but all large specimens 
reported have been taken in bottom trawls, 
usually at depths in excess of 100 meters 
(Lowe, 1962; McKenney, 1961; Poll, 1959). 
Ariomma apparently schools, for single net 
hauls contain numerous specimens (Poll, 

The pharyngeal sacs, but not the stom- 
achs, of most specimens dissected contained 
grit and mud, indicative of a bottom-feed- 
ing habit. This habit could account for the 
relatively high number of gill-rakers, around 
30, found in Ariomma. The stomachs of 
specimens examined by Poll ( 1959 ) con- 
tained bits of crustaceans and unidentifi- 
able meat. The thick adipose tissue on the 
head probably protects the eyes and nos- 
trils as the fish scoops its prey from the sea 

Most Ariomma mature around 160 to 180 
mm SL (A. regula [McKenney, 1961]; A. 
Ieda7wisi [Poll, 1959]; A. indica; A. nigriar- 
gentea), and probably do not grow much 
larger than this. Very large specimens are 
known from the Red Sea (800 mm, Klun- 
zinger, 1884), Japan (356 mm SL, Abe, 
1954), and Hawaii (635 mm, Fowler, 1923). 
The Japanese species, at least, does not 
seem to mature before reaching this size. 

Species. The problem of delineating the 
species of Ariomma is one of the most per- 
plexing in the entire suborder. Fourteen 
have been described, but all of these, from 
the Gulf of Mexico to Hawaii, have almost 
the same numbers of finrays and gill-rakers. 
Reports of lateral line scale counts vary 
slightly, but the scales are so deciduous in 
Ariomma that this information must be 
viewed with caution. 

The species of Ariomma, with one inter- 
mediate exception, are either elongate, with 
the maximum depth less than 30 per cent of 
the standard length, or deep-bodied, with 

the maximum depth greater than 40 per 
cent of the standard length. It is possible 
that these two groups warrant subgeneric 
recognition, but, pending further study, this 
action is deferred. 

The nominal species in Ariomma are: 

Ariomma regidiis (Poey, 1868) = Psenes 
regidus Poey. Gulf of Mexico to British 
Guiana, type locality Cuba. Deep-bodied. 
D XI, I 15. A III 15. P 21-24. Gill-rakers 
usually 7 + 1 + 15. ( From McKenney, 
1961.) Spotted. Teeth not cusped. 

Ariomma indica (Day, 1870) = *P.sene.s 
indicus Day. India to southern Japan, type 
locality Madras. Deep-bodied. D XI-XIl, 
14-15. A III 14-15. P 21-23. Gill-rakers 
usually 7 + 1 + 15. Vertebrae 12 or 13 + 
18. The pectoral fin of this silvery species 
becomes produced, up to 35 per cent of the 
standard length, with growth. The teeth 
in the hind part of the lower jaw are three- 
cusped; the rest are simple. Psenes extra- 
neiis Herre, 1950, known from a single 
Philippine specimen, is very likely a syn- 

Ariomma brevimanus (Klunzinger, 1884) 
= Ctibiceps brevimanus Klunzinger. Red 
Sea. Elongate, known from a single speci- 
men 800 mm long. D XI, 15. A II 15. P 24. 
( From Klunzinger, 1884. ) 

"^Ariomma lurida Jordan and Snyder, 1904. 
Hawaii. Elongate. D XI-XII, 14-15. A III 
13-14. P 20-21. Gill-rakers 9 + 1 + 20. Ver- 
tebrae 14 + 18. This species is distinguished 
from *A. cvcrmanni Jordan and Snyder, 
1907, by the large eye, greater than 30 per 
cent of the length of the head, and fewer 
pectoral finrays. 

* Ariomma evermanni Jordan and Snyder, 
1907. Hawaii. Elongate. D XI-XII, 15. A 
III 14. P 25. Gill-rakers 9 + 20. Vertebrae 
13 + 18. Eye is less than 28 per cent of the 
length of the head. Attains a large size; 
Cubiceps thompsoni Fowler, 1923 (type 
635 mm long), is a probable synonym. 

Ariomma africami (Gilchrist and von 
Bonde, 1923) = '^Psenes africanus Gilchrist 
and von Bonde. South Africa. Deep-bodied. 
D IX-X, 15. A III 16. P 22. Gill-rakers 8 


Bulletin Muscutn of Comparative Zoology, Vol. 135, No. 2 

+ 1 + 16. Vertebrae 13 + 18. Very similar 
to A. re<iu1m (Poey, 1868). Spotted. Teeth 
not eusped. 

Ariomma dollfusi ( Chabanaud, 1930) = 
'^Cubicc'ps doUfiisi Chabanaud. Gulf of 
Suez. Intermediate, maximum depth of co- 
type 32 per cent of the standard length (112 
mm). D XI-XII, 15. A III 15. P 22. Gill- 
rakers 7 + 1 + 14. Vertebrae 12 + 18. The 
teeth are said to be "comprimees . . . et 
crenelees" (Chabanaud, 1930:520), sug- 
gesting close relationship or synonymy with 
A. indica (Day, 1870). 

'■Ariomma hondi Fowler, 19.30. Grenada, 
British West Indies. Elongate, known from 
the holotype, 79 mm TL. D XI-XII, 14. A 

II 15. Gill-rakers 8 + 15. Lateral line scales 
43. Dark above, light on sides. (From Fow- 
ler, 1930.) Possible synonyms are *A. nigri- 
argcntca and or *A. melana, both of Gins- 
burg, 1954. 

Ariomma (Belloc, 1937) = Para- 
cuhiceps ledanoisi Belloc. West equatorial 
Africa. Elongate. D XI-XII, 14-15. A III 
14-15. P 20-22. Gill-rakers ? + ? + 16-17. 
(From Poll, 1959.) 

Ariomnia niiiriargcntca (Ginsburg, 1954) 
= *Cid)icepii nigriargenteus Ginsburg. Gulf 
of Mexico, Caribbean, and north to Cape 
Cod, type locality Cape Romain, South 
Carolina. Elongate. D XI-XII, 15-16. A 

III 15. P 21-22. Gill-rakers 9-10 + 17-19. 
(From Ginsburg, 1954. ) Vertebrae 13 + 17. 
Said to differ from A. meJana ( Ginsburg, 
1954) by the smaller scales (62 to 68 in 
lateral line), color (blue above, silvery 
below), less scalation on the head, and 
shorter maxillary. These characters, how- 
ever, seem to intergrade. 

Ariomma melana (Ginsburg, 1954) = 
'^Cuhiceps meJanti.s Ginsburg. Gulf of Mex- 
ico, Caribbean, and north to Cape Hatteras, 
tvpe locality Mississippi Delta. Elongate. 
li) XI-XII, 15. A III 14-15. P 21-22. Gill- 
rakers 9-11 + 18-20. Lateral line scales 39- 
56. ( From Ginsburg, 1954. ) Vertebrae ap- 
parently 15 + 16. Uniform dusky l)rown. 

Ariomma midti.'iqiiami.s (Marchal, 1961) 
= *raracuhice]).'i m\dtis(piamis Marchal. 

West equatorial Africa. Elongate. D XI- 
XII, 1.5-16. A III 14-16. P 21-23. (From 
Marchal, 1961.) Gill-rakers 9 + 1 + 18. Said 
to differ from A. ledanoisi (Belloc, 1937) 
by having more scales in the lateral line 
(61-63 t;s. 36-40). 


Type genus: Tetragonurus Risso, 1810 

Tetragonuridiie. Risso, 1826:382 (def. ). Liitken, 
1880:437 (disc, rel. to scombroids). Ramsay 
and Ogilby, 1888:9 (disc, rel. to Atherinidae). 
Resan, 1902:206 (rel. to Stromatcidae). Boul- 
enger, 1904:642 (popular account). Grey, 
1955: 1 ( world-wide revision ) . 

Tetragonurina. Giinther, 1861:407 (def., rel. to 
Atherinidae ) . 

Tetragonuroidei. Berg, 1940:323 (definition); 1955: 
247 (definition). Smith, 1953:53 (review). 

Diagnosis. Elongate stromateoid fishes 
with pelvic fins present in the adults, two 
dorsal fins, teeth on vomer and palatines, 
five or six branchiostegal rays, heavy adher- 
ent keeled scales, and four hypural and two 
epural bones in the tail. Papillae in the 
pharyngeal sacs with rounded bases, not in 

Description. Body slender, rounded. Pe- 
duncle thick, square in cross-section, with 
modified scales forming two prominent 
lateral keels on each side. Two dorsal fins, 
the first with 10 to 20 short spines, folding 
into a groove; the base of the fin as long as 
or longer than the base of the second dor- 
sal. One anal spine, not separated from the 
rays. Soft dorsal and anal fins approxi- 
mately the same length, with 10 to 17 fin- 
rays. Last ray of pelvic fin attached to 
abdomen for its entire length, fin folding 
into a depression. Scales moderate in size, 
ctenoid, with heavy longitudinal ridges, 
very adherent, arranged in a geodesic pat- 
tern around the body. Lateral line slightly 
arched forward, descending to run along 
middle of side and extending onto peduncle; 
no tubed scales. Skin thick; subdermal 
mucous canal system well developed, but 
barely visible. Opercular and preopercular 
margins entire or finely denticulate. Oper- 
cle thick, spines not apparent. Five or six 

Stromateoid Fishes • Haedricii 95 



HYPURAL 4 + 5 


Figure 36. Caudal skeleton of Tefraganurus atlanticus, drawing of a cieored-and-stained preparation from a 66-mm speci- 
men, MCZ 41791. All elements identified in Figure 1. 

branchiostegal rays. Mouth large, maxil- 
lary extending below eye. Teeth moderate 
to large, simple and cusped, uniserial in the 
jaws. Vomer, palatines, basibranchials, and 
usually the tongue with teeth. Supramaxil- 
lary bone absent. Eye large, no adipose 
tissue. Sclerotic bones not well ossified. 
Vertebrae 43 to 58. Caudal skeleton with 
four hypurals and two epurals. Pharyngeal 
sacs with small papillae in upper and lower 
halves; bases of papillae not in bands, 
rounded, central stalk with a few teeth. 
Adults one to two feet in length. Unifonn 
dark brown, with no pattern or counter- 

Distribution. The distribution of the 
Tetragonuridae largely parallels that of the 
nomeids (Fig. 54). Tetragonurids are oceanic 
fishes of tropical, subtropical, and temper- 
ate seas. None have been taken in the east- 
em Mediterranean Sea, the Red Sea, and 
the South and East China seas. 

ReIotionsJu})s. Tetiagomirus, the single 
genus in the family, has teeth on the vomer, 
palatines, and basibranchials. Thus, it is 
affiliated with the nomeid stock. The cau- 
dal skeleton (Fig. 36) is similar to that of 
the nomeids, but is advanced an evolution- 
ary grade in having lost one of the epurals. 
The pharyngeal sacs (Fig. 37) and the 
heavy, keeled scales of Tetragonuriis, how- 
ever, are markedly different from the sacs 
and scales of the nomeids. The pharyngeal 
sacs are exceedingly elongate. The papillae 
are poorly ossified and are very reduced in 
size. The bases of the papillae are rounded, 
as they are in the Ariommidae, but there 
are less than six teeth seated on top of a 
short stalk. The fourth pharyngobranchial 
is very elongate and is fused to the third 
pharyngobranchial. This long bone is 
studded with teeth and extends well back- 
ward into the sac, where it no doubt aids 
both in shredding the food and in support- 
ing the sacs. All nomeids have six branchi- 


Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



\ 'T\ O O 

: 3 1 




^•' "tt ,c 

Figure 37. Branchial region of Tetragonurus cuvieri, drawing of a cleared-and-stoined preparation from a 340-mm specimen, 
CNHM 64218. Elements identified in Figure 2. Tfie upper pfiaryngeal bone is formed by tfie fusion of tfie third and fourth 

ostegal rays; tetragon iirids have either six 
or five. 

The highly differentiated jaw teeth of 
Tetrogominis are very similar to those of 
certain species in the genus Psenes. But 
Tetragonurus has teeth on the tongue, and 
cannot be derived from Psenes. It is very 
likely that Tetragonurus branched off fairly 
early from the nomeid stem and is derived 
from no living nomeid genus. The loss of 
an epural and a branchiostegal ray, the 
heavy keeled scales, the very elongate pha- 
ryngeal sacs with the reduced papillae, and 
the greatly increased number of vertebrae 
are quite divergent from the situation in 
nomeids, and together imply that evolution 
has proceeded independently in these two 
groups for some time. 

The divergent characters of Tetragonurus 
are part of its adaptation to a very particu- 
lar mode of life. Tetragonurus is certainly 
a derived form, and is probably quite unlike 
the ancestral nomeid. The central nomeid 
genus Cuhiceps, like Tetragonurus, has 
teeth on the tongue. The nomeid stock 
from which Tetragonurus arose may have 
been in many respects similar to the pres- 
ent-day Cuhiceps. 

Genus TETRAGONURUS Risso, 1810 

Figure 38 

Tetragonurus, 1810:347. (Type .species: 
Tetragonurus cuvieri Ri.sso, 1810:347, by 
monotypy. Mediterranean. ) 

Ctenodax Macleay, 1885:718. (Type species: 
Ctenodax wilkinsoni Macleay, 1885:718, by 
monotypy. Lord Howe Island, Tasman Sea. 
A synonym of *Tetragonurus atlantieus Lowe, 

The combination of elongate body and 
peduncle, modified scales forming two keels 
on the peduncle, origin of first dorsal 
slightly to well behind pectoral insertion, 
base of first dorsal longer than base of sec- 
ond dorsal, heavy keeled scales, and pecu- 
liar lower jaw with heavy knifelike teeth 
distinguishes Tetragonurus from all other 
stromateoid genera. The name, a mascu- 
line noun, is from the Greek t€t pay ojvo'i, with 
four angles, + ovpd, tail, in reference to the 
shape of the caudal peduncle. 

Description. Body very elongate, maxi- 
mum depth less than 20 per cent of the 
standard length, rounded; musculature firm. 
Peduncle long, thick, with modified scales 
forming two prominent keels at base of 
caudal fin. Two dorsal fins, scarcely di- 
vided. First dorsal originating slightly to 
well behind insertion of pectoral fins, with 
10 to 21 stiff spines folding into a groove, 
the longest spine half the length of the 
longest ray of the second dorsal. Anterior 
rays of the second dorsal the longest, those 
that follow decreasing slightly in length, 10 
to 17 finrays in all. Anus well behind mid- 
body, in a depression. Anal fin originating 
shghtly behind origin of second dorsal fin. 

Stromateoid Fishes • Haedricli 


■—■■■■,,"•" '■■-■-   ■•^~•-■■-■■.^■■■'=^37;i?M9t15^;s<MSsass^ 

Figure 38. Tetragonurus cuvieri, drawing of a 129-mm specimen, from Grey, 1955. 

2 cm 

one short spine preceding the rays. Anterior 
rays the longest, those that follow decreas- 
ing slightly in length, 9 to 15 finrays in all. 
Pectoral tin small, the central rays the long- 
est. Pelvic fins small, inserting behind pec- 
toral fin base and before origin of first dor- 
sal, innermost ray attached to abdomen for 
its entire length. Caudal fin forked. Scales 
moderate in size, with heavy longitudinal 
keels, very adherent, following a geodesic 
pattern around the body. Very small scales 
on bases of median fins. Lateral line usu- 
ally slightly arched anteriorly, descending 
to run along mid-lateral line of body and 
ending on peduncle at origin of keels; no 
tubed scales. Skin thick; sulidennal canals 
cannot be traced. Pores to surface minute. 
Head 30 to 20 per cent of the standard 
length. Top of head and snout naked, small 
pores in naked skin. Scales extending for- 
ward over nape almost to level of posterior 
border of the eye. Eye large, with a series 
of grooves on the posterior rim. Nostrils 
toward top of blunt snout, large, well sepa- 
rated, the anterior round, the posterior a 
slit. Maxillary ending under eye, angle of 
gape well before eye. Premaxillary not pro- 
tractile. Lacrimal bone covering most of 
upper jaw at all times, ventral border of 
maxillary remaining exposed. Lower jaw 
almost completely within upper jaw when 
mouth is closed. Supramaxillary absent. 
Teeth in upper jaw small, pointed, recurved, 
spaced. Teeth in lower jaw large, knifelike, 
close set, with small cusps, deeply em- 
bedded in the gum with only the tips show- 
ing. Strong recurved teeth present on head 
of vomer, and in a single series on shaft of 

vomer and palatines. Small teeth on basi- 
branchials and, usually, profusely scattered 
on tongue. Tongue high-sided, depressed 
in center. Opercle and preopercle thin, 
fleshy, scaled, margins entire in adult, 
spinulose in the young; opercle with two 
very weak spines; angle of preopercle 
slightly rounded, not bulging backward. 
Cheeks scaled. Gill-rakers broad, fleshy, 
shorter than the filaments, toothed on inner 
edge, spaced, 8 to 14 on lower limb of first 
arch; rudimentary rakers often present un- 
der the large pseudobranch. Scapula not 
visible. Vertebrae 43 to 58. Sclerotic bones 
well ossified in adult. Stomach a simple 
sac; intestine long. Pyloric caeca numerous, 
in a large dendritic mass. 

Color in preservative uniform brown, 
ranging from tan to almost black. Fins the 
same color as the body. Inside of mouth, 
gill cavity, and peritoneum dark. 

Natural history. Young Tetragonurus have 
been found associated with medusae ( Man- 
sueti, 1963). There are also reports (Emery, 
1882; Lo Bianco, 1909; Fitch, 1949) of 
young specimens found within salps, usu- 
ally Pyrosoma. Mansueti (1963) felt that 
the association of Tetragonurus with jelly- 
fishes was a chance occurrence, but, since 
jellyfish associations are commonly formed 
by other stromateoids, it is likely that the 
association is actively sought. 

Tetragonurus probably feeds almost ex- 
clusively on coelenterates and ctenophores 
(Risso, 1826; Fitch, 1952). The large sHc- 
ing teeth of the lower jaw and the peculiar 
boxlike jaw seem admirably suited for such 
a diet (Grey, 1955). 


Bulletin Musciini of Comparative Zoology, Vol. 135, No. 2 

Risso (1826) reported that the flesh of 
Tetragonurus was poisonous, attributing 
this quahty to the fish's diet of venomous 
jell\'fishes of the genus Stcphanomic. His 
report has been widely spread, but has only 
recently been reinvestigated. Fitch ( 1952 ) 
analyzed four California specimens and 
found them to be not poisonous. The pos- 
sibility remains that Tetragonunis is poi- 
sonous only during certain seasons, for 
example at the time of spawning ( Petit and 
Amar, 1946). 

In the Atlantic, Tetragonurus cuvieri ap- 
parently spawns in spiing and summer 
(Grey, 1955). Guiglia (1950) reports ma- 
ture females of T. cuvieri taken throughout 
the year in the Mediterranean. T. atlonticus 
spawns during the fall in the eastern and 
northern Atlantic, but in winter and spring 
in the western Atlantic (Grey, 1955). 

Tetragonurus is strictly oceanic (Grey, 
1955). Although young specimens occur 
near the surface with jellyfish, the adults, 
judging from their somber coloration, prob- 
ably are members of the meso- or ]:)athy- 
pelagic faunas. Tetragonurus has been 
considered a rare fish (Fitch, 1949), but 
recent surveys in the North Pacific (Lark- 
ins, 1964) indicate that it is much more 
common than is usually thought, bearing 
out a prediction of Grey ( 1955). 

Species. Tetragonurus has been ably 
treated by Marion Grey (1955), who rec- 
ognized three species in the genus. As she 
has pointed out, each species varies widely, 
and further division may be justified when 
more specimens are available. The species 

T. cuvieri Risso, 1810. Mediterranean Sea, 
Atlantic, Pacific oceans. D XV-XXI, 10-17. 
A I 10-15. P 14-21 (?). Lateral line scales 
97-114. Vertebrae 52-58. (From Grey, 
1955. ) 

*T. atlanticus Lowe, 1839. Atlantic, Pa- 
cific, Indian oceans. D XIV-XVII, 10-13. 
A I 9-12. P 14-18. Lateral line scales 83- 
95. Vertebrae 45-51. (From Grey, 1955.) 
Ctenodax icilkinsoni Macleay, 1885, is a 

T. pacificus Abe, 1953. Pacific, Indian 
oceans. D X-XI, 10-12. A I 10-12. P 15- 
17. Lateral line scales 73-78. Vertebrae 
40(?)-43. (From Grey, 1955.) 


Type genus: Sfromateus Linnaeus, 1758 

Stromatees. Cuvier and Valenciennes, 1833:372 

( in part, descr. ) . 
Stromateina. Giinther, 1860:397 (in part, def. ). 

Gill, 1862:126 (genera listed). 
Stroniateinae. Gill, 1884:669 (def., gen.). Bi\h- 

ler, 1930:62 (digestive system). 
Stromateidae. Jordan and Gilbert, 1882:449 

(descr.). Jordan and Evermann, 1896:964 

(descr., North America). Jordan, 1923:182 

(list, + Pampidae). Berg, 1940:323 (dist.); 

19.5.5:248 (dist.). 

Diagnosis. Deep-bodied stromateoid 
fishes with pelvic fins absent in the adults, 
continuous dorsal fin, toothless palate, four 
hypural and two or three epural bones in 
the tail, and well ossified sclerotic bones. 
The papillae in the pharyngeal sacs not in 
bands, in both the upper and lower halves 
of the sac; bases of the papillae stellate. 

Description. Body deep, compressed. 
Single dorsal and anal fins, with none or 
one to ten flat, bladelike spines and three 
to five slender, graduated spines preceding 
the rays. Median fins about the same length, 
usually falcate; caudal fin deeply forked. 
Pectoral fin long and pointed. Pelvic fins 
present only in young Stromateus; absent 
in all others. Scales small, cycloid, ex- 
tremely deciduous. Lateral line high, fol- 
lowing dorsal profile, and extending onto 
the short peduncle. Opercular and preoper- 
cular margins entire. Opercle very thin, 
with two short, flat, weak spines. Gill mem- 
branes usually free from isthmus, but united 
in Pampus. Five or six branchiostegal rays. 
Mouth temiinal to sub-terminal, small, an- 
gle of gape rarely reaching below eye. 
Teeth very small, laterally flattened, with 
three minute cusps, and uniserial in the 
jaws. Vomer, palatines, and basibranchials 
toothless. Supramaxillary absent. Eye fairly 
small; adipose tissue usually not well de- 
veloped. Sclerotic bones well ossified. Ver- 

Stromateoid Fishes • Haedrich 


tebrae 30 to 48. Caudal skeleton with four 
hxpurals and t\\o epurals, except three 
epurals in Stromafeus fiatola. Pharyngeal 
sacs with papillae in upper and lower halves, 
not in bands; bases of the papillae stellate, 
with teeth seated all along a central stalk. 
Adults usually about a foot in length. Sil- 
very to blue, some with spots. 

Distribution. Stromateids live over the 
continental shelves and in the bays of tropi- 
cal, subtropical, and temperate waters. 
They are found on the east and west coasts 
of North and South America, from the 
Mediterranean Sea to South Africa, and 
from the Iranian Gulf to Japan (Fig. 56). 
None occur near oceanic islands, and none 
have reached Australia. Stromateus, in 
southern South America and western Africa, 
is the only genus that has representatives 
on both sides of an ocean. The genera are 
allopatric except for a small area of overlap 
between Stromateus and PepriJus in South 
America. All stromateids school, and many 
are important commercially. 

Relationships. The Stromateidae are the 
current zenith in stromateoid evolution. 
The reduced number of branchiostegals and 
elements in the caudal skeleton (Figs. 42, 
47), the absence of pelvic fins, the small 
mouth with cusped teeth, the broad stellate 
bases of the papillae in the pharyngeal sacs 
(Figs. 43, 46), and the deepened body are 
all advanced conditions. 

The stromateids were derived from some- 
where near the Seriolella group of the Cen- 
trolophidae, perhaps from a fish very like 
the deep-bodied Psenopsis. In Stromateus 
fiatola the presence of peKic fins in the 
young and the three epurals recalls the 
centrolophid heritage. 

The stromateids may have evolved only 
recently. Though the three genera are dis- 
tinct, little speciation has occurred, but, in 
some cases, seems to be in an incipient 
stage. Almost-sibling species occur on 
either side of the Isthmus of Panama. 

The stromateids are an end-point, and no 
other groups have been deri\ed from them. 
But they are by no means an evolutionar>' 

deadend. Pampus, with its restricted gill- 
opening, elongate pharyngeal sac, and pe- 
culiar spines in some species, is diverging 
rapidly from the central stromateid bauplan 
and is widely successful along the coasts of 
southern Asia. 

Key to Stromateid Genera 

1 (4). Inter- and subopercles not united to 

the isthmus. End of maxillary before 
or at anterior border of eye. Cusps 
on teeth in lower jaw subequal, the 
teeth appearing truncate to the naked 
eye. Spine on end of pelvic bone 
present or absent. In small specimens 
( less than 80 mm SL ) pelvic fins 
present or absent. Six branchiostegal 
rays. 2 

2 (3). One to three flat, bladelike spines 

ahead of median fins. A small spine 
projecting posteroventrally from end 
of pelvic bone. Median fins falcate 
or not. Pelvic fins never present. 30 

to 35 vertebrae. 

Peprilus, p. 103. Figs. 40, 41 

3 (2). No flat, bladelike spines ahead of 

median fins. No spine at end of 
pelvic bone. Median fins never fal- 
cate. Pelvic fins absent in adult, but 
present in some small specimens. 40 

to 48 vertebrae. 

Stromateus, p. 99. Fig. 39 

4 (1 ). Inter- and subopercles broadly united 

to isthmus. End of maxillar>' under 
eye. Central cusp on teeth of lower 
jaw much larger than the other two 
cusps, which can hardly be seen with- 
out extreme magnification. No spine 
at end of pelvic bone. Pelvic fins 
never present. Five branchiostegal 
rays. Pampiis, p. 108. Figs. 44, 45 

Genus STROMATEUS Linnaeus, 1758 

Figure 39 

Stromateus Linnaeus, 1758:248. (Type species: 
Stromateus fiatola Linnaeus, 1758:248, by 
monotypy. Mediterranean. ) 

Chrysostromus Lacepede, 1802:697. (Type spe- 
cies: Chrysostromus fiatoloides Lacepede, 
1802:697, by monotypy. Mediterranean. A 
synon>Tn of Stromateus fiatola Linnaeus, 1758: 

Fiatola Cuvier, 1817:342. (Tvpe species: Stroma- 
teus fiatola Linnaeus, 1758:248, by monotypy. 
Mediterranean. ) 

Seserinus Cuvier, 1817:342. (Type species: "Seser- 
imts Rondelet" [Seserinus rondeleti] Cuvier, 
1817:343, by subsequent designation of Jor- 

100 Bulletin Museum of Comparatwe Zoology, Vol. 135, No. 2 







Figure 39. S/romofeus fiatola, drawing of a 228-mm specimen, from Poll, 1959 


dan, 1923:106. Mediterranean. A synonym 
of Stwmatcus fiatola Linnaeus, 1758:248.) 
Ptcrorhomhus Fowler, 1906:118. (Subgenus. Type 
species: Fiatola fasciata Risso, 1826:289, by 
original designation. Mediterranean. A syn- 
onym of Stwmatcus fiatola Linnaeus, 1758: 

The combination of deep body, small 
eye, moderate pectoral fin, no bladelike 
spines ahead of the median fins, no ventral 
spine on the pelvic bone, spotted body pat- 
tern, and (sometimes) pelvic fins in the 
young distinguishes Stromateus from all 
other stromateoid genera. The name, a mas- 
culine noun, is from the Greek (TTpwfiaTev;, 
a brightly colored quilt or bedding, prob- 
ably in reference to the shape and pattern 
of the fish. 

Description. Body deep, maximum depth 
generally greater than 40 per cent of the 
standard length, compressed; musculature 
firm. Peduncle very short, compressed. Dor- 
sal fin continuous, originating over base of 
the pectoral fins, the anteriormost elements 

usually very small and buried in the skin, 
44 to 53 elements in all. Anal papilla before 
mid-body, in a deep slit. Anal fin origi- 
nating slightly before middle of body but 
behind dorsal origin, the small anteriormost 
elements buried in the skin, 35 to 47 ele- 
ments in all. Anterior rays of the median 
fins produced, two to three times longer 
than the subequal rays of the posterior two- 
thirds of the fin, the lobes rounded and not 
falcate. No bladelike spines preceding me- 
dian fins. Pectoral fin moderate in length, 
broad. Pelvic fins present in the young of 
at least some species, inserting under end 
of pectoral fin base, the fins lost in adult 
but with two dark flaps of skin sometimes 
indicating their former presence. Pelvic 
bone usually not visible on mid-line and 
lacking a ventral spine. Caudal fin stiff, 
deeply forked, the lobes very long. Scales 
small, cycloid, deciduous, minute scales 
covering all fins. Simple tubed scales of the 
lateral line moderately high, following dor- 

Strom ATEOiD Fishes • Hacdrich 101 

sal profile and extending onto peduncle but 
not to caudal base. Skin moderately thick, 
subdermal canals not visible, pores to sur- 
face seem to be wanting. Head around 25 
per cent of the standard length, very deep 
and broad. Top of head naked, small pores 
easily seen, naked skin underlain with nu- 
merous parallel canals projecting slightly 
backward over the nape. Eye small, adi- 
pose tissue around eye well developed and 
extending forward surrounding the nostrils. 
Nostrils moderate in size, the anterior 
round, the posterior a slit, located nearer 
to tip of blunt snout than to eye. Mouth 
broad. Maxillary scarcely reaching anterior 
border of the eye, angle of gape well before 
eye. Premaxillary not protractile. Lacrimal 
bone reduced, scarcely covering top of up- 
per jaw when mouth is closed, end of maxil- 
lary exposed. Supramaxillary absent. Jaw 
teeth minute, uniserial, laterallv flattened, 
with three subequal cusps, close set, cov- 
ered laterally by a membrane; vomer, pala- 
tines, and basibranchials toothless. Gill 
membranes broadly united across the isth- 
mus, divided from level of the back part of 
the eye. Opercle and preopercle thin, 
scaled, margins entire; opercle rounded, 
with two ill-defined, weak spines; angle of 
preopercle broadly rounded, projecting 
backward slightly. Cheek scaled. Gill- 
rakers a little less than half the length of 
the filaments, diminishing in size anteriorly, 
not toothed, fairly close-set, about 12 on the 
lower limb of the first arch; no rudimentary 
rakers under the small pseudolnanch. Six 
branchiostegal rays, four on the ceratohyal, 
two on the epihyal. Scapula not visible. 
Vertebrae slightly variable, usually 16 + 26 
= 42 to 19 + 26 = 45. Sclerotic bones well 
ossified. Stomach a simple sac; intestine 
very long. Pyloric caeca numerous, in a 
long dendritic mass. 

Color in preservative brown or bluish 
with a silvery or \\4iitish overlay, dark 
above, lighter below. Back and sides with 
numerous dark spots. The young may have 
four or five dark vertical bands. Fins 
darker or lighter than the body; pectoral in 

some species blackish. Gill cavity, inside 
of mouth, and peritoneum light. 

Natural history. The young of Stromatetis 
commonly associate with medusae (Padoa, 
1956), and Lo Bianco (1909) observed them 
eating jellyfish. Fish up to five inches in 
length have been reported in association 
(Smith, 1949a), but the majority probably 
desert their coelenterate host before reach- 
ing this size. 

Poll ( 1959 ) found the adult common in 
depths from 12 to 50 meters off the coast 
of West Africa. I have examined adults 
taken by the Guinean Trawling Survey in 
30 meters of water, and adults from Chile 
captured with a trammel net. Adult Stro- 
mateus may rarely descend to deeper water. 

Small Stromatetis fiatolo have a vertically 
barred pattern and small pelvic fins. The 
bars and the pelvics are lost usually before 
the fish reaches 100 mm standard length. 
At this point, the young fish probably 
moves into the adult habitat. Whether or 
not the young of South American Stroma- 
tetis have pelvic fins is unknown. Poll 
(1959) reports a Stromateus fiatola 500 mm 
long weighing 151 grams; this is probably 
near the maximum size attained. 

In the Patagonian region, Stromatetis 
moves shoreward to spawn in earh' summer 
( Hart, 1946 ) . At this time the fishes form 
their maximum concentrations. Following 
spawning, they move offshore during the 
fall and winter, and become widely dis- 
persed. In addition to inshore and offshore 
movement, Hart ( 1946 ) found evidence 
that Stromatetis moves from lower to higher 
latitudes in the summer, and back in the 

Said to be a fish with "delicate flesh and 
fine flavour" (Gilchrist and von Bonde, 
1923:11), Stromateus apparently does not 
receive the attention it deserves. Once de- 
scribed as numerous in the markets at Lima 
(Cuvier and Valenciennes, 1833), it is to- 
day the subject of only small local fisheries 
in parts of Chile and along the African 

Relationships. Stromateus contains the 

102 BiiUetin Museum of Comparative Zoology, Vol. 135, No. 2 

most primitive species of the family Stro- 
mateidae. Stwinatcus fiafola, the African 
species, has three epural elements in the 
caudal skeleton and, when young, has pel- 
vic fins. These characters indicate the cen- 
trolophid heritage of the stromateids. In 
the South American species of Stromoteus, 
there seem to be only two epural elements 
in the tail, and the young may lack pelvic 
fins.^ This situation is typical of the stro- 
mateid grade. 

Stromateiis has a very high number of 
vertebrae, more than forty, an advanced 
condition. This high number may be a re- 
cently acquired characteristic of the genus. 
The number itself is variable, and, in a 
large proportion of cases, fusions of the 
centra occur in the caudal series. Centra 
\\'ith two or three neural and haemal spines 
appeared in four of the fourteen specimens 
radiographed. The variability and high in- 
cidence of fusions suggests a genetic in- 
stability perhaps correlated with recent 

Because of its high vertebral count, Stro- 
mateus cannot be the direct ancestor of 
either of the other two stromateid genera, 
both of which have lower counts. Rather, 
all three must share a common ancestor, 
a fish most like Stromatetis but with a ver- 
tebral count somewhere near 13 + 17. 

Species. Stromateiis lives in temperate to 
tropical waters along the coasts of Medi- 
terranean countries, West Africa, Argentina, 
and Chile. Though numerous species have 
been described, there seem to be no more 
than two, or possibly three, valid ones. Spe- 
cies are widespread along a coastline, but 
none jump ocean barriers. The species in 
the genus are: 

Stromateiis fiatola Linnaeus, 1758. Medi- 
terranean, coast of West Africa south to 
Capetown, type locality Mediterranean Sea. 
D 48-51 (total elements). A 35-38 (total 
elements). P 22-24. Cill-rakers usually 3 

■■• I have seen no small Stromatetis from South 
America, but the adults lack the two dark flaps 
of skin wlu'ch bespeak the former presence of pel- 
vies in adult African Stromateiis. 

+ 1 + 11. Vertebrae 18-19 + 24-26. Speci- 
mens less than 100 mm SL have pelvic fins 
and vertical bars on the sides. The fins and 
the bars are gone in adults. The names of 
the Mediterranean Fiatola fasciata Risso, 
1826, and Seseriniis microchiriis Cuvier and 
Valenciennes, 1833, are synonyms based on 
juveniles. Other synonyms are Chrijsostro- 
miis fiatoloides Lacepede, 1802, from the 
Mediterranean and Stromateiis capensis 
Pappe, 1866, from South Africa. 

Stromateiis steUatus Cuvier, 1829. Pacific 
coast of South America, Chile and rarely 
north to Lima, type locality coasts of Peru. 
D 44-53 (total elements). A 39-44 (total 
elements). P 19-24. Gill-rakers around 4 
+ 1 + 12. Vertebrae 16 + 26-27. *Stroma- 
teus maculatus Cuvier and Valenciennes, 
1833 (=S. advectitius Whitley, 1935), a 
junior synonym, is the name most often 
used for this fish. *S. maculatus is also gen- 
erally applied to the species of Stromateus 
which occurs along the Atlantic coast of 
South America. The two forms are very 
close in appearance but the Chilean form 
is a slenderer fish with a slightly longer 
head. The counts of the two overlap almost 
completely. The Atlantic fonn seems to 
breed near the northern limit of its range 
(Hart, 1946). Nothing is known concern- 
ing the breeding habits of the Pacific fonn, 
but the spawning area is probably well 
north of Tierra del Fuego. It seems un- 
likely that there is any gene exchange be- 
tween the two populations, and I suspect 
that future study will show that sufficient 
difference exists to warrant recognition of 
both at the species level. The available 
name for the Atlantic population is: 

*Stromateus ])rasilicnsis Fowler, 1906. At- 
lantic coast of South America, Tierra del 
Fuego north to Uruguay, type locality Rio 
Grande do Sul, Brazil. D 47-53 ( total ele- 
ments). A 44-47 (total elements). P 19- 
20. Gill-rakers around 3+1 + 12. Verte- 
brae 16-17 + 27-30. The bionomics and 
potential fishery for this fish are the sub- 
ject of an excellent discussion by Hart 

Stromateoid Fishes • HacdricJi 103 

Genus PEPRILUS Cuvier, 1829 

Figures 40, -U 

Rliumhus Lacepede, 1800:60. (Type species: 
Chaetodon alepidotus Linnaeus, 1766:460, by 
monotypy. Charleston, South CaroHna. Pre- 
occupied by Rhombus Humphrey, 1797, Mol- 
lusca. ) 

Pcprihis Cuvier. 1829:213. (Type species: Sfcr- 
noptyx gardcnii Bloch and Schneider, 1801: 
494, by subsequent designation of Gill, 1862: 
126. Charleston, South Carolina. A synonym 
of Chaetodon alepidotus Linnaeus, 1766:460.) 

Powrwtus Gill, 1861:35. (Type species: Stro- 
juatcus triacanthus Peck, 1804:51, by mono- 
typy. Piscataqua River, New Hampshire. ) 

Palometa Jordan and Evennann, 1896:966. (Sul> 
genus. Type species: *Stromateus palometa 
Jordan and Bollman, 1889:156, by original 
designation. Bay of Panama, Pacific Ocean. ) 

Simohrama Fowler, 1944b:2. (Type species: *Seser- 
inus xanthurus Quoy and Gaimard, 1824:384, 
by original designation. Rio de Janeiro. A 
synonym of Stromateus paru Linnaeus, 1758: 

The combination of deep body, large eye, 
long pectoral fin, one to three bladeHke 
spines ahead of the median fins, a ventral 
spine on the pelvic bone, and no pelvic fins, 
distinguishes Pcprihis from all other stro- 
mateoid genera. The name, a masculine 
noun, is from the Greek TrcTrptAos, one of 
Hesychian's many l^dv^ ttoios, unknown fish. 

Description. Body deep, maximum depth 
35 to 70 per cent of the standard length, 
highly compressed; musculature firm. Pe- 
duncle ver\' short, compressed. Dorsal fin 
continuous, originating just behind insertion 
of the pectoral fin; one to three flat, blade- 
like spines, the first spine pointed on both 
ends, preceding the 30 to 40 finrays. An- 
teriormost rays of the median fins pro- 
duced, the fins often falcate, the rays which 
follow much shorter, diminishing very 
slightly in length to the last ray, the shortest. 
Pectoral fins long, winglike, their bases 
slightly inclined. No pelvic fins. Pelvic 
bone visible on midline of bod\' under the 
end of the pectoral fin base; a small spine 
on the end of the bone projecting postero- 
ventrally through the skin. Tip of coracoid 
sometimes projecting slightly underneath 
the head at about level of margin of the 

preopercle. Caudal fin stiff, deeply forked, 
the lobes long and equal. Scales very small, 
cycloid, thin, very deciduous, extending 
onto all fins. Simple tubed scales of the 
lateral line moderately high, following dor- 
sal profile and extending onto peduncle but 
not to caudal base. A branch of the lateral 
line extending upward from the head of the 
hvomandibular in a short, wide, bonv tract. 
Skin very thin; main subdermal canal along 
intermuscular septum and side branches 
conspicuous, pores to surface very small. 
In some species, a row of large conspicuous 
pores in the back under the first half of the 
dorsal fin. Head around 25 per cent of the 
standard length. Top of head naked, pores 
clearly \dsible, naked skin underlain with 
numerous parallel canals projecting back- 
ward over the nape. Eye large. Adipose 
tissue around the eye developed, extending 
forward and surrounding the nostrils. Nos- 
trils small, the anterior round, the posterior 
a slit, located near tip of the obtuse snout 
at level of top of the eye. End of maxillary 
barely reaching to below eye, angle of gape 
well before eye. Premaxillary not protrac- 
tile. Lacrimal bone reduced, scarcely cov- 
ering top of upper ]a.\y when mouth is 
closed, end of maxillary exposed. Supra- 
maxillary absent. Jaw teeth minute, uni- 
serial, laterally compressed, with three sub- 
equal cusps, close set, covered laterally by 
a membrane. Vomer, palatines, and basi- 
branchials toothless. Gill membranes united 
across the isthmus, divided from about le\el 
of the forward part of the eye. Opercle and 
preopercle thin, not scaled, preopercle finely 
striated, opercle smooth, margins entire; 
opercle with two ill-defined flat spines; 
angle of preopercle rounded, not projecting 
backward. Cheek not scaled. Gill-rakers a 
little more than half the length of the fila- 
ments, with fine teeth on the inner edge, 
the rakers close set, about 18 on the lower 
limb of the first arch; no rudimentary 
rakers under the small pseudobranch. Six 
branchiostegal rays, four on the ceratohyal, 
two on the epihyal. Scapula visible. Ver- 
tebrae variable, 13 + 17 = 30 to 12 4- 23 = 

104 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 40. Pepnlus triacanthus, drawing of a 7.5-inch specimen, courtesy of the Smithsonian Institution. 

35. Sclerotic l)oncs well ossified. Stomach 
a simple sac; intestine very long. Pyloric 
caeca verv numerous, in a dendritic mass. 

Color in preservative brownish, often 
with a silvery overlay, dark above, lighter 
below. Back and sides sometimes with 
spots. Median fins darker or lighter than 
the body; pectoral light. Gill cavity, inside 
of mouth, and peritoneum light. 

Naturol history. The species of Pcprihis 
are prized for food wherever they occur. 
The biggest fishery is for P. triaconthus, 
which is found along the east coast of the 
United States. In some years, more than 
two million pounds of this species are landed 
in Massachusetts alone (Bigelow and 
Schroeder, 1953). Considering its commer- 
cial importance, surprisingly little is known 
of the habits of Peprilus. Almost all our 
knowledge of the natural history of the 
genus comes from a few general studies on 
P. triacanthus. The discussion here is based 
largely on the excellent review of Bigelow 
and Schroeder (1953). 

Though young P. triacanthus are some- 
times found with Cyanea, they do not seem 

to associate with medusae as actively as 
some other stromateoids. The small fishes 
are just as often observed swimming inde- 
pendently at the surface or clustered under 
floating Gulf weed. A much stronger as- 
sociation is formed by P. alepidofus with 
the sea nettle Chrysaora in Chesapeake Bay. 
P. alepidotus feeds actively on the medusa 
(Mansueti, 1963). 

Pcprihis triacanthus spawn in Massachu- 
setts waters from summer into early fall. 
Spawning takes place a few miles offshore, 
but, except at this time, the adults are usu- 
ally closer to shore in water less than 30 
fathoms deep. The eggs are pelagic and, 
at a temperature of 65 °F, hatch in less than 
two days. During their first summer, the 
young fish may grow to a length of three 
or four inches. They probably mature when 
about two years old at a length of seven 
inches. A large adult is no more than a foot 

During th(> late fall, schools of Pcprihis 
triacanthus apparently move offshore where 
they winter near bottom in about 100 fath- 
oins. Though sometimes occurring as far 

Stromateoid Fishes • Hacdrich 


Figure 41. Peprilus alepidotus, drawing of a 7.5-inch specimen, courtesy of the Smithsonian Institution. 

north as Newfoundland, P. triacanthus, like 
all members of the genus, is basically a wami 
water fish. 

Relationships. Peprilus is more advanced 
than Stromatcus in that all species lack pel- 
vic fins and have only two epural elements 
in the caudal skeleton (Fig. 42). But it is 
more primitive than Pampus in having six 
instead of five branchiostegal rays and a 
shorter pharyngeal sac (Fig. 43; cf. Fig. 46). 
The few spines before the median fins in 
Peprilus are very similar in fonu to the 
more numerous spines of some Pampus, but 
the similarity is probably due to common 
heritage rather than to direct ancestry. It 
is unlikely that Pampus is derived from 
Peprilus. The most primitive species in 

Pampus has no spines before the median 
fins and shows no trace of the pelvic spine 
so characteristic of Peprilus. Peprilus is de- 
rived from a fish somenhat like Stroma- 
teus, but probably with fewer vertebrae. 

Species. Peprilus is widespread, with a 
number of species along both the Atlantic 
and Pacific coasts of the New World north 
of the equator. One species is found as far 
south as Montevideo on the east coast of 
South America. Peprilus has been divided 
into several nominal genera, but the dif- 
ferences on which these are based — depth 
of body, fins falcate or not, certain pores 
well developed or not — are the differences 
between species, not genera. Osteologi- 

106 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 



HYPURAL 4 + 5 — 

HYPURAL 2 + 3 

Figure 42. Caudal skeleton of Peprilus triacanthus, drawing of a cleored-and-stained preparation from a 36-mm specinnen. 
All elements identified in Figure 1. 

cally, all members of the nominal genera 
are very similar. 

The species differ but little from one an- 
other. There is some question as to whether 
or not certain populations are to be re- 
garded as full species or only as subspecies. 
The marked similarity between species, and 
therefore the doubt as to the rank to be 
accorded certain forms, is probably due to 
the fact that active speciation is occurring 
now in the genus. More variational studies 
and increased knowledge of the natural his- 
tory are needed to resolve these questions. 

The species in Peprilus are: 

Pcprilus parti (Linnaeus, 1758)=S/ro- 
mateus paru Linnaeus. West Indies to Uru- 
guay, type locality Jamaica. D III 38-44. 
A II 3,5-41. P 20-22. Vertebrae 13 + 17. 
This species is distinguished from the very 
closely related P. alcpidotus ( Linnaeus, 
1766) by the lower counts and narrower 
pectoral fin (Hildebrand, MS). Synonyms 
are: '^'Seserinus xcinthuriis Quoy and 

Gaimard, 1824, from Brazil; Rhombus 
crenulotus Cuvier and Valenciennes, 1833, 
from Cayenne; and Rliombus orbicularis 
Guichenot, 1866a, also from Cayenne. 

Pcprilus alcpidotus ( Linnaeus, 1766 ) = 
Chaetodon alcpidotus Linnaeus. East coast 
of North America from Massachusetts to 
Florida and Gulf of Mexico, type locality 
Charleston, South Carolina. D III 43-49. 
A II 39-43. P 21. Vertebrae 13 + 17. This 
species is considered distinct from P. paru 
by Hildebrand (MS), although both are 
often lumped under this name. This is the 
"harvestfish" of the North American fish- 
eries literature. Stromateus longipinnis 
Mitchill, 1815, from New York Bay, is a 

Pcprilus triacanthus (Peck, 1804) = Stro- 
mateus triacanthus Peck. East coast of 
North America from Newfoundland to Flor- 
ida, type locality Piscataqua River, New 
Hampshire. D III 43-46. A II 37-43. P 
19-21. Vertebrae around 13 4- 19. This spe- 

Stromateoid Fishes • Haedrich 107 

Figure 43. Branchial region of Pepnius triacanthus, drawing of a cleored-and-sfained preparation from a 120-mm specimen. 
Elements identified in Figure 2. 

cies is very close to P. biiiti Fowler, from 
which it is distinguished by a slightly 
higher vertebral count (Collette, 1963). 
These two fonns have only recently di- 
verged. This fish is usually known as Poro- 
notus triacanthits, the "butterfish" of North 
American fisheries literature. Stromatcus 
ciyptoms Mitchill, 1815, from New York 
Bay, is a synonym. 

Peprilus simillimiis (Ayres, 1860) = Poro- 
notus similliiims Ayres. West coast of North 
America, British Columbia to Baja Califor- 
nia, type locality San Francisco. D III 45- 
47. A III 39-44. P 20-22. Vertebrae 13 + 
17. This species is a member of the P. tiia- 
canthiis-burti complex. 

Peprilus medius (Peters, 1869)=S7ro- 
matcus medius Peters. Known onlv from 

Mazatlan, Mexico, Pacific Coast. D III 42. 
A III 32. (From Fordice, 1884.) Possibly 
a synonym of P. simiUimus (Ayres, 1860). 

Peprilus polometa (Jordan and Bollman, 
1889) = "^Stromateus palometa Jordan and 
Bollman. Pacific coasts of Panama and 
Colombia. D III 44-48. A II 43-46. P 22- 
23. Gill-rakers 5-6 + 1 + 15-16. Vertebrae 
13 + 20-21. This deep-bodied fish with fal- 
cate median fins is a member of the Atlan- 
tic P. alepidotus-paru group. 

*P. smjderi Gilbert and Starks, 1904. 
Known only from Panama Bay. D III 41- 
47. A III 41-42. P 22-23. Gill-rakers 4 + 1 
+ 14. Vertebrae 13 + 23. This rarely seen 
species is distinguished from P. palometa 
(Jordan and Bollman, 1889) in having more 
vertebrae, a longer snout, and very short 

108 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 44. Pampas chinensii, a species lacking spines before the median fins, drawing of a 4-incfi specimen, from Day, 

lobes on the median fins. P. snijderi ap- 
proaches Stromateiis in the increased num- 
ber of vertebrae and in the very reduced 
spines preceding the rays in the median 
fins. It may be very Hke the stromateid an- 
cestral to Pcprilus and Stromatcus. Inves- 
tigations of its systematic position, geo- 
graphic distribution, and natural history 
should be very instructive. 

Peprihts biirti Fowler, 1944b. Gulf of 
Mexico, type locality Breton Island, Loui- 
siana. D III 43-45. A III 40-41. P 20-21. 

(From Fowler, 1944b.) Vertebrae around 
13+17 (Collette, 1963). This species is 
very close to P. triacanthus (Peck, 1804). 
Caldwell (1961) and Collette (1963) differ 
in their interpretation of its systematic 

Genus PAMPUS Bonaparte, 1837 

Figures 44, 45 

P(iin})iis Bonaparte, 1837:48. ( Subtrenus. Type 
species: Strotmitcus cfinilidiis Cuvier and 
Valenciennes, 1833:391, by subsequent desig- 
nation of Jordan, 1923:187. Malabar Coast. 

Stromateoid Fishes • Haedrich 


Figure 45. Pampus argenteus, a species with spines before the median fins, drawing of a 10.5-inch specimen, from Jordan 
and Metz, 1913. 

A .synonym of Stronmtcus argenteus Euphra- 
,sen, 1788:53.) 

Stromateoides Bleeker, 1851:368. (Type .species: 
Stromateus cinereus Bloch, 1793:90, by sub- 
sequent designation of Gill, 1862:126. A 
synonym of Stromateus argenteus Euphrasen, 

Chondroplites Gill, 1862:126. (Type species: 
Stromateus atous Cu\ ier and Valenciennes, 
1833:389, by original designation. After Rus- 
sell's "atoo-'koia" (1803: plate 21), Viza- 
gapatam. A synonym of Stromateus chinensis 
Euphrasen, 1788:54.) 

The combination of deep body, no pelvic 
fins, fixed maxillary, and gill membranes 
broadly united to the isthmus distinguishes 
Pampus from all other stromateoid genera. 
The name, a masculine noun, is from the 
vernacular of the 19th century East Indian 
Spanish and Portuguese colonials, who gen- 
erally used the term "pampus" (ultimately 
from "pampano") for any silvery, com- 
pressed fish. 

Description. Body very deep, maximum 
depth greater than 60 per cent of the stan- 
dard length, highly compressed; muscula- 
ture finn. Peduncle very short, compressed. 
A continuous dorsal fin; both median fins 
preceded by either none or five to ten flat, 
bladelike spines, pointed on both ends, pro- 
truding but slightly and resembling the 
ends of free interneurals. In forms with 
bladelike spines, dorsal fin originating 
slightly behind end of pectoral fin base, the 
first spine generally over or slightly before 
the pectoral insertion; in forms lacking 
spines, dorsal fin originating over the pec- 
toral fin base. Anal papilla well before mid- 
body, in a slit. Anal fin originating at or 
before middle of body and only slightly 
behind origin of the soft dorsal. Anterior- 
most rays of the median fins produced, the 
fins often falcate, rays which follow shorter; 
in forms with bladelike spines, rays of the 

110 Bulletin Museum of Cotnparative Zoology, Vol. 135, No. 2 

posterior two-thirds of the fin short and 
subequal, the anal fin lobe often extremely 
produced; in forms lacking spines, rays of 
the posterior two-thirds of the fin decreas- 
ing in length evenly to the last ray, the 
shortest. Pectoral fin long, winglike, the 
base of the fin inclined about 45°. No pel- 
vic fins. Pelvic bone not visible on midline 
and lacking a ventral spine. Tip of coracoid 
often projecting slightly underneath head 
at about level of margin of preopercle. Cau- 
dal fin stiff, deeply forked, in forms with 
bladelike spines the ventral lobe often ex- 
tremely produced. Scales very small, cy- 
cloid, thin, deciduous, extending onto bases 
of all fins. Simple tubed scales of the lat- 
eral line fairly high, following dorsal pro- 
file, and extending onto peduncle. Skin 
thin; main subdermal canal along intermus- 
cular septum and side branches usually 
quite apparent, pores to surface seem want- 
ing. Head around 25 per cent of the stan- 
dard length, \'ery deep and broad. Top of 
head naked, subdermal canals visible under 
naked skin but pores not visible, naked skin 
underlain with numerous parallel canals 
projecting backwards over the nape and 
along first part of lateral line. Eye small. 
Adipose tissue around eye developed and 
extending forward around the nostrils. Nos- 
trils large, the anterior round, the posterior 
a long slit, located near tip of the inflated 
snout at level of the top of the eye, nasal 
capsules greatly expanded. Mouth subter- 
minal, curved downward, small, maxillary 
scarcely reaching to below eye and angle 
of gape before eye. Premaxillary not pro- 
tractile. Maxillary immobile, covered with 
skin and united to cheek. Lacrimal bone 
very much reduced. Supramaxillary absent. 
Jaw teeth minute, uniserial, flattened, with 
a large rounded central cusp and two 
shorter auxiliary cusps, close set, covered 
laterally by a membrane. Vomer, palatines, 
and basibranchials toothless. Gill mem- 
branes broadly united to the isthmus. Gill 
opening a straight slit, covered with a flap 
of skin. Gill-rakers small, about one-quar- 
ter the length of the filaments, not toothed. 

widely spaced. Pseudobranch absent. Five 
branchiostegal rays, three on the ceratohyal, 
two on the epihyal. Scapula not prominent. 
Vertebrae variable in species with blade- 
like spines, 14 + 20 = 34 to 16 + 25 = 41; 
in species without bladelike spines, verte- 
brae 14 + 19 = 33. Dermal skeleton soft 
and spongy, but sclerotic bones well ossi- 
fied; skeleton in general fibrous. Stomach 
a simple sac; intestine very long. Pyloric 
caeca numerous, in a small dendritic mass. 

Color in life very silvery with a bluish 
cast on the back. Color in preservative 
brown or bluish with a silvery or whitish 
overlay. Median fins and caudal yellowish 
with dark borders. Plead a little darker than 
the body, with fine speckling. Gill mem- 
branes and inside of mouth dark. Perito- 
neum silvery with black speckles. 

Natural liistonj. Pampii.s is the most 
sought after of all the stromateoid fishes. 
Throughout the Orient, it commands a good 
price wherever it appears. In India, where 
it is known as "pomfret," the 1962 landings 
totaled 25.7 thousand metric tons, more 
than four per cent of the total marine catch 
( FAO 1964 ) . However, despite its com- 
mercial importance, virtually nothing is 
known of the life history of Pampus. 

The young occur in shallow water along 
the coasts, and may even ascend estuaries 
( Day, 1875 ) . The small mouths with cut- 
ting teeth and the long pharyngeal sacs sug- 
gest that soft-bodied coelenterates may 
figure largely in the diet. Most stomachs 
examined seemed to contain the shredded 
remains of these animals, but bits of fish 
were also found. Chopra (1960) found that 
a sudden appearance of numerous cteno- 
phores and medusae in the waters off Bom- 
bay was accompanied by a marked increase 
in the local catch of Pampus. 

Rehitionsliips. Pampus is the most ad- 
vanced stromateid genus. The advanced 
characters are the reduction in the number 
of branchiostegal rays to five, the lengthen- 
ing of the pharyngeal sac (Fig. 46), the 
restriction of the gill opening, the loss of 
th(^ pseudobranch, and the development, in 

Stromateoid Fishes • Hacdrich 111 

Figure 46. Branchial region of Pampus echinogosfer, drawing of a cleared-and-stained preparation from a 180-mm speci 
men, ABE 1743. Elements identified in Figure 2. 

some species, of flat, bladelike spines ahead 
of the median fins. The genus is derived 
from a fish very Hke Stromateus, but with 
fewer vertebrae. The most primitive spe- 
cies in Fampus has 14 + 19 vertebrae, but 
the more advanced may have as many as 
16 + 25. All members of the genus have 
the typical stromateid caudal skeleton (Fig. 

Species. Pampus is widely distributed in 
tropical waters over the continental shelves 
from the Iranian Gulf to Japan. There are 
reports of specimens from Hawaii ( Fowler, 
1938) and from the Adriatic (Soljan, 1948). 
No subsequent records have appeared from 
either place. The two localities are so far 
out of the established range of the genus 
that the records can only have been based 
on specimens brought from elsewhere. 

Gill (1884) divided the genus Stromat- 
eoides {= Pampus) into tsvo groups, which 
he apparently regarded as subgenera. The 
group Stromateoides had falcate fins and 

prominent dorsal and anal spines; the other 
group, CJwndwplitcs, had neither. This 
dichotomy does exist in Pampus, but more 
work is needed to decide whether or not 
the distinction merits subgeneric recogni- 

Numerous species have been described 
in the genus. The majority are probably 
synonyms. Published descriptions provide 
for the most part no clear-cut means of dis- 
tinguishing species. At the present state of 
knowledge, only three can be recognized: 

Pampus cliincnsis (Euphrasen, 1788) = 
Stromateus cJiinensis Euphrasen. India to 
China, type locality "Castellum Chinense 
Bocca Tigris." D 43-50 (total elements). 
A 39-42 (total elements). P 24-27. Ver- 
tebrae 14 + 19. This species lacks the pe- 
culiar flat spines before the median fins 
which are found in the other two species. 
The median fins are not falcate. The fin- 
rays gradually diminish in length posteri- 
orly (Fig. 44). P. chincnsis, the type for 

112 Bulletin Museum of Comparative Zoology, Vol 135, No. 2 



HYPURAL 4 + 5 — 

HYPURAL 2+3 — 

Figure 47. Caudal skeleton of Pampus argenteus, drawing of a cleared-and-stained preparation from a 48-mm specimen, 
ABE 1937. All elements identified in Figure 1. 

Gill's (1884) genus Chondroplites, is cer- 
tainly the most primitive species in Pompus. 
Synonyms are: Stromateus alhus Cuvier 
and Valenciennes, 1833, from Pondichery; 
Strot7uiteus atous Cuvier and Valenciennes, 
1833, from Vizagapatam; and Stromateoides 
atokoia Bleeker, 1852, from Malaysia. The 
trivial name is commonly written incorrectly 

Pampiis argenteus (Euphrasen, 1788) = 
Stromateus argenteus Euphrasen. Iranian 
Gulf to Japan, type locality "Castellum 
Chinense Bocca Tigris." D V-X 38^3. A 
V-VII 34-43. P 24-27. Vertebrae 14-16 + 
20-25. Falcate median fins, preceded by 
flat bicuspid spines, are characteristic of 
this species (Fig. 45). The species is ap- 
parently very wide ranging. Further study 
will no doubt show it to be composed of 
numerous subspecies or even species. This 
fish is the "pomfret" of Eastern fisheries 

literature, and is important commercially in 
India, China, and Japan. Probable syn- 
onyms are: Stromateus cinereus Bloch, 
1793; Stromateus candidus Cuvier and 
Valenciennes, 1833, from Pondichery; Stro- 
mateus securifer Cuvier and Valenciennes, 
1833, from Bombay; Stromateus griseus 
Cuvier and Valenciennes, 1833, from Pondi- 
chery; Stromateus punetatissimus Temminck 
and Schlegel, 1850, from Japan; and Pampus 
simoprosopus Fowler, 1934b, from Siam. 

Pampus echinogaster (Basilewsky, 1855) 
= Stromateus echinogaster Basilewsky. 
China, Korea, and Japan, tvpe locality 
China. D VIII-X 42-49. A V-VII 42-47. 
P 24-25. Vertebrae 14-15 + 24-26. This 
species has more median finrays than P. 
argenteus. Abe and Kosakai (1964) report 
that P. echinogaster has fewer, thicker 
pyloric caeca than P. argenteus, and 3-6 + 
12-15 gill-rakers as opposed to 2-3 + 8-10 

Stromateoid Fishes • Haedrich 113 

in P. orgcnteus. Pompus lighti Evermann 
and Shaw, 1927, from Nanking is a prob- 
able synonym. 


Gosline (1959) and Liem (1963) have 
recently stressed the need for an under- 
standing of functional moiphology in con- 
nection with phylogenetic studies. Natural 
selection acts on efficiencies and abilities, 
and evolutionary change results. If func- 
tion is understood, or at least taken cogni- 
zance of, an attempt can be made to evalu- 
ate observed differences in terms of effi- 
ciences and abilities. Within this framework, 
evolutionary trends can be discussed. Anat- 
omy alone is insufficient; the way of life 
must also be taken into account. 

The functional significance of many char- 
acters, however, is not known. It is hard 
to understand, for example, why selection 
should favor a fish with 15 principal 
branched rays in the caudal fin or with 25 
vertebrae, yet these numbers have appeared 
in many independent phyletic lines. Studies 
of characters of this sort are, nonetheless, 
of much importance. The argument that the 
overall trends observed are real is made 
stronger when these characters change in 
step with characters for which the function 
is at least partially understood. 

The trends in the evolution of the stro- 
mateoid fishes are discussed below, treat- 
ing separately, as much as possible, groups 
of characters which can be considered ana- 
tomical units, i.e., caudal skeleton, branchial 
region, etc. Drawing on what little is known 
of the way of life of stromateoids, func- 
tional significance can be attached to 
changes in the teeth and jaws, the pharvn- 
geal sacs, and the caudal skeleton. But the 
meaning of the changes in the size and 
shape of the body, the fin pattern, the bran- 
chial region, and the number of vertebrae 
cannot be easily interpreted. By and large, 
the discusion under each of these is neces- 
sarily brief and loaded with conjecture. 
Though each unit is treated separately, it 

must be remembered that they have evolved 
together. The organism responds as a whole 
to the environment, and the products of 
natural selection are all interdependent. 

Attention should be drawn to the distinc- 
tion between characters typical of a taxon 
and those typical of the grade of a taxon. 
In the first case, the characters are found 
in all members of the taxon. In essence, 
they are a part of the definition of that 
taxon. Grade refers to the "average" evo- 
lutionary status of the taxon vis-a-vis other 
taxa. The characters typical of a grade need 
not be found in all members of the taxon. 
In fact, because different parts of the ani- 
mal respond to the environment at differ- 
ing evolutionary rates, it is unlikely that 
these characters will be found in all. The 
use of grades pro\'ides a convenient way of 
discussing evolutionary trends without con- 
stantly itemizing the exceptions to the gen- 
eral picture. Thus, while 25 vertebrae char- 
acterize the centrolophid grade, not all 
members of the family Centrolophidae have 
25 vertebrae. Most do have 25 but some 
have 26 and others have near 60. 

Before proceeding to the discussion, let 
us briefly recall the more salient features 
of each of the five stromateoid families. 
The characters of the individual genera are 
presented in Table 2. 

The Centrolophidae are one to four feet 
in length with moderately deep to elongate 
bodies. All have pelvic fins. Their mouths 
and the teeth in their jaws are fairly large. 
There are no teeth on the palate. The pa- 
pillae in the round pharyngeal sacs have 
irregularly shaped bases. There are seven 
branchiostegal rays, a pseudobranch, and 
25, 26, 29, 30, or 50 to 60 xertebrae. The 
caudal skeleton has six hypural and usually 
three epural elements. There are six genera: 
Hyperoghjphe, Schedophilus, CentroJophus, 
Icichthys, SerioIcUa, and Psenopsis. 

The Nomeidae are six inches to over two 
feet in length with deep to elongate bodies. 
All have pelvic fins. Their mouths and the 
teeth in their jaws are small. The palatines 
and the vomer bear teeth. The papillae in 

114 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Table 2. Characters of the stromateoid genera. + denotes presence; — absence. 






Form of 




Epurals -\- 












4 feet 






3 + 6 



3 feet 





25, 26, 29, 30 

3 + 6 



4 feet 






3 + 6 



3 feet 





50 to 60 

2 + 6 


moderate to 

3 feet 






3 + 6 



1 foot 






3 + 6 



1 foot 

— in 





3 + 4 



1 foot 






2 + 4 


very deep 

18 inches 





33 41 

2 + 4 



3 feet 






3 + 4 



1 foot 






3 + 4 


to deep 

1 foot 


simple or 




3 + 4 





3 feet 


<1 foot 


simple or 



3 + 2 



2 feet 


simple and 




2 + 4 

the round pharyngeal sacs have stellate 
bases. There are six branchiostegal rays, a 
pseudoliranch, and 30 to 38, 41 or 42 ver- 
tebrae. The caudal skeleton has four hypu- 
ral and three epural elements. There are 
three genera: Cubiceps, Nomeus, and 

Most species of the Ariommidae are less 
than a foot long, although a few species 
attain more than twice that length. Their 
bodies are either deep or elongate. All have 
pelvic fins. Their mouths and the teeth in 
their jaws are very small. There are no 
teeth on the palate. The papillae in the 
elongate pharyngeal sacs have round bases. 
There are six branchiostegal rays, a small 
pseudobranch, and 30 to 33 vertebrae. The 
caudal skeleton has two hypural and three 
epural elements. There is one genus: 

The Tetragonuridae are less than two 

feet long with very elongate bodies. All 
have small pelvic fins. Their mouths are 
fairly large. In the upper jaw, their teeth 
are small and recurved; in the lower jaw, 
they are large and knifelike. The palatines 
and the vomer bear teeth. The papillae in 
the very elongate pharyngeal sacs are much 
reduced and have small round bases. There 
are five or six branchiostegal rays, a pseu- 
dobranch, and 43 to 58 vertebrae. The 
caudal skeleton has four hypural and two 
epural elements. There is one genus: Tetra- 


The Stromateidae are usually no more 
than a foot long with deep bodies. None 
have pelvic fins when adult. Their mouths 
and the teeth in their jaws are very small. 
There are no teeth on the palate. The pa- 
pillae in the round-to-elongate pharyngeal 
sacs have stellate bases. There are five or 
six branchiostegal rays, either a small pseu- 

Stromateoid Fishes • Hacdrich 115 

dobranch or none at all, and 30 to 48 verte- gonurid is 600 mm long (Fitch, 1951). These 

brae. The caudal skeleton has four hypural highly modified fishes are very slender, 

and usualh' two epural elements. There are with the maximum depth usually less than 

three genera: Stromateus, Peprilus, and 20 per cent of the standard length. The 

Pampus. stromateids rarely exceed 450 mm in length, 

There are two main lineages in the stro- and mature when less than 200 mm long, 

mateoids (p. 51). One is composed of the These fishes are very deep bodied, the max- 

Centrolophidae and their derivative, the imum depth ranging from 35 to over 70 per 

Stromateidae. The other, a looser assem- cent of the standard length, 
blage, is composed of the Nomeidae and The course of evolution in the form of 

their two derivatives, the Ariommidae and the body has been one of diminution in size 

the Tetragonuridae ( Fig. 7 ) . The Centro- and of increase in depth. These t\\'o evolu- 

lophidae and the Nomeidae are the basal tionary tendencies are also displayed in 

stocks. Of these two, the Centrolophidae other teleostean groups (Myers, 1958; Liem, 

are in most respects the most primitive. 1963). 

Familial and generic relationships, to be Fins (Fig. 48). Only one major change 

touched upon only briefly here, have been has occurred in the fin pattern of stromat- 

discussed in the individual accounts of eoids — the loss of the pelvic fins at the 

family and genus. stromateid grade. The presence of pelvics 

Figure 48 summarizes some of the major in young Stromotctis fiafolo and their sub- 
evolutionary trends in the stromateoid sequent loss in the adult are important clues 
fishes. Each group is treated more or less in understanding the phylogeny of stroma- 
as a grade in the diagram. The characters teids. There is a difference between the 
sho\\'n are relative size and shape of the basic fin patterns of the t\\'o stromateoid 
bod>-, fin pattern, presence or absence of lineages. Members of the nomeid line have 
palatal dentition, number of branchiostegal two dorsal fins; members of the centro- 
rays, vertebrae, epural plus hypural ele- lophid line usually have but one. 
ments in the tail, and the shape of the pa- The thoracic pelvic fins of perciform 
pillae in the pharyngeal sacs. These are dis- fishes are used in braking and turning 
cussed in detail in the following accounts. (Harris, 1938). In deep-bodied fishes, how- 
The width of the arrow leading to each ever, the effecti\eness of the fins for these 
grade is proportional to the number of purposes is probably greatly decreased due 
genera in that family. to the change in the hydrodynamic profile 

Body (Fig. 48). The largest stromateoids of the fish. In this situation, selective pres- 

are members of primitive centrolophid sure may favor loss of the fins. This has 

genera. McCulloch ( 1914) reports a 1,072- apparently been the case in stromateids, 

mm Hyperoghjphe porosa weighing 41 and is also observed in Parasfromatcus, 

pounds from the Great Australian Bight, Monodactyhis, Psetfus, and a number of 

and I have seen an 1,195-mm Centrolophus other unrelated deep-bodied teleosts. 
niger taken south of New England on a In the great majority of stromateoids, the 

long-line. The maximum depth in most anterior rays of the median fins are no more 

centrolophids is within 25 to 30 per cent than two or three times the length of the 

of the standard length and never exceeds posterior rays. In the stromateids, however, 

50 per cent. In the nomeids and in the the median fins have become falcate and 

ariommids, there are a few species whose the anterior rays are very much produced, 

maximum length approaches a meter but In Stromateus this tendency is but little 

most are smaller. The maximum depth in pronounced. The deeper-bodied species in 

these families is from about 25 to 40 per Peprilus have very falcate fins. The anterior 

cent of the standard length. A large tetra- finrays of the anal fin in these species are 

116 BiiUrtin Museum of Comparative Zoology, Vol. 135, No. 2 


Figure 48. Evolutionary trends in the Stromoteoldel, showing relative size and shape, fin pattern, and (within the outline 
from left to right) presence or absence of palatal dentition, and numbers of branchlostegals, vertebrae, and epurals + 
hypurals. Inset shows a papilla. Width of arrows proportional to number of genera in the family. See text. 

seven or eight times longer than the pos- fin may be ten to 20 times longer than the 

terior rays. In Pampu.s both the anterior posterior finrays. Lacking observations on 

rays of the anal fin and of the lower caudal living Pampus, it is difficult to know what 

lobe are produced, and those in the anal advantage these elongate fins confer. 

Stromateoid Fishes • Hacdrich 117 

Teeth and jaws (Fig. 48). In most cen- flattened, cusped teeth are more suited to 

trolophids the angle of the gape may he slicing the tissues of coelenterates. With 

helow the eye hut in all other stromateoids such animals, there is little need for catch- 

the angle of the gape is hefore the eye. The ing and holding. Nonetheless, the tetra- 

ariommids and stromateids have the small- gonurids, existing almost entirely on salps 

est mouths of all. In these two families and coelenterates, have conical, recurved 

even the end of the maxillary is before the teeth in the upper jaw and on the palate 

eye. Centrolophids, with the exception of which must hold the prey firmly while the 

Psenopsis, ha\e a small supramaxillary bone, long, knifelike teeth of the lower jaw slice 

but this is gone in all other stromateoids. off mouthfuls. 

The presence of teeth on the palate is In the course of evolution, the jaws of 

usually considered primitive (Liem, 1963). stromateoids have become shorter and the 

The palatine and vomerine teeth in the supramaxillary bone is lost. The conical 

nomeids indicate that this group branched teeth have become flattened, cusped, 

off early from the ancestral stock. Both the smaller, and more closely set. In the no- 

centrolophids, in most respects the most meid lineage, the palatine dentition is lost, 

primitive of stromateoids, and their deriva- Changes in the dentition by and large re- 

tive, the stromateids, lack these teeth. Pala- fleet increasing specialization in the food 

tal dentition is well developed in the tetra- habits. 

gonurids, which are derived from fish an- Pharyngeal sacs (Figs. 49, .50). The sacs 

cestral to the nomeids. The ariommids, of centrolophids are higher than they are 

which are probably derived directly from a long, and the papillae are arranged in ten 

nomeid, have lost these teeth. or more elongate patches (Fig. 49A). The 

In the majority of primitive centrolophids bases of the papillae ( Fig. 50A ) are irregu- 
and in many nomeids, the jaw teeth are lar in shape, with the teeth seated all over 
relatively large, strong, spaced, and often the inner face; the base is often humped up 
slightly recurved. The advanced centro- to fit over a ridge of muscle in the sac. In 
lophids, stromateids, and ariommids have the nomeids, the sac is not so high in re- 
much smaller, close-set, straight teeth. The ^pect to its length, and the papillae are in 
teeth of all stromateids and of some ariom- ^bout five longitudinal patches (Fig. 49B). 
mids are laterally flattened and bear minute ^he papillae are verv different from those 
cusps. Tetragonurids and some species in ^^ ^j^^ centrolophids. 'The bases ( Fig. SOB ) 
the nomeid genus Psenes have two sorts of ^^^ ^^^^^^^^ ^^^^ ^j^^ ^^^^j^ ^^^ concentrated 
teeth m the jaws. Tliose m the upper ,aw ^^^^^ ^^^ ^^^^ ^^ ^ ^^^j^ ^^^ ^^^ .^^ ^^^^^^_ 

are conical, spaced, and recurved; those m . . , , t-- Ar,r^ \ • i. i i. i -^ • 

^, , . ^ , , .r ,., , teids (Fig. 49C) is at least as long as it is 

the lower jaw are long, knitelike, very close r,- i, • p /t^- 4f-\ i. • . .j j 

set, and often bear verv minute cusps. ,° ' „ ' v &• I h  

The structure of the teeth and jaws is ^he papillae are m only two ill-defmed 

certainlv a function of the diet. In stromal- P^tdies in the top and bottom halves of the 

eoids, which possess a masticatory organ sac. As in the nomeids, the bases (Fig. 50C) 

in the pharyngeal sacs, the jaw teeth are are stellate, but they are in general larger, 

primarily for catching and holding prey. and the teeth are seated all along the long 

The diet of most centrolophids and no- central stalk instead of only near the end. 

meids is fairly diverse and often includes The sacs in both the ariommids ( Fig. 49D ) 

rather large animals. The strong, conical, and the tetragonurids are longer than high, 

slightly recurved teeth are ably suited to markedly so in the latter (Fig. 36). The 

the catching and holding of fairly vigorous large papillae of ariommids (Fig. SOD) have 

prey. The stromateids feed rather exten- round bases, and the small teeth are seated 

sively on jellyfishes. Their smaller jaws and all along the central stalk. The papillae are 

118 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 




5?aki .. ;. I ^ ,■•,'- 73^"t »•! 'J'. ^ 

Figure 49. Comparison of bronchial regions in four stromateoid families. A. Centrolophidoe, Hyperoglyphe, from Figure 9. 
B. Nomeidoe, Nomeus, from Figure 25. C. Stromateidae, Peprilus, from Figure 43. D. Ariommidae, Ariommo, from Figure 
31 . See text. 

in a single patch, in the upper half of the 
sac only. In tetragonurids, the small papil- 
lae are widely separated and are not in 
bands. They are rounded, and there are a 
few weak teeth on the end of a short stalk. 
The nomeids and the stromateids do not 
share a direct common ancestor. The no- 
meids are derived from a pre-centrolophid 
form and the stromateids are derived from 
an advanced centrolophid. Nonetheless, 

there is a great similarity in the stellate 
papillae found in both families (Fig. SOB, 
C), but this similarity is due to parallelism. 
The centrolophid fishes are unspecialized 
in their diets. They feed on other fishes, 
on squids, on crustaceans, on jellyfishes 
and, sometimes but certainly not customar- 
ily, on garbage. The large sacs are capable 
of admitting fairly large objects. The crude 
papillae do shred the prey to some extent. 

Stbomateoid Fishes • Haedrich 




A, A,, B, C 


Figure 50. Comparison of papillae in the pharyngeal sacs of four stromateoid families. A. Centrolophidae, Hyperoglyphe, 
from preparation shown in Figure 9, large papilla. A). Same, small papilla. B. Nomeidae, Nomeus, from preparation 
shown in Figure 25. C. Stromateidae, Pepri/us, from preparation shown in Figure 43. D. Ariommidae, Ariomma, from prep- 
aration shown in Figure 31. See text. 

but never so much as to render stomach 
contents completely unrecognizable. 

Little is known of the feeding habits of 
nomeids. Fish and jelhfish remains have 
been found in their stomachs. The fairly 
small sacs and the papillae with their stel- 
late bases firmly seated in the muscular 
wall of the sac combine to make a good 
shredding organ, and stomach contents are 
often difficult to identify. 

Stromateids may feed very largely on 

small crustaceans and medusae. The rela- 
tively smaller and more elongate sacs, the 
papillae with greatly extended bases, and 
the teeth ranged all along the central stalk 
of the papillae make a very efficient shred- 
ding organ, ably suited to rendering the 
rubbery tissues of medusae. The shredded 
stomach contents of stromateids are almost 
impossible to identify. 

Too little is known of the natural history 
of ariommids to be able to understand the 

120 Bulletin Musctim of Comparative Zoology, Vol. 135, No. 2 

structure of the peculiar pharyngeal sacs 
found m this group. In almost every speci- 
men examined, the sacs were filled with 
mud and silt. Is it possible they perform 
some sort of filtering function? 

Tetragonurids may live largely on jelly- 
fishes. The sacs of these fishes are very 
elongate, as might be expected, but the 
papillae are very reduced and are probably 
not very efficient shredders. The upper 
pharyngeal bones, houever, are studded 
with teeth and extend verv far backward 
into the sacs (Fig. 46). The pharyngeal 
bones are capable of considerable back-and- 
forth motion (Grey, 1955) and, in tetra- 
gonurids, may perform the shredding action 
for which the papillae do not seem suited. 

The main changes that have occurred in 
the pharyngeal sacs of stromateoids have 
been elongation of the sac, reduction in the 
number of bands of papillae, and increase 
in complexity of the papillae. These changes 
are correlated with a change from more or 
less omnivorous feeding habits to increasing 
utilization of jelly fishes for food. 

Branchial rciiion (Figs. 4<S, 49). Seven, 
blunt-ended branchiostegals and a large 
ceratohyal fenestra are found at the centro- 
lophid grade (Fig. 49A). The advanced 
centrolophids ScriolcUa and Pscnop.sis have 
pointed branchiostegals, and the first one is 
reduced in size (Figs. 20, 23). In nomeids 
(Fig. 49B) and ariommids (Fig. 49D) there 
are six tapering branchiostegals, and the 
ceratohyal fenestra is much smaller, or, in 
some species, closed. At the stromateid 
grade ( Fig. 49C ) there are six tapered 
branchiostegals and the ceratohyal fenestra 
is closed. The stromateid genus Potnpus, 
perhaps the most advanced of stromateoids, 
has but five branchiostegal rays. Within the 
tetragonurids, the number of branchioste- 
gals is either six or five. 

A pseudobranch is present in all stromat- 
eoids with the exception of Pampas. Its 
loss may be correlated with the unification 
of the gill-covers to the isthmus. In most 
stromateoids, the pseudobranch is very well 
developed and the gill-covers are cleft well 

fonvard. In the stromateid genera Stronia- 
teus and Pcprilus the pseudobranch is small 
and the gill-covers are united across, but 
not to, the isthmus. Finally, in Pampiis the 
pseudobranch is gone and the gill-covers 
have become broadly united to the isthmus. 

In the hyal series of stromateoids, two 
changes have occurred. The branchiostegal 
rays ha\'e become more slender and one 
ray is lost, and the ceratohyal fenestra be- 
comes closed. The pseudobranch, well de- 
veloped in most stromateoids, is lost in 

Axial skeleton (Fig. 48). Most centro- 
lophids have 10 -I- 15 vertebrae, the well- 
know n basic perciform number. The excep- 
tions are in some Schedophilu.s with 10 4- 
16, 12 + 17, or 10 + 20, and Icichthys with 
a total of 50 to 60. In the nomeids both 
numbers have increased; there are 13 to 15 
precaudal, and 17 to 23, 26, or 27 caudal ver- 
tebrae. The stromateids have 12 to IS pre- 
caudal, and 19 to 27 caudal vertebrae, and 
within any one species the number may be 
quite variable. Ariommids, derived from a 
nomeid stock, usually have 12 or 13 + 17 
or IS vertebrae. The tetragonurids, also 
derived from the nomeids, have continued 
the increase, to a total of 43 to 5S vertebrae. 

It is difficult to understand the selective 
pressures responsible for an increase in ver- 
tebral niunber. In Nomeu.s, however, the 
situation is reasonably clear. This genus has 
41 vertebrae and is elongate in fomi. The 
high number of vertebrae allows the fish 
to move in a very sinuous manner, and to 
turn in a very small radius. The pelvic fins 
are also much enlarged and aid in the turn- 
ing. This ability enables the fish to avoid 
more easily the stinging tentacles of the 
PJujsalia under which it lives. 

In general, as the number of vertebrae 
has increased, the relative length of each 
individual vertebra has decreased. Fishes 
with an increased number of vertebrae have 
more neural and haemal spines than fishes 
with fewer vertebrae, and these are closer 
together. Hence there is a stronger frame 

Stromateoid Fishes • Hacdrich 


Figure 51. Comparison of caudal skeletons of four stromateoid families. A. Centrolopfiidae, Hyperoglyphe, from Figure 10. 
B. Nomeidae, Nomeus, from Figure 24. C. Stromoteidae, Peprilus, from Figure 42. D. Ariommidae, Ariomma, from Figure 
33. See text. 

for attachment of the muscles. This prob- 
ably has permitted, or even encouraged, 
the evolution of the deep, firm body char- 
acteristic of the stromateid grade. 

With the increase in vertebral number, 
the number of median finravs increases as 
well. This, of course, might be expected, 
but need not necessarilv follo\\ . The ariom- 
mids and the tetragonurids, both with in- 
creased numbers of vertebrae, have very 
decreased numbers of median finrays. In 
Psenes and in the stromateids, the increased 
number of anal finrays has resulted in a 
forward swing of the first interhaemal so 
that it forms an abrupt angle with the 
haemal spine of the first precaudal verte- 

bra. This tends to support and protect the 
belly of the fish. In the stromateids, the 
enlargement and extension of the pelvic 
bones, which bear no fins, and of the post- 
cleithrum almost complete this trend to- 
wards support and, possibly, protection. 

There has been a general tendency to- 
^^'ards increase in the number of \ertebrae 
in the evolution of the stromateoids. The 
number of caudal vertebrae has tended to 
increase the most, but the number of pre- 
caudal vertebrae has been affected as well. 
Secondary increase in the number of ver- 
tebrae from a basic number near 10 + 15 is 
of common occurrence in teleosts ( Gregory, 
1951; Liem, 1963). In many stromateoids. 

122 Bulletin Museum of Comparative Zoology. Vol. 135, No. 2 

there has been a concomitant increase in 
the number of median finrays. 

Caudal skeleton (Figs. 48, 51). The gen- 
erahzed percifonn type of caudal skeleton 
with six hypurals and three epurals is t\ pi- 
cal of the centrolophid grade (Fig. 51A). 
With the fusion of hypurals 2 + 3 and 3 + 
4, the number is reduced to fovu" in the no- 
meids (Fig. 51B), stromateids (Fig. 51C), 
and tetragonurids. Three epurals are pres- 
ent in nomeids, but one of these is lost at 
the stromateid grade (Fig. 51C). In the 
ariommids (Fig. 51D), the fusion of hy- 
purals 1 + 2 + 3 and 4 + 5 + 6 fonns two 
solid blocks. In both the ariommids and 
some stromateids there is a tendency to- 
wards further fusion of hypural elements 
with the urostylar vertebrae. The ariom- 
mids have three epurals but the second one 
is very reduced in size, and is probably on 
its way to becoming lost. The tetragonurids 
have two epurals. Fusion of the uroneurals, 
both with each other and with the urostylar 
vertebrae, has occurred in several stromat- 
eoid genera. 

Most of the centrolophid fishes spend at 
least the first part of their lives hovering 
quietly under floating objects. These fishes 
are able to hang almost motionless with a 
slight fanning of the pectorals and strong 
rotary motion of the caudal fin. This rotary 
motion is possible because of the nimierous 
elements in the caudal skeleton. In the 
advanced centrolophids — fast-swimming, 
schooling fishes such as Seriolella violaceo — 
partial fusions in the hypural series result 
in a more rigid tail. 

Many of the nomeids are hovering fishes. 
At this grade, fusions in the hypural series 
tend to make the tail fairly stiff. This may 
be counteracted by the long, well-developed 
autogenous haemal spines, which may be 
moved laterally to produce a rotary motion 
in the fin. The long pectoral fins of no- 
meids, too, may aid their hovering. Obser- 
vations on living fishes are, however, lack- 
ing, and are sorely needed. 

The consolidated tail of stromateids al- 
lows for little rotarv motion. These school- 

ing fishes probably do not hover as much 
as nomeids or centrolophids but may swim 
fairly constantly. Specimens of Feprilus 
triacanthus observed in the Woods Hole 
Aquarium never remained still, but moved 
slowly forward, bouncing up and down with 
beats of their long, broad pectoral fins. 

There are no observations of living ariom- 
mids or tetragonurids. From the structure 
of their caudal skeleton and fin, it can be 
assumed that the former at least are very 
strong, fast swimmers. Living near the bot- 
tom, they may not need to hover, but may 
cruise over the sea floor buoyed up by their 
well-developed air bladder. 

The evolution of the caudal skeleton in 
stromateoids is marked by a reduction in 
the number of elements. In the hypural 
series, this reduction is accomplished by a 
series of fusions; in the epural series, an 
element is lost. The tendency toward con- 
solidation and reduction of elements, ulti- 
mately resulting in a fused hypural plate, is 
a general phenomenon found in numerous 
percifonn lineages (Gosline, 1961a). The 
changes in the stromateoid tail coincide ap- 
proximately with a change from hovering 
to swimming fairly constantly in schools. 


Distributional data for stromateoid fishes 
are at best scanty. Nonetheless, all avail- 
able data tend to support the conclusions 
based on anatomical data, that is, that the 
centrolophids arose first, followed by the 
nomeids, tetragonurids, and, most recently, 
the stromateids and ariommids. Because of 
the scantiness of the data, the map figures 
accompanying this section must be con- 
sidered approximate only. In general, the 
distributions have been extrapolated from a 
few records. Although I am fairly sure of 
the general picture presented, fine details 
of the distribution of stromateoids are lack- 

CentroJophidae. The major features of 
the centrolophid distribution are disconti- 
nuity, bipolarity, endemism, and sympatry 
of genera. The first three are found in the 

Stromateoid Fishes • Haedrich 


Figure 52. Distribution of ttie soft-spined Centrolophidae. 

more primitive members of the family, the 
soft-spined centrolophids (Fig. 52). Schedo- 
philus is found mainly in the Atlantic 
Ocean. The presence of two isolated popu- 
lations, in the China seas and in the en- 
virons of the Tasman Sea, indicates that the 
former range of the genus was once much 
wider. The distribution of the two Pacific 
area populations of SchedopJiihis and the 
distribution of Centrolophus are bipolar, 
again indicative of a shrinking range. 
IcichtJujs, fonnerly considered an endemic 
element of the North Pacific fauna, has re- 
cently been found off New Zealand ( Haed- 
rich, in press), and is thus bipolar. Icichthys 
and Centrolophus probably share a com- 
mon ancestor; the characteristics of each 
genus may well have developed in the iso- 
lation provided by an ancestral relict dis- 
tribution. In the hard-spined centrolophids 
(Fig. 53), Hyperoglyphe, like SchcdopJiihts, 
is bipolar in the Pacific but widespread in 
the Atlantic. The most primitive species in 
this genus, H. antarctica, is found only 
south of 30° S. Seriolella, a relatively ad- 
vanced genus, is widespread in the higher 
latitudes of the Southern Hemisphere, 
where it is no doubt endemic. The most 
recently evolved centrolophid genus, Pse- 

nopsis, may be spreading out from the wa- 
ters of the East Indian region. By and large, 
centrolophid species are oceanic or found 
near the edge of the continental shelf. Some 
species of the soft-spined centrolophids may 
even be meso- or bathypelagic. The ad- 
vanced genera Seriolella and Psenopsis, 
however, commonly occur in shallow wa- 
ter, and some species may even enter estu- 
aries. Some overlap with at least one other 
genus occurs within the ranges of all cen- 
trolophid genera. Four of the six genera 
occur in Australia and New Zealand. 

Nomeidae. In the distribution of the no- 
meids (Fig. 54) there are no relicts, no bi- 
polar species, and no regional endemism. 
For the most part, the three genera seem 
broadly sympatric, but records are too few 
to discuss the limits of each genus with 
precision. In the North Atlantic, however, 
Nomeus is found in the western parts, but 
has never been reported from Madeira, 
where its companion Physalia is common. 
Cuhiceps, though it occurs in the western 
Atlantic, is much more common in the east- 
em portions and the Mediterranean. Most 
nomeid species are oceanic; a few species 
in Psenes seem to be mesopelagic. In gen- 
eral confined to more tropical waters, a 

124 Bulletin Museum of Comparative Zoology. Vol. 135, No. 2 

Figure 53. Distribution of the hard-spined Centrolophidae. 

number of species, such as Nomeus ^ronovii 
and Psenes cyonoplinjs, are found in all 
oceans. All genera are found in Australia 
and New Zealand. 

Tctruiiomuidae. The distribution of the 
tetragonurids is very poorly known. In gen- 
eral it seems to approximate the distribu- 
tion of the nomeids ( Fig. 54 ) . 

Ariommidae. The family Ariommidae, a 
nomeid derivative, is found mainly in tropi- 
cal waters (Fig. 55). One deep-bodied spe- 
cies occurs off South Africa. All members 
of the single genus Arionima seem to be 
engybenthic in deep water over continental 
shelves or near islands. The deep-bodied 
and elongate forms of Ario77rma occiu- to- 
gether in the New World, but tend to be 
allopatric elsewhere. Apparently there are 
no representatives on the west coast of Cen- 
tral America, although the genus is wide- 
spread throughout the Gulf of Mexico and 
the Caribbean, and elongate species occur 
in Hawaii. The latter are undoubtedly de- 
rived from Japanese forms. The most ad- 
vanced species in the genus, A. indica, is a 
deep bodied silvery species with cusped 
teeth, found from the Gulf of Iran through- 
out the East Indian region to the East China 

Sea. No ariommids occur in Australia or 
New Zealand. 

Sticmmteidae. The stromateid distribu- 
tion is characterized by continuity, wide- 
spread species, restriction to continental 
shelves, a trans-Isthmian genus in the New 
World, and allopatry of genera. The dis- 
tributions of each of the three genera ( Fig. 
56) are more or less continuous. In S^ro- 
matciis, one species is found from the Medi- 
terranean to South Africa. Stromatctis is 
the only genus that has managed to cross 
an ocean. This has been accomplished 
across the shortest possible gap, from Africa 
to South America, and in the direction of 
the prevailing winds and currents. The ad- 
vanced StromateiLS of the east and west 
coasts of southern South America are very 
little differentiated from one another, and 
may be speciating at the present time. The 
genus Feprilus, apparently derived from 
Stromateu.s through a species such as the 
west coast P. snijderi. has spread on both 
coasts of North America and southward 
along the east coast of South America to 
Uruguay, where it occurs sympatrically 
with Stwmatetis. This is the only place 
where two stromateid genera are found to- 

Stromateoid Fishes • Haedrich 125 

Figure 54. Distribution of the Nomeidae: Cubiceps, Nomeus, and Psenes. 

gether. The most advanced stromateid 
genus, Pampiis, occurs from the Gulf of 
Iran to Japan. Both an advanced species, 
P. orgenteits, and a more primitive species, 
P. chincnsis, occur through most of the East 
Indian region. No stromateids have crossed 
Wallace's line into Australia or New Zea- 

Discussion. The two most recently 
evolved families are the ariommids and the 
stromateids. In the ariommids, a single 
genus is widespread. Containing two in- 
cipient genera, one elongate and one deep- 
bodied, AriotJima has apparently had insuf- 
ficient time for characteristics worthy of 
generic division to develop. Three genera 
have evolved in the stromateids, but in 
general each genus is restricted to a sepa- 
rate continental area. Peprihis has spread 
in a classical circular pattern, from Pacific 
South America across the Isthmus of Pan- 
ama and south to Uruguay, to re-encounter 
the ancestral Strornatcus stock. This small 
region in the western South Atlantic is the 
only area where stromateid genera occur 
sympatrically. Speciation is currently active 
in both families. 

The ariommids and the stromateids, in 
contrast to the other stromateoid families. 

are restricted to near land. The ariommids 
live in deep water over the shelves and in 
the vicinity of Hawaii. The stromateids may 
prefer quite shallow water, and occur in 
large schools in wide embayments. Be- 
cause of this relationship with the land, it 
is possible to examine and possibly date the 
emergence of the two families in the light 
of past tectonic activity. 

The present stromateid distribution 
throughout Asia (but not the Red Sea), 
the Mediterranean, West Africa, and the 
New World is strongly suggestive of an 
ancestral Tethyan distribution. The stro- 
mateid ancestor could have been a member 
of the warm water shelf fauna which ex- 
tended uninterrupted across this region in 
Tertiary times. In the Upper Eocene or 
Oligocene, the emergence of land in the 
Near East divided this fauna in two. The 
ancestral stromateid isolated in the East 
gave rise to Pampus., the form in the West 
was the central Stwmafetis stock. In the 
Pacific Panamanian region, separated from 
the southern stock of Strornatcus by long 
coastlines unsuitable for stromateids, Pep- 
rihis evolved. Found today in both oceans, 
this genus must have been established be- 

126 Bulletin Museum of Comparative Zoology, Vol. 135, No. 2 

Figure 55. Distribution of the Ariommidoe. 

fore the emergence of the Isthmus of Pan- 
ama in the lower PHocene. 

The ariommids are not so tightly bound 
to the coasts as the stromateids. Oceanic 
dispersal may be facilitated by pelagic 
juveniles, a few of which have been taken 
at Bermuda and in the tropical Central At- 
lantic. Widespread in the Gulf of Mexico 
and the Caribbean, no ariommids occur on 
the west coast of the New World. This 
suggests at least a late Pliocene dispersal. 
This fact, plus the remarkable homogeneity 
of the group and the structure of the pha- 
ryngeal sacs and the fused hypural fan, are 
strong evidence for considering the ariom- 
mids the most recently evolved stromateoid 

The remaining stromateoid families, the 
centrolophids, nomeids, and tetragonurids, 
are by and large all oceanic. Changes in 
the configuration of the land would not 
have affected these fishes as they did the 
ariommids and stromateids. Since the major 
ocean basins have probably been a stable 
feature since well before the Cretaceous, 
the period of the great flowering of the 
teleosts, it is unlikely that tectonic activity 
has been an important isolating mechanism 
in the evolution of these groups. 

The centrolophid distribution bears all 
the earmarks of an older group. Disjunct 
distributions, including bipolarity, are char- 
acteristic of an old group which has passed 
its peak. Another indication of the age of 
the centrolophids is their diversity. There 
are six genera in the family, and the spe- 
cies inhabit a wide range of en\'ironments. 
In Australia and New Zealand, where no 
stromateids occur, the advanced centro- 
lophid genus Seriolella lives in shallow wa- 
ters near the coast, the typical stromateid 
habitat. Numerous ebbs and flows have 
occurred in the distribution of the centro- 
lophids, for numerous genera are found to- 
gether. Lacking fossils, it is impossible to 
date the emergence of the centrolophids. 
Nonetheless, they certainly antedate the 
stromateids which had their beginnings in 
the mid-Tertiary. The centrolophids, then, 
probably arose in the early Tertiary, or per- 
haps even in the late Cretaceous. 

The nomeids probably arose concurrently 
with, or perhaps a little after, the centro- 
lophids. The genera occur together through- 
out the range of the family. Little specific 
differentiation seems to have developed, al- 
though the apparent commonness of cir- 
cumtropical species in this group may only 

Strom ATEOiD Fishes • Haedrich 127 

Figure 56. Distribution of the Stromateidae. 

reflect the premium placed on a particular 
phenotype in the rigorous oceanic environ- 

The great divergence from the nomeid 
condition of a derived family, the tetra- 
gonurids, suggests that they branched off 
at an early stage. The tetragonurids have 
become very specialized, and may be an 
evolutionary dead-end. There are appar- 
ently only three species in the single genus. 

Two general features of the stromateoid 
distributions are of interest. The first is the 
tendency for the more primitive taxa to be 
found in higher latitudes. Included in this 
group are: CentroJophus\ Icichthys, Hijpero- 
ghjphe antarctica, Stromateus in the New 
World, and, perhaps, the giant nomeid 
Cubiceps capensis. The Ariommidae and 
Stromateidae, both advanced, have not 
reached the Australian region. The second 
feature is that the most advanced or most 
recently evolved taxa have their centers of 
distribution in the East Indian region. Ex- 
amples are the centrolophid genus Psenop- 
sis, the stromateid genus Tampus, and the 
ariommid Ariomma indica. 


In the course of this studv I have been 

supported by an Emerson Fellowship from 
Harvard University and a Summer Fellow- 
ship and Predoctoral Fellowship from the 
Woods Hole Oceanographic Institution. 
The National Science Foundation spon- 
sored my participation on Cruise 6 of the 
ANTON BRUUN, as a part of the U. S. 
Program in Biology, International Indian 
Ocean Expedition, and also permitted me 
to visit Japan under the joint U. S. -Japan 
Cooperative Program in Science (OF- 147 
to Harvard University). Funds from the 
Harvard Committee on Evolutionary Biol- 
ogy covered my visits to natural history 
museums in Washington, London, Paris, 
and Copenhagen. Incidental support has 
been derived from National Science Foun- 
dation Grant GB-543 to the Woods Hole 
Oceanographic Institution. 

This study would not have been possible 
without the kind cooperation of numerous 
curators who have freel\- sent me specimens 
from far-flung collections. For these ser- 
vices my thanks go to Frank J. Talbot of 
the Australian Museum, Sydney; William 
J. Richards of the Bureau of Commercial 
Fisheries Biological Laboratory, Washing- 
ton, D. C; Alfred W. Ebeling of the Bing- 
ham Oceanographic Laboratory, Yale Uni- 

128 Bulletin Muscuttt of Comparative Zoology, Vol. 135, No. 2 

versitv; Frank Williams of the Guinean 
Trawling Survey, Lagos; M. J. Penrith of 
the South African Museum, Capetown; 
George S. Myers of the Natural History 
Museum, Stanford University; Richard H. 
Rosenblatt and Joseph F. Copp of the 
Scripps Institution of Oceanography, La 
Jolla; Werner Ladiges, Zoologisches Mu- 
seum, Hamburg; and J0rgen G. Nielsen of 
the Zoological Museum, Copenhagen. On 
my visits to various institutions, which also 
supplied specimens for the study, I received 
the very best of treatment from James Tyler 
of the Academy of Natural Sciences of 
Philadelphia; C. C. Lindsey and Norman 
J. Wilimovsky of the University of British 
Columbia, Vancouver; P. H. Greenwood 
and A. C. Wheeler of the British Museum 
(Natural History), London; Loren P. Woods, 
Marion Grey, and Pearl Sonoda of the Chi- 
cago Museum of Natural History; M. 
Bauchot and M. Blanc of the Museum Na- 
tional d'Histoire Naturelle, Paris; and 
Daniel M. Cohen, Bruce B. Collette, Robert 
H. Gibbs, Jr., Nathaniel Gramblin, and 
Leonard P. Schultz of the United States 
National Museum, Washington, D. C. 
Frank J. Mather, HI of the Woods Hole 
Oceanographic Institution allowed me free 
access to collections made in connection 
with his pelagic fish studies and also ex- 
amined the types of SerivIeUa in Paris. 
Especial thanks go to Erik Bertelsen of the 
Danish Carlsberg Foundation who provided 
the nucleus of specimens which initiated 
this study and who has encouraged it 
throughout, and to Tokiharu Abe who pro- 
vided numerous Japanese specimens, who 
has increased my understanding through 
discussions, and who was to me a most 
gracious host during my stay in his country. 
Especial thanks also go to the staff of the 
Museum of Comparative Zoology — particu- 
larly Myvanwy M. Dick, Josie DeFalla, 
Elaine Kelley, and Charles Karnella — and 
to the staff of the Marine Biological Labo- 
ratory Library — particularly Jane Fes- 
senden — for help cheerfully given. 

I gratefully acknowledge the particular 

services rendered by a number of people, 
all friends and colleagues at the Woods 
Hole Oceanographic Institution. Louise 
Russell made the drawings of the gill arches 
and pharyngeal sacs. Clifford Hinton, Jr. 
made the final plates, and through discus- 
sion gave me much insight into the prob- 
lems of photographic reproduction and of 
technical illustration. Barbara Jones, Mar- 
tin Bartlett, and George Grice have been 
good listeners, and have lent assistance at 
many points. James Craddock has criticized 
parts of the manuscript, in particular the 
keys. And Jane Peterson has never faltered 
in her cheerful typing, retyping, and typing 
again of the manuscript. 

Finally, I owe a special debt of thanks 
to two people who have discussed, criti- 
cized, and encouraged this work in all of 
its stages. The first of these is Richard H. 
Backus, Senior Scientist at the Woods Hole 
Oceanographic Institution. The second is 
Giles W. Mead, Curator of Fishes at the 
Museum of Comparative Zoology, my ad- 
viser at Harvard University, and the person 
who first introduced me to the stromateoid 


Known to the ancients, and investigated 
by such able ichthyologists as Giinther, Gill, 
and Regan, the development of the stromat- 
eoid classification has a long history. Only 
recently, however, has sufficient material 
become available to clarify the confusion 
surrounding the systematics of these fishes. 

The percifonn suborder Stromateoidei 
is diagnosed by the possession of toothed 
pharyngeal sacs and small uniserial teeth 
in the jaws. Comparative study of the na- 
ture of the pelvic and dorsal fins, the tooth 
pattern, the number of vertebrae and bran- 
chiostegal rays, and, in particular, the 
structure of the caudal skeleton and the 
pharyngeal sacs suggests a separation of the 
suborder into five families and fourteen 
genera. These are: Centrolophidae — 
Ihjpcroiijyphc, Schedophilus, Centrolophus, 
Icichthijs, ScrioIcJla, Psc)ioi).sis-, Nomeidae — 

Stromateoid Fishes • Haedrich 129 

Cuhiceps, Noirwus, Psenes; Ariommidae 
(fam. nov. ) — Aiiommo; Tetragoniiridae — 
Tetmg,onunis-, and Stromateidae — Stroma- 
teiis, Pcprilus, Fiimpus. The Centrolophidae 
are the most primitive in the suborder, and 
have given rise directly to the Stromateidae. 
The Nomeidae have evolved parallel to the 
centrolophid-stromateid line, and have 
gi\en rise to the Tetragonuridae and the 

Within the suborder, evolutionary trends 
from the generalized to the highly evolved 
condition are marked. The maximum size 
attained becomes smaller, and, in the stro- 
mateids, the relative depth of the body in- 
creases. The pelvic fins are lost. The mouth 
becomes smaller, the jaw teeth become 
cusped, and the palatine dentition may be 
lost. The phar)ngeal sacs become more 
elongate and the structure of the papillae 
within them becomes more complex. The 
number of branchiostegal rays is reduced 
from seven to five. The pseudobranch is 
lost. The vertebrae increase in number from 
a basic 25. The number of elements in the 
caudal skeleton is reduced through losses 
and fusions. 

The major features of the centrolophid 
distribution are discontinuity, bipolarity, 
endemism, and sympatry of genera. Four 
of the six genera occur in Australia and 
New Zealand. The three nomeid genera are 
broadly sympatric in temperate and tropi- 
cal oceans, and there are no relicts, no bi- 
polarity, and no regional endemism. The 
tetragonurid distribution is very poorly 
known, but is similar to that of the nomeids. 
The ariommids are found in deep water 
over the edge of the continental shelves 
from the east coast of the New World to 
Japan, and near Hawaii. The stromateid 
distribution is characterized by discontinu- 
ity, widespread species, restriction to con- 
tinental shelves, and allopatry of genera. 
None occur in Australia and New Zealand. 
The distributional data support the conclu- 
sions based on comparative morphology. 

The relationships and natural history of 
the stromateoid taxa are discussed. Svn- 

onymies, keys, and, under each genus, lists 
of nominal species are included. 


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u.':ri'i>.€ V?'-'JV 

^uitetin OF THE 

Museum of 



Morphology and Relationships of the Holocephali 
with Special Reference to the Venous System 


MwiiOMm of rnmpar flljye Zoology. Ca mbridge. Massachusetts 
J st i tuto di Bt o togter-Generak^UDraexsUa-df-PtsurlTaty — 



JANUARY 27, 1967 





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Reprint, $6.50 cloth. 

Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In- 
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Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth. 

Lyman, C. P. and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- 
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Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 15. ( Price list on 
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Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae 
(Mollusca: Bivalvia). $8.00 cloth. 

Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution 
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@ The President and Fellows of Harvard College 1967. 




Introduction __, 141 

Materials and Methods 143 

Description of the Venous System 144 

The Subcutaneous Veins 145 

The Deep Veins 

Precardiac Group 147 

Postcardiac Group ._ 151 

The Hepatic Portal System 154 

Discussion of the Significance of the Circula- 
tory System to the Problem of Holoce- 

phalian Relationships 155 

Discussion of Phylogenetic Clues from Other 

Organ Systems 162 

The Nervous System 162 

The Skeletal System _ 167 

The Muscular System 176 

The Urogenital System 181 

The Digestive System 186 

Conclusion 190 

Acknowledgments ._ 193 

References Cited 193 


Although the evolutionary relationships 
of the Holocephali have been under con- 
sideration for years, no one theory of their 
descent has appeared so satisfactory that 
the question may be laid to rest. This paper 
is the result of the continuing search for 
progress in clarifying the position of these 
fishes. To this end an investigation of the 
venous system of Chimacra colliei Lay and 
Bennett (Hydrolo^tis colliei ) has been un- 
dertaken, and a reassessment of the anatomy 
of the Holocephali has been made, taking 
into consideration both the structural ar- 
rangements revealed by the new dissection 
and current paleontological knowledge. 

The problem of classifying the holo- 
cephalian fishes (the living genera of which 
are: Chimacra, CaUorhijnchus, Rhinochi- 
maera, and Harriotta) has become more 
and more difficult as the understanding of 

^ St. Anselm College, Manchester, N.H. 

Bull. Mus. Comp. 

the history of fishes has grown. In contrast 
to Linnaeus who set the Holocephali down 
in the same group with the sharks, rays, 
sturgeons, and lampreys because of their 
cartilaginous skeleton, modem scholars are 
giving much thought to the propriety of 
even including them with the elasmo- 

The day has passed, too, \\'hen an inves- 
tigator could seek to solve the problem by 
focussing upon a single structure and sug- 
gesting that it indicates a probable evolu- 
tionary relationship. This point bears men- 
tion because the spotting of isolated simi- 
larities has generated several hypotheses, 
concerning the evolution of the Holocephali, 
which have proven untenable \\'hen an in- 
tensive examination is pressed. Based upon 
reasoning of this sort is the idea that the 
Holocephali might possibly be allied to the 
lungfishes through the common possession 
of autostyly and cutting toothplates. When 
the idea was tested by further study, it was 

Zool., 135(3): 141-213, January, 1967 141 

142 BiiUctiu Museum of Comparative Zoology, Vol. 135, No. 3 

shown that the nature of the palatoquadratc 
fusion was different, that tlie toothphites 
were surely not homologous structures, and 
that other anatomical characteristics were 
not alike. When paleontological evidence 
is considered, the probability of a relation- 
ship between the Holocephali and Dipnoi 
recedes still further. Despite the large 
amount of cartilage in the skeleton, lung- 
fishes have definitely sprung from ances- 
tral bony fishes, which sets them far from 
the holocephalians. The latter fishes arose 
probably from forms more nearly, though 
not necessarily very closely, allied with the 
ancestors of sharks than with the predeces- 
sors of the Osteichthyes. Assuming the 
truth of this statement, one can cast aside 
the hypotheses which link the Holocephali 
to fishes like Latimcrio and Pohjptenis 
whose position as bony fish is well estab- 
lished, and also those which embed the 
holocephalians in the line of fishes leading 
to tetrapods. 

Currently only two possibilities of holo- 
cephalian origin are receiving serious atten- 
tion. One opinion holds that holocephalians 
are aberrant off-shoots from ancient carti- 
laginous fishes. Specifically, this school 
favors the idea that the Holocephali are 
descendants of the bradyodonts, an extinct 
group of presumed shark relatives distin- 
guished by nonreplaceable teeth of a pecu- 
liar histological structure. Although no 
wealth of fossil material exists, paleontol- 
ogists have speculated that at least some 
of the bradyodonts were autostylic, as are 
the Holocephali. A leading advocate of the 
bradyodont origin of the Holocephali, Moy- 
Thomas ( 1936 ) has studied one of the rare 
bradyodont fossils which consist of more 
than teeth and spines and found in it many 
resemblances to holocephalian design. This 
specimen, the cochliodont Ilclodus simplex, 
dates from Carboniferous times. If it is 
ancestral to the Jurassic chimaerids, one 
must assume that all the distinctive holo- 
cephalian characteristics which Helodus 
does not possess were evolved in the inter- 
vening years. Other bradyodonts such as 

Menaspis and Omcantlius have been dis- 
cussed in relationship to the problems of 
holocephalian origin, but they have either 
possessed structures like the spines on the 
head of the former which makes one hesi- 
tate to place them in the direct ancestral 
line or they have been, like the latter, in too 
fragmentary a condition to allow a thorough 
comparison. In a recent paper, Patterson 
( 1965 ) concludes that the bradyodonts are 
closely enough related to the Holocephali 
to be grouped with them in the class Holo- 
cephali, but abandons the idea that Helodus 
or any other bradyodont is ancestral to the 
holocephalian line. 

The second possible source of the Holo- 
cephali is an older one. Amongst the 
ptyctodonts, a placodenu group, have been 
found several fossil forms that show charac- 
teristics which could be ancestral to those 
of holocephalians. The resemblances were 
recognized early (Pander, 1858), but ne- 
glected after the ptyctodonts were alhed 
with the arthrodires, and after Moy-Thomas 
offered, in Helodus, a bradyodont ancestor 
for the holocephalians which had long been 
classified in a general category with sharks. 
The idea of a ptyctodont ancestor has re- 
turned to favor, however, as the magnitude 
of the differences between holocephalians 
and sharks has been revealed. It seems now 
most attractive to find a stock, traceable 
far back into the Devonian, which could be 
ancestral to the Holocephali. There are a 
number of fossils ( in a more complete state 
than many of the cochliodont forms ) which 
have been used as a basis for comparison 
with extinct and Recent chimaerids. Of 
these forms, students of holocephalian evo- 
lution cite most often RliampJwdopsis, 
Pttjetodus, and Ctenuiella. The last is 
considered by 0rvig (1962) to show a re- 
markable number of similarities to the Holo- 
cephali. However, the ptyctodont-holo- 
cephalian relationship, while possible, is 
far from proved. There are still serious 
questions to be solved. One must suppose, 
for instance, if the relationship is a fact, 
that over the countless generations which 

Morphology and Relationships of Holocephali • Stahl 143 

separated the Devonian ptyctodonts from 
the Jurassic chimaerids the animals lost 
their distinctixe pattern of dermal armor, 
their pectoral spines, and their internal 
bone. While changes of this nature are not 
impossible, there is no fossil evidence to 
prove that they did take place. 

In trying to decide whether it is more 
likely that holocephalians originated from 
ptyctodonts than from a group closer to the 
shark line, one tiuns normally to the data 
available from embryological studies. In 
the case of the Holocephali, very little 
embryological work has been done. Since 
the holocephahan fishes lay their eggs, al- 
ready fertilized and enclosed in a case, in 
deep water, the embryos are not often ob- 
tained. There have been only two studies 
made of embryonic forms: that of Schauins- 
land on CoUorhynchus (1903) and that of 
Dean on Chimacia (1906). Although both 
studies were elegant pieces of work, a lack 
of certain stages resulted in the absence of 
observations of the fusion of the upper jaw 
and the step-by-step fomiation of the hyoid 
arch, for example. An understanding of 
these two points would shed great light 
upon the evolutionary question. 

Besides the paleontological and embryo- 
logical approaches, there is a third useful 
avenue of investigation. The contribution 
from the area of comparative anatomy can- 
not be omitted in assembling e\'idence 
which bears upon the problem. Although 
holocephahan fishes have been dissected 
numerous times, the work upon the anat- 
omy of these fishes is not completed. In 
early anatomical inxestigations the dissector 
often placed his emphasis upon structures 
which are not the best keys to the evolu- 
tionary problem. There is no information 
available concerning some of the areas 
which are of great interest from the com- 
parative point of view. 

One such area, that of the venous system, 
has been completely untouched. Although 
there have been publications concerning 
the distinctive portions of the arterial path- 
way, there is nothing in the literature about 

the pattern of vessels returning blood to the 
heart. It was in the hope that the venous 
system would show special features which 
might serve as clues to a better understand- 
ing of holocephahan evolution that this 
study was undertaken. 


Since it was desirable to avoid describ- 
ing as the general occurrence an anomalous 
vessel in a single fish, dissections were re- 
peated until it appeared certain that a par- 
ticular pattern was a normal and not an 
abnormal feature. The relatively large num- 
ber of specimens available made this method 
possible. The first specimen to be dissected 
was a female Chimaera coUiei, uninjected, 
which had been preserved in formalin and 
transferred to alcohol. A group of twelve 
specimens of Chimaera colliei were obtained 
fresh-frozen from Vancouver, B. C, through 
the kindness of Dr. Norman J. Wilimovsky. 
The procedure used with these animals 
was to defrost them, inject immediately with 
latex, preserve first in formalin, and after 
five days to transfer them in several steps 
to 70 per cent alcohol. The last six speci- 
mens of Chimaera coUiei, four females and 
two males, were received already injected 
with latex through the kind efforts of Dr. 
Richard Snyder. For comparative purposes 
one specimen of CaUorhynchus and one 
specimen of Rhinoehimaera were examined. 
Several methods of injection \\'ere tried. 
Because of the delicate nature of the vein- 
walls and the consequent similarity be- 
tween veins and strands of connective tis- 
sue in some areas, nothing was interpreted 
as a vein unless it was observed filled with 
an injecting material or remnants of brown- 
colored agglutinated blood. In the first 
uninjected specimen described above, a car- 
mine suspension was injected in area after 
area as the dissection proceeded. India ink 
was also tried. In the defrosted specimens, 
as noted, latex was used. Finallv, to fill 
certain empty areas in the professionally 
injected specimens, ordinary poster paint 

144 Bulletin Mu.scinii of Comparative Zoology, Vol. 135, No. 3 

was employed in its regular concentration 
and also in a slightly diluted form. 

All these media were introduced through 
a glass-barreled syringe fitted with a num- 
ber 23 needle inserted into an inch-and-a- 
half-long piece of polyethylene tubing, size 
50. The tubing was tied into the vessel 
through which the injection was made. 

The routes that were available for injec- 
tion were limited. Injection via a sinus 
proved impractical, because the injecting 
apparatus could not be tied tightly to the 
delicate sinus- wall. Very fine veins disin- 
tegrated under the most careful handling. 
Only large veins of well-defined cylindrical 
shape were useful. Injecting through them 
was hampered only by the presence of 
valves which restricted the amount of in- 
jection material able to pass beyond into 
tributary vessels. In particular, this diffi- 
culty arose in getting material to pass from 
the common cardinal vein forward into the 
anterior cardinal sinus and also in filling 
the deep veins of the fins. Although various 
vessels were tried as the dissection ad- 
vanced, for the initial attempt to fill as 
much of the venous system as possible, two 
veins were relied upon. To inject the he- 
patic portal system, the posterior dorsal in- 
testinal vein was employed. To fill the 
systemic vessels, injection was made into 
the lateral cutaneous vein immediately pos- 
terior to the scapula. This vein could be 
uncovered easily over a considerable dis- 
tance by removing the skin just below the 
lateral line. Injection was made through 
this vessel first in an anterior and then in a 
posterior direction. 


The veins return blood to the two com- 
mon cardinal vessels which lie medial to 
the anterior edge of the scapular process 
of the pectoral girdle and empty into the 
lateral corners of the sinus venosus. On 
either side, three veins meet at the point at 
which the last pharyngobranchial cartilage 
articulates with a facet of the scapula, to 
create the common cardinal of that side. 

These three are the anterior cardinal, the 
posterior cardinal, and the lateral cutaneous 
veins. The first comes from a fonvard di- 
rection, the second comes from the pos- 
terior region, and the third runs downward 
and slightly caudad to meet the other two. 
Into the upper end of the common cardinal, 
the inferior jugular vein opens. More ven- 
trally, the brachial sinus opens into the 
common cardinal from the posterior side 
(PI. 5, B, C). 

Each of the major venous trunks will be 
described with its tributaries and the areas 
which they drain. So that the description 
may be more easily understood, the pattern 
of the venous system is presented first in 
concise, outline, form: 

I. The Subcutaneous System 
Lateral eiitaneous vein 
Caudal tributary 
Axial tributaries 

Pelvic anastomotic area 
Clasper veins 
Ventro-lateral tributary 
Postscapular tributary 

Dorso-lateral axial branch 
Dorsal fin branch 
Prescapular tributary 

Dorsal ceplialic branch 
Anterior subcutaneous tributary 
Ventral cephalic branch 
Opercular branches 
Subscapular tributary 
II. The Deep Veins 

A. Precardiac vessels 
Anterior cardinal sinus 

Inferior jugular vein 
Posterior cerebral vein 
Postorbital vein 
Hyoid tributary 
Orbital sinus 

Maxillo-facial vein 
Preorbital branch 
Deep labial branch 
Orbito-nasal vein 
Posterior palatal vein 
Superior adductor mandibidar 

Anterior cerebral \ ein 

Posterior cerebral tributary 
Anterior cerebral tributary 
Ethmoidal \'ein 

B. Postcardiac vessels 
Brachial sinus 

Posterior brachial vein 

Morphology and Relationships of Holocephali • Stahl 145 

Anterior brachial \ e in 
Posterior cardinal sinus 

Ventro-anterior parietal vein 
Deep epaxial veins 
Dorsal fin sinus 

Medial dorsal vein 
Anterior epaxial \'ein 
Spino-basal vein 
Esophageal veins 
Anterior parietal veins 
Veins of the reproductive tract 
Renal veins 
Femoral vein 

Rectal tributary 

Dorsal fin tributaries 

Ventral fin tributaries 
Hepatic veins 
Renal portal veins 
Caudal vein 
Parietal veins 

Ventro-posterior parietal \ ein 
Hepatic portal vein 
Intra-intestinal vein 

Anterior dorsal intestinal tribu- 
Anterior ventral intestinal vein 
Mesenteric vein 

Dorsal posterior intestinal vein 

Ventral posterior intestinal vein 

Auxiliary splenic veins 

Lieno-pancreatic vein 

Auxiliary pancreatic veins 

The subcutaneous system is shown in 
Plate 1. The deeper veins are represented 
diagrammatically in Plates 2 and 3. 


There is an extensive system of sub- 
cutaneous drainage (PI. 1). The vessels 
which form it lie in the loose connective 
tissue under the skin. Although their path- 
ways vary somewhat in different specimens, 
the basic pattern of flow is generally the 
same. Assigning names to the vessels of 
this system is a hazardous business because 
of the numerous anastomotic connections 
which are present, but there are several 
principal trunks \\'hich can be specifically 

The chief collecting trunk deserves the 
name lateral cutaneous vein, for it courses 
anteriorly, paralleling the lateral line. In 
the caudal region it is located about a half 
inch belo\\' the lateral line canal, but at the 

level of the base of the pelvic fin it bends 
dorsally somewhat and can be followed for- 
ward into the trunk region where it is to be 
found just ventral to the lateral line. At its 
anterior end it continues forward lateral to 
the muscle-covered dorsal extension of the 
scapular cartilage, bends medially around 
the anterior edge of this cartilage, and then 
runs ventrally for a short distance to form, 
with the anterior and posterior cardinals, 
the common cardinal vein. As it passes ven- 
trally on the medial side of the scapula, it 
enlarges sufficiently to merit the name of 
subscapular sinus. Where it approaches the 
upper end of the common cardinal it is 
flanked by passing nerves, the anterior 
nerve trunk containing fibers of the cervical 
plexus which innervate the hypobranchial 
muscles, and the posterior trunk containing 
branches of the first through third spinal 
nerves. ( There is also in the anterior trunk 
a small group of visceral vagus fibers. ) 

The lateral cutaneous vein, as the prin- 
cipal trunk of the subcutaneous system, has 
the firmest wall of any vein involved in the 
superficial drainage. The toughness of the 
wall is due primarily to an ensheathing 
layer of dense connective tissue. This vein 
receives many tributaries which will be de- 
scribed below, beginning with those bring- 
ing blood from the most posterior regions. 

Although, in the caudal region, the lateral 
cutaneous runs forward a short distance 
ventral to the lateral line, there is another, 
smaller vein which accompanies the sensory 
canal. This caudal tributary turns ventrally 
to empty into the lateral cutaneous vein at 
the point along the length of the body 
which is on a level with the posterior limit 
of the pelvic fin attachment. 

As the lateral cutaneous vein courses for- 
ward, it collects blood returning from the 
superficial regions of the axial musculature 
dorsal and ventral to it. The axial tributaries 
are arranged in an orderly but not a rigidly 
segmental pattern. The dorsal tributaries 
are relatively short and in the region of the 
trunk posterior to the dorsal fin spine have 
as their source a network of little veins 

146 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 

which forms a narrow band dorsal to the 
lateral line and parallel to it. The ventral 
tributaries collect blood from a much 
greater area and in the pelvic region are 
considerably enlarged. There, they draw 
from an anastomotic network of veins. As 
part of that network, a vein can be seen 
running along the line of origin of the su- 
perficial levator muscle of the fin. The 
location of this line may be described as 
being about halfway between the lateral 
line above and the base of the pelvic fin 
below^ Into this vein run tributaries from 
the levator muscle, from the axial muscle 
medial to the levator, and from the axial 
muscles which are posterior and ventral to 
the pelvic region. These tributaries have 
connections, also, with two veins which to- 
gether encircle the base of the fin. One 
runs around the base laterally; the other 
runs around it medially, thus edging the 
anal region. These two vessels receive veins 
draining the fin web and the superficial 
muscles of the fin itself. In the male Chi- 
maera, the veins of the clasper, which re- 
ceive blood from the erectile tissue in the 
clasper tips, become superficial as they 
course proximally and empty into the ve- 
nous ring at the fin base (Pi. 4, A). The 
chief clasper veins are two which appear on 
the ventral side of the clasper. One drains 
each prong, and they merge shortly before 
emptying at the posterior edge of the fin 

From the anterior corner of the venous 
network in the pelvic region there flows 
forward a vessel of rather large size which 
gathers blood from the skin and superficial 
axial musculature ventral to the field served 
by the axial tributaries to the lateral cuta- 
neous vein. This vessel meanders forward 
over a slightly wavy pathway, finally curv- 
ing dorsally behind the pectoral region to 
empty into the lateral cutaneous trunk just 
before the latter turns inward around the 
anterior edge of the scapula. The name 
ventro-lateral tributary seems appropriate 
for this vein. In one specimen which had 
been injected with India ink, small veins 

were seen entering it from the posterior 
edge of the operculum dorsal to the gill 
opening and from the ventral part of the 
trunk immediately behind the opening from 
the gill chamber. The veins in this area 
were not injected successfully in any other 

The lateral cutaneous trunk receives two 
sizable tributaries bearing blood from dor- 
sal regions. The first one to be described 
begins lateral to the muscle-covered pos- 
terior tip of the scapular cartilage which is 
bound against the epaxial muscles at the 
base of the dorsal spine. This vein, called 
the post.scapular tril)iitani, receives blood 
from the dorsal fin branch, draining the 
web and muscles of the dorsal fin. Halfway 
along its course to the lateral cutaneous 
vein, the postscapular tributary receives the 
(lorso-Iateral axial branch. The latter vessel 
is a long one, running parallel but dorsal to 
the lateral cutaneous vein. It collects blood 
returning from the superficial epaxial mus- 
cles which lie dorsal to those drained by the 
axial tributaries of the main lateral trunk. 
Some of the branches which join the dorso- 
lateral axial branch can be seen to connect 
also with a median dorsal vessel whose 
blood flows eventually into the posterior 
cardinal sinus. These connections represent 
one of the few anastomoses between the 
subcutaneous and the deep venous drain- 
age systems. 

Far dorsally, near the base of the dorsal 
fin spine, there are prominent vessels which 
form an anastomosis between the postscap- 
ular vein and the second of the two sizable 
tributaries from the dorsal region. The 
second one, the prescapular tributary, 
courses ventrally just in front of the an- 
terior edge of the scapula to join the lateral 
cutaneous vein at the point at which it 
turns medially to meet the common cardi- 
nal. Shortly before emptying into the lat- 
eral cutaneous, the prescapular tributary 
receives the dorsal cephalic brancJi carrying 
blood from the flattened triangular-shaped 
dorsal surface of the head. Atop the head, 
the dorsal cephalic branches of the left and 

Morphology and Relationships of Holocephali • StaJil 147 

right sides are connected through anasto- 
mosing venules. As it runs toward its meet- 
ing with the prescapular, the dorsal 
cephalic follows the posterior portion of the 
supraorbital sensory canal, collecting blood 
from fine venules which parallel the mucous 
canals above the eye. A small vein draining 
the skin immediately above the orbit may 
empty into the dorsal cephalic branch or 
may be connected to the tributary next to 
be described. 

This tributary, called the anterior sub- 
cutaneous, empties into the lateral cutane- 
ous vein at the same point at which the 
prescapular enters it. Approaching that 
point, it courses dorso-posteriorly, approxi- 
mately paralleling the posterior quarter of 
the suborbital sensory canal. This vessel 
receives several opercular branches (some 
of which may anastomose with the ventro- 
lateral tributary near its anterior end). It 
receives also a ventral cephalic branch 
which drains veins collecting forward and 
ventral of the orbit and fine venules which 
parallel the group of mucous canals an- 
terior and ventral to the eye. The ventral 
cephalic branch may also receive blood 
from the region just posterior to the lower 
jaw, but in no specimen could the injection 
medium be made to penetrate that far for- 

The last tributary to the lateral cutane- 
ous vein \\'hich remains to be mentioned is 
the subscapular. This one is really a small 
sinus, lying against the medial surface of 
the scapular cartilage. It receives venules 
from the cartilage itself and from two fine 
veins which follo\A' the posterior border of 
the cartilage, one coming from a xentral 
and the other from a dorsal direction. The 
subscapular tributary is the last one to join 
the lateral cutaneous vein before it empties 
into the common cardinal vein. 


Precardiac Group 

The return of blood from the deep por- 
tion of the bodv anterior to the heart takes 

place through the anterior cardinal sinus. 
This sinus is exposed by lifting the dorsal 
constrictor muscle which covers the gill 
area. As the connective tissue beneath the 
muscle is cleared away dorsal to the oper- 
culum, the scalpel falls into the sinus. The 
blood-space lies lateral to a muscle origi- 
nating under the subocular shelf and insert- 
ing posteriorly upon the last pharyngobran- 
chial cartilages. This muscle, the trapezius 
internus of Vetter (Vetter, 1878), covers a 
portion of the branchial branches of the 
vagus nerve. The latter are visible through 
the medial wall of the anterior cardinal 
sinus for a short part of their pathway ven- 
tral to the muscle-band. The sinus is situ- 
ated dorso-laterally with respect to the 
efferent branchial arteries and entirely dor- 
sal to the branchial skeleton. 

Just as the anterior cardinal sinus, at its 
posterior end, curves slightly ventrad to join 
the common cardinal, it receives the inferior 
jufj,ular vein. This vein, which enters the 
sinus from the ventral side, has so broad a 
mouth that it might be interpreted as open- 
ing partially into the common cardinal it- 
self. The inferior jugular originates far 
anteriorly behind the lower jaw (Pi. 5, A ) . 
Although its main branch comes from within 
the hyoid "tongue" which protrudes from 
the floor of the mouth, branches also reach 
it from the thyroid gland, the ventro-medial 
fibers of the ventral constrictor muscle, and 
the anterior portion of the coracomandibu- 
laris. Veins from these sources were actu- 
ally seen, but it is also possible that there 
exist venules which failed to be injected, 
draining all the tissues located posterior to 
the mid-ventral portion of the mandible. 

About a centimeter behind the mandible, 
the inferior jugular vein turns medially and, 
running dorsal to the coracohyoideus mus- 
cle, almost meets its fellow of the opposite 
side. Without actually doing so, however, 
the vein turns posteriorly and takes a path 
lateral to the insertion points of the coraco- 
branchial muscle fibers upon the branchial 
cartilages. The vein follows the coraco- 
branchial insertion line, flaring widely from 

148 BiiUctin Museum of Comparative Zoology, Vol. 135, No. 3 

the ventral midline and curving dorsally as 
it does. This route leads the inferior jugular 
to the postero-ventral corner of the anterior 
cardinal sinus as described above. In its 
course along the inserting border of the 
coracobranchial, it receives blood from the 
lateral and medial sides of that muscle- 

The drainage of the coracomandibularis 
and coracohyoideus muscles is only partly 
accounted for by the inferior jugular vein. 
Although no other veins in this area were 
injected, dissections suggest that there may 
be a deep vein immediately ventral to the 
ventral aorta which provides additional 
drainage ( PL 4, B ) . From it blood may re- 
turn through small veins in the dorsal peri- 
cardial wall to the common cardinal or 
possibly over a more ventral course to a 
pair of veins, to be described below, which 
run through a channel in each side of the 
pectoral girdle. 

Farther forward than the entry-point of 
the inferior jugular the anterior cardinal 
sinus receives into its dorsal side the po.s- 
terior cerelyraJ vein (PI. 6, A). This vessel 
collects l)lood from fine veins over the cere- 
bellum and from membranes in the dorsal 
part of the cranial cavity. Since there is 
little likelihood, from the position of this 
vein, that it returns blood from any part 
of the cerebrum of Chimacra, the use of 
the teiTH "cerebral" in naming the vessel is 
technically incorrect. The adjective has 
been retained merely as a convenience to 
indicate that this vessel is the posterior of 
two draining the brain region. There is a 
possibility that fine veins which connect 
with the posterior cerebral may also con- 
nect with the orbital sinus via an anasto- 
mosing vein that passes through the wall of 
the orbit with the trochlear nerve. The 
existence of a vein traveling with that nerve 
was not clearly demonstrable, however, and 
so is best left in question. 

The posterior cerebral vein is formed as 
a median vessel in the dorsal portion of 
the cranial cavity over the medulla oblon- 
gata between the endolymphatic ducts. In 

addition to the blood from the brain and 
associated membranes, the posterior cere- 
bral receives tributaries from the inner ear 
of each side. These veins pass through the 
wide opening by which the cavity of the 
inner ear communicates with that of the 
brain and hence do not pierce cartilage. 
Immediately posterior to the point of its 
formation, by the confluence of the small 
vessels described, the posterior cerebral 
vein widens, over the rear part of the me- 
dulla, to fonn a small sinus. Into the pos- 
terior end of this sinus run several little 
tributaries carrying blood forward from the 
spinal cord. The blood collected in the 
sinus leaves it through two veins which may 
be considered as paired posterior cerebrals, 
continuing from the median vessel. Each 
passes directly into a long, ventrally directed 
channel in the cartilage on its own side of 
the chondrocranium. Each channel, occu- 
pied solely by the paired portion of the 
posterior cerebral vein, tenninates by pass- 
ing dorsal to the vagus nerve (which is 
also traversing the cartilage at that point) 
and opening ventro-laterally, anterior to 
the foramen of the latter. The posterior 
cerebral vein runs forward close under the 
otic region of the chondrocranium and then 
tiuns laterally at the level of the posterior 
limit of the semicircular canals to join the 
anterior cardinal sinus. 

At the anterior end of the anterior cardi- 
nal sinus, lies the opening of the postorbital 
vein (PI. 6, A). This vein, which travels 
through the posterior wall of the subocular 
shelf, in a ventral direction, with the hyo- 
mandibular branch of the seventh nerve, 
forms a bridge between the orbital sinus 
and the anterior cardinal. As it enters the 
latter, dorsal nutrient veins from the gill 
septa were seen, in one specimen, to send 
a common stem dorsally to this blood chan- 
nel. In no other specimen were these little 
veins detected. 

The postorbital vein was examined with 
care, for it was expected that the hyoid 
sinus should open into it or nearby. How- 
ever, no evidence was found in any speci- 

Morphology and Relationships of Holocephali • Stahl 149 

men of the existence of a shark-like hyoid 
sinus. There was only a small vein, the 
hyoid tributary (PI. 7, B), which could be 
traced ventrally to the dorsal tip of the 
ceratohyal cartilage and no further as a dis- 
sectable vessel. Posterior to the ceratohyal 
cartilage and anterior to the afferent bran- 
chial artery, in the position of the selachian 
hyoid sinus, it was possible to trace an un- 
injected vein in specimens with a favorable 
distribution of agglutinated blood. Al- 
though a connection \\'ith the above-de- 
scribed small vein was not clearly seen, it 
is possible that there was one and that this 
entire blood pathway is homologous to the 
selachian hyoid vessel. 

In each dissection of the region ventro- 
anterior to the postorbital vein, the subocu- 
lar shelf and the cartilaginous bar which 
runs to the mandibular articulation were 
removed after examination of the bordering 
tissues. Beneath the cartilage and immedi- 
ately dorsal to the skin of the roof of the 
mouth was a layer of loose connective tis- 
sue. The veins running through it were 
visible only when they remained filled with 
blood, as the injection mass never pene- 
trated to them. They anastomosed with 
each other and one ran to join the post- 
orbital as it emerged from beneath the sub- 
ocular shelf (PI. 7, B ) . It seems possible 
that the vessel traced from the postorbital 
vein to the tip of the ceratohyal may have 
connections with the veins of the connec- 
tive tissue layer via a fine vessel which 
passes forward, dorsal to the ceratohyal, in 
company with the efferent pseudobranchial 
artery. This artery passes dorsally, pierces 
the chondrocranium, and splits into the 
cerebral and optic arteries. ( In its dorsal 
course it runs along the posterior edge of 
the lymphomyeloid mass [Kolmer, 1923] 
dorsal to the skin of the palate. ) Although 
it is difficult to discern, it is probable that 
a vein travels with the artery. Judging 
from the pathway of the artery, this vein 
might have connections to the network of 
veins in the connective tissue just described 
and to the orbital sinus as well. It is also 

possible that some drainage from the base 
of the brain might be carried to the orbital 
sinus or to the postorbital vein via the path- 
ways which exist through the connective 

The orbital sinus receives all the blood 
returning from the head except that which 
passes through the subcutaneous vessels, 
the inferior jugular, and the posterior cere- 
bral veins. The sinus encircles the orbit 
medial to the nerves running through it. 
Intimately connected with this sinus is 
lymphomyeloid tissue. This tissue, which 
seems to be situated in the lateral edges of 
the blood-space, is present in such quantity 
at the ventro-anterior corner of the orbit 
that it bulges laterally in two sizable masses 
which are visible as soon as the skin is re- 
moved from that area. When the skin, con- 
nective tissues, and mucous canals^ are 
removed from the head in front of and be- 
low the orbit, the largest tributary to the 
orbital sinus can be seen. This vessel, the 
maxillo-facial vein, coursing dorsally over 
the posterior palatoquadrate region and up 
over the subocular shelf, enters the ventral 
side of the orbital sinus anterior to the point 
at which the postorbital vein leaves it (Pi. 
6, A ) . As the maxillo-facial vein approaches 
the sinus it assumes a position medial to 
the nerves which run out of the orbit. The 
vein carries blood from the deep portions 
of the overlying mucous canals, from the 
dermis of the upper lip and the area above 
it lateral to the labial cartilages, and from 
the muscles of the facial region which in- 
sert upon the labial cartilages and the lower 
jaw. As the maxillo-facial vein approaches 
the orbital sinus, it is joined by the pre- 
orbitaJ branch, draining the muscle tissue 
anterior to the eye, and by small veins 

1 In the head region of Chimaera there are two 
groups of tubules lying under the skin which exude 
mucus through pores at their posterior ends. The 
dorsal set of six parallel tubules lies above the orbit 
and extends behind it. The ventral tubules, ap- 
proximately the same in number and arrangement, 
cover an area of the face ventral to the eye and 
anterior to it. 

150 Bulletin Museum of Coinjxiidtive Zoology, Vol. 135, No. 3 

which come from tissues lying just posterior because its course Hes over that area. Its 

to the maxillo-facial vein itself. relation to the suprapalatal lymphomyeloid 

Since the maxillo-facial vein was nearly tissue suggests that it plays a part in drain- 
empty of blood in the frozen-and-thawed ing it. This vessel is never filled with the 
specimens and did not proxe amenable to injection mass. Its presence is demonstrable 
injection, its anatomy was studied in the only because of the blood left in it. 
fish which had been injected with latex and There are also veins which enter the 
preserved immediately in formalin. In these orbital sinus in its antero-dorsal comer. To 
animals the maxillo-facial vein was ob- reach the orbital sinus at this point, the 
served only upon the left side. The right veins must traverse the posterior part of 
side showed what appeared to be a large the ethmoid canal. The canal is a large, 
sub-surface pool of agglutinated blood — cartilage-roofed, median space dorsal to 
surely an artifact. Since the veins on the the portion of the cranial cavity occupied 
left were entirely empty, it is probable that by the elongated telencephalon. It is sep- 
the fresh-caught fish were stored right-side- arated from the brain cavity by a cartilag- 
downward, causing the blood to accumu- inous partition. The ethmoid canal, which 
late and to obliterate the vessels on that is filled with lymphomyeloid material, en- 
side. Although the vessels of the left side closes the ophthalmic nerves as they pass 
of the face were not filled with latex, it was from the orbital region towards the snout, 
possible to trace them by injecting poster- One of the veins which passes through the 
paint into the orbital sinus and expressing ethmoid canal on its way to the orbital sinus 
it into the facial veins by pressing gently is the small .superior adductor mandibidor 
upon the eye. vein. It drains the most dorsal portion of 

A small deep labial branch of the maxillo- the deep adductor muscle. Leaving the 

facial vein brings blood from the lower lip muscle, the vein passes inward through the 

and jaw, the upper jaw region medial to wall of the ethmoid canal and joins the 

the labial cartilages, and the nasal capsule path of the superficial ophthalmic trunk, 

(PI. 7, A). traveling with it through its foramen into 

There are two other veins which enter the orbit, 
the orbital sinus from the ventral side, but A second vein, the anterior cerebral, en- 
both of them pierce the subocular cartilage ters the orbital sinus by passing through a 
to do it. The orbito-na.sal vein passes through foramen in the cartilage between the back 
its own foramen. As it travels toward the of the ethmoid canal and the front edge of 
foramen from the nasal region, it lies the orbit (PI. 5, D). The foramen, which 
against the dorsal surface of the supra- is medial to the departure-point of the 
palatal lymphomyeloid mass. Tracing this ophthalmic profundus from the orbit, trans- 
vein anteriorly, one finds that it can no mits only this vessel. The anterior cerebral 
longer be separated from the lymphomye- vein brings blood back from the anterior 
loid tissue where the anterior tip of the end of the brain. It is formed as a median 
mass abuts the posterior side of the nasal vessel within the cranial cavity by the union 
capsule. of a posterior and an anterior cerebral trib- 

The second vein which reaches the or- utary. The posterior tributary runs from the 
bital sinus by piercing the subocular shelf tip of the long epiphysis (which extends 
has already been mentioned. This is the forward to a position above the inter- 
vessel which accompanies the efferent pseu- orbital area ) ventrad in a course which 
dobranchial artery along the posterior edge follows the curving posterior edge of the 
of the lymphomyeloid mass and thence interorbital septum. The anterior cerebral 
through its subocular foramen. The vein tributary, which drains the telencephalic 
has been called the posterior pcdatal vein lobes, follows a dorsal pathway posteriorly 

Morphology and Relationships of Holocephali • Stahl 151 

through the cranial cavity to meet the pos- 
terior tributary just below the ventral edge 
of the interorbital septum. The anterior 
cerebral vein produced by the union of 
the two tributaries passes antero-dorsally 
through the edge of the interorbital parti- 
tion and then through a short channel in 
the cartilage to enter the posterior end of 
the ethmoid canal. There it bifurcates. 
Each branch turns posteriorly to enter the 
orbital sinus on its own side. Just before 
it leaves the ethmoidal canal each portion 
of the bifurcated anterior cerebral vein 
receives an ethmoidal vein. The ethmoidal 
veins bring blood back through the ethmoid 
canal from the most rostral part of the 
snout. These vessels enter the anterior end 
of the canal through the same pair of 
foramina through which the superficial 
ophthalmic nerves issue. 

Postcardiac Group 

Ventral to the confluence of the anterior 
cardinal, lateral cutaneous, and posterior 
cardinal trunks, there is an opening into 
the posterior side of the common cardinal 
vein from the brachial sinus. That sinus, 
which receives all the blood returning from 
the pectoral fin, lies behind the base of the 
fin in the angle between it and the body 
wall. There is an extension of the sinus 
ventrally along the posterior side of the 
pectoral girdle which meets its pair in the 
midline. The entrance of the brachial 
sinus into the common cardinal is edged 
by a shaip fold which acts as a valve. It 
is this valve, apparently, which prevents 
good injection of the pectoral veins. 

The brachial sinus receives blood from 
two sources. The larger contributor is the 
posterior brachial vein. It borders the 
posterior edge of the muscle mass of the 
fin. In dorsal view it can be seen running 
along the posterior side of a deep levator 
of the fin (Pi. 5, B). Although neither 
Vetter (1878) nor Shann (1919) gives a 
specific name to this muscle, it can be 

recognized easily through its origin from 
the postero-medial surface of the scapula, 
its strap-hke shape, and its insertion upon 
the metapterygial cartilage. The brachial 
nerves run from the body wall through the 
axial region toward the posterior brachial 
vein. Upon reaching it, they divide into 
dorsal and ventral branches, the former 
passing over the vein and the latter under 
it. The posterior brachial vein receives 
blood from the deep portions of the posterior 
half of the pectoral fin. 

The second source of the blood collected 
by each brachial sinus is the anterior 
brachial vein. This vein emerges from a 
channel in the cartilage of the pectoral 
girdle to pour its contents into the sinus. 
The channel, which for most of its length 
contains the brachial artery as well as the 
anterior brachial vein, is a long one, 
piercing the cartilage in the coracoid re- 
gion at a point close to the ventral midline 
and running dorsally through the girdle to 
open on the medial edge of the scapular 
process near the brachial sinus. Between 
its beginning in the coracoid area and its 
temiination adjacent to the sinus, the chan- 
nel opens to the surface twice more: there 
is a foramen facing ventro-laterally anterior 
to the articulation of the fin and another 
facing posteriorly dorsal to the base of the 
fin. Although the most ventral opening of 
the channel is sizable and set in the anterior 
side of the coracoid bar, no veins could be 
seen entering it from the coracomandibular 
muscle which originates from that surface 
of the girdle. Since the veins draining the 
muscle fibers in that area remained un- 
injected in every specimen, it is possible 
that such veins do exist but \\'ere not 

Between the entrance to the channel in 
the coracoid area and the ventro-lateral 
foramen mentioned above, the channel is 
filled with lymphomyeloid tissue like that 
in the head region. If the passage does 
carry a \'ein from the area of the hypo- 
branchial musculature, the vessel would 
undoubtedlv have connections with the 

152 Bullet hi Museum of Comparatwe Zoology, Vol. 135, No. 3 

vascular network of the lymphomyeloid 
substance. The first vein which appears 
certainly in the channel, however, is the 
tributary draining the deep, anterior ven- 
tral part of the fin. This vessel enters the 
passage through the ventro-lateral opening 
and follows the path of the channel dorsad. 
A tributary from the deep part of the an- 
terior dorsal half of the fin enters the chan- 
nel next, through the posterior foramen, 
and merges with the tributary from the 
ventral part of the fin to form the anterior 
brachial vein. It is this vein which leaves 
the channel at its dorsal termination to 
enter the brachial sinus. 

Of the major trunks which empty into 
the common cardinal vein, the only one 
which remains to be described is the pos- 
terior cardinal .sinii.^. Although this vessel 
is paired, there are numerous, sizable com- 
munications between the left and right 
sides, and posteriorly, at the origin of the 
trunk between the kidneys, there is a single 
median portion. There are four constant 
features concerning the anterior portion of 
this sinus which should be noted. Firstly, 
the entrance into the common cardinal of 
each side is cavernous. An injection mass 
introduced into the lateral cutaneous vein 
always descends and turns posteriorly into 
the posterior cardinal sinus rather than en- 
tering the smaller opening of the anterior 
cardinal. Just as the sinus approaches the 
common cardinal, the subclavian artery and 
two spinal nerve branches cross through it. 
Secondly, there seems to be a connection 
between the posterior cardinal and the 
brachial sinus. The connecting passage 
runs from the ventro-lateral edge of the 
posterior cardinal to the brachial sinus 
dorsal and posterior to its opening into the 
common cardinal. Thirdly, the left and 
right posterior cardinal sinuses extend 
ventro-laterally to meet each other in the 
ventral midline. This midline communica- 
tion parallels that of the brachial sinuses 
and is separated from it by a sheet of con- 
nective tissue. It is to this part of the 
posterior cardinal sinus that the left and 

right ventro-anterior parietal veins bring 
blood from the deep anterior ventral and 
anterior ventro-lateral axial musculature. 
Fourthly, the sinus of each side extends 
dorso-medially as a blind pouch forward 
of its point of union with the anterior 
cardinal sinus. Thus, a cross-section made 
just in front of the anterior edge of the 
scapula shows the left and right pouches 
close to the midline above the branchial 
region and the anterior cardinal sinus of 
each side lying in a more ventro-lateral 

Since the posterior cardinal sinus runs 
retroperitoneally against the dorsal body 
wall between the dorsal aorta and the more 
laterally placed kidney, the veins from the 
deep epaxial muscles surely empty into it. 
These fine deep epaxial veins were not 
injected and so remained invisible, but 
their presence may be predicated with 

Besides this drainage and that from 
superficial regions of the dorsal muscula- 
ture via the subcutaneous svstem, there is 
one other route to be mentioned: between 
the left and right epaxial muscle groups 
in the trunk region can be found a median 
dorsal vein (PI. 6, B). This vessel was 
injected successfully and seen to collect 
from the most dorsal parts of the muscula- 
ture. Anastomoses existed between its trib- 
utary veins and those of the subcutaneous 
system. The median dorsal vein carries its 
blood forward to a median fin sinus 
set behind the base of the dorsal fin. This 
sinus has a single anterior opening on each 
side through which blood leaves it. These 
openings lead to the posterior cardinal 
sinuses. The blood returned from the 
median sinus by this route enters the pos- 
terior cardinal far forward, flowing into 
the blind pouch which extends anteriorly 
and dorsomedially into the anterior tip of 
the coelom. 

Posterior to the opening of this sinus, on 
a line with the base of the dorsal spine, an 
anterior epaxial vein enters the posterior 
cardinal on each side, bearing blood from 

Morphology and Relationships of Holocephali • Stahl 


deep muscles forward of the dorsal fin. 
Medial to its point of entry is the cartilage 
plate which supports the dorsal spine. the side of this plate runs a vessel, 
the spino-bosal vein, which connects the 
subcutaneous veins at the base of the fin- 
spine with the posterior cardinal sinus deep 

The remaining tributaries to the posterior 
cardinal enter it more ventrally. There are 
several which come from the esophageal 
wall, leaving it as the gut tulie makes its 
entry into the anterior end of the body 
cavity. In the region of tlie trunk anterior 
to the kidneys, anterior parietal veins on 
each side contribute l)lood from the most 
dorsal portions of the hypaxial musculature. 
The gonads and the ducts of the reproduc- 
tive tract send their blood to the posterior 
cardinal, too. The blood from these struc- 
tures seems to collect in sinuses between 
the double walls of the suspending dorsal 
mesentery. The sinus parallelling the ovi- 
duct of the mature female is quite spacious. 
There are veins running medially from it 
over the short distance to the posterior 
cardinal sinus. Around the anterior end of 
the functional kidne\ there is a wider com- 
munication between the two sinuses. The 
male fishes available for dissection were 
small and apparently not fully mature. The 
vas deferens was very fine and bound 
closely to the lateral border of the gland 
of Leydig (the transformed anterior end 
of the kidney ) . The venous drainage of the 
duct was invisible. 

The renal veins run their usual short 
course, leaving the kidneys ventro-medially 
and entering the posterior cardinal sinus. 
Posterior to the entrance of all but a few of 
the renal veins the posterior cardinal sinus 
in Chimaera receives a vein which does not 
usually empty into it in cartilaginous fishes. 
That vessel, the large, firm-walled femoral 
vein (PI. 7, C), is situated posterior to the 
femoral artery and, at the fin-base, is 
formed from dorsal and ventral fin tribu- 
taries, branches which come from the deep 
dorsal and ventral surfaces of the fin. Al- 

though the femoral vein is strong enough to 
withstand the injecting process, it proved 
impractical to use it as a route for injecting 
the deep veins of the pelvic fin because 
of a valve at its distal end which prevents 

A short distance proximal to the fin-loase, 
the femoral vein receives the rectal trib- 
utary. This vessel collects blood from a 
capillary network in the wall of the rectum 
and also, in the female, from the problem- 
atical glandular "seminal receptacle." 

As the femoral vein runs toward the pos- 
terior cardinal sinus it passes the lateral 
edge of the kidney and turns ventral to it. 
There is, however, a small branch which 
leaves the femoral, passes over the lateral 
edge of the kidney to the dorsal side and 
connects with the renal portal vein. The 
connection, the iliac vein (PI. 3), is a 
delicate one and possibly not uniformly 
present. Because of the small size of the 
vessel its functional significance is doubtful. 

The last contributor to the posterior car- 
dinal sinus is a strange one for any verte- 
brate. Without a doubt, the hepatic veins 
( PI. 9, A and B ) empty into this dorsal 
channel instead of passing forward through 
the transverse septum to enter the sinus 
venosus. Inspection of the posterior wall 
of the sinus venosus showed a pair of open- 
ings so reduced as to be incapable of carry- 
ing the entire bloocbflow from the liver. 
When the liver was cut transversely through 
the small area which is bound to the back 
of the transverse septum, there were no 
sinuses to be seen in the interior of the 
tissue. In Chimaera, however, the anterior 
attachment of the liver to the back of the 
transverse septum is not the only bridge 
between that gland and surrounding tissues 
across which a hepatic vein might travel. 
The front portion of each lobe is attached 
dorsally to the underside of the posterior 
cardinal sinus and thus to the roof of the 
coelom. The line of attachment extends 
along the dorsal edge of the right lobe of 
the liver for about a third of its length and 
along that of the shorter left lobe for half 

154 BuUeiin Mu.scmn of Comparative Zoology, Vol. 135, No. 3 

of its length. When these areas of at- 
tachment were explored, it was found 
that hepatic veins of varying sizes passed 
through them, carrying blood from the liver 
directly to the posterior cardinal sinuses. 
The largest of these veins was one which 
drained the posterior two-thirds of the 
right lobe and entered the right posterior 
cardinal at a point immediately anterior 
to a mesentery strap extending from the 
midline, between the sinuses, to the tip 
of the pancreas. If the sinus wall is cut 
and deflected at that point, the dissector 
can look through a fenestrated membrane 
separating the sinus from the interior of 
the hepatic vein. This is the largest of all 
the hepatic veins and drains the entire free 
end of the large right lobe of the liver. For 
the most part, the branches feeding this 
hepatic vein lie dorsal to those from the 
hepatic portal vein which carry blood into 
the right lobe. Anterior to the point at 
which this large hepatic vein enters the 
posterior cardinal sinus, smaller hepatic 
veins enter the sinus separately. Although 
the left lobe of the liver is smaller, it is 
similarly drained. There are two main 
hepatic veins that leave it to enter the 
posterior cardinal sinus on the left side. 

To conclude the description of the sys- 
temic venous drainage, one turns to the 
rcnml portal veins (Pi. 8, A). The blood 
entering the kidneys from the renal portal 
veins comes largely from the deep portions 
of the axial musculature. The muscles of 
the tail region send ])lood to the caudal 
vein which bifurcates behind the body 
cavity to become the left and right renal 
portal veins. These run forward along the 
dorsal surface of each kidney, medial to 
the mesonephric duct. At segmental inter- 
vals the renal portals receive veins from 
the body wall. The renal portals extend 
far forward, even beyond the region of the 
functional kidney, collecting the segmental 
parietal veins and finally dwindling to 
nothing about a centimeter behind the pec- 
toral fin. At the level of the anterior edge 
of the pelvic girdle the renal portal receives 

the ventro-posterior parietal vein. This ves- 
sel can be seen through the peritoneum, 
lying parallel to the long axis of the body. 
It begins in the middle region of the trunk 
and carries blood posteriorly from the deep 
hypaxial muscles. Immediately anterior to 
the pelvic girdle it curves dorsally to join 
the renal portal. The only tributaries to the 
renal portal which do not return blood from 
axial musculature are the small veins from 
the posterior end of the mesovarium. For 
a short distance beyond the end of the 
oviducal sinus in the crowded posterior part 
of the body cavity, these vessels from the 
lower end of the oviduct enter segmental 
parietal veins as they are about to join the 
renal portal. 

The Hepatic Portal System 

Since the nature of the digestive tract 
and the arrangement in the body cavity 
of the associated glands are distinctive in 
chimaerid fishes, the pattern of the veins 
draining the system is also singular. Before 
trying to visualize the path of the vessels, 
one must understand several anatomical 
points. Firstly, there is no stomach in these 
animals. Instead of a long J-shaped struc- 
ture there is a short continuation of the 
esophagus which reaches the beginning of 
the spiral intestine. Secondly, the spleen 
is not attached to the gut tube by mesen- 
teries. The mesenteries are exceedingly 
reduced and the spleen is fairly free, bound 
only to the posterior end of the pancreas. 
When a fresh fish is opened, the pancreas 
and spleen appear to lie ventrally in the 
body cavity. Because the spleen has no 
relation to a stomach, the term "gastro- 
splenic" or "lieno-gastric" is not applicable 
to any vessel in the hepatic portal system. 
It is well to keep these facts in mind when 
pondering possible homologies between the 
vessels of chimaerid fishes and any others. 

The he]Hitie portal vein is formed against 
the surface of the liver posterior to the base 
of the gall bladder by the confluence of 
the intra-intestinal, the anterior ventral in- 
testinal, and the mesenteric veins (Pi. (S, B). 

Morphology and Relationships of Holocephali • Siahl 155 

It runs immediately into the right lobe of 
the liver in one direction and, in the other, 
sends a large division along the posterior 
edge of the midventral portion of the liver. 
The blood carried in this vessel is distrib- 
uted to the left lobe of the liver. 

The first of the vessels which deliver 
blood to the hepatic portal, the intra- 
intestinol vein, drains the spiral valve and, 
as it emerges from the intestine wall, re- 
ceives anterior dorsal intestinal trilmtaries. 
These vessels collect blood not only from 
the anterior wall of the spiral intestine but 
also from the posterior region of the esoph- 
agus which connects with it. These trib- 
utaries anastomose \\'ith others which con- 
verge to form the anterior ventral intestinal 
vein (PI. 10, A). The intra-intestinal and 
the anterior ventral intestinal leave the 
intestine wall from points lying close 
against opposite sides of the bile duct. 
Running closely apposed to the duct, these 
vessels finally reach the hepatic portal vein. 

The last of the vessels which contribute 
blood to the hepatic portal, the mesenteric 
vein, is fonned by the confluence of the 
two posterior intestinal veins. The pos- 
terior dorsal intestinal vein, which drains 
the lower end of the intestine as far as 
the beginning of the rectum, receives trib- 
utaries which can be seen on the surface 
of the intestine wall. These tributaries 
anastomose with others which converge on 
the opposite side of the intestine to form 
the posterior ventral intestinal vein ( PI. 10, 
A). Both posterior intestinal veins leave 
the surface of the intestine to run free to 
a position against the side of the pancreas 
where they merge and are bound down. 
In their free portions, the dorsal and ventral 
veins have a different appearance: the 
ventral one is narro\\'er and runs through a 
band of mesentery; the dorsal one is very 
wide in diameter and absolutelv uncon- 
fined. At the point at which they merge 
and are tied to the pancreas, there is a 
thin mesentery strap which leaves to reach 
the dorsal midline behind the dorsal attach- 
ment of the liver. 

The mesenteric vein, thus formed, re- 
ceives several auxiliary splenic veins (the 
spleen is tied to the posterior end of the 
pancreas just behind the point where the 
two intestinals reach it) and then receives 
the relatively large lieno-pancreatic vein. 
The latter vessel travels through the length 
of the spleen and the portion of the pan- 
creas which lies posterior to the origin of 
the mesenteric. After receiving the lieno- 
pancreatic vein, the mesenteric turns an- 
teriorly and runs in company with the 
pancreatico-splenic artery, collecting from 
auxiliary pancreatic veins in its course. The 
mesenteric enters the hepatic portal in con- 
junction with the intra-intestinal vein (Pi. 
10, B). 

Additional Observations 

One specimen of Callorhynchus, a small 
female, was examined after the dissections 
of Chimaera colliei were completed. It was 
found that the pattern of the confluence of 
the major venous trunks to form the com- 
mon cardinal agreed with the findings in 
Chimaera. The hepatic veins were also 
found to enter the posterior cardinal sinus. 
An inspection of the posterior wall of the 
sinus venosus showed extremely small aper- 
tures that were similar to the reduced 
hepatic openings in Chimaera. 


The venous system of the chimaerids 
bears a greater resemblance to that of 
sharks both in the structure of its vessels 
and their arrangement than it does to the 
system of any other group of extant fishes. 
There are certain deviations from the 
selachian plan, however, which are cer- 
tainly clues to the separate evolution of 
the holocephalian line. A consideration of 
the significance of the venous system to 
the question of holocephalian relationships 
necessitates first, recognition of the resem- 

156 Bulletin Museum of Conjparativr Zoology, Vol. 135, No. 3 

blances, and then evaluation of the differ- 
ences which exist. 

The veins in both groups of animals, like 
those of all fishes, possess little muscular 
tissue. Their walls are therefore exceed- 
ingl\ delicate and difficult to differentiate 
from connective tissue in gross dissection. 
Even the main vessels may be opened by 
a chance touch of a scalpel tip. It is the 
combination of fragile walls and the pres- 
ence of valves which gives rise in both 
holocephalians and selachians to the diffi- 
culties experienced in injection procedures. 
The lack of detailed description of the 
drainage of the head region stems directly 
from the inability of investigators to intro- 
duce substances into the veins which empty 
into the orbital sinuses. The valves, which 
are mere folds of the lining of the vein 
wall, seem to have a like distribution in 
chimaerids and sharks if non-penetration 
of injection media can be taken as a guide 
to their location. The dissector can see 
that the entrance into the common cardinal 
from the anterior cardinal is valved but 
that the opening from the posterior cardinal 
is not. This arrangement is also shown for 
Hcptanchus by Daniel (1934). 

The main venous channels in holoce- 
phalians as in sharks are sinuses. In both 
types of fishes the largest ones are held 
together from within by a network of con- 
nective tissue trabeculae. The position of 
these sinuses relative to each other is not 
distinctive in the Holocephali. 

A great part of the basic arrangement of 
the venous system of selachians is dupli- 
cated in Chimaera colliei. The orbital sinus 
is the major collecting point for blood re- 
tinning from the tissues of the head. As 
in Ueptanchus, Mu.stcJus, and ScylUtim, it 
receives the orbito-nasal and anterior cere- 
bral veins. Although Daniel (1934), Parker 
(18S6), and O'Donoghue (1914), who in- 
vestigated the three sharks named, respec- 
tively, do not describe in detail the specific 
structures drained by the orbito-nasal vein, 
the vessel in Chhiuwra is probably exactly 
comparable except that it receives blood 

from the palatal lymphomyeloid mass 
which the sharks do not possess. The an- 
terior cerebral vein of Chimaera drains the 
same regions of the brain as the selachian 
vessel, although there is no reception of 
an ethmoidal vein in sharks, which lack an 
ethmoidal canal. The dissection of Chi- 
maera revealed a maxillo-facial vein and 
two smaller vessels which also empty into 
the orbital sinus. That there are no com- 
parable vessels shown for sharks is prob- 
ably due to their having been uninjected 
and unreported rather than to their absence. 

A postorbital channel exists in both holo- 
cephalians and sharks to carry blood from 
the orbital sinus to the anterior cardinal. 
Although its location relative to the two 
blood spaces it connects is the same in both 
types of fishes, its associations with skel- 
etal and nervous elements are not identical. 
In Chimaera the postorbital vein passes 
through a foramen in the posterior comer 
of the orbit accompanied by the hyoman- 
dibular branch of the seventh nerve. In 
sharks, the vessel merely lies in a post- 
orbital groove and the hyomandibular 
nerve, which never enters the orbit, does 
not share this anterior pathway. It seems 
likely that the postorbital veins of the two 
types of fish are homologous and that the 
different nervous and skeletal arrangements 
are due to the autostyly and forward com- 
pression of the cephalic structures in Chi- 
maera. The presence of a foramen rather 
than a groove, for example, is due to the 
fusion with the cranium proper of an otic 
process extending from the jaw joint to the 
ear region. This cartilage provides the en- 
tire lateral wall of the foramen. The for- 
ward course of the hyomandibular nerve 
can be explained by the anterior displace- 
ment of the tissues it serves and by the 
absence of a spiracle and hyomandibular 
cartilage behind which it \\'Ould normally 

The chimaerid anterior cardinal sinus 
carries blood over the gill region to the 
common cardinal in the shark-like manner, 
receiving in its course the posterior cerebral 

Morphology and Relationships of Holocephali • Stahl 157 

vein. The way in which the anterior car- 
dinal and the other large veins of sharks 
meet to fomi the common cardinal vessel 
shows a degree of variation sufficient to 
preclude the interpretation of the holo- 
cephalian arrangement as worthy of special 
note. Even the connection of the lateral 
cutaneous vein via the subscapular sinus 
to this confluence of vessels has its proto- 
type amongst the selachians. 

In the postcardiac region the similarity 
of the selachian and holocephalian plan is 
still evident. The renal portal and posterior 
cardinal vessels are substantially the same 
in both groups. Although the hepatic portal 
system will receive special consideration 
below, it may also be generally described 
as more like that of sharks than that of 
other fishes. It is in the drainage of the 
pelvic fins, the ventral body wall, and the 
liver that significant differences do appear 
in the chimaerids. 

The possession of a subcutaneous system 
of veins which run unaccompanied by 
arteries is a final point of likeness which 
should be mentioned. In sharks as well 
as chimaerids, the chief vessels in the net- 
work, the lateral cutaneous veins, receive 
segmental tributaries and finally lead to 
the subscapular sinuses. There are connec- 
tions elsewhere with deeper vessels in the 
region of the dorsal and pelvic fins. 

That the similarities between the venous 
systems of holocephalians and selachians 
do signify an evolutionary relationship is 
strongly suggested by the fact that their 
common pattern sets them off distinctly 
from the bony fishes. Neither ray-finned 
forms nor lungfishes show the development 
of spacious sinuses. Allen's (1905) excel- 
lent description and beautiful drawings of 
the circulatory system of the teleost, Ophio- 
don, reveal a complex arrangement of veins 
of small bore whose homology to vessels in 
cartilaginous fishes would be difficult to 
prove. In Ophiodon, fine facial vessels 
empty into a pair of jugular veins directly, 
since there are no orbital sinuses. The 
jugular veins, which carry blood posteriorly 

over the gill region, are thought by Van 
Gelderen (193S) not to be homologous to 
the anterior cardinals in the Chondrich- 
thyes. The posterior cardinal vessels of 
Ophiodon, like those of cartilaginous fishes, 
receive blood from the renal capillaries, 
but in other bony fishes there may be 
direct connections with the caudal vein 
which never persist in sharks or chimaerids. 
No actinopterygians that have been investi- 
gated have subcutaneous veins except the 
Thunnidae (and here the veins are ac- 
companied by arteries). As an adjunct to 
the venous system, teleosts have a well- 
developed set of lymphatics which all 
cartilaginous fishes (except possibly Tor- 
pedo) lack. A glance at the pattern of 
veins in dipnoans shows an even greater 
departure from the arrangement found in 
selachians and holocephalians, since there 
appear in the lungfishes vessels which are 
similar to those of amphibians. 

Despite the broad similarities between 
the selachian and holocephalian venous 
systems there are points of apparent dif- 
ference whose significance must be con- 
sidered. A dissection of the hypobranchial 
region, for instance, does not reveal a pair 
of easily recognizable, shark-like inferior 
jugular veins. The area is laterally com- 
pressed and occupied by the massively 
developed hypobranchial muscles whose 
arrangement leaves no straight channel for 
these veins to follow. Beneath the muscles, 
and ventral to the ventral aorta, there exists 
a mass of loose connective tissue which 
probably does have fine veins draining 
l^lood posteriorly toward the sinus venosus. 
Allis, who made a preliminary sketch of 
this area in 1916,- drew a pair of veins here 
which he interpreted as inferior jugulars. 
Not finding a hyoid sinus in the usual loca- 
tion, he ga\e that name to a large vein 
which begins behind the mandible and 

- This and other unpubHshed sketches of the 
anatomy of Chimaera coUiei were given by the son 
of the hite E. P. AHis to Dr. A. S. Romer and are 
in his hbrary in the Museum of Comparative Zool- 
ogy at Harvard University. 

158 Bulletin Museitm of Comparative Zoology, Vol. 135, No. 3 

curves dorsally to enter the back corner of 
the anterior cardinal sinus. It seems more 
Hkely, ho\\'e\'er, that any veins lying deep 
and near the midline represent auxiliary 
drainage and that the more lateral, dorsally 
cur\ing vessel is the inferior jugular vein of 
CJumoera. Although the curious position of 
this vein suggests that it might be a dif- 
ferent vessel from that of sharks, its rela- 
tion to the base of the gill arches is quite 
similar and its entry point into the anterior 
cardinal sinus near the union with the com- 
mon cardinal is not very different from 
that shown by Parker ( 1886 ) for Mustchis. 
It seems that the inferior jugular vein has 
been shifted dorsally as the head became 
laterally compressed, and is not so different 
after all from that of sharks. It contrasts 
markedly with the inferior jugular of tele- 
osts which is squeezed medially and in 
some forms, like Ophiodon, exists as a 
single median ventral vessel for most of 
its length. 

The absence of a hyoid sinus in Chimacra 
may also be due to the relative reduction 
of the branchial apparatus. In contrast to 
the large channel which connects the an- 
terior cardinal sinus with the inferior jug- 
ular vein in selachians, no major vein 
appears on either side of the ceratohyal 
cartilage. Examination with a dissecting 
scope showed in two specimens a fine 
vessel which lay anterior to the afferent 
branchial artery, but the vein was more 
easily traced along a pathway with the 
artery out upon the opercular flap than to 
a connection with the postorbital or an- 
terior cardinal channels. If it is correct 
to assume that main vessels develop in the 
embryo as the result of dominance of par- 
ticular pathways through the initial capil- 
lary net, then it is not surprising to find 
that chimaerids lack a shark-like hyoid 
sinus. In sharks, where the epihyal cartilage 
enlarges and develops a close association 
with the cranium, it is possible that a vein 
would appear behind it with a connection 
to the lateral head vein above. In Chi- 
maera, where the epihyal never departs 

from its serial alignment with other parts 
of the visceral skeleton, the absence of a 
special hyoidean vein or sinus might be 
expected. If one follows this inteipretation 
and accepts this assumption that the non- 
suspensory hyoid is primitive, it follows 
logically that the absence of a hyoid sinus 
may represent an original character rather 
than a secondary loss. If the holocephalians 
sprang from an ancestral stock which 
lacked a suspensory hyoid and a distinctive 
hyoid vein and then evolved the laterally 
compressed, short head of the extant ani- 
mals, neither the space nor the stimulus 
for the development of a hyoid sinus would 
have existed. 

In pointing out singularities in the pre- 
cardiac drainage of Chimacra, one should 
not omit mention of the situation of the 
anterior cardinal sinus. Although it has 
generally the same location as the sinus 
in selachians, its route does show one varia- 
tion: the vein passes lateral to a muscle 
which Vetter ( 1878 ) calls the trapezius 
intenius. In sharks, the sinus lies medial 
to the entire levator (trapezius) series. 
Again, this difference could arise from a 
shift of the musculature rather than the 
development of a new vein, but it is also 
possible that there has been emphasis upon 
an alternate embryonic blood pathway in 
Chimacra. Leaving to one side the possi- 
bility of a mistake in the identification of 
the muscle (the fibers run posteriorly from 
the subocular shelf to insert upon the an- 
terior edge of the scapula just beyond the 
last pharyngobranchials), it seems that 
either a shift in the arrangement of the 
muscles or the vein must be admitted. How 
great a significance should be attached to 
such an alteration remains an open question. 

In the postcardiac part of the venous 
system of Chimacra there are differences 
from the selachian pattern that are more 
clear-cut if not easier to inteipret. The 
lateral abdominal veins are absent, and 
the blood which they would have collected 
is differently distributed. From the pelvic 
fins and also from a rectal capillary net- 

Morphology and Relationships of Holocephali • Stahl 159 

work it is sent directly into the posterior 
end of the posterior cardinal sinus. A small 
part of the returning blood may even find 
its way to the renal portal by a fine con- 
nection from the femoral vein. Further 
forward, the blood from the deep part of 
the ventral body wall is collected by the 
ventro-posterior parietal vein which flows 
posteriorly to a confluence with the renal 
portal. Finally, the blood from the pectoral 
fins, instead of joining the flow from a 
lateral abdominal, enters the common car- 
dinal vein alone. The drainage from the 
posterior half of the pectoral fin enters 
what may be more exactly described as 
a brachial sinus than a brachial vein at the 
base of the fin and is joined there by the 
blood from the anterior half of the fin 
which returns by a vein that passes through 
a channel in the pectoral girdle. There are 
two possible explanations for the derivation 
of this peculiar venous pattern. The first, 
which cannot be flatly dismissed, is that 
the ancestral stock of the Holocephali pos- 
sessed this arrangement of vessels. The 
second and perhaps more probable sug- 
gestion is that the lack of lateral abdominals 
is secondary in chimaerids as it seems to be 
in the teleost fishes. The lateral abdominals 
and their homologues, the ventral abdom- 
inal and the umbilical veins, play too large 
a part in vertebrate development to allow 
one to dispose of them lightly. Their dis- 
appearance might be imagined to have 
required the longest possible evolutionary 
course. That these veins have not been 
described in cyclostomes complicates rather 
than solves the problem. If their absence 
represents a secondary loss, one has still 
no clue to the reason for their absence in 
holocephalians. If, on the other hand, the 
lack of lateral abdominals is a primitive 
vertebrate characteristic, which holocepha- 
lians are presumed to have retained, one 
must then question the homology between 
the lateral abdominals which selachians 
have developed and the ventral abdominal 
vein which appears in the first tetrapods. 
The most distinctive feature of the ve- 

nous drainage which might merit the same 
interpretation is the lack of hepatic sinuses 
opening into the sinus venosus. The an- 
terior portion of the liver, just behind the 
transverse septum, is very thin, contains no 
sinuses, and seems to send no blood forward 
into the heart. The back wall of the sinus 
venosus, on the other hand, seems entire, 
but may have vestiges of hepatic openings. 
Since no injection material can be made 
to pass through, one might suppose that 
no passages exist. Slight indentations are 
visible, however, in a likely location, and 
a needle can be made to pass, after some 
probing, without seeming to pierce tissue. 
If traces of old entrances into the sinus 
venosus are present, they prove that the 
lack of hepatic sinuses in the usual location 
is secondary. Even if it is not certain that 
such openings are there, it would seem from 
the design of the hepatic drainage that a 
secondary arrangement has arisen. The 
liver is bound to the underside of the pos- 
terior cardinal sinus, in its forward portion, 
and sends to that channel one main vein, 
two or three other large ones, and several 
minor auxiliary vessels. In contrast to this 
pattern, the forward flow from liver to 
heart is characteristic of every other jawed 
vertebrate, embryo and adult. In the most 
deviant pattern, the hepatic sinuses of rays 
open into the common cardinal veins rather 
than into the more medial sinus venosus. 
If the holocephahan pattern were to be 
judged primitive, it would have to be sup- 
posed that it was the sole remaining ex- 
ample of a distinctive circulatory arrange- 
ment which existed in ancient times among 
ancestral vertebrates — a not too likely pos- 
sibility. As in the case of the absence of 
the lateral abdominals, it seems more sen- 
sible to suggest that the liver drainage rep- 
resents a great deviation from the usual 
vertebrate condition and may well have 
been the product of a long independent 

The search for differences between se- 
lachians and holocephalians should be ex- 
tended to the hepatic portal system, too, 


BiiUetiii Museum of Comporative Zoology, Vol. 135, No. 3 

because the homologies between these ves- 
sels of sharks and chimaerids are not at 
once apparent. Upon close scrutiny it seems 
that the chimaerid veins are more simpli- 
fied and abbreviated than those of sharks. 
All of the gastric veins are absent, of course, 
as are those which drain the spleen and run 
through mesentery. A short lienopancreatic 
vein and some small splenics join the vessel 
returning blood from the posterior regions 
of the intestine. Consequently, only one 
vein runs forward to join the intra-intestinal. 
The union of these two vessels and a smaller 
one from the anterior intestine and neigh- 
boring esophagus creates the hepatic portal. 
Although it is probable that the lack of 
mesenteries and the removal of the spleen 
from its usual place are secondary changes, 
it is not necessary to assume that the entire 
pattern of portal tributaries has been only 
recently evolved. The existence of two 
sizable branches draining the posterior in- 
testine, an arrangement which has no coun- 
terpart in sharks, may not have arisen from 
any selachian forerunner. It would be 
tempting to add the lack of gastric veins 
as another possible preselachian character, 
because the absence of a stomach has been 
supposed to be a primitive arrangement; 
but it cannot be assumed definitely that 
the lack of a stomach and the minimal 
development of the spiral valve are prim- 
itive rather than degenerate developments. 
The hepatic portal system, one must con- 
clude, does bear a resemblance to the 
selachian pattern but may be somewhat 
secondarily modified. Despite resemblances, 
on the one hand, and late modifications, on 
the other, however, it is not impossible that 
there might be some elements of an older 
independent pattern still included in the 

The association of lymphomyeloid tissue 
with the venous system in the Holocephali 
is another characteristic which distinguishes 
these fishes from the shark group. Kolmer 
( 1923), who studied this tissue in Chimaera, describes it as consisting of a 
mass of lymphatic cells of varying sizes 

mixed with red blood cells in all stages of 
formation. These cells are supported by 
a network of fine connective tissue fibers 
which merge with the adventitia of the 
wide veins and small arteries that ramify 
within the mass. If the tissue found in the 
esophageal wall of sharks and rays is com- 
parable to that which exists in a much 
expanded state in the chimaerids, one might 
predicate the presence of this material in 
the early placoderms. It may have been 
carried in several lines as a hemopoietic 
organ, its different location and extent in 
holocephalians and selachians indicating 
separate evolutionary pathways. It is 
harder to imagine that this tissue, which 
is Vvddespread in sharks and batoids in its 
esophageal location, should disappear from 
that place and appear in the head and 
girdle regions of the Holocephali as they 
branched off from a shark stem. It is also 
possible that the tissue of the two groups, 
although it looks similar under the micro- 
scope, may not have a common origin. If 
that be the case, the hypothesis that chi- 
maerids have been derived from early 
sharks would have another point against it. 
A study of the remaining portions of the 
circulatory system reinforces the idea that 
holocephalians and sharks probably arose 
from placodenn stocks possessing some 
characters which both groups of fishes have 
carried to the present day. One has only to 
dissect the heart in each animal ( cf., for 
example, Lankester, 1878, and Hyman, 
1942: 329) to be convinced that chimaerids 
and sharks, while distinct lines, cannot be 
widely separated from each other on the 
evolutionary tree. The hearts of the two 
types of fishes are identical in their gross 
anatomy, and markedly distinct from the 
heart of lungfishes or that of ray-finned 
forms. "^ The only characteristic which dis- 
tinguishes the chimaerid heart from the 
shark structure is its relatively small size 

^ Lankester (1878), besides describinn; the heart 
of Chiiiiacni, makes a visil)ly futile attempt to point 
out homoloj^ies between its arrangement of valves 
and tliat in the heart of dipnoans. 

Morphology and Relationships of Holocephali • Stahl 161 

(Fig. 4B). Although no measurements 
were made, it seems that the heart of 
Chimaera would have a smaller capacity 
than that of a shark of the same size. If 
one wishes to suppose that sharks and 
holocephalians have long traveled upon 
separate evolutionary paths, one must pos- 
tulate that their type of heart represents 
the primitive gnathostome structure which 
has been retained in all cartilaginous fishes 
and highly modified in bony ones. 

The one salient difference in the arterial 
pattern occurs in the head region. The 
Holocephali show a type of blood supply 
to the brain which differs from both 
selachians and bony fishes in that the 
pseudobranchial efferent alone reaches the 
cranial cavity (AlHs, 1912). The hyoidean 
efferent, which feeds the internal carotid 
in elasmobranchs, forms in holocephalians 
only a commissure with no continuation 
running forward to the brain. Even if the 
investigator assumes that there were in the 
primitive state two pairs of efferent arteries 
which sent blood forward to the brain, and 
that in the Holocephali the more posterior 
pair has degenerated, he has not disposed 
of the entire problem posed by the cephalic 
arteries of the Holocephali. There is also 
an unusual mandibular artery for which to 
account. In sharks the lower jaw is sup- 
plied by a vessel which leaves the ventral 
end of the first efferent arterial loop ( Hy- 
man, 1942: 324); in holocephalians the 
mandibular artery runs ventrally from the 
efferent pseudobranchial, itself a more dor- 
sal branch from the loop. The suggestion 
has been made by Allis ( 1912 ) that the 
holocephalian vessel, which follows the line 
of the jaw, represents the \entral portion 
of the afferent mandibular artery and that 
the retention of this vessel, which has gen- 
erally disappeared in other vertebrates, is 
a primitive feature. If this supposed homol- 
ogy is correct, it would not be possible to 
derive the chimaerids from a shark group 
in which the mandibular afferent had al- 
ready disappeared. However, Marples' 
( 1936 ) discovery of a similar mandibular 

artery in Sqiiatina, and his statement of 
the existence of the same type of vessel 
in Polyodon and certain teleosts, makes 
questionable Allis' interpretation and any 
evolutionary theories which might be based 
upon it. 

The only other portion of the arterial 
system which deviates from the selachian 
pattern is the efferent branchial series of 
vessels. In contrast to the distinct loops 
created in selachians (Hyman, 1942: 324) 
by the union of well-formed pre- and post- 
trematic arteries, the poor development of 
the chimaerid pretrematic branches creates 
discontinuities in the posterior three collec- 
tor loops. Allis ( 1912) believed that he saw 
four complete loops, but Allen ( 1905 ) and 
Parker ( 1886 ) failed to find any. Without 
doubt, the pretrematic vessels seem second- 
ary to the posttrematic ones. In Chimaera 
coUiei, the ventral ends of the pretrematics 
diminish in size and the distinct ventral 
commissures which close the loops were 
visible only in well-injected specimens. 
Again, two possible explanations present 
themselves: either the chimaerid situation 
represents an early step toward loop-forma- 
tion or it is the result of a modification 
associated with the reduction of the gill 
arch apparatus and its concealment beneath 
the operculum. There is no way of deciding 
which theory is more probable. 

In conclusion, then, one recognizes that 
the holocephalian circulatory system resem- 
bles that of selachians in numerous ways 
and vet differs from it distinctlv in certain 
characteristics. In some of its nonselachian 
features the chimaerid system parallels the 
structure of bony fish while in others it 
is apparently unique. 

Among the similarities to selachians can 
be listed the structure of the heart, the 
general design of the arterial system, the 
presence in the venous system of great 
sinuses, and the arrangement of the prin- 
cipal venous sinuses. Pursuing further sim- 
ilarities in the venous system, one must 
mention the existence of a pair of inferior 
jugular \essels which are more like the 


Bulletin Muscuin of Comparai'wc Zoology. Vol. 135, No. 3 

inferior jugulars of sharks than hke those 
of other fishes, of similar renal portal sys- 
tems, and of a hepatic portal system that 
is certainly closer to the type of sNsteni 
found in selachians than to that found in 
bony fish. Like the selachians, the holo- 
cephalians have a system of cutaneous veins 
and lack lymphatics. As a last point of sim- 
ilarity, even the valves in the chief vessels 
seem to be located at the same places. 

Characters which distinguish the holo- 
cephalian circulatory system from that of 
sharks are found in both the arterial and 
venous pathways. It is perhaps significant 
that features of the arterial system which 
are unique to the Holocephali are all found 
in the head, a region which is as a whole 
very highly specialized. Although the in- 
complete collector loops and the absence 
of the anterior extension of the internal 
carotid arteries can be viewed as relatively 
small modifications of the selachian plan, 
the significance of the branching of the 
mandibular artery from the afferent pseudo- 
branchial is not so easy to interpret. 

In the venous system, the absence of 
lateral abdominal veins represents a great 
divergence from the selachian pattern and 
a resemblance to that of ray-finned fishes. 
With those vessels missing, the brachial 
veins empty into the common cardinals 
directly, as is the case in teleosts, and the 
femoral veins open into the posterior car- 
dinal sinuses. The entrance into these sinuses 
of blood from the liver is surely a peculiar- 
ity developed in the holocephalian line, as 
such a route is unheard of in any other 
group of fishes. Whether the absence of 
a hyoid sinus is also to be interpreted as a 
secondary development or as a primary 
arrangement is not clear. The resolution of 
this question may depend upon the solu- 
tion of the problem of the status of the 
hyoid arch with which the hyoid sinus is 

In surveying the circulatory system of the 
Holocephali, one receives the impression 
that the resemblance to the bony fish, 
which exists chieflv in the absence of the 

lateral abdominal veins, is fortuitous rather 
than representative of ancestral connec- 
tions. Since the resemblance to the circula- 
tory system of sharks is more general, the 
question seems to be whether the holo- 
cephalian system is a derivative of the 
selachian one or whether it has developed 
in its own path from a system which char- 
acterized a placodenn group ancestral to 
both holocephalians and selachians. An 
opinion as to which of these alternatives 
is the more probable could be more strongly 
supported if the structural arrangements of 
other systems were brought into evidence. 
Should they be found to contain characters 
too primitive to have been derived from 
the more specialized homologues of sharks, 
resolution in favor of descent from separate 
placoderm ancestors would be indicated. If 
the other systems seem entirely shark-like 
or differ from the shark pattern in minor 
ways only, a direct descent from some shark 
group cannot be ruled out. To extend the 
comparison between holocephalians and 
selachians beyond the confines of the cir- 
culatory system, then, a review of the 
nervous, skeletal, muscular, urogenital, and 
digestive systems is undertaken in the fol- 
lowing pages. The study of these systems 
also provides a check upon the assumption 
made here that, although similarities be- 
tween structures of holocephalians and 
actinopterygians do appear, there is little 
likelihood of an ancestral affiliation be- 
tween the two groups. 


The Nervous System 

In any study in which the relationships 
of the Holocephali are reviewed, the ner- 
vous system receives primary attention. Its 
gross anatomy has been studied in detail 
(Braus, 1898; Cole, 1896; Carman, 1904; 
Fiirbringer, 1897; Nicol, 1950; Wilder, 
1877), and microscopic work has been 
attempted in some areas (Kappers, 1911, 
1912; Backstrom, 1924; Johnston. 1910; 

Morphology and Relationships of Holocephali • Stahl 163 

Nicol, 1950). From the evidence presented 
in these papers, it seems that the holo- 
cephahan ner\ous system retains some char- 
acteristics that must surely he primitive hut 
exhihits pecuHarities which are generally 
interpreted as secondary phenomena. 

Although the form of the spinal cord is 
quite regularly shark-like, the hrain is of an 
extraordinary nature. In its posterior por- 
tion it resemhles closely the selachian organ, 
but the telencephalon is unique in the ani- 
mal kingdom. In Chimacm and CoUorliyn- 
chus\ this anterior region of the brain 
extends forward as a long stalk beneath the 
interorbital septum. It widens finally into 
a subdivided telencephalic lobe just behind 
the olfactory bulbs associated with the nos- 
trils. The uniqueness of the arrangement 
argues for its secondary development. Kap- 
pers and Carpentier (1911) have considered 
the elongation of the telencephalon and 
feel that it has taken place as a corollary 
to the enlargement of the eyes. Since the 
eyes have encroached upon the medial 
region normally occupied by the cerebral 
hemispheres, the forepart of the brain has 
been displaced. Rather than being com- 
pressed posteriorly, in the ancestors of the 
Holocephali, alone among all the animals 
that have developed enormous eyes, the 
telencephalon became displaced anteriorly. 
The result of the forsvard gro\\'th of the 
telencephalon has been the creation of long 
brain stalks through which regular connec- 
tions with the diencephalon are maintained. 
RJiinochimaera, in which the eyes are 
smaller, the brainstalks not quite so elon- 
gated, and the olfactory tracts more sela- 
chian-like, may represent an earlier stage 
in the evolution of this curious arrangement. 

Another characteristic of the telencepha- 
lon which lends itself to comment of a 
phylogenetic nature is the development of 
the pallium. Holmgren, who has studied 
forebrain moq^hology in lower vertebrates 
(1922), has investigated the pahial region 
in holocephalians, selachians, ganoids, tele- 
osts, and lungfish. He points out that the 
selachian pallium is inverted to a greater 

/. supopK. n. opt. I. cbl. 

3T.r\. I I / 

med obi. 

Fig. 1. Chimaera coliiei. Brain, lateral view, cbl., Cere- 
bellum; eth.c, ethmoid canal; hyp., hypophysis; med.obl., 
medulla oblongata; oll.b., olfactory bulb;, 
ophthalmicus profundus nerve; opt. I., optic lobe; sup.oph.n., 
superficial ophthalmic nerve; tel., telencephalon;, 
telencephalic stalk; //, optic nerve. (After Gorman.) 

degree than is the case in the other fishes. 
By inversion he means a rolling medially 
of both left and right edges of the embry- 
onic neural i^lates, resulting in their contact 
dorsally if the two masses of tissue reach 
the midline as they do in selachians. E vagi- 
nation of the more lateral portions of the 
developing forebrain wall gives rise to 
paired cerebral hemispheres. If nerve cells 
mass dorsally over the ependymal layer, the 
dorsal brain wall thickens and the dividing 
furrow between the hemispheres may be 
more or less obliterated. The developmental 
mode which occurs in the Holocephali, how- 
ever, consists of a lateral rather than a dor- 
sal concentration of nerve cells. The brain 
roof is then left relatively thin. In extreme 
manifestations of this latter tendency, the 
lateral brain walls grow exceedingly thick 
and may actually evert, leaving the roof to 
be covered by an expanded tela. This is 
the case in actinopterygians. A less extreme 
and perhaps more primitive version of the 
same condition is found in the lungfish 
Cemtodus (Holmgren and Horst, 1925). 
The dipnoan has a broad thin roof over the 
pallium but the moderately thick cerebral 
walls are not everted. 

In holocephalians the laterally thickened 
pallium is inverted but never becomes con- 
tinuous over the dorsal midline. There is 
always a small strip of ependyma bridging 

164 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 


qpc he 


Fig. 2. Transverse section of the forebrain in A, Acanthias, B, Chimaera, C, Protopferus, D, Lepidosfeus, to show position 
of the palhum. epnd., Ependyma; g.p.c, general pallial cortex; /i.e., hippocampal lobe; 1. 1. p., lateral limit of pallium; 
n.olf.l., nucleus olfactorius lateralis; pal., pollium; p.c, pyriform cortex; sub. p., subpallium. (After Holmgren.) 

the dorsal gap. The palHum is evaginated 
to form two separate olfactory lobes which 
carry separate ventricles forward of the 
foramen of Munro. The pallium is rather 
small, being confined to the anterior, en- 
larged portion of the telencephalon. The 
brainstalks which connect the anterior en- 
largement with the remainder of the brain 
are composed entirely of subpallial tissue. 

Kappers, who reviewed holocephalian 
brain structure in his compendium on the 
nervous system of vertebrates ( 1936 ) , 
grouped the Holocephali with the lower 
actinopterygian fishes as intermediate be- 
tween selachians and teleosts. He regarded 
the inversion of the pallium as carried over 
from the former and the eversion of the 
brainstalk walls as presaging the great pal- 
lial eversion of the latter. Holmgren dis- 
agrees with the conclusion of Kappers, 
however. He feels that eversion of the sub- 
pallial tissue of which the brainstalks con- 
sist cannot be regarded as an early stage of 
the pallial eversion seen in bony fish. In 
making his interpretation of the phylo- 
genetic position of the Holocephali, Holm- 
gren considers only the true pallium whose 
limits he has determined by histological 
study. He reasons that the holocephalian 
pallium resembles most nearly, in its degree 
of inversion and evagination, what must 
have been the type ancestral to that of ex- 
tant cartilaginous and bony fishes. The 
development of greater inversion with re- 
sulting fusion across the dorsal midline 
would lead to the selachian condition, 
whereas the development of thicker and 

more widely separated walls would lead to 
the lungfish-lower actinopterygian-teleost 

Observations upon the microscopic struc- 
ture of the holocephalian brain have been 
more fragmentary, and no clear-cut indica- 
tions of phylogenetic position arise from 
them. Kappers (1912) has mapped the ar- 
rangement of the motor nuclei in Chimaera 
monsfrosa and he and several other workers 
have determined the course of some of the 
brain tracts in the chimaeroids ( Kappers, 
1911; Biickstrom, 1924; Johnston, 1910). 
One example of the quandary to which these 
studies have led should suffice. The sela- 
chians, with which investigators have sought 
to compare the holocephalians, are charac- 
terized by three telencephalic tract decus- 
sations — one dorsal and two ventral. Since 
the left and right pallial masses of holo- 
cephalians do not fuse in the dorsal mid- 
line, the dorsal decussation is absent. It is 
not known whether the fibers which cross 
dorsally in selachians are channeled through 
the ventral commissures in holocephalians 
or whether these fibers are wholly or par- 
tially absent. In speaking of the ventral 
decussation, Biickstrom goes on to say, "It 
is, however, possible that a number of fibre 
connections in this decussatio existing in 
Chimaera are lacking in selachians or vice 
versa" (Biickstrom, 1924: 232). 

The arrangement of the cranial nerves 
has also been examined by a worker with 
the phylogenetic question in mind. Cole, 
who has dissected these nerves in Chimaera 
in detail (1896; Cole and Dakin, 1906), was 

Morphology and Relationships of Holocephali • Stohl 165 

especially interested by the emergence from were farthest removed from the beginning 

the brain and the distribution of nerves V, of the evolutionary line. He had detemiined 

VII, and X. He points out that there is no that the number of occipital nerves coming 

trigemino-facial complex in Chimaera as through foramina at the back of the skull 

there is in sharks. Nerve V emerges by two ranged from five in notidanid sharks to one 

roots anterior to \'II and underneath the or none in rays. At first, after finding five 

buccal branch of the latter. It sends sen- such nerves in Chimaera and four in Cal- 

sory and motor fibers to the usual destina- lorhynchus, Fiirbringer was ready to place 

tions without e\'er mingling with portions these fish on a level with the notidanids. 

of VII. Contact between these two cranial On closer observation, however, he noticed 

nerves is limited to a variable degree of that only the first two resembled the occipi- 

binding together of their superficial oph- tal nerves of the selachians in appearance 

thalmic branches. Nerve VII can be divided and in their course to the hypobranchial 

into a small motor portion and a larger muscles. The remainder looked very much 

lateral line component. Cole recognizes like the succeeding spinal nerves and, like 

that the isolation of the lateral line fibers the latter, sent fibers to the brachial plexus, 

from the rest of the cranial nerve is also Fiirbringer distinguished these nerves as 

characteristic of other fishes and of am- spino-occipital nerves, explaining that in 

phibians which ha\e a lateral line system the Holocephali two or three of the anterior 

and so has no special significance. The vertebrae have been incorporated into the 

tenth cranial nerve of Chimaera is distinc- skull bringing their segmental nerves with 

tive, though, in having its four parts in addi- them. Thus these spino-occipital nerves are 

tion to the laterahs component ( three bran- not homologous to the posterior occipital 

chial branches and one visceral) completely nerves of simple selachians but are proof 

separate: each arises separately from the that the holocephalians belong to a "hohere, 

brain and each has its own ganglion. Cole mehr specialisirte Abtheilung" ( Fiirbringer, 

dwelt upon the evidence of the primitive 1897: 446). 

position of Chimaera, which the separation The true spinal nerves of holocephalians 
of the posterior cranial nerves suggests, and bear a greater resemblance to those of 
concluded: "The discrete nature of the fifth, selachians than to those of bony fishes in 
seventh, and lateral line nerves makes Chi- that the dorsal and ventral roots retain the 
maera a very unique fish as regards its large degree of independence which is 
cranial nerves, and it is to be presumed that characteristic of the former group. In Chi- 
such a simple condition is more primitive maera, one can see in each segment of the 
than the more complex fusions and inter- trunk two roots emerge, give off dorsal 
minglings that obtain in other fishes. This rami, and then, as ventral rami, gradually 
separation may, ho\\'e\'er, be purely second- come together. The segmental nerve formed 
ary, just as the form of the brain of Chi- by their union soon divides, and the nerve 
maera undoubtedly is, but on the other continues its lateral course as a double- 
hand the vagus is also in a very simple and stranded structure. If the two strands rep- 
unfused condition in Chimaera, and the resent the reseparation of dorsal and ven- 
same may be said of its cranial nerves gen- tral root fibers, the holocephalians would 
erally" (Cole and Dakin, 1906: 599). then show a very limited association of dor- 
While Cole was upholding the primitive sal and ventral root elements— an arrange- 
position of Chimaera suggested by the ar- ment seemingly closer to the primitive state 
rangement of its cranial nerves, Fiirbringer of complete separation than that shown 
(1897) was concluding from his compara- even by selachians. At the level of the pel- 
tive study of the occipital nerves of sela- vie fin, Davidoffs dissections (1879) show 
chians and holocephalians that the latter a separation of the strands of the spinal 

166 Bulletin Miisetim of Compnraiwe Zoology, Vol. 135. No. 3 

nerves followed distally by a recombination ganglia in the tail ) ; but these ganglia are 

involving the posterior strand in one seg- haphazardly connected by a network of 

ment and the anterior strand from the seg- nerve fibers and communicate with the 

ment behind. In Davidoff's opinion, this spinal nerves by white rami only. Since 

arrangement as well as further connections there are minor differences between the 

lietween the first two nerves which supply systems of selachians and Chimaera, Nicol 

the fin suggest the beginnings of a more is of the opinion that the chimaeroids split 

complex peKic plexus than is present in from the selachian line and have evolved 

other fishes. The holocephalians' lack of a in the final stages on a separate path, 

collector nerve in the pelvic region, accord- A study of the sense organs produces little 

ing to Van der Horst {in Bolk, Vol. II, evidence which can be brought to bear 

1934), also sets them apart from selachians, upon the phylogenetic question. The olfac- 

dipnoans, and lower actinopterygians. The tory, optic, and otic structures in holoce- 

pU'xus at the level of the pectoral fin, how- phalians are similar to those of sharks, and 

e\er, resembles that of selachians and there are no fossil remains of the first two 

actinopterygians in being of a cervico- types of sense receptors from which their 

brachial nature. The holocephalians and hereditary history could be learned. The 

these fishes are distinguished in this feature design of the inner ear in early vertebrates 

from the dipnoans which have, like tetra- has been revealed through cranial casts, 

pods, two separate plexuses in this region, however, making comparisons possible. 

Assessing the various characteristics of the Stensio's ( 1963 ) cast of the cavities in the 

spinal nerves and the plexuses in which cranium of the arthrodire Kujdanowiaspis 

they are involved, one might conclude that shows a general arrangement of the laby- 

the holocephalians may show the retention rinth which still characterizes both holo- 

of a relatively primitive arrangement which cephalian and selachian fishes. Even the 

has been modified to form a unique pattern endolymphatic duct appears, rising to open 

in the pelvic region. upon the dorsal surface of the head. The 

The last remaining part of the nervous only point of difference between chimaerids 

system to be discussed, the autonomic divi- and sharks, of which Stensio speaks, con- 

sion, has been described thoroughly by cerns the structure of the utriculus. That 

Nicol (1950). His study, it must be men- of selachians is divided, while the utriculus 

tioned, was based only upon Chimaera col- of holocephalians is not. Stensio thinks that 

liei. Without attempting to repeat Nicol's the undivided state is more primitive and 

description, one may say that he found a that the divided utriculus has appeared in 

very close similarity between the autonomic certain orders of arthrodires and in elasmo- 

systems of selachians and holocephalians branchs through parallel development. If 

and substantial contrasts between their type Stensio's speculation is correct, the ances- 

of system and that of bony fishes. For ex- tors of holocephalians and early sharks 

ample, he notes that teleosts have well- would have been separate but related 

ordered sympathetic chains connected to stocks. 

the spinal nerves by Ijoth gray and white Both Stensio (1947) and Holmgren 

rami. Although the dipnoans show a less (1942a) have included a study of the lateral 

well-developed pair of ganglionated chains, line system of the Holocephali in their sur- 

the presence of a delicate chain-structure veys of lateral line systems in fishes and 

differentiates even these fishes from the amphibians. Although these authors dis- 

selachians and from Chimaera. In the car- agree as to whether a general pattern of 

tilaginous forms there is a more or less seg- head canals can be defined, they state in 

mental arrangement of sympathetic ganglia concert that no explanation of the evolution 

throughout the trunk (and an absence of of the holocephalian pattern is possible at 

Morphology and Relationships of Holocephali • Stahl 167 

tliis time. Holmgren finds it improliable 
tliat the holocephalian arrangement could 
be derived from that of selachians or vice 
versa. He suspects that the holocephalian 
system has been reduced from a more 
elaborate pattern although the absence of 
embryological studies prevents his specu- 
lating upon what the ancestral state might 
have been. He is forced to abandon the 
problem with the statement that the holo- 
cephalian head canals "could not be identi- 
fied with lines in any other vertebrate" 
(Holmgren, 1942a: 21). Stensio would have 
liked to have made a comparison between 
the head canals of holocephalians and those 
of ptyctodonts, but unfortunately the latter 
have not been preserved. The anatomy of 
the sensory canals of holocephalians reveals 
no more to the investigator than their ar- 
rangement. As Garman ( 1888, 1904 ) and 
Reese (1910) have shown, the sensory cells 
lie in open grooves in Chimaera, in slit 
tubules in Rhinodiimaera, and in closed 
tubules in CaUorhynchiis. It is not possible 
to detennine which of these arrangements 
is primitive or if any one of them is. 

Although no single characteristic of the 
nervous system serves as a key to the evolu- 
tionary history of the Holocephali, it is pos- 
sible to make a reasonable speculation based 
upon the group of anatomical features dis- 
cussed above. The survey of the holoce- 
phalian nervous system has shown that no 
portion of it resembles that of any bony 
fish. Although the anterior extension of the 
telencephalon, the ramifications of the 
spinal nerves, and the pattern of the sen- 
sory canals are unique, the posterior parts 
of the brain, the sensory organs, and the 
autonomic nervous system are strikingly 
like the shark structures. In drawing con- 
clusions based upon the nature of the ner- 
vous system, then, one must lay aside the 
possibility that the Holocephali might be 
allied to any line leading to bony fishes 
(despite Holmgren's view that the dipnoan 
pallium might be derived from the holo- 
cephalian type) and predicate, instead, 
some degree of relationship to the early 

sharks or their ancestors. If one agrees with 
the assumption that the partially inverted 
pallium is more primitive than the strongly 
inverted selachian structure and also \\'ith 
the interpretation of the cranial nerve ar- 
rangement as more primitive than that 
found in sharks, it follows that the Holo- 
cephali could not have evolved from early 
sharks in which the more complex selachian 
organization was already established. One 
is left with the hypothesis that the Holo- 
cephali have descended from an ancestral 
group separate from that of selachians but 
allied to it. This hypothesis allows, firstly, 
for the retention in holocephalians of the 
structures assumed to be primitive even 
though these elements are modified in 
sharks. Secondly, it provides an explana- 
tion for the presence of similar nervous 
structures in both types of cartilaginous 
fishes, since these elements may have been 
characteristic of the larger group to which 
both ancestral stocks belonged. Thirdly, the 
hypothesis suggests that the holocephalians 
developed along a separate line long enough 
to permit the evolution of the special struc- 
tures which are unique to them. The alter- 
native theory, that holocephalians are de- 
scended from an early shark group, \\'ith its 
corollary that the structures of the nervous 
system must all be derived by modification 
of the selachian plan, seems less likely than 
the above hypothesis which predicates no 
such close relationship between the two 
extant groups. 

The Skeletal System 

Extant holocephalians, like selachians, 
have a skeleton constructed entirely of car- 
tilage, their only hard parts being isolated 
placoid scales and the large dorsal fin spine. 
In the Jurassic forms Squaloraja and Myria- 
canthus, ho\\'ever, the dermal elements are 
more extensive. The fact that there is a 
greater amoimt of hard tissue in extinct 
holocephalians than there is in modern ones 
gives added support to the idea that the 
cartilaginous skeleton characteristic of the 
Chondrichthyes is not the primitive verte- 


BuUetin Museum of Comparative Zoo/ogr/, Vol. 135, No. 3 

brate framework but a secondary develop- 
ment. Although this hypothesis does not 
disallow the possibility that the Holocephali 
split from the selachian line after the dis- 
appearance of bone, it invites one to specu- 
late that the reduction to cartilage may have 
occurred in the two lines separately. 

Comparative studies of the skeleton pro- 
vide some evidence which can be used in 
trying to determine where the holoce- 
phalian and selachian fishes diverged, but 
one feels the lack of sufficient fossil data at 
every turn. Fossils of early sharks are not 
abundant and among those which have 
l)een studied, there is not one whose char- 
acteristics suggest that it might have served 
as an ancestor for the holocephalian line. 
Moy-Thomas ( 1936 ) has offered the coch- 
liodont, Helodiis simplex, as an ancestral 
type, but the bradyodonts are themselves 
distant from selachians. Watson (1938) and 
0rvig ( 1962 ) have both suggested that the 
fiolocephali have been derived from pty- 
ctodonts, and thus they take the stand that 
holocephalians have never shared the sela- 
chians' evolutionary pathway. A review of 
the holocephalian skeleton can at best, then, 
only attempt to define the degree of simi- 
larity between it and that of selachians and 
can try to determine whether a relationship 
to ptyctodonts or cochliodonts is possible 
only where comparable structures ha\'e been 

A review of the studies of the skull shows 
that relatively few workers have tackled the 
head skeleton in its entirety. Only Allis 
(1917, 1926), DeBeer and Moy-Thomas 
(1935), and Holmgren (1942b), have 
looked much beyond the labial cartilages. 
In making their more inclusive studies, they 
complain of the lack of data concerning em- 
bryonic development: literally nothing 
exists except the examination of a few 
embryos by Dean ( 1906) and Schauinsland 
( 1903). It is a pity that the breeding places 
of these laboratory-shy fishes are not well- 
known, for a careful review of a series of 
embryos from the earliest stages would go 

far toward settling some of the questions 
which Allis, DeBeer, and Holmgren raise. 

The first of these questions concerns the 
developmental interrelationships of the eyes 
and the cranial cavity. As has been men- 
tioned before, Kappers felt that the depres- 
sion of the telencephalic space occurred be- 
cause of the dorsomedial expansion of the 
eyes. With this conclusion Holmgren would 
agree. Holmgren surmises from this point 
that the ancestors of the Holocephali must 
ha\'e been slightly flattened fonns with 
rather dorsally placed eyes. Otherwise, 
Holmgren reasons, it would not be likely 
that expansion of the orbits would force 
the brain downward. It follows, in Holm- 
gren's thinking, that even a more broad- 
headed cochliodont than Helodus would be 
a likely ancestor for the holocephalians. 

In speaking of the structure of the cranial 
cavity, both Holmgren and Allis take issue 
with the opinion of DeBeer, Moy-Thomas, 
and Watson. The latter workers believe 
that the cranial cavity does not include the 
passage known as the ethmoid canal, 
through which the superficial ophthalmic 
nerves run forward after leaving the orbits. 
DeBeer and Moy-Thomas ( 1935 ) state that 
this canal is roofed over by a dorsal exten- 
sion of the orbito-nasal lamina beyond the 
true cranial roof, and they present a series 
of drawings of hypothetical evolutionary 
stages from the uncovered to the covered 
condition of this supracranial space. In his 
pul)lication of 1936 in which he presents 
the case for the descent of the Holocephali 
from the cochliodont Helodus simplex, Moy- 
Thomas stresses the fact that Helodus al- 
ready shows a dorsally-flared orbito-nasal 

Allis and Holmgren both hold that since 
the ethmoid canal is continuous with the 
cranial space, it is, therefore, a part of it. 
Allis ( 1926) suggests that the cranial space 
anterior to the orbits was cut off indirectly 
through the pressure of a mysterious em- 
bryonic "vesicle' which appears between 
the midbrain and the forebrain. As the fore- 
brain is pressed downward, the trabeculae 

Morphology and Relationships of Holocephali • StaJiI 169 

are squeezed outward, eventually rising up 
and inward to cut the cranial cavity in two. 
According to Allis' theory, the trabeculae 
form the floor of the ethmoid canal and the 
roof of the telencephalic enclosure. The 
floor under the telencephalon is composed 
of intertrabecular tissue with perhaps a con- 
tribution from the fused palatoquadrate. 

Holmgren's interpretation is based more 
on anatomical examination and less upon 
flights of fancy. In his study of the heads 
of fishes (1942b), Holmgren presents photo- 
graphs of six transverse sections through 
the orbital region of Chimaera monstrosa 
in which he points out a fine channel, run- 
ning from the main cranial space forward 
over the interorbital septum to open into 
the ethmoid canal. He remarks that in 
Rhinochimaera, which seems to be the most 
primitive holocephalian, this channel is 
much wider, making even clearer the con- 
tinuity of the two spaces. To explain the 
presence of a floor to the ethmoid canal 
which divides it from the telencephalic 
space beneath, he suggests that this carti- 
lage may be a neomorph, citing its very 
late chondrification as shown by Schauins- 
land's study (1903) of a CallorJiyncJius em- 
bryo. He goes on to hypothesize that, as in 
some sharks, the superficial ophthalmic 
nerves of the ancestors of the Holocephali 
may ha\"e run in left and right preorbital 
canals \\'hose lateral \\'alls were formed by 
extensions of the supraorbital crest carti- 
lages. Just as these nerves of Pristiophonis 
come inside the cranial space intennittently 
because of deficiencies in the walls medial 
to the preorbital canals, so in the Holoceph- 
ali, through complete disappearance of the 
cranial walls in this area, the two preorbital 
canals may have merged with the anterior 
brain cavity. It is by such a change, accord- 
ing to Holmgren, that the superficial oph- 
thalmic nerves may have come to run \\'ithin 
what he considers to be the anterior part of 
the cranial cavity in the holocephalians. In 
disavowing the existence of an ethmoid ca- 
nal as a unique holocephalian character, 
Holmgren removes one of the structures 


Fig. 3. Rhinochimaera pacifica. Neurocranium, lateral view. 
Cranial cavity with brain outlined. anf.cer.v., Anterior 
cerebral vein; com.c, communicating channel between 
cranio! cavity and ethmoid canal; cr.cov., cranio! cavity; 
endl.d., endolymphatic duct; eth.c, ethmoid conol; int. orb. - 
sept., interorbital septum; oH.b., olfactory bulb;, 
olfactory tract; orb.o., orbital artery; sup.oph.n., super- 
ficial ophthalmic nerve; tel., telencephalon; //, optic nerve. 
(After Holmgren.) 

upon which DeBeer and Moy-Thomas 
leaned heavily in associating the Holoceph- 
ali with the cochliodonts. 

On the basis of what has been said about 
the interrelationship between the eyes and 
the cranial cavity, it becomes plausible to 
conclude that the chondrocranium probably 
surrounded a brain space of quite ordinary 
dimensions in the ancestors of the Holo- 
cephali and that the enlarging eyes press- 
ing an interorbital septum between them 
gradually reduced the median cavity to its 
present divided condition. If this reason- 
ing is correct, the Holocephali must have 
long been upon a separate evolutionary 
pathway, leading from a form like Wiino- 
chimoera to one like Callorhynchus and fi- 
nally to the chimaerids. This sequence of 
e\olution is supported by the fact that the 
interorbital septum in Chimaera is even 
more extensive than that of CaUorhijnchus 
( Hubrecht, 1877 ) . One would expect the 
area of the septum to be largest in the 
group which shows the greatest median 
expansion of the eyeball. 

170 BtiUetin Miiseinii of Comparotive Zoologtj, Vol. 135, No. 3 

The otic region of the holocephalian 
braincase is short, the ear capsule being 
pressed close against the back of the orbit. 
Many selachians also show a relatively 
short otic portion of the skull. If, however, 
the selachians are descended from primitive 
fomis with an elongate otic and occipital 
region as Romer believes (Romer, 1964) it 
becomes less probalile that the holoce- 
phalian fishes di\ erged from early selachian 
stock. The alternative suggestion, that they 
diverged from the shark line after shorten- 
ing of the otic region had occurred, places 
the origin of holocephalians very late, per- 
haps in the Permian or even in the Triassic 
period. If Dean (1904) is correct in his 
identification of Menaspis — a Pemiian form 
apparently not in the selachian line — as an 
early chimaerid, it would be better to seek 
a separate ancestral group for the Holo- 
cephali among the Devonian placoderms 
in which the posterior part of the skull was 
already short. 0rvig ( 1962 ) suggests the 
pt\'ctodonts as such a group. In particular, 
he describes the ptyctodont CtcntircJki as 
possessing a short otic region set behind 
large orbits. Since the ethmoid region of 
CteniircJJa slopes downward anterior to the 
eyes, the general form of the skull does bear 
a resemblance to that of the holocephalians. 
The holocephalians are unlike the sela- 
chians in having no cartilage wall separat- 
ing the otic from the cranial cavity. Fossil 
remains are not sufficiently abundant to 
indicate whether the presence of a parti- 
tion was primary, but Stensio (1963) .states 
that in the arthrodire Kiijdanoiiias))i.s the 
two cavities were separated b\' a thick wall. 
If the condition in Kujdanoickispis was the 
general one in arthrodires as it is in modem 
selachians, one must assume that the con- 
fluence of the ear and brain cavities in the 
Holocephali is a secondary development. 
Although some teleosts show a confluence, 
it seems that in each group the modifica- 
tion arose separately. 

Another characteristic of the posterior 
end of the braincase which is very probably 
secondarv is the consolidation \\'ith the oc- 

cipital region of two or three vertebral ele- 
ments. Rays and also durophagous fishes 
( with disproportionate development of the 
head and shoulder region ) show a fusion 
of skull and vertebral elements similar to 
that of the Holocephali. Such a modifica- 
tion has surely obscured the ancestral con- 
dition in both groups of fishes. 

In comparing the braincase \\ ith that of 
sharks, Holmgren mentions particularly the 
location of foramina. The entryway into 
the orbit for the superficial ophthalmic nerve 
is separate from that of the other nerves in 
both groups, as Moy-Thomas also claims it 
to be in the cochliodont HcJodtis. The fora- 
men for the entrance of the internal carotid 
artery, however, is farther forward in the 
Holocephali. The hypophysis grows ven- 
trally into a depression that has an open 
passage in its floor in the Holocephali, and 
the notochord, \\'hich runs toward it in the 
base of the cranium, lies in a groove rather 
than being completely embedded in the 
cartilage as in sharks. The position of the 
notochord shows, in fact, not only a dif- 
ference from that of sharks, but also a 
similarity to that of certain arthrodires 
described by Stensio ( 1963 ) . In his recon- 
struction of KujcJanowiaspis he shows the 
notochord lying in a groove on the dorsal 
surface of the cranial floor as it does in the 
holocephalians. This similarity to the arthro- 
dires (if it proves to be general) and con- 
trast to the selachians would favor the idea 
that the Holocephali have direct arthrodiran 

The holocephalians also differ from 
sharks, Recent and fossil, in the possession 
of a palatoquadrate cartilage which is fused 
with the chondrocranium rather than being 
suspended in amphistylic or hyostylic fash- 
ion. Since autostyly is known in extant fish 
only in dipnoans, the palatoquadrate in the 
Holocephali represents a remarkable de- 
parture from the piscine plan. DeBeer, 
Moy-Thomas, and Holmgren have each de- 
voted attention to the holocephalian palato- 
quadrate, and, doubtless due to the scarcity 
of the embrvonic material available to them. 

Morphology and Relationships of Holocephali • Stahl 171 

their opinions as to its nature are divided. 
DeBeer and Moy-Thomas ( 1935 ) see it as 
an elongate structure extending posteriorly 
to the mandibular joint b\' an otic process 
which fuses to the cranium against the ear 
capsule. Holmgren ( 1942b ) recognizes the 
cartilaginous lamina between jaw-joint and 
ear region but is not so sure that it is truly 
a part of the palatoquadrate. It may be, 
he feels, a separate cartilage in the early 
embryo. If it is, then the Holocephali are 
characterized by a very short palatoquad- 
rate element, and only fossil fishes having 
a short structure should be sought as pos- 
sible ancestral stock. One might regard 
both the ptyctodonts and the cochliodont 
Hclodus as having a short palatoquadrate 
if that element terminated at the jaw-joint 
and the extension to the otic region devel- 
oped separately, as Holmgren believes it 
does in holocephalians. 

Even though the complete fusion of the 
palatoquadrate to the chondrocranium 
seems so distinctive a feature, Holmgren 
minimizes the distance that this fusion puts 
between holocephalians and sharks. He 
points out that in shark embryos the palato- 
quadrate is connected to the trabeculae by 
a membrane, parts of which chondrify. One 
of the chondrifications attaches itself to the 
palatoquadrate as the orbital process; an- 
other fuses to the trabeculae to fonn the 
subocular shelf. He asks whether, if the 
entire membrane should chondrify, firmly 
uniting the palatoquadrate with the cra- 
nium, this process would be so far from 
what occurs in selachians. A glance at 
Schauinsland's ( 1903 ) ilkistration of the 
developing skull of the 65 mm CaUorhijn- 
chus embryo, however, suggests that Holm- 
gren's speculation here may be wide of the 
mark. At that stage, true cartilage already 
extends from the palatoquadrate area near 
the mandibular joint upwards to include 
the lower half of the orbit. The region in 
front of this smooth mass of cartilage, as 
well as that of the problematic extension 
to the otic capsule, is still in precartilaginous 
form. It seems that if the holostyly of the 

sup oph n 


Fig. 4. Callorhynchus antarcticus. Skull of 60mm embryo, 
lateral view. True cartilage, stippled; precartilage, white, 
mand.. Mandible; pq., palatoquadrate; rpm., medial rostral 
process; rpp., paired rostral processes; sup. oph. n., super- 
ficial opfithalmic nerve. (After Schauinsland.) 

Holocephali was developed through further 
chondrification of a shark-like arrangement 
of the palatoquadrate and the trabecula, 
some indication of the separate nature of 
these elements should appear in this early 
stage before the palatoquadrate is devel- 
oped completely. On the other hand, it is 
possible that the developmental step for 
which we are searching has been gradually 
abbreviated to the point of disappearance. 
Thus, there are two alternatives: either the 
separate palatoquadrate never existed even 
as an embryonic structure in fishes at the 
holocephahan level, or its development was 
suppressed later as the line evolved to the 
present day. Both of these answers imply 
an evolutionary path long separate from 
that which led to modem sharks. 

It is not inconceivable that holocephalians 
might ha\e inherited their autostyly from 
an earlier gnathostome group. That condi- 
tion was evolved, according to Stensio 
( 1963 ) in several groups of arthrodires and 
apparently was not a rare occurrence. 0rvig 
admits, however, that in CtenurcUa (the 
ptyctodont that he regards as being closely 
allied to the holocephalian line ) the palato- 
quadrate was not fused to the neurocra- 
nium. Moy-Thomas, in advocating a coch- 
liodont ancestor for the Holocephali, points 
to the autostylic suspension of the palato- 

172 BuUetin Museum of Comparative Zoology, Vol. 135, No. 3 

quadrate of Helodus as an important simi- 
larity between that fonn and the holoce- 

A second pecuHarity of the visceral arch 
skeleton in holocephalians concerns the dor- 
sal portions of the hyoid arch. As one might 
expect from the autostylic suspension of the 
palatoquadrate, no part of the second arch 
is enlarged as a hyomandibular. The epi- 
hyal and pharyngohyal resemble their serial 
homologues in the successive branchial 
arches. The question arises with regard to 
these elements of the hyoid arch as to 
whether their state is truly primitive or 
whether they have been reduced from a 
specialized, suspensory condition to mimic 
the simple arrangement of the posterior 
arches. Holmgren agrees with DeBeer and 
Moy-Thomas in regarding the non-suspen- 
sory condition of the hyoid arch as primi- 
tive. DeBeer and Moy-Thomas have ex- 
amined the holocephalian hyoid arch in 
detail. In their opinion they have located 
all its parts, including the pharyngohyal, 
and judge it to be unmodified. They argue 
against the possibility that any portion of 
the hyoid could be fused to the cranium 
and so lost to view. To make this supposi- 
tion, one would have to allow the migration 
of the cartilage dorsal to the lateral head 
vein, leaving all the other visceral cartilages 
properly ventral to it. Judging from the 
unanimity of opinion amongst these anato- 
mists, then, it would appear that a non-sus- 
pensory hyoid is one of the primitive char- 
acteristics that the holocephalians have 
carried in their hereditary baggage from 
early gnathostome times. As an early 
gnathostome source for the non-suspensory 
hyoid arch, Watson would have offered the 
placoderms generally, since he believed 
them to be aphetohyoidean. Stensio, how- 
ever, is of the opinion that the early placo- 
denns possessed a suspensory hyoid and 
that the hyomandibular was reduced to a 
non-suspensory bar in groups in which developed. It would not be 
feasible, according to Stensio's inteqoreta- 
tion, to seek a placoderm ancestor for the 

Holocephali if their hyoid arrangement is 
truly a primary one. 

One would expect, in a fish with an ele- 
mentary hyoid, to find a full gill slit an- 
terior to the hyoid arch like that which 
Watson predicated for aphetohyoidean 
placoderms. Although a slit does appear 
in the embryo, it is dorsally placed and soon 
disappears. The space between the hyoid 
and the mandible is later crossed by three 
ligaments: not only is the spiracle absent, 
then, but the area has been completely re- 
built. It is probable that this change is a 
modification connected with the forward 
displacement of the visceral skeleton as a 
whole and, one might add, of the pectoral 
girdle behind it. The palatoquadrate is set 
far forward and fused to the cranium, as 
we have seen; the hyoid is close behind the 
mandible and firmly tied to it by the above- 
mentioned ligaments; and the remaining 
five arches are crowded up under the pos- 
terior end of the cranium. The last two 
pharyngobranchials and epibranchials are 
squeezed to a fusion with each other, cre- 
ating a small flat disc against which the 
scapula abuts. The entire gill apparatus is 
reduced and covered by an operculum. This 
arrangement of the visceral skeleton con- 
trasts sharply with that of extant sharks 
which have five arches, or in the notidanids 
and ChlaimjdoscJachus more than five, in 
an extensive pharyngeal region. Fossil forms 
with a short pharyngeal region (and bran- 
chial arches crowded forward beneath the 
posterior end of the braincase) did exist, 
and might be a more logical choice as a 
group ancestral to the Holocephali than the 
early sharks. The ptyctodonts have been 
figured by Watson (1938) and by 0rvig 
( 1962 ) as having only a small branchial 
area, and Moy-Thomas ( 1936 ) describes 
the cochliodont Helodus as having the pec- 
toral apparatus set close behind the head. 
The possession of a single median rostral 
cartilage also distinguishes the Holocephali. 
In Cltimacra the rostral cartilage is short; 
in Callurhynchus it is longer and bent ven- 
trally; in RhinucJiimaera it is longest and 

Morphology and Relationships of Holocephali • Stalil 


Fig. 5. He/odus simplex. Restoration of fish, lateral view. (After Moy-Thomas.) 

extends directly forward. Carman ( 1904 ) 
believes that the longest cartilages are the 
most primitive. This supposition seems 
reasonable as Rhinochimaeni, with the 
longest rostrum, also shows several other 
characters in what is apparently their 
earliest form. The fossil holocephalians 
Sqiialoraja and Myriacantluis show well 
developed rostral cartilages, the former 
exhibiting some calcification of the element. 
There, however, the trail ends mysteriously. 
Ischyodiis, another extinct form, is figured 
by Dean ( 1895 ) as having a short, blunt 
head, and the earlier possible ancestors, the 
cochliodont HcJodus and the ptyctodont 
RJiampJiodopsis, are not known to have 
possessed rostral structures. CtemireUa, 
according to 0rvig, has a pair of rostral 
processes but not a medial one. It may be 
that such structures were not preserved, 
but in any case the rostral cartilages cannot 
now be used as Ariadne's thread to reach 
the light. 

Holocephalians, like sharks, have paired 
labial cartilages. Howe\'er, in their number 
and form the labial cartilages differ from 
the simple, slim bars — an upper and a lower 
one on each side — which meet at the angle 
of the jaw in selachians. At the mouth 
angle in holocephalians, on each side, there 
are two labial cartilages which meet, but 
the small superior maxillary element and 
the larger, flattened inferior maxillary are 
often fused in the adult. Against the an- 
terior end of the lower jaw there may be a 

premandibular labial cartilage (it is absent 
in Chimaeni coUici); beside the upper jaw 
there are always a large prelabial and a 
smaller premaxillary element. 

The labial cartilages of the Holocephali 
were studied for two reasons. Comparative 
anatomists examined them hopefully as pos- 
sible clues to the history of the descent of 
modem chimaeroids, and workers inter- 
ested in the transition from agnathous to 
gnathous fish sought in them the remains of 
the premandibular visceral arches. Despite 
the descriptions given bv Allis ( 1926), Dean 
( 1906), Carman (1904),' Holmgren ( 1942b), 
Hubrecht (1877), Luther ( 1909), and others, 
the significance of these cartilages has not 
been surely decided. Their early fossil record 
is dubious. 0rvig finds some in CtenureUa 
which he thinks resemble those of holo- 
cephalians rather than those of sharks. 
Holmgren suggests that they might be rep- 
resented in three small elements in Rliam- 
pJwdopsis which Watson ( 1938 ) had iden- 
tified as parts of the hyoid arch. To the 
suggestion that these elements are modified 
premandibular arches there are at least two 
objections: firstly, they are lateral to, rather 
than medial to, the branchial arteries; and 
secondly, they show no close resemblance in 
number or design to visceral arches. Only 
their position against the upper and lower 
jaws argues for the assumption. Taking these 
objections into consideration, Luther (1909: 
32) suggests that "Diese Stiickchen stellen 
aller Wahrscheinlichkeit nach einer ciino- 

174 Biillefin Museum oj Comparaiive Zoology, Vol. 135, No. 3 


Fig. 6. Reconstruction of the head and shoulder girdle of two ptyctodonts; lateral view. A, Rhamphodopsis trispinofus 
Watson; B, Ctenurella glodbochens/s 0rvig. o.d., Dorsal arcualio; a. v., ventral arcualia; has., basal; bro., branchial arch; 
c.hy., ceratohyal; d.sp., dorsal spine; ep.hy., epihyal; lab. cart , labial cartilage; Mk., Meckel s cartilage;, nnandib- 
ular toothplate; n.c, notochord; ph.hy., pharyngohyal; pq., palatoquadrate; pq./p., upper toothplate; rpm., medial rostral 
process; rpp., paired rostral processes;, scapulocoracoid ossification; sp., spinale. (A after Watson; B after 0rvig.) 

genetischen Erwerb dar, der speciellen 
mechanischen Bediirfnissen entsprang." All 
that can really be said with certainty is that 
the laliial cartilages are quite different from 
those of sharks in their number and fonn, 
and in having muscles inserted upon them, 
and that their present condition implies a 
long, separate evolution. 

The remainder of the axial skeleton is 
very much simpler to analyze than the skull, 
but no more directly indicative of the holo- 
cephalians' ancestry. The vertebral column 
presents certain distinctive characteristics 
which may be listed in a straightforward 
manner. Anteriorly, it is consolidated rad- 
ically — not only are the first two or three 
vertebrae fused with the cranium, but the 
first seven elements posterior to the occip- 
ital articulation are broadly fused with each 
other to support the strong dorsal spine and 
an accompanying basal fin-cartilage. True 
centra are never present; in CaUoilu/ncJiu.s 
the notochord is unconstricted; in Wiino- 
chimaera and Chirnaero cartilaginous rings 
develop within the notochordal sheath. 
Rabinerson (1925), who studied the com- 
parative anatomy of the vertebrae of carti- 

laginous fishes, was of the opinion that the 
Holocephali were distinct from the selachi- 
ans in the development of these elements. 
He recognized that the supra- and hypo- 
chordal arch elements of holocephalians 
bore a greater resemblance to those of 
selachians than to those of bony fish, but 
still he held that the similarity was due 
to convergence rather than to close relation- 
ship. Although the holocephalians have 
retained the primitively unconstricted noto- 
chord and in some forms surrounded it with 
a variable number of skeletal rings, they 
share with the selachians the tendency to 
develop arches and intercalary arches above 
and below it. If Rabinerson is correct in 
his opinion that the location of the foramina 
for the spinal nerves differs in holocepha- 
lians and in sharks and that the arch units 
in the two types of cartilaginous fishes have 
been laid down in a different arrangement, 
it would seem likely that the holocephalians 
and selachians evolved separately from a 
group in which only the general nature of 
the arch elements was defined. 

Among the fossil forms which have been 
suggested as belonging at the base of the 

Morphology and Relationships of Holocephali • Stahl 175 

holocephalian line, only the ptyctodonts characterizes the bony fishes, too, it may 

seem to show any vertebral structures have been a common occurrence in early 

which bear a special resemblance to holo- gnathostomes. Watson describes a pelvic 

cephalian characters. 0rvig ( 1962 ) has girdle of this type in the ptyctodont 

stated that CtenureUa possessed a fusion of Rhamphodopsis and Moy-Thomas also at- 

arch elements and an enlarged basal of tributes such a structure to Helodiis. The 

the dorsal fin. Although Watson (1938) holocephalian girdle also contrasts with that 

did not find a synarcual in R]iomp]wdo})sis, of selachians in developing a strong dorsal 

he does figure an element which he be- process and foramina for the passage of 

lieved to be the enlarged basal. nerves to the fin. One cannot seize upon 

In reviewing the appendicular skeleton of these differences as demonstrating a sig- 

the Holocephali, one is struck first by the nificant separation of the holocephalian 

fact that its general structure is similar to line, however, as Dean ( 1909 ) describes 

that of selachians and quite unlike that of in Cladoselache, in the position of the 

bony fish. In both t)'pes of cartilaginous pelvic girdle, a structure with separate left 

fishes the pectoral girdle takes the forni of a and right portions. 

large and firm U which embraces the body The pterygiophores of holocephalians 
from the ventral side. Articulated with this and selachians, although similar in their 
girdle and with the smaller one in the pelvic general extent, do differ from each other, 
region are basipterygia to which are attached The basals of the former group are some- 
jointed fin radials that extend halfway out what more compact, there being two rather 
upon the fin. The remainder of the fin than three in the pectoral fin and one rather 
is supported by demial rays. On closer than two in the pelvic fin. The radials in 
inspection of the holocephalian skeleton, both forms are jointed, although those of 
however, distinctive features do appear, holocephalians show a tendency to fuse at 
The pectoral girdle is extraordinarily mas- their proximal ends. Males of both groups 
sive and contains a pair of channels within bear pterygiophores modified as claspers. 
it for the passage of blood vessels. Its If one accepts Clodoselache, with its broad- 
scapular process extends extremely far dor- based fins, long, unjointed radials, and 
sally. Whether the form of this girdle rep- probable lack of claspers, as typical of the 
resents a modified selachian type or a ancestors of modem selachians, clearly one 
different development is not possible to must derive the holocephalians from se- 
decide. Fossil evidence concerning the lachians later than Cludoselache in which 
deep elements of the skeleton in the shoul- the modern type of fin was already estab- 
der area is meager. Moy-Thomas believed lished or predicate a remarkable convergent 
that the pectoral girdle of the cochliodont evolution in the two groups. Again fossil 
Helodus retained separate left and right data is too scanty to back either alternative 
haKes and if so \\'Ould not have evinced the convincingly. Both Watson and 0r\dg claim 
consolidation characteristic of the holo- that the ptyctodonts they have examined 
cephalian structure. Neither Watson nor probably possessed claspers, and, consider- 
0rvig describes the internal pectoral girdle ing the wide variety of clasper-designs 
of the ptyctodonts. Since the demial annor among cartilaginous fishes shown by Leigh- 
of the shoulder was elaborate, however, Shaq^e (1920 ff. ), it is not impossible to 
one may speculate that inner, non-demial, imagine their having evolved from more 
skeletal elements were not extensively than one source. There is no evidence of 
developed. pterygiophores in ptyctodonts, although 

The pelvic girdle differs from that of Watson speculates that the pelvic fins in 
selachians in consisting of separate left and Rhamphodopsis were probably narrow- 
right halves. Although this arrangement based and freely movable. 

176 Bulletin Museum of Coniparotwe Zoology, Vol. 135, No. 3 

The conclusion to which this review of 
tlie skeletal s>'stem leads is twofold. First, 
although the skull, xertebral column, and 
appendicular structures of holocephalians 
have distinctive features, there exists a suf- 
ficient similarity between the general plan 
of the holocephalian skeleton and that of 
selachians to suggest that the two groups 
are related in some way. Second, it seems 
obvious that the holocephalians have very 
little in common with the bony fishes. 
There are occasional similarities — the ab- 
sence of a partition between the otic and 
cranial cavities and the existence of sep- 
arate halves of the pelvic girdle — but no 
really firm basis exists for postulating a 
relationship between the two lines. 

The nature of the relationship between 
holocephalians and selachians demands 
analysis. Certainly, the cartilaginous nature 
of the skeleton in both is a factor to be 
considered, but the possibility of its having 
been evolved separately removes the obli- 
gation to derive the holocephalians from 
an already established selachian line. It is 
not necessary to adhere to the improbable 
theory that the holocephalian braincase, 
with its downward-sloping ethmoid and 
short otic regions, was derived from the 
early selachian chondrocraniiun. If the non- 
suspensory hyoid is truly primitive, a non- 
selachian origin for it must be sought. If 
it is a secondary development, the feasi- 
bility of its dedifferentiation from the ex- 
panded selachian hyomandibular is still 
questionable. The palatoquadrate is also 
different in its proportions from the se- 
Uichian structure if the point of articulation 
with the mandible marks its posterior limit. 
Its fusion to the braincase seems to have 
been an early event rather than a recent 
modification if its already cryptic embry- 
onic development has any significance. 
Finally, labial cartilages are structures in 
the head which it is difficult to visualize 
as having been derived from their counter- 
parts in selachians. Since the labial carti- 
lages are regarded as vestigial in the latter 
group, it is not Hkely that they would have 

redeveloped to become the elaborate ap- 
paratus of the holocephalians. The median 
rostral cartilage is harder to assess. The 
structure is unique and may be a neomoq^h. 
The postcranial skeleton of the Holo- 
cephali shows two features which are dis- 
tinct from their selachian counteqoarts and 
difficult to imagine as having been derived 
from them. The circumchordal elements in 
chimaerids may be independent develop- 
ments rather than merely reduced versions 
of selachian centra. The absence of any 
type of centra or ring-like structures around 
the notochord in CaUurJiijnclnis is possibly 
a primitive character. The same may be 
said of the separate halves of the pelvic 
girdle found in all holocephalians. 

The Muscular System 

The muscles of the Holocephali have 
been described by several investigators 
interested in evolutionary relationships 
among fishes. Maurer (1912) made a sur- 
vey of trunk musculature, whereas Edge- 
worth (1935), Kesteven (1933), Shann 
(1919), and Vetter (1878) confined their 
attention to the muscles of the head and 
shoulder regions. Vetter provided the most 
exhaustive description of these muscles and 
assigned names to them. His paper is ac- 
companied by a handsome set of drawings 
which are helpful in interpreting the text. 

In surveying, first, the trunk musculature, 
one is forced to recognize the similarity of 
its structure in all fishes. The overriding 
demands of locomotion as perfomied by all 
but a relatively small number of specialized 
fonns have been met by the visibly seg- 
mented, more or less zigzagged myotomes 
which run from the back of the skull and 
pectoral girdle to the caudal fin. Holo- 
cephalians share this general arrangement 
of the trunk muscles with other fishes but 
show one specialization which is apparent 
as soon as the skin is removed: the anterior 
portion of the ventral hypaxial musculature 
has become a non-segmented sheet which 
rises to the level of the lateral line, covering 
the more dorsal hypaxial bundles. This 

Morphology and Relationships of Holocephali • Stahl 177 

> OS- 

Fig. 7. Trunk musculature: anterior part, lateral view. A, C/i/omydose/ochus anguinens; B, Chimaera monstrosa. a,b,c,d. 
Divisions of hypaxial musculature; /., lateral line;^, inferior oblique; o.s., superior oblique; R.p., rectus profundus; line 
xy, dorsal limit of inferior oblique. (After Maurer.] 

sheet inserts, as one would expect, upon the 
pectoral girdle. Maurer (1912), who di- 
vides the hypaxial muscles into superior 
oblique, median oblique, and inferior 
oblique groups, regards the holocephalian 
sheet as being a modification of the in- 
ferior oblique portion. For Maurer, the 
state of the inferior oblique in the Holo- 
cephali represents a more highly evolved 
condition than exists in any other carti- 
laginous fish. In the arrangement which 
Maurer believes is primitive — that seen in 
Chlamydoselaclie and Heptanchus — there 
is a discontinuity between the inferior 
oblique and the median oblique ( line x-y 
in his figures) which is set quite far ven- 
trally, leaving much of the median oblique 
visible. In the course of evolution, the level 
of the discontinuity rises. The inferior 
oblique overlaps the median oblique and 
the latter is gradually reduced. Maurer 
relates this change to the growing dom- 
inance of the pectoral apparatus to which 
the inferior oblique is attached, and states 
that the Holocephali represent the extreme 
expression of this tendency. (He considers 
sharks but not batoids.) In Maurer's opin- 
ion, the Holocephali are also advanced in 
lacking a ventral rectus muscle of the sort 
that Chlamydoseloche shows. That shark 

has the two most ventral muscle bundles 
(c and d in Maurer's figures) rolled medi- 
ally to fonn a band bordering the midline. 
In the sharks, which Maurer regards as 
more highly developed, and in holocepha- 
lians this band does not appear. Through- 
out his paper, Maurer emphasizes the pro- 
gression from primitive selachians to Holo- 
cephali. It is clear that he regards this pro- 
gression as having taken place separately 
from the evolution of the bony fishes. 

Shann (1924) noted that fibers of the 
trunk musculature of fishes are diverted to 
hold the pectoral girdle in place. Although 
Shann doubts that it is possible to draw 
homologies between the various shoulder 
muscles with absolute accuracy in every 
case, he does see a basic likeness between 
the muscles of holocephalians and elasmo- 
branchs. Shann points out, however, that 
the shoulder muscles of the Holocephali 
show a far greater differentiation. In 
sharks, the scapular process is held firm by 
the antagonistic action of the hypaxial mus- 
cles and the cucullaris. The former insert 
upon the posterior border of the scapular 
cartilage and the latter upon its anterior 
edge. In holocephalians, however, both of 
these groups of muscles are subdivided into 
external and internal portions. The origin, 

178 BttUefin Mus-eiiw of Comparative Zoology, Vol. 135, No. 3 

rd.p. rlvp.e. gdci. 

Fig. 8. Chimaera colliei. Muscles of the left pectoral region, lateral view, add.s., Adductor superficialis; d. const. m., dorsal 
constrictor muscle; l-v.m., latero-ventral muscle; p.d.p., protractor dorsalis pectoralis; r.d.p., retractor dorsalis pectorolis; 
r.l.v.p.e., retractor latero-ventralis pectorolis externus; r.p.i., retractor pectoralis superior; scop., scapula; tr.exf.m., trape- 
zius externus muscle. 

insertion, and fiber direction of each differ 
slightly, clearly a more specialized arrange- 
ment. Since the scapular process of holo- 
cephalians rises above the level of the 
horizontal septum, there are also epaxial 
fibers which insert upon it. In sharks the 
epaxial muscles are not involved in the 
shoulder musculature. 

In contrast to the more highly differen- 
tiated state of the holocephalian shoulder 
groups, the muscles which are associated 
with the coracoid region may be simpler 
than those of sharks. The bases of the 
coracobranchials are not fused into common 
coracoarcuals as they are in elasmobranchs. 
The coracohyoid muscles actually originate 
on the coracoid cartilage rather than on the 
fascia over the muscles anterior to it. These 
aspects of the hypobranchial musculature 
outweigh, in Shanns mind, the seemingly 
special, massive development of the cora- 

comandibularis, and he emphasizes his im- 
pression that the Holocephali are in these 
structural arrangements more primitive than 
the sharks and rays. 

From the musculature of the paired fins 
few inferences may be drawn concerning 
the relationships of the Holocephali. Again, 
in principle, the fin muscles of all fishes are 
much alike. To raise, depress, and twist the 
fins all that has proved necessary are a 
dorsal and a ventral muscle mass, some 
fibers of which are drawn into the fin 
over an oblique course. The holocephalians 
present l)ut one modification of the general 
scheme. The proximal portion of the dorsal 
muscle mass associated with the pectoral 
fin is differentiated into discrete bands 
rather than existing as a simple sheet of 
parallel fibers. The most superficial band 
originates on fascia at the level of the lat- 
eral line and inserts upon the anterior edge 

Morphology and Relationships of Holocephali • StaliJ 179 

of the fin through a small tendoi). From the comparable to the quadrato-mandibularis 

girdle another band of fibers extends to of sharks and, anterior to it, a second part 

the front edge of the fin and another to the which is regarded as homologous to the 

posterior edge. Between the latter band selachian preorbitalis. In contrast to the 

and the former two, which insert anteriorly, relative size of these muscles in sharks, 

the deeper fibers which cover the fin- however, the posterior part of the adductor 

radials lie exposed. The distal fibers of in holocephalians is smaller than the exten- 

the dorsal muscle mass are unmodified and sive preorbitalis. The preorbitalis has spread 

resemble those of sharks. A dissection of upward over the broad wall of cartilage 

the remaining fin muscles in either the created in front of the eye, by the fusion 

pectoral or pelvic region shows that the of the palatoquadrate cartilage to the neuro- 

superficial fibers originate upon fascia or cranium, and the development of the high 

upon parts of the girdle and insert upon cartilage wall in the ethmoid region. The 

connective tissue over the fin basals and levator and constrictor elements associated 

radials in the usual way. The deeper fibers with the selachian mandibular arch are not 

originate and insert upon the fin itself as present in holocephalians. The muscles 

they do in sharks. which insert upon the holocephalian labial 

The muscles associated with the anterior cartilages, however, appear in no other 

dorsal fin of holocephalians bear special group of fishes. 

mention. They consist of a proximal and The muscles of the hyoid and successive 

a distal group of fibers on each side. The arches contrast sharply with those of 

proximal muscle mass originates on the sharks. The levator fibers in holocepha- 

plate formed by the anterior vertebral Hans are grouped in external and internal 

fusion, inserts upon the base of the dorsal divisions, as was mentioned above, rather 

fin spine, and acts to elevate the spine. The than existing as a unified cucullaris. The 

distal fibers arise from the broad basal individual constrictor muscles of the pos- 

cartilage of the fin and insert at the base tenor arches, identifiable in sharks, have 

of the dermal fin rays, allowing the web disappeared. Only the hyoid constrictor 

of the fin to be drawn laterally. This com- remains, and this element is expanded to 

bination of proximal and distal muscles, provide the musculature of the operculum, 

which is not found in any other cartilagi- In the possession of a hyoid constrictor of 

nous fishes, may have been present among this kind and in the reduction of the 

the ptyctodonts if 0rvig's interpretation of musculature associated with the branchial 

the skeletal elements of Ctenurella is cor- arches covered by the operculum, holo- 

rect. In Ctenurella, he finds a synarcual cephalians bear a resemblance to the bony 

element beneath the dorsal fin and a basal fishes. Kesteven (1942-1943), who ac- 

piece which could have served as sites of cepted this resemblance as evidence of 

origin for the proximal and distal fibers, evolutionary relationship, was led into the 

respectively. construction of an evolutionary scheme 

Much more has been written about the which is untenable in the face of recent 

musculature of the head and pharyngeal paleontological findings. It might be more 

region than about that of the trunk and correct to suppose that the similarities 

fins. From Vetters ( 1878 ) description of which do exist between holocephalians 

the branchial muscles of the Holocephali, and bony fish have come about through 

one sees that the mandibular arch group convergence. 

resembles the selachian type, lacking the One could assume, then, that the holo- 

complex subdivision shown by that group cephalian branchial musculature, with its 

in bony fishes. The adductor mass in holo- distinctive specializations, developed in cor- 

cephalians consists of a portion which is relation with the crowding forward and the 

180 Btdletin Museum of Comparative Zoolof^y, Vol. 135, No. 3 

Fig. 9. Chimaera monsfroso. Muscles of the head, lateral 
view. C.max., Maxillary cartilage; C.plb., prelabial car- 
tilage; C.pmnd., premandibular cartilage; l.a.o.a., levator 
anguli oris anterior; l.a.o.p., levator anguli oris posterior; 
ic.p., levator of prelabial cartilage; M.a.m., adductor 
mandibulae; M.I. a., labialis anterior muscle; M.l.i., lobiaiis 
inferior muscle; M.I. p., labialis posterior muscle;, 
preorbitalis muscle; n.cap., nasal capsule. (Adapted from 

fusions which took place within the visceral 
and cranial skeleton during the independent 
evolution of the Holocephali. As the gill 
arches became compressed under the oc- 
cipital region and the extrabranchial carti- 
lages spread to form an opercular cover, 
the branchial constrictor muscles gave way 
in favor of an expanded hyoid constrictor 
sheet. The branchial levators, adductors, 
and interbranchials all became reduced in 
accordance \\'ith the reduction and com- 
pression of the cartilages of the arches. 
Since the mandible is short in holocepha- 
lians and forms only a shallow curve, the 
ventral portion of the hyoid constrictor 
(which reaches the midline in sharks as 
the interhyoideus ) apparently shifted the 
origin of its most anterior fibers forward 
to the connective tissue on the posterior 
ventral edge of the mandible. There being 
no division between the palatoquadrate and 
the ethmoid region of the cranium, the 

muscles innervated by the trigeminal nerve 
spread over the entire anterior region of 
the head. The divisions of this muscle 
which insert upon the labial cartilages 
would seem to be late developments. If the 
branchial muscles of the Holocephali 
evolved as suggested here, it would be 
logical to seek an ancestral stock in which 
the shortening of the head region had 
already begun. The ptyctodonts show such 
a condition and may thus be a better choice 
as ancestral material for the holocephalians 
than the longer-headed cochliodont Helo- 
cJus or any early selachian. 

In sum, then, one recognizes in the 
muscular system of the Holocephali a num- 
ber of similarities to the system of sharks, 
many characteristics which are certainly 
specializations peculiar to the group, and 
certain features which are comparable to 
those of bony fishes. Among the holo- 
cephalian muscles, which show some re- 
semblance to selachian counterparts, are 
the trunk and fin muscles, the hypobran- 
chial muscles, and the adductor muscles of 
the mandibular arch group. Within each 
of these groups of muscles, however, some 
unique arrangement appears: the sheet- 
like, nonsegmented inferior obhque among 
the axial muscles; the special nature of 
the proximal pectoral extensors among the 
fin muscles; the great expansion of the pre- 
orbitalis in the mandibular arch group. Be- 
sides these peculiarities, the complexity of 
the shoulder musculature, the anterior dor- 
sal fin muscles, and the muscles which 
insert upon the labial cartilages must be 
regarded as singular and non-selachian in 
nature. The sole resemblance of the holo- 
cephalians to the bony fish lies in the pres- 
ence of an expanded hyoid constrictor and 
reduced musculature of the posterior bran- 
chial arches. In assessing this similarity as 
evidence of convergence rather than rela- 
tionship, one may well be on solid ground. 
Estimating the significance of the similar- 
ities between holocephalians and selachians 
is more difficult, however. Since the mus- 
culature of the holocephalians shows no 

Morphology and Relationships of Holocephali • Stahl 181 

characteristics which are clearly more prim- release eggs into the body cavity, but the 

itive than those of any shark — unless the ostium of the oviduct may be located more 

absence of the common coracoarcuals be posteriorly than it is in the cartilaginous 

so considered — the possibility of its evolu- fishes and the oviduct itself never shows 

tion from a generalized selachian pattern the specialized areas characteristic of the 

cannot be ruled out. On the other hand, oviducts in Chondrichthyes. In species 

the axial and branchial musculature shows which are descended from the earlier parts 

many specializations which are closely of the bony fish line ( Pohjpterus, Acipenser, 

allied to the design of the skeleton. If one Amia, Lepisostcus), the ovary is unenclosed 

considers the evolution of the muscular but is either more elongated or located 

system in correlation with that of the skel- more posteriorly. The oviduct in these 

eton, it seems more logical to suppose that forms differs in design from that in carti- 

it developed, as the skeleton seems to have laginous fishes. Admittedly, the position 

done, from a more ancient root than the of the gonads and ducts in the female 

early selachian fishes. And if one leans lungfish corresponds more nearly to that 

toward the idea of descent from a ptycto- of the Holocephali. The lungfish ovary is 

dont rather than from a selachian group, it much longer, however, and the oviducts 

may be perhaps because it is easier to are unspecialized and have separate ostia. 

imagine building holocephalian muscula- The specialized regions of the holocepha- 

ture upon a ptyctodont frame, especially lian oviduct resemble closely the selachian 

in the head region, than it is to derive it type. Prasad, who made a series of histo- 

from shark-like origins. logical studies of such specialized areas, 

said, ". . . the nidamental glands of Hydro- 

The Urogenital System lagtis coUiei exhibit a structure very similar 

Little research has been done on the uro- ^^ that of a typical oviparous elasmo- 

genital system of the Holocephali. Studies t)ranch . . . (Prasad, 1948: 57). One could 

of its development are lacking and the ^^y, m view of the similar reproductive 

histologv of its component organs has re- habits of oviparous elasmobranchs and hol- 

ceived onlv cursorv attention ( Burlend, ocephahans, that their similarly specialized 

1910; Leydig, 1851). Its gross anatomv, oviducts were a parallel development, but 

which is known, is almost exactlv like that ^^^^^^ ^-^ "» evidence to disprove the idea 

of sharks and quite different from that of ^1^'^^ t^^^^ fi"^^^^''* '"'^y l^^^e inherited both 

bonv fishes habits and the structures from an ear- 

A glance 'at the reproductive organs of li^'-even a very much earlier-common 

the female holocephalian reveals an ar- ^ ^^ ' 

rangement which is exactly like that of , ^'\ searching for differences between 

many selachians. Both ovaries, equally ^^arks and holocephalians, one might seize 

well developed, are set far forward in the ^1^^" ^^^^ ^f * *^^'^* f^''^\ female sharks have 

body cavitv. The oxiducts run lateral to ^ ^^^^^^ ^^^^^^^ *^^^^ holocephalian coun- 

the ovaries' to open xxith a common ostium ^^'^f"'^' ^^ "f • However, the importance 

in the extreme anterior end of the coelomic ^ ^^''^ P"")^ diminishes when one sees that 

space. The shark-like nature of this ar- the young female holocephalian has at least 

rangement is emphasized if one reviews ^ ^^^P urogenital sinus which disappears 

the female genital svstem of other types ^^ the uteri enlarge and press outward in 

of fishes: in almost all teleosts the oviduct the maturing animal. The one unique struc- 

is continuous with the ovary so that the ture possessed by the female holocephalian 

eggs, which are produced in large numbers, is the so-called seminal vesicle. Hyrtl, who 

are at no time free in the coelom. In a reported in 1850 on the indented blind sac 

few forms like the trout, the ovar\' does which opens just posterior to the anus. 


Bulletin Museum of Comparative Zoologij, Vol. 135, No. 

thought that it functioned as a "Samen- 
tasche," but Rurlend (1910) showed that it 
was glandular. Redeke ( 1898 ) sa\\' in this 
sac a possible homologue of the digitiform 
gland of sharks: if the rectum of the holo- 
cephalian were pulled inward from the sur- 
fact>, drawing the "seminal receptacle" in 
\\ ith it, the latter structure would be in the 
same relation to the hindgut as the gland 
of the shark. It is probable that, whatever 
its mode of formation, the blind sac, which 
is not found in any other xertebrate, repre- 
sents a minor specialization which has 
occurred in the later evolution of the Holo- 

The reproductive system of the male hol- 
ocephalian is as shark-like as that of the fe- 
male. In both types of fishes the testis is 
connected by vasa efferentia to a highly 
coiled epididymis through which sperm are 
conducted to the more posterior and wider 
poi-tion of the vas deferens. The vasa ef- 
ferentia of the shark represent transformed 
anterior kidney tubules which lead into the 
embryonic Wolffian duct, and it is pre- 
sumed that the efferent ductules of the 
Holocephali are homologous structures. The 
anterior part of the kidney in immature 
sharks and chimaerids has glomeruli in it, 
but these disappear during growth toward 
sexual matiuity. The anterior part of the 
kidney transforms itself from an excretory 
to a secretory organ and is then known as 
Leydig's gland. In holocephahans, as in 
sharks, its secretion, which passes through 
short ducts to the epididymis and vas def- 
erens, serves as a fluid matrix for the sus- 
pension of the sperm. The posterior portion 
of the kidney in both kinds of fishes re- 
mains excretory, sending urine through one 
or more ureters which empty into a urogeni- 
tal sinus. In commenting upon the arrange- 
ment of pathways in the male system. Van 
Oordt says, "hinsichtlich, der Abfiihrung 
der Spermien stimmen die Holocephalen 
mit den Selachiern iiberein" (Van Oordt, 
in Rolk, 1938, Vol. V: 750). In resembling 
the selachian system so closely, the male 
reproductive system of the holocephalians 

is markedly different from that of the bony 
fishes. In the latter group one finds either 
a duct for sperm which is separate from the 
original archinephric duct or the tendency 
to develop such an arrangement. Even in 
Acipenser, where the expression of this 
tendency is minimal, the urogenital system 
is distinguished from the selachian and hol- 
ocephalian types by lacking a secretory 
portion derived from the anterior end of 
the kidney. No bony fish develops an ac- 
cessory organ comparable to Leydig's gland. 

Given the great degree of similarity be- 
tween male selachians and holocephalians, 
investigators have tried to define the rela- 
tively small points of difference which do 
exist. It has been observed, for instance, 
that the number of vasa efferentia varies. 
In contrast to one in ScylUum, Chimacra has five or six. Borcea (1906: 
349), who made an extensive study of the 
urogenital system of elasmobranchs, con- 
siders that "le nombre des vaisseaux ef- 
ferents est plus eleve et le canal longitudi- 
nal de I'epididyme est plus long chez les 
types les plus primitifs." In making this 
statement, Borcea had in mind the fact that 
the batoids are characterized by a few or 
only one vas efferens. 

Another minor difference concerns the 
posterior region of the vas deferens which 
is enlarged to form an ampulla (Van den 
Brock's term) or a sperm vesicle (Rurlend's 
tenn). In both sharks and chimaerids, the 
inner wall of this structure is thrown into 
folds which divide the lumen of the duct. 
In sharks like Scyllium, however, the par- 
titions are as simple as septa in a mushroom 
cap, whereas the inner walls in a large sec- 
tion of the chimaerid ampulla run into one 
another in a more complex fashion, cutting 
up the space \\'ithin the passage into inter- 
connecting compartments. One feels, upon 
studying these septa, that their different 
design is less important than the fact of 
their presence in both holocephalians and 
selachians. The appearance of these struc- 
tures is a remarkable point of similarity in 
two forms whose lines ( in consideration of 

Morphology and Relationships of Holocephali • StaJil 183 

other organ systems) seem to have sepa- 
rated far back in time. 

This same idea may be emphasized in the 
matter of the claspers of the male. Before 
descril)ing the differences \\'hich exist be- 
tween these structures in sharks and holo- 
cephaHans, one must dwell a moment upon 
the fact that claspers, which are not a com- 
mon vertebrate characteristic, do appear in 
a generally similar form in both of these 
groups of fishes. It would seem, at first, 
that the possession of such claspers is signal 
proof of the close relationship of sharks and 
holocephalians. The major obstacle to the 
acceptance of this idea lies in the fact that 
CladoseJaclic, a fonn apparently anteced- 
ent to modern sharks, shows no claspers. 
If it really had none, then the holocepha- 
lians must have developed their claspers 
independently. That they did so is not an 
impossible assumption. It appears that 
claspers may not be as peculiarly elasmo- 
branchian a character as one would assume 
from a study of extant fishes. If Watson 
(1938) and 0rvig (1962) are correct in 
postulating the presence of claspers in 
Rhamphodopsis and Cteniirelki, respec- 
tively, it may be that these structures were 
possessed by a number of placoderm groups. 
If that was the case, holocephalians and 
selachians might bear claspers inherited 
from separate ancestral stocks. In support 
of this hypothesis one might cite 0rvig's 
finding of a pair of dermal spines anterior 
to the pelvic girdle of CteiuireUa. He be- 
lieves that these spines may have been as- 
sociated with anterior claspers, adjuncts to 
the reproductive system found in holoce- 
phalians but not in elasmobranchs. 

The elaborate array of claspers charac- 
teristic of holocephalians sets these fishes 
apart from other cartilaginous forms. No 
other type of fish has either the aforemen- 
tioned anterior claspers in front of the pel- 
vic fins or the strange median frontal clasper 
or tenaculum set upon the dorsal surface of 
the head. In all extant holocephalians the 
anterior claspers are represented as small, 
gripping structures which are concealed in 

a pouch when not in use. Leigh-Sharpe 
( 1922 ) believes that the prepubic processes 
found in Squaloraja supported anterior 
claspers in that Jurassic form. There are 
no reports of these structures in earlier fos- 
sils, however, except for 0rvig's mention of 
the spines in CtemireUa. Since 0rvig found 
no trace of a tenaculum in CtenurcUa, the 
earliest form of that structure is known 
from Squaloraja and its contemporary, 
Myriacanthiis. The tenaculum in those 
fishes was a long pointed protuberance. In 
living holocephalians, the tenaculum is 
smaller and rounded at its distal end. 

In a lengthy series of papers Leigh- 
Shaqoe ( 1920 ff. ) presents a review of elas- 
mobranch and holocephalian claspers. He 
describes the claspers of Chimaera and 
CaUorhijnchus as having two branches and 
suggests that these branches represent the 
ultimate and penultimate pelvic fin radials. 
He believes that claspers of this type are 
primitive. However, Rhinochimaem, which 
is thought to be the most primitive holo- 
cephalian in terms of its other systems, has 
an unl)ranched clasper more nearly like that 
of sharks. Leigh-Sharpe ( 1922 ) includes a 
drawing of a clearly preserved clasper of 
the fossil Squaloraja which shows a single 
but unusually broad structure terminated 
by a group of small, dermal hooks. Since 
the clasper of Squaloraja is unique in form, 
and since Squaloraja lived in Jurassic times 
when the holocephalian line was already 
established, one cannot be sure that the 
claspers of this fish give evidence of the 
original nature of the holocephalian struc- 

In his classification of the cartilaginous 
fishes according to the type of clasper they 
show, Leigh-Sharpe sets the Holocephali 
amongst the primitive forms for still another 
reason. They have not developed the ab- 
dominal structures — a pair of muscular 
cavities called siphons — which play a role 
in sperm passage during the copulation of 
most elasmobranchs. Holocephalians do 
ha\e a different sort of cavity, though, lo- 
cated in the proximal portion of the clasper. 

184 BtiUetin Museum of Comparative Zoology, Vol. 135, No. 3 


ani cl 

Fig. 10. Claspers of various holocepholian forms. A, Squalorajo; B, Chimaera monstrosa; C, Rhinochimoero atlantica. ant. 
c)., Anterior clasper; bos., basal; cl., clasper;, pelvic girdle; ppb., prepubic processes; r., fin-rays; v.c, vertebral 
column. (After Leigfi-Sharpe.) 

Leigh-Sharpe interprets this cavity as ho- 
mologous to that of Chlamijdosdachc and 
so brackets these fishes together. Surely a 
common category for these forms stands on 
shaky ground. The Holocephali should 
probably be set apart even here if the pres- 
ence of their curious frontal and anterior 
claspers is taken into consideration. 

Although the kidneys have not been 
thoroughly examined histologically, their 
gross anatomy and their relationship to the 
genital organs have been well described 
(Burlend, 1910; Leydig, 1851). There is no 
doubt that these organs, too, are like those 
of elasmobranchs and quite different from 
those of other fishes. Unlike the kidneys 
of the cartilaginous forms, those of bony 
fishes never become closely involved with 
reproductive structures in the male and, in 
both sexes, are generally unifonTi in tubule- 
structure throughout their length, under- 
going neither transformation nor degenera- 
tion at the anterior end as the animal 
reaches maturity. It is not necessary to lean 
entirely upon structural resemblances to 
predicate a possible relationship between 
the Holocephali and the Selachii either. 
The excretory systems of both groups bear 
the same distinctive functional earmark: 
the kidneys resorb urea selectively and 

maintain that substance in the bloodstream 
in unusually high concentration. 

In adult holocephalians, as in sharks, 
urine is produced in the posterior regions 
of the kidney and drained by specially de- 
veloped ureters. This arrangement contrasts 
with that of bony fish in which urine is 
produced throughout the entire kidney and 
is removed through the opisthonephric duct. 
In cartilaginous fish of the male sex the 
anterior kidney and the Wolffian duct be- 
come part of the reproductive system as 
was mentioned before. In females, despite 
there being no secondary use for the an- 
terior region of the kidney, that portion 
degenerates and the Wolffian duct stretches 
forward and ends blindly. In the animals 
of both sexes the kidney gives some hint of 
its originally segmented nature. Especially 
in the anterior region traces of segmental 
divisions remain. The segmental blocks are 
particularly noticeable in the male, because 
ducts leave the gland of Leydig at segmen- 
tal intervals. 

Borcea ( 1906 ) , in the study to which 
reference has already been made, is plainly 
of the opinion that the elasmobranchs rep- 
resent the primitive vertebrates from which 
all the others have descended. Although 
most students of evolution no longer agree 
with that premise, they still admit the pos- 

Morphology and Relationships of Holocephali • Stahl 185 

sibility that certain characteristics of car- relationship between the testis and the kid- 

tilaginous fish may have been carried over ney similar in principle to that which ap- 

from their primitive ancestors at the placo- pears in the cartilaginous fishes. This idea 

derni level. With this idea in mind and in is supported further by the emphasis, again 

consideration of the similarity of the uro- in all vertebrates except bony fishes, upon 

genital systems in holocephalians and elas- the posterior portion of the kidney as the 

mobranchs, one may find interesting the part chiefly responsible for excretory func- 

following comment of Borcea: "C'est le tion. 

groupe des Elasmobranches, qui nous To summarize the foregoing points, one 

montre la succession de ces trois stades (of may state that the urogenital system of the 

the evolution of the vertebrate kidney) avec holocephalians resembles the selachian sys- 

la plus grande nettete. Dune part, ils tem closely. In the position of the gonads, 

presentent I'etat nephridioide . . . plus nette- the specialization of the accessory ducts, 

ment que n'importe quel autre groupe de the nature of the kidney, the development 

Vertebres. D'autre part, ils sont parmi of accessory ureters, and the possession of 

ceux-ci, les animaux les plus primitifs chez claspers on the pelvic fins of male animals, 

lesquels les glandes genitales entrent en re- the two groups of cartilaginous fishes are 

lation avec le rein et son uretere primaire remarkably alike. The type of urogenital 

et alors la serie des changements se montre system they share is distinct in all of these 

d'une fa9on tres manifeste. Chez les Elas- features from that of bony fishes. The kid- 

mobranches la division de I'uretere primaire neys of holocephalians and selachians are 

est tout a fait nette. Chez les plus primitifs set apart from those of all other vertebrates 

d'entre eux ce n'est qu'a I'etat adulte (en by their ability to resorb urea selectively 

relation avec la maturite sexuelle), qu'on and return it to the circulating blood. The 

constate la modification du rein superieur" major point of difference between holo- 

( Borcea, 1906: 251). cephalians and selachians lies in the pos- 

Disregarding Borcea's use of the term session by the former of claspers anterior 

"etat nephridioide " which summons up an to the pelvic fins and of a median tenacu- 

argument quite apart from the subject of lum. 

this paper, one can still see in his statement Although the remarkable similarity of the 
reasons to support the thesis that the elas- urogenital system of holocephalians to that 
mobranch urogenital system is primitive of selachians could be cited as evidence of 
rather than secondarily simplified. If the the evolution of the Holocephali from the 
system is primitive, then there is an alter- selachian line, there appears to be an alter- 
native to the theory that the holocephalians, native to that hypothesis. Since it seems 
whose urogenital organs seem shark-like, possible that the urogenital system of car- 
must therefore have diverged from the tilaginous fishes is truly primitive and if so 
elasmobranchs relatively late. It is possible may have existed in a number of early 
to speculate that, as evidence drawn from gnathostome groups, holocephalians and 
other structures suggests, the holocephalian selachians could have evolved from two 
and elasmobranch lines did split far back separate ancestral stocks. Both types of 
among their placoderm forebears, and that cartilaginous fishes could have retained the 
both groups of fish ha\'e carried to modern urogenital system in its ancient fonn. This 
times the type of urogenital system which theory presumes that the holocephalian and 
those early vertebrates possessed. That a selachian claspers were not derived from 
system of this type may have become wide- the same source. The possibility that pty- 
spread in primitive gnathostomes generally ctodonts possessed claspers allows one to 
is suggested by the development in all ver- believe that there may have been more than 
tebrates except the bony fishes of an inter- one source of those structures at the placo- 


Bulletin Museum of Comparative Zoolop^xj, Vol. 135, No. 3 

derm level. The presence of anterior and 
frontal claspers in living and fossil holo- 
cephalians but not in selachians increases 
the probability of the existence of a sepa- 
rate placoderm ancestor for the holocepha- 

The Digestive System 

The search of the digestive system for 
e\idence of hereditary relationships turns 
up a profitable thread or two and also re- 
veals several alleys which end blindly. As 
might be imagined, an investigation of the 
structure of the teeth gives rise to specula- 
tions based on firmer ground than does an 
examination of the digestive tract itself or 
its associated glands. 

Holocephalians have three pairs of tooth- 
plates. The smallest, called vomerine plates, 
are located in the anterior portion of the 
upper jaw immediately in front of the larger 
palatine pair. The mandibular plates of the 
lower jaw are the largest, being equal in 
length to the other two combined. A com- 
parison of the sections of these teeth fig- 
ured by Brettnacher ( 1939 ) with those of 
cochliodonts shown by Nielsen ( 1932 ) sug- 
gests that the two tooth-types are not 
similar, as Moy-Thomas ( 1936 ) had main- 
tained. A difference if it does exist, is im- 
portant, because the structure of the tooth- 
plates was one of the main supports of the 
theory that the Holocephali arc descended 
from bradyodonts. If the teeth of the two 
groups are truly unlike, and if the presence 
of holostylic jaw suspension in both groups 
is not as important a factor as Moy-Thomas 
thought it was, then the case for close rela- 
tionship becomes very much weaker. 

The discrepancies in tooth-type become 
apparent when descriptions of the internal 
structure of the teeth of each are set side 
by side. Eigil Nielsen (1952: 34) gives 
the now classic description of the bradyo- 
dont type: "This Bradyodont structural 
type is especially characterised by possess- 
ing a system of numerous, more or less 
parallel vascular canals ascending through 
the greater part of the crown, but ending 

blindh' just below the tritoral surface. The 
ascending canals are lined with layers of 
dentine, and the dentine around each canal 
is separated from that around the other 
canals by a hard tissue, described as enamel 
by me in 1932." 

The chimaerid toothplate has been ex- 
amined, described, and figured by Barg- 
mann (1933) and Brettnacher (1939). Their 
accounts of the histology of the toothplates 
agree, although the terminology that they 
use in their descriptions is not exactly the 
same. The outer surface of the crown of 
each plate as well as its embedded portion 
consists of a type of dentine which Brett- 
nacher calls "Hiillendentin" and Bargmann 
calls "Manteldentin." In areas where epi- 
thelium comes in contact with the tooth- 
plate, there is a superficial layer of very 
hard material which, for Brettnacher, is 
true enamel, and for Bargmann merely a 
specially transformed part of the "Mantel- 
dentin." In the interior of the tooth, accord- 
ing to both men, there is a meslncork of 
dentin trabeculae rather than parallel den- 
tinal tubules. Brettnacher gives these tra- 
beculae the special name of "Balkendentin" 
(because they form supporting beams), al- 
though he docs state that they are formed 
by an extension of the odontoblast layer 
which creates the "Hiillendentin." Barg- 
mann uses the temi "Manteldentin" to 
embrace the trabeculae as well as the pe- 
ripheral layer. The spaces in the trabecular 
region are pulp channels which Bargmann 
says are slowly obliterated in the pressure- 
receiving parts of the plate by deposition 
of circumpulpar dentin. 

Jacobshagen, who relies upon Brett- 
nacher's work, has included the chimaerid 
toothplate in his review of the structure of 
selachian teeth (1941). As he presents his 
figures and comparative descriptions, one 
sees that there could be logic in his reason- 
ing that the internal arrangement of the 
holocephalian plate is a primitive variant of 
the dentinal pattern still in existence in 
extant elasmobranchs. Both holocephalians 
and selachians show the outer "Hiillenden- 

Morphology and Relationships of Holocephali • Sfahl 187 

tin " covering an inner trabecular mesh- ( 1951 ) would classify as "tubular dentin" 

work. The categories that Jacobshagen and in Nielsen's figures look singularly dif- 

establishes depend upon the thickness of ferent from anything produced by the Holo- 

the outer dentin layer and the amount and cephali. The "Balkendentin" which fills the 

distribution of the inner "Balkendentin." chimaerid toothplate seems more akin to 

Jacobshagen does make a separate cate- 0rvig's osteodentine in its arrangement and 

gory for the toothplates of the Holocephali, its apparent mode of development, 

not only because of their plate-like struc- If it is not correct to associate holocepha- 

ture, but also because they contain a unique lian and cochliodont teeth with each other, 

material which both Brettnacher and Barg- one is free to seek other relationships. It 

mann describe. Brettnacher calls it "pri- seems not unreasonable to connect the chi- 

mary dentin" and Bargmann uses the old maerid structure with that of ptyctodonts. 

tenn "Kosmin" to refer to it. This substance Ptyctodont plates have been studied histo- 

is found within the pulp channels in several logically most recently by Gross ( 1957 ) and 

regions within each plate. Sometimes the 0rvig (1957). Gross found very little dif- 

Kosmin appears in pearl-like masses strung ference between the teeth of Rhynchodus 

in rows; in some teeth the "pearls" seem and Ptijctodus, and his general description 

coalesced to form an elongated bar. All the reveals a surface layer of dentin supported 

investigators who have discussed Kosmin from within by dentinal trabeculae which 

regard it as an ancient vestige. Schauins- formed a network. Against these internal 

land thought it represented the remains of trabeculae in tritoral areas, what Gross calls 

fused cylindrical teeth. Bargmann discards a secondary dentin was laid down. It would 

this idea, however, for the teeth of younger have been interesting if Gross had referred 

specimens show Kosmin in its undivided to the work of Brettnacher and Bargmann. 

bar-like form. The rather periodic, pearl- Without such a reference one cannot be 

like division, he feels, is a later manifesta- sure whether Gross considered the dentin 

tion. Bargmann has his own theory: he material which he mentions equivalent or 

compares the structure of Kosmin to the similar to that of the Holocephali. It is 

structure of the surface knobs on Cepha- impossible from Gross' description, for in- 

hspis plates, and speculates that in the e\'0- stance, to tell whether he saw something 

lution of the Holocephali this early type of like Kosmin. It appears that he did not. 

hard tissue may have sunk inward. 0rvig's description of PoIeomyJus is more 

Brettnacher and Jacobshagen point out puzzling. He states that the PaJeomyJus 

that dentin in general may have evolved toothplate is much like those of Ptyctodus 

from a relatively soft substance, penetrated and Rhynchodus, and in the number of its 

by widely spaced, branching tubules to a tritoral columns even more like the Mesozoic 

much harder material with close-ranked and Cenozoic Holocephali. But he describes 

parallel tubules. With this idea in mind these tritoral columns as being separated by 

they both consider that the dentin-tissue in acellular bone, while in holocephalians they 

the Holocephali is of the primitive type, the are separated by an interstitial substance 

toothplate deriving its strength from the "not unlike enamel." He refers to the chi- 

arrangement of the dentinal trabeculae maeroid columns as being of a peculiar tu- 

rather than from the hardness of the dentin bular dentin ".s?// ficncris." Since describing 

itself. the Pcdcomylus toothplate in 1957, however, 

As these workers describe and discuss the 0rvig has revised his terminology. For hard 

structure of holocephalian toothplates, it tissues which grow inward toward the basal 

seems less and less likely that these plates region of the toothplate, including tritoral 

have much in common with cochliodont columns, he has introduced the name 

teeth. The latter consist of what 0rvig "pleromic hard tissue." Although he does 

188 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 

not mention Paleoinyhis specifically, in a 
forthcoming book he indicates similarities 
between the pleromic hard tissue of ptycto- 
dont arthrodires and holocephalians. He 
emphasizes the difference in arrangement 
of the pleromic tissues of ptyctodonts and 
holocephalians, on the one hand, and of 
]:)radyodonts, on the other, by classifying 
the pleromic material of the former as 
columnar and of the latter as coronal. 

Although it is usual to analyze the histo- 
logical structure of teeth in an effort to 
derive evidence of phylogenetic signifi- 
cance, it might be well to keep in mind the 
possibilit)' that convergent evolution could 
have brought about similar structure where 
no relationship exists. Radinsky (1961), 
who has found similar patterns in the den- 
tin of bradyodonts, batoids, selachians, and 
dipnoans, is of the opinion that the internal 
structme of teeth may be adaptive and that 
classification should therefore not be based 
entirely upon it. Despite this consideration, 
however, the results of a comparison of 
cochliodont, holocephalian, and ptyctodont 
teeth seems useful. The difference between 
holocephalian teeth and those of cochlio- 
donts, although the latter fishes were ap- 
parently durophagous, should be kept in 
mind. The resemblance between the struc- 
ture of ptyctodont and holocephalian teeth 
may be significant in combination with 
other evidence. 

One should not leave a discussion of chi- 
maerid toothplates without mentioning the 
problem of their origin. Their plate-like 
structure is unusual and has dictated com- 
parisons between the Holocephali and other 
vertebrates like Dipnoi that also possess 
plate-like formations in the mouth. These 
comparisons founder, however, upon one 
point. The folates of lungfish, the teeth of 
most cochliodonts, and the pavement denti- 
tion of rays, all can be shown to be com- 
pounded of units which arise first as 
separate entities. In holocephalians no 
amalgamation of individual denticles is de- 
monstrable. Even in the early embryos 
which Schauinsland studied there were no 

indications of a fusion of teeth or tooth 
buds. It is possible that the Holocephali 
descended from forms whose teeth lost 
their discrete nature and that, as the group 
evolved, ontogenetic evidence of fusion was 
suppressed. Since it has not been demon- 
strated that all fossilized toothplates evolved 
through a compounding of individual units, 
however, it may be that holocephalian 
toothplates were derived from pre-existing 
integral structures. As antecedents of holo- 
cephalian toothplates, ptyctodont plates 
might be preferable to large cochliodont 
teeth produced through fusion. 

In turning from the toothplates to the 
digestive tract, one reaches a series of struc- 
tures whose evolutionary history is even 
harder to define. All the Holocephali show, 
beyond the mouth and pharynx, an esopha- 
gus which leads to the intestine directly, 
without the intervention of a differentiated 
stomach expansion. The obvious question — 
is the lack of a stomach a primitive or a 
degenerate character? — has found no sure 
answer. Since the stomachless condition is 
found in a number of unrelated fishes, one 
could argue that it represents a common 
type of degeneration which has occurred 
independently in several lines. On the 
other hand, the absence of a stomach in 
cyclostomes may be a remnant of the ear- 
liest vertebrate plan. At least one worker, 
Fahrenholz (1915), assumes that this is true 
in the case of the Holocephali. Since one 
answer seems as logical as the other, neither 
can be relied upon to carry much weight 
in the solution of the evolutionary problem. 

The same may be said about the holo- 
cephalian spiral intestine. All the chimaerid 
fishes show an intraintestinal fold which 
takes one slow turn throughout the greatest 
part of the intestinal tube and then makes 
two and a half tighter turns at the posterior 
end. The edge of the fold is free in the 
loosely coiled forepart and caught up in the 
center of the corkscrew tunis at the end. 
This arrangement seems to be a combina- 
tion of the "gerollte" type which Jacobshagen 
(1915) described as existing in a few sharks 

Morphology and Relationships of Holocephali • Stahl 189 

and the "gedrehte" type which he dechiied 
to be much more common amongst the 
selachians. The pecuhar nature of the spiral 
valve can be interpreted in either of two 
ways. Firstly, as Fee (1925) and Dean 
( 1906 ) see it, the viscera of the chimaerid 
fishes, believed by them to be modified re- 
latively late from sharks, have been crowded 
into a shortened body cavity. The stomach 
dilation fails to develop and "the intestinal 
valve, instead of undergoing the further 
spiral development of sharks, makes but a 
few tunis (about four) . . ." (Fee, 1925: 
179). The view of the valve arrangement 
as secondary, as set forth here, might be 
supported by Jacobshagen's contention that 
reduction in the intestinal fold always takes 
place from the anterior end. In fishes 
which bear a degenerate spiral valve or a 
vestigial one, the parts of it that remain are 
in the posterior region of the intestine. 
Secondly, the holocephalian valve might 
be held as primitive, especially in its histo- 
logical structure. Evidence for this conten- 
tion has been presented by Jacobshagen 
(1934), who has made a detailed compara- 
tive study of the spiral intestine in sela- 
chian, dipnoan, ganoid, and jawless fish. 
He points out that the valvular infolding 
in sharks includes only the mucosa and the 
muscularis mucosae. Since the ammocoetes 
larva shows inclusion of circular muscle as 
well, Jacobshagen suggests that the primi- 
tive fold was an indentation of the whole 
intestinal wall which lay within the envel- 
oping serosa. Significantly, the holocepha- 
lians are the only fish that show portions 
of the main circular muscle of the intestine 
still included in the adult valvular fold. Of 
course, Jacobshagen's idea may be incor- 
rect, and the inclusion of the muscle may 
not be a primitive condition in either ani- 

As one advances to a consideration of the 
glands associated with the digestive tract, 
one finds less and less information avail- 
able. Scammon, who has studied the sela- 
chian liver, reports in his account of it that 
"the histology of the adult elasmobranch 

liver was first briefly described by Leydig 
from observations on Chimaera" (Scammon, 
1915: 245). Since Scammon does not even 
think to distinguish the holocephalian from 
the selachian organ, it is apparent that their 
characteristics must be very much alike. 
Scammon holds that the elasmobranch liver 
differs from that of other vertebrates by its 
unique type of lobulation, its accumulation 
of fat within the hepatic cells, and its com- 
paratively slight development of the bile 
duct system. It is impossible to decide 
whether these characteristics are peculiar 
to the shark line or whether they arose deep 
within the placoderm stock. 

The holocephalian pancreas has appar- 
ently not been studied. Siwe, writing in 
1926, does not mention the chimaerid struc- 
ture in his paper on the comparative anat- 
omy of that gland. The only other glandular 
organ associated with the digestive tract of 
the Holocephali that has received attention 
is an intraparietal mass of tubules located 
at the posterior end of the spiral valve. 
Citterio ( 19.32 ) discusses this gland, first 
described by Leydig, suggesting that it 
might be homologous to the digitiform 
gland of selachians and more primitive in 
its intraparietal location. 

Another structure which may have a sela- 
chian homology is the mass of lymphomye- 
loid tissue dorsal to the skin of the palate. 
Extant sharks and rays have a pair of struc- 
tures, similar in their histology, built into 
the sides of the esophagus (Fahrenholz, 
1915 ) . The tissue itself seems of a like con- 
struction in the Holocephali and the sela- 
chians: both show several different types 
of myeloid cells set in a fibrous stroma 
which is highly vascular. Kolmer ( 1923 ) 
who examined the tissue in Chwmero mon- 
strosa regarded it as hemopoietic. Its dis- 
tribution in the Holocephali is singular. 
There is none in the esophageal wall, but 
it exists in a large mass not only over the 
palate but also within each orbit and in the 
ethmoid canal. The tissue masses are con- 
nected by strands which run through foram- 
ina from one area to another. There seems 


Bitlk'tin Museum of Coiuparativc Zoolofiy, Vol. 135, No. 3 

to be a relatively small mass of it, isolated 
from the rest, within a pair of ventral chan- 
nels in the pectoral girdle. Kolmer, im- 
pressed by the fact that much of this tissue 
was surrounded by cartilage, refers to it as 
"knockenmarkahnliche Gewebe." However, 
all of it seems to be external to the peri- 
chondrium. The presence of this tissue 
raises more questions than it answers. No 
one has dared to guess whether it is, in its 
present extent in the Holocephali, a spe- 
cialization lately developed or another 
primitive vestige. 

Conclusions from the nature of the diges- 
tive tract are difficult to draw. The Holo- 
cephali are extraordinary in the structure 
of their teeth, the lack of a stomach, the 
design of the intestinal valve, and the pres- 
ence in association with the gut of unique 
masses of glandular and lymphomyeloid 
material. Examination of these character- 
istics, however, does not produce extensive 
evidence of value in solving the phylo- 
genetic problem. Some clues may be 
gleaned, nevertheless. The greater resem- 
blance between the internal structure of 
holocephalian and ptyctodont teeth than 
between those of holocephalians and coch- 
liodonts suggests, if such similarities are at 
all significant, that there is more likelihood 
of a relationship between the Holocephali 
and the fonner than the latter group. The 
contrast between the integral structure of 
holocephalian toothplates and the tendency 
towarcl fusion of teeth which Moy-Thomas 
( 1936 ) describes as being exhibited by the 
cochliodont Helodus makes it seem im- 
probable that this type of cochliodont was 
ancestral to the Holocephali. 

A hint of similarity to selachians lies in 
the likeness of the liver in the two groups 
of cartilaginous fishes. The affinities of the 
remaining soft parts of the digestive system 
of holocephalians defy analysis. It is impos- 
sible to determine whether the lack of a 
stomach and the minimal development of 
the spiral valve are primitive or secondary 
conditions. The evolution of the glandular 
mass at the posterior end of the intestine 

and of the lymphomyeloid matter in the 
pharyngeal region is equally obscure. One 
must admit, then, that little can be derived 
from an analysis of the digestive organs to 
reinforce either the theory of a selachian or 
a non-selachian origin of the Holocephali. 


The study of the venous system of Chi- 
maera coUiei was undertaken in an attempt 
to clarify the evolutionary history of the 
Holocephali. The fishes of this group have 
been long regarded as an offshoot from 
the shark line and as such have been placed 
with selachians, bradyodonts, and batoids, 
in the class Chondrichthyes. The non- 
replacement of their toothplates resulted in 
their association with the bradyodonts, and 
through the work of Moy-Thomas (1936) 
the theory was established that they might 
have descended from a cochliodont of that 
group. Of late, however, 0rvig ( 1962 ) has 
argued that the Holocephali are more prob- 
ably derived from a ptyctodont ancestor 
and so only distantly related to sharks. 

In an effort to re-evaluate the position of 
the Holocephali, the anatomy of the venous 
system was examined for similarities and 
differences between it and that of other 
fishes. Undoubted resemblances to the 
selachian system were found in the pres- 
ence and arrangement of sinuses and in the 
existence of a subcutaneous network of 
veins. The hepatic portal system, while not 
exactly like that of sharks, resembled the 
selachian system more nearly than that of 
bony fishes. The two main points of differ- 
ence from selachians lay in the absence of 
lateral abdominal veins and the opening of 
the hepatic veins into the posterior cardinal 
sinuses. Further examination of the cir- 
culatory system brought forth no similar- 
ities to the bony fishes but a heart of the 
selachian type, and a unique arrangement 
of arteries in the head region. It was 
obvious from the study of the circulatory 
system that holocephalian structure agreed 
with that of bony fishes only in the lack 
of lateral abdominal veins, and that it bore 

Morphology and Relationships of Holocephali • Stahl 


a much greater resemblance to the selachian 
type. The peculiarities of holocephalian 
vessel arrangement gave no clue as to their 
derivation. It was impossible to detemiine 
whether they represented modifications 
from the selachian plan or whether they 
had been inherited from a non-selachian 

A review of the holocephalian nervous, 
skeletal, muscular, urogenital, and digestive 
systems was made in the search for char- 
acteristics whose derivation could be more 
clearly interpreted. Since each system dis- 
played distinct differences from the com- 
parable system of bony fish, and the sim- 
ilarities to selachian structure were often 
marked, the degree and the implications 
of the resemblance to selachians became 
the focal problem. 

A strong similarity between holocepha- 
lian and selachian structure allows the 
possibility of the origin of the former from 
the latter group but does not necessitate it. 
The possession of similar structures might 
also have occurred through their inheri- 
tance from a common ancestor at a lower 
level of the vertebrate line. In the case of 
a single structure, its presence may be the 
result of parallel evolution. The existence 
of characters which seem unlikely to be 
derived from selachian structures or of 
those which seem more primitive than their 
homologues in sharks might be less equiv- 
ocal. If it can be shown that a structure 
is basically unlike its selachian counterpart 
or that it is not a secondary simplification 
of a form which exists in a more specialized 
state in sharks, one could conclude that the 
Holocephali should logically be traced back 
to placoderm stock by an independent line 
rather than to an early shark group. 

The review of the nervous system re- 
vealed likenesses to selachians in the ar- 
rangement of the autonomic fibers and 
the anatomy of the sense organs and pos- 
terior regions of the brain. Although the 
unusual form of the telencephalon could 
have originated as a modification from the 
selachian plan, it does not appear likely that 

the structure of the pallium itself or the 
simple arrangement of the cranial nerves 
could have been so derived. It appears 
doubtful too, that the pattern of the sen- 
sory canals came from a selachian source. 

The fact that the skeleton of both holo- 
cephalians and sharks is completely carti- 
laginous was once thought to be indicative 
of close relationship, but it has become 
apparent that that conclusion is not the 
only possible one. Since it seems, now, that 
a transition from bone to cartilage occurred 
in several vertebrate lines, one must allow 
that the cartilaginous skeletons of sharks 
and holocephalians may have developed 
independently. If one can look beyond the 
similarity of the skeletal material, holo- 
cephalians can be seen to have several 
skeletal characters that would be difficult 
to derive from early sharks. Their form of 
autostyly is distinctive. Although it ap- 
pears that autostyly has developed several 
times among vertebrates, it is hard to be- 
lieve that the arrangement in the Holo- 
cephali could be a modification of selachian 
structure. If it were, one would expect to 
find a longer palatoquadrate element rather 
than a short one with a process extending 
postero-dorsally in finger-like fashion to 
reach the otic region. Also, the hyoid would 
be expected to show some sign of its former 
involvement in the jaw suspension. In holo- 
cephalians it does not, being to all appear- 
ances exactly like the succeeding arches 
even in its dorsal part. In addition to the 
difference of the palatoquadrate and hyoid 
elements from the shark type, the presence 
of elaborate labial and rostral cartilages and 
the general proportions of the skull, with 
its short otic and steeply sloped ethmoid 
areas, distinguish holocephalians from early 

A study of the muscular system produces 
less that is clearly significant. The similar- 
ity of the musculature of fishes generally 
and the difficulty of ascertaining homol- 
ogies are obstacles to meaningful analysis. 
Peculiarities in holocephalian axial, appen- 
dicular, and branchial muscles are appar- 

192 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 

ent, but there is nothing to indicate whether 
they were or were not derived from the 
selachian plan. There seems to be no sure 
ground for denying that they could ha\'e 

The urogenital system of holocephalians 
resembles that of sharks very closely in the 
nature of kidneys, the gonads, the accessory 
ducts, and the interrelationship between 
those structures. Because that interrelation- 
ship is characteristic of most extant verte- 
brates (bony fish are the cardinal excep- 
tion ) , it is possible to interpret the arrange- 
ment as one which was widespread among 
early gnathostomes and so obviate the ne- 
cessity of deriving the holocephalian system 
from a specifically selachian source. If one 
is free to seek its forerunner in a wide 
variety of early vertebrate groups, one 
might consider the ptyctodonts as having 
had a system which could have been ances- 
tral to the holocephalian type. Although no 
evidence of soft organs remains, it seems 
that ptyctodonts may have had, associated 
with the reproductive system, accessory 
claspers similar to those of holocephalians. 
No trace of those structures appears in any 
other fossil group. 

The digestive system of the Holocephali 
is unlike that of selachians in its lack of 
a stomach and poor development of the 
spiral valve. Among the soft organs, the 
liver is the only structure which bears a 
striking resemblance to its selachian coun- 
terpart. While the evolutionary history of 
the soft parts of the digestive system is not 
clear, the holocephalian toothplates, which 
show no evidence of having developed 
through a fusion of separate teeth, seem 
not to ])e derived from any known shark 

The general conclusion to be drawn from 
this study is that, although similarities be- 
tween holocephalians and selachians are 
numerous, holocephalians possess certain 
characteristics which suggest that these 
fishes evolved from other than a selachian 
stock. The existence in sharks and holo- 
cephalians of like structures does not con- 

tradict this hypothesis, since such structures 
may have been carried over from a common 
ancestor or developed convergently. Even 
the derivation of the Holocephali from the 
bradyodont sharks can be questioned. Al- 
though the cochliodont HeJodus shows, 
according to Moy-Thomas, a number of 
similarities to holocephalians, the teeth of 
that fish show a tendency toward fusion of 
which there is no hint in the Holocephali. 
Hclodus was apparently autostylic, as are 
the holocephalians, but autostyly has arisen 
repeatedly in vertebrate groups and cannot 
be considered as weighty evidence in favor 
of the holocephalian-cochliodont relation- 
ship. There is as good, or better, evidence 
in favor of a relationship between holo- 
cephalians and ptyctodonts. Although the 
ptyctodont palatoquadrate was not fused 
to the cranium, the toothplates appear to 
have been integral structures, and the body 
form, with the large, short head, was sim- 
ilar to that of holocephalians. If one will 
concede that the dennal skeleton of the 
ptyctodonts could have disappeared as the 
evolution of the group continued, then the 
presence of labial cartilages, rostral proc- 
esses, anterior and pelvic claspers, a synar- 
cual, and a dorsal fin supported by radials 
posterior to the dorsal spine, stand forth 
as a substantial and therefore possibly sig- 
nificant number of characteristics suggest- 
ing linkage between the ptyctodont and 
holocephalian lines. 

In sum, one may assume from available 
evidence that holocephalians are not de- 
rived from selachians or bradyodonts but 
have evolved along an independent line. 
However, anatomical similarities between 
extant holocephalians and selachians which 
set both groups apart from the bony fishes 
suggest that these cartilaginous forms 
shared a common ancestor. Tliis ancestral 
stock must have existed at the placoderm 
level or even earlier among unknown ante- 
cedent forms. Although the specific group 
of placoderms from which sharks originated 
is imknown, the ptyctodonts may represent 
the root of the holocephalian line. 

Morphology and Relationships of Holocephali • Stahl 193 


I want to express my gratitude to Dr. 
Alfred S. Romer of Harvard University for 
the guidance and encouragement that he 
has given me throughout my research and 
the preparation of this paper. I am in- 
debted, also, to Dr. Richard Snyder of the 
University of Washington and to Dr. Nor- 
man J. Wilimovsky of the University of 
British Columbia through whose efforts I 
obtained the specimens of CJiimacra colJiei 
that I used. Instruction in the technique of 
latex-injection was given me by Dr. Richard 
Thorington who was, at the time, a grad- 
uate student at the Harvard Biological 
Laboratories. Mrs. Myvanwy Dick allowed 
me to preserve and store my fishes in the 
Fish Department of the Museum of Com- 
parative Zoology and was kind enough to 
lend me a specimen of CaUorhijnchus. The 
list of people who answered my letters of 
inquiry is very long. I owe thanks especially 
to Dr. D. L. Gamble of Ward's and to Mr. 
Thomas E. Powell, Jr. of Carolina Bio- 
logical Supply Company who sent advice 
about injecting frozen-and-thawed material. 
Lastly, I should like to thank Dr. David G. 
Stahl \\'ho encouraged me to undertake this 
work and whose patience and good \\\\\ 
enabled me to finish it. 


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Stefiostoma tifiriniim Cm. und der Holo- 
cephalen. Acta Soc. Sci. Fennicae, 37: 1-60. 

Marples, B. J. 1936. The blood vascular system 
of the elasmobranch fish Sciuatina squatimi 
(Linne). Trans. Rov. Soc. Edinburgh, 58: 

Maurer, Fr. 1912. Die ventrale Rumpfmusku- 
latur der Fische. lena. Z. Naturwiss., 49: 

Moy-Thomas, J. A. 1936. On the structure and 
affinities of the Carboniferous cochliodont 
Helodus simph'x. Ceol. ^h^g., 73: 488-503. 

NicoL, J. A. C. 1950. The autonomic ner\'ous 
system of the chimaeroid fish, Hijdrola^us 
coUiei. Quart. J. Micros. Sci., 91: 379-400. 

Nielsen, E. 1932. Permo-carboniferous fishes 
from East Greenland. Medd. om Gr0nland, 
92(3): 1-63. 

. 1952. On new or little-known Edes- 

tidae from the Permian and Triassic of East 
Greenland. Medd. om Gr0nland, 144(5): 

O'Donoghue, C. H. 1914. Notes on the circula- 
tory system of elasmobranchs, I. The venous 
system of the dogfish (Sei/Uium cauietda). 
Proc. Zool. Soc. London, 19i4: 435-455. 

0RVIG, T. 1951. Histologic studies of placo- 
derms and fossil elasmobranchs, I. The endo- 
skeleton with remarks on the hard tissues of 
lower xertebrates in general. K. Svenska 
Vetenskap.-Akad., Ark. Zool., Ser. 2, 2(2): 

. 1957. Notes on some Paleozoic lower 

vertebrates from Spitzbergen and North Amer- 
ica. Norsk Geol. Tids.skr., 37(3-4): 285-353. 

. 1962. Y a-t-il une relation directc entre 

les arthrodircs ptyctodontides et les holo- 
cephales? Problemes Actuels de la Paleon- 
tologie, Centre Nat. Recher. Sci., No. 104: 

Morphology and Relationships of Holocephali • Stahl 195 

Pander, C. H. 1858. Uber dir Ctenodipterinen 
des devonischen Systems. St. Petersburg, 
65 pp. 

Parkeh, T. J. 1886. On tlie blood vessels of 
Mustelus (into ret if us. Phil. Trans. Roy. Soc. 
London, Ser. B, 177: 685-731. 

Patterson, C. 1965. The phylogeny of the 
chimaeroids. Phil. Trans. Roy. Soc. London, 
Ser. B, 249: 101-219. 

Prasad, R. R. 1948. Observation on the nida- 
mental glands of Hydrohigus colliei. Raja 
rhiiui and Plati/ihinoidis tri.scriatti.s. Copeia, 
No. 1: 54-57. ' 

Rabinerson, a. 1925. Beitriige znr vergleichen- 
den Anatomie der Wirbelsaule der Knorpel- 
fische. Anat. Anz., 59: 433-454. 

Radinsky, L. 1961. Tooth histology as a taxo- 
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Morphol., 109: 73-92. 

Redeke, H. C. 1898. Onderzoekingen betref- 
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en Holocephalen. Amsterdam, 85 pp. 

Reese, A. M. 1910. The lateral Hue system of 
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RoMER, A. S. 1964. The braincase of the Paleo- 
zoic elasmobranch Tamiohatis. Bull. Mus. 
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Scammon, R. E. 1915. The histogenesis of 
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ScHAUiNSLANu, H. H. 1903. Beitriige zur Ent- 
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Shann, E. W. 1919. The comparative myology 
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fishes. Trans. Roy. Soc. Edinburgh, 52: 531- 
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SiWE, S. A. 1926. Pancreasstudien. Morphol. 
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Stensio, E. 1947. The sensory lines and dermal 
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K. Svenska Vetenskaps.-Akad. Handl., Ser. 3, 
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Akad. Handl., 9(2): 1-419. 

Van den Broek, A. J. P. 1938. Gonaden und 
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Van Gelderen, Chr. 1938. Venensystem, mit 
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tarkreislauf. 7/i: Bolk, L. et al., Handbuch 
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Vol. 6: 685-744. 

Van der Horst, C. I. 1934. Spinalnerven. In: 
Bolk, L. et al., Handbuch der vergleichenden 
Anatomie der Wirbeltiere, Vol. 2: 505-540. 

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muskeln der Fische. II Teil. Jena. Z. Natur- 
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Watson, D. M. S. 1938. A ptyctodont from the 
middle Old Red Sandstone of Scotland. Trans. 
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Wilder, B. 1877. On the brain of Chinuiera 
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1877: 219-250. 

(Received 27 Septend)ci\ 1965.) 

196 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 



























































































































— ' 











— ; 





























































































1 — 
























Morphology and Relationships of Holocephali • StaJiI 197 

a.v. int.v. 


v. in+.v. 

Plate 2. The hepatic portal system of Chimaera colliei. Diagrammatic view. X 0.75., Anterior dorsal intestinal 
tributary; oux. spl.v., auxiliary splenic vein; a. v. Inf. v., anterior ventral intestinal vein; h.p.v., hepatic portal vein; i-i.v., intra- 
intestinal vein; /-p. v., lieno-pancreatic vein; mes.v., mesenteric vein; p.d.tnt.v., posterior dorsal intestinal vein; p.v./nt.v., 
posterior ventral intestinal vein. 

198 Bulletin Museum of Comparative Zoology. Vol. 135, No. 3 

Plate 3. The systemic and renal portal veins of Chimaera colliei. Diagrammatic view. X 0.5., Anterior brachial 
vein; ant. card., anterior cardinal sinus; onf.cer.o., anterior tributary of the anterior cerebral vein; ant.cer.p., posterior 
tributary of the anterior cerebral vein; ant.sbct.v., anterior subcutaneous vein; br.i., brachial sinus; br.s.mid-v.exf., mid- 
ventral extension of brachial sinus; coud.v., caudal vein; efh.v., ethmoidal vein; iem.v., femoral vein; h.v., hepatic vein; 
inl.jug.v., inferior |ugular vein; il.v., iliac vein; lot. cut. v., lateral cutaneous vein; m-f.v., maxillo-facial vein; o-n.v., orbito- 
nasal vein; orb.s., orbital sinus; ov.s., oviducol sinus; par. v., parietal vein;, posterior brachial vein; post. card., 
posterior cardinal sinus; post. card. mid-v.exf., mid-ventral extension of posterior cardinal sinus; post.cer.v., posterior cere- 
bral vein; postorb.v., postorbital vein; preorb.v., preorbital vein; prescap.f., prescapular tributary; rect.trib., rectal tributary; 
rn.v., renal vein; r.p.v., renal portal vein; sbsc.s., subscapular sinus; sbsc.tnb., subscapular tributary; s.v., sinus venosus; 
v-a.par.v., ventro-anterior parietal vein; v-p.par.v., ventro-posterior parietal vein. 

Morphology and Relationships of Holocephali • Stahl 


on^.cer. a. 
o-n. V. 
orvt-cer p. 
orb. s. 





iiaf JLLci.v. 


to airuxm 


br. s. 

br. s.mui-v. ejd. 

p ost . card . m ui- V. ext . 

post. cord. 


oar. V. 

from V\.W. 
ov. S. 

V-p. parv. 
or. V, 




200 BiiUetin Miisciiu} of Comparative Zoology, Vol. 135, No. 3 

Plate 4. A, The subcutaneous veins of the closper and pelvic fin. Ventral view. X 1- B, The heart and vessels of the 
hypobranchial region. Ventral view. Coracomandibuloris muscle and right half of pectoral girdle removed. X 1. ob.p., 
Abdominal pore; aff.brn.a., afferent branchial artery; onf. cl., anterior closper; br.o., brachial artery; br.n., brachial 
nerve; br.s., brachial sinus; c.o., conus arteriosus; c-brn.m., coracobranchialis muscle; c-h.m., coracohyoideus muscle; cl.v., 
closper vein; c-m.m., coracomandibuloris muscle; com. card., common cardinal vein; cor.c, coracoid cartilage; hy.c, hyoid 
cartilage; hyp.m., hypaxial muscle; hypobrn.n., hypobranchial nerve; ml. jug. v., inferior jugular vein; mand.c, mandibular 
cartilage; m.w.g.c, medial wall of gill chamber; pect.l., pectoral fin; post. card., posterior cardinal sinus;, sub- 
cutaneous veins of pelvic fin; s.v., sinus venosus; frib./nf.|ug., inferior jugular tributary; v.o., ventral aorta; x, fine vein 
accompanying ventral aorta. 

Morphology and Relationships of Holocephali • Stahl 201 






aff brn.Q, 

C-brn. m. 





cor.c. (cui) 

br. s 

202 Bulletin Muscuiu of Comparative Zoologij, Vol. 135, No. 3 

Plate 5. A, Origin of right inferior jugular vein, showing drainage of tissues immediately posterior to mandible. Ventral 
view. Coracomandibuloris muscle cut and deflected toward midline. XI- B, The brachial veins. Postero-dorsal view of 
right pectoral fin, proximal region. XI- C, The systemic veins entering the sinus venosus. Diagrammatic view. X 0.5. 
D, The anterior cerebral vein and its tributaries. Lateral view. Cartilage removed to show ethmoid and cranial cavities. 
X 1- Anterior brachial vein,- ant. card., anterior cardinal sinus; ant.cer.a., anterior tributary of the anterior cere- 
bral vein; ant.cer.p., posterior tributary of the anterior cerebral vein; onf.cer.v., anterior cerebral vein; ont.v.const.m., 
anterior ventral constrictor muscle; a-v., antero-ventral; far.o., brachial artery; br.s., brachial sinus; cor/., cartilage; cfa/., 
cerebellum; cer.o., cerebral artery; c-m.m., coracomandibuloris muscle; com. card., common cardinal vein; ent.orb.s., en- 
trance to orbital sinus; ep., epiphysis; eth.v., ethmoidal vein; hyp.m., hypaxial muscle; inf. jug. v., inferior jugular vein; 
Inf.hy.m., interhyoideus muscle; inf. orb. sept., interorbital septum; lev.m., levator muscle; lig., ligament; lym., lymphomyeloid 
tissue; mond.c, mandibular cartilage; n.cop., nasal capsule; nos., nostril; o-n.v., orbito-nasal vein; opt. a., optic artery; 
opt. I., optic lobe;, posterior brachial vein; peel. I., pectoral fin; pect.gir., pectoral girdle; poit.card., posterior car- 
dinal sinus; psb.o., pseudobronchial artery; sfasc.s., subscapular sinus; scop., scapula; sp.n., spinal nerve; sup.oph.n., super- 
ficial ophthalmic nerve; s.v., sinus venosus; tel., telencephalon; thy.gld., thyroid gland; tr.inl.m., trapezius internus muscle; 
V. const. m., ventral constrictor muscle; //, optic nerve; III, oculomotor nerve; IV, trochlear nerve; X, vagus nerve. 

Morphology and Relationships of Holocephali • Stahl 203 

insertion c-m.m 
orx^.v. const, m 

V. const. m. 

^' 1//!/" ' ^ int-Wym. 

WW//////;^ C-m.m. 






5ca^. (cjui.) 

to a-v. »ul£ 
of pecl.T. 
channel through 
Lev. m. 
p. br V. 



scop- (cut) 
irtP. juq.v. 
com. card. 



Ini. orb. sept. SUp.opb.a 

204 Bulletin Museum of Comparative Zoology, Vol. 135, No. 3 

Plate 6. A, Systemic veins and related structures in the postero-dorsai region of the head. Lateral view. X 1- B, 
Veins draining dorsal region of trunk. Lateral view. Epaxial muscles cut and partially removed. Scapular cartilage cut 
and deflected ventrally. X 1- o.ep.v., Anterior epaxial vein,- anf.cord., anterior cardinal sinus; a.v.s.c, anterior vertical 
semicircular canal; brn.n., branchial nerve; chcr., chondrocranium; com. cord., common cardinal vein; d. const. m., dorsal 
constrictor muscle; d.f.s., dorsal fin sinus; d.sp., dorsal spine; endl.d., endolymphatic duct; ep.m., epaxial muscle; lot. cut. v., 
lateral cutaneous vein; lym., lymphomyeloid tissue; m.d.v., median dorsal vein; m-l.v., maxillo-faciol vein; of. cop., otic 
capsule; p.cbr.s., posterior cerebral sinus; post. card., posterior cardinal sinus; posf.cer.v., posterior cerebral vein; pos/orb.v., 
postorbital vein; post.scap.fnb., postscapular tributary; sbsc.tnfa., subscapular tributary; s-b.v., spino-basal vein; scap., scap- 
ula; sp.n., spinal nerve;, trapezius internus muscle; V, trigeminal nerve; VII, facial nerve; VII,hyo., hyomandlbular 
branch of facial nerve; IX, glossopharyngeal nerve; X, vagus nerve. 

Morphology and Relationships of Holocephali • Stahl 205 




tr. inlm 



to post. 



post. cand. 
porl.cer. V. 
ot. cap. 


to com. canal. 

d. const. m. 

scop. Uid) 
sbsc. -brib. 

206 Bulletin Museum of Coniixiidtive Zoology, Vol. 135, No. 3 

Plate 7. A, The maxillo-facial vein and its tributaries. Lateral view. Lower portion of adductor mandibulae muscle re- 
moved. XI- B, Deep veins associated with lymphomyeloid tissue dorsal to mouth cavity. Lateral view. Palatoquodrate 
cartilage cut and partially removed. XI- C, The femoral vein. Lateral view. Right side. X 0.75. cart., Cartilage; 
d.lob.v., deep labial vein; etf.rn.v., efferent renal vein; ex.ov.op., external oviducal opening; fern. a., femoral artery; 
lem.v., femoral vein; hy.c, hyoid cartilage; lab. cart., labial cartilage; /ofa.s., labial sinus; lym., lymphomyeloid tissue; 
mond.arf., mandibular articulation; m-f.v., maxillo-facial vein;, mandibular foothplate; n.cop., nasal capsule; o-n.v., 
orbito-nasal vein; orb.s., orbital sinus; ov.s., oviducal sinus; pel.gtr., pelvic girdle; post. card., posterior cardinal sinus; 
postorb.v., postorbital vein; pq., palatoquodrate; preorb.m,, preorbitalis muscle; preorb.v., preorbitol vein; psb.o., pseu- 
dobranchial artery; rect.trib., rectal tributary; r.p.v., renal portal vein; som., samentosche; v. const. m., ventral constrictor 
muscle;, vomerine toothplate; y, hyoid tributary; z, possible venous pathways; V, trigeminal nerve; VII, facial nerve; 
V//,hyo., hyomondibular branch of facial nerve; VII, pal., palatine branch of facial nerve. 

Morphology and Relationships of Holocephali • Stahl 






r p.v. 




pd. qIt. 

208 Bulletin Musctim of Comparative Zoology, Vol. 135, No. 3 

Plate 8. A, The renal portal vein. Lateral view. Left side. XI. B, The hepatic portal system. Dorsal view. X 1-, anterior dorsal intestinal tributary; aif.rn.v., afferent renal vein; a. v. int. v., anterior ventral intestinal vein; b.d., 
bile duct; esoph., esophagus; iem.a., femoral artery; fern. v., femoral vein; g.b., gall bladder; b.p.v., hepatic portal vein; 
hyp.m., hypaxiol muscle; i-i.a., intra-intestina! artery; i-i.v., intra-intestinal vein; il.v., iliac vein; k., kidney; mes., mesen- 
tery; mei.v., mesenteric vein; ov.s., oviducal sinus; pan., pancreas; pan.d., pancreatic duct; par.v., parietal vein; p. d. int. v., 
posterior dorsal intestinal vein; pel.gir., pelvic girdle; p.mes.o., posterior mesenteric artery; post. cord., posterior cardinal 
sinus; p. V. int. v., posterior ventral intestinal vein; r.p.v., renol portal vein; spl., spleen. 

Morphology and Relationships of Holocephali • Stahl 209 

.V. int.v. 

210 Bulletin Museum of Comparative Zoology. Vol. 135, No. 3 

Plafe 9. A, Hepatic veins. Lateral view. Right side. XI- B, Hepatic veins. Lateral view. Left side. X 1- br.s. 
Brachial smus; epid., epididymis; ien.mem., fenestrated membrane; g.b-, gall bladder; h.p.v., hepatic portal vein; h.v. 
hepatic vein;, Leydig's gland; mes., mesentery; pect.gir., pectoral girdle; post. card., posterior cardinal sinus; sem.ves. 
seminal vesicle; test. a., testicular artery; t.s., transverse septum; v.d., vas deferens. 

Morphology and Relationships of Holocephali • Stahl 211 

.card, sem.ves. 

L.qLN ^v.d. 

eft epid. 

212 Bulletin Miiseiivi of Comparative Zoology, Vol. 135, No. 3 

Plate 10. A, The hepatic portal system: veins draining the intestine. Xl- B, The hepatic portal system: veins draining 
the pancreas and the spleen. X 1-, Anterior dorsal intestinal tributary; aux.spl.v., auxiliary splenic vein; a. v. int. v., 
anterior ventral intestinal vein; b.d., bile duct; coei.a., coeliac artery; esoph., esophagus; g.b., gall bladder; h.o., hepatic 
artery; h.p.v., hepatic portal vein; i-i.a., intra-intestinal artery; t-i.v., intra-intestinal vein; l-p.v., lieno-pancreatic vein; 
mei.v., mesenteric vein; pan., pancreas; pon.d., pancreatic duct; pan. v., pancreatic vein; p. d. int. v., posterior dorsal intes- 
tinal vein; p.mes.a., posterior mesenteric artery; p-s.a., pancreatico-splenic artery; p. v. in/. v., posterior ventral intestinal 
vein; spL, spleen; spl.v., splenic vein. 

Morphology and Relationships of Holocephali • Staid 213 




p V. trtt.v. 


rectum . 



p.v. ini.v. 




seum o 

■tXiJ^'UV--^' h!K 

A Review of the Mesochrysinae and 
Nothochrysinae (Neuroptera: Chrysopidae) 




FEBRUARY 24, 1967 





Bulletin 1863- 

Breviora 1952- 

MEMoms 1864-1938 

JoHNSONiA, Department of Mollusks, 1941- 

OccASiONAL Papers on Mollusks, 1945- 

Other Publications. 

Bigelow, H. B. and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
Reprint, $6.50 cloth. 

Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In- 
sects. $9.00 cloth. 

Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth. 

Lyman, C. P. and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- 
malian Hibernation. $3.00 paper, $4.50 cloth. 

Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 15. (Price list on 
request. ) 

Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae 
(Mollusca: Bivalvia). $8.00 cloth. 

Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution 
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Proceedings of the New England Zoological Club 1899-1948. ( Complete sets 
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Publications of the Boston Society of Natural History. 

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Harvard University 

Cambridge, Massachusetts 02138, U. S. A. 

© The President and Fellows of Harvard College 1967. 




In this preliminary subfamilial classifica- 
tion of the Chrysopidae, Mesochrysopidae 
is reduced to subfamilial rank. Keys to sub- 
families and to genera of Nothochrysinae 
are presented. The Nothochrysinae, as 
newly constituted, is characterized by re- 
tention of jugum and frenulum, lack of alar 
tympanal organ, archaic pseudomedia (ex- 
cept in Nothochn/sa and Dyspetochrysa) 
and little sclerotized prosternum. It includes 
Trihoch ryso, Dictyoch rysa, Triploch rysa, 
HypocJirysa, Kimochrysa, Pamochrysa, Pi- 
niachrysa, and Nothochryso. The living 
species of Nothochrysinae are cataloged, 
and the North American species described 
and illustrated. 

New fossil taxa are: Archaeochrysa, new 
genus for Paleochrysa creed i Carpenter, 
fracta (Cockerell) and parancrvis n. sp. 
(Florissant, Colo.); Dyspetochrysa, n. gen. 
for Trihochrysa vetuscuht Scudder. New 
Recent species are: Pimaclirysa albicostoles-, 
Baja Calif., and Arizona; fiiseo, intermedia 
and nigra, southern Calif. 


This study comprises the first section of 
a taxonomic revision of the North American 
Chrysopidae. The concept of the subfamih' 
Nothochrysinae as herein employed is en- 

^ California State College, Fullerton. 

Bull. Mus. Comp. 

tirely new, necessitating a preliminary re- 
classification of the family.- 

There appears no justification for retain- 
ing in a separate family the Mesozoic forms, 
which are placed in the family Meso- 
chrysopidae. These already have achieved 
the alignment of the inner gradate veins 
which precedes the pseudomedia of the 
later forms. In addition, they show the fun- 
damental chrysopid characteristics: many 
straight, unforked branches of Rs, which 
diverge from R at a wide angle; two reg- 
ular gradate series; MPl and MP2 few- 
branched, intersecting the wing margin 
near the base, so that most of the discal 
area is occupied by the R-Rs-MA system. 
These basic features of proportion, while 
difficult to define, readily separate the 
Chrysopidae from all other families. 

The Apochrysinae, a distinctive and ho- 
mogeneous group, has recently been revised 
by Kimmins ( 1952b ) . The Chrysopinae 
constitutes a receptacle for the remaining 
vast and \arious assemblage of forms, 
doubtless requiring subdivision when bet- 
ter understood. 

All described genera of Mesochrysinae 
and Nothochrysinae are included in the 
keys, and all species have been cataloged. 
Descriptions and illustrations have been 
included for all living North American 

- Although this paper was submitted tor publica- 
tion prior to the appearance of Tjeder's paper, 
1966, it has been possible to include his new taxa. 
He delimits the Nothochrysinae (=Dictyochr>sinae) 

ZooL, 135(4): 215-238, February, 196' 


216 Bulk'iin Miiscuni of Comparative Zoology, Vol. 135, No. 4 

species, and for such fossil or Old World 
species as are of particular interest. I have 
not examined material of Dictyochrysa or 
TripJoclin/sa, which were reviewed b\' 
Kimmins (1952a). 

It is interesting that the southwestern 
United States should have such a rich rep- 
resentation (two genera, six species) of 
this archaic group, which has changed but 
little since the Miocene (Adams, 1957). 
This concentration of these relics is rivaled 
by South Africa, with two genera and four 
species, and Australia and Tasmania, also 
with two genera and four species. Such a 
distribution contributes to the mounting 
evidence that the southwestern United 
States has served as an evolutionary re- 
fugium for the Neuroptera. 


Grateful acknowledgment is made of the 
help and encouragement of F. M. Carpenter 
and P. J. Darlington, Jr., of the Museum 
of Comparative Zoology (MCZ), Harvard. 
Ellis MacLeod, of the Biological Labora- 
tories, Harvard, has made many valuable 
criticisms, and has given generously of his 
time during the preparation of the manu- 
script. D. E. Kimmins has kindly examined 
type material and has made useful sugges- 
tions. Material has also been made avail- 
able through the courtesy of C. D. MacNeil, 
California Academy of Sciences (CAS), J. 
D. Powell, University of California, Berke- 
ley, California Insect Survey (CIS), J. N. 
Belkin, University of California, Los Angeles 
(UCLA), P. R. Timberlake, University of 
California, Riverside (UCR), L. Stange, 
University of California, Davis (UCD), J. 
Lattin, Oregon State College (OSC), E. I. 
Sleeper, Long Beach State College, J. E. H. 
Martin, Entomology Research Branch, Can- 
ada Department of Agriculture, Ottawa 
(CNC), Hugo Rodeck, University of Colo- 
rado Museum, and Floyd Werner, Univer- 
sity of Arizona. 

Financial assistance has been provided 
through grants-in-aid from Sigma Xi-RESA, 
the American Academv of Arts and Sci- 

ences, and the University of California, 
Santa Barbara. 


The wing-coupling apparatus consists of 
a large jugal lobe on the fore wing, and 
a frenulum, bearing several long setae, on 
the hind wing; this is essentially the same 
apparatus as in Hemerobiidae, etc. The 
Chrysopinae may have a weak frenulum, 
but the jugal vein is thin, and there is no 
jugal lobe. Loss of the wing-coupling ap- 
paratus appears associated with narrowing 
and strengthening of the wing base, and 
is probably of great adaptive significance. 
A similiar modification has occurred in 
the evolution of the Myrmeleontidae and 
Ascalaphidae from an osmyloid ancestor 
( Adams, 1958 ) , and in the Mantispidae. 

In the Nothochrysinae, there is no obvi- 
ous tympanal organ (Friedrich, 1953; Er- 
hardt, 1916) in the base of R in the fore 
wing, and the stem of M is easily visible 
extending in a straight line adjacent to R. 
In the Chrysopinae, the tympanal organ 
fonns a conspicuous bulge in R, at the 
point where Cu diverges; the base of M 
usually is coalesced with R, but if visible, 
makes a detour posteriorly around the tym- 
panal organ. Probably this is an auditory 
organ (Adams, 1962). 

In No])i]inus ( Apochrysinae), the tym- 
panal organ involves a large area between 
R and M, but is longer and does not form 
a bulge on the underside of R, as in 

Another character, probably of great 
adaptive significance, is the pseudomedia, 
which differs fundamentally in the more 
primitive chrysopids, and in the Chryso- 
pinae — Apochrysinae. In Pimachnjsa (Figs. 
1, 2), Ili/ (Fig. 5), and in most 
of the fossil genera, Psm is composed of 
crossveins alternating with the branches 
of Rs + MA, and is merely a basad exten- 
sion of the inner gradate series. The course 
of the longitudinal veins and composition 
of the jirimitive pseudomedia is particularly 
clear in I h/pochrysd. In the Chrysopinae 

Mesochrysinae and Nothochrysinae • Adaim 217 

(Fig. 45), and Apochrysinae (Fig. 44), and 
in Nothoch)-yso (Fig. 3), Psm, at least 
basally, is composed oi overlapping zig- 
zagged branches of Rs and M, with no 
crossveins between them. ( Some specimens 
of N. fulviceps show no overlap. ) In a few 
genera of Chrysopinae, such as Ytimachrysa 
and Chrysopielki, there is a transition from 
the primitive arrangement apically, with 
no overlap, to the more advanced arrange- 
ment basally, with overlapping veins. 

Primitively, the longitudinal veins forked 
at the posterior wing margin, except for 
the anals in both wings, and CuP in the 
hind wing. This condition may be seen in 
the hind wing of Archacochnjsa (Figs. 40, 
41). There is a tendency for the point of 
furcation to move proximally until it reaches 
the outer gradate crossveins; when this 
occurs, the pseudomedial cells appear each 
to give rise to two marginal veinlets. This 
process begins at the wing base, and pro- 
ceeds apically; the sequence is best seen 
in the fore wing of Archaeochryso para- 
nervis (Fig. 40). Frequently the longi- 
tudinal veins fail to fork at all. 

The pseudocubitus has evolved like the 
pseudomedia, by alignment of longitudinal 
veins and gradate crossveins. In the Notho- 
chrysinae it usualh- is more strongly de- 
veloped than Psm, especially in the fore 
wing. In the Chrysopinae, where the longi- 
tudinal veins may overlap at Psc, it is im- 
possible to detemiine, in most cases, which 
marginal veinlets have arisen from which 
longitudinal veins. For this reason the over- 
lap at Psc shown in Figures 44 and 45 is 
conjectural, although consistent with the 
tracheation of Psc in Chrysopa sipmta 
Walker, as demonstrated by Tillyard 

Evolution of the basal Banksian cell of 
the hind wing has been described by Car- 
penter ( 1935 ) . The archaic condition is 
exemplified by Archacochrysa, where MPl 
is connected to Rs + MA by the sectorial 
crossvein, which intersects MPl in a Y- 
formation. More advanced genera exhibit 
slight basad migration of MF, and fusion 

of MPl with Rs + MA. This evolutionary 
sequence can be seen by comparing Figures 
41, 5, 4, and 2. 

In Chrysopinae and Apochrysinae, the 
wing flexes along a line immediately an- 
terior to Psm (dotted line, Figs. 44, 45). 
To facilitate this flexion in the fore wing, 
the first sectorial crossvein (first crossvein 
distal to the base of Rs + MA ) and the 
branches of Rs + MA are interrupted or 
articulated at the point of intersection with 
the pseudomedia. In the Nothochrysinae, 
the sectorial crossvein is always interrupted, 
but the branches of Rs + MA never are so 
(except in Notliochiysa). Flexion along the 
pseudomedia is accomplished, in Hypo- 
chrysa, by folds traversing the crossveins 
of Psm (Fig. 5). In Pimachrysa, there 
appears to be no distinct line of flexion, ex- 
cept perhaps in P. nigra (Fig. 4). In this 
species the veins show no obvious weaken- 
ing or articulation, but most specimens 
have a slight wrinkle in the membrane 
parallel and anterior to Psm. A similar line 
of flexion occurs between MA and MP in 
most other families of Neuroptera. In 
Chrysopidae, this line has been lost early in 
evolution; the pseudomedial fold is its func- 
tional, but not morphological, equivalent. 
As an interesting example of parallel evo- 
lution, the shape of the wing, and course of 
the pseudomedial fold in Chrysopinae is 
strikingly like that of the smaller Nymphi- 
dae ( e.g. Osmylops, Nesydrion ) . In the 
Nodita-Leucochrysa complex (Chrysopi- 
nae), Psm curves up to join the outer gra- 
date series; this evolutionary line has cul- 
minated in the Apochrysinae, of which the 
nymphid, MyiodactyJiis, is a counterpart. 

Another striking feature of the Notho- 
chrysinae is the near uniformity in thickness 
of most of the veins at the wing base. In 
Chrysopinae and Apochrysinae, R is ex- 
panded basally, and Cu is always inflated 
near the intersection with the first medial 

The Apochrysinae appear to be special- 
ized derivatives of the Nodita-Leucochrysa 
complex ( Chrysopinae ) . The closest resem- 

218 Bulletin Museum of Comparaiwe Zoology, Vol. 135, No. 4 

blance is to forms such as Gonzaga, from 
which the less speciaHzed Apochrysinae, 
such as Synthochn/sa (Fig. 44), differ 
only in the loss of the basal subcostal cross- 
vein, less distance between Psm and Psc, 
condition of MP2, and slight changes in the 
proportions of the wing. 

In Gonzaga (Fig. 45), MP2 bends 
close to CuA, then up toward MP, thence 
down to CuA again, which it joins to form 
part of Psc. If MP2 were to shift posteriorly 
so that it joined CuA directly, eliminating 
the zig-zag, a configuration like that in the 
Ajx)chrysinae would result. It may be noted 
that the second apparent medial crossvein 
of the Apochrysinae ( Fig. 44 ) lies in ex- 
actly the same relationship to the first 
sectorial (sxv) and cubital crossveins as 
does the medial fork in Gonzaga. This is 
significant, since, if MP2 had instead moved 
anteriorly to coalesce with MPl, the ap- 
parent second medial crossvein would be 
expected to intersect CuA at the middle of 
the second cubital cell. 

The degree of overlap of the branches of 
Rs -I- MA in Apochrysinae is the same as 
in Chr\'sopinae, and may be verified by 
matching branches of Rs + MA with the 
veinlets between Psm and Psc, working 
basally from the distal end of Psm. Such a 
count would not, of course, bear on the 
question of whether MP2 had disappeared 
by moving forward and coalescing with 
MPl, or whether it may have moved pos- 
teriorly and fused with CuA. But it pre- 
cludes the possibility that the first median 
crossvein is MP2, as suggested by Kimmins 
( 1952b), since in that case one would have 
to account for an extra crossvein between 
MP and CuA. 

Investigation of male genitalic structures 
in a wide variety of neuropterous types has 
led me to believe that in the Planipennia, 
the ninth gonocoxites primitively articulate 
on the arms of the gonarcus. In the Chry- 
sopidae, the "parameres" of most authors 
appear to be the gonocoxites and are re- 
ferred to as such below. 

Key to the Siibfainilies of Chrtjsopidae 

1. Media posterior 1 runs in an even curve to 

margin, not annulate at intersections with 
l)asal inner and outer gradate crossveins. 
These crossveins intersect MP at nearly 
a right angle, so that MP does not appear 
interrupted by either Psm or Psc ( Meso- 
zoic) Mesochrysopinae. 

- Media posterior 1 usually definitely angu- 

late at intersections with liasal outer and 
inner gradates, so that it appears inter- 
rupted by a definite pseudocubitus, and 
(usually) a pseudoniedia 2. 

2. Jugal lobe of fore wing large; frenulum 

present on hind wing; tympanal organ of 
fore wing absent Nothochrysinae. 

- fugal lobe of fore wing and frenulum of 

liind wing reduced or absent; tympanal 
organ of fore wing present ( reduced in 
Apochrysinae) 3. 

3. In fore wing, basal subcostal crossvein and 

intramedian cell present; space between 
Psm and Psc relatively wide (Fig. 45) 

- In fore wing, basal subcostal crossvein and 

intramedian cell absent; space between 
Psm and Psc relatively narrow ( Fig. 44 ) 
__ Apochrysinae. 

Subfamily MESOCHRYSOPINAE Handlirsch 

Mesochrysopidae Handlirsch, 1908, Die fossilen 
Insekten: 612, pi. 48, fig. 14. Type: Meso- 
chnjsupa Handlirsch 1908. 

Genus MESOCHRYSOPA Handlirsch 

Mc.wehnjsopa Hamllirsch, 1908, Die fossilen In- 
sekten: 612. Type (by monotypy): Hagen- 
ioteimes zitteli Meunier, 1898. 

Mesochrysopa ziffeli Meunier 

Ihigeniotc'iines zitteli Meunier, 1898, Arch. Mus. 
Teyl. (2) 6: 34, pi. 2, fig. 2 (reference from 
Handlirsch ) . 

Mesochrysopa zitteli: Handlirsch, 1908, luc. cit. 
Tillyard, 1916, Proc. Linn. Soc. N.S. Wales 
41: 245-248, text-fig. 8. Martynova, 1949, 
Trudy Palcontol. Inst., Akad. Nauk SSSR 
20: 169. Adams, 1957, Psyche 63: 72. 

Discussion. From the Jurassic limestone 
of Bavaria. A myrmeleontid appearance de- 
rives from the long slender wings, with Sc 
and R apparently fused apically, and with 
many branches of Rs. But the absence of 
pectinate branching of MP and CuA pre- 
cludes relationship with the osmyloid- 

Mesochrysinae and Nothochrysinae • Adams 


myrmeleontoid families. Although the inner 
and outer gradate series are somewhat ir- 
regular, the basal inner gradates are strongly 
inelined, forming a well-defined pseudo- 
media, which does not include MP. The 
multiplication and irregularity of veins is 
likely a function of the insect's large size 
(fore wing 36 mm long); the arrangement 
is essentially like that of the following 

Genus MESYPOCHRYSA Martynov 

Mesijpochnjsa Martynov, 1927, Izvestia Akad. Xauk 
SSSR 21: 764. Type species ( liy monot\p\ ) : 
Mesijpochnjsa hitipentiis Mart\no\ . Mart>no\a, 
1949, Trudy Paleontol. Inst. Akad. Xauk 
SSSR 20: 169. 

Mesypochrysa laiipennis Martynov 

Mesijpochnjsa latipcnnis Martyno\', 1927, Izvestia 
Akad. Nauk SSSR 21: 765, figs. 10-12 (hind 
wing and body structure). Martynova, 1949, 
Trudy Paleontol. Inst. Akad. Nauk SSSR 20: 
169. Type from the Jurassic of Kara-tau ( Tur- 
kestan), Paleontol. Inst. Acad. Sci. USSR, not 

Discussion. Martynov's figure (redrawn 
as Fig. 37) shows the body and hind \\'ing 
in some detail. The course of MP is prob- 
ably drawn too straight; one would expect 
slight zig-zagging at the intersection of the 
basal gradate crosswing. Probably two cross- 
veins were present between the branches 
of MP, as in nearly all chrysopids. 

This fossil is of particular interest in 
that the hind \\'ing is exactly what one 
would postulate in the ancestor of Archae- 
ochnjsa. The basal Banksian cell is similar, 
but the inner crossvein is longer and more 
strongly inclined in Mesijpochnjsa. The 
basal branches of Rs + MA are only slightly 
zig-zagged, not strongly bent as in Archac- 
ochnjsa. It differs from more advanced 
chrysopids in the short, transverse pro- 


Nothochrysinos i\a\as, 1910, Broteria 9: .38-59. 

Type: NotJiochnjsa McLachhm. 
Nothochrysini Navas, 191.3, Ann. Soc. Sci. Bruxel- 

les .37: 303. Type: NotJuxlinisa McLachlan. 

Dictyochrysinae Esben-Petersen, 1918, Ark. Zool. 
11 (26): 26. Tjeder, 1966, p. 246. Tvpe: 
Dicttjochnjsa Esben-Petersen. NEW SYNON- 
YNH (subjective). 

Description. Wing veins of nearly uni- 
form diameter near base, tapering e\enly 
apically; MP extends adjacent to R, but is 
not fused with it; no tympanal organ ap- 
parent on \entral side of R in fore wing; 
jugal lobe of fore wing conspicuously pro- 
duced; Psm (except in Nothoclinjsa and 
DyspetocJinjsa) composed of branches of 
Rs alternating with crossveins. Prosternal 
area largely membranous. Color rarely 

Discussion. Navas (1910) based his 
tribal division of the Chrysopidae on rela- 
tively trivial venational characters. His 
tribe Nothochrysinos included all the forms 
which had the intramedian cell (the cell 
immediately distal to the fork of MP) 
either (juadrangular, or absent. This re- 
sulted in the inclusion of NotJiochnjsa ( most 
species of which belong in Italochnjsa), 
LeucocJinjsa, and Nesochiysa. With the 
exception of three species of Nothochnjsa, 
all of these are members of the Chrysopi- 
nae. Hijpochnjsa was placed in the "Chry- 

There is some question as to the validity 
of Navas' 1910 names. The taxa were clearly 
designated as tribes and names properly 
constructed but for the colloquial plural 
endings. These names bore correct termina- 
tions in the 1913 paper, and have priority 
from that date at latest. While Navas' usage 
of the name makes it nomenclatorially avail- 
able, the concept of the subfamily here pro- 
posed is entirely new. 

The Tertiarv' genera differ from living 
forms in position of the basal subcostal 
cross\'ein onh. Furthermore, Kimochnjsa 
africana (Recent) has the crossvein as in 
the fossils. Two Tertiary chrysopids appear 
to ha\e been described from Eurasia, both 
belonging to Recent genera {Nothochnjsa, 
Chnjsopa). Martynova (1949) reports hav- 
ing collected a series of wing imprints from 
the Miocene of Stavropol (North Cau- 
casus), but these ha\e not been described. 

220 Bulletin Mitseitni of Coniparative Zoology, Vol. 13.5, No. 4 is included here because oi 
its many archaic features, not found in any 
member of the Chrysopinae. However, the 
development of the pseudomedia is more 
advanced than in an\' other nothochrysine 
(except Dy.spetuclinj.m), and does not dif- 
fer from that of the Chrysopinae. Scleroti- 
zation of the prosternum is intermediate 
between that of Pimachrysa and Ilypo- 
clirysa, and Chrysopinae. These are both 
important evolutionary ad\ances over the 
rest of the Nothochrysinae, and tribal sep- 
aration might be advisable, but for the 
small number of genera involved. 

Biolo'^y. Biological information on this 
subfamily is meagre. Nothochrysa (Killing- 
ton, 1937; Toschl 1966) and Uypochrysa 
( Principi, 1958) lay stalked eggs. The larvae 
of Hypochrysa (Brauer, 1867) and Notho- 
chrysa (Killington, 1937; Toschi, 1966) 
have been described. Both N. capitata and 
N. fiihiceps may occasionally carry trash, 
but have exceptionally small thoracic tu- 
bercles. There is some indication of pref- 
erence by adults for ancient host-plants: 
Nothochrysa fulviceps associates with oak, 
and N. capitata with pine (Killington), N. 
calif ornica with both conifers ( W. Wade, 
pers. comm.) and oak. Pimachrysa inter- 
media feeds on willow pollen; other species 
of Pimachrysa have been taken in bait 
traps (fusca, albicostales) or on flowers 
{nifi,ra, on Ccanothus) and may be pollen 
or nectar feeders. 

In Nothochrysa and llyj)Ochrysa nohilis, 
a pale mass of material is deposited on the 
dorsal surface of the female abdomen. This 
has been erroneously interpreted as a 
spermatophore (Killington, Principi). How- 
ever, Toschi, 1966, has observed that in 
female specimens of N. californica bearing 
such a dorsal mass, the spermatophore is 
to be found internally in the bursa. In 
Pimachrysa fusca, and presumably the other 
species, a sac-like spermatophore ( Fig. 21, 
sp) may olten be found protruding from 
the gonocoxites ( "gonapophyses laterales") 
which marginally bear hooked hairs, prob- 
ably for its retention (Fig. 20). 

Key to the Genera of Nothochrysinae 

1. Basal subcostal crossvein of fore win^ 

arises distally to origin of Rs + MA, ap- 
proximately opposite the medial fork 
(MF, Fig. 42); mostly fossil genera -— 
„ 2. 

- Basal subcostal crossvein of fore wing 

arises basally to origin of Rs + Ma, or 
subcostal crossveins numerous; living 
genera 6. 

2. Innermost branch of Rs + MA coalesced 

with MP 1+2 Dyspetochrysa gen. n. 

- Innermost liranch of Rs -|- MA connected 

to MP 1 + 2 by a crossvein 3. 

3. In fore wing Rs -|- MA arises basally, 

nearer the first medial crossvein than 
to MF; in hind wing MP and Rs joined 
by a crossvein Archaeochrysa gen. n. 

- In fore wing, Rs -|- MA arises more api- 

cally, nearer MF than to first medial 
crossvein; in hind wing MP coalesces 
with Rs for a short distance 4. 

4. Inner gradate series forms a smooth curve 

continuous with Psm _. 5. 

- Apical inner gradate series closer to 

Rs + MA than is Psm, thus forming 
a broken curve, with basalmost inner 
gradate of apical series arising prox- 
imally to most distal inner gradate of 
basal series Trihoclmjsa. 

5. Subcosta and costa fused apically; living 

species 11. 

- Subcosta and costa not coalesced; fossil 

species Paleochrysa. 

6. More than two series of gradates in both 

wings 7. 

- Only two series of gradates in both wings 


7. Three regular series of gradates 

- Triplochrysa. 

- Six or seven irregular series of gradates; 

discal area divided into a meshwork 
of polygonal cells Dictyochrysa. 

8. Proximal branch of Rs + MA usualh 

coalesced for a short distance with MP 
1; in living species, pseudomedial fold 
follows entire length of Psm in both 
wings Nothochrysa. 

- Proximal branch of Rs + MA never co- 

alesced with MP 1, so that Psm consists 
of alternating longitudinal veins and 
crossveins; pseudomedial fold intersects 
first sectorial crossvein above intra- 
medial cell, and may cross some basal 
gradates of the Psm, but never extends 
the full length of Psm 9. 

9. Hooked hairs on ninth gonocoxites ("gon- 

apophyses laterales") of female; basal 
crossveins of Psm not interrupted by 
pseudomedial fold Pimachrysa. 

Mesochrysinae and Nothochrysinae • Adains 221 

— No hooked hairs on ninth gonocoxite ot 

female; basal crossveins of Psni inter- 
rupted l)y pseudoniedial fold 10. 

10. In fore winji, 2A and 3A fused apically; 

male epiproct fused with ninth termite, 
with ventral processes Hijpochnjsa. 

— In fore winji, 2A and 3A connected by a 

cross\ein; male ectoproct demarked 
from ninth tergite, with no ventral pro- 
cesses 11. 

11. Intramedian cell rhomboid, almost as high 

as long; fore pterostigma shorter tlian 
half wing width Pamochnjsa. 

— Intramedian cell about twice as long as 

high; fore pterostigma as long as two- 
thirds wing width Kimochnjsa. 

Genus DICTYOCHRYSA Esben-Petersen 

Dictyochnjsa Esben-Petersen, 1917, Proc. Linn. 
Soc. New South Wales 42: 214-215 (type, 
by original designation, D. fulva Esben-Peter- 
sen); 1918, Arkiv for Zool. 11(26): 1-37. 
Kimmins, 1952, Ann. Mag. Nat. (12)5: 
70-72 ( key to species ) . 

Dictyochrysa fulva Esben-Petersen 

D. fulva Esben-Petersen, 1917, Proc. Linn. 
Soc. N.S. Wales 42: 214-215, pi. 13, fig. 10. 
Holotype, from Queensland, Australia, Frog- 
gatt Collection, CSIRO, Canberra (not seen). 
Kimmins, 1952, Ann. Mag. Nat. Hist. (12)5: 

Dictyochrysa peferseni Kimmins 

D. peterseni Kinmiins, 1952, Ann. Mag. Nat. 
Hist. (12)5: 70-71, fig. 2 (wings, head, pro- 
notum). Holotype, from Mt. Kosciusko, New 
South Wales, 6000 ft., 12 December 1931, 
R. J. Tillyard, coll. In Brit. Mus. (Nat. Hist.), 
not seen. 
Additional distribution. Mt. Maria, Tasmania? 

Dicfyochrysa latifasciata Kimmins 

D. latifasciata Kimmins, 1952, Ann. Mag. Nat. 
Hist. (12)5: 71-72, fig. 3 (male genitalia). 
Holotype male, from Mt. Wellington, S. Tas- 
mania, 1300-2300 ft., 12-21 March 1913. 
Abdomen in balsam. Brit. Mus. (Nat. Hist.), 
not seen. 

Triplochrysa pallida Kimmins 

T. pallida Kimmins, 1952, loc. cit., fig. 1 
(wings). Holotype female, from Bunya Mt., 
Queensland, Australia, 27 January 1951, in 
Brit. Mus. (Nat. Hist.), not seen. 


Hijpochnjsa Hagen, 1866, Stettiner Entomol. Zeit. 
27: 377. Type species (by monotypy): 
Chrysopa nohilis Schneider. 

Hypochrysa nobilis (Schneider) 

Figures 5, 32, 33 

Chrysopa elc^ans Burmeister, 1839, Handb. d. 
Entomol.: 981. Type locality "Harze," Saxe- 
sen collector, probably in the Zoological Mu- 
seum, Univ. of Halle (not seen). This name 
has priority, but is a nomen oblitiim under 
Rule 231). Not to be confused with Hcm- 
erohius elegans Ciuerin 1838, which was trans- 
ferred to Chrysopa by Schneider in 1851, and 
probably is a Gonzafia; this name is pre- 
occupied by Henicrobiiis elegans Stephens 

Chrysopa nohilis Schneider, 1851, Symb. ad 
Monogr. Gen. Chry.sopae: 142-144, pi. 51. 
( Name replaces elegans Burmeistei . ) 

Hypochrysa nobilis: Hagen, 1866, Stettiner 
Entomol. Zeit. 27: 377. Brauer, 1867, Verb. 
Zool.-Bot. Ces. Wien 17: 27-29, pi. 9, fig. 1 
(larva). Navas, 1913, Insecta 28: 129-1.30, 
fig. 1; 1915, Arx. Cien., Barcelona 3(2): 
88. Principi, 1956, Atti Acad. Sci. Inst. Bolo- 
gna, Rend. (XI)3: 1-3, pi. 1 (Ethology, 
photographs of adult, egg, habitat); 1961, 
Mem. Mus. Civ. Stor. Nat. Verona 9: 109. 

Description. Male ninth tergite fused 
with ectoprocts, not articulated with ninth 
sternite. Eighth and ninth sternites not 
fused, but moveable. Ectoprocts (Fig. 32, 
ect. ) each posteroventrally bearing heavily 
sclerotized toothed process. Gonarcus flat- 
tened, heavily sclerotized, black; mediuncus 
bilobed; gonocoxites ( "parameres" ) weakly 
sclerotized, dorsolateral to mediuncus lobes. 

Di.itri])ution. Widespread in southern 

Remark. Hypochrysa arg.entina Navas, 
1911, is a Chrysopa (Tjeder, 1966). 


Triplochrysa Kimmins, 1952, Ann. Mag. Nat. Hist. 
(12)5: 69-70 (type, by original designation, 
7'. pallida Kimmins). 

Genus KIMOCHRYSA Tjeder 

Kiniochrysa Tjeder, 1966, S. African Anim. Life 
12: 254. Type .species (by original designa- 
tion): Kimochrysa inipar Tjeder. 


Bulletin Miiscmii of Comparative Zoology, Vol. 135, No. 4 

Kimochrysa impar Tjeder 

Kii)n>chnj.sii impar Tjeder, ibid.: 256—259, figs. 
820-834. Holotype male, Kleinmond, Cape 
Province, S. Africa, in the South African 
Museum, Cape Town (not seen). 

Di.scussio)}. This is the only species oi 
the Nothochrysinae with numerous sub- 
costal \ ('inlets in the fore wing. In the male, 
the ninth abdominal tergite is not hinged to 
the eighth and ninth sternites, and these 
sternites are completely fused. 

Kimochrysa africana (Kimmins) 

Hypochrtjsa africdiui Kimmins, 1937, Ann. Mag. 

Nat." Hist. (10) 19: 307-308, fig. Holotype 

female, from Worcester, Cape Province, S. 

Africa, in Brit. Mus. (Nat. Hist.) (not seen). 
Kimocliriisci africana: Tjeder, 1966, S. African 

Anim. Life 12: 259-261, figs. 835-838. 

Kimochrysa raphidioides Tjeder 

Figures 10, 29-31 

Kimochrysa rapJiidioide.s Tjeder, 1966, ibid.: 261- 
262, figs. 839-844. Holotype female, from 
"Cap Drege" ( abl^rexiation for "Cape of 
Cood Hope, Drege, collector"), in the MCZ 
( examined ) . 

Description. Head marked with dark 
as in Figure 10. Pronotum transverse, a 
thin transverse black line at outer edge of 
furrow, a short longitudinal line at posterior 
corner. Subcostal crossvein distal to origin 
of Rs + MA. Male ninth tergite hinged 
to ninth sternite (Fig. 31); fusion of eighth 
and ninth sternites demarked by internal 
sclerotized ridge. Gonocoxites ( Figs. 29, 30, 
gcx ) paddle-shaped. 

Discussion. In addition to the type, a 
male with no data is also in the MCZ. This 
species is doubtfully distinct from africana, 
from which it differs only by the position 
of the subcostal crossvein (proximal to the 
origin of Rs + MA in africana), and by 
greater sclerotization of the spermatheca 
(which may be due to different degrees of 
maturity in the specimens). Tjeder dif- 
ferentiates these two species principally on 
the position of the second medial crossvein. 
Hilt in africana, it varies in position from 
distal to jMoximal to the fork (Kimmins, 

pers. comm.). In the male of raphidioides, 
the crossvein intersects MP at the fork. In 
the specimen of K. impar figured by Tjeder, 
the second medial crossvein is proximal to 
the fork on one wing, and distal on the 

The coloration of the two species is sim- 
ilar, although the frontal markings below 
the antennae are absent on the type of 
africana, and faint on the second specimen; 
the interantennal marks are faint on both 
specimens of africana ( Kimmins, pers. 
comm. ) . 

Genus PAMOCHRYSA Tjeder 

Pamochrijsa Tjeder, 1966, S. African Anim. Life 
12: 248. Type species (by original designa- 
tion): stcUata Tjeder. 

Pamochrysa sfellata Tjeder stcllata Tjeder, ibid.: 250-253, figs. 
797-815. Holotype male, from Cathedral 
Peak, Forestry Reserve, Indumeni River, 
Drakensberg, Natal, S. Africa, March 1959, 
in the Natal Museum, Pietermaritzburg, not 

Genus NOTHOCHRYSA McLachlan 

Nothochrysa McLachlan, 1868, Monograph of the 
British Neuroptera-Planipennia. Trans. En- 
tomol. Soc. London (1868): 195. Type 
species, Chrysopa fidviceps Stephens, desig- 
nated by Banks, 1903, Trans. Amer. Entomol. 
Soc. 29:' 142. Tjeder, 1941, Entomol. Tidskrift 
(1941): 30-31. Principi, 1946, Bol. Inst. 
Entomol. Univ. Bologna 15: 86. 

Nathanica Navas, 1913, Trans. 2nd Intern. Congr. 
Entomol., Oxford, 1912, 2: 181 (type [by 
subsequent designation of Tjeder, 1941, ibid.]: 
Hemcrobius capitatii.s Fabricius). 

As McLachlan failed to designate a type 
for this genus. Banks' listing of N. fulviceps 
as type constitutes a valid type designation. 
Principi (1946) restricted Nothochry.'ia to 
include fulviceps and capitata only, erect- 
ing a new genus, IfalocJirysa, for N. italica 
Rossi. It appears that the great majority of 
Old World species formerly included in belong instead to Italochnjsa. 

Description. Head wide, vertex low, 
eyes small, labrum emarginate. Antennae 

Mesochrysinae and Nothochhysixae • Adams 223 

about as long as fore wing; scape about as 
long as wide, flagellar segments longer than 
wide. Prosternum small (Fig. 35). Tarsal 
claw either with basal enlargement {fulvi- 
ceps) or simple {copitata and californica). 
Fore wing (Figs. 3, 43): basal subcostal 
crossvein opposite midpoint between first 
medial crossvein and medial fork. Rs origi- 
nates basad of medial fork; two rows of 
gradates. Psm formed by overlapping lon- 
gitudinal veins, may extend beyond basal 
inner gradates; pseudomedial fold present; 
MP2 parallels MPl, connected by a cross- 
vein; second medial crossvein intersects 
MP2 midway between Mp and CuA. Psc 
parallels Psm; runs into outer gradate series. 
lA forked apically; 2A and 3A simple. Jugal 
lobe prominent. Hind wing: frenulum well 
developed, MP coalescent with Rs for a 
moderate distance; two rows of gradates. 
Psm \ve\\ developed, but longitudinal veins 
do not overlap ( except basally in fiilviceps ) ; 
instead each intersects the next where it 
bends shaiply apicad, so that the veins be- 
tween Psm and Psc are exactly opposite the 
basal branches of Rs ( a very conspicuous 
and characteristic feature ) . Pseudomedial 
fold present. Psc runs into outer gradate 

Male: (Fig. 17) ninth abdominal tergite 
heavily sclerotized anteroventrally, with ir- 
regular margin; fused with ectoproct. Eighth 
sternite distinct or only weakly fused with 
ninth sternite. Cuticular gland openings 
present. Gonarcus (Figs. 18, 19) bears wide 
triangular mediuncus; gonocoxites small. 
Female: No hooked hairs on ninth gono- 
coxites; eighth sternum bilobed, lobes fit- 
ting into cuplike ninth vaKulae ("divertic- 
ulos linguiformes" of Principi ) so as to close 
the genital cavity anteriorly, as in Chryso- 

Key to the Recent Species of Nothochrysa 

1. Proiiotum membranous medially; Psm in 
lore wing extends beyond inner gradate 
series; head orange and lilack; i\orth 

American californica. 

Pronotum sclerotized medially; Psm in 
fore wing runs into inner gradate series; 

head concolorous, orange or orange- 
brown; European -_ 2. 

2. Thorax with median pale band, tarsal 
claws with basal expansion; several mar- 
ginal veinlets from Psc forked; large 
species ( expanse 37-48 mm ) .... fulviceps. 

— Thorax orange-brown, no median band; 
tarsal claws simple; marginal veinlets 
from Psc rarely forked; smaller (expanse 
27-36 mm) capitata. 

Nofhochrysa fulviceps (Stephens) 

Cbrysopa fulviceps Stephens, 1836, lUustr. Brit. 
Entomol. Mand., 6: 101. Types: 4 syntypes 
in the Brit. Mus. (Nat. Hist.), not seen. 

Notlwchiysa ftilvice})s: McLachlan, 1868, Trans. 

Entomol. .Soc. London (1868): 207. 
Nathanica fulviceps: Navas, 1913, Trans. 2nd 

Intern. Congr. Entomol., Oxford, 1912, 2: 

181. Killington, 1937, Monogr. Brit. Xeuropt. 

2: 236-242, fig. 114 (male genit.), fig. 115 

(female genit.), pi. 27, fig. 2 (wing), pi. 30, 

fig. 3 (larva). 
Nathanica fulviceps var. flavida Navas, 1919, Bol. 

Soc. Entomol. Espana 2: 55. 
Nothochn/sa fulviceps: Tjeder, 1941, Entomol. 

Tidsicrift (1941): 30-31. 

Di,stril)tdion. Europe. 

Nofhochrysa capitafa (Fabricius) 

Hemerobius capitatus F'abricius, 1793, Entomol. 

Syst. 2: 82. Type locality "Germania. Dom. 

de Paykull." Zool. Mus., Univ. of Kiel, not 

Chrysopa capitata: Curtis, 1834, Brit. Entomol. 

" pi. 520. 
Nothochrysa capitata: McLachlan, 1868, Trans. 

Entomol. Soc. London (1868): 207. 
Nathanica capitata: Navas, 1913, Trans. 2nd 

Intern. Congr. Entomol., Oxford, 1912, 2: 

181. Killington, 1937, Monogr. Brit. Neuropt. 

2: 242-246, fig. 115c (female genit.), pi. 27, 

fig. 3 (wings); pi. 30, fig. 4 (larva). 
Nothochrysa capitata: Tjeder, 1941, Entomol. 

Tidskrift (1941): .30. 

Distrihtition. Europe. 

Nofhochrysa californica Banks 

Figures 3, 8, 17, 18, 19, 22, .35 

Nothochrysa californica Banks, 1892, Trans. Amer. 
Entomol. Soc. 19: 373. Male holotype from 
Los Angeles, Calif., MCZ No. 11406, examined. 
1903, Trans. Amer. Entomol. Soc. 29: 142, pi. 
2, fig. 3 (wing base); 1904, Cat. Neuropt. In- 

224 BtiUcfiii Museum of Comparative Zoology, Vol. 135, No. 4 

sects I'.S., Philadelphia: 26. MacGillivra\ , 
1894, Canad. Entoniol. (1894): 171. Smith, 
1932, Ann. Entoniol. Soc. Anier. 35: 582, pi. 
1, fig. 5 (body and wings, color). Bickley 
and MacLeod, 1956, Proc. Entoniol. Soc. 
Washington 58: 182-183. 

Description. Head (Fig. 8): antennal 
socket.s large, margin.s blaek. A black line 
extending down .suture from antennal 
.socket to anterior tentorial pit; black line 
connecting pit.s bent posteriorly in middle; 
dark mark on clypeu.s connected to this 
line laterally; black genal .spot, near man- 
dibular articulation. Vertex elevation slight; 
median and lateral stripes connected to 
circumantennal marks. Antennae black; 
scape wider than long. Palpi dark, pale at 

Pronotum divided by median longitudinal 
pale membranous area; lateral sclerites dark 
umbraceous, lateral margins pale. Alinotum 
dark, mesoprescutum with posterior pale 
band interrupted by black median longitu- 
dinal suture; mesoscutum pale anteriorly. 
Propleuron and sternum dark. Meso- and 
metapleura dark, pale posteriorly. Coxae 
dark, trochanters dark basally; femora with 
dorsal and ventral longitudinal dark stripes, 
confluent on hind femur. Tibiae pale with 
dark basal and apical marks; tarsi fuscous, 
dark apically. Claws lack basal tooth. 

Fore wing ( Fig. 3 ) : veins black, except 
C, R, 3A and posterior marginal vein pale 
basally; sensory area posterior to apex of 
3A and posterior marginal vein pale basally; 
sensory area posterior to apex of 3A, and 
jugal lobe, fuscous; stigma fuscous, spaces 
between apical subcostal crossveins ivory. 
MPl desclerotized at intersection with basal 
sectorial crossvein; Psm continues beyond 
basal inner gradates. Hind wing: venation 
dark, R white to stigma; Rs and MPl white 
basally; CuP white; stigma as in fore wing. 

Abdomen: tergites black, narrowly pale 
apically; sternites may be pale both basally 
and apically. Female ninth tergite with 
pale spot posterolaterally; tenth tergite pale, 
callus cerci and posterior margin black. 
Eighth sternum (Fig. 22) bilobed. Dorsal 
postcopulatory mass white or creamy yel- 

low. Male ninth sternum pale vcntrally. 
Callus cerci fused to ninth tergum; ninth 
sternum partially coalescent with eighth 
(Fig. 17). Cuticular gland openings lack- 
ing on heavily sclerotized anteroventral area 
of ninth tergum, and tenth tergum. Medi- 
uncus (Figs. 18, 19) broad, hoodlikc; gono- 
coxites small, platelike. 

Measurements ( mm ) : Fore wing length 
12.5-15. (13.5); width 4.5-5.5 (5.1); an- 
tenna 10-12 (11.2). 

Distribution: CALIFORNIA: Alameda 
Co.: E. C. Van Dyke (USNM); Berkeley, 
28-IV-12, 9, J. C. Bridewell (USNM); Oak- 
land, 17-IV-15, E. P. Van Duzee (MCZ); 
Hills back of Oakland, 30-IV-1911, 6, 9- 
V-09, 6, E. C. Van Dyke (CAS). Contra 
Costa Co.: Near Orinda, 21-IV-50, W. Wade 
(CNHM). Marin Co.: Mill Valley, 28-11- 
26, 9, 6,7-111-26, 2 9, E. P. Van Duzee 
(CAS), 25-III-52, 2 6, 19, H. B. Leach 
(CAS), 8-V-56, E. S. Ross (CAS); Muir 
Woods, 23-IV-1911, 9, E. C. Van Dyke 
(CAS); San Geronimo, 20-IV, O. Sacken 
( MCZ ) . Mendocino Co. : Caspar Lumber 
Camp, 12 mi. E. Fort Bragg, 20-VI-38, 9 , 
Van Dyke ( CAS ) ; Van Damme State Park, 
18-V-47, i, H. Welsh (CAS); Yorkville, 
l-V-24, 9, E. P. Van Duzee (CAS). Santa 
Clara Co.: Stevens Cr. 16-III-41, 9, E. S. (CAS); Palo Alto, 27-V-92 (MCZ); 
Hills back of Palo Alto, 29-IV-1928, 9, 
(CAS); Stanford Univ. 5-IV-04, 9, (MCZ). 
San Mateo Co.: King's Mt., 5-VII-46, c^, 
E. S. (CAS). 

OREGON: Benton Co.: Corvallis, 19- V- 
45, 9, Marge Johnson (OSC); McDonald 
For., 5 mi. N. Corvallis, 26-V-56, 9, J. R. 
Mori (OSC); Sulfur Spgs., 6 mi. N. Cor- 
vallis, ll-V-56, 9, J. F. O'Brien (OSC); 
Rock Cr., 12 mi. W. Corvallis, 4-V-56, 9, 
N. E. Johnson (OSC); 1 mi. N. Dawson, 
2-V-62, 4 9 , 1 c5 , W. Barnett, ( OSC ) ; 2 mi. 
S.E. Summit, 12-IV-57, c? , J. Lattin (OSC). 

WASHINGTON: Kittitas Co.: Easton, A. 
Koebele (USNM). 

20-VI-55, 1 <^, 1 9, J. W. Brown (CNC); 
So. Pender Is., 30-\'-50, on Douglas Fir, 

Mesochrysinae and NoTHOCHRYSiNAE • Aclcims 225 

1 5, (CNC); Vancouver, 31-V-31, on snow, 
4700 ft., Seymour Mt., H. B. Leech (MCZ); 
Victoria, 4-V-lS, W. B. Anderson (Smith, 

Diagnosis. This species differs markedly 
from N. fulviccps and capitata in head 
coloration, in Psm extending beyond the 
basal inner gradates, and in the peculiar 
separation of the lateral pronotal sclerites 
by a median membranous area. It is re- 
tained in Notliochnjsa because it shares 
with the other species the well developed 
Psm and Psc. 

Discussion. The paucity of material from 
Oregon, Washington, and western Canada 
probably indicates less collecting activity, 
rather than scarcity. In the San Francisco 
Bay area it sometimes is locally fairly nu- 
merous. W. Wade (pers. comm.) collected 
eggs on conifers in the Orinda, California, 
area. D. Breedlove (pers. comm.) has noted 
adults abundantly on oak near Oakland, 
California. Throughout its range, it appears 
to be restricted to moist forest areas, not 
far from the coast. The type is probably 
mislabelled; in spite of intensive collecting 
around Los Angeles since 1892, no addi- 
tional specimens have been taken. 

Nothochrysa praeclara Statz 

Figure 43 

Nothochrysa praeclara Statz, 1936, Bechiana 93: 
215-216, fig. 3 (photograph of fore wing), 
fig. 4 (drawing of fore wing). Holotype, 
from Rott am Siebengebirge, West Germany, 
mid-Tertiary, in the Los Angeles County 
Museum ( examined ) . 

This species, known from a single fore 
wing, differs from capitata only in having 
fewer overlapping veins in Psm. In all the 
specimens of capitata examined, at least 
four branches of Rs + MA coalesce with the 
next proximal vein at the Psm, whereas 
only one basal branch is so coalesced in 
praeclara. This may not be significant; in 
fulviccps the venation is very irregular, and 
some specimens show no overlap at all. 
Praeclara differs from fulviccps in smaller 
size (fore wing length 16.33 mm, 16 
branches of Rs -f MA ) . 


Pimachrysa Adams, 1957, Psyche 63: 67-70. Type 
( by original designation ) : P. Virata Adams. 

Description. Vertex moderately elevated; 
anterior tentorial pits large; labrum barely 
emarginate. Antennal length variable: scape 
short, little swollen; apical flagellar seg- 
ments more than twice as long as wide. 
Tarsal claws simple, without basal enlarge- 
ment. Prosternum unsclerotized anteriorly 
to sternal pits (Fig. 34). Male with ninth 
tergite clearly separated from ectoproct; 
ninth sternite clearly demarked from eighth, 
posterior margin acute (Figs. 11, 14). 
Mediuncus slender, apex angulate (Figs. 
12, 13, 15, 16). Cuticular gland openings 
present. Female with enlarged ninth gono- 
coxites bearing marginally a band of spoon- 
shaped setae (Figs. 20, 21). Eighth sternite 
(subgenital plate) and ninth valvulae lie 
on membrane of genital opening, so that 
it is permanently closed anteriorly. Sper- 
matophore ovoid, protruding from gonocox- 

Fore wing: costal area narrow; basal sub- 
costal crossvein slightly basal to origin of 
Rs; Rs arising basally to cubital fork. MP2 
not fusing with MPl, but turning to wing 
margin at apex of intramedian cell. Second 
mediocubital crossvein near MF, usually 
distal to it. Two gradate series, extending 
smoothly into Psm and Psc (except in P. 
nigra); no overlap of adjacent longitudinal 
veins on Psm or Psc. Pseudomedial fold 
undeveloped except for weak spot at inter- 
section of first sectorial crossvein and MPl. 
Anals neither branched nor anastomosed. 
Jugal lobe prominent, with jugal vein. 

Hind wing: frenulum present. Costal area 
narrow. Rs and MP merely touching, or 
fused for a short distance; basal Banksian 
cell large. Psm and Psc similar to those of 
fore wing. Anals neither forked nor anas- 

Discussion. This genus is most similar 
to H\ipochnjsa, but differs in having en- 
larged ninth gonocoxites, bearing modified 
hairs, probably serving to hold the sper- 
matophore in place. In Hijpochnjsa, a 

226 BiiUctiii Miisciini of Compdiative Zoology, Vol. 135, \o. 4 

postcopulatory mass is placed on the dorsal 
surface of the female abdomen, the second 
mediocubital crossvein is more basal, and 
some anal veins are anastomosed; tl\e eyes 
are smaller in relation to the remainder 
of the head, than in PimacJirysa. 

Key to the Species of Pimachrysa 

1. A slight break lietween inner gradates and 

Psni; only 1 or 2 Ijranches of Rs forked 
marginally; color yellow-orange and 
black P. ninra sp. n. 

- Inner gradates merge smoothly with Fsm: 

nearly all branches of Rs forked mar- 
ginally 2. 

2. Head unmarked except for interantennal 

mark; body mostly pale yellow; stigma 
white P. ^rata Adams 

- Head with dark marks on face and xertex; 

subcostal veinlets of stigma conspicu- 
ously dark-bordered; body gray-prui- 
nose - 3. 

3. Discal veins of hind wing mostly black 

P. inicrmcdia sp. n. 

- Veins of hind wing (except Sc) all pale 4. 

4. Costal \einlets of fore wing pale; in hind 

wing MPl touches Rs + MA at one 
point only P. alhicostales sp. n. 

- Costal veinlets of fore wing black; in liind 

wing MPl coalesced with Rs + MA as 
far as length of first radial crossvein 
P. fusca sp. n. 

P'tmachrysa grata Adams 

Figures 9, 23 grata Adams, 1957, Psyche 63: 67-70 
(1956), figs. 1-5 (head and pronotum, geni- 
talia, wings ) . Holotype female, from Madera 
Canyon, Santa Rita Mts., Arizona, 26 August 
1949, at light. P. Adams, col. MCZ No. 29624, 

Description. Head (Fig. 9): straw yel- 
low, marked with black; flagelhim black. 
Pronotum ivory, with three fuscous stripes. 
Meso- and metanota yellow, scuta infus- 
cate posterolaterally. Meso- and meta- 
pleurae bright yellow, with fuscous marks. 
Legs ivory, femora with wide apical fus- 
cous band. 

Wings (Adams, 1957, Fig. 1) more acute 
th;in in fiisca; inner gradate series sinuous, 
so that in middle of series, the distance be- 
tween inner and outer gradates equals half 
or less the distance from inner 2;radates to 

Rs. Second medial crossvein of fore wing 
basal to MF. Most marginal veinlets forked. 

Fore wing: costa, subcosta and R white, 
stigma white; other veins fuscous except at 
wing base. 

Abdomen ( 9 ) short, slender, terminal 
segments greatly enlarged. Gonocoxites 
larger than in other species, with more 
hooked setae. Subgenital plate (Fig. 23) 
ligulate, with transverse ridge on ventral 
(anterior) surface. (The illustration, from 
Adams, 1957, shows a ventral view, with 
the distal margin uppermost. In the other 
species a posterior view is shown, with the 
distal margin below. ) 

Measurements (mm): Antennae 16; fore 
wing length 14. 

Diagnosis. Immediately distinguishable 
from fnsca by its pale body and venation. 

The following three closely related species 
are grey with prominent stigmas, and look 
much alike. 

Pimachrysa albicostales sp. n. 

Figures 27, 28 

HoJotijpe. Male from Mexico; Raja Calif., 
22 mi. N. of Punta Prieta, 9-XII-58, H. R. 
Leech, col., in the California Academy of 
Sciences. The specific name refers to the 
white costal veinlets. 

Description. Eight or nine black spots 
on apical subcostal crossxeins of fore wing, 
six or seven in hind wing. Intramedian cell 
of fore wing about four times as long as 
high, branches of Rs + MA more sharply 
inclined to Psm than in fusca, and cells be- 
t^\'een Rs and Psm longer. MPl of hind 
wing merely touches Rs + MA, not co- 
alesced over a long distance, as in fusca. 
Wing setae black; on dorsum of fore wing 
equal to length of longest costal veinlet. 

Abdomen pale, tergites 2-8 \\'ith triangu- 
lar dark marks (apices anterior); 9 with 
small mediodorsal dark rectangle, ectoprocts 
fuscous. Sternites light fuscous, paler medi- 
ally; ninth dark-lineate basally, dark api- 
cally. Apical beak-like extension of ninth 
sternite longer than in fnsca. Cuticular 
glands of basal segments sparser than in 

Mesochrysinae and Nothochrvsinae • Adams 227 

fusca. Gonarcus shorter, mediuncus (Figs. 
27, 28) longer, spoon-shaped distally, with 
apieal tooth. 

Measurements (mm). Fore wing 10.0- 
11.3 (10.6). 

Diagnosis. Head and thorax marked sim- 
ilarly to fusca, but paler. Wing veins colored 
as in fusca, but costal veinlets of fore wing 
pale. Membrane not brown in center of 
cells as in fusca. 

Record (Paratype). ARIZONA, Tucson, 
Tumamoc Hill, week ending 5-XI1-61, <5 , 
in ethylene glycol pollen trap, Sandra Ray 
Johnson, col. (MCZ, in alcohol). 

Pimachrysa fusca sp. n. 

Figures 1, 7, 11-13,21,24 

Holotype. Male from L. Covington Flat, 
Joshua Tree Nat. Mon., Riverside Co., 
Calif., 19-III-61, E. L. Sleeper, col. In the 
California Academy of Sciences. The name 
refers to the dark appearance of this species. 

Description. Head pale, marked with 
black ( Fig. 7 ) ; scape pale with medial and 
lateral black stripes; pedicel and flagellum 
fuscous. Pronotum about as long as broad, 
lateral margins strongly deflexed; no trans- 
verse groove; pale ivory with fuscous me- 
dian and marginal stripes; a short narrow 
longitudinal black stripe each side between 
the fuscous bands, connected to the median 
stripe posteriorly by a narrow transverse 
black band; black spot in posterolateral 
corner. Meso- and metanota with median 
and lateral fuscous stripes; mesoscutum 
pale near wing base, with black spots ( areas 
devoid of microtrichia ) on inner and outer 
margins of lateral stripe. Pleurae fuscous. 
Membrane anterior to prosternum pale. 
Legs pale, femora with dark preapical band; 
tibiae with dark bands at base, about two- 
fifths of distance to apex, and at apex. Tarsi 

Wing venation as in Figure 1. Fore wing: 
veins black except for C, apex of Sc, R, 
and bases of Cu and anals, white. Stigma 
pale, with 4-6 apical subcostal crossveins 
conspicuously black-bordered. Marginal 
area posterior to 3A black; membrane 

hyaline, usually brown-tinted in central 
portion of all cells. Intramedian cell about 
3 times as long as high. Setae on upper sur- 
face black, long, stiff, sparse. Hind wing: 
veins, except Sc, all pale; posterior mar- 
ginal black at intersections of marginal 
veinlets. MPl coalesced with MA + Rs for 
a short distance. Macrotrichia about one- 
third length of those of fore wing. 

Abdomen in male dark fuscous. Ninth 
sternite with a beak-like posterior projec- 
tion, fused with eighth sternite medially 
(Fig. 11). Pores of cuticular glands sparse 
on apical segments, on ninth sternite con- 
fined to anterolateral area. Mediuncus ( Figs. 
12, 13) widened between attachment of 
membrane and apical tooth. Female abdo- 
men mostly fuscous; ninth tergite pale lat- 
erally. Ninth gonocoxites ( Fig. 21 ) not so 
large as in P. grata, and with fewer hooked 
setae. Eighth sternite ( Fig. 24 ) with blunt, 
anterodorsally directed horn; a basal frag- 
ment present, shaped as a narrow trans- 
verse band. Ninth valvulae appear as slen- 
der rods. Spermatophore ( Fig. 21, sp ) ovoid, 
with cuplike apical indentation, and tubular 
process extending into genital opening. 

Measurements (mm). Fore wing length 
8.8-11.1 (10.3); maxillary palpus (apical 
three segments ) 0.67-0.83 ( 0.73 ) ; scape 
0.24-0.32 (0.28). 

Diagnosis. In shape of head and wings, 
and in venation, this species is closer to P. 
grata than to P. nigra. It may be recognized 
by the gray-pruinose body, smoky wings, 
prominent dark pterostigma, dark costal 
veinlets of fore wings, and pale veins in 
the hind wing. 

Records ( Paratypes ) . CALIFORNIA : 
Riverside Co., Joshua Tree National Monu- 
ment, L. Covington Flat #1, 19-III-61 6; 
#2, 5-III-61 9; #4, 25-XI-60 6, 5-XI-60; 
U. Covington Flat #1, 5-XI-60 i; Smith- 
water Wash #1, 25-XI-60 9, 4-I1I-61 9; 
all collected by E. L. Sleeper in molasses 
bait traps (Long Beach State College, MCZ, 
Adams Collection). Los Angeles Co.: Aliso 
Can. chapparal, 26-X-58 9, R. X. Schick 

228 Bulletin Museum of Comparative Zoology, Vol. 135, No. 4 

Pimachrysa infermedia sp. n. 

Figures 2, 20, 25 

Holotypc. Female from Snow Creek, 
1500', White Water, Rixerside Co., Calif., 
8-III-1955, W. R. M. Mason, col. Canada 
Department of Agriculture, Ottawa. The 
name means intermediate. 

Description. Head marked with black 
similarly to fmca, but dark border of an- 
tennal socket not connected to lateral ver- 
tex stripe, median \ertex mark larger, and 
genae black-bordered posteriorly. Maxillaiy 
palpi and scape both shorter than in fiisca. 
Pronotum longer than in fusca, pale, with 
lateral and median red-fuscous stripes, 
broader posteriorly; between these each 
side a brown stripe, extending to furro\\' 
(which is barely perceptible). Meso- and 
metanota dark; yellow spots laterally on 
prescuta and scuta. Propleura and cervical 
sclerites pale fuscous, prosternum dark- 
fuscous; membrane anterior to sternum pale, 
with median and lateral fuscous stripes, 
overlaid with a rufous transverse band. 
Meso- and metapleurae fuscous. Legs pale, 
femora with a broad preapical dark band; 
tibiae with thin dark band at about one- 
third distance from base to tip; fore and 
mid tibiae slightly infuscated, hind with 
two narrow dark stripes on ventral surface. 

Wing venation as in Figure 2; few mar- 
ginal veinlets forked. R (basal two-thirds) 
and 3A in both wings, and bases of lA, 2A, 
and Cu in hind wing, pale; other veins all 
dark. Stigma white, with dark clouds on 
apical subcostal crossveins; membrane prox- 
imal to end of 3A in fore wing fuscous. 

Abdomen fuscous, segments narrowly 
pale at anterior and posterior margins; mus- 
cle scars on tergites black, surrounded by 
pale areas. Female external genitalia sim- 
ilar to those of fmca; subgenital plate bears 
knob on basal margin, basal fragment much 
wider, and less sclerotized than in fusca 
(Fig. 25). 

Measurements ( mm ) . Fore wing 9.2; 
apical three segments, maxillary palpus, 
0.49; scape 0.21. 

Diagnosis and discussion. In the features 

by which this species differs from typical 
fusca — dark venation, smaller number of 
gradate cells, fewer forked marginal vein- 
lets, short palpi, color and shape of pro- 
notum — it grades toward nigra. Further- 
more, it was taken geographically close to 
a nigra locality. It may be a unique hybrid 
individual, or a representative of an intro- 
gressed population. 

Biology. The crop of the type is filled 
with Salix pollen (identified by D. Walk- 
ington, California State College, Fullerton). 

Pimachrysa nigra sp. n. 

Figures 6, 14-16, 26, 34 

Holotypc. Male from Gavilan, Riverside 
Co., Calif. 19-III-36, on Ceanothus crassi- 
folius, Timberlake, col. (California Acad- 
emy of Sciences ) . The name is descriptive 
of the dark coloration. 

Description. Head long, eyes small; color 
shades from ivory anteriorly, to orange on 
vertex; dark-fuscous markings as in Figure 
6; mark between antennae extending an- 
teriorly to frontal suture, and broadly over 
antennal sockets and vertex. Maxillary palpi 
short, dark, pale at joints. Occipital fora- 
men broadly dark-margined. Scape pale, 
broad medial and lateral dark stripes; ped- 
icel and flagellum dark. Pronotum long, 
brown-fuscous with t\\'o narrow submedian 
pale stripes. Meso- and metanota dark, two 
disconnected yellow stripes. Propleurae, 
sternum, and cervical sclerites dark; rufous 
median area on membrane anterior to pro- 
sternum. Meso- and metapleurae dark. Legs 
dark; femora pale basally and apically; 
tibiae (especially metathoracic) paler ex- 
teriorly, with apical dark band. 

Fore wing veins dark; bases of C, R, 
3A, and marginal vein in jugal area, pale. 
Stigma black; conspicuous ivory spots on 
membrane, between apical subcostal cross- 
veins. Hind wing similar, but R pale for 
two-thirds length. 

Venation ( Fig. 4 ) : A break in alignment 
of inner gradates and Psm in both wings; 
weak pseudomedian fold in both wings 
(especially hind), but no definite articula- 

Mesochrysinae and Nothochrysinae • Adams 229 

tions, as are found in NotJiochrysa; usually 
only 1 or 2 branches of Rs forked margin- 
ally ( 3 and 4 in hind wings of one female ) . 

Abdomen dark, segments narrowh' pale- 
banded posteriorly. Male 8th and 9th 
sternites, and 9th tergite-ectoproct more 
strongly fused than in fusca (Fig. 14). Many 
cuticular gland openings, except on ecto- 
proct. Margin of mediuncus between apical 
tooth and attachment of membrane, nearly 
straight ( Figs. 15, 16 ) . Female terminalia 
similar to those of fusca; 8th sternite (Fig. 
26) preceded by broad basal fragment, 
ventrally with blunt horn. Ninth valvulae 
reduced to dotlike sclerites. 

Measurements (mm). Fore wing 7.5-9.0 
(8.0) long; apical three segments, maxillary 
palpi, 0.48-0.53 (0.51); antennae 7-8 (7.5). 

Diagnosis. This small species may im- 
mediately be distinguished from all other 
species of Pimachrysa by its brown-black 
and orange markings. It strikingly resem- 
bles Nothochnjsa calif omica Banks, for 
which it has been mistaken by several 
workers, including the writer. In shape of 
head, and condition of pseudomedia, it is 
closer to Nothoclirysa than is any other 
species of Pimachrysa. However, these 
genera are structurally so different that the 
colorational resemblance is undoubtedly co- 

Records (Paratype). CALIFORNIA: 
Gavilan, l-IV-38, 2, on Ceanothus crassi- 
foUus, Timberlake, col. (MCZ). Pinnacles, 
San Benito Co., 23-111-40, R. L. Usinger, 
col. (CIS). Lebec, Kern Co., l-IV-39, 9, 
E. S. Ross (CAS). 

Figure 42 

Type species. Tribochrysa vetuscida 
Scudder. The feminine name means "diffi- 
cult-chrysopid," referring to the intricacy 
resulting from the superimposition of fore 
and hind wings in the fossil. 

Description. Basal subcostal crossvein 
distal to origin of Rs, opposite MF; Rs + 
MA originates far basal of MF. First sec- 
torial crossvein intersects MPl verv near 

MF. Intramedian cell long. Proximal branch 
of Rs -I- MA coalesces with MPl, and sec- 
ond branch coalesces with proximal branch, 
so that basal portion of Psm contains no 
crossveins. Psc strongly developed, two 
marginal veinlets from third medial cell, 
basal one forked; at least four additional 
non-forked marginal veinlets from Psc. Hind 
wing with zig-zagged Psc. 

Discussion. The fore wing base is similar 
to that of ArcJiaeochrysa, except for the 
overlap of branches of Rs + MA forming 
Psm. It is most interesting that the wing 
should demonstrate such a generalized 
structure of Rs + MA and the intramedian 
cell, while having a pseudomedia which, at 
least basally, resembles that of Nothochrysa. 
In this regard, Dyspetochrysa is more ad- 
vanced than all other Nothochrysinae, save 
Nothochrysa itself. 

Dyspeiochrysa vefuscula Scudder 

Figure 42 

Tribochrysa vetuscida Seudder, 1890, U. S. Geol. 
Geogr. Survey Terr., Kept., 13: 170, pi. 14, 
fig. 9. Holotype: Florissant, Colorado, Mio- 
cene; MCZ No. 245, tip of abdomen and distal 
half of fore wing missing: hind wing venation 
mostly not visible, examined. 

Paleochrysa vetusciila: Cockerell, 1908, Canad. 
Entomol. 40: 90. 

Paleochrysa stricta: Carpenter, 1935, J. Paleontol. 
9: 263 (not Scudder). 

Description. Head large, interocular 
width 1.0 mm, eye 0.5 mm. Scape short, 
flagellar segments 0.2 X 0.1 mm. Pronotum 
wide, anterior margin straight, corners 
slightly angulate; dark, a four-lobed pale 
central area; width 1.44 mm; length, ex- 
treme 1.0 mm, midline 0.7 mm. Costal area 
narro\\', tallest cell 0.63 X 0.55 mm. Intra- 
median cell 0.35 X 1.5 mm, pseudomedial 
area as wide as pseudocubital area. 


Figures 40, 41, 46, 47 

Type species. Paleochrysa creedei Car- 
penter. The feminine name means "old- 
chrysopid" referring to the archaic \enation. 

230 Bulletin Miificuw of Contpanitive Zoology, Vol. 135, No. 4 

Description. Fore wing: Basal subcostal 
crossvein distal to origin of Rs + MA op- 
posite MF. Rs + MA arises nearer to basal 
medial crossvein than to MF. Sectorial 
crossvein intersects MPl slightly distally to 
MF. Proximal branch of Rs + MA not 
coalesced with MPl, but connected by a 
gradate crossvein; Psm strongly zig-zagged. 
Psc well developed. 

Hind wing: Base of Rs + MA moderately 
long; not coalesced with MPl. Psm zig- 
zagged, with no overlap of veins. Psc poorly 
developed, strongly zig-zagged, so that the 
gradate series apparently extends to MP2; 
marginal veinlets from Psc all forked. 

Pronotum broad, squarish; cervical scle- 
rites and prosternum as in Pimachrysa. Ter- 
minal abdominal segments of male broad, 
short; ectoprocts forcipate (Fig. 46). 

Discussion. This is the most archaic of 
the Miocene genera. It is unique in its lack 
of fusion of MPl and Rs + MA in the hind 
wing. The unspecialized Psc of the hind 
wing is found elsewhere only in Faleo- 
chnjsa icickhami Cockerell. 

Archaeochrysa creedei (Carpenter) 

Figure.s 41, 46, 47 

Paleochnisa creedei Carpenter, 1935, J. Paleontol. 

9: 265, lis. 3; 1938, Psyche 45: 108. Holo- 

type: Creede, Colorado, Miocene, A. Caplan, 

coll.; MCZ No. 4316, examined. 
Paleochrijsa .striata: Carpenter, 1938, Psyche 45: 

108, fiji. 1. 

Description. Costal area narrow, 0.25 mm 
wide, cells about as high as long. Ten 
branches of Rs + MA in fore wing, and 7-8 
in hind wing. Gradate cells about twice 
as long as wide. Intramedian cell narrow, 
about 0.375 X 1.5 mm. Flagellar segments 
0.1.3 X 0.21 mm. Pronotum (Fig. 47) squar- 
ish, anterior margin not clear; 0.94 mm 
wide, 0.91 mm (0.625 mm on midline) long. 
Ectoprocts (Fig. 46) lateral, forcipate; 
gonarcus small. 

Discussion. The lateral margins of the 
conspicuous cervical sclerites apparently 
correspond to the pronotal margins in Car- 
penter's (1935) figure. 

The prosternum apparently was un- 
sclerotized. The basal Banksian cell is nor- 
mal for; on the type, the 
proximal branch of Rs + MA is very in- 
distinct, and was not shown by Carpenter, 
but is present on all the other specimens 
of this species. 

Records. Creede, Colorado, Miocene; 
MCZ No. 4462-4471. 

Archaeochrysa paranervis sp. n. 
Figure 40 

Paleochrysa vetiiseula Cockerell, 1908, Canad. 

Entomol. 40: 90. (Erroneous identification.) 
Paleoehnjsa .siricta: Carpenter, 1935, J. Paleontol. 

9: 263, fi.u. 1. (Erroneous identification.) 

Holotype. Univ. Colo. Mus. No. 4419, 
Florissant, Colorado, Miocene, expedition 
1907, examined. The name means "beside- 
vein," referring to the juxtaposition of the 
radial crossveins and the branches of the 
radial sector. 

Description. Venation as in Figure 40; 
fore wing broad; costal area broad; Sc ap- 
proaches C near stigmal base; 12 branches 
of Rs + MA, several branches opposite 
radial crossveins; longest gradate cell 2.5 
times as long as broad. Fore wing length: 
14.2 mm. 

Diagnosis. Differs from creedei in the 
length and arrangement of branches of Rs, 
and in width of the costal space. This speci- 
men was the basis for Carpenter's ( 1935 ) 
figure and redescription of FaJeochrysa 
strict a. 

Archaeochrysa fracfa (Cockerell) 

Paleochrysa fracta Cockerell, 1914, J. Geol. 22: 
716, fig. 2. Holotype: Florissant, Colorado, 
Miocene, H. F. Wickham, col.; MCZ No. 4501 
(formerly No. .3349), isolated fore wing, 
examined. Carpenter, 1935, J. Paleontol. 9: 
264, fig. 7, p. 268. 

Description. Costal area moderately 
wide, the cells 0.95 mm X 0.5 mm; veinlets 
inclined about 10°, 17 branches of Rs + MA, 
Rs hardly zig-zagged. Longest gradate cells 
3.1 times as long as wide; intramedian cell 
2.0 X 0.5 mm, acute proximally, distal cross- 
vein oblique. lA forked. 

Mesochrysinae and Nothochrysinae • Adams 231 

Diagnosis. This species differs from 
crecdei in the shape of the subcostal area, 
more branches of Rs + MA, longer gradate 
cells, and forked lA. 

Carpenter's figure of this specimen was 
inadvertently transposed with that of Tri- 
hochnjsa firmata, and appeared on page 
268, as his Figure 7. 

Genus PALEOCHRYSA Scudder 

Figures 38, 39 

Paleochnjsa Scudder, 1890, U. S. Geol. Geogr. 
Survey Terr., Rept., 13: 166. Type species 
(by nionot\pv') : Paleochnjsa stricta Scudder. 
Cockerell, 1908, Canad. Eutomol., 40: 90. 
Carpenter, 1935, J. Paleontol. 9: 262. 

Lithochnjsa Carpenter, 1935, J. Paleontol. 9: 265. 
Type species ( by original designation ) Paleo- 
chnjsa icickhami Cockerell. xew synonymy. 

Description. Basal subcostal crossvein 
distal to origin of Rs + MA; opposite MF. 
Rs + MA arises near MF; proximal branch 
of Rs + MA not coalesced with MPl, hence 
no overlapping veins in Psm. Gradate series 
run into Psm and Psc in an even curxe in 
both wings. Psc of hind wing either reg- 
ularly zig-zagged with forked marginal 
veinlets basally (P. wickhatni), or with 
furcations more proximal, so zig-zagging is 
irregular, as in Pimachnjsa, Hijpochnjsu, 
etc. (P. stricta). 

Discussion. The type of P. stricta has 
all four wings superimposed, making inter- 
pretation excessively difficult. But the short 
distal stem of Rs + MA, in the fore wing 
and the coalescence of Rs + MA with MPl, 
forming a quadrangular Banksian cell in 
the hind wing, are clear; thus this species 
has the essential characteristics of Litho- 
chnjsa Carpenter. Paleochrysa icickhami, 
the type of Lithochrijsa, does not differ 
from stricta in any important respect save 
the more archaic structures of Psc in the 
hind wing. The species of Pimachnjsa show 
such variation in the site of furcation of the 
marginal veinlets (e.g., cf. Figs. 1 and 2) 
that I do not consider this difference in 
structure of Psc of sufficient importance to 
warrant maintaining a generic distinction. 

Paleochrysa sfricta Scudder 
Figures 38, 39 

Paleochnjsa stricta Scudder, 1890, U. S. Ceol. 
C;eogr. Survey Terr., Kept., 13: 166, pi. 14, 
figs. 13, 14. Holotype: Florissant, Colorado, 
Miocene, S. H. Scudder; MCZ No. 242 a-b, 

(not) Trihochnjsa vettiscttla: Cockerell, 1908, 
Canad. Entomol. 40: 90. Carpenter, 1935, J. 
Paleontol., 9: 263. 

Description. Costal area moderately wide, 
cells 0.69 mm long, 0.875 mm tall. Twelve 
branches of Rs + MA. Gradate cells 0.5 X 
1.0 mm; intramedian cell 0.35 X 1.25 mm; 
8 unbranched veinlets from Psc in fore 
wing. In hind wing, 5 unbranched veinlets 
from Psc to wing margin; CuA 3-branched, 
the distal branch with a marginal fork. 
Fore wing length, 16.0 mm, width 5.0 mm. 

Paleochrysa wickhami (Cockerell) 

Paleochnjsa wickhami Cockerell, 1914, J. Geol. 
22: 717, fig. 3. Holot>pe: Florissant, Colo- 
rado, Miocene, H. F. Wickhani, col.; MCZ 
No. 4499, e.xamined. 

Lithochnjsa wickhami: Carpenter, 1935, ]. Paleon- 
tol., 9: 265, fig. 4. 

Description. Costal area moderately 
broad, cells about 0.46 x 0.5 mm. Winlets 
inclined about 15 degrees. Nine branches 
of Rs + MA in fore wing, and hind wing. 
Intramedian cell 0.313 X 0.88 mm. Gradate 
cells 5.25 X 0.75 mm. Costal space nar- 
rows abruptly before stigma, as in Llypo- 
chnjsa nobilis- C and Sc may be coalesced 
for a short distance. Radius runs in a 
smooth curve at wing apex, not angulate 
near stigma as in Hijpochnjsa. 

Eye fairly large (0.44 mm diam.). Pro- 
notum rounded anteriorly, sides straight, 

Paleochrysa concinnula Cockerell 

Paleochnjsa concinnula Cockerell, 1909, Canad. 
Entomol. 41: 218, fig. 5. Holotype: Floris- 
sant, Colorado, Miocene; Colo. Uni\ . Mus., 
not examined. 

Lithochnjsa concinntila: Carpenter, 1935, J. Pa- 
leontol. 9: 266, fig. 5. 

Paleochrysa ferruginea Cockerell, ibid.-. 218, fig. 6. 
Holotype: Florissant, Colorado, Miocene; Colo. 

232 Bulletin Museum of Ciunpaiative Zoology. Vol. 135, No. 4 

I'liiv. Mils., not ixaiiiiiR'cl. (^aipt'iitt r. Inc. 
(it.: 266-267. 

Genus TRIBOCHRYSA Scudder 

TrihtnluiiMi ScucUIlt, LS85, in Zittrl-Banois, Traite 
PaU'ont. 1 : 777. Type species ( by monotypy ) : 
Tiil>(>chrt)sa iudciinalis Scudder. Scudder, 
1890, U. S. Ceol. Ccogr. Surv. Terr., Kept. 
13: 168. 

Description. Eyes large (0.5 mm); .scape 
large, flagellar segments 0.15 x 0.25 mm. 
I'roiiotuin short, broad, margin arcuate. 
.Subcostal crossNciu obscured in all speci- 
lucns. Hs -f M.\ arises slightly basally to 
Ml'\ \o oNcrlap of veins in Inner 
gradates in two series, so that the fourth 
gradate (numbered from base) is much 
closer to Rs than is the second or third. Psc 
well de\eloped in fore wing. In hind wing, 
Hs + MA coalesces with MPl, and Psc is 
less well developed than in fore wing. 

Discu.ssion. The irregularity in the inner 
gradate series is a necessary precondition 
for separation from Psm, as has taken place 
in man\- Chrysopinac. It is, therefore, not 
surprising that .several other taxa exhibit a 
similar, but less emphasized break in the 
inner gradate series [Fimachrijsa ni<^ra (Fig. 
4), lliipoclinisa (Fig. 5), Notlioclinjsa cali- 
f arnica ( Fig. 3)]. 

Tribochrysa inaequalis Scudder 

Trihoclin/.sa ina((j\iali\ Scudder, 1S85, ;'/i Zittel- 
Barrois, Traite Paleoiit. 1 ; 777, fijf. 982. Holo- 
t\pe: Floris.sant, Colorado, Mioceue; MCZ 
.\o. 24.3, examined. Scudder, 1890, U. S. 
Ceol. Ceoyr. Surv. Terr., Rept. 13: 170. Coek- 
erell, 1908, Canad. Entouiol. 40: 90. Carpen- 
ter. 1935, J. i^deoutol. 9: 267, fig. 6. 

Tribochrysa firmafa Scudder 

Tribochrysa finiidta Si^udder, 1890, U. S. Ceol. 
Ceogr. Siu\. Terr., Kept. 13: 172, pi. 14, figs. 
6, 7, 10. 11. Cotypes: Florissant, Colorado, 
Miocene- MCZ No. 241, Xo. 4127, examined. 
Carpenter. 195.5. j. Paleontol. 9: 267, fig. 7. 


AuA.Ms, P. A. 19.57. .\ new genus and new 
species of Chr\sopidae from the western 
United States, with remarks on the wing 
xcnatinn of tlie famil\'. i'svehe 6.'i: 67-74. 

— . 1958. Studies in the Neuroptera, with 
special reference to wing structure and evolu- 
tion in the Osmyloidea. Ph.D. thesis, Har- 
vard University (unpubl.). 

1962. A stridulatory structure in Chry- 

sopidae. Pan-Pac. Entomol. 38: 178-180. 
Bk.\uer, a. 1867. Larve von Hypochnjsa nohilis 

Heyd. Verhandl. Zool.-Bot. Ces. Wien 17: 

27-29, pi. 9, fig. 1. 
CARPENTER, F. M. 1935. Tertiary insects of the 

family Chry.sopidae. J. Paleontol. 9: 259-271. 
Ehh.'\hi)t, E. 1916. Zur Kenntnis der Inner- 

vierung und der Sinnesorgane der Fliigel von 

Insekten. Zool. Jahrb. ( Anat. ) 39: 295-3.34, 

pis. 17-18. 
I'lUEDRicH, H. 1953. Neuroptera. Bronns Klas.sen 

u. Ordnungen des Tierreichs, 5 Band, 3 Al)t. 

Xll Buch, Teil a. Leipzig. 148 pp. 
KiLLiNGTON, F. J. 1937. A monograph of the 

British Neuroptera. Ray Society, London. 269 

pp., 15 pis. 
Ki.Mxnxs, D. E. 1952a. Some new Australian 

Chrysopidae. Ann. Mag. Nat. Hist. (12) 5: 

. 1952b. A revision of the genera of the 

Ap()chr\sinae ( fam. Chrvsopidae ). Ann. Mag. 

Nat. (12) 5: 929-944. 
M.AHrvxovA, O. 1949. Mesozoic lacewings 

( Neuroptera ) and their bearing on concepts 

of phylogeny and systematics of the order. 

(In Russian.) Trudy Paleontol. Inst., Akad. 

Nauk SSSR 20: 150-170. 
Nav.As, L. 1910. CriscSpidos nuevos. Broteria 9: 

. 1913. Les Chrysopides (Ins. Nevr. ) du 

Musee de Londres. Ann. Soc. Sci. Bruxelles 

37: 292-330; 38: 73-114. 
Phixcipi, M. M. 1946. Contributi alio studio dei 

Neurotteri Italiani. IV. Ni)tlu> italica 

Rossi. Boll. 1st. Entomol. Univ. Bologna 15: 

. 1958. Neurotteri dei Monti Sibillini. 

Mem. Mus. Ci\ . Storia Nat. Verona 6: 175- 

Scudder, S. H. 1890. The fossil insects of 

North America, with notes on some European 

.species. II. The Tertiary in.sects. Rept. U. S. 

Ceol. Ceogr. Surv. Terr.' 13: 1-734, pis. 1-28. 
Tn.EYAHi), R. J. 1916. Studies in Australian 

Neuroptera. No. 3. The wing-venation of the 

Chrysopidae. Proc. Linn. Soc. New South 

Wales 41 (2): 221-248, pis. X, X bis, XI. 
TjEUER, Bc). 1941. Some remarks on the generic 

names of the British Neuroptera. Entomol. 

Tidskrift (1941): 24-31. 
. 1966. Neuroptera-Planipennia. The lace- 
wings of Southern Africa. 5. Chrysopidae. 

South African Animal Life 12: 228-534. 
T()sc:iTi, C. A. 1966. The taxonomy, life his- 
tories, and mating beha\'ior of the green lace- 

Mesochrysinae and Nothochrysinae • Adams 233 

wings of Strawberry Canyon. Hilgardia 36: 

(Received 21 October 1965) 


b — basal Banksian cell; CUA — cubitus anterior; 
CUP — cubitus posterior; CV — cervical sclerite; CX 
— coxa; ect — ectoproct; fr — frenulum; g.c. — gradate 
cell; gcx — 9 , nintb gonocoxite ( gonopophyses 
laterales ) , i , gonocoxites ( = "paranieres," "ento- 
processus"); gs — gonarcus; hyp. i. — hypandriuni 

internum; i.g. — inner gradate crossvein; im — intra- 
median cell; J— jugal vein; jl— jugal lobe; MA— 
the proximal branch of the fused media anterior 
and Rs, presumably media anterior; memli — mem- 
brane; MF— medial fork, fork of MP; MP— media 
posterior; mu — mediuncus; o.g. — outer gradate 
crossvein, PL — propleuron; pni — paramere; psc — 
pseudocu!)itus; psm— pseudomedia; R— radius; RS' 
— proximal branch of radial sector; sp — spermato- 
phore; ST — sternum; SXV — first sectorial cross- 
vein; lA, 2A, 3A— anal veins; Im— first medial 

234 Bulletin'uni of Conijxiiative Zoology, Vol. 135, No. 4 

Wings: Fig. 1. Pimachryio lusca sp. n. Fig. 2. P. inlermedia sp. n. Fig. 3. Nothochrysa californica. Fig. 4. P. nigra sp. n. 
Fig. 5. Hypochryso nob/7is. Heads: Fig. 6. P. nigra. Fig. 7. P. fusca. Fig. 8. N. ca//7ornica. Fig. 9. P. gra(o. Fig. 10. 
K. raphidioides. 

Mesochrysinae and Nothochrysinae • Adatm 235 

Figs. 11-13. Pimachrysa fusco. Fig. 11. Mole abdomen, lateral view. Fig. 12. Gonorcus, lateral view. Fig. 13. Same, 
posterior view. Figs. 14-16. P. nigro. Fig. 14. Male abdomen, lateral view. Fig. 15. Gonorcus, posterior view. Fig. 16. 
Some, loterol view. Figs. 17-19. Nofhochryso calilornica. Fig. 17. Male abdomen, lateral view. Fig. 18. Gonorcus, lateral 
view. Fig. 19. Some, posterior view, and hypandrium Internum. Fig. 20. P. intermedia, setae on margin of right ninth 
gonocoxite, lateral view. Pig. 21. P. fusca, female abdomen, with spermotophore. Figs. 22-26. Female eighth sternum or 
subgenitoi plate. Fig. 22. N. californica. Fig. 23. P. grata. Fig. 24. P. fusco, anterior view below, posteroventral above. 
Fig. 25. P. intermedia, onteroventrol view. Fig. 26. P. ni'gro, onteroventral view. 

236 Bulletin Museum of Comparative Zoology, Vol. 135, No. 4 

Figs. 27, 28. P/machrysa olbicosfa/es, gonarcus, lateral and ventral views. Figs. 29-31. Kimochrysa ahicana. Fig. 29. 
Gonarcus, ventral view; Fig. 30. Same, lateral view. Fig. 31. Male abdomen, lateral view. Figs. 32, 33. H. nob///s. Fig. 
32. Male abdomen, lateral view. Fig. 33. Gonarcus, dorsal view. Figs. 34-36. Prothorax, ventral view, membranous areas 
stippled. Fig. 34. P. nigra. Fig. 35. N. calilornica, left coxa removed to show sternal apophysis. Fig. 36. Chrysopiella 
sp. (Chrysopinae). 

Mesochrysinae and NoTHOCHRYsiNAE • Acluuis 237 

— ■'::q =^=^-4=^ / V / 1 1 / 1 T ^ 

Fig. 37. Mesypochryso, hind wing (after Martynov). Figs. 38, 39. Paleochrysa ilricta, fore and hind wing bases. Fig. 40. 
Archaeochryio paranervis type, wings. Fig. 41. A. creedei, wings (from type, and from MCZ specimens No. 4464 and No. 

238 BiiUetin Museum of Comparative Zoology, Vol. 135, No. 4 

Fig. 42. Dyspefochfysa veluscula, type, fore wing base; arrows indicate position of basal subcostal crossvein, and point of 
fusion of MPl and MA. Fig. 43. Nothochrysa praechra, fore wing base. Fig. 44. Synfhochryso (Apochrysinoej, fore wing 
base, showing overlap of veins at Psnn, probable overlap at Psc and probable course of MP2 (modified from Kimmins, 1952b). 
Fig. 45. Gonzogo |Chrysopinae), fore wing base, showing overlap of veins at Psm, and course of MP2. Fig. 46. Archaeo- 
chryso creedei, tip of male abdomen, showing forcipate gonarcus (from MCZ specimen No. 4466). Fig. 47. A. creedei type. 

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Abstract 240 

Introduction 240 

Systematic Section 242 

Family Leptosomatidae 242 

Genus Anticoina 242 

Oxystomatidae 246 

Halalaimits 246 

Tripyloididae 247 

Halanonchus __ 248 

CijtoIaiDiium 249 

Bathylaimus 250 

Phanodermatidae 251 

Phanodcrmopsis 251 

Enoplidae 251 

Enoploides .._ 251 

Mesacanthoides 254 

Oncholaimidae 255 

Oncholaimus 255 

Oncholainihim 255 

Metoncholaimus 256 

Prooncholdimus 258 

Viscosia 258 

Enchelidiidae 260 

Eiinjstomina 260 

Illiiim 261 

Pohjgastrophora 262 

Cyatholaimidae 263 

Pomponcma 264 

Longicyatholaim us 265 

Xijzzors 266 

Paracanthonchus 267 

Paracyatliolaimus 268 

Halichoanolaimus 269 

Neotonchus 270 

Desmodoridae 270 

Spirinia 271 

S. (Perspiria) 272 

Chromaspirina 272 

Metachronuidoid 273 

1 Zoologisches Institut, Universitat Wien, Wien, 

- Nematology Section, Entomology Research In- 
stitute Research Branch, Canada Department of 
Agriculture, Ottawa. 

Paradesmudora 276 

Dcsmodoia 277 

D. ( Pseudochromadora ) 277 

Monoposthia 277 

Monoposthioides 278 

Microlaimidae 279 

Pcirainicmhiimus 279 

Chromadoridae 279 

Hypodontulaimus 279 

Rhips 281 

Chromadora 281 

Tinimia 281 

SpilophoreUa 282 

ProchromadorcUa 282 

Chiomadorella 282 

Euchrotuadora 284 

Atrochromadora 287 

Comesomatidae 287 

Mesonchiuin 287 

Sahatieria 288 

A.xonolaim idae 289 

Axonolainitis 289 

OdontopJiora 290 

Purodontophora 290 

Leptolaimidae 292 

Ahiimella 292 

Camacolaimidae __ 292 

Camacolaimits 292 

Diplopeltidae 292 

Parataivaia 292 

Didelta 293 

Linhomoeidae __ 293 

Terschellingia __ 293 

Monhysteridae 295 

PanimonJiystera 295 

Stcincria 295 

Thcristiis _____ 296 

T. ( Penzancia ) 296 

T. ( Daptonema ) 298 

T. ( Trichoiheristus ) _ _____ 298 

T. (Cyliudrotheristii.s) 300 

Monhystcra __ 302 

Scaptrella ___. 303 

Xenolainius 303 

Acknowledgments ___ 303 

References Cited 304 

Table 1, List of collecting sites in Florida __._ 242 
Talkie 2, List of species reported from Florida _ 244 

Bull. Mus. Comp. ZooL. 135(5): 239-344, April, 1967 239 

240 Bulletin Museum of Comparative Zoology, Vol. 135, No. 5 


A beginning is made on a series of 
papers in which the nematode fanna in- 
habiting the east coast of North America 
from Florida to Newfonndland is to be de- 
scribed. The present paper, deahng with 
90 species, extends the number of species 
known from Florida to 118. One new genus, 
Paratarvaia within the Diplopeltidae, and 
48 new species, are described, viz.: Hala- 
laimus meyersi, Bathyhimus aithropappus, 
Enoploidcs ])isulciis, E. ij^ryplius, Mcsa- 
canthoidcs fiJ)uJatiis, M. psittacus, Mcton- 
cholaimus intermedins, M. simplex, M. 
scis.m.s, Prooneholaimus hosiatus, Viscosia 
onchohimelloides, lUium lihidinosum, Poly- 
il(i.stro})liora eddx, Pomponema tesselotum, 
Loniiieyatholaimus annae, Xyzzois inii,Iisi, 
Paracanthonehus platypus, Paracyatholai- 
muspesavis, Spirinia hamata, Chromaspirina 
inaurita, Metochwmadora pulvinota, M. 
meridiana, Paiadesmodora toreutes, Mono- 
posfJiioides mayri, Paramiewhiimus luna- 
tus, Uypodontolaimus intcrruptus, Chwma- 
dorella tiilix, C. vanmeterae, Euchromadoni 
pectinata, E. meadi, Atrochwmadom dentic- 
idata, Sabatieria paradoxa, S. poracupida, 
AxomtJaimus Iwxapihis, Odontophoni varia- 
bilis, Paratarvaia seta, Tersehellingia Jongi- 
spieuhita, T. monohystcra, Paramonhystera 
canicida, Steineria ampullaeea, Thcristus 
calx, T. osientator, T. floridanus, T. e rectus, 
T. galeatus, T. fistidatus, T. tortus, and T. 
.xyaliformis. The Ualanonchinae n. sub- 
fam., with UaJanonchus Cobb, 1920, Rhab- 
docoma Cobb, 1920, and Cytolaimium 
Cobb, 1920, is created within the Tripyloidi- 
dae. The Halanonchinae, via Trefusia, is 
considered to link the Tripyloididae with 
the Oxystomatidae. 


The nematodes from the coasts of the 
American continents are poorly known, a 
fact that is regrettable since this group 
represents such an important component 
of the fauna of every marine habitat. 

To improve this situation we decided to 

collect material towards a monograph on 
the marine nematodes inhabiting the east 
coast of North America. In order not to 
overburden this monograph with descrip- 
tions of new species and with taxonomic 
discussions, we shall publish the results of 
our investigation in a numbers of papers 
each dealing with a portion of coastline 
between Florida and Newfoundland. In 
each paper, descriptions and figures will be 
given for all the species found except for 
those in which agreement between our 
specimens and representations in the litera- 
ture were considered very good. In addi- 
tion, a list containing all species reported 
from the particular region will be included. 

In the monograph, the information con- 
tained in the special reports will be con- 
densed. Short clescriptions and figures will 
be given of all species, together with keys 
to most of the genera and families of ma- 
rine nematodes. 

A portion of the material for this under- 
taking was collected separately by us on 
excin\sions to the eastern seaboard during 
the period 1958-1961, or it was sent to us 
by various colleagues whose cooperation 
we shall acknowledge in the relevant sec- 
tions of this series. The bulk of the ma- 
terial, however, was collected by us be- 
tween May 7 and June 8, 1963, on a trip 
in which we covered the area from south- 
ern Florida to Maine. 

The following remarks should suffice to 
clarify our approach to the whole project: 

Habitats. — At each location we tried to 
cover the important types of habitat that 
could be reached by manual sampling. 
Shallow water dredge or core sampling 
was carried out in Miami, Florida, Beau- 
fort, North Carolina, Lewes, Delaware, and 
Woods Hole, Massachusetts. In addition 
we received shallow or deep water samples 
from various sources which we shall 
acknowledge later. 

Methods. — In general, nematodes were 
extracted from the substrate utilizing a 
combination of the sieving and decanting 
processes. Live nematodes were relaxed in 

Florida Marine Nematodes • Wieser and Hopper 241 

an oven maintained at 54 "C prior to fixa- 
tion in 27c formaldehyde. As the number 
of specimens recovered in most instances 
was considerable, only a representative 
fraction of the total was picked out and 
processed into dehxdrated glycerine. The 
remainder of the material was also pre- 
served and is being kept for reference in 
the Nematology Section, Entomology Re- 
search Institute, at Ottawa. 

The dehydration process used was an 
adaptation of Seinhorst's methyl alcohol 
method (Seinhorst, 1959). The preserved 
specimens were placed in a B.P.I, watch- 
glass containing 107f glycerine in methyl 
alcohol. The watchglass was then placed 
in an oven maintained at 54°C to hasten 
the evaporation of the methyl alcohol, a 
process which took less than 30 minutes. 
The watchglass containing the specimens 
was then transferred to a desiccator for 
24-48 hours to insure complete dehydration 
of the glycerine prior to the construction of 
slides. Nearly all marine nematode species 
do well when subjected to this dehydration 
process. Sporadic distortion of varying in- 
tensitv results with certain members of the 
Desmodoridae and a few isolated genera 
of other groups. 

Systcmotics. — When we write the planned 
monograph we hope to be able to arrange 
the species and genera of marine nematodes 
into a more satisfactorv classification than 
has been in use so far. For the special re- 
ports, however, we shall adhere to the old 
Filipjev-Micoletzky system as used by 
Schuurmans-Stekhoven (1935), with a few 
improvements suggested by Chitwood, in 
Chitwood and Chitwood (1950), and Chit- 
wood ( 1951 ) . We shall proceed only from 
the family downward, ignoring the position 
and status of higher systematic categories. 

Descriptions. — W^e feel that too much 
\^'eight has been attached in the past to the 
use of formulas in the descriptions of nema- 
tode species. Purely relative formulas like 
those suggested by de Man or by Cobb are 
of little value. Formulas in which absolute 
dimensions are used suggest a degree of 

precision in the construction of nematode 
species, which is fairly unrealistic. We 
shall, therefore, in our descriptions give as 
many absolute measurements as possible 
of body and organ dimensions but refrain 
from assembling them into formulas. In- 
stead, we would like to stress the need for 
giving as precise a representation as pos- 
sible of the morphology of the animal, es- 
pecially of the stmctiu-e of the male genital 

Various authors (i.e., Wieser, 1955, Chit- 
wood, 1960, Inglis, 1962) have drawn at- 
tention to the necessity of the presence of 
a male specimen of a species as a prerequi- 
site to a valid description of the species. 
Inglis (1962) goes as far as to state, ". . . 
I will not, in general, accept identifica- 
tions based solely on females or larvae and 
I will insist on treating almost all species 
based originally on females or larvae alone 
as species chi])iae." We are in complete 
sympathy with this view so far as the "lar- 
vae" are concerned. With regard to the 
female, however, we must support the con- 
cept with certain limitations. If such a rule 
were to be followed with no exceptions, 
some genera would receive little or no at- 
tention, e.g., Illiiim in this paper. Males 
are scarce or unknown in certain groups, 
particularly the Plectoidea, and the ad- 
vancement of our knowledge of these 
groups could be curtailed if Inglis' sug- 
gestion were adhered to without reserva- 
tion. Therefore, we feel that the action 
suggested by Inglis might best be limited 
to those genera in which males are com- 
monly known to occur. 

In additi(m to species descriptions and 
figures, general discussions and keys will 
be supplied wherever necessary. 

Florida collection. — The Florida coast- 
line is one of the least known with respect 
to the nematode fauna of all the regions 
investigated. Only Cobb (1920, 1922), 
Chitwood (1951,' 1956), and Hopper 
(1961a, 1966) have reported a few species, 
mainlv from Miami and Kev \\est. Our 

242 Bulletin Museum of Comparative Zoology, Vol. 135, No. 5 

Table 1. Lisi of Collecting Sites in Florida 




Kt.'\- Biscayne, Bear Cut area; about low water level; seaweeds scraped off mangrove 

roots. Canadian National Collection of Nematodes No. 4066. 

Key Biscayne, Bear Cut area; shallow water (about 20 cm at low tide), close to 

submerged patch of'm and Syrini^odium; fine sand and debris. C.N.C. of N. 

No. 4067. 

Key Biscayne, Bear Cut area; flat around high-water level, with Uca; fine to medium 

sand, debris. C.N.C. of N. No. 4068. 

Off Rickcnbacker Causeway; very soft mud, stagnant water at low tide. C.N.C. of 

N. No. 4069. 

Virginia Key; beach on grounds of Marine Institute; clean sand, mid-tide level. 

C.N.C. of N. No. 4070. 

Everglades National Park, Florida Bav near Flamingo; clavish mud with a little 

sand; below low-tide level. C.N.C. of N. No. 4071. 

Everglades National Park, Florida Bay near Flamingo; upper part of shore; sandy 

mud. C.N.C. of N. No. 4072. 

Biscayne Bay, dredgings in Tliala.ssia beds, about four meters deep; sand, shells and 

mud; several subsamples. C.N.C. of N. No. 4073. 

Lauderdale-by-the-Sea; verv exposed beach around mid-tide level; clean, coarse 

sand, shells. C.N.C. of N. No. 4074. 

Vero Beach. Sheltered, muddy sand, with Uca and mangroves nearby. C.N.C. of 

N. No. 4075. 


























collection increases the total of species 
known to 118. 

Most of onr collecting was carried out in 
Biscayne Bay, in the vicinity of the Insti- 
tute of Marine Science of the University of 
Miami, Virginia Key, Miami; additional 
collecting sites were in the Everglades, 
near Lauderdale-by-the-Sea, and near Vero 
Beach. A list of the collecting sites is given 
in Table 1, and a list of all the species 
found, together with the species reported 
in the literature, in Table 2. 

ANTICOMA Bastion, 1865 

Type species.— Anf/coma eberfhi Bastion, 
1865: 141, pi. 11, figs. 143-145. 

In this genus it appears possible to dis- 
tinguish two groups of species character- 
ized by the position of the excretory pore 

^ Abbreviations used in this paper are as fol- 
lows: a.b.d., anal body diameter: c.b.d., corre- 
sponding l)od\- dianictcr: L, length; \'u, \'ul\a: 
w, widtli. 

and the length of the terminal excretory 
duct. The correlation between these two 
characters is fairly well established, al- 
though, due to a few insufficient descrip- 
tions, there remains some doubt as to its 
general apjDlicability. The two groups (A 
and B ) coincide, respectively, with group 
A and groups B + C in Wieser (1953). 
Filipjev ( 1927 ) had previously arranged 
the species of this genus in a similar man- 

Characteristics of Species of 
Anticoma Group A 

Excretory pore situated between poste- 
rior cervical setae and nerve ring; terminal 
excretory duct short ( never longer than the 
width of the excretory ampulla ) . 

Key to Species of Anticoma Croup A 

1. Gubernaculum absent -.. A. daJiIi Wieser, 19.53 

A. major .\hrwson, 1956 
Gubernaculum present 2 

2. Excretory pore a short distance behind 

cervical setae — 3 

E.xcretory pore about halfway between cervi- 
cal setae and nerve ring 4 

Florida Marine Nematodes • Wicscr and Hopper 243 

Excretory pore a short distance in front of 
nerve ring 6 

3. Tail short A. camphcUi Allgen, 1932 

Tail long, filiform ._ A. filicaiida Mawson, 1956 

4. Longer cephalic setae 24-25 ij. = about one 

head diameter long 

— A. eberthi Bastian, 1865 

Cephalic setae less than 10 ^ = one-half to 
two-thirds of head diameter long 5 

5. Cuticularized portion of spicula ( excluding 

velum ) more or less parallel 

A. lata Cobb, 1898 

Cuticularized portion of spicula strongly- 
dilated near proximal end 

A. cohimha Wieser, 1953 

Syn. A. aiistrulls Mawson, 1956 

6. Supplement opening posterior to proximal 

end of spicula ___ , A. Utoris Chitwood, 1936 
Supplement opening distinctly anterior to 
proximal end of spicula 7 

7. Amphids one-third to one-fourth of cephalic 

diameter wide A. trichina Cobb, 1891 

Amphids one-sixth of cephalic diameter wide 8 

8. Tail short (4-5 a.b.d.) 

A. insulaealbae Filipjev, 1927 

Tail long (8-10 a.b.d.) 

A. tenuicaudata Filipjev, 1946 

Others. A. typica Cobb, 1891 is related 
to the above three species but too incom- 
pletely known for its position to be deter- 
mined with certitude. 

A. kergiielensi^ Mawson, 195Sb is closely 
related to A. lata, perhaps even identical. 

Characteristics of Species of Anticoma 
Group B 

Excretory pore situated on a level with 
or in front of cervical setae; terminal excre- 
tory duct long ( at least as long as the width 
of the excretory ampulla). 

The species within this group will not be 
differentiated in this work. However, for 
the reader's convenience, a list of the recog- 
nizable species, including the most recent 
synonymy, is given: 

A. acuminata (Eberth, 1863) (Syn.: 
Odontobius acuminatus Eberth, 1863; 
Stenolaimus lepturus Marion, 1870; Anti- 
coma JimaJis Bastian, 1865 pt.; A. tijrrhenica 
de Man, 1878; A. caheti de Rouville, 1903; 
A. pontica Filipjev, 1918; A. zosterae 
Schulz, 1932; A. similis Cobb, 1898 [see 
Gerlach, 1962]; and A. profunda Mico- 

letzky, 1930 [see Gerlach, 1962]); A. pel- 
hicida Bastian, 1865 (Syn. A. hmilis Bastian, 
1865 pt.); A. subsimilis Cobb, 1914 (see 
Mawson, 1958); A. arctica Steiner, 1916 
(Syn. A. procera Micoletzky, 1930 [see 
Gerlach, 1962]); A. minor Filipjev, 1927; 
A. murmanica Filipjev, 1927; A. extcnsa 
Wieser, 1953; A. stekhoveni Wieser, 1953 
(Syn. A. acuminata of Schuurmans-Stek- 
hoven, 1950, nee Eberth, 1863); A. iciescri 
Mawson, 1958 (Syn. A. stekhoveni Maw- 
son, 1956, nee Wieser, 1953). 

AnVicoma lata Cobb, 1898 
Plate I, fig. 1, a-e 

Anticoma lata Cobb, 1898: 384, 385. 

Anticoma ditlevseni Micoletzkv, 1930: 255-258, 

fig. 2. 

L = 2.04-2.06 mm; w = 58 /x; esophagus 
390-412 ii; tail 215-220 ^ ( 9 , 6.5 a.b.d., 
6, 5.5 a.b.d.). Cephahc setae 4-4.5 + 3- 
3.5 /x. Buccal cavity conical. Amphids 3-4 
p. wide, 11 JUL behind anterior end. Cervical 
setae 33 fx behind anterior end. Excretory 
pore 115 /J. from anterior end. Spinneret 
delicate and pointed, directed dorsally 
(always?). Spicula 61-70 ^u, dorsal and 
ventral contours nearly parallel, \'elum 
present. Gubernaculum 21-24 fi. Supple- 
ment 56-70 IX in front of anus. 

Representation in samples studied. — M- 
1, Key Biscayne. 

Gcoiiraph ical distribution. — Australia 
(Cobb, 1898), Sunda Islands (Micoletzky, 
1930), Maldives (Gerlach, 1962), Red Sea 
( Gerlach, 1958c ) . 

Remarks. — The spicula are somewhat 
longer than reported by Micoletzky and 
by Gerlach ( 61-70 /j. as against 40-47 p. ) , 
but our specimens seem to agree in all other 

Anticoma frichura Cobb, 1898 
Plate I, fig. 2, a-d 

Anticoma trichura C()l)b, 1898: 385, 386. 

L = 3.01-3.04 mm; w = 49-52 ^; esopha- 
gus 390-445; tail in 9 , 530-565 /x ( 18 a.b. 
d.), in i, 460-480 p. (14 a.b.d.). Cephalic 
diameter 18-19 /x. Cephalic setae 15 + 8 /x. 

244 Bulletin Museum of Comparative Zoology, Vol. 135, No. 5 

Table 2. List of Free-lring Nematodes Reported from the Coasts of Florida (Numbers of 

Specimens found are given in parentheses)^ 


Auticoma lata Cobb, 189S 

A. trichura Cobb, 1898 

Ualalaimus cf. fleichcri Mawson, 1958 
//. mcyersi n. sp. 

H. pachydermotus (Cobb, 1920) Syn. Tijcnodora ]>. 
Poiocoina strkita Cobb, 1920 
Litiniuin acqitalc Cobb, 1920 

Hcihinonrlius niacruni.s- Cobb, 1920 
Ciitolaimium exile Colli), 1920 
Bathylaimus australis Cobb, 1893 

B. arthropappus n. sp. 

Thanodermopsis longisetae Chitwood, 1936 

Enoplnide.s l)i.sulciis ii. sp. 
E. gryphus n. sp. 
Mesaeanthoides fihulattis n. sp. 
M. psittacus n. sp. 

Tri.ssoiirhulus oceami.s Cobb, 1920 

Anoplostoma heterunmi (Cobb, 1914) 

Syn. Oncholaimelhi^s- h. 
A. vwiparum ( Bastian, 1865 ) 

Syn. Symplocostoma v. 
Oiiehohiiniiis dujdrdinii de Man, 1878 
Onchokiiiiiiuin iippeadiculatiim Cobb, 1930 
O. domesticuni (Chitwood & Chitwood, 1938) 
Metoiuholaimiis inienuedius n. sp. 
M. s-hnplex n. sp. 
M. .S'cm-u.y n. sp. 
Pwourholaimus- hastaius n. sp. 
Vlscosia onchohiimelhndes n. sp. 
\". papillata Chitwood, 1951 
V''. macramphida Chitwood, 1951 

Eurysiomina mintitisctdae Chitwood, 1951 
lUium exile Cobb, 1920 
/. lihidiitosiim n. sp. 
Pnlygasirophora edax n. sp. 
Calyptronema eohhi W'iescr, 1953 

Syn. Caldhdiuu-s aeuminatus Cobb, 1920 

Ponipoiieiiia tcs.selatum n. .sp. 
Longieyathohiiiinis annae n. sp. 
Xyzzors inglisi n. sp. 
Paracanthnnchus ]il(itypu.s ii. sp. 
P. trtincatus (Cobb, 1914) 

Syn. Cyatholavmus t. 
Pararydtholdirnus pesdri.s n. sp. 
II(di(li()diii)lainiu.s cpidlhiordecittipapilLittis 

Chitwood, 1951 
H. duodecimpapillatiis Timm, 1952 
Neotonchiis liitosus Wiescr and Hopper, 1966 
Syiiouehiinn ohtu.sum Cobb, 1920 





Cobb, 1920, Key West 
Cobb, 1920, Biscayne Bay 
Cobb, 1920, Miami 

M-8(2), Cobb, 1920, Biscavne Bay 
M-2(l), Cobb, 1920, Biscayne Bay 



M-2 (13) 

Cobb, 1920, Miami 

Chitwood, 1951, Ocala 
Hopper, 1961a, Panama City 


M-l(l), M-2(l) 






M-3(4), M-8(16) 

M-7(5), M-8(l), V(6) 

M-2(S), M-8(23) 

M-l(4), M-7(l) 
M-8( 1 ) 
M-8(6), V(3) 

Cobb, 1920, Biscayne Bay 


M-3(13), M-8(l) 



Chitwood, 1951, Silver Springs 


M-2(3), M-6(7), M-8(l) 



Cobb, 1920, Miami 

Florida Marine Nematodes • Wieser and Hopper 245 


Spirinia parasitifera ( Bastian, 1865) 

S. hamata n. sp. 

CJtronui.spirina inaurita n. sp. 

Metachromadora piilvinata n. sp. 

yf. onijxoides Cliitwood, 19.36 

M. nicridUina n. sp. 

Paradesnwdora toreutes n. sp. 

Desmodora qiiadripapiUata { Daday, 1899) 
Syn. Fseiidocdiromadora (j. 

Xennella cephaluta Cobb, 1920 

LcptoncmcUa cincta Cobb, 1920 

Monopu-sthia mirahilis Schidz, 1932 

Monoposthioides maijri n. sp. 

ParamicroJaunus lunatiis n. sp. 

Hypodontolaimus interruptus n. sp. 

H. pandispicuhtiis Hopper, 1961 

Rhips ornata Cobb, 1920 

Actinonemo pachijdermata Cobb, 1920 

Chromadora nuiciolaimoides Steiner, 1915 

Timmio parva (Timm, 1952) 
Syn. Parachromadora p. 

Spilophorclh paradoxa (de Man, 1888) 

ProchromadorcUa mcditerranca ( Micoletzky, 1922) 

Chwmadorella filifonnis (Bastian, 1865) 

C. trUix n. sp. 

C vanmeterac n. sp. 

Euchromadora gaulica Inglis, 1962 

E. pectinata n. sp. 

E. mcadi n. sp. 

Atrocliioinadora denticulata n. sp. 

Mesonchiitm peUiicidum (Cobl\ 1920) 
Syn. Pepsonema p. 

M. poriferum Cobb, 1920 

Sahaticria paradoxa n. sp. 

S. paracupida n. sp. 

Laimella longicauda Cobb, 1920 

Nanunlaimus giittatu.s Cobb, 1920 

Axonolaimus hexapihis n. sp. 

Odontophom variabilis n. sp. 

Parodontophora hrevamphida (Timm, 1952) 

Araeolaimus punctatus (Cobb, 1920) 
Syn. Coinonema p. 

Alainiclla cincta Cobl?, 1920 

Cyniira iiniformis Cobb, 1920 

Haliplectus floridanus Cobb in Cliitwood, 1956 

H. hickncri Chihvood, 1956 

Camacolainnis prijtherchi Chitwood, 1933 

Onchium oceUatum Cobb, 1920 

O. metocellatum Wieser, 1956 

Syn. Onchidella ocellata Cobb, 1920 

Neiirelki simplex Cobb, 1920 

M-4(10), V(.55) 




M-2(l), V(103) 


M-2(l), M-3(l) 
Cobb, 1920, Key West 
Cobb, 1920, Miami 


M-l(3), M-3(2), M-5(17) 

M-3(25), M-7(2) 

M-5(29), Cobb, 1920, Miami, Key West 

Cobb, 1920, Key West 

M-l(.30), M-4(9), M-5(l), M-7(l), M-8(3; 

















, V(2) 

M-3(l), M-6(5), M-7(2), 



Cobb, 1920, Key West 

M-7(2), V 


Cobli, 1920, Biscayne Bay, Key West 

Cobb, 1920, Biscayne Bay 




Cobb, 1920, Biscayne Bay, Key West 

M-2(l), Cobb, 1920, Biscayne Bay 

Cobb, 1920, Miami 

Chit\\ood, 1956, Long Key 

Clritwood, 1956, Atwood Gro\e, Ellenton 


Cobb, 1920, Key West 

Cobb, 1920, Biscayne Bay 
Cobb, 1920, Key West 


Bulletin Miiscuni of Comparaiive Zoology, Vol. 135, No. 5 

loncnui cohhi ( Steiner, 1916) 

Syn. loiicDui ocrllatiiiii Colili, 1920 
Nemclla uccllatu Cobb, 1920 

Pdnitarvaia seta n. gen., n. sp. 

Didclta maculatinn Cob):), 1920 

Terschclliii^id lon<iicaud(it(i dc NFan, 1907 

r. lon^ispiculata ii. sp. 

T. iijonohijstera n. sp. 

Aiifici/dtlnis tcntiicandatiis Colili. 1920 

lluUuciua spinosuin Cobb, 1920 

Pandinhomoeus fuscacephalum (Colil), 1920 i 
Syn. Crysttdloneina f. 

P. .simdc (Cobb, 1920) 
Syn. Cry.stallonema s. 

Linhomocihi rxilis Cobb. 1920 

Paramonhy.stera canicula n. sp. 

Stcincrifi ampidlacca n. sp. 

Thoi.stus tiictaflcvcn.sis Gerlach, 1955 

T. calx n. sp. 

T. poliichactophilus Hopper, 1966 

T. ostcntator n. s\}. 

T. floridaniis n. sp. 

T. erectiis n. sp. 

T. galcatus n. sp. 

T. oxyuroides ( Schuurmans-Stekhoven, 1931' 

T. fistidatus n. sp. 

T. tortus n. sp. 

T. xyaliformi.s n. sp. 

Monhystcra parva (Bastian, 1(S65) 

SvaptrcUa c'lucta Cobl), 1917 

Xenohiiniii.s .stiiatus Col)b, 1920 

GreefficUa dasyura Colib, 1922 

Cobb, 1920, Biscayne Bay 



Cobb, 1920, Key West 

M-4(l), M-6(13), M-7(89), M-S(l 

M-7(24), M-8(l) 



Colib, 1920. Miami 

Cobb, 1920, Biscayne Bay 

Colili, 1920, Miami 

Cobb, 1920, Miami 

Cobb, 1920. Biscayne Bay 

M-l(l), M-3(6) 



M-4(3), V(33) 

Hopper, 1966, Virginia Key 



M-2(3), M-3(S3), M-4(l), 



M-2(l), M-4(19), M-5(l), 





M-l(7), M-7(l) 


M-2(9), Cobb, 1920, Bisca) 

/ne Bay 

Cobb, 1922. Biscayne Bay 

1 The niiinber of specimens indicated in this table represents only those that were present in the fraction of the total 
sample examined. In this manner the relative abundance of each species is somewhat suggested. In some cases, where 
a species was represented by only a few specimens, an effort was made to locate additional material from the preserved 
remainder of the sample. These additional specimens are not recorded in this table. Species for which the number of 
specimens is not indicated were not iiresent in the original fraction examined. These were subsequently recovered from 
the remainder of the sample. 

Amphids 5 /x wide = one-third of body 
diameter. Buccal cavity conical. Cervical 
setae 35 /x behind anterior end. Excretory 
pore 170 p. behind anterior end, i.e., 85% 
of distance anterior end to nerve ring. 
Spicula 59-63 ^, proximally bent, dorsal and 
ventral contours nearly parallel, with slight 
hump near proximal end. Velum present, 
smooth (not striated as figured by Gerlach, 
1962). Cubernaculum about 20 /x. Supple- 
ment 55-60 ji in front of anus. 

Representation in samples studied. — M-2, 
Key Biscayne. 

Geoii^raphical distribution. — Australia 
(Cobb, 1898), Sunda Islands (Micoletzky, 

1930), Maldives (Gerlach, 1962), Antarc- 
tic, Subantarctic (Mawson, 1958b, Allgen, 

HALALAIMUS de Man, 1888 

Type species.— Ho/a/a/mus gracilis de Man, 
1888: 3, 4, pi. 1, fig. 1. 

A key to the species of this genus, ex- 
cluding the subgenus PacJu/odora, was given 
by Mawson ( 1958b ) . 

There are three species which are set 
apart from the rest by the occurrence of a 
distinct circle of labial setae, i.e., H. papil- 

Florida Marine Nematodes • Wieser and Hopper 247 

lifer Gerlach, 1956, //. fletcheri Mawson, 
1958, and H. filicoUis Timm, 1961. In the 
first species the tail is rounded at the tip, 
in the latter two the tail is filiform and its 
tip is bifid. The Miami material contained 
one species \\'hich seems to correspond in 
all essential features with H. fletcheri, al- 
though the indistinctness of the amphids 
and the somewhat sketchy figures of the 
type leave room for doubt. 

The second species in our material is 
closely related to H. supercirrhatiis Gerlach, 
1955, and //. longiseto^us Hopper, 1963, but 
is distinguished by the much more elon- 
gated, filiform tail, the spicula which have 
a ventral hump, and the gubernaculum with 
its lateral guiding pieces. Examples of H. 
longisefosus have been recovered from the 
Charleston, South Carolina, samples, and 
its status will be clarified in the paper deal- 
ing with the specimens collected from that 

Halalaimus (H.) cf. fletcheri Mawson, 1958 

Plate II, fig. 3, a-c 

Halalaimus (H.) cf. fletclwri Mawson, 19.58: .332. 
fig. 13, a, b. 

L = 2.6-2.37 mm; w = 26-35 ju.; tail in <^ , 
310 ^ (18 a.b.d.), in 9 275 ,x. Head at level 
of first cephalic setae about 4 /x wide and 
6 ^ high. Six labial setae, 2 /x long. Cephalic 
setae 6 /x long, arranged in two circles ( 6 + 
4) about 4 fx apart. Amphids beginning 
about 10 /,<, from anterior end, indistinct, 
particularly the posterior end which appears 
to run into lateral alae. Its length in one 
specimen is probably 17 p.. Anterior portion 
of neck exceedingly drawn out, narrow for 
about two-thirds its length. Tail with bifid 
tip, each prong 10 /j. long. Spicula 27 jx 
long, with velum. Gubernaculum strongly 

Representation in samples studied. — M- 
2, Key Biscayne. 

Geograpliical distribution . — Macquarie 
Isl, Kerguelen Isl. (Mawson, 1958b). 

Halalaimus (H.) meyersi new species 
Plate II, fig. 4, a-c 

L = 2.26-2.49 mm; w = 17 /x; Vu = 47%; 

tail in i , 540 /x. Head about 4.5 p. wide, 7 
/x high. Cephalic setae 20 /x long, arranged 
in two circles ( 6 -f 4 ) . Cuticle finely 
striated. Amphids 38-40 /x long, beginning 
20-30 fx from anterior end. Excretory pore 
50 p. from anterior end, ampulla at base of 
amphids. Terminal excretory duct 8-10 /x 
long. Narrowed portion of neck about one- 
fourth its total length. Spicula 21 /x long, 
with ventral swelling at the end of its 
proximal third. Gubernaculum with lateral 
guiding pieces. Tail very thin, elongated, 
with narrow tip. 

Holotypespecimen.~Mc\\e; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa Collection Num- 
ber 4067, Type slide No. 60. Type locality, 
M-2, Key Biscayne. 

Representation in samples studied. — M-2, 
Key Biscayne. 

Remarks. — This species is named in 
honor of Dr. Samuel P. Meyers, our Miami 

HALANONCHINAE new subfamily 

Type genus.— Ho/anoncfius Cobb, 1920: 

Diagnosis. — Tripyloididae, with three 
large, well-separated lips, six labial papillae, 
and 10 cephalic setae in two circles, the 
anterior circle consisting of six jointed, 
mostly elongated setae; large, unarmed 
conical or cylindrical buccal cavity without 
partitions; spiral or tubular amphids; spicula 
short, bent; gubernaculum plate-shaped, 
simple, without apophysis; pre- and some- 
times postanal supplements present in 

Discussion. — In Cobb's paper of 1920 
one finds the description of three closely 
related genera, the systematic position of 
which has been doubtful ever since. These 
genera are: 1) Cijtolaimium [with the 
species C. exile Cobb, 1920, and C. obtusi- 
caudatum Chitwood, 1936, the latter, ac- 
cording to Gerlach ( 1962 ) , being a synonym 
of the former], 2) Rliabdocoma [with the 

248 Bulletin Museum of Coiiii>ar(itivc Zoology, Vol. 135, No. 5 

species R. americana Cobb, 1920, R. artic- 
iilata Gerlach, 1955, R. brevicauda Schuur- 
mans-Stekho\'en, 1950, R. cyJindricaiida 
Schiuirnians-Stekhoxen, 1950, R. macnira 
Cobl), 1920], and 3) Hcdanonchiis (syn. 
Latilai))iii.s Allgen, 1933) [with the species 
//. macramphidus Chitwood, 1936, and H. 
macrurus Cobb, 1920]. 

All tliree genera are characterized by a 
smooth cuticle, an arrangement of cephalic 
sense organs in three circles of 6 + 6 + 4, 
the second circle consisting of the longest 
and distinctly jointed setae, three large, 
deeply cut lips, a buccal cavity which is 
either conical or cylindrical, amphids which 
are either spiral with a single turn and 
a posterior break (Cytolaimiiim, RJuiJydo- 
coma), or more tubular or pocket-shaped 
with a circular opening (Halanonchus), 
simple spicula, plate-shaped, simple guber- 
nacula, and the occurrence in males of 
many supplements, not only preanally but 
also postanally and even in the cervical 

The number (though not the arrange- 
ment) of the cephalic setae, the deeply cut 
lips, the shape of the amphids (although 
the situation in Halanonchii.s- is not quite 
clear), and the presence of a large buccal 
cavity suggest relationship with the Tripy- 
loididae, to which family Halanonchus and 
Rlial)d()coma ha\c been referred by Filipjev 
( 1934 ) . On the other hand, as Schuurmans- 
Stekhoven ( 1950 ) and Gerlach ( 1955, 1962) 
have pointed out, both Rliahdocoma and 
Cyfohimium have a number of featiues in 
common with Trcftt.mi de Man, 1893, which 
has no buccal cavity, pocket-shaped or spiral 
amphids and no supplements, and which 
so far has been considered an oxystomatid 
of somewhat uncertain position. Finally. 
Chitwood (1936, 1951) placed Cytohimium 
and Rhahdocoma with the Monhysteridae. 

We consider that by virtue of the deeply 
cut lips, the jointed setae, the large Iniccal 
cavity (particularly in Halanonchus), and 
the spiral amphids (in Cytokiimium and 
RhaJydocoma ) , the three genera mentioned 
belong to the family Tripyloididae. How- 

exer, because of the simplicity of the buccal 
cavity, the arrangement of the cephalic 
setae in two circles (instead of one circle), 
and the different structure of the male 
genital armature, a distinct subfamily should 
be created for them, for which we propose 
the name Halanonchinae new^ subfamily, 
with Halanonchus Cobb as the type genus. 

This new subfamily, via Trcfusia, links 
the Tripyloididae with the Oxystomatidae 
and thus allows a more satisfactory place- 
ment of the former family which so far has 
occupied a rather isolated position either 
within the order Araeolaimoidea (Schu- 
lu-mans-Stekhoven, 1935), or the super- 
family Chromadoroidea (Chitwood, 1951). 

\\'ithin the genus Trcfusia and the sub- 
family Halanonchinae it seems as if a transi- 
tion of the amphidial shape from spiral to 
pocket-shaped or tubular had taken place, 
thus stressing the intennediary position of 
this group of genera between the Oxysto- 
matidae and the Tripyloididae. This is dem- 
onstrated not only by a comparison of 
Halanonchus with Cyfolaimium and Rhah- 
docoma, but also by Trcfusia varians Ger- 
lach, 1955, in which the juxeniles have spiral- 
shaped, and the adults tubular-shaped am- 
phids not too different from the shape 
which we observed in H. macrurus (see 
below). Further proof of the intennediary 
position of the new subfamily might be 
the fact that Rhahdocoma is reported to 
have just one posterior ovary (as is the 
case in many oxystomatids), whereas Cy- 
folaimium has two ovaries (like the Trip- 
yloidinae ). 


Type spedes.— Halanonchus macrurus Cobb, 

1920: 266, fig. 51. 

Halanonchus Cobb, 1920: 266. 
Lotihimus Allgen, 1933: 90. 

Halanonchus macrurus Cobb, 1920 
Plate III, fig. 5, a-c 

Halanonchus macrurus Cobb, 1920: 266, fig. 51. 

L = 2.00 mm; w = 40 /x; esophagus = 265 
/A. Head diameter (on level of 4 cephalic 

Florida Marine Nematodes • Wiescr and Hopper 249 

setae) 19 fi. Three large, deeply cut lips, 
on each lip 2 thin labial setae and, shortly 
behind, 2 larger, t\vo-jointed cephalic setae, 
measuring 3-3.5 /x. Further behind, there 
are the four setae of the second cephalic 
circle, measuring about 5 ^ and being non- 
jointed. There are many yellowish granules 
which are scattered throughout the epider- 
mis of the body, although a certain arrange- 
ment into longitudinal rows can be dis- 
cerned. The amphids give the impression 
of sawed-off and slightly bent pieces of 
tubing with an indistinct, more or less 
circular orifice; they are 8 /x long and 
situated 20 p. behind the anterior end. Cobb 
shows the amphids more pocket-shaped 
with a distinctly circular orifice. Each lip 
seems to be supported by a large oval 
structure which apparently was mistaken 
for the amphids by Allgen ( 1933 ) in his 
description of Lotikiiiniis zosferac. In the 
cervical region one can distinguish 5-6 
ventral bumps which might be papillae. 
The same organs have been observed in 
undcscribed species of Rhahdocoma and 
HaJammchus by Gerlach (1962). Buccal 
cavity with strong walls, 20-22 jx long. 
Esophagus weakly dilated posteriorly, no 
bulb. A small triangular cardia is present. 
Spicula slender, 32 jx long, gubernaculum 
either absent or consisting of a thin lamella, 
lying parallel to the spicula. There are 
about 12 preanal supplements, the posterior 
six being more distinct than the more an- 
terior ones. Tail 610 jx long, a.b.d. 28 /x. 

Rcpresenfafion in samples studied. — M-8, 
Biscayne Bay. 

Geographical distribution. — Biscayne Bay 
(Cobb, 1920). 


Type species.— Cy/o/o/'m/um ex/7e Cobb, 

1920: 251, fig. 31. 
Cytolaimium exile Cobb, 1920 

Plate III, fig. 6, a-d 

Cyfokiimium exile Cobb, 1920: 251, fig. 31. 
Cytolaimium obtiisicauclatiim Chitwood, 1936: 13, 
"fig. 3, J-L (cf. Gerlach, 1962). 

L = 3.2 mm; w = 32 fx; diameter at base 

of esophagus 30 /x. Esophagus 250 /x long. 
Head diameter 20-25 /x. Head with six 
setose labial papillae, six segmented ce- 
phalic setae and four subcephalic setae. 
Cephalic setae in female 20 /x long, in male 
somewhat longer but distorted so as to 
render precise measurement impossible. 
Amphid 8 /x wide and located 20-23 /x from 
anterior end. Subcephalic setae 10 /x long 
in male, 5-6 /x in female and located 35 /x 
from anterior end. Body with several short 
(3-4 ij.) cervical setae beginning 90 /x from 
anterior end and ceasing at nerve ring and 
with a few, very thin, somatic setae 10-15 
IX in length. Female didelphic, amphidelphic, 
ovaries reflexed. Male diorchic, testes out- 
stretched. Spicules 37 /x long, their chord 
28 /x. Male preanally with 9 pairs of discoid 
supplements and 2 pairs of fleshy, setose 
papillae and postanally with 3 pairs of 
discoid supplements and 5 pairs of fleshy 
setose papillae. Tail length extremely vari- 
able, 15 /x long for one female, 420 /x for 
the other, and 740 /x for the male. 

Representation in samples studied. — M-2, 
Key Biscayne. 

GeograpJncal distribution. — Biscayne Bay 
(Cobb, 1920). Beaufort North Carolina 
(Chitwood, 1936), Maldives (Gerlach. 1962). 

Remarks. — Tlie data for our specimens 
( 1 6 , 2 9) lend support to the view of 
Gerlach (1962) that C. obtusicaudatum 
Chitwood, 1936, is a synonym of C. exile 
Cobb, 1920. The tail length in our three 
specimens ranged from 15 ^ in one female 
to 740 fx in the male. In the specimens with 
shorter tails, the terminus appears abnonnal 
and suggests the phenomenon of \\ound- 
healing as discussed for this species by 
Gerlach. Even in the case of the longest- 
tailed specimen there is the possibility of 
a missing portion, as no typical spinneret 
can be recognized. 

The setose papillae associated with the 
male tail appear to be similar to the papillae 
seen on the discoid supplements, the only 
difference being the lack of the disc. 

250 BuUctiii Mus( iini of Comparative Zoology, Vol. 135, No. 5 

Type spedes.—Bafhyloimus australis Cobb, 
1893: 409, 410, fig. 9, l-IV. 

Our material contained two species of 
the genus BafliyJaimus, one of wliich ap- 
pears to be the type species, B. amtialis 
Cobb, 1S93. The other represents an un- 
described species closely related to B. 
co}xicoms Hopper, 1962, from which it can 
be distinguished b\ lia\'ing the shorter of 
the submedian cephalic setae equal in 
length to the basal segment of the longer. 
In B. capacosus the shorter of the pair is 
approximately half as long as the basal 
segment of the longer. 

Bathylaimus ausfralis Cobb, 1893 
Plate IV, fig. 7, a-e 

Bathylaimus australi.s Cohh, 1893: 409-410, fig. 

9, I-I\'. 
Bathi/laiinus assimilis de Man, 1922b: 119, 120, 

fiu. 2-2e. NEW SYNONYMY. 

h= 6 , 1.5-2.2 mm, 9 , 1.4 mm; w = 45- 
53 II- diameter at base of esophagus 41-44 /x. 
Esophagus 270-,370 ^. Vu = 51%. Tail 9S- 
135 /x (3-3.5 a.b.d.). Head diameter 22-24 /x. 
Labial setae about 4 ix. Cephalic setae 20 + 
9 /x, the longer set comprised of four seg- 
ments. Cervical region bearing eight rows 
of somatic setae; those anterior to nei-ve 
ring being 5 /x long. Buccal cavity 37-39 /x 
long, bipartite (29-31 + 8-9 ^); without 
armature. Amphid 20 /x from anterior end, 
positioned over posterior half of the anterior 
portion of buccal cavity; internal amphidial 
ponch 5 /;, wide, its orifice approximately 
3 /-. Spicula 4(S-50 /x long, gubernaculum 
50 /x long. Female tail without setae, male 
tail bearing .setae as illustrated (Fig. 7, b, 


Representation in .samples- studied. — ^V, 

Vero Beach. 

Gc()'j,i(i))hical distribution. — Cosmopoli- 

Remarks. — In considering the synonymi- 
zation of i^. assimilis de Man with B. 
au.^tralis Cobb, the following facts are per- 
tinent. \\'i(scr. 1956, separated the two 

species on the basis of different values for 
the lengths of both the buccal cavity and 
the spicula. According to the information 
presented by that author, B. au.stralis has 
a buccal cavity of 33 /x and spicula of simi- 
lar length, while in B. assimilis the figures 
were 50-55 fx and 45 ^, respectively. Fur- 
ther, W'ieser states that: "B. australis and 
B. assimilis are very closely related and the 
differences in the length of both spicula 
and buccal cavity are the only ones I can 
find." An examination of the original de- 
scription of B. australis shows that Cobb's 
animal had a buccal cavity of 50 /x and 
spicula of approximately 40 fx. These fig- 
ures closely approach those presented by 
de Man in his original description of B. 
assimilis (buccal cavity 40-43 ^, spicula 37 
fi). On this basis we consider B. as.similis 
de Man, 1922b, along with its synonyms, to 
be a junior synonym of B. australis Cobb, 

B. setosicaudatus Timm, 1961, while ex- 
tremely close to B. au.stralis.. can be sepa- 
rated by the fact that the spicular cephali- 
zation is reduced while that of B. au.stralis 
is prominent. Also the spicula are uni- 
formly bent in B. seto.sicaudatus, while in 
B. aiLsiralis most of the cur\'ature is limited 
to the mid region. 

Bathylaimus arfhropappus new species 
Plate IV, fig. 8, a-f 

L = 3.1-3.3 mm; w = 6 , 52-55 /x, 9 , 62 
fx. Diameter at base of esophagus, $ , 44- 
48 /x, $ , 53 IX. Esophagus i , 687-750 ix, 
9 , 820 IX, Vu = 56% . Tail 6 , 118-130 ^ (3.4- 
3.7 a.b.d.), 9, 220 il (5.4 a.b.d.). Head 
diameter, 34-37 ^. Labial setae about 20 ^. 
Cephalic setae 47-52 + 1^19 /x. Both the 
labial setae and longer cephalic setae are 
segmented, with the former having three 
segments and the latter four. Cervical re- 
gion bearing eight rows of somatic setae 
which are more or less arranged in circles. 
First circle, 6 ix long, near base of amphid. 
Second circle, 17 /x long, half the distance 
from the anterior end to the nerve ring. 
Third circle, 8 /x long, just anterior to nerve 

Florida Marine Nematodes • Wieser and Hopper 251 

ring. Remaining two circles, 8-9 ix long, 
posterior to nerve ring, the last 70 /x poste- 
rior to nerve ring. Buccal ca\'ity 28-35 p- 
long, without armature. Amphid in i , 12- 
13 |U, in 9 , 8 /x wide, located posterior to 
buccal cavity, 37-38 fx from anterior end. 
Spicula 50-57 /x long, proximally cepha- 
lated. Gubernaculum 50 jx long. Male tail 
bearing setae as illustrated ( Fig. 8, f ) ; sub- 
terminal setae 40-45 ix long. Four gland 
cells are associated with the spinneret ap- 
paratus in both sexes. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4070, Type slide No. 61. 
Type locality, M-5, Virginia Key. 

Representation in samples studied. — M- 
5, Virginia Key. 

Geo^iraphica] disiribufion. — The species 
also occurs at Gulf Shores, Alabama (un- 
published observation). 

Retnarks. — The region of the esophagus 
directly posterior to the buccal cavity is 
constructed in such a manner that, if the 
esophageal musculature were to exert a 
pull in this region, a cavity might arise 
which could be mistaken for a second com- 
partment of the buccal cavity. This subject 
was also brought up in the description of 
B. capacosus Hopper, 1962. 

PHANODERMOPSIS Ditlevsen, 1926 

Type species.— Phanodermopsis groenland- 
ica Ditlevsen, 1926: 13, 14, pi. 7, figs. 
1, 2; pi. 8, fig. 5. 

Our material contained typical represen- 
tatives of P. longisetoe Chitwood, 1936. 

The following species have been de- 
scribed since a key to the species of this 
genus was given b\- ^^'ieser (1953): P. 
conicauda Filipjev, 1946, and P. in^irami 
Mawson, 1958, belonging to W'ieser's group 
A, and P. ohtusicauda Filipjev, 1946, be- 
longing to group B. P. necta Gerlach, 1957, 
does not appear to belong to this genus, 
since it has a well de\eloped supplement 

and spicula as in Phanoderma. P. suecica 
Allgen, 1953, is a species inqiiirenda. 

Mawson ( 1958a ) raised the question of 
the position of this genus since Ditlevsen 
(1926) did not designate a type species. 
However, Filipjev ( 1927 ) in the appendix 
to his paper, established synonymy of his 
genus Galeonema with Phanodermopsis 
and designated P. fi,roenIandica Ditlevsen 
as the type species. 

Phanodermopsis longisetae Chitwood, 1936 
Plate V, fig. 9 a-c 

PJianodermopsis longisetae Chitwood, 1936: 209, 
210, pi. 26, fiss. 16-19. 

L = 3.85 mm; w = 80 /.; tail 250 /x. Head 
diameter 15 /i, capsule weakly developed. 
Labial papillae distinct, conical. Cephalic 
setae 15 + 10 jx long. Amphids 6 ix wide = 
40'yr of head diameter, 10 ^ behind anterior 
end. Excretory pore 55 /x behind anterior 
end. Spicula 360 ix. Gubernaculum 56 fx. 
Caudal setae arranged in characteristic 
pattern, the setae being of two types: one 
fleshy and S-shaped, the other slender and 
straight or slightly curved. 

Representation in samples studied. — V, 
\'ero Beach. 

GcograpJi ical distribution. — Beau- 
fort, North Carolina (Chitwood, 1936). 

ENOPi.O/DES Saveljev, 1912 

Type species.— Enop/o/c/es fypicus Saveljev, 
1912: 115. 

In this genus, classification is possible 
only on the basis of the male genital arma- 
ture. Consequently, we have to insist that 
all species known from juveniles or by fe- 
males only are to be regarded as species 
inquirendae. This includes, in addition to 
the doubtful species mentioned by Wieser 
( 1953), the following: E. labiatus BiitscWi, 
1874 [Synonymy of this species with E. 
spiculohamatus Schulz, 1932, cannot be 
proven in an\- way and should be aban- 
doned, as advocated by Brunetti, 1950.], 
E. tridentatus Sa\'elje\-, 1912, E. brevis 

252 Bulletin Museum of Comparative Zoology, Vol 135, No. 5 

Filipjev, 1918, E. hratistrorni, E. parakibio- 
tus, E. icdiictu.s, and E. lon^icaudatus all 
Wieser, 1953, E. oU<iotricha Mawson, 1956 
(syn. E. oli'j^ochiictus Mawson, 1956), E. 
ptcro^^natlius Mawson, 1956, and E. ker- 
'^ucJcnsc Mawson, 1958. 

The remaining species can be separated 
into two gronps, one with short spicula, the 
other with extremely long spicula. The 
former group comprises only two species, 
viz., E. ciirhatus Filipjev, 1918, and E. 
tyrrhenicus Brunetti, 1949 (cf. Gerlach, 
1952), for which most likely a new genus 
or subgenus should be established. 

The group with long spicula is very uni- 
fonn. Since the gubernaculum represents 
one of the best distinguishing characters, 
the shape of this organ in all the species 
belonging to this group (except E. ti/picus, 
of which no figures were given) is shown 
in Text-figure 1. A number of species can 
be separated immediately by the shape of 
the gubernaculum, viz., E. hirsiitits Filipjev, 
1918, E. hnmettii Gerlach, 1952, E. vcctis 
Gerlach, 1957 (syn. E. brunettii var. vcctis), 
and E. harpax \\'ieser, 1959. In the remain- 
ing species the gubernaculum is more or 
less S-shaped, although differences in shape 
between the species can be found. The 
species comprising this remaining group 
can be separated by use of the following 

Key to Species of Enoi'I.oides Possessing 
s-shaped gubernacula 

1 . Cephalic setae of equal length 

E. ccplialophorus ( Ditlevsen, 1919) 

Cephalic setae of unequal length 2 

2. Longest cephalic setae about 1.2 head diam- 

eters long, shorter setae measuring l:{ of 
longer ones. Gubernaculum with cliar- 
acteristic ventral knob ., £. ^njphtts n. sp. 
Longest cephalic setae measuring not more 
than one head diameter, shorter ones 
about 'j that length. Cmbernaculum 
rather smoothly S-shaped — 3 

3. Tip of gubernaculum apparently 3-pronged; 

supplement measiuing about half the cor- 
responding body diameter (description 

and figures not (luite clear) 

_„ E. spictiloluimatus Schnlz, 1932 

Tip of gubernaculum 2-pronged; supplement 
much shorter 4 

4. Distal end of spicula with moljile spine; 

mandibles 55-60 m long 

E. amphioxi Filipjev, 1918 

Distal end of spicula without mobile spine; 
mandil^les 45 m <'i" less 5 

5. Spicula smooth, distal end pointed, then 

dilated; mandibles about 45 fj. (extrapo- 
lated from Southern's figmes and text) 

E. labrosiriatiis (Southern, 1914) 

Spicula vertically striated, in their distal 
half with a diagonal break (Fig. 10, c); 
mandibles 30-35 m long - E. I)isulcus n. sp. 

The type species, E. typicus Saveljev, 
1912, is poorly known since no figures were 
given. It seems to differ from all other 
species by the small dimensions of its 
organs ( cephalic setae only 8 ^a long, buccal 
ca\ity only 9 /x deep! ) . 

Enoploides bisulcus new species 
Plate V, fig. 10, a-d 

L = 3.5-4.2 mm; w = 115-120 //; diam- 
eter at base of esophagus 95-105 ^a,. Esoph- 
agus 750-800 fx; nerve ring at 25%. Vu = 
52.5%. Eggs 165x75 /x, one per uterus. 
Tail in 6 , 170-200 /x (4 a.b.d.), in 9 , 215- 
225 IX (3.7-4.5 a.b.d.), in juvenile 4.8 a.b.d. 
long. Head diameter 50-56 /x. Labial setae 
25 pi. Cephalic setae 45-50 -f 25-28 /x. 
Mandibles 30-35 /x long, deeply notched 
anteriorly, with strong apophyses and mus- 
cles that link them to the buccal capsule. 
Subventral teeth small, nearly parallel to 
mandibles and thus difficult to obsei-ve in 
lateral view. Their presence can best be 
ascertained in en focc view. Dorsal tooth 
seemingly absent. Peribuccal portion of 
esophagus strongly de\eloped, muscular. 
Spicula 420-475 /x, knobbed proximally, 
vertically striated throughout, with a diag- 
onal break in their distal end that runs from 
dorsal to ventral, tip pointed. Gubernacu- 
lum S-shaped, its proximal end open, ill- 
defined; distal end forked. Supplement 20 
/x long, 120-150 /x preanal. Tail with pair 
of fleshy, S-shaped spines and a number 
of setae. Setae in anal area and paired 
terminal setae, 17 y. long. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 

Florida Marine Nematodes • Wicscr and Hopper 253 


a=^ &<:. 

cF=y. (r^ 

Figure 1. Gubernacula of several species of Enop/oides. a — E. hirsutus: b — E. brunettii; c — E. vecfis; d — E. harpax; 
e — £. amphioxi; f — E. labrostriatus; g — E. cepho/ophorus; hi — E. sp/cu/ohomofus; i — E. b/su/cus; k — E. gryphus. All 
gubernacula copied from original descriptions of respective species. 

mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 62. Type 
locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Enoploides gryphus new species 

Plate II!, fig. lie; plate V, fig. 1 1 a, b 

L = 3.0-3.7 mm; w = 8.5-100 /^; diameter 
at end of esophagus 70-85 fi. Esophagus 
800-900 ix. Tail 190-240 /x; a.b.d. 45-60 ix. 
Head diameter 45-57 fi. Labial setae 28-32 
/._. Cephalic setae 60-80 + 20-27 ix. Man- 
dibles 26-28 fx. Apart from the dimensions, 
the structure of the buccal armature is the 
same as in E. hisidciis. Spicula 230-260 /x, 
6-7 /x wide, knobbed proximally, vertically 
striated throughout, without break, tip 
pointed. Gubernaculum hook-shaped, with 

characteristic ventral knob. Supplement 
80-110 fx preanal. Tail with pair of slender 
postanal spines (not S-shaped as in the 
foregoing species) and several setae. (Note: 
All the somatic and caudal setae on this 
species are shorter and more slender than 
on E. bisidciis.) 

HoJotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomol- 
ogy Research Institute, Ottawa, Collection 
Number 4070, Type slide No. 63. Type 
locality, M-5, Virginia Key. 

Representation in .samples .studied. — M- 
5, X'irginia Key. 

Remarks. — This species resembles the 
foregoing one in most respects. The main 
distinguishing features are the longer 
cephalic setae and the shorter spicular ap- 

254 Bulletin Museuni of Conipanitivc Zoology, Vol 135, No. 5 


Type species.— Mesacanf/io/des scufpfilis 

Wieser, 1953: 86, fig. 46, a-d. 

Through the addition of two new species 
this genus is becoming more heterogeneous 
as far as the male genital appaiatus is con- 
cerned. Howexer, its most characteristic 
feature, the shape and texture of the man- 
dibles, suffices to separate it clearly from 
related genera. Classification is largely 
based on the genital armature ( see \\' ieser, 
1959). Consequently, M. wieseri Mawson, 
1956, has to be considered a species in- 
quirenda, although it appears to be closely 
related to M. Jatignaihus. 

Key to Species of Mesacanthowes 

1. (;ubernaculum reduced, no supplement 

M. latignathus ( Ditlevsen, 1919) 
Gubernaculum and supplement present 2 

2. Supplement small, tubular - 3 

Supplement \erv large, "wrench-like" 

M. sculptilis Wieser, 1953 

3. Spicula more than two anal diameters long, 

tail filiform, with flagellum 

M. fiJjuhifus n. sp. 

Spicula about one anal diameter long, tail 
plump or elongate, never filiform 4 

4. Gubernaculum more or less plate-shaped; 

head with four circles of cephalic setae___ 

M. caputmediisae (Ditlevsen, 1919) 
Cubcrnaculuni S-shaped; head witli 2-3 

circles of cephalic setae 5 

.5. Implantation of cephalic setae near middle 
of cephalic capsule; spicula strongly arcu- 
ate, tip pointed -^ M. sinuosus Wieser, 1959 
Implantation of cephalic setae at posterior 
edge of cephalic capsule; spicula nearly 

straight, tip elaborately armed 

M. p.sittacus n. sp. 

Mesacanthoides fibuiatus new species 
Plate VII, fig. 12a-c 

L = 4.3 mm; w = 75 /x. Esophagus = 675 
//. Tail = 350 /v.. Head diameter 33 p. 
Labial setae 10 /i. Longest cephalic setae 
42 ij., setae of submedian pairs sticking to- 
gether. One circle of subcephalic setae in 
male. Cephalic capsule deeply lobcd. Ce- 
phalic organ present, in front of lateral 
cephalic seta. Mandibles 18 X 10 /x, solid 
as ill Knoploides but with an additional 

transverse bar near the anterior end; more- 
over, the sclerotization is not uniform but 
gives a mottled impression. The tips of the 
claws are darker than the rest of the man- 
dibles. Teeth well developed, about half 
as long as the mandibles. Stomodeal ring 
forming three "brackets" around the man- 
dibles which serve as muscular attach- 
ments. Spicula 125-135 p, about 4 anal 
diameters, cephalate proximally, with a 
break just before the distal sixth. Guber- 
naculum slightly S-shaped, 15 p long. Sup- 
plement small, about 90 p in front of anus. 
Tail at first conical, then abruptly attenu- 
ated and drawn out into an extremely long 
and whip-like flagellum, about five times 
the length of the spicula. In the circumanal 
region there are scattered setae. 

Holotijpe specimen. — Male; Canadian 
Xationai Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4069, Type slide No. 64. 
Type locality, M-3, Key Biscayne. 

P.epresenfation in samples studied. — M- 
3, Key Biscayne. 

Mesacanthoides psittacus new species 
Plate VI, fig. 13a-e 

L = 2.32 mm; w = 50 p; esophagus = 540 
IX- tail = 222 p.. Head diameter 35 p. Lips 
plump, labial setae stout, 15 p. Cephalic 
setae: lateral 62 p, submedian 56 + 27 p, 
implanted near posterior edge of cephalic 
capsule. The latter with straight edge, 17 
p. high. Mandibles 19 X 11 p., powerful, 
plump, of typical shape and texture, each 
claw with an oval apophysis on its 
"shoulder." Teeth well de\eloped, about 
half as high as the mandibles. Spicula 
plump, nearly straight, 50 /x long, their tips 
broacl, each with a three-pronged process 
on the caudal edge. Gubernaculum S- 
shaped, about 26 p. long, powerful. Supple- 
ment small, 90 p in front of anus. Tail 5 
anal body diameters long, with scattered 

HoIntj/))c specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute. Ottawa, Collec- 

Florida Marine Nematodes • Wiescr and Hopper 255 

tion Number 4074, Type slide No. 65. Type 
locality, L, Lauderdale-by-the-Sea. 

Representation in samples studied. — L, 

ONCHOLAIMUS Dujardin, 1845 

Type species.— Onc/io/a/mus aftenuatus Du- 
jardin, 1845: 236. 
Keys to the species of Oncholaimiis can 
be found in the works of Kreis ( 1934 ) and 
Wieser (1953). 

Oncholaimus dujardinii de Man, 1878 
Plate VII, fig. 14, a, b 

Oncholaimus dujardinii de Man, 1878: 94, pi. 7, 
fig. 4, a-c. 

L = 2.4-2.95 mm; w = 42-45 //; esopha- 
gus 350 jji. Yu = 769f . Tail, in 9 , 67 /x 
long (2.6 a.b.d.). Head 18-20 ,j. wide, 
bearing 10 short, subequal cephalic setae, 
the longest 4 /x long. Buccal cavity 22-24 X 
12 fx anned with 3 strong teeth of which 
the left subventral is the more prominent. 
Amphids in male 7-9 jj. wide ( = 37-459f of 
c.b.d.). Anterior end of esophagus with a 
pair of dark pigmented masses (cf. de Man, 
1878). Excretory pore 45-50 ^ behind an- 
terior end. Renette cell prominent, 90-100 
fj. long, located approximately 200 ^ poste- 
rior to base of esophagus. Spicula 27-31 /j. 
long, proximally cephalated. Gubernacu- 
lum absent, although a dorsal thickening of 
the cloacal lining may give the impression 
of the presence of such a structure. The 
thickened area appears to be a point of 
attachment for muscles associated with the 
spicula. Male tail 40 /x long, ventrally 
curved; with 12-16 stout, circumcloacal 
setae. Distal extremity of male tail slightly 
enlarged and bearing two pairs of short, 
stout, subventral setae and a pair of slen- 
der, subdorsal setae. 

Representation in samples studied. — M- 
1, Key Biscayne. 

Geographieal disiribution. — Cosmopoli- 

Discussion. — While our species appears 
to be O. dujardinii de Man, 1878, there re- 

mains some doubt as to the identity of this 
species. SchuuiTnans-Stekhoven (1950) and 
Inglis (1962) reason that the typical O. 
dujardinii is devoid of a gubernaculum, 
whereas O. dujardinii de Man sensu Steiner, 
1915 ( and other authors ) possesses such an 
organ and thus represents a different spe- 
cies. However, we are of the opinion that 
the dorsal thickening of the cloacal lining 
described in our specimens has been oc- 
casionally misinterpreted as a gubernacu- 
lum and we regard the questioned accounts 
of O. dujardinii as representing de Man's 


Type species.— Oncfio/a/m/um oppendicula- 

tum Cobb, 1930: 227, figs. 2, 3, 6, v, 

8, 9. 

A discussion of this genus, with a key to 
species, was given recently by Chitwood 
(1960). We feel that separation of this 
genus from Oncholaimus should be based 
mainly on the presence of a Demanian 
system and not so much on that of the pre- 
anal papillae (not postanal, as erroneously 
stressed by Wieser [1953] and subsequent 
authors), as rudiments of preanal papillae 
can also be found in representatives of the 
latter genus as well as in other oncholaimid 
genera. This would suggest that the two 
short-spiculed species of MetoneJioJaimus 
described by Mawson, i.e., M. brevispicu- 
Jum Mawson, 1957, and M. thysanouraios 
Mawson, 1958, actually ought to be re- 
ferred to Oncholaimium. The diagnosis of 
Cobb ( 1930 ) has to be emended so as to 
include, in this genus, species in which the 
Demanian organ is provided with exit 

Both species found in Florida have been 
previous!}' reported from the east coast of 
the United States. 

Oncholaimium appendiculatum Cobb, 1930 
Plate VIII, fig. 15, a-c 

Oncholaimiuni appcndicuhiluin Cobb, 1930: 227, 
figs. 2, 3, 6, V, 8, 9. 

256 BuUctin Museum of Conipanitive Zoology, Vol 135, No. 5 

L = 2.27 mm; \v = 44 /x; esophagus = 360 
/x; nerxc ring 2(S() // l^'hind anterior end. 
Head diameter 26 ji. Lal)ial eapsule lobed, 
eonspicuous. Labial papillae conical. Ce- 
phalic setae 9 /x, equal. Buccal cavity 30 X 
18 /.. Teeth 25 + 15-16 /x, the longest one 
reaching to about the level of implantation 
of the cephalic setae. Amphids 11 /x = 40% 
of c.b.d. wide. Scattered cervical setae. 
ExcretoiA pore 30 fx in front of nerve ring. 
Spicula 65 fx (one tail length). No guber- 
naculum. Anal diameter 25 /x. One large, 
"prehensile" preanal papilla. Twenty-four 
to 26 circum- and postanal setae, about 13 
/x long. Tail with small xentral papillae at 
the beginning of the distal third. 

Representation in samples studied. — M- 
1, Key Biscayne. 

Geo<iraphieal distribution. — Woods Hole, 
Massachusetts, ? Beaufort, North Carolina 
(Pearse, Humm and Wliarton, 1942). 

Oncholaimium domesficum Chitwood and 
Chitwood, 1938 

Plate VIM, fig. 16a-d 

Oncholaimium di>iucsiiciitii Chitwood and Chit- 
wood, 1938. 

Oncholaimium oxijuiis var. domcsticus Chitwood 
and Chitwood, 193S: 458, 459, fig. 1, f-h: nee 
Timm, 1952. 

L = 3.3-3.4 mm; w = <i , 65 /x, 9 , 85 /x; 
esophagus = 525-550 /x. Tail = 70 /x. Vu = 
64%. Head diameter 37 /x. Labial capsule 
lobed, inconspicuous. Six labial papillae. 
Cephalic setae subequal, 8 jx. Buccal cavity 
37-38 X 25-26 p.. Teeth 25 + 21 ix. Am- 
phids 7-8 /x. = 20% wide ( both sexes ) . Ex- 
cretory pore 70-75 /x behind buccal cavity. 
Demanian organ well developed, uvette 
about 400 fx posterior to vulva, 1-2 adanal 
openings on each side. Spicula 45 jx (one 
a.b.d.). No gubernaculum. Preanal papilla 
with two setae. Postanal papillae at the 
beginning of the distal third of the tail. 
Circumanal setae present. Tail 60-70 /x, 
with terminal swelling. 

Representation in samples studied. — M- 
1, Key Biscayne. 

Geoiiraphieal distribution. — New York 

(Chitwood and Chitwood, 1938), California 
(Chitwood, 1960). 

Reinarks. — The characters distinguishing 
this species from O. oxyure (Ditlevsen, 
1911) are the short teeth, the well-devel- 
oped preanal papilla with setae and the 
stout spicula. The specimens described by 
Timm ( 1952 ) deviate in all these charac- 
teristics from O. domesticum and seem to 
be representatives of O. oxyure. 

METONCHOLAIMUS Filipjev, 1918 
Type species.— Oncfio/a/mus demani Zur 
Strassen, 1894: 460, pi. 29, fig. 2. 

A key to this genus was given recently 
by Chitwood ( 1960 ) . We are of the opin- 
ion that, because of the shortness of their 
spicula, M. brevlspieulum Mawson, 1957, 
and M. thysanouraios Mawson, 1958, be- 
long to Oncholaimium. M. haplotretos 
Mawson, 1958, is considered as doubtful 
since only females are known. 

Our material contained three new spe- 
cies, all of which are characterized by rela- 
tively short spicula ( < 180 fx as against 
250-750 IX in other species) and by the 
absence of a gubernaculum [which is also 
lacking in M. albidus ( Bastian, 1865) but 
is present in all other species]. Moreover, 
M. intermedius and M. simplex are dis- 
tinguished by short and plump tails, M. 
simplex by the presence of only one De- 
manian exit pore, xA/. scissus by two ventral 
papillae on the 6 tail and by the slit-like 
openings of the Demanian organ. The rela- 
tive position of osmosium and uvette serves 
as a further character distinguishing M. 
simplex and M. intermedius. 

Mefoncholaimus intermedius new species 
Plate IX, fig. 17 a, d, e; Plate X, fig. 17 b, c 

L = 2.2-2.7 mm; w = 38 /x; esophagus = 
350-400 /x; Vu = 66-74%. Head diameter 
26 /x. Labial papillae small but distinct. 
Labial capsule deeply lobed. Cephalic 
capsule weak. Cephalic setae 8 /x, sub- 
en [ual. Amphids 6 ^ in 9 , 8-9 /x = 30-33% 
of c.b.d. in 6 . Buccal cavitv 25-28 X 16-18 

Florida Marine Nematodes • Wieser and Hopper 257 

fj.. Longest tooth 19-20 /x. Ventral gland 
150-200 /x behind esophagus; excretory 
pore 80-90 /x behind anterior end. In some 
specimens large coelomocytes. Demanian 
organ well developed, uvette 325 /x behind 
vuKa, osmosium anterior to uvette, two 
exit pores 70-100 /x preanal. Eggs 90 X 40 
/x. Spicnla 70-77 ix long, no gubernaculum. 
Preanal elevation with short, stout seta. 
Fourteen circumanal setae. One pair of 
postanal elevations with setae, near be- 
ginning of distal third of tail. Anal diam- 
eter 25 /x. Tail in 9 , 30 /x, in S 48 /x 
long. Caudal glands 350, 490, and 560 /i 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4070, Type slide No. 66. 
Type locality, M-5, Virginia Key. 

Representation in samples studied. — M- 
5, Virginia Key. 

Mefoncholaimus simplex new species 

Plate IX, fig. 18 a; Plate X, fig. 18 b-e 

L = 1.95-2.32 mm; w = 6 , 40, 9 , 43-53 
/x; esophagus = 330-360 /x; Vu = 63-70%. 
Head diameter 24-27 /x. Labial papillae 
small. Labial capsule indistinct. Cephalic 
capsule relatively (for this family) well 
developed. Cephalic setae 8-9 fx, subequal. 
Amphids 9 /x in 9 , 10 /x = 40% of c.b.d. in 
S . Buccal cavity 29-30 X 16-18 /x. Excre- 
tory pore 90-100 /x behind anterior end. 
Demanian organ well developed, uvette 
360-430 /x posterior to vulva and 200-280 
ji anterior to anus; osmosium posterior to 
uvette; one exit pore, opening on level of 
anus at dorsal side of body. Moniliform 
glands of varying grades of distinctness in 
different specimens. Eggs 107-120 x 40-45 
fx. Spicula 120 /x long; no gubernaculum. 
Twelve to 14 circumanal setae and perhaps 
two preanal, indistinct pores. Tail conical, 
37 fx long in 6 , 40-46 ^a in 9 . Anal diam- 
eter 26 IX in 6 , 30-32 /x in 9 . lu the single 
male a subterminal dorsal structure was 

seen on the tail which might represent the 
opening of two of the three caudal glands. 
In the female all caudal glands definitely 
open through the spinneret. 

HoJotiipc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion ^Number 4070, Type slide No. 67. 
Type locality, M-5, Virginia Key. 

Representation in samples studied. — M- 
5, Virginia Key. 

Mefoncholaimus scissus new species 

Plate IX, fig. 19 a, b; Plate X, fig. 19 c 

L = 4.3-4.5 mm; w = c5 , 64, 9 , 72 /x; 
diameter at base of esophagus <^ , 59, 9 , 
69 IX. Esophagus 600-640 ^ long. Vu = 
65%. Head diameter 36-40 /x. Head with 
six small labial papillae. The nerves in- 
nervating the labial papillae and the points 
at which they pass through the cuticle are 
more prominent that the papillae them- 
selves. Cephalic setae 13-15 /x, subequal. 
Amphid 10-11 p. wide. Buccal cavity 45- 
48 x 25-27 IX. Longest tooth 35-38 fx, 
shorter teeth 25-28 ju. Excretory pore 105- 
125 IX from anterior end. The most con- 
spicuous structures of the Demanian organ 
are the moniliform glands. These glands 
are approximately 225 /x. long and open to 
the exterior via 17 /j. wide slits (not pores), 
which are located 157-172 /x preanal. Eggs 
115-140 X 55-60 IX, six seen in the uterus of 
one specimen. Spicules 175-180 /x long. 
Dorsal wall of cloaca thickened, but not 
fomiing a distinct gubernaculum. Tail in 
9 , 200-215 IX, in S , 220-230 ,x long. Male 
with 5 pairs of circumanal setae, 6-7 pairs 
of subventral setae and two prominent ven- 
tral papillae. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4070, Type slide No. 68. 
Type locality, M-5, Virginia Key. 

Representation in samples studied. — M- 
5, Virginia Key. 

258 Bulletin Mim'imt of Comparative Zoology, Vol. 135, No. 5 

PROONCHOLAIMUS Micoletzky, 1924 
Type species.— Oncho/a/mus megasfoma 
Eberth, 1863: 26, pi. 1, figs. 18-20. 

This genus is easily recognized by the 
large bubble-like cells ( "Trabekula-Struk- 
tur," in Gennan literature) that occur in 
the pseudocoeloniic caxity between the 
longitudinal chords and the intestine. P. 
armi^cr Gerlach. 1955, does not possess 
these cells and is, therefore, of uncertain 
status. Moreover, since the spicular appa- 
ratus turns out to be of prime importance 
for classification, a number of species 
known only as females or juveniles have to 
be considered species inqidrendae. These 
are: P. keiemis, P. longisetosus, and P. 
ohtusicaiidutus, all Kreis, 1932. 

The remaining species fonn an extremely 
closely related group and are difficult to 
separate on the basis of existing informa- 
tion. We agree with Inglis (1962) that the 
shape of the distal end of the spicula might 
serve as an important taxonomic character, 
and use it as such in the differentiation of 
our species, although we realize that the 
data in the literature on which some of our 
conclusions are based probably are insuf- 

L/.sY of species of Prooxciiolaimus {and 
their synonyms) 

P. meiiastoma (Eberth, 1863) [syn. 
Oncholaimus mcii^astoma Eberth, 1863, Pro- 
oncholaimiis mcditernineiis Schuurmans- 
Stekhoven, 1943, nom. iiov. for P. mcga- 
.itoma Micoletzky, 1924 nee Eberth, P. 
me<^astovui var. neapoUtanus Micoletzky, 
1924, P. neapoUtanus (Micoletzky) Kreis, 
1934]; P. eherthi (Fihpjev, 1918) [Syn. 
Metoncholaimus ehertlii Filipjev, 1918]; 
P. ornatus Kreis, 1932; P. Chit- 
wood, 1951; P. hanijidensis Inglis, 1962; 
P. hastatus n. sp. 

Our new species, P. hastatus, is most 
closely related to P. hamjulensis, in that the 
distal extremity of the spicula possesses a 
distinct barb which is separated from the 
subterminal swelling of the spicula by a 

"handle." The two can be separated as 


Handle between subterminal swelling of spieula 
and barb very short; length of spieula 119- 

135 m; tail with long terminal setae 

P. hanijidensis Inglis, 1962 

Handle between subterminal swelling and barb 
about the same length as the barb itself; 
length of spicula 90-95 p.- terminal setae very 
short P- hasiatiis n. sp. 

Prooncholaimus hasfoius new species 
Plate XI, fig. 20 a-e 

L = 2.10-2.53 mm; w = 80-100 ^; esoph- 
agus: 6 , 310-375 11, 9 , 425 /x; tail: 6 , 
120-150 /., 9, 175 ,x; Vu = 73%. Head 
diameter 28 /x. Labial capsule well devel- 
oped, lolled. Cephalic setae 5 ^, subequal. 
Amphid in i , 10-11 ^ = 307^ of c.b.d., in 
juvenile 9 = 7 /x. Buccal cavity 40 X 21 /x, 
teeth 31 + 23 /x. Excretory pore on level 
of cephalic setae in adults. Esophagus with 
posterior pyriform swelling. Spicula 90-95 
/x long, funnel-shaped proximally, distally 
with subterminal swelling, handle and 
well-developed barb. Gubernaculum 15 /j. 
long, rather stout. There is an indication 
of a bursa and there are four pairs of 
adanal setae. More setae can be found 
anterior to the anus and subventrally along 
the tail. Anal body diameter 33 /x. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4073, Type slide No. 69. 
Type locality, M-8, Biscayne Bay. 

Representation in samples studied. — M- 
8, Biscayne Bay. 

VISCOSIA deMan, 1880 
Type species.— Oncho/a/Vnus viscosus Bas- 
tian, 1865: 136, pi. 11, figs. 131-133. 

There is a group of species in this genus 
in which the cephalic setae are reduced to 
papillae or even to barely visible shallow 
pits in the cuticle. This morphological 
feature, in addition to the fact that in this 
genus the spicula offer hardly any dis- 
tinguishing characters, renders classifica- 
tion particularK' difficult. Stress has to be 

Florida Marine Nematodes • Wiescr and Hopper 259 

laid on size of aniphids, shape of buccal 
cavity and teeth, arrangement of male cir- 
cumanal organs like papillae and bursa, 
and shape of tail. 

Key to Species of Viscosia Hamng 
Cephalic Papillae 

1. Buccal ca\ it>' dixided In' strong cuticular 

ring into two chamliers; one side of buc- 
cal ca\ity weakh' cuticularized ( resem- 
bling the condition in Oncholaimellus); 
S amphids 607r of c.b.d.; pharyngeal 
valve about three times its own length 

behind buccal ca\ity 

V. oucholaimclloides n. sp. 

Buccal cavity not divided by cuticular ring 
( at most a faint line can be seen ) ; buccal 
wall well de\eloped all around; amphids 
not more than 50% of c.b.d. wide; 
phar\Tigeal valve not more than its own 
length l:)ehind buccal cavity 2 

2. Male with bursa (or circumanal "alar mem- 

brane" sensu Chitwood, 1960, who was 
the first to point out the importance of 

this character) .3 

Male without bursa 4 

3. Male amphids % of c.b.d.; walls of buccal 

cavity strongly cuticularized; S tail 6 

a.b.d. V. papillata Chitwood, 1951 

Male amphids 40-45% of c.b.d.; walls of 
buccal ca\itv normally developed; i tail 

3.4^.6 a.b.d. 

V. papillatoicles Chitwood, 1960 

4. Nhile amphids at most 33% of c.b.d. wide __ 5 
Male amphids 4.5-50% of c.b.d. wide 7 

5. Cephalic papillae distinct; long tooth not 

quite reaching to anterior end of buccal 
cavity; i with 3 preanal and 3 postanal, 

small, setose papillae ..^ 

V. keiensis Kreis, 1932 

Cephalic papillae indistinct, long tooth 
reaching to anterior end of buccal cavity; 
c^ with not more than 1 preanal and 1 
postanal papilla 6 

6. Male amphids 33% of c.b.d.; 1 preanal 

papilla; spicula open proximally 

V. nuda Kreis, 1932 

Male amphids 25% of c.b.d.; 1 preanal and 

1 postanal papilla; spicula knobbed 

V . mcridionalis Kreis, 19.32 

7. Cephalic papillae distinct; longest tooth not 

quite reaching to anterior end of buccal 

cavity 8 

Cephalic papillae indistinct to seemingly ab- 
sent; longest tooth reaching to anterior 

end of buccal ca\ity 

V^. cdinleyensis Kreis, 1932 

8. Male tail cylindrical, 5 a.b.d. long 

- V. nicaraguensis (Gerlach, 1957) 

Syn. V . papillata var. nicaraguensis 
Gerlach, 19.57 

Male tail filiform, 8-13 a.b.d. long _... 9 

9. Male with onh' traces of circumanal papil- 
lae V. glabra (Bastian, 1865) 

Male with 6 pairs of setose circumanal papil- 
lae V. macramphida Chitwood, 1951 

Remarks. — The relationship of the two 
latter species is vm certain because no good 
figures of the male amphids in V. gjahra 
have been published. However, the figures 
of female heads as given by De Coninck 
(1944) and Schuunnans-Stekhoven (1950) 
show the amphids to be V?, to % of the c.b.d. 
in width, or nearly as wide as the stoma, 
from \\'hich it may be concluded that the 
male amphids should be just as wide as those 
of V. macramphida. The only good differ- 
ence we could find between the two species 
in question seems to be the size and arrange- 
ment of the genital papillae in the male. 
We consider this difference to be of spe- 
cific value for the time being since figures 
of the male genital region in V. glabra, as 
given bv two such excellent obsen'ers as 
de Man (1890) and Micoletzky (1924a), 
fail to show anything that can be compared 
with the distinct setose papillae that Chit- 
wood ( 1951 ) and we ( see below ) found 
in V. macramphida. 

Doubtful species are: V. linstowi (de 
Man, 1904), V. pscudogjahra Kreis, 1932, 
V. duhiosa Kreis, 1932, V . fatigans Filipjev, 
1946, all of which are known only as juve- 
niles or females, and V. pelliicida (Cobb, 
1898) nee Allgen, 1959, of which no figures 
were given. Tlie statement in the key by 
^^'ieser ( 1953 ) , copied from Kreis ( 1934 ) , 
to the effect that in V. linsfoici the excre- 
tory pore lies only two stomatal lengths 
from the anterior end, was based on an 
erroneous translation of de Man's text. 

Viscosia oncholoimelloides new species 
Plate XI, fig. 21 a-c 

L = 1.95 mm; w = 22 /x; esophagus = 300 
/x; tail: 6 , 140 p.. Head diameter 12-13 jx. 
Lips distinct; cephalic papillae distinct. 
Buccal cavity separated into two chambers 

260 Bulletin Museum of Compomtive Zoology, Vol. 135, No. 5 

by cuticular ring; anterior portion 5 /x, pos- 
terior portion S-9 /x deep. The large sub- 
ventral tooth is anchored to the buccal 
wall from the cuticular ring back to about 
the middle of the posterior chamber; from 
there on the buccal \\'all is weakly devel- 
oped, resembling the condition in Oncho- 
luimcUus. The two smaller teeth are nor- 
mally developed as in other species of 
Viscosia and not reduced as in Onchohimcl- 
Iiis. Large subventral tooth 10 /i long. 
Pharyngeal valve 7-8 /x behind buccal cav- 
ity. Scattered papillae in cervical region. 
Excretory pore just behind nerve ring, 165 
Ij. behind anterior end. Male: spicula dag- 
ger-shaped, 17 IX long (1 a.b.d.). There 
are five pairs of circumanal papillae and 
one more pair of preanal and postanal 
papillae, respectively. Scattered papillae 
on tail. 

HoJotijpc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4073, Type slide No. 70. 
Type locality, M-8, Biscayne Bay. 

Representation in samples studied. — M- 
8, Biscayne Bay. 

Viscosia popillafa Chitwood, 1951 
Plate XII, fig. 22 a-f 

Viscosia })(i]uU(it(i Chitwood, 1951: 627, fij 


L= 6, 1.18-1.24 mm, 9, 1.04 mm; w = 
39-47 /x; diameter at base of esophagus 39- 
40 p.. Esophagus 235-250 p. long. Tail 120- 
12.5 fji long. Vu = 67%. Head diameter 16 /x. 
Labial capsule well developed, cephalic 
papillae distinct. Amphid in male 6 /x, in 
female 5 jj. wide, % of corresponding body 
diameter. Buccal cavity 21-22 /x deep, its 
walls particularly strong. Excretory pore 
130-145 /x from anterior end, 1.5-20 /x behind 
ner\'e ring. Excretory cell 30 /x long, 40 /x 
behind base of esophagus. Spicula 24-25 
IL long. Male with circumanal ala contain- 
ing six pairs of papillae, the posterior pair 
the prominent. In addition a pair of 
preanal setae is present. 

Representation in samples studied. — M-7, 

Everglades National Park; V, Vero Beach. 
Geographical distribution. — Copano Bay, 
Texas (Chitwood, 1951), Chesapeake Bay, 
Maryland (Timm, 1952). 

Viscosia macramphida Chitwood, 1951 
Plate XII, fig. 23, a-d 

Viscosia macramphida Chitwood, 1951: 627, fig. 1, 

L = 1.5-1.8 mm; w = 40 /x; diameter at 
base of esophagus 34-39 /x. Esophagus 260- 
305 /x long. Tail 180-195 /x long. Head 
diameter 1.3-16 /x. Labial capsule in- 
distinct, cephalic papillae distinct. Amphid 
7 IX wide. Buccal cavity 20-21 ix deep. 
Excretory pore 30 /x. behind nerve ring. 
Spicula 23-25 fx long. Male with 4-5 pairs 
of setose papillae associated with circum- 
anal ala as in V. papillata. 

Representation in samples studied. — M-2, 
Key Biscayne; M-8, Biscayne Bay. 

Geographical disiribution. — Aransas Bay, 
Texas (Chitwood, 1951). 

EURYSTOMINA Filipjev, 1918 
Type species.— Eurysfomo specfobile Marion, 
1870: 20, 21, pi. E, figs. 1-1 b. 

This genus has been reviewed by Inglis 
( 1962 ) , who bases his classification almost 
entirely on the shape of the gubernaculum. 
While we agree that quite generally in 
marine nematodes more emphasis should 
be placed on the structure of the male 
genital armature, we feel this feature should 
not dominate to such an extent that other 
characters are ignored. Thus, Inglis con- 
siders E. americana Chitwood, 1936, and 
E. minutisculae Chitwood, Timm, 
1952, to be conspecific because of the simi- 
larity in the shape of their gubemacula, al- 
though (in Inglis' own words): "Chitwood 
reports only one row of denticles in the 
buccal cavity and the absence of ocelli 
while Timm, in describing his E. minutis- 
culae, mentions three rows of denticles and 
ocelli." Moreover, it could be added, Chit- 

Florida Marine Nematodes • Wiescr and Hopper 261 

wood shows an almost rectangularly bent 
spiculum, Timm a semiciicular one, Chit- 
wood a short terminal excretoiy duct, Timm 
a long one, Chitwood a tail provided with 
setae, Timm a naked one, etc. On the other 
hand, Inglis considers E. americana Chit- 
wood, 1936, noi to be conspecific with E. 
americana of Chitwood, 1951, because of a 
slight difference in shape between the two 
gubernacula in question, although all other 
characters in the two descriptions seem to 

This approach ignores the possibility of 
small variations in structural features and 
overrates differences taken from the illus- 
trations of authors of different reliability. 

Our material contained one species which 
is sufficiently close to £. miniitiscidae as 
described by Chitwood, 1951, and by Timm, 
1952, to identify it with tliis species. The 
gubernacula of our male specimens are 
very similar to that figured by Timm, 
whereas the spicula more closely resemble 
Chitwood's figure. The heads of the speci- 
mens described by Chitwood, by Timm, 
and by us are so similar that it would seem 
unwarranted to refer them to different 
species, although we feel that Chitwood 
and not Timm was correct in the intei*preta- 
tion of the excretory gland. 

Eurysfomina minutisculae Chitwood, 1951 
PlateXIII,fig. 24, a, b 

Eunjstomina minutisculae Chitwood, 1951: 629, 
fig. 3, d-g. 

L = 3.1-3.3 mm; w = 45 /x; diameter at 
base of esophagus 40-45 jx. Esophagus 660- 
670 /I long. Tail 114-122 /x (3 a.b.d.) long. 
Head diameter 20 /x. Cephalic setae 9 + 5 
IX. Buccal cavity 17-18 /x long, separated 
into two chambers by three rows of denti- 
cles. Amphid aperture transversely oval, 
displaced dorsally. Excretory pore on level 
of amphids; terminal excretory duct short. 
Ocelli 58 jx from anterior end. Gland-like 
structures ( ? ) present in anterior neck re- 
gion. Spicula 63-66 /x long, apophysis of 
gubernaculum 26-31 /x long. Supplements 
70-75 IX and 130-155 /x preanal. Three pairs 

of preanal setae present, one subventral and 
longer that the two submedian pairs; cuticle 
in vicinity of anterior pair thickened. 

Representation in samples studied. — M-1, 
Kev Biscayne; M-7, Everglades National 

Geographical distribution. — Aransas Bay, 
Texas (Chitwood, 1951), Chesapeake Bay, 
Maryland (Timm, 1952), ? San Salvador 
(Gerlach, 1955: E. aff. minutisculae). 

ILUUM Cobb, 1920 

Type species.— ////um ex/7e Cobb, 1920: 
261,262, fig. 45. 

In 1920 Cobb described a genus, Illium, 
which has never been found again until noN\'. 
Our material contains two species of which 
one seems to be Cobb's I. exile, while the 
other is new. So far no males are known 
in this genus, for which reason we have 
to deviate from our rule and base the fol- 
lowing two descriptions on females only. The 
shape of the buccal cavity is very much 
as in Symplocostoma or Fohjgastrophora 
except that there are no teeth. This might 
also be the reason for the weaker develop- 
ment of the cuticular rings surrounding the 
buccal cavity. Our two species are easily 
distinguished by the dimensions of head, 
amphids, and buccal cavity, as well as by 
the position of the amphidial "sensilla" with 
respect to the buccal cavity. 

iWium exile Cobb, 1920 
PlateXIII, fig. 25a-d 

lUium exile Colib, 1920: 261, 262, fig. 45 

L = 2.02-2.22 mm; w = 23-30 /x; esopha- 
gus = 438—492 ix; nerve ring = 240 /x behind 
anterior end; tail = 110-122 ,x; Vu =60%. 
Head rounded, with six lips and, perhaps, 
minute labial papillae. There might be a 
second circle of cephalic papillae, but all 
these organs are extremely difficult to see. 
Buccal cavity consisting of vestibulum and 
two chambers, separated by two cuticular 
rings; the two chambers measure 7 /x in 
length, greatest width is 3 /x. Amphids con- 

262 BuUctin Museum of Compavalivc Zoology, Vol 135, No. 5 

sisting of opening, about 3 /x wide, pouch, 
duct and an unusual structure which we 
call "sensilla." There is no indication of an 
ocellus as assumed by Cobb. Distance of 
sensilla from anterior end = 12 /x, c.b.d. = 
10 fx. Excretory pore a short distance be- 
hind nei-ve ring. Anal body diameter 17 /x. 

Representation in samples studied. — M-S, 
Biscayne Ra>'. 

Geo<iraphieal distribution. — Jamaica 
(Gobi), 1920). 

////urn libidinosum new species 
Plate XIII, fig. 26, a, b 

L = 2.28 mm; w = 70 /x; esophagus = 570 
/x; nerve ring = 258 p. behind anterior end; 
tail = 200 /-.; Vn = 60?r . Head as in the fore- 
going species but much larger. Buccal 
cavity 10 X 5 /x, walls more cur\ed than in 
the foregoing species. Amphids 5 /x wide, 
sensilla 15 fx behind anterior end, filled with 
fragments of a dense material. Excretory 
pore 25 fx behind nerve ring. Anal body 
diameter 33 p.. 

Ilolotype specimen. — Female; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4073, Type slide No. 71. Type 
locality, M-8, Biscayne Bay. 

Representation in samples studied. — M-8, 
Biscayne Bay. 

POlYGASJROPhORA de Man, 1922 
Type species.— Po/ygasfrop/iora aiienua\a 
de Man, 1922a: 131, 132. 

In this genus four species each possessing 
seven esophageal bulbs have been described, 
to which we shall add a fifth one. Of these 
five species, P. tenuicoUis (Allgen, 1951) 
is best considered a species inquirenda as 
advocated by Chitwood ( 1960 ) , and not 
a synonym of P. heptabulha as suggested 
by Wieser ( 1953 ) . The remaining four 
species are separated mainly by the arrange- 
ment of their cephalic setae, but we are 
not so sure whether this character will turn 
out to be as stable as we assume at present. 

List and Short Characterization of Valid 

PoLVGASTROPHORA spp. Fosscssiug, Seven 

Esop]uig,eal Bulbs 

P. maior Sehulz, 1932: Submedian ce- 
phalic setae in both sexes very unequal in 
length (1-2 and ^f> of c.b.d., respectively). 
Excretory pore 130-150 /x from anterior ex- 
tremity. Spicula length = % of tail. Guber- 
naculum conical. 

P. heptabulba Timm, 1952: Submedian 
cephalic setae reduced in £ , one head 
diameter long and subequal in 9 . Excretory 
pore 44 /x behind anterior end. Spicula 
length = Vj of tail. Gubernaculum absent. 

P. septemhulba Gerlach, 1954: Subme- 
dian cephalic setae subequal ( 7-6 /x ) in S , 
very unequal in 5 (shorter ones about 2 /x). 
Excretory pore 41-52 fx from anterior end. 
Spicula length = % to % of tail. Guber- 
naculum alxsent (Gerlach, 1954) or small 
(Chitwood, 1960). 

P. eda.x n. sp.: Submedian cephalic setae 
very unequal in length in both sexes (8- 
7 + 2 /x ) . Excretory pore 48-50 ^ behind 
anterior end. Spicula length = ^-i of tail. 
Gubernaculum plate-shaped. 

Our new species P. eda.x is related to P. 
septembulba from which it can be distin- 
guished by the characters presented in the 
above list. 

Polygostrophora edax new species 

Plate XIV, fig. 27, a-c 

L = 2.65-2.90 mm; w = 105-110 ^; esoph- 
agus = 550-600 p.; tail: <i , 175 /x, 9 , 165 
fx (3.5-4 a.b.d.); Vu = 53%. Head diam- 
eter 10 /<. Mouth opening crenate. No 
labial papillae. Cephalic setae in 6,7 + 2 
^, in 9,8 + ?, indistinct. Amphids oval, 4.5 
IX wide in S  Buccal cavity 14 X 7 /x, with 
two faint cuticular rings at about its middle 
and one strong basal band that is resolvable 
into elongated cuticular bodies. Refractory 
bodies (ocelli?) 18-20 ^a behind anterior 
end. Excretory pore 48-50 /x behind an- 
terior end, ampulla about 72 fx. Male: 
Spicula 80 IX, knobbed proximally. Guber- 
naculum faint, plate-shaped. Subventral 
circumanal setae and along the tail. 

Florida Marine Nematodes • Wieser and Hopper 263 

, TIHI -. 




Figure 2. Types of preanal supplements in the Cyatholaimidae. a — Pomponemo (type A); b — Longicyatholaimus (type 
B); c — Paracanthonchus (type C-1); d — Paracyatholaimus (type C-2, two forms). 

Holotypc specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection 
Number 4075, Type slide No. 72. Type 
locality, V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 


The genera within the subfamily Cy- 
atholaiminae can be divided into t\vo groups 
based on the presence or absence, in the 
male, of preanal supplements. In those 
genera in which the males possess preanal 
supplements, a further division is possible 
utilizing the structure and arrangement of 
these organs. \\'ieser (1954) presented a 
key to the genera of this subfamily, a key 
that still remains useful, but requiring sev- 
eral additions and emendations. 

The genera with preanal supplements 
can be separated into the following three 
basic types (A, B, C) of which the last 
is subdivided into forms with "tuboid" sup- 
plements (C-1) and fonns with "setose" 
supplements (C-2) (see Text-fig. 2). 

Type A ) Supplements large, complicated, 
consisting of several elements; cuticle be- 
tween supplements lamellated. Genera in- 
cluded: Fomponema Cobb (syn. Endolai- 
miis Filipjev), Niimmoceplialus Filipjev 
(syn. Haustrifera Wieser), Craspodema 
Gerlach (syn. Kraspedonema Gerlach) and 

Aiui.xuncJiiiim Cobb. Tentatively included 
is Dispira Cobb. 

Type B ) Supplements cup-shaped, large 
to minute, with narrow ducts leading to 
the cups. Genera included: Longicyatholai- 
mus Micoletzky, Xyzzors Inglis, and 
Biarmifer Wieser. 

Type C-1 ) Supplements tubular, large to 
medium. Genera included: Paracanthon- 
chus Micoletzkv, Acanthonchus Cobb, 
SeuratieJhi Ditlevsen and ParaseuraticUa 

Type C-2) Supplements "setose," small. 
This type can probably be derived from 
type C-1, the "setose" papillae described by 
some authors most likely being minute 
tubuli with very narro\\' ducts. Genera in- 
cluded: Paracyatholaimus Micoletzky (in 
Text-fig. 2, represented by two forms), and 
Paraeyatholaimoides Gerlach. 

Those genera that definitely do not 
possess preanal supplements are: Cyatholai- 
mus Bastian, XenocyatJwkiimus Gerlach, 
MetaeyathoJuimus Schuurmans-Stekhoven, 
and PhyUohimus Murphy. 

Other genera are considered doubtful. 

There are two main difficulties in classi- 
fying genera of this subfamily: first, separa- 
tion of type C-1 from type C-2, since the 
difference between small tubuli of the 
Paracanthonclius-type and the so-called 
"setose papillae" of Paracyatholaimus may 
be only slight; second, deciding whether 
a species is devoid of supplements or pos- 


Bulletin Muscuin of Couiixiiativc Zoology, Vol 135. No. 5 

sesses small, cup-shaped papillae of the 
type B. A case in point is Lon<i,icycit1ioIaimiis, 
in which species with and species without 
supplements ha^•e been deserib(>d (see be- 
low ) . 

The genus Choniokiimus, referred to this 
subfamily by many authors and included in 
his key by Wieser ( 1954 ) , will have to be 
transferred to the Choanolaiminae since the 
redescription of the type species, CJi. papil- 
latiis Ditle\'sen, bv Cerlach ( 1964), and the 
description of Ch. jHuiiciis- Gerlach, 1957, 
have made it clear that in this genus the 
buccal cavity is of a shape (luite different 
from oth(n- cyatholaimids. Consequently, the 
species referred to Clioniolaimus on the 
strength of their preanal supplements but 
with a typical cyatholaimid buccal cavity, 
i.e., Ch. macrodcntatus Wieser, 1959, and 
Ch. iciescri Inglis, 1963, are to be trans- 
ferred to Lon0cijathoJ(iimus (see below, 
p. 265). CyathoJaimu.s fautraensis Allgen, 
referred to Chonioloimus by Wieser ( 1954) 
is better placed with Niimmocephalm ( see 
Gerlach, 195Sb). 

POMPONEMA Cobb, 1917 
Type species.— Pomponema mirobile Cobb, 
1917: 118, fig. 3. 

This genus is characterized by the strong 
developmcMit of the l:)uccal cavity, in which 
the vestibular ribs seem to function as 
particularly mobile clasping organs; the 
dorsal tooth is powerful and opposed either 
b\ two strong subventral teeth or by a 
great number of denticles. Further char- 
acteristics are the heterogeneous cuticular 
ornamentation, the lateral differentiation 
of the latter, and the peculiar male supple- 
ments. Closely related to Fomponeina is 
the genus Nt(mmocej)haIus Filipjev (syn. 
Housirifera Wieser) which has less well 
developed vestibular ribs and teeth, and 
in which the cuticular markings are more 
uniformly dot-like. Lon^icijatholmmus line- 
dins Gerlach, 1952, which was referred to 
Pomponema by Wieser (1959), is perhaps 
better placed with Nummoceplialus. 

Key to Species of Pomponema 

1. Lateral differentiation beginning on level 

with buccal ca\ity; in the cervical region 
the 2 longitudinal rows are half the c.b.d. 
apart __.- P. multipapillatum (Filipjev, 1922) 
Lateral differentiation beginning approxi- 
mately at end of esophagus; 2-4 longitu- 
dinal rows which are far less apart than 
half the c.b.d. 2 

2. Cephalic setae in two circles, 4 -f 6 

P. segregatuin Wieser, 1959 
Cephalic setae in one circle 3 

3. Suliniedian pairs of cephalic setae very un- 

equal in length. Dorsal tooth opposed by 
groups or rows of smaller teeth or 

denticles 4 

Submedian pairs of cephalic setae subequal. 
Dorsal tooth opposed by two subventral 
teeth 5 

4. Male amphids 6-6.5 turns, 50% of c.b.d. 

wide P. stomachor Wieser, 1954 

Male amphids 4.5 turns, 35% of c.b.d. wide 
P. pohjdonta Murphy, 1963 

5. Male amphids 6-6.5 turns, 90% of c.b.d. 

wide P- mirahilc Cobb, 1917 

Male amphids 4.5 turns, 65% of c.b.d. wide .. 
P. tcsscldtuin n. sp. 

Pomponema tesselatum new species 
Plate XIV, fig. 28, a-d 

L = 1.53 mm; w = 42 /x; esophagus = 360 
/x. Head diameter 25 /x. Labial setae 14 ^. 
Cephalic setae 16 + 13 ^. Buccal cavity 
spacious, vestibulum protrusible, its ribs 
giving the appearance of prehensile clasp- 
ing organs which are linked by joints to 
the buccal wall; dorsal tooth large, hollow, 
opposed by two similar though smaller 
subventral teeth. Amphids 19 ^i = 68% of wide, describing 4.5 turns. Cuticular 
ornamentation complex and heterogeneous; 
in cer\'ical region each annule at high focus 
with slit-like markings, at low focus of 
tesselated appearance; what are seen as 
dots in lateral view are actually short 
columns between the outer and the inner 
layer of the cuticle; these columns become 
rather thin from the mid-cervical region on, 
and the dots, consequently, smaller. There 
are two rows of dots per annule. A lateral 
differentiation in the form of four longitu- 
dinal rows of larger and more widely spaced 
dots begins at about the end of the esopha- 

Florida Marine Nematodes • Wiescr and Hopper 265 

gus. Many pores are arranged in more or 
less regular longitudinal rows all along the 
body. Spicula 45 /x, gubernaculum 36 /x, 
consisting of two portions, the distal one 
with lateral projections. There are 15 pre- 
anal supplements of characteristic shape. 
Between the supplements the cuticle gives 
a lamellated appearance. One preanal seta 
and many setae on the tail in four longitu- 
dinal rows. Tail 132 /x, a.b.d. 36 /x. 

Holofype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4068, Type slide No. 73. Type locality, 
M-3, Key Biscayne. 

Representation in samples studied. — M-3, 
Key Biscayne. 

Type species.— Cyafholaimus longicaudatus 

de Man, 1878: 111, 112, pi. 10, fig. 

16, a-c. 

This genus is characterized by the spicu- 
lar apparatus, which is of a general shape 
found in some species of Faracanihonchus 
and Paracyatholaimiis, in combination with 
the fact that the preanal supplements are 
never setose or tubular. In the type species, 
L. longicaudatus (de Man), indistinct papil- 
lae of type B (see above, p. 263 were ap- 
parently seen by de Man ( 1878 ) and by 
Kreis ( 1928 ) but not by Schuurmans-Stek- 
hoven (1943). Since then other species have 
been described in which supplements were 
not mentioned. It would simplify matters 
if one were permitted to assume that in all 
these cases the small cup-shaped supple- 
ments were so indistinct as to have been 
overlooked, but the possibility that there 
exist species truly devoid of supplements 
must be considered. In such cases dif- 
ferentiation from other genera without sup- 
plements, particularly from Cijatholaimus, 
would have to be based on the shape of the 
spicular apparatus and, to a lesser extent, 
on the shape of the tail. 

Further characteristics mentioned by 
Micoletzkv are the lateral differentiation of 

the cuticular ornamentation and the long, 
filiform tail. However, species with fairly 
short, though always slender, tails have been 
described, which in other respects fit the 
generic diagnosis. 

Classification of the species is rendered 
difficult by the uncertainty as to how the 
reputed absence of preanal supplements is 
to be judged. For the present, we shall 
accept this feature at its face value. This 
provides for an immediate separation of 
the species into two groups. Group A con- 
taining species described with supplements, 
and Group B, containing species in which 
no supplements were reported. The species 
contained within these groups are as fol- 
lows : 

Longicyatliolaimus species group A: L. 
longicaudatus (de Man, 1878); L. minor 
(Cobb, 1898); L. effilatus (Schuurmans- 
Stekhoven, 1946); L. .stekhoveni Wieser, 
1954 (syn. L. effikitus Schuurmans-Stek- 
hoven, 1950 nee 1946); L. quadriseta Wieser, 
1954 nee 1959;^ L. macrodentatus (Wieser, 
1959) new combination (syn. Choniohimus 
macrodentatus) and L. tvieseri (Inglis, 
1963) new combination (syn. Choniolaimus 

Longicyatholaimus species group B: L. 
trichurus (Cobb, 1898); L. zo.sterae Allgen, 
1933; L. ehoanolaimoides (Schuunnans- 
Stekhoven, 1942); L. continus Filipjev, 
1946; L. duhius Filipjev, 1946; L. trichocauda 
Gerlach, 1955, and L. dayi Inglis, 1963. 

Species of doubtful status are: L. heteru- 
/•j/.s (Cobb, 1898); L. tenuicaudatus (Saveljev, 
1912) and L. fiJicaudatus Schuurmans-Stek- 
hoven, 1950. 

As the species we found belongs to group 
A, a key to the species of this group is 

^ Note: The species described as L. quadriseta 
Wieser liy Wieser, 1959, is equipped with setose 
and not cup-shaped supplements. Since there are 
other differences from the type (smaller amphids, 
shape of gubernaculum) we consider L. quadriseta 
Wieser sensu Wieser, 1959, to be a different species 
and refer it to Paracijatholaimus under the name 
Paracyatholaitniis pugetteiisis new name and 
new combination. 

266 Bulletin MimiiDi of Comparative Zoology. Vol. 135, No. 5 

Key to Species of Longicyatholaimus Group A 

1. Lateral differentiation of cuticle in 4 longitu- 

dinal rows L. minor (Cobb, 1898) 

Lateral differentiation of cuticle irregular . 2 

2. Posterior portion of tail filifomi, much longer 

than conical portion 

L. lon^icaudattis (de Man, 1878) 

Posterior portion of tail cylindrical, not longer 
than conical portion 3 

3. Gubernaculuni distall> truncate or notched: 

3 preanal supplements — L. effilatus 

(Schuunnans-Stckhovcn, 1946) and L. stck- 
hoveni Wieser, 1954 
Gubernaculum distally with well dexeloped 
teeth or processes; 6-7 preanal supple- 
ments 4 

4. Male amphids about 60% of c.b.d. wide; 

gubernaculum distally witli three digiti- 
form processes of rather unequal shape „___ 

L. wieseri (Inglis, 1963) n. comb. 

Male amphids 30-40% of c.b.d. wide: guber- 
naculum distally widi 3-4 equal-sh:iped 
teeth 5 

5. Cephalic .setae digitiform, 8-10 + 5-6 /n long. 

Preanal supplements 5 + 2, the 5 anterior 
ones regularly spaced: gubernaculum with 

3 distal teeth 

___ L. macrodentatus (Wieser, 1959) n. comb. 
Cephalic setae conical, slender, subequal, 
12 -f 13 II long; preanal supplements 4 + 2, 
the first one almost three times as far 
from the 2nd as the latter from the 3rd; 

gubernaculum with 4 distal teeth 

L. annac n. s-p. 

Longicyatholaimus annae new species 
Plate XV, fig. 29, a-c 

L = 1.96 ami; w = 70 /x; esophagus = 300 
/J.. Head diameter 33 /x. Labial papillae 
setose, stout, 4.5 /x long. Cephalic setae 
13 + 9 ix. Buccal cavity deep, with one large 
dorsal tooth, two small subventral teeth 
and cuticular ridges. Amphids in i 13 
/x = 31% of c.b.d. wide, 4.5 turns. A group 
of dorsolateral cervical setae a short dis- 
tance behind amphids. Cuticle with annules 
and homogeneous rows of dots; no lateral 
differentiation except on tail; there is one 
row of dots per annule in the anterior cervi- 
cal region, but two rows can be found on 
the remainder of the body. Many pores ar- 
ranged in more or less longitudinal rows. 
Spicula 70 /x, somewhat S-shaped, with 
velum in distal half, knobbed proximally. 

Gubernaculum 64 ^, stout, distally with 4 
equal-sized teeth. There are 6 small but 
distinct cup-shaped preanal supplements, 
the anteriormost one at a distance of 210- 
230 fj. preanally; the distance between the 
first and the second papillae is about as 
great as that between the latter and the 
anus. Tlie two posteriormost papillae are 
more closelv spaced than all the others. 
Tail 420 /x, a.b.d. 60 /x. 

Holotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4068, Type slide No. 74. Type locality, 
M-3, Key Biscayne. 

Representation in samples studied. — M-3, 
Key Biscayne. 

Remarks. — This species is named after 
Mrs. Ann Hopper, friendly hostess to wan- 
dering nematologists. 

XYZZORS Inglis, 1963 

Type species.— Xyzzors fifzgeraldae Inglis, 
1963: 544-546, figs. 25-29. 

According to Inglis ( 1963 ) Xyzzors is 
characterized by irregular lateral differen- 
tiation of the cuticle, cup-shaped preanal 
papillae, large and well-developed buccal 
armature and some structural peculiarities 
of the spicular apparatus. However, none 
of these characters separates Xyzzors un- 
equivocally from Longicyatholaimus. The 
cuticle and the supplements are of the same 
type as found in the latter genus, the 
gubernaculum is somewhat larger than in 
most species of Lon<^icyathoJaimus but, 
e.g., L. dubiiis Filipjev, 1946, has a guber- 
naculum of exactly the same shape. More- 
over, the figures given by Inglis do not 
justify his statement that "the spicules are 
much more elaborate than is usual in 
species of the Cyatholaimidae." The two 
featiues which could perhaps serve as char- 
acters distinguishing Xyzzors from Longi- 
cyatJiolaimus are the nearly conical tail 
and the buccal annature which indeed 
seems to be somewhat more elaborate than 
observed in the latter genus. 

In oiu" new species the spicular apparatus 

Florida Marine Nematodes • Wiescr and Hopper 267 

is simpler than in X. fitz^i^eraldac Inglis, 
and the proximal ends of the spicula are 
not doubled. There are 3 cup-like supple- 
ments followed posteriorly b\' two indistinct 
ducts, whereas Inglis reports 6 cup-like sup- 
plements. The amphids describe 4 turns in 
our species as against 6.75 in A', fitzgeraldae. 

Xyzzors inglisi new species 
Plate XV, fig. 30, a-c 

L = 1.25 mm; w = 54 ^; esophagus = 240 
/x. Head diameter 28 /x. Labial papillae 
stout. Cephalic setae 12 + 10 /x. Buccal 
cavity deep, with one large dorsal tooth, 2 
pairs of subventral teeth and one conspicu- 
ous ridge that surrounds the ventral half of 
the buccal cavity. Amphids 16 /i = 487c of 
c.b.d. wide, 4 turns. Cuticle as in the fore- 
going species. Spicula 52 /x, with xelum; 
proximally the inner edges are more strongly 
cuticularized, distally the outer edges. 
Gubernaculum 38 fx, distally expanded and 
dentate. There are 3 preanal supplements 
that consist of a cup-shaped portion ( pro- 
truded in Fig. 30, b, c) and a duct leading 
to the latter. The distances are: 25 ix from 
anus to posterior papilla, 45 ix from this to 
the next one, 30 /x to the anterionnost one. 
Between the last papilla and the anus, two 
minute, indistinct ducts can be discerned. 
Tail conical, 108 /x, a.b.d. 44 /x. 

Uolotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4075, Type slide No. 75. Type locality, 
V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 

PARACAN7HONCHL/S Micoletzky, 1924 
Type species.— Cyafho/a/mus coecus Bastion 

sensu de Man, 1889b: 204-207, pi. 

7, fig. 10, a-g. 

The species of this genus were grouped 
by W'ieser ( 1954 ) who used the shape of 
the gubernaculum as the main distinguish- 
ing feature. The shapes of gubernacula 
representing species belonging to groups 

A, B, and C, respectively, in Wieser's key, 
are shown in Text-figure 3. There are, how- 
ever, species which do not fit readily into 
these three categories, for example, those 
that have a large gubernaculum subtermi- 
nally dilated and tapering towards a pointed 
or spoon-shaped distal tip. This type of 
gubernaculum represents a transition be- 
tween groups A or B and C, and is also 
shown in Text-figure 3. Species with such 
a gubernaculum are mainly P. mens Wieser, 
1954, and the new species to be described 
below, but P. anii^idatiis ( Schuurmans-Stek- 
hoven, 1950), P.hatidus Gerlach, 1957, and 
P. miitatus \\'ieser, 1959, come rather close. 
P. platypus n. sp. is separated from its 
closest relative, P. mens \\^ieser, 1954, 
mainly by the number, size, and arrange- 
ment of the preanal tubuli, and by the size 
of the spicular apparatus. 

Paracanthonchus platypus new species 
Plate XVI, fig. 31, a-c 

L = 1.18-132 mm; w = 44-48 jx; esopha- 
gus 190 /.; tail: 6 , 135 /x, 9 , 105 ^; Vu = 48%. 
Head diameter 21-23 jx. Labial papillae 
conical. Cephalic setae 4 -h 5 /x. A short 
cephalic capsule present. Buccal cavity 
with well-developed vestibular ribs and 
with medium-sized triangular tooth. Am- 
phids in £,llix — 40% of c.b.d., in $ , 9 
/x = 32% of c.b.d. wide. Excretory pore 27- 
32 jx behind anterior end. Ocelli 47-50 fx 
behind anterior end, with fibrils running 
from the pigment spot forward and back- 
ward, a bit reminiscent of the structures 
described for Acanthonchus rostratus by 
Murphy ( 1963 ) , but a proper lens was not 
seen. Cuticular ornamentation homoge- 
neous. Spicula 36 ^i, gubernaculum 35 /x, 
spoon-shaped, with subterminal dilation and 
tapering towards the tip; there is a lateral 
projection. Preanally there are 4 tubuli, 
22-23 /x long, one large spine (Fig. 31, c), 
and two subventral rows of slender setae. 
On the tail there are short setae and a 
characteristic ventral pair of long setae. 
Anal body diameter 43 /x. 

Holotype specimen. — Male; Canadian Na- 


Bulletin Museum of Coiujxirdiivt' Zoology, Vol. 135, No. 5 

Figure 3. Types of gubernacula in Paracanthonchus. a — P. strandensis (after Schuiz, 1932); b — P. coecus (after Timm, 
1952); c— P. cochlearis (after Gerloch, 1957); d— P. platypus n. sp. 

tioiial Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4073, Type slide No. 76. Type locality, 
M-8, Biscayne Bay. 

Representation in samples studied. — M-8, 
Bisca>ne Bay. 

PARACYATHOLAIMUS Micoletzky, 1921 
Type species.— Cyaf/io/a/mus dubiosus BUt- 
schli, 1874: 284, pi. 7, fig. 31, a, b. 

This genus is characterized by the simple 
giibernaculum which is supposed to hardly 
expand distally, and in the diagnosis by 
Micoletzky ( 1924b ) was described as being 
"ohne Dornen," and by the male supple- 
ments which Micoletzky called "Borsten- 
papillen." These supplements probably are 
not true setae l)ut represent ducts of the 
ParacanthoncJiiis-type with the lumina so 
narrow as to give the impression of setae. 

Gerlach ( 1955 ) described a species, P. 
paucipapilhitus, in which the gubernaculum 
expands distally to form a plate covered 
with rasp-like denticles. The supplements 
consist of strongly protruding conical papil- 
lae with cuticularized, narrow ducts. The 
first character links Gerlach's species to 
ParacantJioncliiis, the second character sets 
it apart from all other species of Paracan- 
tli())icliiis and ParacyalJiokiiuiiis with the 
exception, perhaps, of P. diii^itatus Gerlach, 
1957, in which similar supplements have 
been described and the gubernaculum is 
also rather strongly dentate. Our material 
contained representatives of what at first 

we held to be P. paiicipapillatus. However, 
on closer examination we noted the follow- 
ing differences: 1) distal end of guber- 
naculum with distinct teeth, numbering 
from 4-8, rather than with a rasp-like field 
of denticles, 2) ducts of supplements more 
elaborate, and 3) buccal cavity with one 
large dorsal tooth and three small sub- 
ventral teeth, whereas Gerlach speaks only 
of one dorsal tooth. 

We consider these differences important 
enough to establish a new species, P. pesams 
n. sp., named after the appearance of the 
distal end of the gubernaculum which re- 
sembles a spread bird's foot. This species 
and the two described by Gerlach men- 
tioned above form a rather distinct group 
within the two genera Paracanthonchus and 
Parac\iatholaimus . 

Paracyafholaimus pesavis new species 
Plate XVI, fig. 32, a-e 

L = 1.08-1.17 mm; w = 35-44 ^u; esopha- 
gus = 195-200 /x. Head diameter 18-20 {x. 
I^abial papillae distinct. Cephalic setae 
7-8 + 5-6 /x. Short cephalic capsule. Buc- 
cal cavity with one large pointed dorsal 
tooth and three small subventral teeth. 
Amphids 11 ^a = 44% of c.b.d. wide, ap- 
proximately 5 turns. Cuticular ornamenta- 
tion with slight lateral differentiation in 
cervical and anal region. Spicula 29-31 ix, 
with velum. Gubernaculum 22-23 jj., ex- 
panding distally to a plate which in one 
specimen carried 8, in another 4 small 

Florida Marine Nematodes • Wieser and Hopper 


teeth (PL XVI, fig. 32, c, d ). There are two 
large conical papillae at a distance of 25- 
28 and 50-53 /x, respectively, from the anus. 
The papillae are penetrated by cuticular- 
ized ducts which show some distal elabo- 
rations; their openings are posteriorly di- 
rected. There is a very faint third papilla 
( 10-13 fx in front of the anus ) which might 
actually consist of two closely spaced 
minute tuliuli as found in other species of 
Favucijatholaimus. Tail 80-100 jx long, nar- 
rowing abruptly in distal third; a.b.d. 35- 
40 IX. Spinneret 6-7 ^ long. 

Holotijpe specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4071, Type slide No. 77. 
Type locality, M-6, Everglades National 

Representation in samples studied. — M- 
6, Everglades National Park; V, Vero 

Remarks. — The closely related species, P. 
pancipapiUatus, was originally described 
from the Pacific coast of San Salvador, and 
later from the Congo estuary, and from 
Brazil ( Gerlach, 1957a, b, c). The possi- 
bility that the Atlantic specimens are ac- 
tually representatives of our new species 
is intriguing but remains to be proven. 

HAL/CHOANOM/MUS de Man, 1888 
Type species.— Sp///pf)era robusta Bastion, 
1865: 166, pi. 13, figs. 226, 227. 

Our material contained two closely allied 
species which appear to represent H. qiiat- 
tuordccimpapillatus Chitwood, 1951, and 
//. duodecimpapiUatiis Timm, 1952. As 
neither author specifically stated the tail 
length for their respective species, it was 
necessary to make use of the de Man c 
value in identifying the two populations. 
The two species can be separated on the 
basis of the tail length as well as the shape 
of the spicules and gubernaculum. As the 
heads of both species are very similar, only 
one has been figured, H. duodecimpapiUa- 

Halichoanolaimus quattuordecimpapillatus 
Chitwood 1951 
Plate XVII, fig. 33, a-c 

Halichoanolaimus quattuordecimpapillatus Chit- 
wood, 1951: 639, fig. 7 c. 

L = 6 , 1.7-2.0, 9 , 2.0-2.4 mm; w = 6 , 
60-65, 9 , 70-90 fx; diameter at base of 
esophagus, 6 , 55-70, 9 , 70-83 p.. Vu = 
43-45%. Esophagus in 6 , 270-300 p, in 
9 , 310-340 p long. Excretoiy pore in S , 
150-155 p, in 9 , 162-180 p from anterior 
end. Head 35-40 p wide, with an internal 
circle of 6 labial papillae and an external 
circle of 10 cephalic papillae of which the 
laterodorsal and lateroventral pairs are 
setose. Amphid 15 p wide, 16-19 p from 
anterior end, spiral with 3.75-4 turns. Buc- 
cal cavity typical, containing 3 posterior 
apophyses, the anterior margins of which 
bear comb-like ribs and a medial, retrorse 
tooth. Spicules 88-90 p long, weakly ceph- 
alated proximally and narrowing gradually 
distally. Gubernaculum paired, 40-45 p 
long, with characteristic proximal cephali- 
zation (bottle-cap-opener). Male with 11- 
14 papilloid supplements. Tail initially 
truncate-conoid, then filifomi, in 6 , 265- 
300 p long, in 9 , 315-370 /x long. Spinneret 
10-12 p long. The male has a postanal ven- 
tral depression just prior to the filiform 
portion of the tail. The de Man c value for 
both sexes is 5.8-6.6. 

Representation in samples studied. — M- 
2, Key Biscayne, Everglades National Park. 

Gcog^raphical distribution. — Aransas Bay, 
Texas (Chitwood, 1951), Congo estuary, 
^^'est Africa (Gerlach, 1957b), Cananeia, 
Brazil (Gerlach, 1957c). 

Remarks. — An entire specimen of Spi- 
lophorella paradoxa was found within the 
gut of one female. 

Halichoanolaimus duodecimpopillafus Timm, 
Plate XVII, fig. 34, a-d 

Halichoanolaimus dtioclecimpapillatus Timm, 1952: 
26-28, pi. 5, fiff. 44. 

L = ^ , 1.9-2.2. 9 . 2.1-2.5 mm; w = $, 

270 BuUciin Mii.scuiii of Cotuparatwc Zoology, Vol. 135, No. 5 

68-72, 9 , 80-97 /x; diameter at base of 
esophagus, 6 , 62 /a, 9 , 65-87 fj.. Vu = 40- 
47%. Esophagus in i , 250-280 /x, in $ , 
270-310 ,x. ExcretoiN' pore 130-147 /x from 
anterior end. Head, eephahc sense organs 
and buccal cavity as described above for 
H. quattuordecimpapiUatus. Spicules 90- 
100 IX long, proximal cephalization, broad 
and flat. Distally the spicules narrow more 
abruptly than in the foregoing species. 
Gubernaculum about 50 jx long, of char- 
acteristic shape. Male with 11-13 papilliod 
supplements. Tail elongate-conoid, then 
filifonn, in i , 180-190 //long, in ?, 165- 
230 )x long. Spinneret 9-10 ix long. The 
de Man c value for the i is 10.1-11.3 and 
for the 9 , 9.1-14.3. 

Representation in .'samples studied. — V, 
Vero Beach. 

Geo<i,r(iphical distribution. — Chesapeake 
Bay, Maryland ( Tinim, 1952 ) . 

NEOTONCHUS Cobb, 1933 
Type species. Neotonchus punctatus Cobb, 
1933: 87. 

Comesa Cerlach, 1956: 94. 

Neotonchus lufosus Wieser and Hopper, 
Plate XXVI, fig. 56, a-d 

L = 0.87-1.02 mm; w = 42 /x; esopha- 
gus = 120 IX. Head diameter 20 /x. One 
circle of six minute labial papillae, one 
circle of six short cephalic setae, 3 ix long, 
and four submedian setae of about ecjual 
length. Scattered short cervical setae. Am- 
phids 10 IX = 507c of c.b.d. wide, 4-4.2 
turns. Cuticle with homogeneous puncta- 
tion, the lateral dots somewhat larger and 
more widely spaced than the submedian 
ones. Buccal cavity 14 /x long, with one 
large hollow dorsal tooth and two small 
subventral teeth. Esophageal bulb pyri- 
form, 30 X 25 /x, with two weak interrup- 
tions. Excretory pore 40 fx from anterior 
end. Spicula of shape typical for genus, 
bent at beginning of distal third, 29 /x long. 
Gubernaculum plate-shaped, dilated dis- 

tally. One stiff preanal seta and 20 large, 
complicated supplements. Tail 75 /x long, 
a.b.d. 28 /x. 

Representation in samples studied. — M- 
4, Rickenbacker Causeway. 

Geographical distribution. — Restricted to 
above locality. 


The Desmodoridae was first subdivided 
by Chitwood ( 1936 ) . The original group- 
ings, however, do not appear to be entirely 
satisfactory, especially with regard to their 
generic composition and systematic posi- 
tion. Uncertainties regarding the shape of 
amphids in the Stilbonematinae present 
further difficulties. A key position within 
the family is assumed by S})irinia (syn. 
Spirina ) which not only links the Meta- 
chromadorinae with the Desmodorinae — 
perhaps via Chromaspirina ( see Gerlach, 
1963) — but also shows close affinities to 
the Microlaimidae and the Linhomoeidae 
(Wieser, 1954; Timm, 1962). The discus- 
sion of many genera belonging to the 
Desmodoridae by Gerlach ( 1951b, 1963b ) 
has been a valuable aid in the classification 
of the family. The characters of each sub- 
family are briefly outlined below: 

Brief Characterization of Subfamilies of 

A) Metachromadorinae (Chitwood, 1936): 
Cuticle always finely striated (striation 
sometimes so indistinct as to impart a 
smooth appearance to the cuticle); head 
not sharply set off from striation, non- 
rigid; striation always surrounding am- 
phids; cuticle not tiled; esophageal bulb 
always present, either roimd or elongated; 
buccal cavity typically with well-developed 
annature, except in Spirinia where there 
are only minute teeth. ( For further classi- 
fication, see below. ) 

B) Richtersiinae Cobb, 1933: Cuticle 
striated, with many longitudinal rows of 
spines or hooks; head non-rigid; buccal 
cavity small or wide, unanned; esophagus 

Florida Marine Nematodes • Wiescr and Hopper 271 

cylindrical, without bulb. With the genera: 
Richtersia Steiner, 1916 (syn. RichtersicUa 
Kreis, 1929), and Ptcwniiim Cobb, 1933. 

C) Desmodorinae Micoletzky, 1924: 
Cuticle heavily annulated; head rigid, 
sharply set off from annulation; amphids 
not surrounded by annulation (exceptions 
are Paradesmodora and Metadesmodom in 
which the heavy annulation serves as dis- 
tinguishing characters from the Metachro- 
madorinae); amphids loop-shaped or spiral; 
cuticle not tiled (except, occasionally, on 
the head); esophageal bulb round or 
elongated; buccal cavitv alwavs amied 
with distinct teeth. (For further classifica- 
tion, see below.) 

D) Stilbonematinae Chitwood, 1936: 
Cuticle striated or annulated, not tiled or 
longitudinally broken; buccal cavity shal- 
low-conical or absent, unarmed or with 
minute teeth; head always well defined, 
amphids wholly outside striation, spiral- 
shaped, but apparently sometimes sunk 
into the cuticle so that only the slit-like 
opening is visible; esophageal bulb round 
to pyriform. With the genera: Eiibosfii- 
chus Greeff, 1869 (syn. Catancma Cobb, 
1920, ?Lcixm Cobb, 1893), Laxonema Cobb, 
1920, Lcptonemella Cobb, 1920, Stilboncma 
Cobb, 1920, Robbed Gerlach, 1956, and 
Sqiianenui Gerlach, 1963. 

E) Ceramonematinae Cobb, 1933': Cuti- 
cle heavily annulated, tiled or longitudinally 
broken by spined alae; head well set off 
from annulation; buccal cavity minute or 
absent, unarmed; amphids obscurely spiral 
to shepherd's crook; esophagus. With the 
genera: Cemmonema Cobb, 1920, Xenclla 
Cobb, 1920, DasynemeUa Cobb, 1933 (syn. 
Dasiincma Cobb, 1920), Pristioncmo Cobb. 
1933, PseJkmcma Cobb, 1933, Dosync- 
moidcs Chitwood, 1936, MetodasyncmcUa 
de Coninck, 1942, and Pteriji;oncma Ger- 
lach, 1954. 

F) Monoposthiinae Filipjev, 1934: Cuti- 
cle coarselv annulated, broken longitudi- 

^ This subfamily may not belong to the Des- 
modoridae. Gerlach (1957) considers it related 
to the Axonolaimidae and the Halaphanolaimidae. 

nally by alae; head well set off from annu- 
lation, rigid; amphids circular, surrounded 
by annulation; buccal cavity well armed; 
esophageal bulb barrel-shaped, ^^'ith the 
genera: Monoposfliia de Man, 1889, 
Niidora Cobb, 1920, Rhinema, Cobb, 1920, 
and Monoposiliioides Hopper, 1963. 

Key to Genera of Metachromadorixae 
( Based on classification of Gerlach, 1951 ) 

1. Male supplements heavih' cuticularized, 

large and tubular 2 

Male supplements indistinct or conical or 
consisting of narrow ducts, not large and 
tul)ular; only in M. vivipara and M. 
quadrihuiha are the supplements heavily 
cuticularized but not tubular 4 

2. Supplements strongly S-shaped, heavily 

cuticularized _ 3 

Supplements faintly S-shaped, cuticulariza- 
tion light ..— Onyx Cobb, 1891 

3. Cephalic and subcephalic setae present 

Sigmophora Cobb, 1933 

Cephalic setae only present 

Polysigrna Cobb, 1920 

4. Teeth absent or minute 

..„ Spirinia Gerlach, 1963 

Teeth well developed 5 

5. Esophageal bulb weakly developed, round 

to pyriform; cuticular lining faint 

CJiromaspirina Filipjev, 1918 

Esophageal bulb well de\eloped, usually 
elongate, sometimes "barrel-shaped," rarely 
cla\ate and indistinctb' set off from esoph- 
agus; cuticular lining usually distinct 

Metachromadora Filipjev, 1918 

Fseudomeiachioinadora Timm, 1952, is a genus 
of doubtful position. The esophagus is barely 
enlarged posteriorly, the cuticular striation is all 
but absent, the amphids are situated near the lips 
and the buccal cavity is cylindrical and strongly 

SP/R/N/A Gerlach, 1963 
Type species.— Sp/ra parasitifera Bastion, 
1865: 159, 160, pi. 13, figs. 201-203. 

Spira Bastian, 1865, nee Brown, 1844, and Sp/c/no 
Filipjev, 1918, nee Kayser, 1889. 

Our material contained two species, one 
of which undoubtedly is S. parasitifera 
( Bastian, 1865 ) . A redescription of this 
cosmopolitan species with discussion of its 
synonymy was recently given by Gerlach 
( 1963b ) . Our second species is closely re- 


Bulletin Mu.scuin of Conijxirative Zoology, Vol. 135, No. 5 

lated to S. .striaiicaudata (Timm, 1962) 
from which it can be distinguished b\' the 
hook-shaped proximal end of the spicula 
and the presence of small but distinct teeth 
in the conical buccal cavity. These two 
species are separated from all other mem- 
bers of the genus by the flagellate tail 
which in both sexes is much more distinctly 
and coarseK' striated than the rest of the 
body. This is such a conspicuous and char- 
acteristic feature that we feel justified in 
establishing a new subgenus on it. 

Spirinia (S.) parasitifero (Gerlach, 1963) 
Plate XVII, fig. 35, a-e 

S))irinia (S.) paiasififcia ( Bastian, 1865) Cer- 
lath, 1963b: 67. 

Spira pcirasififcm Bastian, 1865: 159-160, pi. 13, 
figs. 201-203; Spirilla parasitifcra auct.; Spi- 
lophora oxyccphala ButsL-lili, 1874: Spirina 
nidro.siensis Allgen, 1933; S. zosterae Filipjev, 
1918; and S. rotivillei Schnnrmans-Stekho\en, 

L = 1.57 mm; w = 55 /x; esophagus = 140 
/x; nerve ring 77 fi from anterior end; tail = 
140-155 /x. Head diameter 22 /j,; cephalic 
setae 5 /x, on level of amphids. Cervical 
setae beginning at short distance behind 
amphids, rather scattered. Amphids 6 /x 
wide. Buccal cavity small, with 3 minute 
teeth. Esophageal bulb 40 X 36 /x. Spicula 
50 IX, knobbed proximally, with velum, tail 
conical, a.b.d. 24-28 /x. 

Rcprcseniaiion in samples .studied. — M- 
4, Rickenbacker Causeway, V, Vero Beach. 

Geographicol distribution. — Baltic, North 
Sea, North Atlantic, Black Sea, Mediterra- 
nean, Barents Sea, Indian Ocean (Mal- 
dives ) . 

Spirinia iPerspiria) new subgenus 
Type species.— Sp/r/n/a iPerspiria) hamafa 
new species. 

Diffcrcnlifd diaiino.sis. — Perspiria n. subg. 
is distinguished from S})irinia sensu siricto 
by the more prominently striated and 
flagellate tail. In Spirinia sensu stricto the 
tail is conoid and the striations are fine — no 
coarser than those on the remainder of the 

In addition to the new species described 
below, Spirinia striaticaudata (Timm, 1962) 
(syn. Spirina striaticaudata) is also in- 
cluded in the new subgenus. 

Key to Species of SpiRhyiA ( Perspiria ) 

1. Stoma without teeth; proximal end of spicula 

knobbed, rounded 

S. (P.) striaticaudata (Timm, 1962) 

Stoma with minute teeth; proximal end of 

spicula hook-shaped 

____ S. (P.) hamata new species 

Spirinia [Perspiria) hamata new species 
Plate XVIII, fig. 36, o-c 

L = 2.04 mm; esophagus = 160 /x. Head 
diameter 16-21 /i. Cephalic setae 4-5 jx. 
Very few short, scattered cervical setae. 
Amphids 5-6 p.. Buccal cavity small, with 
three minute teeth. Esophageal bulb 45 X 
36 fx. Excretory pore between bulb and 
nerve ring. Cuticular striation distinct but 
weak. Spicula 37-43 /x, gubemaculum 16 
fx. Tail 230-340 /x, from beginning of sec- 
ond fifth on coarsely striated, tip unstriated. 
Anal body diameter 32-43 /x. 

Holotypc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4068, Type slide No. 78. 
Type locality, M-3, Key Biscayne. 

Representation in samples studied. — M- 
3, Key Biscayne. 

CHROMASPIRINA Filipjev, 1918 

Type species.— C/iromosp/r/na pontica Filip- 
jev, 1918: 229, 230-234, pi. 7, fig. 
45, a-c. 

Mesodoni.s Cobb, 1920: 325. 

As previously suggested by Gerlach 
( 1963b ) , this genus probably links the 
Metachromadorinae with the Desmodori- 
nae. We do not agree, however, with Ger- 
lach that the intermediate position of this 
genus is sufficient justification for merging 
the two subfamilies together. The position 
of Chromaspirina becomes less problemati- 
cal if some of Gerlach's new combinations 
are returned to thcnr former status. Thus 

Florida Marine Nematodes • Wieser and Hopper 273 

we cannot accept the transfer of Dcsmo- 
dora inflexa Wieser, 1954, and D. dimorpha 
Hopper, 1961, to Chromaspirina. Both spe- 
cies are characterized by heavy anniilation 
and a rigid head with the amphids situated 
outside the annulation (although in the 
latter species, admittedly, the position of 
the amphids with respect to the cuticular 
annulation is a bit doubtful ) and thus most 
likely represent true members of Desmo- 
dorci. The same would seem to apply to 
D. rahosa Gerlach, 1956, which was also 
tranferred to Chromaspirina by Gerlach. 
However, this species is known only from 
one female and thus remains doubtful. 

Other doubtful species are C. paiicispira 
Schuurmans-Stekhoven, 1950 ( $ only) and 
C. robusta Wieser, 1954 (juv. only). Two 
more species are known only as females, 
i.e., C. crinita Gerlach, 1952, and C. pellita 
Gerlach, 1954, but in their cases the pilosity 
of either head (crinita) or body (pellita) 
is probably sufficient to recognize them as 
good species. The following are regarded 
as good species of CJiromaspirina: C. cy- 
lindricollis (Cobb, 1920) (syn. Mesodorus 
ci/lindricoUis), C. indica Gerlach, 1963, C. 
madagascaricnsis Gerlach, 1953, C. para- 
pontica Luc and De Goninck, 1959, C. 
pontica Filipjev, 1918, and C. thicnji De 
Goninck, 1943. 

The conspecificity of C. pontica Filipjev 
sensu Gerlach, 1951, with Filipjev's species 
is questionable, as, in Gerlach's specimens, 
the cephalic setae are 9-11 ^u, long (as 
against 5 //), the amphids are relatively 
larger, and the gubernaculum is shorter 
and of a somewhat different shape. 

From the above mentioned species, C. 
inaiirita n. sp., is separated by the indis- 
tinct cephalic papillae, the large and oval 
amphids which are about as long as the 
head is wide, the shape of the gubernacu- 
lum, and the presence of ventral papillae 
on the tail. 

Chromaspirina inaurifa new species 
Plate XVIII, fig. 36, a-c 

L = 1.18-1.35 mm; w = 29-35 /x; esopha- 

gus = 105-115 p.; tail: c^ , 90-95 /x, 9 , 90 
ix; Vu = 49%. Eggs 30-33 x 60-70 /x. Head 
diameter 14 /x. Six minute cephalic papil- 
lae. Four cephalic setae, 5 /j. long. In $ 
two ( dorsal and ventral ) subcephalic setae, 
in both sexes scattered short cervical setae. 
Amphids in 6 , 12-14 + 10-12 ,i, loop- 
shaped, inner circle more heavily cuticular- 
ized, 6 X 5.5 //, more narrowly coiled. Gu- 
ticular annulation distinct. Buccal cavity 
with three small teeth (these teeth are 
smaller than in most other species of 
Chromaspirina and stress the proximity of 
the genus to Spirinia). Esophageal bulb 
pyriform, 25 X 20 /x. Spicula 27-28 /x, with 
faint velum, hooked proximal end. Guber- 
naculum sickle-shaped, 15 fx. The preanal 
ventral cuticle is crenate over a long dis- 
tance. Immediatelv in front of the anus 
there is a stout spine. On the tail there are 
four ventral supplements each accompanied 
by a pair of setae. Subventrally and sub- 
dorsallv more setae can be seen. A.b.d. 

Holotijpe specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4068, Type slide No. 79. 
Type locality, M-3, Key Biscayne. 

Representation in samples studied. — M- 
3, Key Biscayne. 

METACHROMADORA Filipjev, 1918 

Type species.— Mefoc/iromoc/oro macroufera 

Filipjev, 1918: 218, 219-225, pi. 6, 

fig. 42a; pi. 7, fig. 42, b-h. 

Key to Species of Metachromadora 

1. Male supplements conical, each consisting 

of three ( 1 central, 2 lateral ) cuticular- 

ized pieces subgenus Chroma- 

doropsis Filipjev, 1918 2 

Male supplements indistinct or narrow 
ducts the openings of which are differ- 
entiated into button-shaped or conical 
bodies 3 

2. Esophageal bulb, 2-sectioned 

M. vivii)cira (de Man, 1907) 

(syn. Chrornadora civipaia de Man, 1907) 

Esophageal bulb, 4-sectioned 

M. quadrihidha Gerlach, 1956 

3. Somatic setae arranged in 10 dense longi- 

tudinal rows subgenus Metomjx 

274 Bulletin Miiscmii of Coin])(initwe Zoology. Vol. 135, No. 5 

Chitwood, 1936 

M. horrida Chitwood, 1936 

Somatic setae not arranged in 10 dense 
longitudinal rows 4 

4. Head with pronounced longitudinal stria- 

tion subgenus Mctachrumadora 

Filipjev, 1918' ...- 5 

Head without pronounced longitudinal 
striation 6 

5. Length 2.4-2.6 mm; male with 26-48 pre- 

anal supplements 

M. luacrotitem Filipjev, 1918 

Length 1.0-1.4 mm; male with 12-14 pre- 

anal supplement 

M. chandkri (Chitwood, 1951) 

(syn. Ichthiiodesmodora chandleri 

Chitwood, 1951. 
and MetacJiroiuadora 
parasitifera Timm, 1952 ) 

6. Posterior portion of esophagus clavate; 

bulb not well set off, partitions absent 
or indistinct; cuticle smooth to indis- 
tinctly striated. (Subgenus doubtful, 
perhaps a new one to be established ) 7 

Posterior portion of esophagus bulbular; 
bulb well set off, partitions distinct: 
striations of cuticle distinct 9 

7. Male without preanal supplements 

M. .s/)//y;/(.v Ccrlach, 1955 

Male with distinct preanal supplements _ 8 

8. Male with 3 knob-like preanal supple- 

ments M. chivata C.erlach, 1957 

Male with 19-21 conoid preanal supple- 
ments M. serrata Gerlach, 1963 

9. Posterior cephalic setae absent or stout 

and short; amphids (at least in male) 
on thick cuticularized plates; cuticle 

with lateral wings subgenus Mcta- 

chromadoroklcs Timm, 1961 10 

Posterior cephalic setae slender; amphids 
not on thick cuticularized plates — . 13 

10. I'^sophageal bullj, 2-sectioned _.-. 

M. remanei Gerlach, 1951 

Esophageal bulb, 3-sectioned 11 

11. Male without preanal supplements and 

without ventral caudal "bumps" or 

"warts" M. vuh^arls Tinnn, 1961 

Male with 17-23 preanal supplements and 
with 2 ventral caudal "bumps" or 
"warts" 12 

12. Cephalic .setae present 

M. pidvinata new species 

Cephalic setae absent 

., M. complexa Timm, 1961 

13. Lateral wings present subgenus 

' M. ctjfito.'ieirae Filipjev, 1918, also belongs to 
this subgenus. However, it appears to have Iieen 
described on the basis of a single female, and, as 
such, its systematic position is doulitful. 

Neonyx Cobb, 1933 14 

Lateral wings absent _ _ _ subgenus Bradij- 
laimus Schuurmans-Stekhoven, 1931 ___ 18 

14. Circles of cephalic setae in typical ar- 

rangement (6 + 6-f 4) 15 

Circles of cephalic setae in atypical ar- 
rangement, 6 + 6 -(- 8, through fusion 
with subcephalic setae ( Cobb mentions 
this condition in rather vague fashion 
for M. cancellata) 17 

15. Esophageal bulb, barrel-shaped; preanal 

supplements, 8; body shape, obese, de 
Man a value := i, 16-24, 9, 9.5-11.5 

M. ohcsa Chitwood, 1936 

Esophageal bulb, elongate; preanal sup- 
plements, 12; body shape more slender, 
de Man a value = <^ , 33-44; 9, 34 ____ 16 

16. Subcephalic and cervical setae shorter 

than the longest cephalic setae 

M. pscudocampijcoma Hopper, 1961 

Sulxephalic and cervical setae longer than 

the longest cephalic setae 

M. canipijcoma (Cobb, 1933) 

( doubtful species ) 
( syn. Neonijx campijcoma Cobb, 1933 ) 

17. Buccal ca\ itv with denticles 

'.._. M. cancdlaia (Cobb, 1933) 

(syn. Neomjx cancellata Cobb, 1933) 

Buccal caxity without denticles 

M. tncridiana new species 

18. Esophageal bulb, 3-sectioned 19 

Esophageal bulb, 2-sectioned 21 

19. Buccal cavity with denticles; head with 

several circles of long subcephalic and 
cervical setae posterior to cephalic setae 

M. setosa Hopper, 1961 

Buccal cax'ity without denticles; head with 
only a single circle of long subcephalic 
setae posterior to cephalic setae 20 

20. Male with 9-10 preanal supplements 

M. onyxoidcs Chitwood, 1936 

Male without supplements 

M. asiipplementa (Crites, 1961)" 

(syn. Neotiyx a.sti))j)lcmi'nta Crites, 1961) 

21. Esophageal bulb elongate, length more 

than twice width — 22 

Esophageal bull) ovate, length less than 
twice width 23 

22. Amphid 7.5 /u wide, less than one-third 

c.b.d. M. ^crlachi new name 

( syn. M. onyxoide.s .scu.sii Gerlach, 1955, 
nee Chitwood, 1936) 
Amphid 15 ^j, wide, more than two- 
thirds c.b.d. __ M. spectaii.s Gerlach, 1957 

23. Amphid 12 fx wide, more than one-half 

c.b.d. M. pncuniaticd Gerlach, 1954 

Amphid 7-8 /j. wide, about one-fourth 

c.b.d. M. .^ticcica (Allgen, 1929) 

(syn. Oi.'itolaimti.s .succiciis Allgen, 1929) 

"Possiblv a svnon\ni of M . oiujxoidcs. 

Florida Marine Nematodes • Wieser and Hopper 275 

Metachromadora {Mefachromadoroides) 
pulv'mafa new species 

Plate XIX, fig. 38, a-c 

L = 1.72 (juv. 9 =1.11) mm; w = 95 
(juw 9 =65) /jl; esophagus = 315 (juv. 
9 =220) /x; tail: juv. 9 = 80 /x; Vu 647^. 
Head diameter on level of amphids 40 fi. 
Lips cushion-like, demarcated from head 
by distinct groove. Labial papillae conical. 
Cephalic setae stout, 8 /x long in male. 
Short cervical setae. Somatic setae up to 
18 jji in posterior part of body. Amphids 
in 6 , 22 X 18 /x. on cuticularized plates, 
ring-shaped, in 9,8x6 /j,, loop-shaped. 
Cuticular striation distinct, reaching to 
base of lips. Lateral differentiation not 
very pronounced, beginning around end of 
esophagus, ending at some distance in front 
of anus. Buccal cavity strongly cuticular- 
ized, in 6 , 50 /t long, with large dorsal 
tooth, 2 small subxentral teeth and an ad- 
ditional tooth at the bottom of the buccal 
cavity; vestibulum with ribs. Esophageal 
bulb tripartite, \\ ith heavy cuticular lining, 
about 105 X 62 ^. Spicula 55 /x long, 15 /x 
broad. Gubemaculum 27 fj.. Preanally the 
\entral cuticle is extended and forms a 
striated membrane which is traversed by 
about 23 narrow ducts, each duct ending 
in a button-shaped body. Extended cuticle 
reaching 540 fx preanal. Immediately in 
front of anus one strong spine. Tail 110 
/x = 2 a.b.d. long, in 6 with two ventral, 
conical, cuticularized warts. Many long 
spines in longitudinal rows. 

Ilolofype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4075, Type slide No. 80. 
Type locality, V, Vero Beach. 

Representation in sampJes studied. — V, 
Vero Beach. 

Remarks. — The only other species in this 
subgenus with postanal ventral warts is M. 
complexa Timm, 1961, which, however, is 
devoid of cephalic setae. 

Metachromadora {Bradylaimus) onyxoides 
Chitwood, 1936 

Metachromadora (Bradylaimus) onyxoides Chit- 
wood, 1936: 5, Hr. 1, v-x. 

Our specimens appear to be typical rep- 
resentatives of M. onyxoides as described 
by Chitwood (1936) and Hopper (1961a). 
The amphids in our male measured 7-8 /x. 

Representation in samples studied. — M- 
5, Virginia Key. 

Geogra ph ica 1 distrib ution . — B eauf ort , 
North Carolina (Chitwood, 1936), Gulf 
Shores, Alabama (Hopper, 1961a) and ? 
Pemambuco, Brazil ( Gerlach, 1956 ) . 

Remarks. — Gerlach's (1955) identification 
of specimens from San Salvador as M. 
onyxoides is doubtful. In the table below 
some dimensions of our own specimens are 
compared \\'ith those of the animals de- 
scribed by Hopper ( 1961a ) and by Gerlach 
( 1955 ) . It follows that Gerlach's speci- 
mens belong to a different species for 
which the name M. fs^erlachi new name is 









Length of 







Length of 







Length of 



60 m 


Length of 





40 m 





of par- 



20+25+25 M 


Metachromadora (Bradylaimus) gerlachi 

new name 

Metachromadora onyxoides Chitwood Ger- 
lach, 1955. 

276 BiiUctiii Mtisctint of Couipaidlivc Zoology, Vol. 135, No. 5 

Mefacliromadora ( BradyJciimus) '^crhchi 
new name is related to M. omjxoidcs Chit- 
wood, 1936, from which it is distinguished 
by ha\ini:; a two-sectioned esophageal bulb, 
the bulb in M. omjxoidcs being three-sec- 

hAeiochromadora {Neonyx) meridiana new 
Plate XIX, fig. 39, a-d 

L = 0.95-1.05 mm; w = ^ , 47, 9 , 55-62 
fj.; esophagus 175-185 //. Lips prominent. 
There are three distinct circles of cephalic 
sense organs: in front two circles, each 
composed of 6 conical, setose papillae, 
followed by one circle of 8 slender setae, 
each measuring 6-7 /x. It is assumed that 
this circle consists of the t\'pical 4 cephalic 
plus 4 subcephalic setae. Close to the am- 
phids there are two more subcephalic setae 
on each side of the body, measuring 8 /x. 
Cervical setae in the anterior region up to 
10 II, in posterior region up to 15 /<. long. 
Amphids spiral, 7 /x = 30% of c.b.d. wide. 
Cuticular striation reaching to anterior end 
of amphids; lateral alae starting around 
middle of cervical region. Buccal cavity 
with rather thin \\'alls, very strong dorsal 
tooth and small subventral projections. 
Esophageal bulb tripartite, with heavy cu- 
ticular lining, 60-70 X 28-32 /x in ^ , 80 X 
35 /x in 9 . Eggs 70-75 X 40-50 /x. Spicules 
arcuate, 58 /x long, gubernaculum 29 /x long. 
Male with 9-10 thin preanal supplements, 
traversing the raised ventral cuticle. Tail 
80-95 /x long. Female a.b.d. = 28 /x, male = 
35 /I. Caudal setae on male tail arranged 
as illustrated in Plate XIX, figure 39c. 

Il()l()ti/))c specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4075, Type slide No. 81. 
T\\)v locality, V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 

Remarks. — At first we thought we had 
found M. ohesa. However, Chitwood's 
original description is rather poor and all 
other authors who subs(Yjuently described 
this species (Timm, 1952; Hopper, 1961b; 

Crites, 1961) mention the typical arrange- 
ment of 6 short and 4 long cephalic setae, 
whereas in our specimens there is definitely 
a circle of S long cephalic setae. 


Key to Genera of Desmodorinae 

1. Esophageal bulb elongate, tripartite 2 

Esophageal bulb round to pyrifonn ..._ 3 

2. Head with large plates in posterior portion 

(head "jointed" or tiled) 

Acanthophoryngoides Chitwood, 1936^ 

Head simple, without plates 

AcantJiopIwrynx Marion, 1870 

syn. Xanthudora Cobb, 1920 

3. Amphids half or completely surrounded by 

annulation 4 

Amphids not surrounded by annulation 

Desmodora de Man, 1889 

(see discussion by Gerlach, 1963b) 

4. Amphids half surrounded by annulation, not 

situated on cuticularized plates 

Paradcsmodora Schuurmans-Stekhoven, 1950 
Amphids completely surrounded by annula- 
tion, situated on cuticularized plates 

Metadesmodora Schuurmans-Stekhoven, 1942 

PARADESMODORA Schuurmans-Stekhoven, 

Type species.— Poroc/esmoc/ora cephalafa 

Schuurmans-Stekhoven, 1950: 117, fig. 

67, a-e. 

This genus contains the following spe- 
cies: P. campheUi (Allgen, 1932) Gerlach, 
1963 (syn. Spirina campheUi), P. immersa 
Wieser, 1954, P. punctata Gerlach, 1963, 
and P. toretites n. sp. The type species, P. 
ccpJwlata Schuin-mans-Stekhoven, 1950, is 
known from a juvenile female only and is 
considered a species inquirenda. 

Our new species can be distinguished 
from all other species of the genus by the 
shape and arrangement of male supple- 
ments, the hook-shaped spicula and the 
cuticular differentiations of the head. 

Paradesmodora toreutes new species 
Plate XX, fig. 40, a-f 

L = 1.62 mm; w = 21 /x; esophagus = 114 

^ In this genus the cuticular annulation is not 
quite as pronounced as in other genera of this 

Florida Marine Nematodes • Wiescr and Hopper 277 

fx; head diameter 14 /x. Lips distinct, papil- 
lae minute. Cephalic setae 6 /x. Cuticle in 
posterior portion of (adult) head very 
much enlarged and forming plates which 
surround the anterior portions of the am- 
phids. Amphids 6 /x == 40% of c.b.d. wide, 
one circular loop. There are a few short 
cervical setae. Cuticular annulation coarse. 
Esophageal bulb pyriform, 22 X 15 fx. Spic- 
ula semicircular, 20 /x, proximal end hooked. 
There are 10 preanal supplements each 
consisting of a ventral bump and a thicken- 
ing of the cuticle. The row of supplements 
extends 220 fx preanally. Bet\veen two sup- 
plements there are 13-18 cuticular annules. 
Tail 95 /J. long, a.b.d. 20 /j.. Juvenile tail 7 
a.b.d. long. 

Holofype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4068, Type slide No. 82. Type locality, 
M-3, Key Biscayne. 

Representation in samples studied. — M-3, 
Key Biscayne. 

DEShAODORA de Man, 1889 

Type species. — Sp//opfiora comments 

Butschli, 1874: 282, 283, pi. 5, fig. 27, 

a, b; pi. 7, fig. 27, c, d. 

Gerlach (1963) has recently reviewed this 
genus, relegating several kno\\'n genera to 
subgeneric rank (i.e., Pseudochromadora, 
Xenodesmodoia, Croconema, Bolhonema, 
Desmodorella and Zalonema). Excellent 
discussions, with keys, for both the sub- 
genera and the species within each sug- 
genus are presented in Gerlach's paper. 

Gerlach's (1963b, p. 84) discussion of 
Xenodesmodora makes synonymization of 
BJa Inglis, 1963, with the former subgenus 
inevitable. The type species, Bla nini Inglis, 
1963, therefore, is transfen^ed to the 
genus Desmodora and becomes Desmodora 
(Xenodesmodora) nini (Inglis, 1963) new 

The genus Desmodora was represented 
in our material, in samples M-2 and M-3 
from Key Biscayne, by a single, well-known 
species, Desmodora {Vseudochromadora) 

quadripapillata (Daday, 1899) Gerlach, 1963 
(synonyms Fseudocliromadora quadripapil- 
lata Daday, 1899, Micromicron cephalata 
Cobb, 1920, and M. luticola Timm, 1952). 

Pertinent measurements from a male 
specimen are as follows: L = 0.74 mm. 
Head diameter 14 /x. Amphids 6 /x. Spicula 
28 /x. 

Representation in samples studied. — M-2, 
Key Biscayne, M-3, Key Biscayne. 

Geographical distribution. — New Guinea 
(Daday, 1899), Costa Rica (Cobb, 1920), 
Chesapeake Bay, Maryland (Timm, 1952), 
and Cananeia, Brazil ( Gerlach, 1957 ) . 

MONOPOSTHIA de Man, 1889 
Type species.— Sp///p/iora cosfofo Bastion, 
1865: 166, 167, pi. 13, figs. 228, 229. 

We prefer not to follow Gerlach ( 1963 ) 
in his synonymization of Monoposthia and 
Nudora. Thus in Monoposthia only species 
are retained in which the spicula are absent 
and the single gubernaculum is not con- 
spicuously enlarged proximally. \\'ithin the 
genus there is a group of species character- 
ized by the enlarged second annule. A key 
to this group reads as follows: 

Key to Species of Monoposthia 

1. Botli first and second annule enlarged; am- 

pliids between the two annules 

M. thorakista Schulz, 1935 

Only second annule enlarged; amphids on 
this annule 2 

2. Cuticle with 12 longitudinal rows of V- 

shaped markings ____ 

M. dtiodecimalata Chitwood, 1936 

Cuticle with 6 longitudinal rows of V-shaped 

markings 3 

3. Cephalic setae measuring less than % of 

head diameter; amphids % of c.b.d. wide _ 

M. mielcki Steiner, 1916 

Cephalic setae approximately one head diam- 
eter long; amphids about ^f, of c.b.d. wide . 

M. mirabilis Schulz, 1932 

syn. M. longiseta Allgen, 1935 

Our material is representative of M. 
mirabilis Schulz, 1932, agreeing in all es- 
sential points with the excellent redescrip- 
tion given of this species by Luc and De 

278 Bulletin Museum of Coiiipdidfive Zoology, Vol. 135, No. 5 

Coninck (1959). M. ovnata Tinini, 1952, 
described on [he basis of one jiuenile, is 
either a synonym of M . iniidhilis or else a 
■sjiccics i)Hjiiiicnchi. 

Monoposfhia mirabilis Schuiz, 1932 
PlateXX, fig. 41,a, b 

Moiii>i)().sflii(i iiiirahilis Sclmlz, 1932: 3S0-382, fiji. 
2fi, a-g 

Male. — L = 1.6 mm; w = 57 /x. Diameter 
at base of esophagns, 54 ^. Esophagns 210 
/L long. Head diameter 19 /x, bearing six 
labial papillae and four, 19 /x long, cephalic 
setae. Cuticle with 6 longitudinal rows of 
\'-like markings. Reversal of V's at mid- 
bod). Bod\- bearing 4 rows of somatic 
setae which carry on to the tail. Buccal 
cavity armed with dorsal tooth and opposed 
by a number of denticles. Gonad single, 
outstretched. Gubernaculum 38 /x long. Tail 
110 /x long, a.b.d. 39 /x. A prominent double 
pair of fleshy papillae are positioned pre- 
anally, surrounded by a cuticularized ridge. 

Representation in samples studied. — M-2, 
Key Biscay ne. 

Gen<iraphical distiihtition. — European At- 
lantic coasts, Mediterranean Sea (Gerlach, 


Type species.— A^onoposf/i;o/c/es anonopos- 

ihia Hopper, 1963: 850-852, figs. 


This genus is differentiated from Mo)U)- 
posthia by the shape of the gubernaculum 
("spiculum" in the interpretation of other 
authors), the long spine attached to the 
latter, the presence of two testes and the 
absence of V4ike markings in the anterior 
portion of the body. 

Our specimens agree with this diagnosis 
except for the absence of the gubernacular 
spine. No trace of it could be detected in 
the four males examined. Further dif- 
ferences from the type, M. anonoposthia, 
are: the cephalic setae measure only about 

one head diameter as against 1.5, and in 
the gubernaculum it is the dorsal rather 
than the ventral arm of the proximal exten- 
sion which is the longer, ^^'e thus consider 
our specimens to represent a new species 
which we call M. mayri n. sp. in honor of 
Dr. Ernst Mayr of the Museum of Com- 
parative Zoology, Harvard University. 

Monoposthioides mayri new species 
Plate XX, fig. 42, a, b 

L = 6 , 1.72, 9 , 1.4-1.5 mm; w = 40 /x; 
esophagus = 1S5 /x; Vu = 90-92%. Head 
diameter 16 /x. Lips distinct, with 6 setose 
papillae. Cephalic setae 17 jx. Cuticle with 
broad rings in the anterior cervical region 
whence they gradually become narrower. 
The second annule is even more enlarged 
than the adjacent ones and measures 7 /x 
in width. There are at least 12, probably 
14, longitudinal rows of V-like markings 
which start about 50 ^u from the anterior 
end. The reversal of the V's takes place 
in the male at 140 /x behind the posterior 
end of the esophageal bulb, in the female at 
240-250 1^1. (in M. anonoposthia the V-shaped 
markings are reversed in the male at a point 
opposite the anterior margin of the bulb 
and in the female at a point approximately 
one and one-half tail lengths anterior to 
the anus ) . Buccal cavity cylindrical, mea- 
suring 22 /x from tip of tooth to base, with 
one large dorsal tooth and small subventral 
projections. Amphids more or less pocket- 
shaped and sunk into the cuticle, somewhat 
irregular in outline. Esophageal bulb 47 X 25 
/x. Single gubernaculum 42 /x long, axe-like, 
proximal end 23 ^ wide, gliding in an anal 
sheath as reported for M. anonoposthia. 
Tail: i , 140 /x, 9 , 100-105 /x long; a.b.d. = 
30 /x. 

Jlolotijpe specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4067, Type slide No. 83. Type locality, 
M-3, Key Biscay ne. 

Representation in samples studied. — M-3, 
Kev Biscavne. 

Florida Marine Nematodes • Wiescr and Hopper 279 

Type species.— Param/cro/a/mus primus 
Wieser, 1954: 64, fig. 135, a-c. 

In this genus the first circle of (6) 
cephahc sense organs is not papilloid but 
consists of slender setae which, in the three 
species so far described, are more than half 
as long as the (4) cephahc setae of the 
second circle. In our new species, the setae 
of the first circle are considerably shorter 
than those of the second. Thus the arrange- 
ment of cephalic organs is very much like 
that of several species of Microlaimiis. How- 
ever, further distinguishing features of 
Poromicroloimus are the transversely oval 
amphids and the distinct preanal papillae in 
the male. The following species are known: 
P. primus \\'ieser, 1954 (9 only!), P. 
papiUatus (Gerlach, 1954) (syn. Micro- 
laimiis papilhtus), P. spindifer Wieser, 
1959, and P. htnatus n. sp. In addition to 
the shorter cephalic setae, P. lunatiis is 
characterized by the semicircular spicula. 

Paramicroloimus lunafus new species 
Plate XXI, fig. 43, a-c 

L = 1.2-1.3 mm; w ^ 29-33 ,x. Head 
diameter 11-13 ^. Labial papillae distinct. 
First circle of cephalic setae short, at best 
3 fx long. Second circle of cephalic setae, 
7 IX long in (^ , 11 /x long in 9  Amphids 
transversely oval, 10 fx wide (% c.b.d.) in 
S ,8 IX wide ( -.^ c.b.d. ) in 9 ; located 10-13 
IX behind anterior end. Cuticular striation 
distinct, extending anteriorly only to base 
of cephalic setae. Buccal cavity wide, coni- 
cal, \\'ith one large dorsal tooth, 2 sub ventral 
teeth and projections at the base. Esopha- 
gus 145-155 IX long, temiinated by esopha- 
geal bulb 28 X 22 ix. Tlie musculature of 
the esophageal bulb is indistinctU- divided 
into two parts by a weak transverse break. 
Spicula 45 ix, regularly bent. Gubernaculum 
simple, 19 /x long. The male bears 5 prom- 
inent preanal supplements which appear to 
be tubular in structure. The cuticle in the 
area of each supplement is somewhat swol- 

len. The 3rd and 4th preanal supplements 
are always closer together than are any of 
the remaining ones. Tail conical, 85-90 jx 
long, bearing 6 pairs of subventral setae; 
a.b.d. = 25-27 ,x. 

Holotype specimen. — Male; Canadian Na- 
tional Collection of Xematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4066, Type slide No. 84. Type locality, 
M-2, Key Biscayne. 

Representation in samples studied. — M-2, 
Key Biscayne. 


The genera in this subfamily are arranged 
as in ^Vieser ( 1954 ) ; that is, the primary 
division is into genera with hollow or with 
solid teeth. \\''ithin each of these two 
groups further separation is based on the 
structure of the cuticle (homogeneous or 
heterogeneous and with or without lateral 
differentiation ) . 

Type species.— Sp///pfiera inaequalis Bas- 
tian, 1865: 166, pi. 13, figs. 223-225. 

Group A in W'ieser's key (1954) contains 
species with very long somatic setae and 
up to now is represented by six species. 
Since classification of these species leans 
heavily on features of the spicular ap- 
paratus, //. lieymonsi (Steiner, 1922), known 
from 1 9 only, is considered a species 
inquirenda. A key to the species reads as 
follows : 

Key to Species of HypoDO-\TOLAiMUS 
(Group A of \\'ieser, 1954) 

1. Cephalic setae twice the head diameter. 

Pharyngeal bull") not \'ery powerful, dorsal 
tooth weakh' S-sl:aped, not pushed into 
\entral Isuccal wall. Indistinct preanal 
papillae present . H. (Biitschli, 1874) 
Cephalic setae measurintj; about 1 head diam- 
eter. Phar>ngeal bulb powerful, dorsal 
tooth strongly S-shaped, pushed into ven- 
tral buccal wall. \o preanal papillae 2 

2. Distal portion of tail with ventral cur\ature. 

First circle of ( 6 ) cephalic sense organs 


Bulletin Miisciiiii of Coiiipdidlivc Zoology, Vol. 135, No. 5 

papilloid. Esophageal bulb iiiclistinctb' 

set off from esophagus — 

H. colesi Inglis, 1962 

Distal portion of tail with dorsal curvature. 
First circle of (6) cephalic sense organs 
setose. Esophageal bulb well set off 3 

3. Gubernaculum with hook-shaped apophysis 

H. steineri Wieser, 1954 

Cubemaculum without apophysis 4 

4. Gubernaculum well de\elopcd, spoon-shaped, 

half the length of the spicula — - 

.... H. scJtuunnansstc'khovcni Cerlach, 1951 
Gubernaculum reduced to a short plate with 
thin lamella between the 2 spicula. Free 
portion of gubernaculum much shorter than 
half the length of the spicula 5 

5. Ciubernacular lamella with dorsal extension. 

Spicula evenly cur\ed, of nearly equal 
width throughout. Tail without break .... 

H. soliv(ip.u.s Hopper, 1963 

Gubernacular lamella minute, without ex- 
tension. Spicula tapering unevenly and of 
characteristic shape (see Fig. 44, d). 

Tail with distinct break in cuticle _._ 

H. iiilcrnipttis n. sp. 

Group H of Wieser's key, comprising 
species without elongated somatic setae, 
includes a few species characterized by the 
possession of a double bull). This group, 
which has been referred to as a distinct 
subgenus, PtijcholaimelJus Cobb, 1920, by 
Gerlach ( 1955 ) , is represented by the fol- 
lowing 4 species: 

H. carinotufi (Cobb, 1920), distinguished 
1)\ long cephalic setae but imperfectly 
known. The specimens described under 
this name by Timm ( 1952 ) almost certainly 
do not belong to //. carinatiis but more 
likely to one of the following two species: 

H. ponticiis- Filipjev, 1922. (For synonyms 
and description see Gerlach, 1951a.) 

//. pandispicuhius Hopper, 1961. On the 
basis of our own material we can confirm 
Hopper's conclusion that this species is 
separated from //. ponticus by the .shape 
of spicula and gubernaculum and by the 
shorter cephalic setae. 

H. macro dent at us Timm, 1961, character- 
ized by knob-like swellings at the base of 
the dorsal tooth, and by tlie shape of the 

Hypodontolaimus (H.) interruptus new spe- 
Plate XXI, fig. 44, a-d 

L = 0.58-0.59 mm; w = 23-25 ^; esopha- 
gus = 95-100 /x. Head diameter 15 /x. Lips 
large. Cephalic setae: first circle of six = 
2.5 IX, second circle of four = 15 fx. Cervi- 
cal and somatic setae up to 35 fx long. The 
somatic setae are arranged in two sub- 
lateral rows on each side of the body. The 
same holds for the cervical setae but some 
submedian setae can also be seen. Cuticle 
annulated, with dots between the annules; 
lateral differentiation consisting of two 
longitudinal rows of larger dots and faint 
wings. Distance of longitudinal rows = 4 ^ 
in mid-body, 5 ^ in cervical region. Buccal 
cavity with powerful dorsal tooth. Pharyn- 
geal bulb well developed. Esophageal bulb 
20 X 16 IX. Amphids elliptical. Spicula 21 
/(. long, tapering unevenly, tip obliquely 
truncate. Gubernaculum reduced to a small 
plate with minute lamella between the 
spicula. Tail 85-87 jx long, with break in 
cuticle shortly before the middle; a.b.d. 22 /x. 

Holotijpe specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4070, Type slide No. 85. Type locality, 
M-5, Virginia Key. 

Representation in samples studied. — M-5, 
Virginia Key. 

Hypodontolaimus (Pfycholaimellus) pondi- 
spiculatus Hopper, 1961 
Plate XXI, fig. 45, d, e; Plate XXII, fig. 45, 
a-c, f 

H t/podontolaimus {Ptijcholaimellus) pandispictdatus 
'Hopper, 1961: 360, 361, figs. 1-4. 

L = 1.14 mm; w = 28-30 /x; esophagus = 
ISO II. Head diameter 15 ix. Cephalic setae 
(only second circle of fom- seen) 5-6 fx. 
Lips distinct. Cervical and somatic setae 
short, in two sublateral rows. Cuticular 
ornamentation typical; longitudinal rows of 
dots 5 IX apart in mid-body, 4 /x on level 
of esophageal bulb. Buccal cavity with 
medium-sized dorsal tooth and subventral 

Florida Marine Nematodes • Wiescr and Hopper 281 

pro|ections. Pharyngeal bulb not veiy large. 
Esophageal bulb 40 X 28 fi, barrel-shaped, 
double, the anterior portion smaller than 
the posterior one. Excretory pore 22 /x be- 
hind anterior end. Spicula 35 /x long, with 
xelum. Two gubernacula, 12 fj. long, proxi- 
mally expanded. Tail 110 /x long, a.b.d. 35 fi. 

Representation in samples studied. — M-3, 
Key Biscayne. 

Geop-aphical distribution. — Gulf Shores, 
Alabama ( Hopper, 1961 ) . 

Remarks. — Our specimens differ from the 
type in the more posterior position of the 
excretory pore, but there is agreement on 
all other essential points. 

RHIPS Cobb, 1920 

Type species.— Rfi/ps ornata Cobb, 1920: 
339, 340, fig. 118, a-c. 

This genus comprises two species, R. 
ornata Cobb, 1920, and R. Jon^cauda Timm, 
1961, the latter being characterized by the 
elongated tail. W^e found Cobbs species 
in our material and shall give a figure of 
the head end, the spicular apparatus being 
well described in Cobb's original publica- 

Rhips ornata Cobb, 1920 
Plate XXII, fig. 46, a, b 

Rhips ornata Cobb, 1920: 339, 340, fig. 118, a-c 

L = 1.46 mm. Head diameter 12 /x. Lips 
distinct, with 6 setose labial papillae. Ce- 
phalic setae 5 + 3 ^u,, in two circles. Amphids 
10 /x wide. Head with sLx triangular plates 
that probably serve as supports. Cuticular 
annules resolvable into basketwork-like 
structvires in the cervical region, into elong- 
ated, more or less hexagonal structures in 
remainder of body. V-shaped lateral dif- 
ferentiations in posterior portion of body. 
A few cervical setae, measuring up to 15 /x. 
Buccal cavity with large dorsal tooth and 
two small subventral teeth. Spicula 43 + 28 
/J. long. Gubemaculum (lateral pieces) 19 
/x. Tail 130 ,M long, a.b.d. 23. 

Representation in samples studied. — M-5, 
Virginia Key. 

Geographical distribution. — Ocean Beach, 

Florida (Cobb, 1920), Brazil (Gerlach, 
1957), and ? Campbell Islands (Allgen, 

CHROMADORA Bastion, 1865 

Type species.— C/iromac/ora nudicapitata 

Bastian, 1865: 168, pi. 13, figs. 230- 


This genus was discussed by Wieser 
( 1954, 1955 ) . We found what is probably 
C. mocrolaimoides Steiner, 1915, as in our 
specimens the distal end of the guber- 
naculum seems to be more strongly ex- 
panded than that figured by Steiner or 
Chitwood ( 1951 ) . However, this portion 
is weakly cuticularized and difficult to see. 
The species is characterized by the large 
esophageal bulb, the two small and rather 
faint preanal papillae, the weak cui"vature 
of the spicula and the long spinneret. 

Chromadora macrolaimoides Steiner, 1915 
Plate XXIi, fig. 47 

Chromadora macrolaimoides Steiner, 1915: 234- 
237, fiffs. 23-27; Wieser, 1955 (nee Steiner, 
1922: Allien, 1927). 

Chromadorella macrolaimoides, — Filipjev, 1918; 
Chitwood, 1951. 

Spicula 21 /x long, weakly curved, with 
velum. Gubernaculum 15 /x long, strongly 
expanded distally, edge slightly serrated. 
There are two preanal supplements, rather 
faint and of the usual cup-like shape but 
surrounded by an additional cuticular dif- 
ferentiation. The distance of the posterior 
supplement from the anus is 20 /x. Spin- 
neret 6 p. long. 

Representation in samples studied. — M-1, 
Key Biscayne, M-4, Rickenbacker Cause- 

Geographical distribution. — Sumatra 
(Steiner, 1915), Texas, Culf Coast (Chit- 
wood, 1951), and Japan (Wieser, 1955). 

TIMMIA Hopper, 1961 

Type species.— Parac/iromac/ora parva 
Timm, 1952: 24, 25, figs. 38, 39. 

Parachromadora Tinini, 1952, nee Micoletzky, 1914, 
nee Schulz, 1939. 

This genus is distinguished from Chro- 

282 Bulletin Musntin of Conipmatwc Zoology, Vol. 135, No. 5 

madorina Filipjev, 1918, solely by the oc- 
currence of a tubular supplement in addi- 
tion to the usual preanal papillae in the 
male. Our material, from \'ero Beach, con- 
tained representatives of the type species, 
Timmia parva (Timm, 1952) Hopper, 1961. 
In addition to the present locality, the 
species is known from Chesapeake Bay, 
Maryland (Timm, 1952), and Gulf Shores, 
Alabama (Hopper, 1961). 

SPILOPHORELLA Filipjev, 1918 
Type species.— Sp/7op/iora paradoxa de 
Man, 1888: 45-47, pi. 4, fig. 19. 

We seem to have typical repiesentati\'es 
of the cosmopolitan species, SpilopliorcUa 
paradoxa, in our material. 

Spilophorelia paradoxa (de Man, 1888) 
Filipjev, 1918 

Spilo])hora paradoxa de Man, 1888: 45-47, pi. 4, 

fiti. 19. 
Spilophorelia paradoxa (de Man, 1888) Filipjev, 

1918: 2,59. 

L = 0.73-0.80 mm; w = 31 /-,; esopha- 
gus = 145 /x; Vu = 447c. Head diameter 11 
/x. Cephalic setae 5 fx. Esophageal bulb 
double, typical. Spicula 36-43 ^; guber- 
naculum 30-36 ^. Tail 120-130 p. long, 
spinneret 20-22 /x. 

Representation in samples siudied. — M-2, 
Key Biscayne, M-6, Everglades National 

Geographical distribution. — Cosmo- 

PROCHROMADORELLA Micoletzky, 1924 
Type species.— Cfiromoc/ora neopolitana de 
Man, 1878: 113, 114, pi. 9, fig. 17, 
Prochromadorella mediterranea (Mico- 
letzky, 1922) 
Plate XXII, fig. 48, a-c; Plate XXIII, fig. 
48, d, e 

Chromadora mediterranea Micoletzky, 1922b; 
Chromadorella pontica Filipjev, 1922; and ? 
Hypodontolaimus arahieiis Cnlib, 1891. 

L = 0.5-0.67 mm; w = 17-21 ^a; esopha- 
gus = 100-105 /x; Vu = 46%. Head diam- 

eter 10-11 fx. Lips and labial papillae 
distinct. Cephalic setae in two circles, 6 
short ones (about 1-1.5 /x), 4 longer ones 
(5 /x). A few cei-vical setae up to 10 /x, 
amongst which one characteristic circle of 
four sublateral pairs, two on each side, at 
about 20 jj. from the anterior end. Cuticle 
typical, with at first dots, then elongated 
hexagonal bodies and rods between annules. 
Amphids faint though large, oval. Excretory 
pore on level of cephalic setae. Buccal 
cavity with three solid, subequal teeth. 
Esophagus enlarged posteriorly. Spicula 
semicircular, chord 18 /x long. Guber- 
naculum 10 /x, distally slightly expanded, 
with two or three teeth. Two faint preanal 
supplements 15 and 27 /x from anus, respec- 
tively. Tail in c^ , 75-100 /. ( = 6-6.7 a.b.d.), 
in 2, 110 /x (9 a.b.d.). 

Representation in .samples studied. — M-1, 
Key Biscayne. 

Geographical distribution. — Mediter- 
ranean Sea, Black Sea, Red Sea (Gerlach, 
1958), Bay of Bengal (Timm, 1961). 

Remarks. — Our specimens are in perfect 
agreement with the type and with material 
from the Mediterranean. 

CHROMADORELLA Filipjev, 1918 
Type species.— C/iromac/ora filiformis Bas- 
tian, 1865: 169, pi. 13, figs. 242-244. 

Since our material contained three 
species, two of which are new, we shall 
provide a new key to the genus (see also 
Wieser, 1954). 

Key to Species of Chromadorella 

1 . Cuticular ornamentation always consisting 

of dots and rod-like markings 2 

Cnticular ornamentation in anterior cervical 
region consisting of solid liands with 
crenate contour, or of fused hexagonal 
bodies 5 

2. Foiu" longitudinal rows of dots on each side 

of body C. circa tnflexa Wieser, 1954 

Two longitudinal rows of dots on each side 
of body 3 

3. Lateral differentiation beginning with cutic- 

ular annulation, there measuring % to % 
of c.b.d. 

Florida Marine Nematodes • Wieser and Hopper 283 

C. parapoecilosoma ( Micoletzky, 1922)^ 

Lateral differentiation beginning a short dis- 
tance behind cuticular annulation, there 
measuring not more than ^ik of c.b.d. ____ 4 

4. Longitudinal rows Yiq to M2 of c.b.d. apart; 

membrane present 

C memhranata Micoletzky, 1924 

Longitudinal rows i(; to Vis of apart; 

membrane absent 

C. filiformis (Bastian, 1865) 

5. Twelve preanal supplements 6 

Five to 6 preanal supplements 7 

6. Lateral differentiation irregular 

C. edmondsoni Wieser, 1959 

Lateral differentiation two longitudinal rows, 
widely spaced in anterior cer\ical region ._ 
C. galeata Wieser, 1959 

7. Esophageal bulb distinctly set off, barrel- 

shaped, short. Spicula nearly semicircularly 

curved C. parabolica Wieser, 1954 

Esophageal bulb not so well set off, more 
elongated. Spicula rectangularly bent -— 8 

8. Three longitudinal rows, in anterior cervical 

region Yn) of c.b.d. apart C. trilix n. sp. 

Two longitudinal rows, in anterior cervical re- 
gion % of c.b.d. apart C. vanmeterae n. sp. 
We consider C iinjtilicoJa Filipjev, 1918, as 
doubtful since only females are known. 

Chromadorella filiformis (Bastion, 1865) 
Plate XXIII, fig. 49, a, b; Plate XXIV, fig. 
49, c, d 

Chromadora filiformis Bastian, 1865; DicJiroinadora 
tcnuicauda Schuurmans-Stekhoven, 1950; CJiru- 
madorella filiformoides Chitwood, 1951. 

L = 0.77 mm; w = 26 /x; esophagus = 115 
fx. Head diameter 14-15 /x. Lips distinct. 
Six short cephahc setae and 4 long ones, 
measuring 10-11 p.. Ocelli and pairs of 
cervical setae 20 /x behind anterior end 
( ocelli sometimes indistinct ) . Cuticle an- 
nulated, with transverse rows of dots be- 
tween annules which become elongated in 
the posterior cen'ical region. Lateral dif- 
ferentiation through larger dots \\'hich ar- 
range themselves into distinct longitudinal 
rows at about the level of the ocelli. Be- 
tween esopliageal bulb and anus the longi- 

^ This species is not well known. There are 
doul^ts concerning the cuticular ornamentation 
since Micoletzky (1922) and Schuurmans-Stek- 
hoven (1943) mention only dots and rod-like 
markings, whereas Wieser (1951) figures solid 
liands with crenate contoiu'. 

tudinal rows are from 3.5-4.5 fx apart, that 
is, Vc, to Vk of c.b.d. Buccal cavity with three 
solid, subequal teeth. Esophageal bulb 
elongated, 30-34 X 15-16 fi. E.xcretory pore 
on level of nerve ring, 60 /x behind anterior 
end. Spicula 5 /x wide, strongly curved; 
true length = 30 fx, chord = 22 /x. Guber- 
naculum 19 /x long, consisting of a piece be- 
tween the two spicula and a caudal p)late 
which distally ends in a three-pronged pro- 
jection. There are 5 large preanal supple- 
ments. Tail 110 /x long, a.b.d. 2,5 fx. 

Representation in samples studied. — M-1, 
Key Biscayne. 

Gcoii,raphica] distribution. — Atlantic, 
Mediterranean, Black Sea, Red Sea, Suma- 
tra, Japan, Sargasso Sea, Texas. 

Remarks. — As has been noted in other 
species, the appearance of the ocelli in dif- 
ferent specimens is variable, probably due 
to the action of the preserving fluid. Con- 
sequently, Chit wood's species, C. filifor- 
moides, cannot be maintained. 

A further fact that should be mentioned 
is that in our specimens the longitudinal 
rows of dots seem to be more widely spaced 
than indicated by de Man (1890) in his 
excellent description of European repre- 
sentatives of this species. Comparative data 
on this point would be desirable. 

Chromadorella trilix new species 

Plate XXIII, fig. 50, a-c; Plate XXIV, fig. 
50, d, e 

L = 0.95 mm; w = 28-30 ix\ esophagus = 
135 /x. Head diameter 13 /x. Lips distinct. 
Labial papillae and first circle of cephalic 
setae not seen. Four cephalic setae 5-6 jx 
long. Cuticular ornamentation consisting of 
solid bands with crenate contour in anterior 
cervical region which further posterior be- 
come resolved into rod-like markings. Lat- 
eral differentiation beginning with annula- 
tion, at first consisting of round markings, 
1.5 fj. apart, then of two rows of larger dots. 
About 30 fx behind the anterior end, one of 
the two longitudinal rows of dots moves 
into the middle of the lateral fields and is 
replaced sublaterally by a new row of dots. 

284 Bulletin Museum of Comparative Zoology, Vol. 135, No. 5 

The result is a lateral differentiation con- 
sisting of three longitudinal rows which 
run to approximately the level of the anus 
where the middle row drops out. In mid- 
body the outer rows are 3.5-4 /x apart. 
Buccal cavity with three solid, subequal 
teeth. Esophagus elongated, with three 
fairh- distinct interruptions. Spicula 26 /x 
long. Gubernaculum expanded distally. 
Five large supplements, extending to 90 /x 
preanal. Tail 91 ix, a.b.d. 22 /x. 

1 1 olotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4067, Type slide No. 86. Type locality^ 
M-2, Key Biscayne. 

Representation in .samjyh's studied. — M-2, 
Key Biscayne. 

Cbromadorello vanmeterae new species 
Plate XXIII, fig. 51, a-c; Plate XXIV, fig. 
51, d, e 

L = 1.25 mm; w =^ 35 ^u; esophagus = 160 
IX. Head diameter 20 /x. Head slightly swol- 
len. Lips and labial papillae distinct. First 
circle of cephalic setae not seen, second 
circle of four setae, 5-6 /x. Cervical setae 
about 35 fx, behind anterior end, somatic 
setae along lateral fields. Cuticular orna- 
mentation consisting of solid bands with 
erenate contour in anterior cervical region, 
resolving into elongated markings further 
posterior. Lateral differentiation by larger 
dots, forming two longitudinal rows, 4-6 /x 
apart. Buccal cavity typical. Esophageal 
bulb elongated, with distinct plasmatic in- 
terruptions. Spicula 31 jx long. Guber- 
naculum simple. Six preanal supplements, 
extending to 113 /x preanal. Tail 150 /x, a.b.d. 
23 IX. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 87. 
Type locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Remarks. — This species is dedicated to 
Miss Nancy Van Meter who guided us 

safely to the muds of Florida Bay, Ever- 
glades National Park. 

EUCHROMADORA de Man, 1886 

Type species.— Cfiromadora vulgaris Bas- 

tian, 1865: 167, 168, pi. 13, figs. 233- 


This genus is rather difficult because it 
seems to contain a great number of species 
distinguishable by subtle characters of the 
cuticle and the spicular apparatus. Most 
descriptions, however, are too vague to 
permit comparison on a sufficiently de- 
tailed level. We agree with Inglis (1962) 
that the highly developed cuticle possesses 
a number of features that could be used for 
taxonomic purposes. Probably the most 
suitable is the structure of the annules 
underlying the variously shaped blocks, 
rods, and "basketwork"-types of differen- 
tiations. These annules have, along the 
lateral line of the body, anterior or poste- 
rior projections which may be straight, 
notched, fenestrated, or even entirely sepa- 
rated from the annules, thus fomiing small 
cuticular pieces between the latter. All 
these differentiations may be specific but 
more comparative studies are required. 

A grouping of the genus seems to be 
possible along the lines indicated by Wieser 
( 1954 ) . Thus, there is a group of species 
in which the dorsal tooth is relatively small, 
poorly cuticularized, forward pointing and 
not embedded in pharyngeal tissue. This 
group contains the following species: E. 
amokurae (Ditlevsen, 1921), (syn. Spi- 
Io))Jiora amokurae Ditlevsen, 1921); E. 
arctica Filipjev, 1946; E. hiederitzi Steiner, 
1918 (somewhat doubtful); and E. medi- 
terranea Allgen, 1942. All other species 
referred to this group by Wieser ( 1954, 
group B ) are insufficiently described and 
are considered species inquircndac. 

In the second group, the dorsal tooth is 
large and heavily cuticularized, its base 
embedded in pharyngeal tissue. Two of 
the included species are characterized by 
a well-developed, oval bulb, distinctly set 
off from the esophagus, \'iz.: E. Joricata 

Florida Marine Nematodes • Wieser and Hopper 285 

(Steiner, 1916) (synonyms: Spilophora 
loricata Steiner, 1916, E. archaica Steiner 
and Hoppli, 1926), and E. tijrrlienica 
Brunetti, 1951. 

The remaining species are difficult to 
separate. They are Hsted below, together 
with a short characterization of their dis- 
tinguishing features, but the original de- 
scriptions should be consulted in each case. 

E. vulgaris (Bastion, 1865) 

Chromadom vulgaris Bastian, 1865; E. frUlcntata 
Allgen, 1929. 

Spicula asymmetrical. 

E. striata (Eberth, 1863) 

Odonfobius striatiis Eberth, 1863, non E. striata 
of Chitwood, 1951 (=E. gaidica). 

Spicula 70-90 /x, stout, central portion 
enlarged; lateral pieces of gubernaculum 
about 50% of spicula. 

E. gaulica Inglis, 1962 

E. striata of Chitwood, 1951, non Eberth, 1863; 
E. chitwoodi Coles, 1965. 

Spicula 42-53 /x, proximal portion slen- 
der, proximal and "crinkled," distal portion 
enlarged; gubernaculum 50-66% of spicula. 

E. parafricana Gerlach, 1958 

Spicula 25-27 /x, stout; gubernaculum of 
even thickness, boomerang-shaped. Ce- 
phalic setae shorter than in above species. 

E. pectinata n. sp. (See below, p. 286.) 

Closely related to foregoing species. 
Spicula just as stout but more heavily 
cuticularized, 37-40 /x; gubernaculum 
sharply bent, with acute tip. Comb-like 
ridges in buccal cavity. Cephalic setae even 
shorter. Cuticular annules transversely 

E. permutabilis Wieser, 1954 

Spicula 104-133 /x. Tail plump (4 a.b.d. 
in 9 ). Gubernaculum sharply curved dis- 

E. tokiokai Wieser, 1955 

Spicula 41-50 ^u, slender throughout. 
Gubernaculum 66% of spicula, slightly 
cui-ved distally. 

E. meadi n. sp. (See below, p. 286.) 

First circle of 6 cephalic setae elongated, 
second circle — at least in adults — absent. 
The absence of the second circle of ce- 
phalic setae distinguishes E. meadi from 
the remaining species of this group. 

In addition to these species there are two 
more with unusually shaped spicular ap- 
paratus which might not belong to the 
genus at all, viz., E. inflatispicuhtm Schuur- 
mans-Stekhoven, 1943, and E. kryptospicu- 
hirn Allgen, 1951. 

All the other species described are here 
considered as doubtful, or have been trans- 
ferred to other genera by \\'ieser ( 1954 ) . 

Euchromadora gaulica Inglis, 1962 

Plate XXIV, fig. 52, a; Plate XXV, fig. 
52, b-d 

Euchromadora gaulica IngHs, 1962: 260. 

E. striata of Chitwood, 1951, non Eberth, 1863. 

E. chituoodi Coles, 1965 ( xew synonymy). 

L = 1.11-1.57 mm; w = 40-52 fx; esopha- 
gus = 245 /x; Vu = 50%. Head diameter 
19 /x. First circle of six cephalic setae short, 
second circle of four, 7-9 /x. Cuticular 
ornamentation consisting, in front, of three 
trans\'erse rows of dots, followed by the 
usual annules covered with hexagonal 
bodies which themselves are interconnected 
by a meshwork of lines. Further posterior, 
these bodies are more elongated and form a 
sort of grid that links the annules. The 
latter are solid and possess lateral projec- 
tions ( "lateral plates" of Inglis, 1962) which 
are directed anteriorly in the anterior half 
of the body, posteriorly in the posterior 
half. Each projection fits into a notch of 
the following annule, thus forming a series 
of joints. This condition has already been 
described by Steiner (1918). Occasionally 
the lateral projections are separated from 
the annules and then can be seen as cuticu- 

286 BuIIclin Museum of Comparative Zoology, Vol. 135, No. 5 

lar i)ieces between the latter. Traces of 
pigment spots occur in the anterior cervical 
region. Buccal cavity with large dorsal 
tooth and sexeral subventral and \entral 
denticles which form a comb-like ridge. 
Pharyngeal bulb well developed, esopha- 
geal bulb not set off, indistinct. Spicula 
42-47 fj. long, proximal end giving a 
"crinkled" impression, distal two-thirds di- 
lated. Lateral plates of gubernaculum 25- 
28 /x long, proximal end straight or round 
depending on focus, distal end acute, with 
two minute subterminal denticles; dorsal 
plate 26-33 ,m. long, with lateral projections. 
Tail 135-150 jj. long, a.b.d. 30 /x in 6  

Representation in samples studied. — M- 
1, Key Biscayne. 

Geographical distribution. — Texas, Gulf 
Coast (Chitwood, 1951), Mediterranean 
Sea (Inglis, 1962), coast of England 
(Coles, 1965). 

Euchromadora pectinata new species 
Plate XXIV, fig. 53, a; Plate XXV, fig. 53, 

L = 1.65-1.97 mm; w = 40-50 [x; esopha- 
gus = 25S-290 im; \u = 487c. Head diam- 
eter 19 fji. Labial papillae indistinct. Ce- 
phalic setae: first circle of six, papillose; 
second circle of four, 3.5-4 i^. Cuticular 
ornamentation beginning, a short distance 
behind the cephahc setae, with two fused 
annules on which faint longitudinal stria- 
tion can be seen, followed by single annules 
which, however, from about the middle of 
the cervical region to the middle of the 
tail, are transversely split. Conseciuently 
each annul(> consists — at least in the lateral 
portion of the body — of two parts of which 
the posterior one is the larger. This poste- 
rior part possesses the same lateral projec- 
tions as described for the foregoing species, 
only less pronounced. Moreover, there are 
transverse "lacunae" on the annules of the 
mid-body. The differentiation superim- 
posed upon the annules consists of the 
usual hexagonal blocks in the anterior 
cervical region, rod-like markings in the 
remainder o\ the body. These markings are 

thinner than in the foregoing species. The 
cuticle shows ventral or subventral differ- 
entiations in the vulvar region, in the anal 
region, and in males at a distance of 120 
fj. preanal and 70 /x postanal, respectively. 
Buccal cavity with medium-sized tooth, not 
as strongly cuticularized as in other species 
of this group, and a series of comb-like struc- 
tures. Esophagus dilated, no true bulb. 
Spicula 6-7 /x wide, strongly cuticularized, 
without velum, 37-40 jj. long. Lateral plates 
of gubernaculum 20-22 ix long, sharply 
bent distallv and with acute tip. Tail in 
6 , 160-180' ij. long, a.b.d. 35 /x, in 9 , 200 
IX long, a.b.d. 27 /x. 

Ilolotijpc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4073, Type slide No. 88. 
Type locality, M-8, Biscayne Bay. 

Representation in samples studied. — M- 
8, Biscayne Bay. 

Euchromadora meadi new species 

Plate XXIV, fig. 54, a, b; Plate XXV, fig. 
54, c-f 

L = 1.50 mm; w = 40 /x; esophagus = 200 
/x. Head diameter 19 fx. Lips and labial 
papillae distinct. First circle of cephalic 
setae 2 /x, no second circle in adults. In 
juveniles four short setae could be seen 
immediately behind the first circle of six 
setae. Amphids fairly distinct, 12-13 /x 
wide. Cuticular ornamentation beginning 
with large annule, longitudinally striated, 
followed by narrower annules which are 
all solid and show no secondary develop- 
ment except the lateral projections de- 
scribed in the species above. Hexagonal 
blocks verv thin even in anterior cei-vical 
region. Buccal cavity with large dorsal 
tooth and at least two subventral projec- 
tions, no comb-like ridges. No distinct 
esophageal bulb. Spicula 45—47 /x long, 
slender, with velum. Lateral plates of 
gubernaculum 22-24 /x long, hammer- 
shaped; dorsal plate 18 /x long, with char- 
acteristic proximal projection. In the anal 
area the cuticular annules are weaklv 

Florida Marine Nematodes • Wieser and Hopper 287 

cuticularized subventrally to form a sort 
of "window." Tail 135 /x long, a.b.d. 35 fx, 
in juveniles 5-6 a.b.d. long. 

Holotypc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 89. 
Type locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Type species.— Sp//op/iora parva de Man, 
1893: 89-91, pi. 5, fig. 5. 

We found what appeared at first to be 
A. parca (de Man, 1893), but closer study 
revealed the following differences: 

parva dcniiculata 
Spacing of longitudinal 

rows (mid-body) 3.5 ^ 6 /^ 

Length of spicula 17-18 ix 34 ^ 

Distal end of gubernaculuni straight denticulated 

Length of spinneret 12 ^ 7-8 ^ 

= V- tail = lis tail 

Afrochromadora denficulafa new species 
Plate XXVI, fig. 55, a-f 

L = 0.79-0.80 mm; w = 31-32 /x; esopha- 
gus = 96-109 ix; Vu = 467f . Head diameter 
11 fx. Lips distinct, labial papillae and first 
circle of cephalic sense organs not seen. 
Cephalic setae 5-6 fx. Cervical setae as in 
A. parva. Amphids distinctly spiral, but 
small. Cuticular ornamentation beginning 
with transverse rows of dots, followed by 
annules. In the lateral region of the body 
a few dots or rod-like markings can be seen 
between the annules, particularly two 
longitudinal rows of larger dots, the spac- 
ing of \\'hich is 5-6 fx between cervical re- 
gion and anus. Buccal cavity with three 
solid, subequal teeth. Esophageal bulb 
round. Excretory pore 37 fx in front of end 
of esophagus, \'entral gland reaching to 48 
fx behind the esophagus. Spicula nearly 
semicircular, 3 /x wide, 34 ^ long. Guber- 
naculum 23 fx long, strongly dilated distally, 
with denticulated end plate. Tail 124-128 
IX long, a.b.d. 28 /x in 6 , 20 /x in 9 . Spin- 

neret 7-8 fx. In 6 there is a slight ventral 
swelling, 45 fx postanally. 

HoJotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 90. 
Type locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 


MESONCHIUM Cobb, 1920 
Type species.— /Vlesoncfi/um poriferum Cobb, 
1920: 294, 295, fig. 76. 

Pepsonema Cobb, 1920: 295. 

The following fixe species are known: 
M. poriferum Cobb, 1920; M. pcllucidum 
(Cobb, 1920) syn. Pepsonema pcllucidum; 
M. nini Inglis, 1961; M. punctatum Timm, 
1961; M. janetae Inglis, 1963. They can be 
separated mainly by the shape of their 
spicula and by the pattern of their lateral 
cuticular differentiations. Moreover, M. 
janetae has shorter cephalic setae, and M. 
nini amphids with more turns than the rest 
of the species. 

Our species agree with the description of 
M. pcllucidum by Cobb (1920) except that 
in the posterior portion of the body we 
observed four longitudinal rows of dots, 
whereas Cobb states that near the tail 
"there are sometimes six or possibly eight 
rows." We cannot agree with Timm's 
(1961) synonymization of M. pcllucidum 
with M. poriferum since in the tvvo species 
the longitudinal rows are differently ar- 
ranged, and in M. pcllucidum the spicula 
are retrorsely barbed (a fact already men- 
tioned by Cobb), whereas in M. poriferum 
they are acute distally. The description of 
M. punctatum Timm is confusing. In the 
text the presence of the usual number of 
four cephalic setae is mentioned, whereas 
the figure shows the head equipped with 
what seems to be eight or ten setae. In the 
text the amphids are said to describe 2.5 
turns; in tlie figure they definitely have 
three turns. Even without these confusing 

288 Bulletin Museum of Comparative Zoology, Vol. 135, No. 5 

discrepancies it would be difficult to give 
unequivocal reasons %\'hy M. punctatum 
and M. porifcrum .should be distinct spe- 

hAesonchium pellucidum (Cobb, 1920) 
Plate XXVII, fig. 57, a-d 

Pepsonema pvUucidiim Col^b, 1920: 295, 296, 

fig. 77. 

L = 1.77-2.00 mm; w = 73-75 ju; esopha- 
gus = 250 IK \'u = 477< . Head diameter 
15 fx. Cephalic setae 12 /j.. Amphids 10 p., 
2.5 turns. Buccal cavity cylindrical, 23 ij. 
long, with three teeth. (The heads of our 
male specimens agree in eveiy respect with 
Cobb's figures.) Cuticular differentiation: 
In both sexes there are four longititdinal 
rows of coarser dots, running from behind 
the amphids to behind the end of the 
esophagus. From there on the two outer 
rows gradually disappear and the whole 
lateral field is raised to form a lateral wing. 
In mid-body there are only two longitudi- 
nal rows of coarser dots. In the posterior 
portion of the body the wings are flattened 
again and the two outer rows of dots re- 
appear. The spaces between the coarser 
dots are resolvable with difficulty into 
transverse rows of minute and closely 
spaced dots which are continuations of the 
usual transverse rows of dots into the 
lateral fields. Esophagus with a barrel- 
shaped posterior bulb and a small cardia. 
Ventral gland opposite end of esophagus, 
excretory pore behind nerve ring. Spicula 
110 ;u, long, proximally cephalate, distally 
retrorsely barbed. Cubernaculum heavy, 
with 30 IX long apophysis. There are about 
16 indistinct tubular supplements. Tail 200 
ij. long, a.b.d. 45-50 /x. 

Representation in .samples studied. — M- 
8, Biscayne Bay. 

Geographical distribution. — Kingson, Ja- 
maica (Cobb, 1920). 

SABAIIERIA de Rouville, 1903 
Type species.— Soba/ier/a cettensis de Rou- 
ville, 1903: 11. 

Two closely related, but distinct, species 
of Sabatieria were found in the sample 

from Vero Beach. Following the key 
given by W^ieser, 1954, they belong to 
the group encompassing S. cupida Bresslau 
and Schuurmans-Stekhoven in Schuurmans- 
Stekhoven, 1935, S. heterura (Cobb, 1898), 
S. rugosa Schuurmans-Stekhoven, 1950, S. 
siinilis (Allgen, 1933), and S. tcnuicaudata 
(Bastian, 1865). 

The two new species, S. paradoxa and S. 
paracupida, can be separated from the 
above-mentioned related species by use of 
the following key. 

Key to Group Based on Male Characteristics 

1. Supplements 28-32 

S. teniiicaudata (Bastian, 1865) 

( doubtful species ) 
Supplements 15-22 2 

2. Cephalic setae more than % of head diam- 

eter in length 3 

Cephalic setae less than % of head diam- 
eter in length 4 

3. Spicules 46-50 ^ long, 1.2 anal body 


S. cupula Bresslau and Schuurmans-Stek- 
hoven in Schuurmans-Stekhoven, 1935 
Spicules 63-68 ix long, 1.7-1.8 anal body 
diameters S. paracupida n. sp. 

4. Spicules 1.6 or more anal body diameters in 

length 5 

Spicules 1.3 or less anal body diameters in 
length 6 

5. Spicules 85 |t long, vv'ithout proximal cepha- 

lation S. heterura (Cobb, 1898) 

Spicules 60-62 m long, proximally distinctly 
cephalated S. paradoxa n. sp. 

6. Amphid with 2.5 turns 

S. rugosa Schuurmans-Stekhoven, 1950 

Amphid with 2 turns 

S. similis (Allgen, 1933) 

SabotieriQ paradoxa new species 
Plate XXVII, fig. 58, a-d 

L = 1.46-1.66 mm; w = 42-44 /.. Diam- 
eter at base of esophagus 3^39 /x. Esopha- 
gus 135-140 IX. Tail 140-142 /x (4.0-4.1 
a.b.d.). Head diameter 13 ^. Labial and 
cephalic papillae distinct. Ceplialic setae 
5-6 IX long, 38-46% of head diameter. Cer- 
vical setae, short, widely dispersed, somatic 
setae shorter and more widely spaced. 
Amphids spiral, with 2.25 turns; 7 jx. wide 
(54% of head diameter). Lateral differen- 

Florida Marine Nematodes • Wiescr and Hopper 


tiation prominent, the transverse rows of 
punctations overlying the lateral chords 
being coarser and more widely spaced than 
on the remainder of the bodv. Spicules 
60-62 /x long (1.7-1.8 a.b.d.),' proximally 
cephalated, distally quite narrowed. Gu- 
bernaculum with lateral guiding pieces. 
Apophyses of gubernaculum 25-27 /x long. 
Preanal supplements minute, 17-19 in num- 
ber. Caudal setae present, arranged as 
illustrated (PI. XXVII, fig. 58, b). 

HoJohipc specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4075, Type slide No. 92. 
Type locality, V, Vero Beach. 

Representation in samples studied. — M- 
7, Everglades National Park, V, Vero 

SabaWer'ia paracupida new species 
Plate XXVIII, fig. 59, a-c 

L = 1.7-1.85 mm; w = 47-52 /x. Diam- 
eter at base of esophagus 44 fi. Esophagus 
180-184 fx. Tail 148-150 /x (4 a.b.d.). Head 
diameter 13-14 fi. Labial and cephalic 
papillae distinct. Cephalic setae 10 fx long, 
71-77% of head diameter. Cervical and 
somatic setae similar to preceding species. 
Amphids spiral, with 2.25 turns; 9 /x wide 
( 64-70% of head diameter ) . Lateral differ- 
entiation as in preceding species. Spicules 
63-68 IX long (1.7-1.8 a.b.d.), lacking proxi- 
mal cephalation. Gubernaculum with lat- 
eral guiding pieces. Apophyses of guber- 
naculum 20 fx long. Preanal supplements 
minute, 19-22 in number. Caudal setae 
present, arranged as illustrated (Pi. XXVIII, 
fig. 59, c). 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4075, Type slide No. 93. 
Type locality, V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 


AXONOLAIMUS de Man, 1889 

Type species.— Anop/osfoma spinosum 

BiJtschli, 1874 sensu de Man, 1888: 

19-21, pi. 2, fig. n. 
Axono/a/Vnus hexapiius new species 
Plate XXVIII, fig. 60, a-e 

L = 6 , 1.8, ? , 1.9 mm; w = i , 28, 9 , 
35 /x; diameter at base of esophagus 28, 31 
ya. Esophagus 172, 137 /x. Vu = 54%. Tail 
128, 138 fx (4.3-4.6 a.b.d.). Head diameter 
13, 14 IX. Head with six labial papillae and 
four cephalic setae (21-24 /x long). An 
additional 21-24 /x long seta is located 
about 5 n behind each amphid. Amphid 
12-13.5 IX long by 6-7 /x wide, slightly to 
distinctly open loop-shaped, 8 /x from 
anterior end. Buccal cavity 14-16 /x deep, 
of which the posterior conoid portion 
makes up about 10 /x. Anterior part of 
buccal cavity with six weak odontia. 
Esophagus clavate, 10 /x wide at base of 
buccal cavity, 12 /x at the nerve ring, ex- 
panding in the posterior % to 20 /x at the 
base. Excretory pore slightly behind 
middle of conoid portion of the buccal 
cavity. Excretory pore ampulla 40 ^ from 
head end, at level of "break" in esophageal 
musculature. Renette cell large, posterior 
to base of esophagus. Pseudocoelomocyte 
immediately behind renette cell. Cuticle 
finely striated. Somatic setae about 5 /x 

Female didelphic, amphidelphic, ovaries 
outstretched; with two eggs observed in 
the posterior uterus (eggs 75-110 /x by 30 
ix). Male diorchic, testes opposed, out- 
stretched. Spicules 39 ix long. Gubernacu- 
lum short, with 12 ix long apophyses. Sup- 
plementary organs glandular, with minute 
exit pores. Male and female tail obtusely 
conoid. Caudal setae arranged as illus- 
trated (PI. XXVIII, fig. 60, c), those near 
terminus longer than the remainder. Spin- 
neret three-lobed, the associated glands 
located in the tail. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes. Ento- 
mology Research Institute, Ottawa, Collec- 

290 Bulletin Mu.scui)} of Coiiijxinitivc Zoology, Vol. 135, No. 5 

tion Number -4070, Type slide No. 94. 
Type loeality, M-5, Virginia Key. 

Representation in samples studied. — M- 
5, X'irginia Ke\-. 

Remarks. — A. Jie.xapihis n. sp. can be 
distinguished from all other A.X(»iokiimus 
species by the presence of a long lateral 
seta associated near the base of each am- 
phid. In possessing this character, along 
with the tri-lobed spinneret, this animal 
has some relationship to the monodelphic 
genus Synodontium Cobb, 1920. 

ODONTOPHORA Butschli, 1874 

Type species. — Oc/onfopfiora morina 

Butschli, 1874: 285, pi. 3, fig. 13. 
Odontophora voriabilis new species 

Plate XXVIII, fig. 61, a, b; Plate XXIX, 

fig. 61, c-e 

L = 1.3-2.2; w = 36-50 /.-; Vu = 53-56%. 
Diameter at base of esophagus 29-33 /x. 
Esophagus 112-135 jx long. Head diameter 
11-15 /x. Head with cephalic papillae, 4 
cephalic setae and 3 circles of subcephalic 
setae. Cephalic setae 14-17 /x long in 6 , 
11-14 IX in 9 . First circle of subcephalic 
setae (paramphidial ) 11-13 fx, second 
circle, S-9 /x, third 4-5 /x long. The sub- 
cephalic setae all occur within the range 
of the buccal cavity. Cervical setae located 
1.5-2.5 /x posterior to base of buccal cavity, 
the first circle the longest, 7-8 /x, the next 
two following closely and being progres- 
sively shorter. Amphid (S-10 /x long. Buccal 
cavity 23-2.5 /x long, the posterior conoid 
portion being 17 /x long. Buccal cavity with 
6 odontia, the odontia with prominent 
apophyses to which are attached longi- 
tudinally directed muscles, an arrangement 
that undoubtedly serves to evert the odon- 
tia. I^xcretory pore at base of odontia. 
Eggs 100-110 /x long by 45 /x wide, two per 
uterus. Spicula sickle-shaped, 47-49 /x long, 
the chord being 32-33 /x. Gubernaculum 
with 12-13 /x long apophysis. Preanal sup- 
plements not observed. Tail length vari- 
able. 70-117 /x long. Male tail with numer- 

ous caudal setae and with tenninal setae 
17-18 IX long. Female tail with fewer 
caudal setae and devoid of terminal setae. 

Holofype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4075, Type slide No. 95. 
Type locality, V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 

Remarks. — The specific name variabilis 
ser\es to indicate the variation found within 
the specimens recovered from the single 
sample taken from the Vero Beach area. 
Body dimensions, such as total length, 
maximum width and tail length, are quite 
variable. Morphological stioictures, how- 
ever, i.e., spicula, buccal cavity, amphid, 
distribution and size of cephalic and sub- 
cephalic setae, were conspicuously in close 
agreement. In particular, the shapes of the 
spicula were nearly identical despite the 
fact that the tails in the three male speci- 
mens observed measured 70 ^, 85 ^i, and 
117 /x in length. 

O. variabilis, in possessing, in addition to 
the cephalic setae, three circles of subce- 
phalic setae which could be interpreted as 
being arranged in four submedian rows, is 
related to O. aniiustilaimoides Chitwood, 
1951, and perhaps may be that species. 
However, no mention is made of the pres- 
ence of cervical setae in O. angiistilaimoides, 
these being a prominent feature in O. vari- 
abilis. In addition, the size of the eggs and 
the number of eggs per uterus differs. 
Moreover, since no males have been de- 
scribed of O. angustilaimoides, this species 
is ot doubtful status. 


Type species.— Pseuc/o/e//a paragranulifera 

Timm, 1952: 45, pi. 9, fig. 78. 
Parodontophora brevomphida (Timm, 1952) 

Timm, 1963 

Pscudoldla brcvanipliida Timm, 1952: 44, 45, pi. 

9, 76, 77. 
Parodontophora ])acifica ( Allgen ) sensu Timm, 

1963: 35, 36, fig. 1, y, h (nf;av synonymy). 

Florida Marine Nematodes • Wicscr and Hopper 291 

L = 1.1-1.4 mm; ^^' = 36-lS /x; Vu = 47- 
50%. Body bearing four rows of somatic 
setae, two on either side of each lateral 
chord. Anteriorly the somatic setae are 
about 4 fx long in the region of the nerve 
ring. At the vuKa their length is reduced 
to less than 2 /x. Except for the anterior- 
most 14 /A, the cuticle is finely striated. One 
dorsal and two ventral setae are located in 
the region of the amphid. Amphid short 
shepherd's crook, with the ventral ann 
longer than the dorsal. Dorsal arm about 
10 11 long, ventral arm 14-17 ^ long. Am- 
phid located on a level with and distinctly 
shorter than the 19 /x long posterior cylin- 
drical portion of the stoma. Head rounded, 
with six lips, six labial papillae, and four 
7-8 IX long, cephalic setae (75% of corre- 
sponding body diameter). There are no 
subcephalic setae. Anterior part of stoma 
with six prominent odontia, posteriorly 
cylindrical, 25-28 /x deep. Esophageal 
diameter increasing posteriorly; without 
basal bulb. Approximately 15 ^a posterior 
to the base of the buccal cavity the tuboid 
esophageal marginal rays commence. Nerve 
ring encircling esophagus at approximately 
65% of its length. Excretory pore not de- 
tected, the ampulla, however, located at 
the base of buccal cavity. Renette cell 65 
/u. long (407c of esophageal length), located 
posterior to base of esophagus. A promi- 
nent pseudocoelomocyte occurs immedi- 
ately posterior to the renette cell. 

Female didelphic, amphidelphic, ovaries 
outstretched. Eggs not observed. Male 
diorchic, testes opposed, outstretched. 
Spicules arcuate, 31-36 ^ long (according 
to angle of view). Gubernaculum 8-10 ^ 
long, arcuate. Male and female tail elon- 
gate-conoid, 135-155 fj- long, the terminal 
Vs nearly cylindrical. Subterminal setae 
present on some specimens. Male with a 
short preanal seta and two subventral rows 
of 7-8 setae. Spinneret present, the gland 
cells located in the anterior portion of the 

Representation in samples studied. — V, 
Vero Beach. 

Geographical distribution . — Chesapeake 
Bay, Maryland (Timm, 1952), Bay of 
Bengal (timm, 1961), Arabian Sea at 
Karachi (Timm, 1962), Maldives (Ger- 
lach, 1962). 

Remarks. — At the present time consider- 
able confusion exists in the taxa that have 
been identified as P. paeifica (Allgen, 
1947) (syn. Odontophora p. Allgen). Ger- 
lach ( 1962 ) considers a variety of species 
as synonyms of Allgen's species. In his list 
of synonyms, Gerlach lumps species in 
\\'hich the amphids are 18 ^ long together 
with species in \\'hich the amphids are 80- 
150 /x long. This action seems a bit prema- 
ture and is considered doubtful. Timm 
(1963) in creating the genus Farodonto- 
phora for species of Odontophora with 
parallel stomatal walls, intimates that P. 
paeifica might best be considered as a 
species inquirenda, validating P. quadri- 
sticha ( Schuurmans-Stekhoven, 1950). We 
agree with Timm's statement and herein 
consider P. paeifica (Allgen, 1947) as a 
species inquirenda. However, in support- 
ing such a move we do not feel that all 
subsequent P. paeifica descriptions refer 
to P. quadristicha and feel that P. brevam- 
phida can be suitably differentiated from 
Schuurmans-Stekhoven's species. 

For the present the following differences 
can be used to separate the two species 
under consideration: 

P. quadristicha [syn. O. paeifica Allgen 
of Wieser, 1956] has amphids 24-32 fx long, 
a buccal cavity 40 /x long, and a renette 
cell 90-100 ^'long (app. 50-55% of the 
esophageal length ) . 

P. brevamphida [syn. Pseudolella pa- 
eifica (Allgen, 1947) of Timm, 1961, 
Odontophora paeifica Allgen of Gerlach, 
1962, and Parodontophora paeifica (Allgen, 
1947) of Timm, 1963] has amphids 12-18 
/x long, a buccal cavity 25-31 jj, long, and a 
renette cell 32-65 ,i long (app. 30-40% of 
the esophageal length). 

292 Bidhiin Mii.scuiii of Comparative Zoology, Vol 135. No. 5 


ALAIMELLA Cobb, 1920 

Type species.— A/a/me//a fruncafa Cobb, 

1920: 234, fig. 7b. 
Alaimella cincto Cobb, 1920 

Plate XXIX, fig. 63, a-c 
AlaimelU cincta Colih, 1920: 233-234, fig. 7a. 

Mu]c,—L = 1.3 mm; w = 16 /x; diameter 
at base of esophagus 13 ix. Esophagus 250 
ja long, \\ itli conoid cardia. Head 7 /x wide, 
bearing six papillae and four 12-13 /x long 
cephalic setae. Aniphid 7 p. wide, with 
central raised "fleck." Cuticle coarsely 
annulated, the annules bearing prominent 
longitudinal markings. Spicules 27 /x long 
(chord 22 /x), proximally cephalated. Gu- 
bernaculum 9 /x long, fail 100 /x long (7 
a.b.d. ), with at least one caudal seta. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Geographical distribution. — Bis- 
cayne Bay, Florida (Cobb, 1920), Aransas 
Bay. Texas (Chitwood, 1951). 

Remarks. — The central raised portion of 
the amphid in our specimens, while illus- 
trated by Chitwood for his example, was 
not originally depicted by Cobb. This 
feature is somewhat suggestive of A. trun- 
cata Cobb, 1920. However, regarding the 
cuticle of A. truncata, Cobb states, "Sec- 
ondary markings of the cuticle faint, if 
any." The Miami specimens, by manifest- 
ing prominent longitudinal markings, can- 
not be regarded as representative of A. 
fruneata in view of this strong statement 
questioning the presence of such markings. 
Thus, even though the amphid on our 
specimens has a central fleck, we feel the 
remainder of the characters support our 

CAMACOLAIMUS de Man, 1889 

Type species.— Camoco/a/mus tardus de 

Man, 1889a: 8. 
Comaco/a/mus prytherchi Chitwood, 1935 
Plate XXIX, fig. 64, a-c 

CamucoUiimus prytherchi Chitwood, 1935: 49, 
50, fig. 7, a-c. 

L = 5 , 1.8, 9 , 1.9; w =: 5 , 33, $ , 39; 
diameter at base of esophagus .30-32 /x; 
esophagus 240-250 /x long. Vu = 54%. Tail 
90-105 ^ (3.1-3.4 a.b.d.). Head 13-14 ix 
wide, bearing six labial papillae and four, 
8-10 /x long, cephalic setae. Somatic setae, 
short and thick, staggered in region over- 
lying lateral chords, widespaced. Cuticle 
finely striated. Lateral surfaces with slight 
"bulges" anteriorly, progressively getting 
higher posteriorly until finally lateral alae 
are formed at a point approximately mid- 
way from vulva to anus, fading away 
quickly on the tail. Dorsal tooth and its 
base, 15 /x long. Esophagus divided into 
three distinct regions — corpus, isthmus, and 
swollen terminal region. Nerve ring en- 
circling isthmus immediately behind the 
coipus. Tenninal region gradually en- 
larged. Cardia 8 /x long and 12 /x wide. 
Spicules 53 /x long. Male tail with caudal 
alae and a pair of postanal setae. 

Representation in samples studied. — V, 
Vero Beach. 

Geographical distribution. — North Caro- 
lina (Chitwood, 1933); Chile (Wieser, 
1956); Maldives (Gerlach, 1962). 

Remarks. — An examination of the type 
specimen of C prytherchi discloses the 
presence of both the caudal alae and the 
postanal setae. We wish to thank Mr. Cur- 
tis Sabrosky for making the specimens 
available for study. 


Some genera of this family have recently 
been transferred to the family Linhomoei- 
dae by Gerlach (1963a). We postpone 
judgment on these changes until the ma- 
terial for our monograph is complete and 
shall adhere, in this paper, to the old clas- 

PARATARVAIA new genus 

Type species.— Paraforvo/a sefa n. sp. 

Definition. — Diplopeltidae (or Linho- 
moeidae?). Head with 6 labial papillae 
and 10 cephalic setae in two circles, 6 + 4. 

Florida Marine Nematodes • Wieser and Hopper 293 

Amphid, doulile spiral, situated on punc- 
tated plaque. Esophagus cylindroid with 
small basal bulb, without valves. Cardia 
small. Female didelphic, amphidelphic. 
Male without supplements. Spicules arcu- 
ate, gubernaculum with medial piece, 
lateral guiding pieces and posteriorly di- 
rected apophyses. 

Remarks. — Parotorvoia possesses charac- 
ters that show affinity with both the 
Diplopeltidae (spiral amphid, amphid on 
plaque ) and the Linhomoeidae ( reduced 
buccal cavity and spicular apparatus as in 
Terschellingio, number of cephalic sensory 
organs ) . However, as we are going to post- 
pone our judgment of the recent proposals 
by Gerlach (1963), we provisionally assign 
Paratorvaia to the Diplopeltidae. Within 
the Diplopeltidae Parafarvoia is related to 
both Tarvaia Allgen, 1934, and Disconema 
Filipjev, 1918. It is distinguished from 
Tarvaia by possessing 10 cephalic setae, as 
opposed to four. From Disconema, which 
lacks a plaque, Paratarvaia is separated by 
possessing a punctated plaque as well as 
by having the 10 cephalic setae separated 
into two circles of 6 and 4. 

Paratarvaia sefa new species 

Plate XXX, fig. 65, a-c 

L =: l.;3-1.4 mm; \\- = 23-24 /x; width at 
base of esophagus 20-21 fx. Head 13 ^. 
wide, bearing an internal circle of six setose 
labial papillae and a divided external circle 
of 10 (6 + 4) setae, 16 + 18 /x in length. 
Amphid a double spiral, 21-23 /x long and 
16-17 fx wide, situated on a punctated 
plaque 25-27 /jl long and 17 {x wide. In 
dorsal-ventral view the plaques are seen 
to be separated by a distance of 2-3 /x. 
Cuticle coarsely annulated, the annules 
about 1.5 IX wide just posterior to the am- 
phids, about 2.0 ix at mid-body and about 
1 IX wide on the tail. Body with fine so- 
matic setae which extend onto the tail. 
Buccal cavity extremely reduced, \\'ith 
minute lips protruding from the head con- 
tour. Esophagus 165-170 ^ long, cylindroid 
to a small, non-valvular, terminal bulb. 
Cardia present, flattened, not prominent. 

Excretory pore not observed. Vu = 54%, 
female with two opposed ovaries. Number 
of testes in male not determined. Spicules 
arcuate, 40 /x long (chord 25 /x). Guber- 
naculum complicated, bearing a medial 
piece between the spicules and two sigmoid 
lateral guiding pieces. Apophyses to guber- 
naculum 10 /x long. Male without supple- 
ments. Tail 150-180 /x long, the anterior % 
conoid, the remainder cylindroid to the 
slightly swollen terminus. Terminus with 
spinneret and two 8 /x long, terminal setae. 

HoJotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 96. 
Type locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 

DIDELT A Cobb, 1920 

Type species.— D/c/e/fa maculatum Cobb, 
1920: 252, 253, fig. 33. 

We found in our material one juvenile 
specimen which in every respect resembles 
Cobb's Didelta maculatum. Since Cobb's 
figure of the head is very good we supply 
only a figure of the tail of our single speci- 

Didelta maculatum Cobb, 1920 
Plate XXX, fig. 66 

Didelta maculatum Cobb, 1920: 252, 253, fig. 33. 

Head diameter 23 fx. Cephalic setae 15 + 
3 II. Amphids ( with plaque ) 25 X 17 /x. 
Esophagus enlarged posteriorly, short car- 
dia (25 11 long, 30 jx wide) present. Tail 
410 /x, a.b.d. 37 p.. No spinneret or caudal 

Representation in samples studied. — M- 
8, Biscayne Bay. 

Geographical distribution. — Off Key 
West, Florida (Cobb, 1920). 


Type species.— Tersche///ng/a communis de 
Man, 1888: 12, pi. I, fig. 7. 

The material from Florida contained 

294 Bulletin Museum of Compamiive Zoology, Vol. 135, No. 5 

three species of TerschcUiuiiia. Excellent 
examples of T. Jongicaudata were found 
both in Biscayne Bay and in Florida Bay, 
with a variant found at Vero Beach. The 
distribution of the cervical setae in the 
specimens from the former habitats is ex- 
actly as depicted by de Man in his original 
figures of the species. The variant from 
Vero Beach has a slightly different arrange- 
ment of the cerx'ical setae, viz., a reduction 
in number and a more posterior location of 
the first circle (compare Pi. XXX, figs. 67a, b, 
and 6Sa, b). These differences are stable 
\\ ithin Florida habitats. Subsequent collec- 
tions from South Carolina contain speci- 
mens, not onlv with the above two diver- 
gent patterns of cervical setae, but with 
intermediary distributions as well. 

A second species, T. monohystem n. sp. 
is unique in that only the anterior ovary 
appears to be well developed, the posterior 
gonad being quite rudimentary. The third 
species, T. lon^spiculata n. sp., can be 
distinguished by the arrangement of the 
cephalic and cervical setae and also by the 
long spicules, which are about 2.5 anal body 
diameters in length. 

Terschellingio longicaudata de Man, 1907 
Plate XXX, figs. 67, a-c, 68, a, b; Plate 
XXXI, figs. 67, d, 68, c-e. 

Terschellin^id lon^icaiuhitd de Man, 1907: 230. 

L = 1.1-1.3 mm; w = 29-32 /x; diameter 
at base of esophagus 27-29 jj,. Esophagus 
90-125 II- nerve ring at 50%. Head with 
four cephalic setae, 4-5 fi long. Cervical 
setae 4-5 p. long and arranged in distinctive 
patterns. The Biscayne Bay and Florida 
Bay specimens have two paramphidial pairs, 
two postamphidial pairs and a somewhat 
broken circle of eight situated about mid- 
wav between the amphid and the nerve 
ring (PI. XXX, fig. 67a). The Vero Beach 
specimens have a circle of four postamphid- 
ial, two sublateral and a dorsal-ventral pair 
(PI. XXX, fig. 6(Sa, b). Amphids circular, 
7-8 fx wide. Terminal esophageal bulb 25 
/x long and nearl\' of equal width. Cardia 

elongate, 14 /x long, 5 /x wide. Excretory 
pore located at a level slightly anterior to 
the anterior end of the esophageal bulb. 
Spicules 38-46 /x long (about 1.4-1.8 a.b.d), 
proximally cephalated. Apophyses of guber- 
naculum 10-12 /x long. Tail 325-365 /x long 
(about 14 a.b.d.). 

Representation in samples studied. — M-4, 
Rickenbacker Causeway; M-6, M-7, Ever- 
glades National Park; M-8, Biscayne Bay; 
V, Vero Beach. 

Geofiraphical distribution. — Cosmopoli- 

Jerschellingia monohystera new species 
Plate XXXI, fig. 69, a-f 

L = 0.97-1.06 mm; w = 33-35 fx; Vu = 
44%. Diameter at base of esophagus 31 /x. 
Esophagus 83-93 [x. Head diameter 13 /x. 
Head with four cephalic setae, 4 /x long. 
Four somatic setae occur in the cervical 
region. The anteriormost lies 8-12 /x behind 
the base of the amphid and is placed ventral 
to the lateral surface of the body. The three 
others are staggered dorsal and ventral in 
respect to the lateral surface. Amphid cir- 
cular, 5 ij. wide, 3-5 /x from anterior end; 
corresponding body diameter 14-15 jj.. Fe- 
male with only the anterior ovary developed 
and producing eggs (195 /m long), posterior 
branch rudimentary (70 ^a long). Spicules 
40 fx long (about 1.3 a.b.d.), without proxi- 
mal cephalization. Apophyses of guber- 
naculum about 13 fx long. Tail 220-225 //. 
long (7.5-10 a.b.d.), of which the posterior 
half is filiform. Caudal setae arranged as 
illustrated (PI. XXXI, fig. 69e). 

JloJotiipe speeimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4075, Type slide No. 99. Type locality, 
V, Vero Beach. 

Representation in samples studied. — V, 
Vero Beach. 

Terschelimgia longispiculafa new species 
Plate XXXI, fig. 70, c, d; Plate XXXII, 
fig. 70, a, b 

L = 2.2-2.3 mm; w = 63-65 /x; Vu = about 

Florida Marine Nematodes • Wieser and Hopper 295 

40%. Diameter at l)ase of esophagus 55- 
57 fj.. Esophagus with six cephalic papillae 
and four cephalic setae, 4-5 ^ long. Two 
circles of prominent cervical setae present, 
four setae in each circle and all setae 4 /x 
long. The first circle occurs at the level 
of the posterior edge of the amphids. In 
addition to the prominent cervical setae, 
setae of lesser stature also occur in the 
anterior neck region. While these could be 
arranged in transverse circles of eight 
(sometimes incomplete), they could also 
be said to be arranged in eight longitudinal 
rows in which the prominent cervical setae 
might represent the anteriormost seta of 
each ro\\'. Amphid circular, 10 ^ wide, 11- 
14 fj. from anterior end; corresponding body 
diameter 3.3-36 fj.. Terminal esophageal 
bulb 50 fx long and 42 ^ wide. Spicules 122 
fx long (about 2.5 a.b.d. ); proximally cepha- 
lated. Apophyses of gubemaculum 25-30 
/A long. Tail 435 ^ long ( about 10 a.b.d. ) . 
Caudal setae arranged as illustrated (PI. 
XXXI, fig. 70, c). 

HoJoUjpc specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4072, Type slide No. 98. Type locality, 
M-7, Everglades National Park. 

Representation in samples siiidied. — M-7, 
Everglades National Park, and M-8, Bis- 
cayne Bay. 




Type species.— Monohystera (Paramonohys- 

fera) megacephala Steiner, 1916: 639- 

641, pi. 32, fig. 37, a-f. 

A key to the subgenera and species of 
Paramonhystera is provided by Wieser 
(1956). Following this author, the sub- 
genus Paramonhystera sensti stricto con- 
tains the following species: P. megacephala 
Steiner, 1916, P. micramphis Schuurmans- 
Stekhoven, 1950, P. biformis Wieser, 1956, 
and P. proteus Wieser, 1956. Of these, P. 
micramphis is known from females and 
juveniles only and is considered a species 

inquircnda. Our new species, P. canicula 
n. sp., is distinguished from the remaining 
species by the long cephalic setae, the 
round amphids and the characteristic shape 
of the gubernaculum. 

Paramonhystera canicula new species 
Plate XXXII, fig. 71, a-d 

L = 1.60 mm; w = 42 /a; esophagus = 250 
IX. Head diameter 25 /x. Lips large, rounded, 
each with two labial setae 5 fx long. Ce- 
phalic setae 23 + 18 /x. Amphids with very 
faint contour, convex, in S , 14 ix = 52% of 
c.b.d. wide. Cervical and somatic setae in 
irregular longitudinal rows, the former 10- 
12 fx long. One short, asymmetrical (dorso- 
lateral) seta between amphid and cephalic 
setae on each side of body. Cuticular an- 
nulation coarse. Spicula 135 fx long, 
knobbed proximally. Gubemaculum rather 
complicated, consisting of a proximal por- 
tion, cylindrical and 20 ix long, and a distal 
portion, IS /x long, dilated, with ventrolateral 
projections, a deep notch and a serrated 
distal edge. Tail 175 /x long, a.b.d. 35 /x. 

HoJotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4068. Type slide No. 100. Type locality, 
M-3, Key Biscayne. 

Representation in samples studied. — M-3, 
Key Biscayne. 

STEINERIA Micoletzky, 1922 

Type species.— A^ono/iysfera polychaeta 

Steiner, 1915: 224-226, figs. 1-3 (new 

subsequent designation). 

MonJiystera setosissima Cobb, 1893, trans- 
ferred to Steineria and regarded as the type 
species by Schuunnans-Stekhoven and De 
Coninck (1933: 10), is herein rejected as 
the type species of the genus Steineria on 
the grounds that it was not one of the in- 
cluded species brought to the subgenus 
Steineria when first established by Micolet- 
zky (1922a: 168). This action is in accord 
with Article 69a (ii) of the International 
Code of Zoological Nomenclature adopted 

296 Bulletin Museum of Coniixiidtivc Zoology, Vol. 135, No. 5 

by the XV International Congress of Zo- 
ology, London, July, 1958. 

In this genus, two species with punctate 
cuticle have been known so far, i.e., S. 
punctata Gerlach, 1955, and S. gcrhchi 
\Meser, 1959. P'rom our Florida material a 
third species can be added that is rather 
closely related to S. punctata but can be 
separated on the following counts: sub- 
cephalic setae in eight groups of 5 and 4 
(instead of 6 and 3), cephalic setae rela- 
tively longer, gubernaculum of different 

Sfeineria ampullacea new species 

Plate XXXII, fig. 72, c; Plate XXXIII, fig. 
72, a, b 

L = 1.44 mm; w = 68 [x; esophagus = 210 
/J.. Head diameter 24 ^. Lips round, with 
small labial papillae. Cephalic setae jointed, 
15 -I- 12 p. Subccphalic setae in eight 
groups, the sublateral ones with 5 setae, 
measuring 20 + 27 + 30 + 40 + 62 ^, the 
submedian ones with 4 setae. Many cervi- 
cal and somatic setae in eight longitudinal 
rows. Cuticle annulated, the annulcs re- 
solvable into dots. Amphids 10 /x in i , 16 
fi behind anterior end. Vestibulum of buc- 
cal cavity vertically striated which might 
be a characteristic separating this genus 
from Thcii.stus, in which the supporting 
structures of the vestibulum give the im- 
pression of a more oblique striation. Ex- 
cretory pore just posterior to nerve ring, the 
whole gland very short, its posterior end 30 
/ji anterior to the end of the esophagus. 
Spicula 55 // long, gubernaculum with 
curved apophysis, 23 /x long. Tail 175 /x 
long, a.b.d. 55 /x, terminal setae 35 fx. 

IloJoti/ix' specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4075, Type slide No. 101. Type locality, 
V, Vero Beach. 

Representation in samples ."itudied. — V, 
\^ero Beach. 

THERISTUS Bastian, 1865 
Type species.— Tfier/sfus ocer Bastian, 1865: 
156, 157, pi. 13, figs. 187, 188. 

Keys to the subgenera and their species 
are provided by Wieser, 1956. Species de- 
scribed subseqvient to this work are con- 
sidered in a more recent paper (Wieser, 

Subgenus Penzancia de Man, 1889 
Type species.— T/ier/sfus ve/ox Bastian, 
1865: 157, pi. 13, figs. 189-191. 

The species of this subgenus may be 
separated into three groups according to 
the presence or absence of a gubernaculum 
and to the shape of the gubernaculum, as 
follows : 

A) Gubernaculum with distal hook or 
triangular plate of characteristic shape 
( see Text- fig. 4 ) : 

T. hipunctatus (G. Schneider, 1906) 
(syn. Monhijstcm hipunctata); T. 
flevensis Schuurmans-Stekhoven, 1935 
(syn. Monhijstera velox Biitschli, 
1874, de Man, 1922 nee Bastian [see 
Gerlach, 1951c]); T. omhvonensis 
Schulz, 1935 (see Gerlach, 1951); T. 
paramhroncnsis Timm, 1952; T. 
macwflevensis Gerlach, 1953; T. 
metaflevensis Gerlach, 1955; T. horosi 
Andrassy, 1958; and T. calx n. sp. 

B ) Gubernaculum conical or blunt: 

T. inermis Gerlach, 1952; T. paivulus 
Timm, 1952 (doubtful species); T. 
hiarcospiculum Timm, 1952; T. tersus 
Gerlach, 1954; T. acrihus Gerlach, 
19.54; T. megalaimoicles Wieser, 1956; 
and T. stranus Gerlach, 1957. 

C) Gubernaculum absent: 

T. megalainui Stewart, 1914 (doubt- 
ful species); T. aeulcatus Schulz, 
1935; r. hetewseanicus Wieser, 1955; 
and T. hamatus Gerlach, 1956. 
All the other species listed by Wieser 
( 1956 ) are here considered doubtful. 

Group A, which was represented by two 
species in our material, is very homogeneous. 
The shape of the spicular apparatus (see 

Florida Marine Nematodes • Wiescr and Hopper 297 

Figure 4. Spicular apparatus of species of Ther;stus flevensis 
group. a — T. ombronensis (after Gerlach, 1951); b — T. 
flevensis (after Gerlacfi, 1951); c — T. bipunctatus (after 
Skwarra, 1924); d — T. macroflevensis (after Gerlach, 1953); 
e — r. borosi (after Andrassy, 1958); f — T. parambronensis 
(after Timm, 1952); g — T. metaflevensis (after Gerlacfi, 1955); 
fi — same species, present material; i — T. calx (present 

Text-fig. 4 ) immediately separates T. horosi 
and T. calx from all other species. As for 
the remaining species, it could be argued 
that T. flevensis, T. ambronensis, and T. 
bipunctatus represent three subspecies of 
T. flevensis, characterized by the position 
of the vulva ( 65-72% ) , the more posterior 
position of the amphids, and the slight 
subterminal protuberance at the inner edge 
of the gubernacular hook (shown by Gerlach 
in 1951, but not in 1957). On the other 
hand, T. metaflevensis and T. parambronen- 
sis have the vulva at 83-877^ and the guber- 
naculum ends distally either in a triangular 
plate or a smooth hook. However, the latter 

species is insufficiently described and the 
figure of the spicular apparatus is obviously 
rather stylized. T. macroflevensis is said 
to be characterized mainly by the long 
somatic setae. In other respects it links 
the flevensis-group with metaflevensis and 

We have identified our second species 
with T. metaflevensis although we saw the 
gubernaculum to be slightly different from 
that described by Gerlach and found only 
one lateral seta instead of three. It is im- 
possible at this time to judge the systematic 
value of these differences. 

metaflevensis Ger- 

Therisfus iPenzancia] 
lach, 1955 
Plate XXXIII, fig. 73 

Theristus (Peuzducia) metaflevensis Gerlach, 1955: 
291-293, fig. 25, a-tl 

L = 2 , 1.36, S , 1.8 mm; w = 34-40 fx- 
esophagus = 230-275 /x; Vu = 83%. Head 
diameter 2 , 19, S , 26 /i. Cephalic setae 9 
/x in female, 14 + 12 ^ in male, only the 
usual ten setae seen, that is, no additional 
lateral setae. Amphids in male 7 /m wide, 
19 fx behind anterior end, in female 6 fx 
and 13 fx, respectively. Spicula 53 fx long, 
typical. Gubernaculum rather large, distally 
with triangular plate in which a hook- 
shaped contour can be discerned. Tail in 
2 , 170 IX, in S , 240 /x long; a.b.d. in the 
latter 42 jx. 

Representation in samples studied. — M-4, 
Rickenbacker Causeway. 

Geograph ical distribution . — San Salvador, 
Brazil (Gerlach, 1955). 

Theristus (Penzancia) calx new species 
Plate XXXIII, fig. 74, a, b 

L = 3.25 mm; w = 70 /x; esophagus = 350 
fx. Head diameter 31 fx. Six setose labial 
papillae, 10 cephalic setae, 16 + 13 /x. Am- 
phids 7 fx = 20% of c.b.d. wide, 23 fx beliind 
anterior end. No cervical setae. Spicula 
118 fx long, of characteristic shape; the 
diagonal list that can be observed in all 
species of this group is here developed into 


Bulletin Mit.sciiin of Comparative Zoology, Vol 135, No. 5 

a lateral plate connecting proximal and 
distal end of the spiculum. The guber- 
naculum is reduced except for the triangular 
plate that also in this species is \er>' prom- 
inent. Tail 300 /x long; a.b.d. 60 /<.. 

Holotijpc specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4069, Type slide No. 102. Type locality, 
M-4, Rickenbacker Causeway. 

Representation in samples studied. — M-4, 
Rickenbacker Causeway. 

Subgenus Dapfonemo Cobb, 1920 
Type species.— Dapfonema {issertdens Cobb, 
1920: 281, 282, fig. 66a. 

In this subgenus there is a group of 
species distinguished by a characteristically 
shaped gubernaculum which gradually en- 
larges in its distal half, has a subterminal 
constriction and ends in a prominent hook 
(see PI. XXXIV, fig. 75, e). Previously this 
group consisted of the following closely 
related species: T. huetschJii Bresslau and 
Schuurmans-Stekhoven, 1940; T. huet.schU- 
oides Chitwood, 1951, and T. parobuetschlii 
Timm, 1961. To this group we add a 
fourth species, T. ostentator n. sp., which 
is separated from the other three by the 
inuch larger male amphids, the shorter 
spicula, the longer cephalic setae and some 
other minor characters. 

Theristus {Dopfonema) ostentafor new spe- 
Plate XXXIII, fig. 75, o, b; Plate XXXIV, 
fig. 75, c-e 

L = 1.76-1.77 mm; w = 30-32 ju; esopha- 
gus = .300-310 m Vu = 63%. Head diameter 
20-23 /x. Lips large, with strongly de- 
veloped framework (labial capsule). Labial 
setae 3 p.. Cephalic setae 20 + 1.5-16 fx, 
with additional lateral setae (1-3). Cervi- 
cal setae short. Amphids, in male, 15 /x = 
66/f of c.b.d. wide, in female, 8 /x and 30%, 
respectively. Cuticular annulation coarse. 
Buccal cavity spacious, with cuticularized 
walls. Spicula 38-42 /x long, cephalate prox- 

iniall\ . Gubernaculum 17-18 ix long, plate- 
shaped in its distal half, with subterminal 
constriction and terminal hook; there are 
also lateral projections. Tail in male, 130- 
155 /x long, a.b.d. 25-26 /x; in female, 175 
IX long, a.b.d. 26 /x. Tenninal setae, $ , 20 
/x long. 

Holotype specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4068, Type slide No. 103. Type locahty, 
M-3, Key Biscayne. 

Representation in samples studied. — M-3, 
Key Biscayne. 

Subgenus Trichotherisfus Wieser, 1956 
Type species.— S/e;ner/a mirabilis Schuur- 
mans-Stekhoven and De Coninck, 
1933: 10, 11, pl. 4, fig. 5; pi. 5, figs. 

Mesothcri.stus Wieser, 1956: 80, 91, NEW SYN- 

We consider it inadvisable to base sub- 
generic division entirely on differences in 
the length of somatic setae and we there- 
fore merge the two subgenera mentioned 
above. The important feature of this sub- 
genus is the occurrence of long somatic 
setae. These setae may be confined to the 
anterior region of the body but they should 
not be confused with the single circle of 
eight groups of elongated cervical setae 
about on level with the amphids, character- 
istic of the subgenus Fseudostcineria. 

Further classification of the subgenus 
may be based on the shape of the guber- 
naculum, the length and arrangement of 
the somatic setae, etc., as set out in the 
following key. 

Key to Species of Subgenus Trichotheristus 

1. Gubernaculum without apophysis 2 

Gubernaculum with apophysis 3 

2. Len^'th of spicula 12.5 m- Setae all over 

body -- T. loniiisc'tosus Schuurmans-Stek- 
hoven and De Coninck, 1933 
Length of spicula 64 /x. Setae in anterior 

half of cervical region only 

T. circumscriptus Wieser, 1959 

3. Apophysis large, set at an angle to the distal 

Florida Marine Nematodes • Wieser and Hopper 


shaft of the gubernacukim, pointing 

dorsocaudally 4 

Apophysis small, in direct continuation of 
distal shaft, pointing dorsally 8 

4. Somatic setae, at least in cervical region, 

measuring 2 c.b.d. or more 5 

Somatic setae not surpassing one c.b.d. in 
length 7 

5. Spicula strongly cephalate proximally, dis- 

tally with large lateral hook. Longest 

setae in mid-cervical region 

- -__ T. sanctimarteni Timm, 1957 

Spicula not cephalate, without lateral hook; 
longest setae in anterior cervical region , 6 

6. Distal half of spicula S-shaped. Additional 

circle of six subcephalic setae in i 

r. floridanus n. sp. 

Distal half of spicula more or less straight. 

No subcephalic setae T. mirahilis 

( Schuurmans-Stekhoven and De 
Coninck, 1933) 

7. Cephalic setae 23—26 m long, male amphids 

13 fi wide, apophysis of gubemaculum 

rod-shaped T. laxus Wieser, 1956 

Cephalic setae 13-16 fj., amphids 5-8 /j., 
apophysis of gubemaculum plate-shaped .. 

T. setosus (Butschh, 1874); 

T. hirtus Gerlach, 1951 
(for differences see Gerlach, 1951) 

8. Somatic setae, at least in cervical region, 

measuring 1.5 c.b.d. or more 9 

Somatic setae not surpassing one c.b.d. in 
length T. erecius n. sp. 

9. Amphids distinctly spiral. Cephalic capsule 

well developed. One circle of four sub- 
cephalic setae in male .. _ T. galeatus n. sp. 

Amphids circular. No cephalic capsule. No 

subcephalic setae 10 

10. Lateral cephalic seta in c? elongated. Am- 
phids 0.5 head diameters behind anterior 
end T. heterus Gerlach, 1957 

Lateral cephalic seta not particularly elon- 
gated. Amphids 1.5 head diameters be- 
hind anterior end 

T. setifer Gerlach, 1952 

Therisfus (Trichotheristus) floridanus new 
Plate XXXIV, fig. 76, a-d 

L = 1.50-1.57 mm; \v = 50-60 ^; esopha- 
gus = 325-350 fx; Vu = 659f . Head diameter 
20-23 /JL. Lips round, labial papillae setose, 
short. Head with 12 cephalic setae, 20 + 16 
jx long; in 6 there is an additional circle of 
6 subcephalic setae. Long and short somatic 
setae, the longest ones measuring 60-70 ^ 
which is nearly 3 c.b.d. in the cervical re- 

gion, about 1.2 c.b.d. in mid-body. Amphids 
in <i , 7 /x, in 9 , 6 /j. wide, 18-23 /a behind 
anterior end. Spicula 37 p. long, distal half 
S-shaped. Gubemaculum complicated, with 
plate-shaped apophysis, 10 /x long. Tail in 
6 225 iJL long = 5 a.b.d., in 9 250 ix long = 
5.5 a.b.d. Terminal setae 60 /x long. 

Holotijpe specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4067, Type shde No. 104. Type locality, 
M-2, Key Biscayne. 

Representation in samples studied. — M-2, 
Key Biscayne. 

Therisfus [Trichotheristus) erecfus new spe- 
Plate XXXIV, fig. 77, b; Plate XXXV, fig. 
77, a, c, d 

L = 1.3 mm; w = 55 ^a; esophagus = 275 
fx. Head diameter 16-24 /x. Lips round, 
labial papillae setose, short. Cephalic setae 
10-17 + S-15 fx long. Short and long 
somatic setae, the longest ones measuring 
34 fx which is about one body diameter in 
the cervical region. There are four char- 
acteristic pairs of sublateral setae behind 
the amphids. Amphids in A 6-7 jx = about 
23% of c.b.d. in width, 16-22 ^x behind 
anterior end. Cuticle with lateral alae 
( incisures? ) which occur, for the most part, 
as t\\o parallel refractive lines 7-8 /x apart 
(about ^7— s c.b.d.). In the posterior neck 
and preanal regions, there frequently occurs 
a third refracti\e line between those nor- 
mally present. Faint cuticular striations 
can be traced over the lateral alae. Spicula 
35-38 fi long, proximal end cephalate, distal 
half slightly S-shaped. distal end with a 
lateral tooth. Gubemaculum 23 ^, with 
small, plate-shaped dorsal apophysis. Tail 
200-210 ,x long, a.b.d. 40 /x. 

Holotijpe specimen. — Male; Canadian Na- 
tional Collection of Nematodes, Entomology 
Research Institute, Ottawa, Collection Num- 
ber 4067, Type slide No. 105. Type locality, 
M-2. Ke\- Biscayne. 

300 Bulletin Mimtim of Comparative Zoology, Vol. 135, No. 5 

Representation in samples studied.— M-2, time. We follow Wieser (1956, 1959) and, 

Kev Biscayne, M-3, Key Biscayne, and M-4, on the basis of the shape of the guber- 

Rickenbacker Causeway. ' naculum, distinguish three groups of species, 

Remarks.— The closest relative of this viz., A) gubernaculum without apophysis, 

species is T. sefosus from which it can be B) with small dorsal apophysis, C) with 

separated by the small and dorsally directed distinct caudal apophysis. No new species 

apophysis of the gubernaculum. have been described since Wieser's paper 

of 1959, but probably more of the species 

T, • . .T • L xL • . \ ; i r^^ included in the grouping by Wieser have 

Thenstus {Tnchofhensfus) galeatus new spe- , . , ? ^ '=' . -^ . . ^ ,^^ 

^ to be considered as species inqimendae, 

ni *^'^^x/N,x/ L- -TO e.g., T. lon^ieaudatus Filipjev, 1922, and 

Pate XXXV, fig. 78, a-c „^ ' . ,. ^ ^,, iqqa 

^ T. naviculworus Cobb, 1930. 

L = 1.02 mm; vv = 32 p.; esophagus = 205 Group A, species without gubernacular 
p.. Head diameter 15 p.. Labial papillae apophysis, contains a number of species in 
setose, short. Cephalic setae jointed, 10 in x\'hich the spicula in their distal fifth or 
number, 15 + 12 /x long. Male with 4 sub- sixth show a very characteristic outward 
cephalic setae. Short and long somatic bend. This additional curvature of the 
setae, the longest ones measuring 65 /x which spicula appears to give, as it were, more 
is about three times the c.b.d. in the cervical room to the development of the distal por- 
region. Cuticular annulation coarse, one i-jon of the gubernaculum which in these 
annule 1.4 /x wide. The head seems to be species shows some complex features that 
strengthened by a cephalic capsule which a,-e difficult to analyze. The situation is 
consists of two portions, an anterior and illustrated by figures 79, c, and 80, c, d on 
a posterior one. Amphids spiral, 6 x 7 /x, plate XXXVI in this paper. To this sub- 
32-36 /x behind anterior end. Spicula 36 /x group belong with certainty T. kornocensis 
long, cephalate proximally, distally with a (Allgen, 1929) sensu Wieser, 1959, T. 
lateral tooth. Gubernaculum with dorsal oxyuroidcs ( Schuurmans-Stekhoven, 1931) 
apophysis and a distal three-pronged piece, and T. fistulatus n. sp., but it cannot be 
In the postanal region there are indications excluded that some more species described 
of 'breaks' in the cuticular annulation. j^ the literature show the same differentia- 
These breaks, however, seem to be rather t^on of the spicular apparatus. T. trecuspi- 
irregular and are differently arranged in datus Wieser, 1959, displays the same char- 
different specimens. Tail 170 /x long, a.b.d. acteristic but, by possessing a small dorsal 
36 /x. apophysis of the gubernaculum, has been 

Holotype specimen. — Male; Canadian Na- assigned to the next group, 

tional Collection of Nematodes, Entomology Group B of this subgenus, comprising 

Research Institute, Ottawa, Collection Num- species with small dorsal gubeniacular 

ber 4070, Type sHde No. 106. Type locality, apophysis, is represented in our material 

M-5, Virginia Key. by two well characterized new species, T. 

Representation in samples .studied. — M-5, toifus n. sp., and T. xyaliformis n. sp. 
Virginia Key. 

Theristus {Cylindrofheristus) oxyuroides 

Subgenus Cy//ndroffiensfus Wieser, 1956 (Schuurmans-Stekhoven, 1931) 

Type species.— Mono/iysfera normandica de , vvv\/i i- to 

.. in/->rt iJLr, 1-71 1 o I- 1 ij Pate XXXVI, tig. /V, a-c 

Man, 1890: 169-171, pi. 3, figs. 1-ld. ^ 

rri . 1 4- 4.1 ,r .4-,...4- ,..-,^K Monkt/stera oxi/uroidcs Schuurmans-Stekhoven, 

This subgenus presents the greatest piob- ^yg^/g^^ 656, fig. 8, a-c. 

lems of classification, and, due to the in- 
sufficiency of many descriptions, no satis- L = 0.9 mm; w = 43 p.; esophagus = 140 
factory treatment is possible at the present p. Head diameter 14 p. Lips round, with 

Florida Marine Nematodes • Wieser and Hopper 301 

short setose papillae. Twelve subequal 
cephalic setae, 9 fj, long. Rows of short 
cer\ical and somatic setae. Amphids 7 fi 
wide, 12 IX behind anterior end. Spicula 28 
fx long, shaipK' bent in middle and with an 
additional uutnard curvature in its distal 
sixth. Gubeniaculum sleeve-like, without 
apophysis, distal half with curved pieces. 
Tail 160 IX long, a.b.d. 29 /x. 

Representation in samples studied. — M-4, 
Rickenbacker Causeway. 

Geograplueal distribution. — Baltic, North 
Sea, Zuiderzee; Chesapeake Bay, Marvland 
(Timm, 1952). 

Remarks. — Our material contained what 
we consider to be fairly typical representa- 
tives of T. oxyuroides ( Schuurmans-Stek- 
hoven, 1931). The characteristic distal 
curvature of the spicula is apparent in 
Schuurmans-Stekhoven's original descrip- 
tion, but both Gerlach (1951c) and Timm 
(1952), as far as one can infer from their 
figures, seem to hold that it is the guber- 
naculum and not the spicula that is curved 
distally. T. kornoeensis (Allgen, 1929) and 
T. fistulatus n. sp. can be separated from 
T. oxyuroides by the more backward posi- 
tion of the amphids and by differences in 
the finer structure of the spicular appara- 

Theristus {Cylindrofherisfus) fistulatus new 
Plate XXXVI, fig. 80, a-d 

L = 0.86-1.04 mm; w = 30-.36 p.; esopha- 
gus = 144-180 IX. Head diameter 15 /x. Lips 
round, bearing short setose papillae. Ten 
to 12 cephalic setae 13-1-11 ix long. Scat- 
tered cervical and somatic setae. Amphids 
8-9 IX = 40-45 per cent of c.b.d. wide, IS- 
IS IX behind anterior end. Spicula 25-27 
IX long, cephalate proximall\% outwardly 
curved in its distal sixth. Gubeniaculum 
sleeve-like, without apophysis, distalh' with 
curved, tubular piece. Tail 210-228 ^ long, 
a.b.d. 2.3-26 ix. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 

mology Research Institute, Ottawa, Collec- 
tion Number 4068, Type sHde No. 107. 
Type locality, M-3, Key Biscayne. 

Representation in samples studied. — M- 
3, Key Biscayne. 

Remarks. — T. fistulatus is characterized 
by the tubular element in the distal portion 
of the gubeniaculum, whereas T. kornoeensis 
is equipped with two lateral projections at 
the distal tip of the gubemaculum. 

Theristus {Cylindrofherisfus) tortus new spe- 
Plate XXXVI, fig. 81, a-d 

L = 0.74-0.76 mm; w = 25-30 ix; esopha- 
gus = 168-192 IX. Head diameter 10-12 ix. 
Lips round, bearing short setose papillae. 
Ten to 12 cephalic setae, 10-12 + 7-9 /x 
long. No cervical or somatic setae. Am- 
phids in 9 , 8 ix = 47% of c.b.d. wide, 24 ix 
behind anterior end, in ^ , 10 /x = 60% of 
c.b.d. wide, 23 ix behind anterior end. 
Thirteen to 15 cuticular annules between 
anterior end and amphids. Spicula slender, 
of even thickness, 27 ^ long, cephalate 
proximally. Gubeniaculum 15-16 ^ long, 
^^'ith short dorsal apophysis, oblique grooves 
in distal half and two conspicuous, laterally 
pointing projections at its distal end. Tail 
in 9 , 175 /x, in 6 , 132-156 ^ long, a.b.d. 

Holotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 108. 
Type locality, M-2, Key Biscayne. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Remarks. — T. tortus n. sp. is distinguish- 
able from all other species of this group by 
the large and posteriorly situated amphids, 
and by the shape of the gubeniaculum. Its 
closest relative is T. resimus Wieser, 1959, 
which has a similar gubeniaculum, although 
without the oblique grooves that occupy 
the anterior part of the gubeniaculum in 
T. tortus. 

302 B 

ulletin Museum of Comparative Zoology, Vol 135, No. 5 

Therisfus {Cylindrofherisfus) xyaliformis new 
Plate XXXVI, fig. 82, a-d 

L = 0.6:3-0.73 mm; w = 21-22 /x; esopha- 
gus = 125-150 iJ. Head round, diameter 
6 /x. Labial papillae indistinet. Ten ce- 
phalic setae, subequal, 5 /x long. No^erv^ 
cal and somatic setae. Amphids 5 ^ - 507^ 
of c.b.d. wide, 20-22 /j. behind anterior end, 
about 30 cuticular annules between ante- 
rior end and amphids. Buccal cavity 
unusuallv deep bv enlargement of the "buc- 
cal ring." Spicula 19-20 ,x long, cephalate 
proximally. Gubernaculum distally with 
small lateral projection, proximally with 
hook-shaped dorsally pointing apophysis. 
Tail 127-145 /x long, a.b.d. 16-19 /x. 

Ilolotype specimen. — Male; Canadian 
National Collection of Nematodes, Ento- 
mology Research Institute, Ottawa, Collec- 
tion Number 4067, Type slide No. 109. 
Type locality, M-2, Key Biscayne. 

Repvesentotion in samples studied. — M- 
2, Key Biscayne. 

Remarks.— In T. xyaliformis n. sp. the 
buccal cavity differs from that of all other 
species of Theristus by its elongation. The 
elongation has come alSont by the widening 
of the "buccal ring" which is in fact the 
prostome (in the sense of Osche, 1952), 
i.e., that part of the buccal cavity that lies 
between the base of the lips and the an- 
terior end of the esophagus. Since the same 
type of elongation is a characteristic feature 
of the subfamily Xyalinae, T. xyaliformis 
can be considered to link this subfamily 
with the Monhysterinae. Moreover, T. 
xyaliformis is characterized by the far 
posteriorly situated amphids and by the 
hook-shaped apophysis of the gubernacu- 

MONHYSTERA Bastion, 1865 
Type species.— Mon/iysfera stagnalis Bas- 
tian, 1865: 97, pi. 9, figs. 9-11. 

The proper status of our species is im- 
possible to determine. It is undoubtedly 
closely related to M. parva (Bastian) but 

the differences of opinion concerning this 
species cannot be resolved at present. De 
Man (1888) figures the spicula of M. parva 
without any teeth or projections but sub- 
sequent authors all show the spicula in 
their proximal third or fourth to be 
ecjuipped with a ventral projection (actu- 
ally the point of attachment of the muscu- 
lar "velum"). De Coninck and Schuurmans- 
Stekhoven (1933) claim that de Man 
overlooked this projection and consider M. 
heteroparva Micoletzky, 1924, to be a 
synonym. Timm ( 1952 ) rejected this claim 
and revived M. heteroparva as a valid 
species, to be separated from M. parva by 
the possession of this ventral tooth. An- 
drassy ( 1958 ) figured the spicular appara- 
tus of M. parva not only with the proximal 
projection but also with a distal "nose" and 
hook. Our species closely agrees with 
Andrassy s description and figure and we 
tend to think that indeed many authors 
overlooked, or misinteipreted, either the 
proximal or the distal projection, or both, 
and that M. parva is a cosmopolitan species 
which needs to be restudied in its type 
habitat. We observed faint preanal supple- 
ments but again have our doubts whether 
this can be considered a real difference 
from previous descriptions. 

Monhystero porva (Bastian, 1865) 
Plate XXXVII, fig. 83, a-d 

TachijhocUtcs parvus Bastian, 1865: 156, pi. 13, 
fifts. 185, 186; Monhystcra parva var. meridUina 
Micoletzky, 1922; M. heteroparva Micoletzky, 
1924; M. kossncmis Paramonon, 1929; M. ant- 
arctica Col^b, 1914. 

L = 0.58-0.72 mm; w = 18-22 /x; esopha- 
gus = 102-115 /x; Vu = 65%. Head diam- 
eter 8-10 /x. No labial papillae seen. Ten 
cephalic setae, 4 + 3 /x long. Amphids in 
6 , 3.5 IX = 36% of c.b.d. wide, 10 ^ behind 
anterior end. Two pairs of submedian 
cervical setae, 25 /x behind anterior end, 
excretory pore 37 /x, nerve ring 63 /x behind 
anterior end. Spicula 26-27 /x long, with 
proximal "handle" and projection on which 
the velum attaches, distally with a triangu- 

Florida Marine Nematodes • Wieser and Hopper 


lar cuticularized piece which forms a sort 
of recurved hook. Gubernacular apophysis 
13 /ji. About 15 indistinct preanal and two 
postanal supplements. In some specimens 
these supplements are hardly visible and 
appear only as darker and lighter portions 
of the cuticle. Tail 95-100 /x long, a.b.d. 
16-18 IX. 
Representation in samples studied. — M- 

1, Key Biscayne. 

Gcoiiraphical distribution. — Cosmopoli- 


SCAPTRELLA Cobb, 1917 

Type species.— Scapfrella cincfa Cobb, 

1917: 119, fig. 4. 
Scaptrella cincta Cobb, 1917 
Plate XXXVII, fig. 84, a-c 

Scaptrella cincta Cobb, 1917: 119, fig. 4. 

L = 1.7 mm; w = 41 /x; diameter at base 
of esophagus 33 fx. Esophagus 220 /x. Tail 
360 IX. Head diameter 20 ix. Labial setae 
6-7 IX long. Cephalic setae 12 ( 6 + 6 ) , the 
lateral pairs 30 + 18 ^, the submedian 50 + 
30 IX long. Amphid circular, \\ith internal 
spiral, 4 , 10 fi, $ , 8 ^u, wide. Buccal cavity 
cylindrical, 30 ix deep. Anteriorly armed 
with 6 jointed odontia. In addition, a blunt, 
weakly-sclerotized, dorsally-positioned on- 
chium is present. Cuticle coarsely striated, 
about 3 n wide in neck region and about 2 
IX in mid-body. Somatic setae numerous, 
very fine, averaging 17 /x long on most of 
the body length. Tail with numerous 
caudal setae and a pair of 20 /j. long termi- 
nal setae. Spicula 34 /x long, proximal!) 
cephalated. Gubernaculum with bidentated 
lateral guiding pieces. 

Representation in samples studied. — M- 

2, Key Biscayne. 

Geofirapliical distribution. — Atlantic 
Coast from Massachusetts to North Caro- 
lina (according to Chitwood, 1951), Per- 
nambuco, Brazil ( Gerlach, 1956 ) . 

Remarks. — Scaptrella eincta is separated 
from S. brevicaudata Gerlach, 1952, bv the 

longer and more filiform tail and by the 
longer cephalic setae. 

XENOLAIMUS Cobb, 1920 

Type species.— Xeno/a/mus striafus Cobb, 

1920: 250, 251, fig. 30. 
Xenolaimus striafus Cobb, 1920 

Plate XXXVII, fig. 85, a-d 
Xenolaimus striatus Cobb, 1920: 250, 251, fig. 30. 

L = 1.06 mm; w = 33 /x; esophagus = 300 
IX. Head diameter 16-17 ^. Lips 7 /x high, 
flap-like. Labial setae 5.5 ix long. Ten 
cephalic setae, the longer ones measuring 
19 /x; the two setae of each submedian pair 
stick together. Buccal cavity wide and 
deep, with two weakly cuticularized teeth 
or cuticular folds projecting from the base 
of the mouth to the base of the lips. Am- 
phids in an enlarged portion of the fifth 
cuticular annule. This enlargement about 
6 IX. First cuticular annule wider than the 
follo\\'ing ones. The head, in the words of 
Cobb (1920: 250), "is protrusile and ap- 
pears as if surrounded by a balustrade 
composed of the anterior annules of the 
cervical cuticle." Cuticular annulation 
coarse, with an unspecified number ( 12? ) 
of longitudinal rows of V-shaped struc- 
tures. Spicula asymmetrical, 28-30 + 32- 
34 /x long. Apophyses of gubemacula also 
asymmetrical, 17 + 21 ^ long. Tail conical, 
145 IX long, a.b.d. 27 /x. 

Representation in samples studied. — M- 
2, Key Biscayne. 

Geographical distribution. — Biscavne 
Bay, Florida (Cobb, 1920). 

Remarks. — The present record represents 
the first time this species has been found 
since Cobb's original description. An un- 
identified example of Xenolaimus was re- 
corded in a list of nematodes from the Gulf 
Coast of Florida by King, 1962 (Xenolaimus 


The collecting trip during the period 
from May 7 to June 8, 1963, was made 

304 Bulletin Museum of Comparative Zoology, Vol. 135. No. 5 

possible through NSF grant GB 498 to one 
of us ( WAV. ) . In this connection, we wish 
to thank Professor Ernst Mayr and Dr. 
Giles Mead of the Museum of Comparative 
Zoology of Harvard University for their 
support and encouragement. Particular 
thanks are due to Mr. R. H. Mulvey, Chief, 
Nematology Section, Entomology Research 
Institute, Canada Department of Agricul- 
ture, Ottawa, in whose Section the slides of 
our collections were mounted and who 
supported our project in many other ways. 
We wish to thank Mr. Curtis W. Sabrosky, 
U.S.D.A., \\'ashington, for making avail- 
able for study several type specimens from 
the United States National Museum. We 
are grateful to Dr. S. P. Meyers, of the In- 
stitute of Marine Science, University of 
Miami, Miami, Florida, who arranged for 
our stay at the Institute and, in addition, 
provided valuable assistance. Miss Nancy 
Van Meter served as an efficient guide to 
collecting sites in the Everglades National 

Many more people were very helpful to 
us during our trip and we shall acknowl- 
edge their hospitality and cooperation as 
we proceed with publication of our reports. 


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. 1936. Some marine nematodes from 

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