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BULLETIN
OF THE
Museum of Comparative Zoology
HARVARD UNIVERSITY
VOL. 133
CAMBRIDGE, MASS. U.S.A.
1965-1966
The Cosmos Press, Inc.
Cambridge. Mass., U.S.A.
CONTENTS
PAGE
No. 1. — GuLAR Musculature in Delphinids. By Barbara
Lawrence and William E. Schevill. May, 1965 . 1
No. 2. — A Revision of the Genus Rhabdephyris in the
Americas (Hymenoptera, Bethylidae). By
Howard E. Evans. May, 1965 67
No. 3. — Nclda.mnrus wrightac, a New Rhachitomous
Labyrinthodont from the Texas Lower Per-
mian. By John Newland Chase. June, 1965 . . 153
No. 4. — The Genera of the Chilocorini (Coleoptera,
Coccinellidae). By Edward A. Chapin. Septem-
ber, 1965 ..." 227
No. 5. — Comments on some Recent Changes in the Clas-
sification OF THE Ciidae ( Coleoptera) . By John
F. Lawrence. October, 1965 273
No. 6. — The Fossil Elephant Shrews (Family Macro-
SCElididae). By Bryan Patterson. November, 1965 295
No. 7. — ^Panamanian Spiders of the Genus T mar us
(Araneae, Thomisidae). By Arthur M. Chicker-
ing. November, 1965 337
No. 8. — The Relationships of Four Small Hispaniolan
Elcuthcrodactyliis (Lei^todactylidae). By Albert
Schwartz. January, 1966 369
PAGE
No. 9. — Two New Fishes of the Myctophid Genus Di-
aphus FROM the Atlantic Ocean. By Basil Naf-
paktitis. January, 1966 401
No. 10. — The Ameiva (Lacertilia, Teiidae) of Hispani-
OLA. II. Geographic A^ariation in Ameiva chryso-
laema Cope. By Albert Schwartz and Ronald F.
Klinikowski. March, 1963 425
No. 11. — A New Attempt to Construct Life Tables for
Kent Island Herring Gulls. By Raymond A.
Paynter, Jr. May, 1966 ..... \ ... 489
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 1
GULAR MUSCULATURE IN DELPHINIDS
By Barbara Lawrence and William E. Schevill
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Bulletin of the Museum of Comparative Zoology
HAEVAED UNIVEESITY
Vol. 133, No. 1
GULAR MUSCULATURE IN DELPHINIDS
By Barbara Lawrence and William E. Schevill
CAMBBIDGE, MASS., U.S.A.
FEINTED FOE THE MUSEUM
May, 1965
Bull. Mus. Comp. Zool., Harvard Univ. 133(1) :l-65, May, 1965
No. 1 — Gular Musculature in Delphinids^
By Barbara Lawrence and William E. Sciievill
TABLE OF CONTENTS
PAGE
Introduction 5
General account of Delphinidae 6
Superficial Layers 7
M. sphincter colli profundus 8
M. sphincter colli primitivus 8
M. auriculolabialis 8
M. orbicularis oculi 9
M. orbicularis oris 9
M. nasolabialis 9
Ear muscles 10
Muscles of the Neck and Throat 11
Interramal and Tongue Muscles 12
Mylohyoid muscle 12
Digastric muscle 12
Geniohyoid muscle 13
M. styloglossus 14
M. hyoglossus 14
M. genioglossus 14
M. palatoglossus IG
Hyoideal Muscles 17
Sternohyoid muscle 17
Sternothyroid muscle 17
Thyrohyoid muscle 18
Occipitohyoid muscle 18
Interhyoid muscle 18
M. hyoepiglotticus 19
Muscles of the Pharynx 19
M. stylopharyngeus 21
M. palatopharyngeus 22
M. pterygopharyngeus 24
Occipitothyroid muscle 25
M. thyreopharyngeus 26
Homologies with Phocoena 26
Discussion 29
1 Contribution No. 14.37 from the Woods Hole Oceanosraphic lustitution
4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Globicephala melaena 50
Muscles of the Xeck and Throat 50
Sternomastoid muscle 50
Mastohumeralis muscle 50
Scalenus muscle 50
Mylohyoid muscle 51
Digastric muscle 51
Geniohyoid muscle 51
M. styloglossus 52
M. hyoglossus 52
M. genioglossus 52
M. palatoglossus 53
Sternohyoid muscle 53
Sternothyroid muscle 53
Thyrohyoid muscle 53
Occipitohyoid muscle 53
Interhyoid muscle 54
Muscles of the Pharynx 54
M. stylopharyngeus 55
M. palatopharyngeus 55
M. pterygopharyngeus 56
Occipitothyroid muscle 56
M. thyreopharyngeus 57
Conclusion 57
Acknowledgments 58
Literature Cited 58
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 5
GULAR MUSCULATURE IN DELPHINIDS
INTRODUCTION
A scattering of authors in the past ninety years have pub-
lished on different aspects of the head and throat musculature
in the odontocetes. No two have gone about their dissections in
the same way, the names used for the muscles vary, and in some
instances the inadequacy of their material has led to conflicting
statements on the number and position of the muscles involved.
Our investigations of sound production in the larynx have
pointed up the need for a series of more detailed dissections of
the entire region, as well as for correlating the descriptions of
some of these early workers with each other and with the condi-
tions that we actually found.
Our primary concern has been to establish the true relation-
ships of the hyo-laryngeo-pharyngeal muscles or, to put it differ-
ently, to study the muscles which lie between the throat car-
tilages and the base of the cranium, medial to the basioccipital
plates and posterior to the palate. Whether or not the great
development of the basioccipital plates is related to the peculiar
structures associated with the ear bones or to the evolution of
an intranarial larynx, the combined result has been a character-
istic arrangement of the muscles in this region which is com-
mon to many odontocetes. Since accurate dissection of these
deeper layers depends on an understanding of the more super-
ficial ones, the latter also have been figured and described.
The muscles external to the pterygoid plates are quite distinct
from those of the throat region, and so are omitted. The
boundary between these two regions is a tough, fibrous mem-
brane which extends from the tip of the pterygoid hamulus to
the tissue investing the bulla on each side, and lies across the
notch in the pterygoid plates. It separates the muscles of the
bony nares from the pterygoid muscles. The complex of air
sinuses and retia external to this has been well described by
Fraser and Purves (1960a, pp. 65-68).
The first section of this paper deals with four rather closely
related genera represented by the following species : Delphinus
delphis Linne 1758, Stenella plagiodon (Cope) 1866 and -S'. styx
(Gray) 1846, Tursiops truncatus (Montagu) 1821, and Lagcno-
rhynchus acutus (Gray) 1828 andL. alhirostris Gray 1846. Herein
they will usually be referred to by generic name only. The pur-
pose of this section has been not to compare the four genera, but
6 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
to establish the common delphiuid arrangement of muscles in
this region. For this reason, the dissections have rarely dupli-
cated each other. ]\Iost of the figures are of Dclphinus, but to
shoAV certain details some figures of Tursiops and some of
Lagenorliynchus are included. The specimens studied are those
reported on in the "Functional anatomy of the delphinid nose"
(Lawrence and Schevill, 1956), and include, in addition, a
specimen of Lagenorhynchiis alhirostris and one of Stenella
styx, as well as a representative of the Phocoenidae, PJiococna
pliocoenn (Linne) 17o8, all collected in the western North
Atlantic.
The second section is a clarification of Murie's classic de-
scription of the throat region in Glohicephala mclacna (Traill)
1809. Although also a delphinid, this is a much larger form
which differs further from the above four genera in its relatively
shorter rostrum and broader skull.
In the following, for ease of description, the muscles consid-
ered are grouped and discussed in three divisions : the superficial
layers, the muscles of the neck and throat, and the pharyngeal
muscles.
GENERAL ACCOUNT OF DELPHINIDAE
In the hope of standardizing muscle names in the cetaceans,
we have not followed one author but have selected what seem
to us the most appropriate terms. Huber's terminology (1934,
pp. 117-119) has been used for the superficial layers. For the
hyo-laryngeo-pharyngeal region, the names are chiefly from
Murie (1873), Kernan and Schulte (1918), and Howeir(1927).
When the names used are Latin adjectival forms, the word
"musculus" is to be understood as preceding the name; when
the adjectives are anglicized, the English word "muscle" may
be understood to follow the name.
In order to avoid confusion between the bones and similarly
named muscles, we have followed Howell (1927) and earlier
authors in using the "-hyal" ending for different components
of the hyoid apparatus, as for instance thyrohyal, while in these
cases we have used "-oid" for muscles, as for instance ihyro-
hyoid. Since the thyro- and basihyal elements fuse early, it is
not possible to tell whether certain muscles attach on both or
on only one or the other. This is an unimportant detail ; never-
theless, it should be remembered that in some instances what
we call a thyrohyal attachment may reallv be a thvrobasihval
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE (
attachment and vice versa. The current names for the laryngeal
cartilages are so universally accepted that the standard forms
thyroid, cricoid, and arytenoid have been used. This is less con-
fusing than in the case of the hyoid apparatus, for in the larynx
parallel names for muscles are not in common use.
To make clear the interrelationship of the muscles as well as
to compare better our findings with those of other workers, the
material was actually dissected in a number of different ways.
The sequence in which the muscles are presented here is chosen
as the easiest for purposes of identification, as they are exposed
in progressively deeper ventrolateral dissection. Additional de-
tails of the more complicated muscles based on other types of
dissection are also illustrated. These are essential if function as
well as identification is to be understood.
Aside from Murie's work on Glohicephala, the most useful of
the earlier dissections of this region in odontocetes have been
made b,y: Schulte and Smith (1918), and Kernan and Schulte
(1918) of Kogia; by A. B. Howell of Neomeris (1927), and
Boenninghaus of Phocoena (1902) ; by Fraser and Purves of
Delphinus (1960a and b) ; and by Hein (1914) of Monodon.
Where our dissections overlap and their findings differ sig-
nificantly from ours, these are discussed ; otherwise their work
is merely noted.
Superficial Layers
Huber's (1931:, pp. 117-120, fig. 4) detailed description of the
facial muscles of Tursiops, published posthumously, is an in-
tegral part of his carefully reasoned series on the evolution of
facial musculature. In another paper (Lawrence and Sehevill,
1956), we have discussed his account of the blowhole muscula-
ture in the light of our own findings. His description of the
superficial facial muscles matched closely what we observed in
the four genera dissected, and the following comments are
mainly supplementary. These outer layers have been considered
by us only because of their relation to the deeper hyopharyn-
geal regions which are our primary concern.
The blubber in the region of the head and throat is closely
bound to the underlying muscle. Some of the outermost layers
of the sphincter colli actually insert in the dense inner layers
of the blubber. Ventral to the eye and surrounded by the
sphincter colli profundus, auriculolabialis, and orhicidaris oris,
there is an extensive area of fat lying directly beneath the
8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
blubber and against the jaw. The remnant of the outer ear lies
buried in this. Distally, this vestige is a very slender tube sur-
rounded by dense fibrous tissue and embedded in fat. Prox-
imally, the tube widens and is partially enclosed by small, curved
cartilages of a characteristic shape.
31. sphincter colli profundus (Fig. 1, spf). This extends in
a thin, transverse sheet from the symphysis of the jaw to the
region behind the flipper, beyond which it was not dissected.
This muscle was studied only in Delphinus and Lagenorhyn-
chus acutus. In these, an aponeurotic area occurs on the throat
over the sternohyoid. Posterior to this, especially laterally
between the forelimb and the ear, the sphincter colli profundus
is thicker. Along each side, at about the level of the eye, it meets
the sphincter colli primitivus. Dorsal to the forelimb, from the
line where these two meet, a small bundle of fibers converges
to insert in the flipper. Anterior to this, a very few fibers
pass ventrally to attach to the fibrous mass surrounding the
auditory tube, and deep to these a few other fibers merge with
those of the auriculolahialis.
M. sphincter colli primitivus (Fig. 2, spt). This lies dorsal
to the sphincter colli profundus and is very difficult to dis-
tinguish from it along the line where the two meet. Only a slight
difference in the direction of the fibers suggests that, instead
of a single thin sheet of muscle wrapping around the side of
the face and throat, there are in reality two muscles. A small,
doubtfully distinct, anterior segment (Fig. 2) attaches in the
dense fibrous tissue investing the slender tube which passes
from the minute orifice of the outer ear to the vestigial ear
cartilages. This may be what Huber (1934, p. 117) refers to as
"a small vestige of the extrinsic postauricular musculature."
This is the only part of sphincter colli primitivus figured.
M. auriculolahialis (Figs. 1, 2, al). This arises mainly along
the rim of the zygomatic process of the squamosal external to
the glenoid fossa and posteroventral to the root of the zygo-
matic arch. Posteriorly, it is partly overlain by the ear cart-
ilage, and a small part of the muscle takes origin in the tissue
here. From its origin, auriculolahialis passes anteriorly, ventral
to orbicularis oculi, diminishing rapidly in size, to insert as
described later. The primary squamosal origin is fairly con-
stant in all the forms dissected, but the origin of the thin layer
which is associated with the vestiges of the outer ear varies.
In Delphinus this is in the fibrous mass surrounding the auditory
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 9
tube, distal to the cartilage; in Stenella plagiodon it is dif-
fusedly iu the tissue investing the ear cartilage itself, while in
Lagenorhy nchus acutus there is no apparent connection with
the cartilage, but the attachment is in the connective tissue
mass between cartilage and skull. The relation of the insertion
of auric ulolabialis to orbicularis oculi, the zygomatic arch, and
the connective tissue beneath the latter varies. In Delphinus
and Stenella plagiodon we found auriculolahialis inserting in
part on the arch itself, while in Lagenorhy nchus acutus the in-
sertion was in the connective tissue ventral to the arch. In all
three there was a certain connection with orbicularis oculi, very
slight in the last genus, more definite in the two former. In
Tursiojjs, Huber says that auriculolahialis passes from the pre-
auricular region to fuse with orbicularis oculi and gives no other
insertion for it. Possibly all of this is individual, not generic,
variation and an indication that auriculolabialis in delphinids
is degenerating. This muscle is the tympano-zygomaticus of
Howell (1927, p. 22).
M. orbicularis oculi (Fig. 2, oc). AVe dissected this in detail
only in Stenella plagiodon. Here, as in Huber 's Tursiops, it
forms a well-developed sphincter around the eye. A few of the
outer fibers anteriorly and posteriorly attach to the margin
of the orbit, while anteroventrally some pass downward into
fibrous fatty tissue between the eye and orbicularis oris.
M. orbicularis oris (Fig. 2, or). This is a weak and very fat
muscle whose fibers radiate from the connective tissue near the
corner of the mouth and end in the fibrous inner layer of blub-
ber.
M. nasolabialis (not figured). The muscles between the eye
and the melon were dissected in detail only in Stenella plagio-
don and did not match Huber 's figure of Tursiops (1934, fig.
4A, p. 118). In this region, he shows a single rather extensive
nasolabialis continuous with the anteriormost fibers of orbicu-
laris oculi. The situation in Stenella was a little different;
here we found two small sheets of fibers which were separated
from the orbicularis oculi by the maxillary bone. The more pos-
terior took origin on a narrow area on the maxillary, dorsal to
the anterior part of the eye, and passed anteriorly to end in the
fibrous mass beneath the melon. The more anterior sheet arose
in front of this, possibly on the lacrymal as well as the maxil-
lary, and passed anterodorsally to end in the same region as
the posterior. Both are poorly developed and no attempt
10 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
has been made to decide of which layers they are vestiges.
Ear muscles (Fig. 2). These were not dissected out in detail,
but an apparently single small muscle was found beneath the
sphincter colli primitivus with fibers directed anteroventrally
to insert in the fibrous tissue on the inner side of the upper
end of the car cartilage and surrounding the slender auditory
tube.
Huber (1934, pp. 134-35, fig. 11) gives a thorough account of
the ear muscles in Monodon, and Murie (1873, p. 250, fig. 29)
discusses them in GloMcephala. Fraser and Purves (1960a,
p. 62) refer to them rather generally in the course of a more
detailed account of other aspects of the ear.
Discussion. Huber 's identification of the muscles in this
region is built on evolutionary studies which have led him to
the conclusion that the platysma and the sphincter colli pro-
fundus derive from the sphincter colli primitivus. In cetaceans
he recognizes both the primitivus and profundus, but says that
the platysyna colli et faciei is lacking. Earlier authors have
tended to include the superficial musculature of the head and
neck in the panniculus carnosus (Murie, 1873, figs. 57, 58; How-
ell, 1927, p. 19; Schulte and Smith, 1918, p. 15). Though Murie
(1873, pp. 272, 273) refers to the anterior portion as the
platysma, the others do not make such a distinction. The dorso-
ventral division into a sphincter colli primitivus and profundus
emphasized by Huber does not show in their figures. This
separation, though not very definite in our dissections, is still
sufficiently apparent to reinforce Huber 's opinion. Kesteven
(1941, pp. 74-75), while recognizing the same muscle arrange-
ment as that found by Huber and ourselves, believes that sphinc-
ter colli primitivus and profundus are actually anterior ex-
tensions of the panniculus carnosus and do not belong to the
superficial facial musculature.
Schulte and Smith's description (1918, pp. 14-15) of the
sphincter colli musculature (called panniculus carnosus) in Tur-
siops does not entirely match Huber 's and our dissections. Their
aponeurotic area on the throat anterior to the pectoralis is as we
found it. We differed, however, in not finding, on the side of
the neck immediately anterior to the forelimb, an aponeurotic
layer with which the ventral aponeurosis was continuous. On
the contrary, in this region the muscle was well developed,
passing ventrally across the sternohyoid, in part to join its fel-
low, in part to end at the pectoralis. Posterior to this and dor-
sal to the forelimb, there is a distinct separation between the
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 11
thicker band of fibers passing anterior to tlie forelimb and the
thinner layer which converges to insert on the forelimb. The
development of an aponeurosis in this area may vary individu-
ally; we found it only in Lagenorliynchus acutus, and here it
was always separated from the aponeurotic throat patch by
the flipper.
Huber 's derivatives of the sphincter colli profundus — the
auriculolabialis (tympano-zygomaticus) , orhicularis oculi, and
orbicularis oris — were all found by Howell. It is probable that
the ''bundles" of Schulte and Smith (1918, p. 14) "caudal to
the angulus oris" are the same as the orhicularis oris.
Muscles of the Neck and Throat
The muscles described in this group are ventral only, and for
the most part associated with the hyoid apparatus. The more
superficial and anterior are described first under the heading
of interramal and tongue muscles, the more posterior and deeper
are described second under the heading of hyoideal muscles.
Correlated with the development of a fusiform body, the neck
in cetaceans has been shortened and thickened and the muscles
of the throat strongly developed. The hyoid elements also are
well developed, with their main portions ossified even in young
animals. In adult individuals, the basi- and thyrohyals form a
single, broadly crescentic bone extending across the throat for
the full width between the basicranial plates. Anteriorly on
each side, a cartilaginous ceratohj^al connects each basihyal with
a three-sided, rod-like stylohyal, which is also ossified. The sty-
lohyal attaches to the exoccipital by means of a cartilaginous tip
which may be a separate element and is sometimes called the
tympanohyal.
Functionally, the expanded hyoid apparatus is important as
an attachment for a number of muscles in this region, where
the typical cetacean relation of jaw and occipitals makes a more
standard mammalian arrangement impossible. The digastric
or depressor mandibulae has its origin here, and a very thick
sternohyoid helps to fix the basi-thyrohyals in place as the jaw
is opened. Counteracting this backward pull is a moderately
strong geniohyoid and the much expanded interhyoid which
draws the thyrohyal up and forward towards the stylohyal,
Avhile the latter is in turn lifted by the palatopharyngeus.
The association of the tongue muscles with the hyoid appa-
ratus is a more normal one and needs little comment here.
12 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Interramal and Tongue Muscles
The muscles of the interramal region and the tongue are
easy to distinguish and not too different from those of other
mammals.
Mylohyoid muscle (Figs. 2, 8, my). This is a thin, coarsely
fibered, transverse muscle that covers the throat from the
symphysis of the jaw to the hyoid region. Anteriorly, a few of
the more external fibers insert in the tough membrane along the
lower margin of the jaAv, while the main mass of the muscle
turns upward to insert in the tissue internal to the lower part
of the jaw. Posteriorly, the muscle passes between the genio-
hyoid and the digastric, becoming increasingly fat towards its
insertion, with the digastric, in the fatty tissue inside the jaw.
Here, the direction of the fibers is dorsal and slightly postero-
dorsal, and the transition from muscle to fat mass is so gradual
that it is not possible to say where the one ends and the other
begins. This is characteristic of the digastric as well. The con-
nection with the hyals is vestigial and lateral only. Here, on
each side, a thin sheet of the mylohyoid passes internal to the
digastric to attach with it via the same aponeurotic sheet on
the thyrohyal. JMedial to this, the mylohyoid ends in an aponeu-
rosis which merges with that attaching the geniohyoid to the
basihyal. Anterior to this aponeurosis, the fibers of each side
meet in an ill-defined raphe which is bound, in the midline, to
the geniohyoid. In Stenella plagiodon, the anterior portion of
the mylohyoid was difficult to separate from the overlying
sphincter colli profundus.
The mylohyoid is identified as such by Schulte and Smith,
Howell, and Hein. Boenninghaus figures but does not discuss it.
Digastric muscle (Figs. 2, 3, 8, 9, d). This arises chiefly from
the thyrohyals and probably in part from the basihyal also;
laterally, a few fibers take origin along a ligament which runs
from the tip of the thyrohyal to the tissue investing the tym-
panohyal cartilage. The digastric becomes increasingly more
fatty as it passes anterodorsally to terminate in the fatty tissue
around the lower margin of the jaw. The same gradual transi-
tion from muscle to fat seen in the mylohyoid makes it not pos-
sible to define exactly the insertion of the digastric. Generally
speaking, it is around the posterior third of the jaw. Some
variation in the exact direction of the fibers showed in the dif-
ferent individuals dissected.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 13
The digastric of this paper is the same muscle as Schulte
and Smith's hyomandihularis and Howell's mono gastric. Hein's
digastric, although figured at insertion only, is clearly the same
as ours.
Geniohyoid muscle (Figs. 3, 8, gh). Situated between the
digastric muscles and internal to the mylohyoid, this muscle is
very distinct from the underlying complex of tongue muscles,
from which it is sej^arated by sheets of loose connective tissue.
It takes origin in an aponeurosis across the basihyal external
to the hyoglossus and internal to the mylohyoid muscles. As
described above, the aponeurotic insertions of both mylohyoid
and geniohyoid are closely bound together. The belly of the
muscle is thick and rather short and the insertion is in a long,
rather thick aponeurosis inside the jaw at the symphysis. A
few of these tendinous fibers are also closely bound to the
'mylohyoid.
Some variation was noted in the different forms dissected.
In Dclphinus, its origin extended a little distance on to the
thyrohyal where it was overlaid by the insertion of the digastric
and its insertion was on the inner surface of each ramus of
the jaw as well as at the symphysis. In Stenella plagiodon and
Lagenorliynchus acutiis there is no overlap of digastric and
geniohyoid at their origin. For the most part this appeared to
be a single muscle, though in Lagenorhynchiis acutus the proxi-
mal end of the belly internally showed a slight separation into
two parts.
Schulte and Smith show and describe (1918, fig. 11, p. 37)
a geniohyoid which is much like ours; in their figure 12, page
39, the origin shown for the genioglossiis is probabh'- that of the
geniohyoid. Howell describes a geniohyoideus wliich differs from
ours in taking origin along the entire cranial borders of the
basihyal instead of transversely across the middle, and suggests
that the insertion is in the tongue, not the mandible. His figures
(1927, fig. 4, p. 10, fig. 9, p. 25) of the geniohyoid, showing an
extensive thyro- as well as a basihyal origin, look more like
our hyoglossus.
Hein found no geniohyoid, but does not take this as proof of
its absence in Monodon. In his specimen only the hyoid attach-
ments of the superficial throat muscles were left, and here, as
described above, the aponeurotic attachments of mylo- and
ge7iiuhyoid are one. For this reason, we agree Avith Hein's sup-
position that a geniohyoid does occur in Monodon, and further
14 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
believe that it is much like the geniohyoid found in our dis-
sections.
The geniohyoid is figured but not discussed by Boenninghaus.
31. styloglossus (Figs. 3, 4, 8, 9, sg). This muscle has its
origin on the anterior surface of the lateral end of the stylo-
hyal, dorsal to the attachment of the interhyoideus. It is charac-
teristically thick at its origin, often blunth" wedge-shaped with
the wide end lateral, and the tapering end sometimes extending
along as much as a third of the length of the stylohyal. From its
origin it passes anteriorly, ventral to the hyoglossus and dorsal
to the geviioglossus to insert in the under surface of the tongue
anterior to, and in part also lateral to, the medial part of the
hyoglossus. Where the two meet at their insertion, they mingle.
Sehulte and Smith, Howell, and Hein all find a styloglossus
which differs little from ours.
M. hyoglossus (Figs. 4, 8, 9, /(). This is a paired muscle, that
of each side being indistinctly divided into two sections (Figs.
3, 8). Its origin laterally is from the anterior border of the
thyrohyal between the mylohyoid and interhyoid, and medially
from the basihyal (processus lingualis) internal to the ge^iio-
hyoid. From its origin the muscle passes forward between the
styloglossus and palatoglossus, radiating in a thin layer of coarse
fibers to surround the oropharynx almost completely. The lateral
section, which is also the most posterior, inserts on the dorsal
surface of the palatoglossus. The medial section sends a few
thin bundles to the sides of the oropharynx, but for the most
part passes internal to the gcnioglossus to mingle with the
styloglossus at its insertion in the under surface of the tongue.
The degree of distinctness of the two parts varied in the genera
examined.
The hyoglossus of Sehulte and Smith as well as that of
Howell is clearly the same as ours, although they differ in
certain respects. The former is described as a smaller muscle
with origin from the ceratohyal. The latter is said to have its
origin on stylo- and ceratohyal; this may be an error, as Hein
in Monodon and we, as well as Boenninghaus, in Phocoena,
found the attachment of the hyoglossus to be much as in the
Delphinidae. Hein's hyopalatinus is the lateral portion of our
hyoglossus.
M. gcnioglossus (Figs. 3, 4, 8, 9, gg). This muscle lies internal
to the geniohyoid and takes its origin medially in a long raphe
from the ventral surface of the tongue and pharynx. From its
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 15
origin it passes anterolaterally, spreading over the insertions
of hyoglossus and styloglossus to insert principally in the lining
of the anterior part of the buccal cavity. A few fibers also mingle
with the mylohyoid to insert in the lower inner margin of the
jaw posterior to the geniohyoid. At its caudal end, a narrow,
scarcely muscular band of the genioglossus lies internal to the
liyals, with which it has almost entirely lost its connection. In
Tursiops, a vestigial connection remains, via a thin aponeurotic
sheet, to the dorsal surface of the basihyals as well as to the
dorsal surface of the geniohyoid. While our failure to find such
a connection in the other three genera does not mean it is
necessarily always absent, the primary posterior insertion of the
genioglossus is medially in the ventral wall of the pharynx.
Where the palatoglossus wraps around the pharynx, the origin
of the genioglossus is from the ventral surface of the palato-
glossus, while anterior to this it is in the root of the tongue.
M. genioglossus is not always clearly distinct at its origin, and,
in Lagenorhynchiis acutus at least, where the lateral section of
the hyoglossus inserts on the palatoglossus, parts of the three
mingle so as to be indistinguishable from each other.
Schulte and Smith as well as Howell find a similar genio-
glossus. Boenninghaus ' genioglossus s. s. (1902, p. 66, pi. 1,
fig. 1) and the anterior part of Hein's genioglossus are homo-
logous with our muscle.
In addition, Boenninghaus describes a posterior extension of
the genioglossus which he calls the genioepiglotticus. Boen-
ninghaus describes and figures this (1902, p. 66, pi. 1, fig. 1)
as a medial, ventral band of muscle passing from the pharynx
to the epiglottis. This in effect would bind the wall of the
pharynx to the epiglottis anteroventrally. We found no such
connection. Posteroventrally, the tongue muscles end with the
insertion of palato- and genioglossus (Fig. 8). Caudal to this,
the pharynx passes dorsal to the cerato- and basihyals and
divides to go between the epiglottid and thyroid cartilages on
each side (Fig. 6). In this region, as described under palato-
pharyngeus, there is a continuity of muscle from pharynx to
epiglottis. This, however, is lateral and posterior to the division
of the pharynx. Medially, where the pharynx divides, the larynx
is strongly bound to the hyals by the hyocpiglotticus (see below)
but this muscle is quite distinct from the pharyngeal muscles.
Boenninghaus further distinguishes a glossoepiglottimis which
he says goes from the dorsal surface of the base of the tongue
16 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
and merpres with genioepiglotticns (1902, p. 66, pi. 1, fig. 1).
In our dissection of Phocoerm a careful search was made for
both of these muscles. Only two extrinsic muscles were found
inserting on the anterior part of the epiglottis, the hyoejnglotti-
cns and the anterior portion of the thyropalatine section of the
'paJafopharyngeus. The arrangement of these did not significantly
differ from that found in the Delphinidae.
In both groups, posterior to the margin of the palatoglossus,
where the pharynx lies dorsal to the hyoid complex, there is
a well defined non-muscular area. Anterior to this are the
muscles of the oropharynx ; posterior to this are the muscles
of the hyoid apparatus and the nasopharynx. Along the lateral
walls of the divided pharynx, a few fibers (see under palato-
glossus) pass between these two regions. They have no con-
nection with the hyoepiglotticus, from which they are separated
by a distinct space. Whether these belong to the palatopharyn-
geal complex or to that of the tongue muscles, or whether they
are vestiges of totally other muscles is not clear. In any event,
they are so poorly develoj^ed as to have little functional sig-
nificance.
Hein (1914), apparently following Boenninghaus, also de-
scribes a complex genioglossus, distinguishing a posterior por-
tion of this as the genioepiglotticus. Where he figures this as
medial (pi. 5, fig. IX, muscle 54), we believe this to be an
error, though possibly it could be part of our hyoepiglotticus.
Where he figures it as lateral (fig. X, muscle 54), it is appar-
ently a better developed part of what we describe below as a
posterior extension of the combined palato- and genioglossus.
Muscle 67 of figure X, also called genioepiglotticus, appears to
be part of the thyropalatine section of our palatopharyngeus.
Hein, like us, was unable to identify a glossoepiglotticus.
31. palatoglossus (Figs. 4, 6, 8, 9, pg). This surrounds the
oropharynx for almost its entire length, and mingles without
sharp boundary with the various tongue muscles which lie
external to it, the hyoglossus laterally and the genioglossus
ventrally. Dorsally, the palatoglossus is closely bound to the
bony palate, and in the midline fibers from opposite sides meet,
but do not form a conspicuous raphe. For the most part, the
palatoglossus is here thinly covered by the lateral section of
the hyoglossus, although anteriorly it extends beyond this latter.
From the posterior margin of the combined palatoglossus and
genioglossus a very few fibers extend, as described above, along
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 17
the lateral walls of the divided pharynx to join a similarly
indistinct section of the palatopJiaryngeus. As stated earlier,
this fiber tract is probably what Ilein has called gc7iioepiglotti-
cus. It may also include the glossocpiglotticns of Boenninghaus.
Schulte and Smith have no muscle corresponding; to this, but
Howell, Hein, and Boenninghaus all report a palatoglossus
which, except as described above, is the same as ours.
Hyoideal Muscles
Identification of those muscles which fix the position of the
hyoid relative to the sternum, and those muscles which fix the
position of the larynx relative to both the sternum and hyoid
apparatus presents no serious problems. Although their arrange-
ment differs in certain characteristic ways from that in other
mammals, homologies are not difficult to understand.
With the deeper layers the situation is different. The great
transverse extension of the thyrohyal bone and the rather nar-
row space separating the thyrohyal from the stylohyal bones
have resulted in rearrangement of the muscles of this region.
A single large mass, presumably equivalent to the cerato-hyoideus
{=interhyoideus) of other mammals, fills the whole interspace
between these two. Of the two other muscles usually found in
this region, the stylohyoideus and occipitohyoideus {—jugulo-
hyoideus), there is only one vestigial remnant. The attach-
ments of this small muscle differ from both of the above, and
whether or not it is the displaced homologue of either, it is not
possible at this time to say. From its position, we tend to be-
lieve it is the occipitohyoid and have so called it.
Sternohyoid muscle (Figs. 1, 2 sli). This is a thick muscle
arising on the anterior part of the sternum, and covering the
thyroid cartilage medially as it passes forward to insert broadly
on the basi- and thyrohyal bones.
There is little disagreement over the name of this muscle
although, by misprint, Howell (1927, p. 26) calls it as well as
the next muscle sternothyroideus.
Sternothyroid muscle (Fig. 4, st). This arises on the anterior
margin of the sternum and, in Stenella plagiodon at least, from
the first rib near its sternal attachment. Thence it passes
anteriorly to insert on the outer surface of the posterior horn
of the thyroid. Schulte and Smith (1918, p. 39), reversing
Howell's error, call this as well as the foregoing muscle ster-
7iohyoid.
]8 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Thyrohyoid muscle (Fig. 4, th). This is a thin, paired muscle
lying ventral to the strong aponeurotic sheet which connects
the thyroid and epiglottid cartilages. The ihyrohyoid takes
origin on the side of the thyroid cartilage anterior and ventral
to the insertion of the sternothyroid. Thence it passes anter-
iorly and a little ventrally, crossing the ventral part of the
thyroid insertion of the occipitothyroid, to insert on the posterior
margin of the basihyal and on the posteromedial border of the
thyrohyal as well. In some instances as in Lagenorhynchus
acutus, there is a well developed fibrous sheet across the angle
between these two, to which the thyrohyoid also attaches.
Schulte and Smith, Howell, and Hein all identify this same
muscle.
Occipitohyoid muscle (Figs. 3, 8, oh). This is a thin, small
muscle which has its origin behind the bulla in the region of
the cartilaginous tip (sometimes called tympanohyal) of the
stylohyal, whence it passes ventrally across this cartilage to
insert on the dorsal tip of the thyrohyal. There is some varia-
tion in exact origin, as the stylohyal and the exoccipital are
closely bound together and invested with tough fibrous tissue
which also covers the bulla. In Delphinns and Lagenorhynchus
acutus, the origin was from this tissue where it overlay the
stylohyal. In Tursiops, we found it in the tissue between the
stylohyal and the bulla.
Schulte and Smith (1918, pp. 36, 37, and 38) describe a
depressor mandihulae which they homologize with the occipito-
hyoideus of Rapp and Stannius. According to them it has its
origin in the dense fibrous tissue in the region of the articula-
tion of the jaw and inserts in the tip of the hyoid (=thyrohyal)
continuously with the mylohyoid. Undoubtedly this is the hom-
ologue of our muscle, although the mylohyoid in our dissections
did not extend this far laterally, and the origin of our occipito-
hyoid was more posterior. Boenninghaus (1902, pp. 61-63, fig. S)
describes and figures this muscle, which he considers to be
homologous with the hyopharyngeus or constrictor medius.
Neither Howell nor Hein mention such a muscle.
Interhyoid muscle (Figs. 3, 4, 8, ih). This paired muscle fills
the space between the hyal bones on each side. It arises from
most of the dorsal surface of the thyrohyal as well as antero-
dorsally on the basihyal, and passes anterodorsally to insert
around the posterior surfaces of stylo- and ceratohyals. Medially,
the two muscles are separated by the hyoepiglotticus. The
lateral extent of the stylohyal insertion of the interhyoid shows
LAWRENCE AND SCHEVILL : GrLAR MUSCULATURE 19
some variation. In Lacicnorlryncliu^ acufus and Drlphrnus, a
small portion attached ventral to the origin of the styloglossus;
in Stcitdla plafiiodon the attachment was more medial with
almost no overlap with the styloglossus.
Schulte and Smith call this muscle ccratohyal or hyoidcus
lotus, Howell calls it ceratohyoidcus, and Hein uses hyostylohj/oi-
deus but gives intrrhrioidcus as a synonym. Boenninghaus does
not refer to it.
M. hyo epiglottic us (Figs. 5, (i. he). This is a strong, single
muscle. Its origin medially on the dorsal surface of the basihyal
separates the left and right interhyoid: it is long anteropos-
teriorly, but rather narrow. In Stenclla plagiodon and Lageno-
rhynclius acntus, its origin extends also onto the ceratohyal,
where it is dorsal to the interhyoid. From its origin the tibers
pass posteriorly and a little ventrally to insert on the anterior
surface of the epiglottid cartilage about one-third of the dis-
tance up from its base. Ventral to this nuiscle between the
posterior border of the basihyal and the lower part of the epiglot-
tis is a strong ligament.
Kernan and Schulte describe a double h yoepiglottic muscle,
while Howell and Hein botli tind it single. Boenninghaus figures
but does not describe a hyocpifiloftieiis whicli looks like ours.
Muscles of the Pharynx
In order that the arrangement of the pharyngeal muscles in the
odontocetes be understood, the relation of the hyoid a])paratus
and the laryngeal cartilages to each other and to the skull
needs to be discussed further. In these mammals the larynx
is permanently intranarial. Consequently, it lies close to the
base of the skull and to the posterior bony nares, into which the
arytenoepiglottid cartilages protrude (.Fig. 10).
While the thyroid cartilage lies posterior to the thyro-
basihyals, the much elongated arytenoepiglottids pass anterodors-
ally internal to the hyoid apparatus, between the basioccipital-
pterygoid plates and into the nares. These plates, which extend on
each side from the exoccipital to the bony nares, project
ventrally as flanges between the basicranial region medially
and the bulla and prebnllar region laterally. This is a typ-
ically odontocete arrangement, as is a corresponding ventral
development of the exoccipital posterior to the bulla. Whether
or not this modification is primarily to help to isolate the bulla
acoustically from the throat region is of peripheral interest for
20 BT'LLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the moment. The signifieant thing is that, coincident with
these changes, the tympanohval has shifted its cranial attach-
ment to the exoccipital behind the bnlla, and as the exoccipital
has moved ventrally, so also has the stylohyal. The shift has
also moved the thyrohyal too far ventrally to articulate with
the thyroid. Such an articulation is further made impossible
by a transverse development of the flattened thyrobasihyals so
great that this complex spans the space between the basiocci-
pital plates (basioccipital crests of Fraser and Purves, 1960b,
p. 24, fig. 7; p. 25, fig. 8). The thyroid cartilage, no longer
sujiported by the hyals, is directly attached to the base of the
skull by strongly developed paired lateral muscles, the occipito-
th 11 raids.
Further rearrangements of the muscles of the pharyngeal
region are caused by the fact that the pharynx, instead of
passing dorsal to the larynx, divides in front of the hyocpiglot-
ticus to pass on each side of this muscle and the elongated
arytenoepigiottid cartilages (Fig. 10). It passes between these
cartilages and the horns of the thyroid on each side and forms
a single ])assage again posterior to the arytenoepiglottids and
internal to the occipiioihyroid muscle. This arrangement of
the pharynx, the position of the hyals, and the attachment of
the thyroid to the base of the skull effectively isolate the pharynx
from any hyal attachments of the constricting muscles.
The bones and cartilages of the base of the skull are not the
only structures that have been modified by the requirements of
an aiiuatic existence. Because of the permanently intranarial
position of the larynx, the functions of those pharyngeal muscles
which arc oft;ni called constrictors differ from those of other
mammals. Primarily, in the odontocetes, they are concerned
with holding the larynx in place in the bony nares. This is
accomplished in two ways : by means of a sphincter around the
tip of the arytenoepigiottid cartilages, and by drawing the
larynx up and forward into the back of the bony nares. Of
the muscles involved, two, the palatopharyngcus and the ptery-
(jopharyngcus, hold the larynx in place by means of a powerful
sphincter. x\ third muscle in this region, the occipitothyroid,
binds the larynx firmly to the ventral surface of the skull. The
only muscle that has a sole constricting function is the thyreo-
pharyngcus. Because of these changes, and without a detailed
study of innervation, it has not been possible to homologize
with any degree of certainty the muscles of this region with
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 21
those of other mammals. For instance, the sphincter muscle of
the posterior nares was found to be made up of two main units,
called here the pterygopharyngeus and palatopharyngeus and
very probably homologous with those of other mammals. The
same is true of the ihyreopharyngius, which might be found
to include the cricopharyugeus. Whether or not the two parts
of the hyopharyngeiis are represented by the entirely differently
placed occipitotJiyroid is uncertain.
Two other mammalian muscles normally found in this region,
the tensor and levator palati, are either missing or much modi-
fied because the soft palate as such has ceased to exist in the
Delphinidae, and their function is obsolete. Instead, there
extends posterior to the hard palate, for a short distance, a
thickened aponeurosis. This is called by some authors (e.g.
Kernan and Schulte, 1918) the velum palati; we have followed
Fraser and Purves (1960a and b) in calling it the palatine
aponeurosis. From its dorsal surface the palatopharyngeus
takes origin in part. Laterally, from the tips of the pterygoids
to the tissue investing the bulla the palatine aponeurosis is
continued as the pterygoid ligament.
.1/. styJopharyngeus (Figs. 6, 7, i), sp). This musch' takes
origin narrowly on the dorsomedial side of the stylohyal near
its cranial articulation and expands somewhat as it passes
anterodorsally across the palatopharyngeus, to mingle with it and
the ventral border of the pterygopharyngeus at its insertion in
the lateral walls of the nasopharynx. It follows closely the course
of the eustachian tube which opens into the nares internal to
the insertion of the stylopharyngeus.
Both Schulte and Smith (1918, pp. 38-39) and Kernan and
Schulte (1918, p. 261) describe a similar stylopharyngeus.
Howell gives no details of this postnarial region. Fraser and
Purves (1960a, p. 68, fig. 85; 1960b, p. 21, fig. 6) call this the
levator palaM.
Boenninghaus describes and figures (1902, pp. 45-17, pi. 1,
fig. 3) a stylopharyngeus in Phocoena with styloid origin as in
our stylopharyngeus, but with the two parts joined and lying
ventral to the pharjmx. This does not agree with our dissec-
tion of Phocoena, in which the stylopharyngeus is paired, and,
except for being less distinct from the pterygopharyngeus at its
insertion, resembles closely that of the Delphinidae. Lack of
knowledge of the Delphinidae and the fact that stylo- and
pterygopharyngeus in Phocoena are more completely merged in
22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the Avails of the pharynx have probably led Boeniiinghaus to a
mistaken identification of the distal ends of these two muscles.
His figure 3 of plate 1, which purportedly illustrates this region,
has certain errors which are further confusing and arc dis-
cussed in more detail below.
M. palatopharijnrjcus (Figs. 4-7, 9, pp, ppt). The palaio-
pharijngcus as identified here is both complex and not well
defined. It is a A'ery thick muscle which has its origin over a
large area extending from the anteromedial wall of the bony
nares to the pterygoid margin of the palate and across the
narrow palatine aponeurosis to the pterygoid ligament and
stylohyal. Within the nares (Fig. f)), its attachment is prin-
cipally on the pterygoids posterior to the vomer and on each
side of a raphe which extends from the vomer to the margin
of the palate and separates the two halves of the muscle. At its
dorsal margin, the paJaiopharyngcus mingles, with no sharp
boundary, with the ventral margin of the ptcriif/opJiarii)ifi(i(s
(Fig. 9). For this reason, it is hard to define exactly the extent
of the insertion of each on pterygoid and palatal bones. From
its origin, part of the palatopharyngeus sweeps more or less
horizontally around the tip of the arytenoepiglottid cartilages
in a strong sphincter (Figs. 7, 9), the outer layers joining those
of the tJii/rcopharyngeus in the mid-dorsal region. Some pass
beneath the thgreopharyngeus to join, via tendinous bands, the
fibers of the opposite side ; a few end on top of the thyreopharyn-
geus. A second part, consisting of the innermost layers of
the paliitopharyngeus, forms the arcus palatopharyngeus. A
third part of the palatopharyngeus, called the pars thyropaJa-
ti)ius, passes posteroventrally to insert between the thyroid and
epiglottid cartilages.
While the main mass of the muscle is internal, it can be
understood best in detail if we examine it as it shows in pro-
gressively deeper lateral dissections. When the thyrohyal bone
and its attached muscles are cut away, the thyropalatine section
of the palatopharyngeus may be seen attaching on the laryngeal
cartilages in the region where the oropharynx divides to pass
around the arytenoepiglottid cartilages (Fig. 5). Its exact
insertion varies somewhat in the genera examined. In Del-
phinus, it is on the lower half of the inner surface of the thyroid
cartilage, the adjacent part of the pharynx and the lower part
of the epiglottid cartilage, with only the suggestion of a division
between the epiglottid and more dorsal portions. In Stenella
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 23
plagiodon the insertion is distinctly double. The posterior part
is on the pharynx and on the inner face of the body of the
thyroid in a narrow vertical band extending ventrally from
the anterior border of the notch. The anterior and also more
ventral insertion is between the thyroid and the epiglottid,
chiefly on the former, but with a few strands to the latter. In
Lagcnorhynclius albirostris there are two separate thyroid attach-
ments as in Stenella plagiodon, although the muscle is not double.
The intervening fibers insert in the tissue between the epiglottid
and thyroid cartilages. In this genus attachments to the wall of
the pharynx are poorly developed.
From the anterior border of the thyropalatine portion of the
palaio pilar yngcus a few fibers (not figured) pass forward be-
tween the interhyoid and the wall of the pharynx, which here
lacks muscle fibers. Some of these fibers end on the dorsal
surface of the interhyoid, to which they are loosely bound by
connective tissue. Anterior to this and somewhat more dorsally,
a few fibers extend on top of the combined genio- and palato-
glossus. This is the portion of the palatopharyngeus referred
to in our discussion of genioglossus.
From its laryngeal insertions, in all three genera, the thyro-
palatine portion of the muscle widens as it passes anterodorsally
towards the posterior bony nares. A posterior segment of the
external portion attaches (Figs. 5, 7) strongly to the stylohyal ;
anterior to this, the external layers pass internal to the stylo-
hyal (Figs. 5, 6), to which they are loosely bound by con-
nective tissue, and attach to the pterygoid ligament. Deep to
these external thyropalatine layers, but not distinct from them,
the main mass of the palatopharyngeus (Fig. 7) passes up the
bony nares internal to the stylopharyngeus.
If the skull is so cut away that the nasopharynx can be laid
open along the midline posteriorly and its mucous membrane
dissected away from the underlying muscle, this latter is ob-
served as a single mass indistinctly divisible on the basis of
texture and direction of fibers. By far the greatest bulk of
this muscle is made up of rather coarse, anteromedially directed
fibers which are the combined palato- and pterygopharyfigeus;
in this inner aspect there is no real separation between the two.
In the midline anteriorly are paired raised bundles of a more
finely fibered layer which is probably Boenninghaus ' pars in-
terim of the palatopharyngeus. At their upper end the fibers
of this layer are as coarse as and indistinguishable from the
24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
undeiiyinf}^ pfenigopliaryvfjcus with which it mingles as the two
converge laterally towards the opening of the eustachian tube.
At their lower end the paired bundles separate to encircle the
arA^enoepiglottid spout in a strong sphincter muscle, which also
is more finely textured than the more external layers with which
it merges. In its pharyngeal aspect this sphincter, the arcus
palafopharyngeus, is separated laterally from the overlying
muscle by deep pocl^ets. The coarser mass external and lateral
to the sphincter is probably what is .sometimes called the pars
externa of the palaiopliaryngcus which medially, in part, passes
ventral to the pars interna to insert in the raphe which is closely
attached to the wall of the bony nares.
Tn Korjia, Kernan and Schulte (1918, pp. 261-262) describe
a palatopharynqens which includes all of the component parts
found in our dissections, although they do not separate a pars
interna or externa. "What we have called a tliyro palatine portion,
they refer to as the "superficial portion of the palato-pharyn-
geus." Their emphasis on the unity of this muscle is equally
applicable to our dissections. In their palatopharyngcus the
blending with pterygopharyngrtis at the insertion of stylopharyn-
geus, the involvement in the arcus palatopharyngeus, and origin
from velum and pterygoid all parallel the situation in the
delphinids. Differences, which include a band of fasciculi pass-
ing beneath our thyreopharyngeiis, as well as no mention of
any stylohyal attachments, are surprisingly slight between fam-
ilies with such markedly unlike crania.
In Dclpliinus, Fraser and Purves' palatopharyngeus pars in-
terna and externa (1960b, pp. 17-22, figs. 3-6) passes up the
nares as does our palatopharyngeus and is probably the same
as our muscle, although in their figure 3 the ventral portion, in
a somewhat misleading fashion, appears to wrap around the
ventral surface of the larynx. Hein's discussion of the pharyn-
geal region is incomplete, but his constrictor pharyngeus su-
perior is our palatopharyngeus.
Connuents on Boenninghaus' Avork will come at the end of
this section.
M. ptery go pharyngeus (Fig. 9, ptp). This muscle lies dorsal
to the palaiopharynge^is and together with it surrounds the tip
of the larjnix. Its origin is from the anterior wall of the upper
part of each bony naris. When the nasopharynx is removed
from the bony nares and the external arrangement of the
muscles examined, the stylopharyngeus is seen partially to
separate the pterygo- and palatopharyngeus laterally.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 25
The pierygopharyngcus passes outward as well as postero-
ventrally to meet its fellow of the opposite side in an ill-defined
raphe in the mid-dorsal line of the nasopharynx. Posteriorly,
in the midline, it is separated from the thyrcopharyngeus and
the transverse fibers of the palatopharyngeus by a small aponeu-
rosis. When the lining of the naso]:)harynx is removed, no
separation between palafo- and pterygopJiaryngeus is found.
Boenninghaus distinguishes a small paired, medial muscle
overlying the pierygopharyngeus dorsallj^ which he calls the
salpingopharyngeus. We found no such distinct layer in this
region, although, at its dorsal insertion, the most posterior
fasciculus of the plerygophuvyng'. iih was distinct i'or a few
millimeters. Internally, when the mucous lining was removed,
a faintly differentiated longitudinal bundle of slightly finer fibers
was observed. This rather finer bundle gradually merged dis-
tally and externally with the surrounding coarser fibers and
was quite distinct in direction as well as structure from the
ventral, internal fiber bundle identified earlier as Boenninghaus'
pars inicrna of the palatopharyngeus. Further comments on
Boenninghaus will come at the end of this section, and are
applicable also to Fraser and Purves, who follow Boenninghaus.
Kernan and Schulte describe a pierygopharyngeus which
closely resembles ours. Neither Howell nor Hein deal with the
nasopharynx.
Occipitothyroid muscle (Figs. 4-7, 9, ot). This strong muscle
draws the thyroid cartilages, and thus the whole larynx, up
towards the base of the skull. Its outer portion has its origin in
the fibrous tissue in the region of the bulla and on the margin
of the basioccipital plate anterior to the hypoglossal canal.
Thence its inner portion extends broadly across the basioccipital
to meet its fellow of the opposite side. Its insertion is on the
external surface of the anterior horn of the thyroid cartilage
and on the adjacent walls of the pharynx. Details appear to
vary in the genera examined, though the main plan is as de-
scribed above. In Delphinus and Lagcnorhynchus alhirostris
some of the posterior fibers wrap around the pharynx, passing
on top of the thyrcopharyngeus in so doing. In Tursiops a small
slip also takes origin on the tip of the stylohyal cartilage. In
Stenella plagiodon the outer and inner sections seem somewhat
more distinct at their origins. Parallel dissections were not
made of each genus, so that complete comparative comments
are not possible.
26 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
In Schulte and Smith (1918, fig. 11, p. 37) the muscle labeled
inferior constrictor is probably the outer part of our occiijito-
thyroid; in Kernan and Schulte (1918, p. 260) it is the rostral
portion of the inferior constrictor (thyreopJiaryngeus) which is
equivalent to our occipitothyroid. The attachment on the thyroid
cartilage is the same, but there is no mention of any cranial
attachment. This is the muscle called by Boenninghaus laryngo-
pharyngeus seu constrictor pharyngis inferior (1902, pp. 59-61).
Fraser and Purves (1960b, pp. 20, 21, 137, fig. 6) identify this
muscle as the constrictor pharyngeus or superior constrictor
muscle.
ilf. thyreopJiaryngeus (Figs. 5-7, 9, tp). This muscle takes
origin on the posterior horn of the thyroid cartilage and passes
anterodorsalh^ across the pharynx, medial and internal to the
occipitothyroid. The thyroid origin of this muscle is the prin-
cipal one in all four genera examined. In Delphinus the origin
extends across the cricothyroid articulation as well, and in
Lagcnorhyncluis alhirostris a few fibers medial to this take origin
on the circular layer of the oesophagus. From its origin, the
thyreopharyngeus passes anteromedially across the circular
muscle of the oesophagus and the palato pharyngeus to meet
the ptcrygopharyngeus. In Delphinus and Lagenorhynchus al-
hirostris, an inscriptio intervenes between the insertion of the
pterygo- and thyreopharyngeus, though not in TursiojJS and
Stenella plagiodon.
This is the muscle identified by Kernan and Schulte (3 918, p.
260) as the caudal portion of the inferior constrictor (thyreo-
pharyngeus). Interpreting Boenninghaus' text and figures in
the light of our dissections of Phocoena as well as of the Del-
l)hiiiida(', it seems clear that his longitudinalis oesophagi is the
same as our thyreopharyngeus . He gives an anterior cranial
attachment for this which we did not find, and which does not
seem possible in view of the way the occipitothyroid passes
dorsal to the thyreopharyngeus in this region. Hein's constrictor
inferior pharyngeus is probably the same as our thyreopharyn-
geus, although it is too incompletely described for certainty.
Homologies with Phocoena. Boenninghaus in his careful work
(1902) on the phar.ynx in Phocoena makes a diligent effort to
find and identify the various muscles found in land mammals,
especially man. This emphasis on homology leads to the recog-
nition of more layers than are apparently distinct in the delphin-
ids, and is one of the reasons why he found it impossible to
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 27
relate his work to Murie's and to Macalister's on Glohicephala.
Our dissection of Pliocoena showed an arrangement of the
pharyngeal muscles very similar to that found in the delphinids,
and one which did not always match Boenninghaus' plates.
For this reason Boenninghaus' discussion of the pars superior
of the pharynx needs to be considered in some detail. In his
introductory comments, Boenninghaus himself emphasizes that
the muscle of the pars superior is essentially a single mass
(1902, pp. 38-41), exactly as we found it to be. If this is not
kept well in mind, his subsequent division of it into as many
parts as he does is misleading.
Primarily, he identifies a constrictor of the pars superior
pharyngis which he says is made up of three distinct muscles in
terrestrial mammals: the ptertjgopharyngeus, palatopharyngeus
(pars externa and interna) and the thyreopalatinus. "We con-
sider the pterygopharyngeus to be better dcA'cloped than does
Boenninghaus, and restrict the palatopharyngens to the more
ventral part of this musculature. This discrepancy between the
limits of his pterygo- and palatopharyngeus and ours stems from
his effort to separate the two on the basis of pterygoid or palatal
attachments. Since the relationship of these bones in Fho-
coena is different from that in the delphinids, while the relation-
ship of the two sections of muscle is much the same in both,
it is better to follow an apparent though slight natural division
in the muscle mass itself.
In the phocoenids, the palatal bones intervene medially be-
tween the pterygoids and extend to the rim of the bony palate,
but in the delphinids these bones are broadly separated from
the margin of the bony palate by the pterygoids. Thus, if
identification were to be based on muscle attachment only,
the positions of pterygo- and palatopharyngeus would be re-
versed between delphinids and Phocoena. Actually, as described
above, a distinct though not pronounced separation between the
two muscles (Fig. 9) occurs approximately at the level of the
insertion of stylopharyngeus. Since in other mammals (cf.,
Sisson, 1917, p. 407) the stylopharyngeus inserts in the wall of
the pharynx between pterygo- and palatopharyngeus, the sep-
aration is best made here. Thus the more posterior part of our
pterygopharyngeus is equivalent to part of Boenninghaus' pars
externa of palatopharyngeus, as well as his associated salpingo-
pharyngeus.
28 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
While it is possible to correlate Boeuuiughaus ' pterygo-, palato-
and thyreopharyngeus with the muscles observed by us in Pho-
coena, his text and especially his figures for salpingo- and stylo-
pharijngeus and the levator veli are confusing. If his figure 3
(pi. 1, 1902) is an anterior, slightly ventral view with the
oropharynx cut away at the entrance to the sinus pyriformis,
then the stylopharyngeus is wrongly identified ; the medial muscle
anteroventral to the epiglottis is the pars externa of the palato-
pharyngeus which, lateral to its insertion on the dorsal surface of
the palatine aponeurosis, attaches also to the stylohyal. The
stylopharyngeus lies dorsal as well as a little lateral to this and fol-
lows the course of the eustachian tube into the bony nares. At
its upper end its course is much like that figured by Boenninghaus
(1902, fig. 3, no 2) for salpingopliaryngcus. We found that the
stijlopharyngem of Phocoena differs from that of the Delphinidae
in merging more completely with pterygopliaryngeus, so that it
was impossible to tell, in the region of the eustachian tube, which
slips were pterygo- and which stylopharyngeus.
At its ventral end, Boenninghaus (1902, p. 42) says his sal-
pingopharyngeus meets in an inscriptio the longitudinalis oes-
phagi (= our thyreopharyngeus). His description of the relation
between these two leaves little doubt that his salpingopharyngeus
is a slip which we could not separate from the main mass of the
pterygopliaryngeus either in the delphinids or in our adult
Phocoena. As far as its insertion at the mouth of the eustachian
tube is concerned, in Phocoena the stylo- and pterygopharyn-
geus were completely merged in this region. In the delphinids,
where the insertion of the stylopharyngeus is more discrete, this
seems to be the muscle surrounding the opening of the tube.
Boenninghaus' levator veli {^levator palati) is that part of
pterygopharyngeus with which the pars interna of palatopharyn-
geus merges. The difference in fiber direction as illustrated in
his plate 1, figure 2, layers 1 and 3, does not indicate distinct
layers, but, as shown in our dissections, merely a gradual curving
around and converging of a single muscle mass towards its
dorsal insertion. At the mouth of the eustachian tube we did
not find the discrete insertion figured by Boenninghaus. For
these reasons, as well as because Boenninghaus himself empha-
sizes the lack of separation of the muscles he discusses in this
region, we have not considered levator veli to be a separate
muscle. No tensor veli (= tensor palati) as described by Boen-
ninghaus was found. Fraser and Purves, although generally
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 29
following Boennmojhaiis, do not agree with his identification of
the tensor palati and believe (1960b, pp. 11, 19, fig. 4) that it
lies lateral to the pterygoid bones and thus external to the area
with which the present paper deals.
Discussion
The highly specific modifications of the gnlar region in the
odontocetes are for the most part centered around changes
in form and in position of the larynx, changes that are directly
related to the needs of an aquatic environment. The advantages
of an intranarial larynx in terms of underwater feeding are
apparent ; to what extent either the intranarial position or the
form of the larynx is also an advantage in phonation is not yet
explained. In any event the laryngeal cartilages both enclose
and pass through the pharynx instead of lying for the most
part ventral to it. The elongated arytenoepiglottid cartilages, on
each side of which the oropharynx passes, have already been de-
scribed. Ventral, as well as anterior to these, lie the postero-
ventrally extended palatal and pterygoid bones. A deep air
sinus divides these into inner and outer laminae, the inner
forming the anterior wall of the bony nares while the outer
extends the roof of the mouth and hence the pharynx downward
as well as posteriorly almost as far as the larynx. Together
these bones separate oro- and nasopharynx. The posterior ex-
tension of the palate, such that a purely ventral movement of
the larynx cannot disengage it from the posterior bony nares,
helps to insure the intranarial position of the larynx. This
development of the palate also directs away from the posterior
nares the food being swallowed.
The rather complicated rearrangement of the pharyngeal
muscles to insure the intranarial position of the larynx may seem
confusing in terms of homologies and comparisons with other
mammals, but functionally is very simple. The thyroid cartilage,
having lost its connection with the thyrohyal, is suspended from
the base of the skull by a very strong muscle, the occipitothyroid.
Lateral to the epiglottis this muscle also draws the walls of
the pharynx apart. Anteriorly and quite distinct from this
muscle is the palatopharyngeus, which lifts the anterior part
of the larynx up and forward into the bony nares. The laryngeal
attachments on each side are near the ventral part of the larynx,
and the muscle passes external to the pharynx.
Both the occipitothyroid and the palatopharyngeus have an
30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
outward as well as an upward pull which insures that the pharynx
is not squeezed shut where it passes on each side between the
arytenoepiglottids and the thyroid.
In addition to providing a strong muscular connection be-
tween the larynx and the skull, part of the palatopharyngeus
forms a complete sphincter around the expanded tip of the
arytenoepiglottids, which further helps to hold the larynx in
position in back of the nares.
While the primary function of these muscles has become a
supporting one, the palatopharyngeus is also involved in shorten-
ing the pharynx during swallowing. Contraction of that part
of the muscle which has its origin in the anterior wall of the
nares and which surrounds the pharynx external to the arcus,
not only grasps the arytenoepiglottids more tightly, but it also
draws the dorsal wall of the pharynx anteriorly. Posterior to
the arytenoepigottids, where the pharynx is again a single pas-
sage, the thyreopharyngeus is, as usual, a constrictor. The func-
tion of the stylopharyngeus is to spread the walls of the naso-
pharynx external to the arcus. It is apparently also involved in
opening the eustachian tube.
Powerful as are the muscles holding the larynx in place, there
are no opposing muscles to exert the posteroventral force which
would be needed to withdraw the larynx from the back of the
bony nares. The only laryngeal muscles with a posterior pull
are the poorly developed sternothyroids. Tension on these spreads
the wings of the thyroid cartilage rather than moving the entire
larynx. Ventral to the larynx, and also with a posterior pull,
lies the strongly developed sternohyoid. Tension on this moves
the hyals only slightly, with no corresponding shift of the
larynx.
Anteriorly, there is only one extrinsic laryngeal muscle, the
hyoepiglotticus. The action of this is to move the epiglottis away
from the arytenoids, thus opening the tip of the larynx. Obvi-
ously, when the larynx is opened it will not at the same time be
withdrawn from the nares, and the anterior slightly dorsal direc-
tion of the fibers from epiglottis to hyals ensures that this will
not happen.
Kecent authors (Lilly, 1961; Brown, 1962) who have assumed
that the larynx in odontocetes can be and is withdrawn from
the nares, have done so on the basis of certain phenomena which
they wished to explain, and not on the basis of the anatomy
involved. Lill}' (1961, p. 237), discussing Tursiops truncatus,
LAWRENCE AND SCHEVILL : GULAB MUSCULATURE 31
states, "The larynx can also be disengaged from the naso-
pharynx entirely by the throat muscles connected to the hyoid
bone and dropped down in order to discharge water into the
mouth from the upper bronchial tree and trachea." Aside from
the improbability of such a need, the absence of a thyroid-
thyrohyal joint, the lack of a suitable muscle connection between
larynx and hyals, and the position of the thyrobasihyals anterior
to the larynx, all prevent the muscles acting on the hyoid from
moving the larynx with it. Further, the position of the strongly
developed sternohyoid immediately ventral to the larynx inhibits
any downward movement of the larynx, especially when this
muscle is contracted as it would have to be if the hyoid were
being pulled posteriorly.
In one of his interesting and informative notes on behavior and
pathology of captive cetaceans, Brown (1962, pp. 63-64) at-
tributed the death of a captive Glohicephala scammoni to "laryn-
geal occlusion induced by inspiration of a stone," although the
stone was not found in the animal. It was picked up after
having fallen out of the severed head and subsequently inter-
preted as the cause of a laryngeal lesion. He supposes the fol-
lowing sequence of events : the stone was swallowed ; the animal
vomited, retracting the larynx as it did so; the stone then
passed up into the nasopharynx ; the larynx was reinserted in the
nasopharynx and the stone inhaled. There are a number of
anatomical difficulties with this explanation. The lack of muscu-
lature for retracting the larynx has already been discussed.
"Within the larynx, the paired arytenoids lie very close to each
other, and to the epiglottid at their base, narrowing the passage
in this region to a slit. When the larynx is opened, this is done
by the hyoepiglotticus, which separates the cartilages at their
tips; there is no additional musculature to enlarge this slit -like
passage. Between the epiglottid and the arytenoids at their base,
and somewhat ventral to the main air passage, is a distensible
pouch. A large object passing down the larynx might lodge
here, and it is difficult to see how it could then be moved down
the main air passage.
As evidence that laryngeal occlusion has occurred, lesions of
the floor and side walls of the larynx in the region of the
thyroid are offered. This is the region where the pharynx passes
on each side of the arytenoepiglottids, separating the thyroid
from the rest of the larynx and trachea. What seems more likely
is that the stone, lodged in the pharynx here, may have been
responsible for the lesions.
32 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Brown further states that a Tursiops whose jaws were forcibly-
held apart convulsed, vomited, and then expelled vomitus
through his blowhole. Though Brown believes this is evidence
of laryngeal retraction, it is far more likely that the vomitus was
passed up the nares between the sphincter and the aryteno-
epiglottids, as is anatomically entirely possible for a liquid.
Brown further theorizes that withdrawing the larynx may be
a means of facilitating "the passage of large items of food."
Actually, even if the larynx is artificially withdrawn from the
nares, the length of the arytenoepigiottids is such that they still
project across the pharynx (Fig. 11). With the larynx in place,
the highly stretchable pharynx on each side of the epiglottis
can pass any mass of food that has passed the isthmus faueium
and that can be accommodated by the oesophagus. This is pos-
sible because of the vertical expansion of the passage and the
lateral spread of the horns of the thyroid.
In a dead animal, it is perfectly possible to disengage the tip
of the larynx by reaching up the throat and into the nasal
passage with the fingers. In a specimen in which the pharyngeal
muscles had not been cut, pulling the sternothyroid muscle as
strongly as possible either ventrally or posteriorly did not with-
draw the larynx from the nares, nor did tilting the larynx force
the tip down and so out of the nares.
The possibility of jaw motion moving the larynx was also in-
vestigated. The attachments of the digastric and other anterior
throat muscles to the hyals and the independence of the hyals
from the thyroid make this seem unlikely. In addition, prying
the jaw of a dead animal open as widely as possible produced
no detectable shift in position of the larynx.
Adapted as the musculature of this region is to holding the
larynx in place intranarially without interfering with the swal-
lowing mechanism, it is not surprising that the larynx is only
with difficulty dislodged. In the absence of any morphological
arrangements to accomplish this and since there is no need for
the animals to do so, it seems clear that the odontocetes do not
voluntarily move the larjaix in and out of the nares.^
Aside from studying the hyo-pharyngeo-laryngeal region as
a functional unit, perhaps the most important result of our
dissections has been our ability to reconcile apparently very dis-
similar dissections by earlier workers, and to establish that in
the five principal odontocete families the arrangement of the
muscles and cartilages in this region is remarkably similar. Few
1 See note on page 35.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 33
investigators of cetacean anatomy have studied this region in
detail. The difficulties involved in the necessary deep dissec-
tions have caused most authors, preoccupied as they usually have
been with the general anatomy of a particular species, to limit
their attention to the more superficial throat and hyoid muscula-
ture. Others, who have studied the larynx in detail, have illus-
trated and described a larynx already severed from the skull.
The most complete account of the throat region in the
Delphinidae is still that of Glohicepliala published by Murie some
ninety years ago. This is discussed in detail in the second section
of this paper. Murie 's earlier work on Lagenorhynchus (1871)
only touches on the region studied here. In this earlier paper,
the function of the musculature of the posterior nares is dis-
cussed (p. 144), but the individual muscles are not described.
Of the hyoid muscles, two are considered in detail, the hyo-epi-
glottic which is the same as ours, and the hyoideus which equals
our interhyoid. Certain other muscles are figured but not de-
scribed. Of these, his geniohyoid (pi. 5, fig. 8) looks like our
genioglossus and his genio-hyoglossus like our hyoglossus. We
have not been able to identify his levator palati. Kesteven's work
(1941, pp. 78-79), which treats of some of the tongue and
hyoideal muscles in Delphinus dclpliis, is of particular interest in
that he describes the innervation of these. His jugulo-hyoideus
{— our occipitohyoid) is supplied by the glossopharyngeal nerve,
which supports Boenninghaus ' belief that this muscle is homo-
logous with the constrictor medius rather than the digastric. His
genio-hyoid, genio-glossus, and stylo-hyoideus {= our interhyoid)
resemble ours. His description and figures of stylo-glossus and
hyo-glossns are reversed ; the long narrow muscle which he calls
the hyo-glossus should have its origin at the end of the stylohyal
and is the styloglossus; the more medial muscle should have
its origin on the hyals and is the hyoglossus. His stylo -pharyngeus
and superior constrictor are not readily identifiable.
More complete accounts have been published of some of the
other odontocetes. For the Phocoenidae, Boenninghaus' account
(1902) of the throat of Flwcoena deals in careful detail Avith the
same region we have studied, and goes considerably further
than we have in discussing development and homology. The
lack of agreement of some of his plates with our dissections
of delphinids led us to check his findings on an adult Phocoena.
The discrepancies were mostly in the pharyngeal region and
have been analyzed earlier. Briefly, it may be said that, where
34 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
differences occurred, the arrangement of these muscles in Fho-
coena was more like that in the delphinids than like Boenning-
haus' plates. Subsequent to Boeuninghaus ' work but without
commenting on it, A. B. Howell (1927) described the myology
of the neck and throat, exclusive of the pharynx, of Neomeris,
another phocoenid. For this reason a synonymy of his muscula-
ture has been included.
The most recent comprehensive studies of this region in the
Physeteridae are those of Kogia by Schulte and Smith (1918)
on the muscles of the neck and throat, and by Kernan and
Schulte (1918) on the muscles of the pharynx. Both of these
studies have already been related to our work. Since these
workers themselves have amply commented on the findings of
earlier investigators, we have not repeated their comparisons.
For the Monodontidae, Hein's account (1914) of Monodon
monoceros is the most pertinent. Though it is primarily of the
larynx, it includes detailed descriptions of the interramal,
tongue, and hyoideal muscles, and a synonymy of his names
has been incuded in our text. His observations on the pharyn-
geal region, incomplete as they are because the hyolaryngeal
complex had been cut away from the skull, suggest an arrange-
ment very similar to that found by us. "Watson and Young's
earlier paper (1880) on Delphmapterus leucas includes the same
interramal, tongue, and hyoideal muscles that we found, with
the exception of the mylohyoid, occipiiohyoid, and hyoepiglot-
ticus. Of these, what they identify as mylohyoid is our digastric,
while the other two as well as our mylohyoid are lacking. Howell
(1930) described the interramal and throat musculature of
Monodon as resembling that of Neomeris and uses the same
names for muscles in both papers.
A recent rather hasty examination of Ziphius (Ziphiidae)
suggests to us that it, too, is much the same.
The intranarial larynx, and its associated structures, seems to
be such a successful adaptation for aquatic life that, once ar-
rived at, it has varied little. Presumably this was a modification
that occurred early in this adaptation. The arrangement of the
structures involved is so peculiar and so different from that in
land mammals that it seems unlikely to have been developed re-
peatedly. Those familiar with cetaceans will know that the
families involved diff'er so widely in skull and external characters
that the similarity in the myology of this region is all the more
remarkable.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 35
NOTE ADDED IN PRESS
In support of his theory that * ' during feeding and swallowing the
larynx is freed, laid in the bottom of the pharynx and food passed over it ' '
(in "Animals in aquatic environments: adaptations of mammals to the
ocean," Chap. 46, Handbook of Physiology — Environment, Amer. Physiol.
Soc, p. 744, 1964), Dr. John C. Lilly (personal communication 9 October
1964) reports of Tursiops that after he has forcibly disengaged the larynx
by hand, the animal re-engages it without assistance.
While this observation is good evidence of the effectiveness of the
muscles drawing the larynx into tlie back of the bony nares, it does not
also show that the animal itself can voluntarily disengage the larynx. It
does suggest that much of the time the muscles lifting the larynx are more
important than the sphincter muscles in holding it in place. This is as
would be expected, because contraction of the sphincter muscles holds the
tip of the larynx shut, and during breathing and probably during phonation
it needs to be open. In swallowing, however, the sphincter muscle would
seem to be the more important, for contraction of this around the tip of
the epiglottis would seal off the nares from the food passage, which here
is directed somewhat dorsally as well as posteriorly, and so passes diagonally
across the entrance to the bony nares. The atypical position of the passage,
more posterior than ventral to the nares, and the absence of a soft palate,
make some such arrangement for keeping the food out of the nose essential.
Dr. Lilly also writes (ibid.), "there is not room between the larynx
and the bone of the lower jaw to pass food of the size which these animals
Qormally swallow. . . . during feeding, one can palpate the throat region
and find the larynx being pulled downward and pressed outwards during
the swallowing act. ' '
Actually the larynx lies postei'oventral to the jaw (see Figs. 4 and 10)
so there is no constriction of the passage between jaw and larynx. Prob-
ably what was felt were the ceratohyals which lie anterior to the larynx and
near the external surface. Because of their close connection with the rest
of the hyoid apparatus to which the tongue muscles are attached, move-
ment of these during swallowing is to be expected. Posteriorly the larynx
itself lies deep in the neck, internal to the thick sternohyoid.
That the larynx could not be "laid in the bottom of the pharynx" is
clear also because the interrelations of aryteno-epiglottids, thyroid, and
pharynx make the postulated action an impossibility. The thyroid and ary-
teno-epiglottid cartilages are firmly attached to each other (see Fig. 17)
so that if they are moved ventrally they move as a unit. Further, since
the pharynx itself is enclosed on each side between the thyroid and aryteno-
epiglottids (Figs. 10, 11), and posterior to this the thyreo-pharyngeus
wraps around the pharynx (Figs. 5-7, 9, 16), if the tliyroid moves
ventrally, it takes the pharynx with it. Dr. Lilly's observations would seem
to show that internal jjressure in the food passage may push the whole
larynx ventrally unless this pressure is counteracted by the occipitotJiyroid
muscle. Moving the larynx ventrally in this manner would not pronate
36
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the aryteno-epiglottids. It is not a muscular action and would be invol-
untary as far as the porpoise is concerned.
Aside from the anatomy involved, it seems unlikely that the intranarial
larynx, an arrangement of such patent utility to an air-breather eating
under water, would be undone and by-passed just when it is most needed.
EXPLANATION OF FIGUEES
The drawings are all of actual dissections of single individuals, except
for Figures 8 and 9. That part of the skull shown by broken lines was
not drawn from the dissection but added later for purposes of orientation.
Figures 1-7 are of progressively deeper dissections of a specimen of
Dclphinus delphis. Figures 8 and 9 are diagrammatic in that they are
based primarily on a dissection of Lagenorhynchus albirostris, but have
certain details added from a subsequent dissection of L. acutus. Figures
10 and 11, of the laryngeo-pharyngeal region, are photographs of Stenella
Styx, but are characteristic of all the Delphinidae. The rest of the figures
are all of Gloiicephala melaena. Figures 12-15 are of progressively deeper
dissections of a single individual; Figures 16 and 17 are of two other indi-
viduals.
ABBREVIATIONS USED IN THE FIGURES
MUSCLES
al auricidolahialis
ca cricoarytenoid
do circular layer of oesophagus
ct cricothyroid
d digastric
gg genioglosstis
gh geniohyoid
h hyoglossus
he hyoepiglotticus
111 hyoglossus lateral portion
hm hyoglossus medial portion
ill. interhyoid
mil mastohumeralis
mps masseter, pars superfidalis
my mylohyoid
oc orbicularis oculi
oh occipitohyoid
or orbicularis oris
ot occipitothyroid
pg palatoglossus
pni. pectoralis major
pp palatopharyngeus
ppt palatopharyngeus, pars fhyropalatinus
ptp pterygopharyngeus
sc scalenus
sg styloglossus
sh sternohyoid
sin sternomastoid
sp stylopharyngeus
spf sphincter colli profundus
spt sphincter colli primitivus
st sternothyroid
t temporalis
th thyrohyoid
tp thyrcopharyngeus
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE
37
STRUCTURES OTHER THAN MUSCLES
A
arytenoid cartilage
NP
AC
auricular cartilage
0
AM
angle of mouth
OP
B
blubber
P
BC
buccal cavity
PA
BH
basihyal bone
PTL
BU
bulla
R
C
cricoid cartilage
S
CH
ceratohyal cartilage
SH
CJ
condyle of jaw
T
CO
occipital condyle
TH
E
eye
TO
EP
epiglottid cartilage
TPH
F
fat
TR
FL
flipper
W
II
humerus
X
J
jaw (in each instance,
right)
Y
]\I
meatal tube and investing fibrous tissue
Z
ML
melon
ZP
nasopharynx
oesophagus
oropharynx
pharynx
palatine aponeurosis
pterygoid ligament
raphe
sternum
stylohyal bone
thyroid cartilage
thyrohyal bone
tongue
tympanohyal cartilage
trachea
see explanation, Fig. 10
see explanation, Fig. 10
see explanation, Fig. 10
zygomatic arch
zygomatic process of squamosal bone
FIG. 1
Delpldnus delphis
Figure 1. Ventral view of gular region with blubber cut away to show
superficial layer of muscles.
38 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Belphinus delphis
Figure 2. Ventrolateral view with sphincter colli profundus and most
of the peetoralis major cut away. Of the sphincter colli pnmitivus, only
the part associated with the ear is shown; the small ear muscle posterior
to this was not identified.
Figure 3. Ventrolateral view of interranial and tongue muscles. The
mylohyoid has been entirely removed. The geniohyoid and digastric have
been folded back at their insertions to show the underlying layers.
LAWRENCE AXD SCHEVILL : GULAR MUSCULATURE 39
";^ M spt
FIG. 2
mh
FIG. 3
40 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Delphinus delphis
Figure 4. Ventrolateral view of the hyoideal muscles. The occipitdhyoid
has been cut away, causing the tip of the thyrohyal to sag ventrally. The
origins of the hyoglossus, styloglossus and genioglossus have also been
removed.
Figure 5. Same view as Figure 4. The thyrohyal bone and the external
muscle layers have been removed to show the deeper layers of the hyolaryn-
geal region, and the hyoid cartilages have been drawn away from the
base of the skull. The pharynx has been dissected away from the bony
palate, and pharynx and tongue folded down as a single mass. The
zygomatic process of the temporal bone, and the frontal bone dorsal to
the orbit, have been partly removed.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 41
FIG. 4
FIG. 5
42 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Delphinus delphis
Figure 6. Same view as Figure 4. The tympanohyal has been cut away
from the occipital posterior to the bulla, and the stylohyal pulled ven-
trally and rotated outward to show the styloplmryngeus going into the
posterior bony nares, as well as the attachments of the palatopharyngeus.
Part of the jaw has also been removed. The right side of the pharynx is
shown passing lateral to the epiglottis.
Figure 7. Details of the palatopharyngeus. All of the hyals except part
of the stylohyal have been removed. The cranial attachment of the
oceipitothyroid has been cut and the larynx has been pulled ventrally. In
order to show the inner layers of the palatopharyngeus which surround
the arytenoepiglottid cartilages, the pars thyropalatinus has been cut
away from the pterygoid ligament and this whole outer part folded down.
For the same reason the styloplmryngeus has been cut near its insertion and
folded up. Part of the thyroid cartilage has also been removed to show
the laryngeal attachments of the pars thyropalatinus. The main mass of
the palatopharyngeus may be seen passing into the bony nares.
LAWRENCE AND SCHKVILL : GULAR MUSCULATURE 43
Fl
FIG. T
44 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Lagenorhynchus albirostris and L. acutus
Figure 8. Ventral view of hyoid apparatus and tongue muscles. The
more superficial layers are on the lower half of the figure ; the area of
insertion of the mylohyoid is shown on this side only and, on this side,
the origin of the hyoglossiis has been cut away entirely to show the
underlying interhyoid. In the upper half of the figure the deeper layers
are shown; parts of the stylo- and hyoglossus have lieen removed and
both muscles folded outward to show their relation to palato- and genio-
glossus at their insertion in the tongue.
Figure 9. Nasopharynx and the entire laryngeo-pharyngeal region, in-
cluding bones and cartilages, cut away from the skull. The relation of
stylo-, palato-, and pterygopharyngeus at their insertion in the walls of
the nasopharynx is shown here. The approximate outline of the larynx
within the nasopharynx is indicated by broken lines. The approximate
boundary internally between palato- and pterygopharyngeus is shown by tlie
straight broken line.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 45
,TPH
FIG. 9
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46
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
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48 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
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LAWRENCE AND SCPIEVILL : GULAR MUSCULATURE 49
50 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
GLOBICEPHALA MELAENA
Ninety years liaA^e gone by since Murie pnblislied his excel-
lent and remarkably detailed morpholooy of Glohiccphala me-
laena based on a single specimen considerably advanced in decay.
Since then few workers have had the patience or the opportun-
ity to re-examine his findings, so Murie 's Avork has remained the
standard for students of odontocete structure. A recent unsuc-
cessful attempt to apply his descriptions to the myology of the
throat and tongue in certain related delphinid genera has led us
to an examination of this region in Glohicephala mdoena. This
has brought to light certain discrepancies in Murie 's work be-
tween text and figures, as well as between his figures and actual
specimens. Some of these are misleading to a student of func-
tion, and the present attempt is to clarify, not derogate from
Murie 's classic. In general, our findings are substantially as
stated in his text, though sometimes a little more detailed. Such
errors as we have found have been largely in his plates. Some
discussion is included of Macalister's work on Glohicephala
(1867), to which Murie often refers.
Ill making this comparison, four fresh-killed individuals were
dissected, each in a slightly different way, a special effort being
made to duplicate the various aspects figured by Murie. This
has shown that some of his figures must have been based on a
reconstruction or interpretation of what he found, not on the
actual dissection, and it is in such figures, particularly numbers
11 and 12 on his plate 31, that the majority of errors occur.
For purposes of orientation, we figure a few adjacent muscles
of the neck and upper thoracic region, but details of their at-
tachments have not been worked out.
Muscles of the Neck and Throat
StcDionidstoid muscle (Fig. 12, sm). The aternomastoid is as
described by Murie (p. 282). Its origin on the mastoid is internal
and slightly ventral to the masfohiimeralis, its insertion on the
sternum is superficial and posterior to the sternohyoid.
Mastohumeralis muscle (Fig. 12, mh). The mastohumeralis
{cephalo-humeral of Murie, p. 274) is essentially as he describes
it. Its origin on the mastoid is external and dorsal to that of the
sternumasioid and its insertion is on the anterior part of the
liead of the humerus.
Scalenus muscle (Figs. 13-15, sc). The relation between
Murie 's obliquus muscles and his scalenus is not entirely clear.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 51
His scalenus (fig. 68) appears to attach medially on the exoc-
eipital, and snbsequent authors (Schulte and Smith [Kogia],
1918. fig. 16, p. 49; Howell \Neomeris], 1927, fig. 3, p. 9) have
called the large muscle attaching in this area scalenus. We our-
selves, having made no dissection of the neck muscles, have fol-
lowed these authors, whose identification of scalenus is confirmed
by Fraser and Purves (in liii.).
Mylohyoid muscle (Fig. 12, my). The tnylokyoid (Murie,
mylo-Jiyoid, p. 251) is a rather thin sheet of transverse fibers,
coarse in structure, and extending from the symphysis of the
jaws to the thyrobasihyal region. Anteriorly, it attaches on each
side between the mandible and the tongue; posteriorly, it passes
between the digastric and geniohyoid to end in the fatty mass
internal to the jaw. Here the muscle itself becomes very fat
and merges with the insertion of the digastric.
The apparent origin of the mylohyoid in Murie 's figure 11 is
misleading, as it seems to arise jointly with the sterno-hyoid
from the thyrohyal cartilage. Actually its connection with the
thyruhyal is internal to the digastric, where the fasciae of both
muscles merge.
Digastric muscle (Fig. 12, d). The muscle which w^e have
called the digastric appears to be that which Murie tentatively
identifies thus (p. 251), even though he describes (p. 252) and
figures (fig. 63) it rather differently. It takes origin broadly
from the thyrohj^al cartilage and in part also from the strong
fascia which runs from the tip of the thyrohyal to the cranial
attachment of the tympanohyal cartilage. This is a fat muscle,
becoming increasingly so towards its insertion in the fatty mass
along the margin and outer surface of the posterior part of
the jaw.
Murie does not include the digastric in his figure 58, but his
styloglossus [sic] of this and of figure 11 apparently takes origin
from the thyrohyal cartilage and bears the same relation to his
genio-hyoid and sterno-hyoid as does our digastric. His styloglos-
sus differs from our digastric only in being inserted too near the
symphysis of the jaw. Probably, then, this and the digastric
of page 251 and figure 63 are the same, as well as being that
which we have called digastric.
Geniohyoid muscle (Fig. 12, gh). The apparently single genio-
hyoid arises via fascia from the anterior part of the ventral sur-
face of the basihyal cartilage, where it is partially covered by
the sternohyoid. Thence it passes anteriorly to the mandibular
52 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
symphysis. Murie describes (p. 252) and figures (fig. 11) paired
genio-hyoidei which otherwise resemble ours.
M. styloglossus (Figs. 13, 14, 16, sg) . The sUjloglossus takes;
origin rather narrowly from the anterior and ventral margin of
the midsection of the stylohyal cartilage. It passes forward be-
neath the digastric, widening as it goes, to insert in the side of
the tongue, external to the hyoglossus and internal to the genio-
glossus. Murie (p. 252) describes a muscle which he tentatively
identifies as the stylo-glossus [sic] as taking origin from "nearly
all the anteroinferior edge of the stylohal," but since he sup-
poses this muscle to be external to the hyoglossus which takes
origin broadly on the thyro- and basihyal cartilage, such a broad
origin for the styloglossus is not possible. On the other hand, his
description of the outer head of the hyo-glossus (p. 252) matches
closely our styloglossus. The discrepancy is one of muscle names,
not numbers of layers, for, while Murie 's text indicates that
there are three distinct sections of muscle, two parts of a hyo-
glossus and a single stylo-glossus, in a region where we only
found two, the left hand side of his figure 11 (right side of dis-
section) shows no more layers than we found. In this figure,
the thyrobasihyal complex is too broad, with a consequent dis-
tortion in the position of the origin of various tongue muscles
relative to each other. If one makes allowance for this and for
the difficulty of determining the actual fiber direction of certain
of the muscles, the figure resembles fairly closely the situation
we found, Hg^ being our styloglossus, Hg- our hyoglossus, while
the muscle he labels Stg is our digastric. In his figure 6, Sg
is apparently the same as our styloglossus.
M. liyoglossus (Fig. 13, h) . The hyoglossus takes origin
broadly along the anterior edge of the thyro- and basihyal
cartilages internal to the digastric and geyiiohyoid, with a few
fibers from the junction of the basi- and ceratohyal cartilages.
It inserts in the side of the tongue internal to the styloglossus.
This is substantially as described by Murie {hyo-glossus p. 252),
although it does not appear to be two-headed and has no
stylohyal attachment. It is the muscle labeled as Hg- in his
figure 11.
M. genioglossus (Fig. 13, gg). The genioglossus on each side
passes upwards, forwards, and outwards from the inferior me-
dian raphe to insert anteriorly in the tissue lining the buccal
cavity. Posteriorly its origin is in the root of the tongue internal
to the hyal cartilages, rather than from the hyals themselves.
This is the muscle called genio-hyoglossus by Murie (p. 252),
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 53
and agrees well with his description except for the absence of
hyal attachments.
M. palatoglossus (Fig. 16, pg). This muscle is much as de-
scribed by Murie (palato-glossi, p. 254), though we found it not
quite so thick. The transverse fibers lining the roof of the mouth
are closely bound to the bony palate and at their insertion
mingle chiefly with the hyoglossus.
Sternohyoid muscle (Fig. 12, sh). The sternohyoid (Murie,
sterno-hyoidei, p. 263) is a thick, strong muscle arising on the
sternum as well as the first costal cartilage, and inserting
broadly on the thyro- and basihyal cartilages posterior to the
digastric. Medially, its fascia partially covers the insertion of
the geniohyoid.
In Murie 's figures 58 and 63, the muscle labeled sterna-thyroid
is too superficial for this and agrees well with our dissection
of the sternohyoid, which it probably is. No such confusion
exists in his figure 11, where the sternohyoid is the more super-
ficial of the two.
Sternothyroid muscle (Figs. 13, 17, st). The sternothyroid on
each side is indistinctly divided into two parts which merge
completely at their sternocostal origin. The more dorsal, which
is the thicker part, takes origin on the anterior margin of the
first rib. The more ventral, more sheetlike portion takes origin
slightly internal to the anterior border of the sternum. Both
insert on the thyroid cartilage with a very thin slip extending
on top of the thyrohyoid. Murie 's description (p. 263) of the
sterno-thyroidei matches our dissections well, while his figure
11 shows the rather narrow appearance of this muscle when
seen from the ventral aspect.
Thyrohyoid muscle (Figs. 13, 14, 15, 17, th) . The paired
thyrohyoid muscles lie internal to the sternohyoid and are es-
sentially as described by Murie {thyro-hyoidei, p. 263). This is
the same muscle which he figures under a different name —
thyroideus — in figure 11.
Occipitohyoid muscle (Figs. 12, 13, 14, oh). The occipitohyoid
is a superficial muscle which takes origin in the tissue near the
articulation of the stylohyal with the skull. It passes across this
cartilage to insert on the lateral tip of the thyrohyal. Murie
describes no such muscle, but refers (p. 264) to a muscle which
takes origin near the auditory canal and inserts halfway down
the stylohyal ; presumably this is the muscle called squamostyloid
in his figure 13. The muscle as figured b}^ him is unlike any
54 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
muscle we found, nor does his figure match Maealister's descrip-
tion of the stylo-keratic (1867, p. 480) or Carte and Maealister's
of the squamostyloid (1868, p. 235), with both of which Murie
homologizes the unnamed muscle described on page 264. The
situation is further confused because both Maealister and Murie
give a stylohyal insertion for their muscle. Our dissections show
a single muscle in this region, which differs from Murie 's and
Maealister's descriptions in its thyrohyal rather than stylohyal
insertion. Material in poor condition could easily give rise
to such an error, and so it can be assumed that our occipitohyoid
is not only the same as the occipitohyoid of Rapp (1837, p. 132)
and Stannius (1849, p. 7), M^hich it matches closely and with
which Murie also homologizes his muscle, but also equivalent to
the above-mentioned stylo-keratic and squamostyloid.
Interhyoid muscle (Figs. 13, 14, ih). The interhyoid, as de-
scribed and figured by Murie {interhyoideus, p. 264 and figs. 11
and 13), fills the space between the combined stylo- and cera-
tohyal cartilages and the thyrohyal cartilage. It is undoubtedly
homologous, as he says, with Maealister's hyo-keratic (1867,
p. 480).
Muscles of the Pharynx
This is a difficult region to understand, even when series of
freshly killed specimens are available for dissection. Murie had
only one far-from-fresh specimen, and so it is not surprising
that his descriptions of the muscles in this region are rather
sketchy, and that there are certain discrepancies between his
text and plates. Neither fit entirely with our findings. We have
found it equally difficult to apply all of Maealister's description
to the various muscles found by us. The situation is further
complicated by these authors' use of many of the same muscle
names, but not always for the same muscles, and bj^ their at-
tempts to identify standard mammalian muscles in this very
non-standard mammal. For this reason it seems best to re-
describe the pharyngeal-postnarial region; Murie 's comments
will be discussed in the light of our findings, not ours as sup-
plementary to his. The arrangement of muscles is remarkably
similar to that found in the delphinids, and the names we use
are those of section one of this paper.
No levator and tensor palati as described by Murie (p. 255)
were found, and the muscle identified as Lp in figure 30, which
is of Lagenorhynchus alhirostris, is probably different from the
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 55
levator palati discussed in the text. The muscle which we found
attaching posterolateral to the fibrous mass around the bulla, as
the figure of Lp appears to show, is the occipitohyoid. Murie's
Lp as figured is too large for this, and there is no indication
of a thyrohyal attachment. Possibly this Lp is the stylo-
pharyngeus, but most probably it is, in reality, one of the neck
muscles, perhaps scalenus. The text states that "the levator
muscle covers the interspace of the pterygoid plates and the
Eustachian enlargement ; it is fleshy forwards on the palate,
narrows posteriorly, and is fixed fibrotendinously near the tym-
panic region." This is the region where the palatopliaryngeus,
pterygopliaryngeus, and stylopharyngeus are found, and no part
of these matches this description. In other words, with the
muscle only partially figured and incompletely described, it
seems best not to identify it with any of the known muscles of
the region. The same is true of the tensor palati, which is said
to be closely associated with the levator.
M. stylopharyngeus (Fig. 16, sp). The stylopharyngeus arises
rather narrowly from the dorsomedial surface of the stylohyal
cartilage, where it is difficult to separate from the palato-
pharyyigeus. It Mddens as it passes anterodorsally across the
palatopharyngeus to merge with the pterygopliaryngeus at its
insertion in the walls of the nasopharynx. This is clearly the
same as Murie's stylo-pharyngeus (p. 255, and fig. 12). In this
same figure Murie shows on the opposite side a stylohyoideus in
much the same relative position as the stylopharyngeus, though
he does not describe this in the text. However, he does (p. 264)
quote Stannius as describing a stylohyoid in Phocoena. Murie
feels that this is equivalent to his interhyoideus. This latter is
quite a different muscle from the stylopharyngeus, and is one
w^hich we also have found. The dorsal dissection of the
pharyngeo-laryngeal region which Murie attempted to show in
his figure 12 is difficult to make and interpret, and the actual
relationship of the different layers is rather different from that
figured. Between the stylohyal and the pharynx we found only
one muscle, the stylopharyngeus, Murie's stylohyoideus on the
other side being very likely a mislabeling of the same.
M. palatopharyngeus (Fig. 16, pp). The palatopharyngeus
takes origin from the pterygoid ligament across the palatine
aponeurosis and into the bony nares. Here, anteromedially, it
merges so completely with the pterygopharyngeus that the exact
origin of each could not be determined. From its origin, part
56 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
of the muscle passes posteroventrally to attach along the stylo-
hyal cartilage, and part of it sweeps back around the naso-
pharynx, passing ventral to the stylopharyngeus which is closely
bound to it, to insert in the posterior wall of the nasopharynx.
Here again its fibers merge with those of the pterygopharyngeus,
and are also partly overlaid by those of the thyreopharyngeus.
Another portion of the palatopharyngeus goes from the stylohyal
cartilage to insert on the internal surface of the thyroid
cartilage. This portion is not shown in the accompanying figures,
but in a lateral dissection of the hyo-laryngeal region after the
thyrohyoid and the thyrohyal bone are cut away, it shows as a
conspicuous mass passing from the inuer surface of the thyroid
to the stylohyal bone. In general, this muscle matches the more
ventral part of Murie's constrictor superior. Murie does not
describe this muscle, but figures it (fig. 12) anterior to the
stylohyal bone ; this must be an error, as we found no muscle
here betAveen the styloglossus and palatoglossus. In addition to
identifying the A'ery complex mass of pterygo- and palato-
pharyngeus as a single muscle, the constrictor superior, Murie
also mentions (p. 254) a palatopharyngeus, referring to
Macalister's account. Macalister's description (p. 480), except
for omitting the stylohyal attachments, matches our palato-
pharyngeus quite Avell.
M. pterygopharyngeus (Fig. 16, p^^)- The pterygopharyngeus
is a powerful muscle which passes doAvn the bony nares to
merge Avith the palatopharyngeus and wrap around the tip of
the arytenoepiglottid cartilages in a strong sphincter. It is in-
timately connected witli the stylopharyngeus also, and takes
origin on the Avails of the bony nares anteriorly; the fibers are
at first longitudinal and then, as described by Murie (p. 254),
"assume an oblique and spiral direction" to surround the tip
of the larynx, and insert in the posterior Avail of the nasopharynx
chiefly anterior to the insertion of the thyreopharyngeus, though
a few fibers mingle Avith this latter. This is that part of Murie's
superior constrictor Avliicli has no thyroid attachment and is
slioAvn in his figure 12 as Cs.
Occipitothyroicl muscle (Figs. 13-17, ot) . Two partially
separate sections are recognizable, an outer AA^hich takes origin
on the stylohyal bone near its cranial attachment and on the
basioccipital medial to the stylohyal, and an inner, larger por-
tion Avhich takes origin on the basioccipital anterior to the
scalenus. Together thej' insert on the anterior and upper por-
tion of the outer surface of the horn of the thyroid cartilage.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE o7
The inner part inserts also more posteriorly on the medial
surface of the horn of the thyroid as well as on the pharynx,
where its fibers mingle with those of the thyreo- and pterygo-
pharyngeiis. The occipitothyroideus of Murie's text (p. 265)
is clearly the same as our occipitothyroid, although we failed to
find the thyrohyal attachment mentioned by him. In our spe-
cimens the muscle passed beneath the tip of the thyrohyal bone
without attaching to it. The muscle labeled occipitothyroid in
Murie's figure 11 is misleadingly small. Adjacent to this, and
not labeled, is a larger muscle, apparently with stylohyal at-
tachments. The relation of the bones and cartilages as shown here
is not quite as we found them, but allowing for a certain error,
this muscle could either be the external part of the occipito-
thyroid or, if the stylohyal attachment is correct, the palato-
pharyngcim. Murie's dorsal view of this region shows no cranial
attachments for any of the muscles here. This is clearh- an
error, but, from the position relatii^e to the adjacent layers of
the muscle labeled constrictor medius, we incline to believe that
it is the inner layer of our occipitothyroid. Murie's brief descrip-
tion of his constrictor medius could easily fit our occipitothyroid.
Macalister's stylopharyngeus (p. 480) is the outer part of our
occipitothyroid, and his hasio-thyro-hyoid is the inner.
M. thyreopharyngeiis (Figs. 14-17, tp) . The thyreopharyngeus
arises chiefly from the outer surface of the horn of the thyroid,
with a few fibers from the inner side. It passes anterodorsally
to insert on the pharynx. Posteriorly, at its insertion, it merges
with the circular layer of the oesophagus, and anteriorly, it
meets the pterygopharyngeus in an aponeurotic sheet. This
matches Murie's description and figure fp. 254, and fig. 12) of
the inferior constrictor. Referring to Stannius' description of
Phocoena, he says (p. 255) that if a thyreopharyngeus exists in
Glohicephala it is probably part of "the constrictor."
Conclusion
The foregoing account is in no sense meant to replace Murie's
work, nor is it intended to be a complete description of the
region studied. Eather it has been the purpose of the authors to
amend Murie's myology in order to make his general and ex-
cellent descriptions of the gross anatomy of the head and throat
more useful to anatomists.
58 BULLETIN: MUSEUM OF COMPARATH^ ZOOLOGY
ACKNOWLEDGMENTS
This work was chiefly supported by the Woods Hole Oceano-
graphic Institution through contracts with the Office of Naval
Research (Biology Branch), and latterly under a grant from
the National Science Foundation.
We are much indebted to the personnel of Marine Studios,
Marineland, Florida, for their kindness in providing a number
of specimens for dissection. We are especially grateful to Dr.
Henry Kritzler not only for procuring material but also for
encouraging and assisting the first of our dissections.
We wish to thank Arctic Fisheries Products, Ltd., and their
manager, Mr. Bernard Andrews, for the opportunity of dissect-
ing Glohicephala melaena at Trinity Bay, Newfoundland, and
Dr. David E. Sergeant of the Fisheries Research Board of
Canada for his help in the field.
Particular thanks go to Dr. Francis C. Fraser and Mr. P. E.
Purves for their critical comments and discussion during the
preparation of the manuscript.
The drawings were made by Miss Jessie H. Sawj^er, the photo-
graphs by Mr. Donald W. Bourne.
LITEEATURE CITED
BOENNINGHAUS, GeORG
1902. Der Eachen von Phocaeua communis Less. Zool. Jahrlj., 17
(1-2) : 1-98, 20 text figs., pi. 1.
Brown, David H.
1962. Further observations on the pilot whale in captivity. Zoologica
(N.T.),47(1): 59-64, 2 pis.
Carte, Alexander, and Alexander Macalister
1868. On the anatomy of Balaenoptera rostrata. Philos. Trans. Roy.
Soc. London, 158: 201-261, pis. 4-7.
Eraser, F. C, and P. E. Purves
1960a. Anatomy and function of the cetacean ear. Proc. Roy. Soe.
London, ser. B, 152(946): 62-77, figs. 82-87, pis. 10-11.
1960b. Hearing in cetaceans. Evolution of the accessory air sacs and
the structure and function of the outer and middle ear in
Recent cetaceans. Bull. British Mus. (Nat. Hist.), Zool., 7(1):
1-140, 34 text figs., 53 pis., frontisp.
Hein, S. a. Arendsen
1914. The larynx and its surroundings in Monodon. Verhand. Kon.
Akad. Wetensch., Amsterdam, sect. 2, 18(3): 4-54, pis. 1-7.
Howell, A. Brazier
1927. Contribution to the anatomy of the Chinese finless porpoise,
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 59
Neomeris phocaenoides. Proc. U. S. Nat. Mus, 70, art 13 :
1-43, 14 text figs., 1 pi.
1930. Myology of the narwhal (Monodon monoceros). Amer. Jour.
Anat., 46(2) : 187-215, 8 text figs.
HuBER, Ernst
1934. Anatomical notes on Pinnipedia and Cetacea. Carnegie Inst.
Washington, Publ. No. 447: 105-136, 12 text figs.
Kernan, John D., Jr., and H. von W. Schulte
1918. Memoranda upon the anatomy of the respiratory tract, foregut,
and thoracic viscera of a foetal Kogia hreviceps. Bull. Amer.
Mus. Nat. Hist., 38, art. 8 : 231-267, 16 figs.
Kesteven, H. Leighton
1941. Some features in the anatomy and later development of the
head of Belphinus delphinus [sic] Linne. Eec. Australian Mus.
Sydney, 21(1) : 59-80, 29 text figs.
Lavtrencb, Barbara, and William E. Schevill
1956. The functional anatomy of the delphinid nose. Bull. Mus.
Comp. ZooL, 114(4): 103-152, 30 figs.
Lilly, John C.
1961. Man and dolphin. Doubleday & Co., Garden City, N. T., 312
pp., ills.
Macalisteb, Alexander
1867. On some points in the anatomy of Globiocephalus svineval
(Gray). Proc. Zool. Soc. London, 1867: 477-482, 1 text fig.
MuEiE, James
1871. Notes on the white-beaked bottlenose, Lagenorhynchus alhiro-
stris, Gray. Jour. Linn. Soc. London, Zool., 11(50): 141-153,
pi. 5.
1873. On the organization of the caaing whale, Globiocephalus melas.
Trans. Zool. Soc. London, 8(4): 235-301, 6 text figs., pis. 30-38.
Kapp, W.
1837. Die Cetaceen zoologisch-anatomisch dargestellt. Cotta, Stuttgart
& Tiibingen, vi -f 182 pp., 8 pis.
Schulte, H. von W., and M. de Forest Smith
1918. The external characters, skeletal muscles, and peripheral nerves
of Kogia breviceps (Blainville). Bull. Amer. Mus. Nat. Hist.,
38(2): 7-72, 21 text figs.
SissoN, Septimus
1917. The anatomy of domestic animals. W. B. Saunders Co., Phila-
delphia & London, 930 pp., 725 figs.
Stannius [Fbiedrich Hermann]
1849. Beschreibung der Muskeln des Tiimmlers (Delphinus phocaena).
Arch. Anat. Physiol, wiss. Med., 1849 (1): 1-41.
Watson, Morrison, and Alfred H. Young
1880. The anatomy of the northern beluga (Beluga catodon, Gray;
BelpMnapterus leucas, Pallas) compared with that of other
whales. Trans. Eoy. Soe. Edinburgh, 29 : 393-435, pis. 7-8.
(Eeceived 16 April 1964.)
60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Glohioephala melaena
Figure 12. Ventrolateral view of head and neck with sphincter colli and
associated layers removed; part of mylohyoid also cut away.
Figure 13. Same view, deeper dissection to show tongue muscles.
For abbreviations used in figures, see pp. 36-37.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE Gl
FIG. 12
sm
FIG. 13
fi2 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Glohioephala melaena
Figure 14. Tougue and sternal muscles cut away to show, in lateral
aspect, the outermost of the muscles binding the hyolaryngeal apparatus
together and to the base of the skull.
Figure 15. Dorsolateral view of same muscles.
For abbreviations used in figures, see pp. 36-37.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 63
FIG. 14
FIG. 15
64 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Globioephala melaena
Figure 16. Dorsal view of pharyngeal region detached from skull by
dissecting the nasopharynx away from the bony nares and by cutting the
tj-mpanohyal cartilage and the occipiiothyroid muscle. Anterior is to the
right of the plate; this is the view figured by Murie (1873, pi. 31, fig. 12).
Figure 17. Lateral view of areas of muscle insertion on the larynx. This
general arrangement is the same in all the odontocetes discussed.
For abbreviations used in figures, see pp. 36-37.
LAWRENCE AND SCHEVILL : GULAR MUSCULATURE 65
TPH
FIG. 16
4— EP
FIG. 17
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 2
A REVISION OF THE GENUS RHABDEPYBIS IN THE
AMERICAS (HYMENOPTERA, BETHYLIDAE)
By Howard E. Evans
With Seven Plates
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
May 26, 1965
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Phylogeny and Evolution of Crustacea. Proceedings of a confer-
ence held at Cambridge, Mass., March 6-8, 1962. H. B. Whittington
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Cloth bound, $6.75.
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Bulletin of the Museum of Comparative Zoology
HAEVARD UNIVERSITY
Vol. 133, Xo. 2
A REVISION OF THE GENUS BHABDEPYRIS IN THE
AMERICAS (HYMENOPTERA, BETHYLIDAE)
By Howard E. Evans
With Seven Plates
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
May, 1965
Bull. Mus. Comp. Zool., Harvard Univ., 133(2) : 67-151, May, 1965
No. 2 — A Revision of the Genus Rliabdepyris in the Americas
{Hymenoptera, Bethylidae)^
By Howard E. Evans
Rhahdepyris is possibly the most generalized of the several
genera which comprise the bethylid tribe Epyrini, and within
it may be seen trends in the directions of many of the other
genera of this tribe. Thus, the hairy-eyed species (subgenus
Trichotepyris) grade almost imperceptibly into Anisepyris>
the small, smooth-eyed, hairy-bodied species (subgenus Rhah-
depyris) differ from Laelius only in their more complete wing
venation ; and the larger, smooth-eyed, smooth-bodied species
(subgenus Chlorepyris) closely approach Epyris and other gen-
era with paired pits on the scutellum. It is important that the
systematics of Rhaidepyris be well understood if progress is to
be made with the remaining genera of this complex. I do not
have enough material to treat the Old World fauna at this
time, but I present here a preliminary review of the 41 known
American species.
The generic diagnosis and key for separation from other
Epyrini which I presented in 1964 (Bull. Mus. Comp. Zool.,
132: 91-96) will suffice for present purposes. A review of the
structure and terminology employed in this series of papers will
also be found there, and the acknowledgments and sources of
material listed there may also be taken to apply to the present
paper. An alphabetical listing of the abbreviations used for
body structures and for museums will be found at the conclu-
sion of this paper. Also appended are a check list and an index
of the American species.
Although in 1964 I placed Trichotepyris and Chlorepyris
in the synonymy of Rhahdepyris, I now find it convenient to
use these names for subgenera. The types of Trichotepyris and
Rhahdepyris, sensu strict o, are Palaearctic species which I have
not seen. It seems probable that our species are consubgeneric
with the Palaearctic species, but I cannot be certain of this.
Kieffer used the name Chloreprjris to apply to species with
paired scutellar pits connected by a narrow groove, but as so
defined Chlorepyris presents no real morphological gap from
such species of Rhahdepyris as, for example, origenus Kieffer.
1 Supported by a grant from the National Science Foundation, GB-1544.
70 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Chlorepyris is here redefined in a broader sense, to include all
glabrous-eyed species in which the body setae are short, pale,
and subappressed.
It is difficult to know which of the three subgenera should be
regarded as most primitive ; actually each is rather generalized
but has at least one specialized feature. All of our species of
Rhahdepyris, scnsu stricto, are small and black, while both
Trickofepyris and Chlorepyris show trends toward larger size,
bright metallic colors, and blotching on the wings. In the
latter subgenus one finds unusual modifications of body sculp-
ture, including a tendency for the scutellar groove to be widened
into pits on the sides and reduced to a thin line medially.
In all three subgenera the wing venation is much alike (see
fig. 63 in Evans, 1964) ; I have therefore made no mention of
venation in the keys and descriptions. The male genitalia are
also relativelj^ uniform in structure. They are not wholly
without characters, but the differences seem to me too small to
justify the difficulties in extracting and mounting these minute
structures.
KEY TO SUBGENERA OF EHABDEPYRIS
1. Eyes covered with short setae ; males with antennal segment three
reduced to a small ring-joint closely consolidated with four (Fig. 18)
B. TRICHOTEPYRIS Kieffer
Eyes glabrous; males with antennal segment three of variable length,
sometimes short and transverse but always well separated from
fourth segment (Figs. 19-24, 71-73) 2
2. Body and major wing veins clothed with coarse, fuscous setae; pronotum
short, its posterior margin paralleled by a foveolate groove; small,
black species A. EHABDEPYRIS Kieffer
Body and wing veins with only fine, pale setae; pronotum fairly long,
its posterior margin not paralleled by a foveolate groove; small to
moderately large species of black or metallic green or blue coloration
C. CHLOREPYRIS Kieffer
A. Subgenus Khabdepybis Kieffer
Rhahdepyris Kieffer, 1904, Bull. Soc. Hist. Nat. Metz, (2)11: 32 (type
species: R. myrmecophilus Kieffer 1904; designated by Kieffer, 1906).
Suhgeneric characters. — Small species of black coloration,
without metallic reflections ; eyes glabrous, but body and major
wing veins clothed with rather large, subappressed to suberect
black setae ; scape and legs with shorter black setae ; middle
EVANS : REVISION OF RHABDEPYRIS 71
tibiae without spines. Mandibles small, in females with two
sharp apical teeth and a series of three smaller teeth basad of
these (sometimes indistinct), in males with five strong teeth;
base of mandibles far removed from bottoms of eyes, the malar
space greater than width of mandibles at their base; antennal
scrobes not margined; males with third antennal segment much
longer than second, nearly as long as fourth segment. Pronotum
rather short, sloping strongly to the collar, its posterior margin
paralleled by a f oveolate groove ; notauli often not reaching
anterior margin of mesoscutum ; scutellar groove strong ; pro-
podeal disc only slightly (1.1-1.3 x) wider than long, with
five to nine discal carinae, transversely striate, the side-pieces
never striate but more or less alutaceous or beaded. Meso-
pleurum rather irregularly ridged and pitted, the foveae not
clearly formed, the lower fovea, when discernible, divided into
two by a vertical or oblique ridge. Claws very weakly dentate,
the tooth tending to slope outward.
Remarks. — I have not seen the type species, myrmecopJiilus,
and consequently use Rhahdepyris as a subgeneric name rather
tentatively for this complex. The six known American species
have all remained undescribed up to the present time. I am
familiar with the males of only two of the six species.
KEY TO SPECIES OF SUBGENUS KHABDEPTRIS
Females
1. Front angle of ocellar triangle less than a right angle (as in Fig. 1)
2
Front angle of ocellar triangle a right angle or slightly greater (as
in Fig. 2) 4
2. Coxae and femora wholly bright nifo-castaneous; propodeum with
nine discal carinae ; front femora robust, about 2.3 x as long as wide
1. mellipes n. sp.
Coxae and femora wholly brown or black ; propodeum with seven discal
carinae ; front femora not quite as robust, measuring 2.5-2.7 x as
long as wide 3
3. LFW 2.2 mm; scape pale castaneous; antennae elongate, third segment
about 1.0 X as long as wide; scutellar groove rather thin and shallow
2. huachucae n. sp.
LFW 1.6-1.8 mm; scape blackish except paler apically; antennae
short, third segment 1.0-1.2 x as long as wide; scutellar groove
relatively wider and deeper 3. muesehecTci n. sp.
4. Head rather long and vertex much produced above eye tops ; WH
0.93 X LH; WF 1.44 x HE; OOL 1.35 x WOT; a larger species.
72
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
LrW 2.0 mm 4. gracilis n. sp.
Head more nearly circular in anterior view, the vertex only moderately
produced above the eye tops (Fig 2); WH 0.98-1.00 x LH; WF not
over 1.25 x HE; OOL less than WOT; very small species, LFW
under 1.8 mm 5
5. Front very narrow, WF 0.90 x HE; third antennal segment wider than
long; scutellar groove wider on the sides than medially; front femora
2.4 X as long as wide 5. minutulus n. sp.
Front wider, WF 1.23 x HE; third antennal segment longer than vride
(Fig. 2); scutellar groove arching, not wider laterally than medially;
front femora 2.9 x as long as wide 6. nigriscapus n. sp.
Males
Front angle of ocellar triangle less than a right angle; head considerably
wider than high 3. muesehecTci n. sp.
Front angle of ocellar triangle approximately a right angle; WH/LH = 1.0
4. gracilis n. sp.
TABLE I. SUMMARY OF SOME CHARACTERS OF FEMALES OF SUBGENUS RHABDEPYRIS
Species
Locality
LFW
(rrim . )
WH/LH
WF/HE
OOL/WOT
Propodeal
carinae
Front angle
ocellar triangle
(degrees, approximate)
1.
mellipes
Fla. (type)
2.0
1.00
1.15
1.33
9
70
2.
huachucae
Ariz, (type)
2.2
0.92
1.06
1.15
7
80
3.
muesebecki
Honduras (type)
1.8
1.01
1.12
1.30
7
70
"Mexico"
1.7
0.98
1.27
1.30
7
80
Guerrero
1.6
0.95
1.12
1.36
7
80
Costa Rica
1.8
0.96
1.09
1.15
7
80
Bolivia
1.7
1.00
1.09
1.16
7
80
4.
gracilis
Calif, (type)
2.0
0.93
1.44
1.35
7
90
5.
minutulus
Peru (type)
1.3
1. 00
0.90
0.80
7
90
b.
nigriscapus
Argentina (type)
1.7
0.98
1.23
0.87
9
100
1. Rhabdepyris (Rhabdepyris) MELLIPES new species
Holotype. — $ , FLORIDA : Orange Co., 22 March 1930 (J. E.
Sadler, Fla. Fruit Fly Trap Survey) [USNM, No. 67,535].
Description of female type. — Length 3.0 mm ; LFW 2.0 mm.
Black ; palpi and mandibles testaceous, the latter with the
teeth rufous; antennae pale castaneous, the flagellum somewhat
dull; tegulae testaceous; legs pale rufo-castaneous except the
front and hind coxae somewhat infuscated ; wings subhyaline.
Clypeus obtusely angulate, more sharply angled on the midline,
the median carina strong, arched in profile. WH/LH = 1.0;
front narrow, WF .57 x WH, 1.15 x HE ; front angle of ocellar
EVANS : REVISION OF RHABDEPYRIS 73
triangle less than a right angle; OOL 1.33 x AVOT (Fig. 1).
Vertex broadly rounded off a considerable distance above eye
tops, distance from eye tops to vertex crest equal to slightly more
than half HE. Front strongly alutaceous, somewhat shining,
punctures shallow and inconspicuous, separated by 2-4 x
their own diameters. First four antennal segments in a ratio of
about 20:6:6:7, segment three (like the following segments
except the last) very slightly longer than thick.
Pronotal disc 1.6 x length of mesoscutum along the midline,
its posterior margin paralleled by a series of large but shallow
foveae ; pro- and mesonota alutaceous and shallowly punctate
like the front ; notauli narrowly tear-shaped, diverging in front ;
scutellar groove quite broad, deflected backward but not en-
larged at each end. Propodeal disc 1.2 x as wide as its median
length ; disc with numerous longitudinal carinae, of which three
are complete, six others nearly so; surface aside from the
carinae transversely striate ; surface of declivity and side-pieces
somewhat beaded. Front femora 2.3 x as long as wide.
Remarks. — This species is known only from the type. It is
the only member of this complex with pale legs.
2. Rhabdepyris (Rhabdepyris) huachucae new species
Holotype. — • $ , ARIZONA : Cochise Co., Huachuca Mts.,
Ramsey Canyon, 22 March 1956 (F. G. Werner & G. D. Butler)
[MCZ, No. 30,936].
Description of female type. — Length 4.0 mm ; LFW 2.2 mm.
Black; palpi and mandibles testaceous, the latter with the
teeth rufous; scape pale castaneous, flagellum dull, light brown
below but much darker on the upper side; tegulae testaceous;
coxae and hind femora black; front femora and to a lesser
extent the middle femora suffused with blackish toward the
middle, the legs otherwise pale castaneous; wings hyaline, the
veins and stigma light brown. Clypeus somewhat rounded
except with a small median tooth formed by the tip of the
strong, arched median ridge. Head higher than wide, WH
.92 x LH; front narrow, WF .59 x WH, 1.06 x HE; front angle
of ocellar triangle slightly less than a right angle, OOL 1.15 x
WOT. Vertex rounded off a distance above eye tops equal
to slightly more than half HE. Front strongly alutaceous,
somewhat shining below but rather dull at the ocellar triangle;
punctures very small, separated by 2-4 x their own diameters.
74 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
First four antennal segments in a ratio of about 26 :9 :10 :12,
segment three 1.6 x as long as wide, all flagellar segments
considerably longer than wide.
Pronotal disc 1.6 x as long as mesoscutum along the mid-
line, its posterior margin paralleled by a shallow, weakly fove-
olate groove ; surface of pro- and mesonota uniformly alu-
taceous, obscurely punctate; notauli slender, slightly attenuate
and divergent anteriorly; scutellar groove forming an arc.
Propodeal disc 1.18 x as wide as its median length; disc with
five parallel carinae, also two additional weaker carinae beside
the median carina, otherwise transversely striate ; declivity
beaded, weakly striate below; side-pieces wholly beaded. Front
femora 2.7 x as long as wide.
Remarks. — This species is known only from the type.
3. Ehabdepyris (Rhabdepyris) muesebecki new species
Holotype. — 9 , HONDURAS : intercepted at quarantine on
banana debris from Honduras, at Galveston, Texas, 18 Febru-
ary 1935 (C. P. Trotter; No. 887) [USNM, No. 67,536].
Description of female type. — Length 3.4 mm; LFW 1.8 mm.
Black ; palpi light brown ; mandibles testaceous except infus-
cated at extreme base ; scape black except apical .3 pale cas-
taneous like the following segment, remainder of antenna dark
castaneous on upper side, light yellowish brown below; tegulae
light brown; coxae and femora dark brown, legs otherwise
testaceous ; wings hyaline, veins and stigma very pale. Clypeus
prominent, obtusely subangulate except with a small acute
median angulation ; median carina high, arched. Head slightly
wider than high, AVH 1.01 x LH; front rather narrow, WF
.58 X WH, 1.12 X HE ; ocelli small, front angle of ocellar tri-
angle less than a right angle, OOL 1.30 x WOT. Vertex broadly
rounded off a distance above eye tops equal to about half
HE. Front evenly alutaceous although moderately shining,
punctures shallow but rather distinct, separated by 2-4 x
their own diameters. First four antennal segments in a ratio
of about 19 :6 :6 :7, segment three 1.1 x as long as thick, segment
eleven not longer than thick.
Pronotal disc 1.4 x as long as mesoscutum; posterior margin
of disc paralleled hy a row of foveae ; pro- and mesonota less
strongly alutaceous and much more shining than the front.
Notauli tear-shaped, diverging toward the front ; scutellar groove
EVANS : REVISION OP RIIABDEPYRIS 75
curved backward and slightly widened on each end. Propodeal
disc 1.20 X as wide as its median length, its features exactly as
described for the preceding species; declivity and side-pieces
uniformly alutaceous, somewhat shining. Front femora 2.5 x
as long as wide.
Allotype. — $ , HONDURAS : La Ceiba, 21 March 1916 (F. J.
Dyer) [USN^I].
Description of male allotype. — Length 2.0 mm; LFW 1.7
mm. Black; palpi and mandibles pale, as in female; antennae
wholly dark brown, scape almost black; legs dark brown, except
front tibiae and tarsi testaceous ; wings hyaline, veins and stigma
brownish. Clypeus angulate, the median ridge subangulate in
profile. Eyes prominent; head wider than high, WH 1.07 x LH;
front fairly broad, the eyes convergent below, WF .61 x WH,
1.35 X HE ; front angle of ocellar triangle slightly less than
a right angle; OOL 1.20 x WOT. Front strongly alutaceous,
rather weakly shining, with shallow punctures distributed about
as in female. Antennae elongate, first four segments in a ratio
of about 13 :5 :9 :11, segment three 1.6 x as long as wide, seg-
ment eleven 1.9 x as long as wide.
Thoracic dorsum somewhat more shining and less alutaceous
than the front; foveolate groove of pronotum, notauli, and
scutellar groove all essentially as in female. Propodeal disc 1.2 x
as wide as its median length, with five discal carinae and distinct
lateral and sublateral carinae, otherwise transversely striate ; side-
pieces alutaceous, somewhat shining.
Paratypes. — MEXICO : 1 $ , with tomato, 30 August 1943,
intercepted at quarantine at Brownsville, Texas, 54653, lot no.
43-11640 [USNIVI] ; 1 6 , "San Rafael Jicoltepec" ( ? = Jilo-
tepec) [USNM] ; 1 $ , 3-6 mi. S Cuernavaca, Morelos, 4000 feet,
17 April 1959 (H. E. Evans) [MCZ] ; 1 9 , Chilpancingo,
Guerrero, 4600 feet, June (H. H. Smith) [BMNH]. COSTA
RICA: 1 9, San Jose, 1940 (H. Schmidt) [Sec. Agri., Sao
Paulo, Brazil]. BOLIVIA: 1 5, Espia, Rio Bopi, July (W. M.
Mann; Mulford Exped., 1921-22) [USNM].
Variation. — The paratypes show only insignificant variation
in size (LFAV 1.6-2.0 mm). The Morelos male has the legs rather
pale, all the tarsi being testaceous, the tibiae only partially
inf uscated ; otherwise there is little variation in color. There is a
certain amount of variation in head shape and width of the
front, the Mexican female taken at Brownsville having a con-
siderably broader front than any other; the Costa Rica and
76 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Bolivia females liave the narrow^est front and also have the
lateral ocelli less far removed from the eye margins (Table I).
The propodeum shows little variation in shape or sculpturing.
It is conceivable, though I think not probable, that I am con-
fusing more than one species under one name.
4. Rhabdepyris (Rhabdepyris) gracilis new species
Holotype. — 9 , CALIFORNIA : Sacramento, 25 August 1932
(no collector given) [CAS].
Description of female type. — Length 3.2 mm; LFW 2.0 mm.
Black, except last abdominal segment suffused with dark reddish
brown ; palpi light brown ; mandibles testaceous on apical half ;
scape dark brown, paler apically, flagellum castaneous, darker
on upper side than below ; tegulae light brown ; coxae and femora
dark brown, legs otherwise light brown ; wings hyaline, veins
and stigma light brown. Clypeus obtusely angulate except acute
at the midline ; median carina low except subangularly produced
toward the base. Head higher than wide, WH .93 x LH; front
broad, AVF .66 x WH, 1.44 x HE ; ocelli small, in a rather broad
triangle, the front angle approximately a right angle; OOL 1.35
X WOT. Vertex produced well above eye tops, distance from
eye tops to vertex crest equal to over two-thirds x HE. Front
strongly alutaceous although moderately shining, the punctures
small and shallow, separated by 2-3 x their oAvn diameters.
First four antennal segments in a ratio of about 19 :7 :7 :8, seg-
ment three about 1.3 x as long as thick, segment eleven barely
longer than thick.
Pronotal disc 1.35 x as long as mesoscutum, its posterior
margin paralleled by a f oveolate groove ; surface of pro- and
mesonota moderately shining, obscurely punctate ; notauli strong
on posterior half of mesoscutum, tapering and diverging an-
teriorly ; scutellar groove rather wide, deflected backward but
barely widened on each side. Propodeal disc 1.10 x as wide as
long, with seven discal carinae, otherwise transversely striate ;
declivity and side-pieces beaded. Front femora 2.8 x as long as
wide.
Allotype. — $ , MEXICO : 10 mi. W Durango, Durango, 12
July 1954 (J. W. MacSwain) [CAS].
Description of male allotype. — Length 2.7 mm; LFW 1.9
mm. Black ; palpi brown ; mandibles testaceous on apical third,
the teeth rufous ; scape black, flagellum dark brown ; legs dark
EVANS : REVISION OF RHABDEPYBIS 77
brown; wings hj-aline. Clypeus broadly subangulate, with a
small median tootli ; median ridge arched in profile. WH/LH
= 1.0; front broad, WF .64 x WH, 1.42 x HE; front angle of
ocellar triangle very slightly exceeding a right angle; OOL 1.33
X WOT ; vertex broadly rounded oif a considerable distance
above the eye tops. Front alutaceous and weakly punctate about
as in the female. First four antennal segments in a ratio of
about 15 :5 :11 :12, segment three 1.8 x as long as wide, segment
eleven about twice as long as wide.
Thoracic dorsum weakly alutaceous, obscurely punctate; no-
tauli very short, strong only on the posterior third of the
mesoscutum; scutellar groove arcuate, slightly widened on each
side. Propodeal disc 1.1 x as wide as long, with five longitudinal
carinae, somewhat beaded beside the median carina but else-
where transversely striate ; declivity and side-pieces beaded.
Remarks. — This association of the sexes seems probable on
the basis of available material, but only a great deal more
collecting will solve this matter finally. I have seen only these
two specimens assignable to this species.
5. Rhabdepyris (Riiabdepyris) minutulus new species
Holotype.— 9, PERU (C.H.T. Townsend Coll.) (no further
data) [USNM, No. 67,537].
Description of female type. — Length 1.9 mm; LFW 1.3 mm.
Head and thorax black; abdomen dark castaneous; palpi and
mandibles testaceous ; scape and following two antennal segments
testaceous, remainder of antenna dark brown except somewhat
paler on the under side; coxae and femora dark brown, middle
and hind tibiae medium brown, legs otherwise testaceous; wings
hyaline, veins and stigma light amber. Clypeus obtusely angu-
late, with small median tooth. WH/LH = 1.0 ; front very
narrow, WF .53 x WH, .90 x HE ; ocelli in a broad triangle,
front angle about a right angle; OOL .80 x WOT. Vertex
rounded off a short distance above the eye tops, distance from
eye tops to vertex crest equal to less than one-third x HE.
Front strongly shining below, more weakly shining and strongly
alutaceous above; punctures indistinct. First four antennal
segments in a ratio of about 14 :5 :2 A, segments three and
eleven both wider than long.
Pronotal disc 1.3 x as long as mesoscutum, its posterior margin
paralleled by a rather weak series of small foveae ; surface
78 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
moderately shining, obscurely punctate. Mesoscutum with the
notauli strong on the posterior half ; scutellar groove fairly
wide, wider on the sides than medially. Propodeal disc 1.14 x
as wide as long, with five longitudinal carinae and two addi-
tional, weaker carinae beside the median carina, otherwise trans-
versely striate ; declivity and side-pieces shining, rather weakly
alutaceous. Front femora 2.4 x as long as wide,
6. Rhabdepyris (Rhabdepyris) nigriscapus new species
Eolotype.— S, ARGENTINA: 5 mi. N Jujuy, 15 Febru-
ary 1951 (Ross & Michelbacher) [CAS].
Description of female type. — Length 2.6 mm; LFW 1.7 mm.
Black ; palpi and mandibles testaceous ; scape black except paler
at tip, antennae otherwise dark brown above, light brown on
under side ; coxae and femora dark brown, legs otherwise pale
castaneous ; wings hyaline, veins and stigma light brown. Cly-
peus broadly rounded apically, with a small median tooth. WH
.98 X LH; front of moderate width, WF .64 x WH, 1.23 x HE ;
ocelli in a broad triangle, OOL .87 x WOT (Fig. 2). Distance
from eye tops to vertex crest equal to slightly less than half
HE. Front rather strongly alutaceous, somewhat shining, with
small, shallow punctures which are separated by 3-5 x their
own diameters. First four antennal segments in a ratio of about
19:7:6:7, segments three and eleven both about 1.2 x as long
as wide.
Thoracic dorsum alutaceous, moderately shining, obscurely
punctate; pronotal disc 1.2 x as long as mesoscutum, its posterior
margin paralleled by a rather strong series of foveae. Notauli
very short, barely longer than wide ; scutellar groove arching,
not wider on the sides than medially. Propodeal disc 1.10 x as
wide as long, with five longitudinal carinae between which are
four other, weaker carinae, otherwise transversely striate;
declivity weakly transversely striate ; side-pieces somewhat bead-
ed. Front femora 2.9 x as long as wide.
Remarks. — This species is similar to the preceding in many
ways, but there are so many minor differences that the two are
unlikely to be conspecific. Both species are known from the
types only.
EVANS : REVISION OP RHABDEPYRIS 79
B. Subgenus Trichotepyris Kieffer
Trichotepyris Kieffer, 1906, m Andre, Spec. Hymen. Eur., 9: 376 (type
species: B. paUidipennis Kieffer, 1906; designated by Muesebeck &
Walkley, 1950) (proposed as subgenus of FJiahdepyris) . — Kieffer,
1914, Das Tierreich, 41: 346 (placed in synonymy of Bhabdepyris) .
Suh generic characters. — Small to medium-sized bethylids ;
black, head and thorax sometimes with metallic reflections, abdo-
men sometimes brownish or in part rufous; ejes densely clothed
with short hairs ; body with short, fine, mostly pale hair, without
the strong dark setae of the preceding group (exception:
nigropilosus) ; middle tibiae with or without spines. Mandibles
large, in the male terminating in five sharp teeth, in the female
with four or five teeth of variable development ; base of mandi-
bles fairly close to bottoms of eyes, malar space less than or
about equal to width of mandibles at their base (exception:
nigropilosus) ; male with third antennal segment very small,
shorter than second segment, closely consolidated with the much
larger fourth segment. Pronotum moderately long, with or
without a foveolate groove paralleling its posterior margin;
notauli usually complete or nearly so ; scutellar groove strong ;
propodeum of variable shape, with from three to seven longitu-
dinal carinae, with or without transverse striations; declivity
and side-pieces with fine striations in most species. Mesopleurum
with an elongate upper fovea (sometimes divided) and a large
lower fovea which is often incomplete above. Claws dentate,
the tooth distinct, erect or sloping outward to some extent.
Rem<arJ{s. — I have not seen the type species of Trichotepyris
and may or may not be employing the name correctly. The type,
from Hungary, has hairy eyes and seems to agree well enough
with the American species so far as the description goes, although
it is only 3 mm long, which is close to the minimum for our
species. The above diagnosis is based entirely on the American
species. One species, nigropilosus, is intermediate in its charac-
ters between this and the preceding subgenus.
KEY TO SPECIES OF SUBGENUS TEICEOTEPYEIS
Females
1. Wings fully developed and unbanded (occasionally somewhat more
clouded around radial vein than elsewhere) ; propodeal disc at least
1.15 X as wide as long 2
80 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Wiiags either (a) very short, not reaching the abdomen, or (b)
strongly twice-banded, the bands separated by a subhyaline streak
at the stigma; small, elongate species, the propodeal disc at most
1.10 X as wide as long (Pulcliripennis group) 12
2. Body setae coarse, fuscous; veins on basal half of fore wing bearing
dark setae which are much larger than the setulae on the wing
membrane; side-pieces of propodeum wholly beaded; LFW not
over 2.1 mm (Nigropilosus group) 7. nigropilosus n. sp.
Body setae fine, pale; veins of fore wing bearing pale setae little if
any longer than those on the membrane; side-pieces of propodeiiui
at least in part striolate or aciculate (Megaceplialus group) 3
3. Antennal scrobes earinate; head very broad, WH at least 1.12 x LH
(Fig. 4) ; propodeal disc at least 1.5 x as wide as long 4
Antennal scrobes not at all earinate; head not as broad, WH at most
1.04 X LH (Figs. 5, 6) ; propodeal disc variable 5
4. Mandibles very large and protuberant, their outer margins strongly
rounded, the fourth tooth much broader than the adjacent teeth
(Fig. 43) ; WF 1.45-1.65 x HE 8. megaceplialus (Ashmead)
Mandibles less prominent, the lower margin much less strongly rounded,
the fourth tooth only slightly wider than adjacent teeth (Fig. 44) ;
WF about 1.40 x HE 9. werneri n. sp.
5. Lower mesopleural fovea well defined, either completely enclosed or
with the upper margin indistinct on the middle third (Figs. 27, 28) ;
front narrow, WF 0.95-1.20 x HE 6
Lower mesopleural fovea poorly defined above, the upper margin very
broadly incomplete (as in Figs. 25, 26) ; front variable 8
6. Front very narrow, WF slightly less than HE ; OOL 1.2 x WOT ; front
femora moderately robust, about 2.1 x as long as wide
16. subaeneus Kieffer
Front somewhat broader, WF slightly exceeding HE (Figs. 6, 7) ;
OOL 1.3-1.4 X WOT; front femora more slender, 2.3-2.5 x as long
as wide 7
7. Lower mesopleural fovea completely enclosed (Fig. 28) ; head and
thorax strongly reflecting green or blue-green . . 15. carolinianus n. sp.
Lower mesopleural fovea with its upper margin indistinct on the middle
third (Fig. 27) ; head and thorax at most faintly aeneous or vio-
laceous 11. texanus n. sp.
8. Lower part of lower mesopleural fovea with some longitudinal striae
(Fig. 26) ; front extremely broad, WF 1.7-1.8 x HE; clypeus rounded
or subtruncate apically (Fig. 5) 10. apache n. sp.
Lower mesopleural fovea without striae ; front less broad, WF not
more than 1.4 x HE .9
9. Head much longer than wide, WH 0.91 x LH (Fig. 8) ; front femora
somewhat swollen, their length about 1.9 x their maximum width;
cutting edge of mandibles unusually strongly oblique (Fig. 48) ....
17. angusticeps n. sp.
EVANS : REVISION OF RIIABDEPYRIS 81
Head slightly wider than long, Wli 1.02-1.03 x LH; front femora
more slender, 2.1-2.4 x as long as wide; cutting edge of mandibles
less strongly oblique 10
10. Front narrow, WF about 1.10 x HE; propodeum elongate, the disc
1.2 X as wide as long, wholly covered with transverse striae ; a minute
species, LFW 2.2 mm 19. plaumanni u. sp.
Front moderately wide, WF 1.25-1.35 x HE; propodeum shorter, the
disc 1.3-1.5 x as wide as long, polished and almost without sculptur-
ing on the sides; LFW 2.4-3.0 mm 11
11. Third antennal segment distinctly longer than wide; legs beyond coxae
light castaneous; scutellar groove very thin medially, connecting a
pair of roimd pits 12. mexicanus n. sp.
Third antennal segment wider than long (Fig. 9) ; femora dark brown;
scutellar groove strong, arching, only slightly widened on each
side; front femora only 2.1 x as long as wide . .13. fortunatus n. sp.
12. Wings very short, not reaching posterior margin of propodeum; pro-
podeal disc elongate, as long as or slightly longer than Avide; OOL
1.4-1.8 X WOT 23. amabilis Fouts
Wings fully developed; propodeal disc slightly wider than long; OOL
1.25-1.45 X WOT 13
13. Head slightly wider than high, WH 1.03 x LH; WF 1.08 x HE;
posterior margin of pronotum not paralleled by a punctate groove
20. pulchripennis n. sp.
Head higher than wide, WH 0.92-0.93 x LH; WF 0.88-0.95 x HE;
posterior margin of pronotum paralleled by a strong, punctate
groove 14
14. Side-pieces of propodeum shining, very finely aciculate; OOL 1.40 x
WOT 21. iridescens n. sp.
Side-pieces of propodeum less shining, with well-defined longitudinal
striae which curve upward posteriorly; OOL 1.25-1.35 x WOT
22. cwpreolus n. sp.
Males
1. Head transverse, very much wider than high (WH at least 1.12 x LH)
(Fig. 17); WF at least 1.30 x HE; propodeal disc at least 1.40 x
as wide as its median length 2
Head subcireular, at most slightly wider than high (WH not more than
1.08 X LH) (Fig. 18) ; front and propodeal disc not usually as wide
as above ^
2. Antennal scrobes not carinate; scutellar groove very thin; OOL about
1.2 X WOT ; lower mesopleural fovea somewhat striate below
10. apadhe n. sp.
Antennal scrobes margined by weak carinae which do not reach the
eye margins (Fig. 17); scutellar groove wider; OOL and WOT
subequal; lower mesopleural fovea without striations 3
82 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
3. Propodeum very short, the disc 1.45 to 1.65 x the median length, more
or less smooth and polished on the sides behind
8. megacephalus (Ashmead)
Propodeum somewhat longer, the disc 1.40 x as wide as the median
length, the surface with fine transverse striations except at the
extreme posterior margin 9. werneri n. sp.
4. Antennal scrobes weakly carinate; front and thoracic dorsmn olive-
green; propodeal disc only slightly wider than long, the width
about 1.15 x the median length 18. olivaceus n. sp.
Antennal scrobes not at all carinate; front and thoracic dorsum not or
but weakly or in small part olive-green; propodeal disc variable .5
5. Femora wholly bright ruf o-testaceous ; scape rufo-testaceous; LPW
3.4 mm; front narrow, WF 1.16 x HE 14. lupus n. sp.
Femora more or less brownish, dull; scape brownish or black; LFW
2.2-3.2 mm 6
6. Median lobe of clypeus strongly angulate, the angle slightly greater
than a right angle except the tip acute ; WH 0.98 x LH ; propodeal
disc wholly covered with rather strong transverse striae
19. plaumanni n. sp.
Median lobe of clypeus broader, obtusely sub-angulate or somewhat
rounded, usually with a median tooth, WH 1.00-1.08 x LH; propodeal
disc with the striae obsolescent at least posteriorly 7
7. Front strongly beaded, rather dull; propodeum strongly alutaceous,
its posterior margin paralleled by a strong row of foveae; OOL 1.25-
1.45 X WOT; side pieces of propodeum weakly or incompletely striate
23. amabilis Fouts
Front alutaceous or moderately beaded, somewhat shining; foveolate
groove along posterior margin of pronotum rather weak ; OOL
1.05-1.20 X WOT; side pieces of propodemn completely covered with
longitudinal striae 8
8. Propodeum short, disc measuring 1.40-1.50 x as wide as the median
length; lower mesopleural fovea not at all defined on its upper side
12. mexicanus n. sp.
Propodeum longer, disc measuring 1.20-1.35 x as wide as the median
length; lower mesopleural fovea defined on the upper side both in
front and behind, the margining ridge generally obsolete in the
middle part of the fovea 11. texanus n. sp.
EVANS : REVISION OF RHABDEPYBIS 83
TABLE II. SUMMARY OF SOME CHARACTERS OF TYPE SPECIMENS OF SPECIES OF SUBGENUS TRICHOTEPYRIS ( ¥? )
Species
LFW
(mm)
WH/LH
WF/HE
OOL/WOT
Propodeal
disc W/L
Front
femora
L/W
Antennal
scrobes
carinate
7. nigropilosus
1.9
1.00
1.00
1.25
1.35
2.5
-
8. megaceplialus
3.8
1.15
1.60
1.25
1.70
2.5
+
9. werneri
3.4
1.17
1.40
1.22
1.50
2.7
+
1 0. apache
4.0
0.94
1.80
1.75
1.60
2.6
-
1 1 . texanus
3.0
0.98
1.10
1.40
1.30
2.5
12. mexicanus
3.0
1.02
1.31
1.10
1.50
2.4
-
13. fortunatus
2.4
1.03
1.28
1.23
1.30
2.1
-
1 5. carolinianus
3.3
0.98
1.10
1.35
1.25
2.4
-
16. subaeneus
2.9
0.92
0.95
1.20
1.25
2.1
-
17. angusliceps
3,0
0.91
1.28
1.75
1.15
1.9
-
1 9. plaumanni
2.2
1.02
1.10
1.20
1.20
2.3
-
20. pulchripeniiis
2.6
1.03
1.08
1.45
1.05
2.0
-
21. iridescens
2.6
0.92
0.93
1.40
1.10
2.3
-
2 2. cupreolus
2.4
0.93
0.95
1.25
1.05
2.4
-
23. amabilis
0.6
0.93
1.05
1.80
1.00
2.3
-
TABLE III. SUMMARY OF SOViE CHARACTERS OF TYPE SPECIMENS OF SPECIES OF SUBGENUS TRICHOTEPYRIS {3cf)
Species LFW WH/LH VVF/HE
(mm)
8. megacephalus 3.0 1.15 1.40
9. werneri 2.9 1.15 1.33
10. apache 3.4 1.12 1.45
1 I . lexanus 2.8 1.04 1.25
12. mexicanus 2.5 1.07 1.35
14. lupus 3.4 1.01 1.16
18. olivaceus 2.5 1.00 1.14
19. plaumanni 2.3 0.98 1.22
23. amabilis 2.3 1.00 1.35
The specimens treated here are either holotypes (lupus, olivaceus), allotypes (werneri, apache, texanus, mexicanus,
plaumanni), or plesiallotypes (megacephalus, amabilis). The males of eight species are unknown.
./WOT
Propodeal
Antennal
Antennal
disc W/L
seg. 4
scrobes
L/W
carinate
0.96
1.60
2.0
+
0.92
1.40
2.2
+
1.20
1.60
2.2
-
1.06
1.20
2.4
-
1.06
1.45
1.9
-
1.23
1.33
2.4
-
1.16
1.15
2.5
+
1.15
1.15
2.5
-
1.45
1.35
2.5
-
84 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
NiGROPILOSUS SPECIES-GROUP
Here I assign a single species which is intermediate in its
characters between this and the precedino: subgenus. The eyes
are hairy, but the setae on the body and wing veins resemble
those of Rhahdepyris, sensu sfricto. It is known from females
only, and it is possible that the male antennae, when known, will
indicate that the species is better placed in Rhahdepyris, sensu
strict 0.
7. Rhabdeptris (Trichotepyris) NIGROPILOSUS new species
Holotype. — 5 , PANAMA : Barro Colorado Island, Canal
Zone, Jan. 1960 (W. L. Brown & E. S. McCluskey) [MCZ,
No. 30,938].
Description of female fi/pe. — Length 2.7 mm; LFW 1.9 mm.
Body black, without metallic reflections; palpi and mandibles
wholly straw-colored ; antennae light brown except basal two-
thirds of scape dark brown ; tegulae light brown ; coxae black,
femora dark brown on outer surface, legs otherwise testaceous ;
wings hyaline, veins and stigma brownish. Head and thorax
with an abundance of black setae of moderate length, most of
them directed strongly backward ; legs also with dark setae, some
of those on the tibiae fully erect ; major wing veins with dark
setae larger than those on the membrane ; abdomen with scattered
setae ventrally and apically. Mandibles with tive teeth, the
basal three teeth small, sharp. Clypeus with its median lobe
moderately prominent, rounded, Avith a small median tooth
formed by the end of the median carina, which is low although
subangulate toward its base. WH/LH = 1.0; front narrow,
WF .57 X WH, 1.0 X HE ; front angle of ocellar triangle much
less than a right angle, OOL 1.25 x AVOT. Antennal scrobes
not carinate; vertex smoothly rounded off a very short distance
above the eye tops. Front shining, moderately alutaceous, the
punctures numerous and fairly large, but so shallow as to barely
interrupt the surface. Antennae compact, first four segments
in a ratio of about 21 :6 :7 :7, segment three barely longer than
wide, outer flagellar segments (except the last) slightly wider
than long. (Fig. 3.)
Pronotal disc rather short and broad, along the midline 1.3
x as long as mesoscutum, abruptly margined both in front and
on the sides, its posterior margin paralleled by a series of small
EVANS : REVISION OF RHABDEPYRIS 85
foveae; surface (like that of the mesoscutum) shining, moder-
ately alutaeeous, the punctures shallow and rather indistinct.
Notauli diverging and much attenuated anteriorly; scutellar
groove strong, deflected backward and slightly expanded on each
end. Propodeal disc 1.35 x as wide as its median length, with
five straight, complete discal carinae as well as well developed
sublateral carinae, the disc also wholly and somewhat irregularly
transversely striate; posterolateral angles foveolate; declivity
strongly beaded, without striae, the median carina strong; side-
pieces wholly beaded, without striae. Mesopleurum with a rather
irregular series of ridges which do not form distinct, depressed
foveae. Front femora measuring 2.5 x as long as wide ; middle
tibiae not at all spinose.
Paratypes. — PANAMA : 2 9 9, El Valle, November 1946
(N.L.H. Krauss) [USNM]. BRAZIL: 3 9 9, Nova Teutonia,
Santa Catarina, July 1953, September 1957, and February 1964
(F. Plaumann) [MCZ; Coll. G. R. Ferguson].
Variation. — The Panama paratypes resemble the type very
closely in size, color, and sculpturing. In both specimens the
head is slightly broader than high (WH 1.02 and 1.04 x LH),
the front also somewhat broader in relation to the eyes (WF
1.07 and 1.12 x HE) ; OOL is 1.18 and 1.35 x WOT. The three
paratypes from southern Brazil, although from a locality over
3000 miles from the type locality, show no important differences
in color or sculpture. All are slightly larger than the Panama
specimens (LFW 2.1-2.2 mm) and two of them have an
unusually broad front (WF 1.30 and 1.50 x HE; OOL 1.25 and
1.40 X HE) ; in all three specimens the head is slightly wider
than high. Presumably this species is widely distributed in
South America.
Megacephalus species-group
To this group I assign twelve species, six of them known
from only one sex. These species lack the strong, dark setae of
the preceding species and also lack the specializations of the
wings of the pidchripennis group. This is a closeh^ knit group,
and the males are rather difficult to separate.
8. Rhabdepyris (Trichotepyris) megacephalus (Ashmead)
Epyris megacephalus Ashmead, 1893, Bull. U.S. Nat. Mus., 45: 61 [Type:
9, CALIFOENIA: Poway (San Diego Co.) (no further data)
(USNM, No. 14,067)].
86 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Rhaidepyris (TricJiotepyris) megacephalus Kieffer, 1908, Genera Insect.,
76: 32.
Bhabdepyris (Bhaidepyris) megacephalus Kieffer, 3914, Das Tierreich,
41: 355.
Rliahdepyris megacephalus Muesebeck and Walkley, 1951, U.S. Dept. Agri.,
Monogr. 2, p. 729 — Evans, 1964, Bull. Mus. Comp. Zool., 132,
figs. 63-67.
Description of female type. — Length 6.5 mm ; LFW 3.8 mm.
Black, except as follows : pronotal collar ferruginous ; apical
abdominal segment castaueous; palpi straw-colored; mandibles
ferruginous ; antennae rather uniformly light castaneous ; tegulae
testaceous ; legs bright castaneous except front coxae blackish ;
wings subhyaline, veins and stigma amber. Mandibles very
stout, their outer margins very strongly rounded ; apical margin
broad, the outer two teeth acute, the third tooth broad and short
but subacute, the fourth tooth very broad and truncate, the fifth
(basal) tooth small, rounded, and weakly separated from the
fourth tooth (Fig. 43). Clypeus very short, very broadly and
weakly subangulate, the median carina subangulate in profile.
Head very broad, 1.15 x as wide as high : front very broad, WF
.67 X WH, 1.60 X HE ; ocelli small, widely spaced, front angle
of ocellar triangle slightly less than a right angle, OOL 1.25 x
WOT. Antennal scrobes carinate, the carinae not nearly reach-
ing the eye margins; sides of head roundly convergent behind
the eyes, the vertex straight across, distance from eye tops to
vertex crest about two-thirds x HE ; head strongly developed
behind the eyes, in lateral view the temples somewhat wider
than the eyes. Front strongly shining, weakly alutaceous, with
abundant punctures which are separated by 1-2 x their own
diameters. Scape long and curved; first four antennal segments
in a ratio of about 26 :6 :5 :7, segment three 1.25 x as long as
its maximum width.
Pronotal disc sloping very gradually to the collar, sides of disc
more abruptly rounded ; collar rugulose ; disc shining, weakly
alutaceous, with small punctures ; median length of pronotal disc
about twice that of mesoscutum ; pronotum with a weak depres-
sion paralleling the posterior margin. Mesoscutum and scutellum
alutaceous, more so than the pronotum and much more so than
the front, punctures smaller than those of the pronotum but well
distributed : notauli linear, diverging toward the front ; scutellar
groove relatively wide and short, broadened and deflected back-
ward on each side. Propodeum very short, the disc 1.7 x as
wide as the median length ; disc with five longitudinal carinae
EVANS : REVISION OF RHABDEPYRIS 87
and some vague indication of other carinae between them, also
transversely ridged between the earinae; posterolateral portion
of disc smooth and polished; sublateral carinae absent; side-
pieces longitudinally striolate. Mesopleurum weakly alutaceous
and punctate, the lower fovea incomplete above. Front femora
2.5 X as long as wide ; middle tibiae very weakly spined above.
Plesiallotype.— S , AKIZONA : Tucson, 13 June 1938 (K. H.
Crandall) [MCZ].
Description of male plesiallotype. — Length 4.0 mm ; LFW
3.0 mm. Head and thorax black, faintly aeneous; abdomen
piceous, fading to dark reddish brown apically; palpi straw-
colored ; mandibles blackish except apical .2 testaceous, the teeth
rufous; antennae very dark brown except fading to medium
brown at the apex ; tegulae testaceous ; coxae dark brown, femora
medium brown, trochanters, tibiae, and tarsi light brown ; wings
hyaline, veins and stigma amber. Clypeus broadly rounded,
with a small angulation formed by the end of the median
carina, the latter very strongly arched in profile. Head broad,
1.15 x as wide as high; WF .64 x WH, 1.40 x HE; ocelli in
about a right triangle, OOL .96 x WOT. Antennal scrobes
carinate, the carinae not reaching the eye margins; vertex
straight across, distance from eye tops to vertex crest equal
to about one-third x HE. Front strongly alutaceous, weakly
shining, the punctures numerous but shallow and inconspicuous.
First four antennal segments in a ratio of about 12 :3 :2 :8,
segment four 2.0 x as long as wide, segment eleven 2.2 x as long
as wide.
Pronotal disc gradually rounded to the plane of the collar,
more abruptly rounded on the sides, the disc alutaceous and very
slightly more shining than the front; disc 1.4 x as long as
mesoscutum along the midline, its posterior margin paralleled
by a distinct, foveolate groove. Mesoscutum moderately shining,
with distinct, small punctures ; notauli linear, strongly diverging
tow^ard the front; scutellar groove strong, roundly expanded
on each side. Propodeal disc 1.6 x as wide as long, with five
longitudinal carinae between which it is somewhat rugulose,
postero-lateral portion smooth and polished; side-pieces longi-
tudinally striolate. Mesopleurum alutaceous, moderately shining,
lower fovea not well defined above.
Specimens examined. — CALIFORNIA : 1 5 , Poway, San
Diego Co. [type, USNIM] ; 1 5 , Imperial Co., 29 May 1912 (On
Helianthus, J. C. Bridwell) [USNM] ; 1 2 , Mojave Desert,
88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Aug. 1937 (R. H. Smith) [USNM]. ARIZONA: 1 9,2 $ $,
Tucson, June [USNM, MCZ] ; 1 9 , 5 mi. W Portal, May (M.
Statham) [AMNH] ; 1 9 , Marana, July (F. Werner) [UA] ;
1 9 , Tubac, June (T. Dees) [UA] ; 1 9 , Huachuca, 1937 (W.
Benedict) [KU] ; 1 9, 1 $, Salmarita, July (G. Butler)
[MCZ] ; 1 9 , Oracle Jet., Pinal Co. (F. Werner) [MCZ] ; 1 9 ,
Empire Mts., 5000 feet. May (A. Nichol) [UA] ; 1 9,2 $ S ,
Santa Rita Mts., May, July [CAS, KU] ; 1 9 , McNeal, July
(C. Williams) [UA] ; 1 9 , Elfrida, July (A. Telford) [UA] ;
1 $ , Maricopa, June (G. Butler) [UA] ; 2 c5 5 , 15 mi. W Ft.
Apache, June (Butler & Werner) [UA, MCZ] ; 1 S , Avra Val,
June [UA] ; 1 $ , Superior, July (G. Butler) [UA] ; 2 $ S ,
Arivaca, July [KU] ; 2 <5 <5 , 30 mi. E Pearce, July (Butler &
Werner) [UA, MCZ] ; 1 S , Continental, July (G. Butler)
[UA] ; 1 S , Theba, July (G. Butler) [UA] ; 2 S $ , Canelo,
July (G. Butler) [UA]. NEW MEXICO: 1 $, Florida, July
(On AcHnella, T. Cockerell) [USNM]. TEXAS: 1 9, Cooper's
Store, Big Bend Park, April (C. Michener) [KU] ; 1 S , Sierra
Blanca, El Paso Co., July [USNM].
Variation. — The females show little variation in size or
standard measurements ; LFW varies from 2.8 to 3.8 mm ; WH
from 1.12 to 1.17 x LH ; WF 1.45 to 1.62 x HE; propodeal disc
from 1.50 to 1.65 x as wide as long. Several specimens have
weak aeneous or violaceous reflections on the front and on the
thoracic dorsum ; several have the middle and hind coxae strongly
infuscated, like the front coxae, and two have the femora more
or less infuscated. The specimens from Tubac and from McNeal,
Arizona, have the front more distinctly alutaceous than any of
the others. None of the variation seems closely correlated with
geography.
The males are somewhat more variable. LFW varies from
2.3 to 3.3; WH from 1.12 to 1.20 x LH ; WF from 1.30 to 1.50
X HE ; propodeal disc from 1.45 to 1.65 x as wide as long. Sev-
eral specimens lack metallic reflections on the head and thorax;
the color of the flagellum varies from wholly light castaneous
to wholly black; in a few specimens the legs are very dark, only
the tarsi being light brown. In some specimens the front and
thoracic dorsum are rather dull, in others moderately shining.
In no case are the punctures of the front conspicuous, but in
several specimens they are more evident than in the plesiallo-
type. Again, none of the variation seems correlated with
geography.
EVANS : REVISION OF RHABDEPYRIS 89
9. Khabdepyris (Trichotepyris) werneri new species
Holotype. — 9 , ARIZONA : Globe, Gila Co., 3600 feet eleva-
tion, 8 July 1949 (mesquite-oak; F. Werner & W. Nutting)
[USNM, No. 67,538].
Description of female type. — Length 6 mm ; LFW 3.4 mm.
Black, except as follows : pronotal collar ferruginous ; apical
rims of abdominal tergites, and all of apical two tergites, brown-
ish ; palpi and tegulae testaceous ; mandibles ferruginous ; an-
tennae light castaneous, slightly paler below than above ; legs
light castaneous except front coxae blackish, middle and hind
coxae and all the femora weakly suffused with brownish ; wings
hyaline, veins and stigma amber. Mandibles with the lower
margin weakly curved, apex with five teeth in an oblique series,
basal two teeth blunt, the fourth tooth only slightly wider than
adjacent teeth (Fig. 44). Clypeus very short, the apex very
broadly and weakly angulate, the median carina rather broad
and ill-defined, not arched or angulate in profile. Head broad,
1.17 X as wide as high; front very broad, WF .67 x WH, 1.40
X HE ; front angle of ocellar triangle slightly less than a right
angle; OOL 1.22 x WOT. Antennal scrobes carinate, the carinae
not reaching the eye margins ; distance from eye tops to vertex
crest equal to about half the eye height ; vertex straight across ;
head wide behind the eyes, but the temples slightly less bulging
than in megacephalus. Front alutaceous, moderately shining,
punctures separated by 1-2 x their own diameters. First four
antennal segments in a ratio of about 40 :10 : 9 :9, segment three
1.35 X as long as its maximum width. (Fig. 4.)
Pronotal disc sloping very gradually in front, sides more
abruptly rounded; collar weakly rugulose; disc 1.65 x median
length of mesoscutum ; surface alutaceous and punctate about
like the front ; posterior margin paralleled by a weak depres-
sion which contains a series of fairly large punctures. Meso-
scutum and scutellum alutaceous like the pronotum, but with
somewhat weaker punctures ; notauli linear, strongly diverging
toward the front ; scutellar groove narrow, bent backward but
not notably broadened on each side. Propodeum longer than in
megacephalus, the disc measuring 1.5 x as wide as the median
length ; disc with five longitudinal carinae and some less dis-
tinct carinae between them, also transversely striolate except
toward the posterior margin ; side-pieces strongly longitudinally
striolate. Mesopleurum alutaceous, weakly punctate, the upper
90 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
fovea complete but the lower fovea not at all defined above
(Fig. 25). Front femora 2.7 x as long as wide; middle tibiae
with several small spines above.
Allotype. — $ , ARIZONA : 19 mi. N Globe, Gila Co., 20
June 1957 (mesquite; F. Werner & G. Butler) [MCZ].
Description of male allotype. — Length 3.5 mm ; LFW 2.9
mm. Black, except as follows: palpi and tegulae light brown;
tips of mandibles rufous; flagellum dark brown, fading to
medium brown apically ; middle and hind coxae and all femora
dark brown, tibiae light brown, tarsi testaceous; wings hyaline,
veins and stigma amber. Clypeus obtusely angulate, with a
small median tooth formed by the end of the median carina, the
latter high and arched. Head broad, 1.15 x as Avide as high ;
WF .65 X WH, 1.33 x HE; ocelli in about a right triangle,
OOL .92 X WOT. Antennal scrobes carinate, the carinae not
nearly reaching the eye margins ; vertex passing nearly straight
across a short distance above eye tops. Front rather strongly
alutaceous although moderately shining, punctures w^eak. First
four antennal segments in a ratio of about 18 :5 :3 :15, segment
four 2.2 X as long as wide, segment eleven twice as long as its
maximum width. (Fig. 17.)
Pronotal disc 1.3 x as long as mesoscutum along midline, the
surface alutaceous about like the front; posterior margin par-
alleled by a shallow, obscurely punctate groove. Mesoscutum
also alutaceous, moderately shining, obscurely punctate ; notauli
linear, diverging toward the front ; scutellar groove widened and
turned backward on each side. Propodeal disc 1.4 x as wide as
long, with five longitudinal carinae, the more lateral ones rather
short, the surface obliquely striolate laterad of the three median
carinae, elsewhere transversely striolate except smooth and
polished on the posterior .2 of the disc ; side-pieces longitudinally
striolate. Mesopleurum moderately shining, the lower fovea open
above.
Paratype. — ARIZONA : 1 ? , Sahuarita, Pima Co., 5 July
1956 (swept from cotton, G. D. Butler) [MCZ].
Variation. — The paratype is smaller than the type (LFW
2.9 mm) and shows faint coppery reflections on the head and
thoracic dorsum. It is closely similar to the type in color and
sculpturing. WF is .66 x WH, 1.39 x HE ; OOL is 1.15 x WOT ;
the propodeal disc is 1.45 x as wide as high.
EVANS : REVISION OP RHABDEPYRIS 91
10. Rhabdepyris (Trichotepyris) apache new species
Holotijpe. — 5 , ARIZONA : Continental, Pima Co., 18 Au-
gust 1960 (sucked from cotton, G. D. Butler) [MCZ, No. 30,-
940].
Description of female type. — Length 7 mm ; LFW 4 mm.
Head and thorax black, thoracic dorsum and to a lesser extent
the head with a weak bluish cast; propodeum black; abdomen
piceous, the venter and the apical tergite suffused with castane-
ous; palpi testaceous; mandibles dark ferruginous, infuscated
toward the base ; scape dark castaneous, infuscated on the upper
side throughout ; tegulae light brown ; legs bright ruf o-testaceous
except all coxae black, front femora mostly blackish on the
lateral surface ; fore wings lightly tinged with brownish, veins
and stigma amber. Mandibles broad, with five teeth, the basal
three teeth somewhat rounded, subequal in size. Clypeus with
a strongly projecting median lobe which is rounded apically;
median ridge arched near the base, then nearly straight to the
apex. Head higher than wide, "WH .94 x LH; front broad,
WF .72 X WH, 1.80 x HE; ocelli small and well separated,
front angle of ocellar triangle less than a right angle, posterior
ocelli removed from vertex crest by a distance slightly greater
than WOT ; OOL 1.75 x WOT, subequal to HE. Head thin, the
temples not developed; antennal scrobes not carinate; vertex
broadly rounded off far above eye tops, distance from eye tops
to vertex crest nearly equal to HE. Front strongly shining, very
weakly alutaceous, punctures small although sharply defined,
separated by 2-4 x their own diameters. First four antennal seg-
ments in a ratio of about 22 :5 :5 :6, segment three 1.2 x as long
as its apical width. (Fig. 5.)
Pronotum rather large, along the midline 1.7 x as long as
mesoscutum; posterior margin not paralleled by a punctate
groove ; surface shining but slightly more evidently alutaceous
than the front, punctures slightly larger and more widely
spaced than on the front. Mesoscutum moderately alutaceous,
with small punctures ; notauli linear, diverging anteriorly ;
scutellar groove relatively long and thin, deflected backward at
each end but only very slightly enlarged there. Propodeal disc
rather short, measuring 1.6 x as wide as its median length ; disc
with five carinae and two more weak, irregular carinae beside
the median carina, otherwise with fine transverse striae except
92 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY
toward the posterior margin ; lateral earinae strong, but sub-
laterals absent; declivity with curved striae which radiate from
the median carina; side-pieces longitudinally striolate. Meso-
pleurum alutaceous, the lower fovea broadly open above, con-
taining some rather distinct longitudinal striae (Fig. 26).
Front femora 2.6 x as long as wide ; middle tibiae with strong
spines above for most of their length.
Allotype. — $ , MEXICO : La Aduana, Sonora, 22 May 1962
(F. D. Parker & L. A. Stange) [MCZ].
Description of male allotype. — Length 5 mm ; LFW 3.4 mm.
Black, head and thoracic dorsum very faintly aeneous, tip of
abdomen dark reddish brown ; palpi light brown ; mandibles
black, flagellum dark brown; legs pale castaneous except all
coxae and femora strongh- infuscated (middle femora only
slightly infuscated) ; wings subhyaline, veins and stigma light
brown. Clypeus obtusely angulate, the median carina nearly
straight in profile. Head broad, 1.12 x as wide as high ; front
broad, AVF .65 x AVH, 1.45 x HE ; ocelli in about a right tri-
angle, OOL 1.20 X WOT. Antennal scrobes not carinate ; vertex
broadly rounded off a distance above eye tops equal to less
than half HE. Front shining, weakly alutaceous, with strong
punctures which are separated by 1-2 x their own diameters.
First four antennal segments in a ratio of about 25 :7 :4 :24,
segment four 2.2 x as long as wide, segment eleven 2.6 x as
long as wide.
Pronotal disc about as long along midline as mesoscutum,
narrowly elevated along posterior margin but without a distinct
punctate groove ; sides of disc rather sharp, front of disc also
sloping rather abruptly to the collar ; surface alutaceous like the
front, the punctures slightly weaker and sparser. Mesonotum
as described for female. Propodeal disc 1.6 x as wide as long,
features much as in female except the transverse striae more
oblique and extending to the transverse carina. Mesopleurum
as in female, but the longitudinal striations somewhat weaker.
Parafi/pe.y. — ARIZONA : 1 9, Avra Val, July- Aug. 1959
(swept from cotton, C. Allen) [USNM] ; 1 9 , Emery Park,
Pima Co., 26 July 1956 (swept from cotton, C. Williams) [UA].
MEXICO : 1 5 , La Aduana, Sonora, same data as allotype
[UCD].
Variation. — The female paratypes are smaller than the type
(LFW 3.0, 3.5 mm) and lack bluish reflections on the head and
thorax. In these two specimens WH is .96 and .97 x LH, WF
EVANS : REVISION OF RHABDEPYRIS 93
1.75 and 1.80 x HE, OOL 1.55 and 1.62 x WOT. In the smaller
specimen, the head is less strongly produced behind the eyes,
the distance from the posterior ocelli to the vertex crest being
slightly less than AVOT ; in this specimen the striations on the
mesopleura are quite weak. The male paratype closely re-
sembles the allotype in size, color, and standard measurements.
Remarks. — Although the male and female here associated
are from localities several hundred miles apart, there are
certain striking features in common, rendering this sex associa-
tion highly probable. These include particularly the striae on
the mesopleura and the very slender scutellar groove.
11. Rhabdepyris (Triciiotepyris) texanus new species
Holotype. — 5 , TEXAS : Big Bend National Park, The
Basin, Chisos Mts., 5400 feet, 8-14 July 1948 (oak honeydew,
H. E. Evans) [MCZ, No. 30,941].
Description of female type. — Length 4.5 mm ; LEW 3.0 mm.
Head and thorax black, the front and thoracic dorsum very
faintly aeneous; abdomen piceous, slightly paler basally and
apically ; palpi and tegulae testaceous ; mandibles bright ferru-
ginous ; scape pale castaneous, fiagellum of this color on the
under side, somewhat darker on the upper side ; legs bright,
pale rufo-castaneous except all coxae strongly infuscated ; wings
subhyaline, veins and stigma amber. Mandibles with five teeth
in an oblique series, the basal three teeth rather broad and
blunt. Clypeus very short, broadly subangulate apically, the
median carina arched in profile. Head slightl}- higher than
wide, WH .98 x LH ; front rather narrow, WE .62 x WH,
1.10 X HE ; ocelli small, front angle of ocellar triangle less than
a right angle ; OOL 1.40 x WOT. Antennal scrobes not carinate ;
temples moderately developed; vertex passing straight across
a distance above eye tops equal to less than half HE. Front
shining, weakly and uniform^ alutaceous, punctures strong,
separated by 1-2 x their own diameters. First four antennal
segments in a ratio of about 35 :9 :8 :11, segment three 1.15 x as
long as its maximum width. (Fig. 6.)
Pronotum 1.5 x as long along midline as mesoscutum, its pos-
terior margin paralleled by a weak, non-foveolate depression ;
surface shining like the front, the punctures slightly more
widely spaced. Mesoscutum and scutellum covered with rather
small punctures, surface moderately shining ; notauli slender,
94 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
diverging and becoming gradually attenuated anteriorly; scu-
tellar groove of moderate width, turned backward and slightly
expanded on each end. Propodeal disc 1.3 x as wide as its
median length ; disc with five strong carinae between which are
four additional weaker carinae, the disc otherwise wholly cov-
ered with fine, transverse striations; side-pieces longitudinally
striolate. Mesopleurum alutaceous but moderately shining, with
scattered small punctures ; upper fovea well formed ; lower fovea
elongate, well formed except on the middle third of the upper
side, where the margin is obsolete (Fig. 27). Front femora
2.5 X as long as wide ; middle tibiae with a series of small spines
above.
Allotype. — $ , MEXICO : 8 mi. SE Elota, Sinaloa, 19 May
1962 (F. D. Parker) [MCZ].
Description of male allotype. — Length 3.7 mm ; LPW 2.8
mm. Entirely black except as follows : palpi and tegulae light
brown; mandibles light brown above and apically, except the
teeth rufous ; scape black, second and third segments light
brown, remainder of flagellum dark brown ; legs dark brownish
fuscous except the middle and hind tibiae medium brown, the
trochanters, front tibiae, and all tarsi testaceous ; wings subhya-
line, veins and stigma light brown. Clypeus obtusely angulate,
with a high median carina which is arched in profile. WH
1.04 X LH; WF .63 x WH, 1.25 x HE; front angle of ocellar
triangle less than a right angle, OOL 1.06 x WOT. Antennal
scrobes not carinate; vertex very weakly arched, distance from
eye tops to vertex crest equal to about one-third x HE. Front
strongly alutaceous, moderately shining, obscurely punctate.
First four antennal segments in a ratio of about 19 :5 :3 :17, seg-
ment four 2.4 X as long as wide, segment eleven 3 x as long as
wide. (Fig. 18.)
Pronotal disc about as long along midline as mesoscutum,
its posterior margin paralleled by a fairly distinct, obscurely
punctate groove ; disc moderately shining, with rather weak
punctures. Mesoscutum alutaceous, obscurely punctate ; notauli
and scutellar groove as described for the female. Propodeal disc
1.2 X as wide as long, with five distinct carinae and two other
weaker carinae close beside the median carina, disc otherwise
with fine transverse striae which are obsolete behind; side-
pieces longitudinally striolate. Mesopleurum alutaceous, indis-
tinctly punctate, lower fovea with its upper margin nearly
complete, but barely perceptible for much of its length.
EVANS : REVISION OP RHABDEPYRIS 95
Paratypes. — TEXAS : 1 9 , Brownsville, 21 November 1911,
palm jungle sweepings [INHS] ; 1 9 , Cedar Lane, Matagorda
Co., 8 September 1928 (J. G. Shaw) [KU]. AKIZONA : 1 9,
S slope Kitt Pk., Quinlin Mts., Pima Co., 22 April 1961 (M. L.
Noller, on composite) [UA] ; 1 5 , S Tucson, 10 June 1962 (F.
Werner) [UA]. MEXICO: 2 S S , Rio Yaqni, near Ciudad
Obregon, Sonora, 19 May 1957 [ENAC] ; 10 S $ , 8 mi. SE
Elota, Sinaloa, same data as allotype except some 19 April
1962 (L. A. Stange) [UCD, CAS] ; 1 5 , 3 mi. N Alpuyeca,
Morelos, 3400 feet, 9 March 1959 (H. E. Evans) [MCZ]
Variation. — In the females LEW varies from 2.5 to 3.1 mm.
The two Texas paratypes have the head and thoracic dorsum
rather strongly shining and without metallic reflections; the
Arizona paratype has these parts rather strongly alutaceous,
moderately shining, faintly aeneous-violaceous. WH varies from
.96 to .98 X LH; WF varies from 1.10 to 1.18 x HE ; propodeum
width varies from 1.25 to 1.30 x median length of disc.
The ten male paratypes from Sinaloa show considerable varia-
tion in size, color, and sculpture. LFW varies from 2.5 to
3.2 mm. In some specimens the second and third antennal seg-
ments are dark like the rest of the antenna, and some variation
can be noted in the color of the flagellum and the tibiae. In
some specimens the front and thoracic dorsum are strongly
alutaceous, almost beaded, while in others these parts are
moderately shining and with distinct small punctures ; in some
specimens the upper margin of the lower mesopleural fovea is
apparently only near the anterior and posterior extremities.
WH varies from 1.01 to 1.05 x LH, AVF from 1.15 to 1.30 x HE ;
the propodeal disc varies from 1.20 to 1.35 x as wide as long.
The two Sonora males show variation in most of these same
characters, but are outstanding because of their large size
(LFW 3.1-3.3 mm) and because of the fact that the second
and third antennal segments are pale castaneous, contrasting
strongly with the remainder of the antennae. On the other
hand, the Morelos male is small (LFW 2.5 mm), although col-
ored like the Sonora males. In this specimen AVH and LH are
equal, WF 1.17 x HE, OOL 1.16 x AVOT. The Arizona paratype
is similar to the allotype in size, color, and standard measure-
ments except that OOL and WOT are equal.
96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
12. Rhabdepyris (Trichotepyris) mexicanus new species
Holotype. — 5 , MEXICO : Nachic, Chiapas [near San Cristo-
bal las Casas], 8000 feet, 27 April 1959 (H. E. Evans) [MCZ,
No. 30,942].
Description of fenmle type. — Length 4.2 mm; LFW 3.0 mm.
Head and thorax black, with faint olive-green reflections, the
pronotal collar dull ferruginous ; propodeum black ; abdomen
black, shining, the last two sternites brownish, the apical tergite
mostly testaceous ; palpi light brown ; mandibles dark ferru-
ginous ; antennae wholly rufo-castaneous except apical segments
weakly infuscated ; tegulae testaceous ; legs light rufo-castaneous
except all coxae infuscated, front pair nearly black; fore wings
lightly tinged with brownish. Mandibles with five teeth, the
basal three teeth somewhat rounded. Clypeus obtusely angulate,
with a median carina which is weakly arched in profile. Head
very slightly wider than high, AVH 1.02 x LH ; WF .65 x WH,
1.31 X HE; OOL 1.10 X WOT. Antennal scrobes not carinate :
vertex broad, nearlj^ straight, distance from eye tops to vertex
crest less than half HE. Front shining although moderately
alutaceous, the punctures sharply defined, separated by 1.5-2.5 x
their own diameters. First four antennal segments in a ratio
of about 26 :9 :8 :9, segment three 1.2 x as long as wide.
Pronotal disc only slightly longer than mesoscutum along
midline, not sharply margined in front or on the sides ; surface
somewhat more alutaceous than the front, its punctures slightly
more widely spaced ; posterior margin paralleled by a weak, non-
foveolate depression. Mesoscutum rather strongly alutaceous,
the punctures small but numerous ; notauli linear, diverging
anteriorly, reaching the anterior margin only as very thin lines;
scutellar groove quite thin medially, roundly expanded on each
end. Propodeal disc about 1.5 x as wide as its median length,
with five distinct carinae and two additional weak carinae
closely paralleling the median carina ; disc irregularly trans-
versely ridged between the carinae, on the sides finely trans-
versely striolate ; declivity with transverse striae, side-pieces with
very fine longitudinal striae. Mesopleurum somewhat alutaceous,
the lower fovea broadly open above. Front femora 2.4 x as
long as wide ; middle tibiae very weakly spinose above.
Allotype. — $ , MEXICO : San Cristobal las Casas, Chiapas,
7500 feet, 26 April 1959 (H. E. Evans) [MCZ].
Description of male allotype. — Length 3.5 mm; LFW 2.5 mm.
EVANS : REVISION OP RHABDEPYRIS 97
Entirely black, the front and thoracic dorsum with a faint
bluish cast ; palpi light brown ; mandibles black except the
teeth ferruginous ; scape black, flagellum dark brown, fading to
medium brown apicallv; coxae and femora dark brown, tibiae
suffused with dark brown basally, legs otherwise light brown ;
fore wings lightly tinged with brownish. Median lobe of elypeus
prominent, somewhat rounded, with a median tooth which is a
continuation of the high, arching median carina. WH 1.07 x LH ;
WF .65 X "WH, 1.35 x HE ; front angle of ocellar triangle less
than a right angle, OOL 1.06 x WOT. Antennal scrobes not
carinate ; vertex broadly rounded off a short distance aboA^e the
eye tops. Front rather strongly alutaceous although moderately
shining, the punctures small and shallow, separated by 2-4 x
their own diameters. First four antennal segments in a ratio
of about 16 :5 :4 :12, segment four 1.9 x as long as wide, segment
eleven 2.2 x as long as wide.
Pronotal disc very slightly longer along midline than meso-
scutum, its posterior margin paralleled by a weakly foveolate
groove ; disc alutaceous and rather weakly shining like the
mesonotum; both pro- and mesonota with numerous but shallow
punctures ; notauli diverging anteriorly ; scutellar groove rather
thin, expanded and deflected backward at each end. Propodeal
disc 1.45 X as wide as its median length ; features of propodeum
essentially as described for female. Mesopleurum alutaceous, the
upper fovea very small, the lower fovea not at all defined on its
upper margin.
Paratypes. — MEXICO : 2 S $ , Canyon de Lobos, near Yau-
tepec, Morelos, 4000 feet, 13-18 March 1959 (H. E. Evans)
[MCZ, USNM].
Variation. — The two male paratypes are dull black, without
any metallic reflections on the head and thorax, and the front
is much more strongly alutaceous and less shining. LFW is
2.5 and 2.7 mm; WH is 1.03 and 1.08 x LH, WF 1.23 and 1.25
X HE, OOL 1.17 and 1.20 x WOT. The propodeum is very
similar to that of the allotype both in shape and in sculpture,
and the mesopleura are also very similar.
13. Rhabdepyris (Trichotepyris) fortunatus new species
Holotype. — 9 , COSTA RICA : Alajuela Prov., Banks of Rio
Fortuna, near La Fortuna, Canton San Carlos, 17 February
1964 (sweeping low vegetation in pasture; H. E. Evans) [MCZ,
No. 30,943].
98 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Description of female iype. — Length 3.7 mm ; LFW 2.4 mm.
Head and thorax black; abdomen black except apical two seg-
ments wholly ruf o-testaceous ; palpi straw-colored; mandibles
testaceous; antennae ruf o-testaceous beneath, segments two and
three wholly of this color, scape and segments 4-13 dark brown-
ish above ; tegulae testaceous ; coxae and femora dark brown,
hind tibiae somewhat inf uscated, legs otherwise light castaneous ;
wings subhyaline, veins and stigma light brown. Mandibles with
five teeth (Fig. 45). Clypeus forming a distinct angle which is
only slightly greater than a right angle ; median carina nearly
straight in profile. WH 1.03 x LH; front moderately wide, WF
M X WH, 1.28 X HE ; OOL 1.23 x WOT. Scrobes not carinate;
vertex very broadly rounded off a short distance above the eye
tops. Front strongly shining, weakly alutaceous, with small
punctures which are fairly close below but very widely spaced
toward the vertex. First four antennal segments in a ratio of
about 25 :7 :5 :8, segment three unusually short, only about .7 x
as long as wide, segment four about 1.2 x as long as wide, seg-
ment eleven barely longer than wide. (Fig. 9.)
Thoracic dorsum slightly more evidently alutaceous than the
front, with small, well-spaced punctures. Posterior margin of
pronotum not paralleled by a groove. Mesoscutum weakly trans-
versly impressed ; notauli diverging and attenuate anteriorly ;
scutellar groove fairly strong, deflected backward and slightly
widened on each side. Propodeal disc 1.3 x as wide as long,
with five discal carinae, transversely ridged between the carinae,
the space between the lateral discals and the lateral carinae
polished and with only weak surface sculpturing; side-pieces
shining, somewhat alutaceous, with a tendency toward the forma-
tion of fine longitudinal striae. Mesopleurum shining, weakly
alutaceous; upper fovea ovoid; lower fovea with its upper
margin broadly obsolete. Front femora 2.1 x as long as wide;
middle tibiae weakly spinose.
Remarks. — This species closely resembles the preceding in
head shape and in most standard measurements. The legs are
darker in color, and the third antennal segment much shorter ;
there are also differences in the sculpturing of the pronotum
and propodeum and in the shape of the scutellar groove. The
species is known only from the type.
EVANS : REVISION OF RHABDEPYRIS 99
1-1. Rhabdepyris (Trichotepyris) lupus new species
Holotype. — $ , MEXICO : Canyon de Lobos, near Yautepee,
Morelos, 4000 feet, 13 April 1959 (H. E. Evans) [MCZ, No.
30,944].
Description of male type. — Length 4.6 mm ; LFAV 3.4 mm.
Black, the head and thoracic dorsum with very faint, dark olive-
green reflections in certain lights, the apical abdominal seg-
ment suffused with light brown ; palpi and tegulae testaceous ;
mandibles rufo-testaceous on the apical half, black basally;
first three antennal segments bright, pale castaneous (except
scape weakly infuscated above), segment four and bej'-ond dark
brown, slightly paler below than above ; legs bright rufo-testa-
ceous except all coxae blackish, the tarsi quite pale ; wings sub-
hyaline, veins and stigma brownish. Clypeus rounded apically,
with a small median angulation formed by the end of the
strong, arched median carina. WH 1.01 x LH; "VVF .62 x WH,
1.16 X HE ; ocellar triangle compact, front angle much less than
a right angle; OOL 1.23 x WOT. Antennal scrobes rather
sharply margined above, but not actually carinate ; vertex pass-
ing straight across a rather short distance above the eye tops.
Front moderately shining although strongly alutaceous, actually
beaded on the lower part, the punctures obscure. First four
antennal segments in a ratio of about 28 :7 :4 :23, segment four
2.4 x as long as wide, segment eleven 2.6 x as long as wide.
Thoracic dorsum more shining and less strongly alutaceous
than the front, also with distinct small punctures ; posterior
margin of pronotum paralleled by a distinct foveolate groove ;
notauli slender throughout, diverging anteriorly; scutellar groove
rather broad, expanded on each side. Propodeal disc measuring
1.33 X as wide as its median length, with seven longitudinal
earinae, the surface with fine, oblique striae except smooth and
polished near the posterior margin ; declivity with transverse
striae, side-pieces with fine longitudinal striae. Mesopleurum
moderately alutaceous, the lower fovea well defined below and
at both ends, but the upper margin broadly discontinuous.
Remarks. — This species is known from the type only. It is
distinctive by virtue of the narrow, beaded front and the brightly
colored legs.
100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
15. RriABDEPYRis (Trichotepyris) carolinianus new species
Holotype. — 9 , SOUTH CAROLINA : Florence, 18 January
1938 (no collector given) [MCZ, No. 30,945].
Description of female type. — Length 6.0 mm; LFAV 3.3 mm.
Head dark olive-green, thoracic dorsum also of this color but
the pronotum with faint coppery reflections, the pleura nearly
black ; propodeum black ; abdomen piceous, except suffused with
reddish brown on the apical third ; palpi testaceous ; mandibles
ferruginous ; antennae light castaneous except upper surface of
flagellum slightly infuscated on the outer two-thirds; tegulae
testaceous ; legs light castaneous except the front coxae strongly
infuscated ; wings very lightly tinged with brownish, more
especially around the radial vein of the fore wing. Body with
abundant whitish to pale golden, short setae. Mandibles with
five teeth, the basal three teeth broad and blunt (Fig. 46).
Clypeus short, broadly rounded except with a faint median
angulation ; median carina very strongly arched in profile.
Head .98 x as wide as high ; front narrow, WF .60 x WH, 1.10
X HE ; OOL 1.35 X WOT ; front angle of oeellar triangle less
than a right angle. Vertex passing straight across a distance
above eye tops equal to less than half HE ; antennal scrobes not
carinate. Front alutaceous, moderately shining, strongly punc-
tate, the punctures separated by 1-2 x their own diameters. First
four antennal segments in a ratio of about 17 :5 :4 :6, segment
three barely longer than its maximum width. (Fig. 7.)
Pronotal disc rounded anteriorly and laterally, about twice
as long along the midline as the mesoscutum, its surface alu-
taceous, moderately shining, with strong punctures which are
separated by 2-3 x their own diameters, posterior margin par-
alleled by a weak, obscurely punctate depression. Mesoscutum
not transversely depressed; notauli diverging anteriorly, of
uniform width throughout ; mesoscutum and scutellum alu-
taceous, punctate throughout, the punctures slightly smaller
than those on the pronotum ; scutellar groove strong, deflected
backward on each side. Propodeal disc 1.25 x as wide as long;
disc with a large, basal, median area filled with longitudinal
earinae connected by cross-carinae ; there are five strong longi-
tudinal carinae, with four additional, somewhat weaker and more
irregular carinae between them, such that nine carinae can be
readily counted ; disc somewhat shining, weakly transversely
striolate on the sides behind; side-pieces longitudinally striolate.
Mesopleurum with lioth upper and lower fovea completely
EVANS : REVISION OF RHABDEPYRIS 101
formed, the lower fovea elongate and only weakly constricted
near the middle (Fig. 28). Front femora 2.4 x as long as wide;
middle tibiae weakly spinose.
Paratype. — 9 , FLORIDA : Gainesville, 5 September 1958
(K. W. Cooper) [USNM].
Variation. — The paratype is smaller (about 5 mm, LFW 2.9
mm ) and is similarly colored except as follows : front with faint
coppery reflections, but thorax without such reflections ; antennae
and legs (except front coxae) darker than in the type, more
or less ferruginous. WH/LH is 1.0; WF is .63 x WH, 1.13 x
HE. Otherwise the resemblance to the type is very close.
Remarks. — I have seen no males which might belong to this
species.
16. Rhabdepyris (Trichotepyris) subaeneus Kieffer
Ehabdepyris subaeneus Kieffer, 1906, Berlin. Ent. Zeitschr., 50: 248 [Type:
9, NICARAGUA: San Marcos (Coll. C. F. Baker) (Pomona College,
Clareniont, Calif.)].
Ehabdepyris (Trichotepyris) subaeneus Kieffer, 1908, Genera Insect., 76:
32.
Ehabdepyris (Ehabdepyris) subaeneus Kieffer, 1914, Das Tierreich, 41:
358.
Description of female type. — Length 4.5 mm; LFW 2.9 mm.
Head and thorax black, with a very weak metallic green luster;
propodeum black; abdomen shining black, apical two segments
and sting-palps suffused with rufous; mandibles wholly rufo-
castaneous; antennae uniformly castaneous; tegulae testaceous;
coxae black, dark brown blotches on the outer side of the front
and hind femora, legs otherwise bright yellowish brown ; wings
hyaline. Mandibles fairly broad, with five teeth in an oblique
series, basal three teeth somewhat rounded, the basal tooth
wider than third and fourth teeth (Fig. 47). Clypeus obtusely
subangulate apically, with a high median carina which is arched
in profile. First four antennal segments in a ratio of about
35:9:9:11, segment three 1.1 x as long as thick, segment eleven
1.2 X as long as thick. Antennae arising well below bottoms of
eyes ; scrobes not margined by carinae. Front shining, uniformly
but rather weakly alutaceous, punctures strong, separated by
about or slightly more than their own diameters. Head higher
than wide, WH 0.92 x LH ; front narrow, WF .59 x WH, .95
X HE ; distance from eye tops to vertex crest equal to only
about .3 X HE. Ocelli small, well separated, front angle of
102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
ocellar triangle less than a right angle; OOL 1.2 x WOT; front
ocellus below a line drawn between eye tops, posterior ocelli
separated from vertex crest by about twice their own diameters.
Pronotum with disc separated from collar by a short, oblique
anterior face ; disc with fairly sharp margins in front and on
the sides, but without carinae, also without a groove paralleling
the posterior margin (merely with a barely depressed line of
punctures) ; disc somewhat shining, alutaceous, punctures some-
what weaker and more widely spaced than on the front. Meso-
scutum strongly alutaceous, rather weakly shining, punctures
small but numerous on posterior half; notauli slender, strongly
diverging anteriorly, absent on anterior third. Scutellar disc
wholly alutaceous, moderately shining ; basal groove fairly broad,
somewhat turned backward on sides. Propodeal disc 1.25 x as
wide as long, with strong lateral and posterior carinae ; disc
with seven carinae, but the two carinae closely paralleling the
median carina rather weak, only the median carina reaching the
posterior margin ; disc otherwise transversely striate, the stria-
tions obsolescent postero-laterally, where the disc is smooth
and polished. Mesopleurum with upper and lower foveae dis-
tinct, the latter not divided although somewhat constricted near
the middle. Front femora 2.1 x as long as wide.
Remarks. — I have seen no specimens of this species other
than the type.
17. Rhabdepyris (Trichotepyris) angusticeps new species
Holotype. — 9 , ARIZONA : Tucson, 18 June 1938 (R. H.
Crandall) [MCZ, No. 30,946].
Description of female type. — Length 5 mm ; LFW 3 mm.
Black, except as follows: pronotal collar ruf o-testaceous ; abdo-
men suffused with reddish brown on the apical third, the
last tergite pale castaneous; palpi straw-colored; mandibles fer-
ruginous; antennae light ruf o-castaneous ; tegulae testaceous;
legs light castaneous except all coxae blackish, front and hind
femora partially suffused with black on their outer faces; wings
subhyaline, weakly suffused with brown around the radial vein
of the fore wing, the veins and stigma amber. Mandibles large,
their lower margin weakly arched, the five teeth in a strongly
oblique series, the basal three teeth very broad and blunt
(Fig. 48). Clypeus very short, very broadly, obtusely angulate,
the median carina arched in profile. Head unusually elongate,
EVANS : REVISION OF RIIABDEPYRIS 103
.91 X as wide as high; WF .66 x WH, 1.28 x HE; ocelli small,
front angle of ocellar triangle less than a right angle, OOL 1.75
X WOT. Sides of head gradually convergent behind the eyes, the
vertex relatively narrow, very weakly concave in anterior view;
distance from eye tops to vertex crest equal to about .7 x HE.
Antennal scrobes not carinate. Front strongly shining, very
weakly alutaceous, with strong punctures which are separated
by 1.5-3 X their own diameters. Scape flattened, curved ; first
four antennal segments in a ratio of about 4:1:1:1; segment
three 1.3 x as long as its maximum width. (Fig. 8.)
Pronotal disc abruptly rounded anteriorly and on the sides,
along the midline 2.4 x as long as the mesoscutum ; pro- and
mesonota more strongly alutaceous than front, moderately shin-
ing, with small, rather evenly distributed punctures ; posterior
margin of pronotum not paralleled by a punctate groove. Meso-
scutum not depressed; notauli linear, strongly diverging toward
the front ; scutellar groove of moderate breadth, deflected back-
ward on each side. Propodeal disc 1.15 x as wide as long; disc
with seven longitudinal carinae, the two carinae close beside
the median carina somewhat weaker than the others; surface
weakly transversely striolate except on the posterior part ; sub-
lateral carinae absent ; side-pieces strongly longitudinally strio-
late. Mesopleurum moderately alutaceous ; upper fovea complete ;
lower fovea elongate, its upper margin obsolete for much of its
length. Front femora broad and flat, measuring 1.9 x as long
as wide ; middle tibiae with a series of small spines on the upper
surface.
Remarks. — The tips of all the tarsi of the type and only
known specimen are missing.
18. Rhabdepyris (Trichotepyris) olivaceus new species
Ilolotype. — 5 , PANAMA : Barro Colorado Island, Canal
Zone, 28 March 1955 (C. W. Rettenmeyer) [USNM, No. 67,539].
Description of male type. — Length 3.7 mm; LFW 2.5 mm.
Head and thorax black, the front and thoracic dorsum Avith
olive-green reflections; abdomen dark brown, shining, slightly
paler toward the base and apex; palpi and tegulae testaceous;
mandibles testaceous except blackish toward the base, the teeth
rufous; antennae dark brown, the scape partially suffused with
black, the flagellum paler beneath than above, antennal seg-
ments 2 and 3 tending to be slightly paler than the others; legs
104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
testaceous except the front coxae rather strongly infuscated,
the femora and the remaining coxae suffused with brownish;
wings subhyaline, veins and stigma light brown. Median lobe of
clypeus prominent, obtusely angulate except the median point
drawn out to an acute angle ; median ridge high and arched.
WH/LH = 1.0; WF .62 x WH, 1.14 x HE; ocellar triangle
compact, front angle less than a right angle; OOL 1.16 x AVOT.
Antennal scrobes margined by a rounded, ill-defined carina
which does not nearly reach the eye margins; vertex elevated
only slightly above the eye tops. Front uniformly and rather
strongly beaded, somewhat shining, punctures obscure. First
four antennal segments in a ratio of about 20 :5 :3 :15, segment
four 2.5 X as long as wide, segment eleven twice as long as wide.
Pronotal disc beaded like the front, quite sharply margined
on the sides, the posterior margin paralleled by a row of fairly
strong foveae. Mesoscutum obscurely punctate, strongly alu-
taceous, the notauli arching, divergent toward the front ; scu-
tellar groove relatively broad, roundly expanded and deflected
baclavard on each end. Propodeal disc measuring 1.15 x as wide
as long; disc with five carinae, between which it is irregularly
transversely ridged, outside the carinae wholly covered with fine
transverse striations; declivity and side pieces covered with
rather weak, irregular striations. Mesopleurum somewhat alu-
taceous although strongly shining, the upper margin of the
lower fovea not at all defined. Claws strongly dentate.
Paratypes. — PANAMA: 1 $, same data as type except 11
February 1955 [KU]. COLOMBIA: 1 5 , La Cumbre, 6000
feet, 23 May 1914 (H. S. Parish) [MCZ].
Variation. — The Panama paratype resembles the type closely
in every respect. The Colombia paratype is very slightly larger
(LFW 2.7 mm) and has the abdomen wholly castaneous, con-
trasting to the head and thorax, and the front is more shining
and with a faint bluish cast (the pronotum, however, is olive-
green). Li this specimen WF is 1.25 x HE, OOL 1.40 x WOT.
In every other respect the resemblance to the type is so close
that there seems little question of their eonspecificity.
19. Rhabdepyris (Trichotepyris) plaumanni new species
Holotype. — 9 , BRAZIL : Nova Teutonia, Santa Catarina,
7 July 1937 (F. Plaumann) [BMNH].
Description of female type. — Length 3.7 mm ; LFW 2.2 mm.
EVANS : REVISION OF RIIABDEPYRIS 105
Head and thorax black, except pronotal collar dark ferruginous ;
abdomen piceous, except the venter and apical two tergites dark
reddish brown ; palpi testaceous ; mandibles ruf o-testaceous, some-
what infuscated toward the base ; antennae light castaneous
except scape and apical few segments somewhat infuscated ; coxae
and femora dark brown, hind tibiae medium brown, legs other-
wise testaceous ; fore wings lightly tinged with brownish, veins
and stigma light brown. Mandibles with five teeth, the basal
three teeth small and rather blunt. Clypeus short and rather
rounded apically, but with a small median angulation formed
by the end of the high, arched median carina. Head very slightl}'
wider than high, WH 1.02 x LH ; WF .60 x WH, 1.10 x HE ;
OOL 1.20 X WOT. Antennae arising well below bottoms of
eyes, the scrobes not carinate ; vertex broadly rounded off a
very short distance above the eye tops. Front shining, weakly
alutaceous, strongly punctate, the punctures for the most part
separated by 1-2 x their own diameters. Antennae rather short,
the first four segments in a ratio of about 20 :7 :5 :6, segment
three slightly wider than long, segments four through twelve
each about as long as thick.
Pronotal disc rather abruptly rounded anteriorly and lat-
erally, measuring along the midline about 1.6 x the length of
the mesoscutum ; surface alutaceous, moderately shining, punc-
tate ; posterior margin paralleled by only a very weak groove.
Mesoscutum alutaceous and with small punctures, the notauli
slightly attenuated and diverging toward the front ; scutellar
groove deep but quite thin in the middle, much expanded on
each side. Propodeal disc 1.2 x as wide as its median length;
disc with 7 carinae, between which it is irregularly transversely
ridged; disc laterad of the carina with fine striations which
extend to the sublateral carinae; declivity with transverse striae
and a median carina ; side pieces with longitudinal striations
on the basal two-thirds, apically merely alutaceous. Mesopleurum
moderately shining and with shallow punctures, the lower fovea
not closed above. Front femora 2.3 x as long as wide; middle
tibiae not spinose.
Allotijpc. — $ , BRAZIL : same data except 2.9 December
1938 [BMNH].
Description of male allotype. — Length 3.6 mm ; LFW 2.3 mm.
Head and thorax black, abdomen dark reddish brown ; palpi
testaceous ; mandibles rufo-testaceous apically and on the upper
margin, elsewhere blackish ; scape black, flagellum dark brown ;
106 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
coxae and femora dark brown, middle and hind tibiae weakly
suffused with brown, legs otherwise testaceous; wings subhya-
line. Clypeus forming a distinct angle medially, the angulation
slightly greater than a right angle except at the mid-point, where
it is acute; median carina very high, strongly arched. WH .98
X LH; WF .63 X WH, 1.22 x HE; ocelli in a compact triangle,
OOL 1.15 X WOT. Autennal scrobes not carinate; vertex evenly
rounded off a short distance above the eye tops. Front uniformly
and rather strongly alutaceous, although somewhat shining,
punctures small and shallow^, separated by 2-3 x their own
diameters. First four antennal segments in a ratio of about
19:6:3:15, segment four 2.5 x as long as wide, segment elcA^en
2.4 x as long as wide.
Thoracic dorsum uniformly alutaceous, moderately shining,
obscurely punctate ; pronotal disc rather abruptly rounded on
front and sides, its posterior margin paralleled by a strong,
foveolate groove ; scutellar groove as described for female. Pro-
podeal disc measuring 1.15 x as wide as its median length, with
five discal carinae, otherwise wholly covered with fine trans-
verse ridges; side pieces longitudinally striate. Mesopleurum
somewhat alutaceous, obscurely punctate, the lower fovea com-
pletelj^ open above.
Paratype. — 1 S , BRAZIL : same data as type except June
1962 [MCZ].
Variation. — The paratype is only 3.0 mm long, LFW 2.0
mm. The mandibles and legs are somewhat darker than in the
allotype, but there are no noteworthy differences in structure.
WF is 1.18 X HE, OOL 1.30 x WOT, the propodeal disc 1.2 x
as wide as long.
PULCHRIPENNIS SPECIES-GROUP
This group includes four specialized members of this sub-
genus, all of them small, slender species exhibiting more or
less unusual color patterns (at least in the females). The wdngs
are conspicuously banded or, in one species, much abbreviated.
The middle tibiae are weakly if at all spinose, the eyes relatively
large and very densely hairy.
20. Rhabdepyris (Trichotepyris) pulchripennis new species
Holotype. — 5 , COSTA RICA : Turrialba, 24 June 1949
(K. W. Cooper) [USNM, No. 67,540].
EVANS : REVISION OF RHABDEPYRIS 107
Description of female type. — Length 3.7 mm; LFW 2.6 mm.
Head and thorax black, with extremely faint coppery reflections,
the pronotal collar ferruginous; propodeum and abdomen shin-
ing black, the tip of the abdomen suffused with reddish brown;
palpi and mandibles testaceous, the antennae also of this color
except the apical segment tipped with fuscous; tegulae tes-
taceous; legs bright testaceous except the front and middle
femora weakly and in part suffused with fuscous ; wings sub-
hyaline except the fore wing with a fuscous cloud over the
outer half of the median and submedian cells and a second
fuscous cloud in and below the marginal cell ; veins brown except
yellowish at extreme base of wing. Body with fairly dense,
short, golden brown setae. Mandibles with five teeth, the basal
three teeth small and sharp. Clypeus sharply, obtusely angulate.
Head 1.03 x as wide as high; front narrow, WF .62 x WH,
1.08 X HE ; OOL 1.45 x WOT. Front alutaceous, rather weakly
shining, with small punctures which are separated, for the
most part, by 1.5-2.5 x their own diameters. Vertex rounded
off a short distance above the eye tops ; autennal scrobes not
carinate. First four antennal segments in a ratio of about 11:
3 :2 :3, segment three considerably wider than long.
Pronotal disc 1.5 x as long medially as mesoscutum, alu-
taceous and punctate much like the front, without a line of
punctures paralleling the posterior margin. Notauli diverging
and attenuated anteriorly ; mesoscutum weakly depressed laterad
of notauli. Scutellar groove turned backward and considerably
widened on each side. Propodeal disc 1.05 x as wide as long;
disc with five strong carinae, the median carina complete and
the others nearly so, transversely ridged between the carinae;
posterior lateral part of disc smooth and polished; side-pieces
not striolate, merely weakly aciculate. Mesopleurum with the
f oveae incompletely defined. Front femora 2.0 x as long as wide ;
middle tibiae spinose.
Remarks. — This species is known only from the type.
21. Rhabdepyris (Trichotepyris) iridescens new species
Holotype. — $ , MEXICO : 4 mi. E Cuernavaca, Morelos,
6000 feet, 25 June 1959 (H. E. Evans) [MCZ, No. 30,949].
Description of female type. — Length 3.6 mm ; LFW 2.6 mm.
Head black, front with strong violet reflections; thorax black,
the dorsum with coppery reflections, the pronotal collar ferru-
ginous; propodeum and abdomen shining black, the extreme tip
108 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
of the abdomen dull brownish ; palpi and mandibles testaceous,
antennae of this color but the outer side of the fiagellum some-
what darker, the apical five segments distinctly infuscated ;
tegulae testaceous ; legs wholly bright ruf o-testaceous ; wings
hyaline except the fore wing with a fuscous cloud over the outer
half of the median and submedian cells and a second cloud
around and below the radial vein ; wing veins brown where the
membrane is clouded, elsewhere (including the stigma) tes-
taceous. Mandibles with five teeth, the basal three teeth short
and rounded. Clypeus obtusely angulate. Head .92 x as wide
as high; front very narrow, WF .59 x WH, .93 x HE; OOL
1.40 X WOT. Front strongly alutaeeous although somewhat
shining, punctures very shallow and inconspicuous. Vertex
rounded off a short distance above eye tops; antennal scrobes
not carinate. First four antennal segments in a ratio of about
14:4:4:5, segment three slightly longer than wide. (Fig. 10.)
Pronotal disc 1.2 x as long medially as the mesoscutum,
alutaeeous and obscurely punctate like the front, with a line of
small foveae paralleling the posterior margin. Notauli linear,
arcuately diverging anteriorly ; mesoscutum not notably de-
pressed on the sides. Scutellar groove slightly broadened and
deflected backward on each side. Propodeal disc 1.1 x as wide
as long ; disc with seven strong carinae, finely transversely striate
throughout, but the striae obsolescent on the sides behind; sub-
lateral carina not distinct ; posterior angles f oveolate ; side-pieces
shining, weakly aciculate. Mesopleurum with the foveae in-
completely defined. Front femora 2.3 x as long as wide ; middle
tibiae only very weakly spinose.
Remarl^s. ■ — This remarkably colored species is known only
from the type, which was collected at honeydew on the foliage
of walnut trees.
22. Khabdepyris (Trichotepyris) cupreolus new species
Holotype. — 9 , BRAZIL : Nova Teutonia, Santa Catarina,
July 1953 (F. Plaumann) [MCZ, No. 30,950].
Description of female type. — Length 3.8 mm; LFW 2.4 mm.
Head and thorax black, the front with coppery reflections, the
thoracic dorsum with obscure coppery to olive-green reflections,
the pronotal collar ferruginous ; propodeum black ; abdomen
piceous, suffused with light brown tow^ard the apex, the last
segment testaceous ; palpi straw-colored ; mandibles rufo-testa-
ceous, slightly darker on the apical half than basally; antennae
EVANS : REVISION OF RHABDEPYRIS 109
wholly testaceous except all segments weakly infuscated on the
upper side ; tegulae testaceous ; legs bright testaceous except
front coxae mostly fuscous, front femora partially infuscated on
the outer surface ; wings faintly luteous, fore wing with two
broad fuscous bands separated by a narrow sub-hyaline band
which includes the stigma, as in the preceding two species.
Mandibles with five teeth, the basal three teeth rather small.
Clypeus broadly, obtusely subangulate, the median ridge arched
in profile. Head .93 x as wide as high ; front very narrow, WF
.57 X AVH, .95 x HE ; ocelli in a compact triangle, OOL 1.25
x WOT. Antennal scrobes not carinate; vertex rounded off a
short distance above the eye tops. Front strongly alutaceous,
rather weakly shining, with shallow punctures which are sep-
arated by 1-3 X their own diameters. First four antennal seg-
ments in a ratio of about 27 :8 :6 :9, segments three and eleven
barely longer than wide, segment four about 1.3 x as long as
wide.
Thoracic dorsum strongly alutaceous, weakly shining, the
punctures obscure ; pronotum with a strong foveolate groove
paralleling the posterior margin ; notauli linear, diverging an-
teriorly; scutellar groove rather wide, roundly expanded at
each end. Propodeal disc 1.05 x as wide as long; disc with five
longitudinal carinae and with two additional weak carinae close
beside the median carinae, otherwise wholly covered with trans-
verse striae ; side-pieces wholly covered with fine longitudinal
striae which tend to curve upward posteriorly. Mesopleurum
strongly alutaceous, the lower fovea elongate, fairly well defined.
Front femora slender, 2.4 x as long as wide ; middle tibiae not
spinose.
Parafypcs. — BRAZIL: 1 9, same data as type [Coll. G. R.
Ferguson] ; 1 9 , Bocaiuva, Parana, 1000 meters elev., May
1963 (F. Plaumann) [MCZ].
Variation. — The topotypic paratype is of the same size as
the type, but differs slightly in color : the coppery reflections
of the front and pronotum are rather strong, and the front also
has rather strong violet reflections, especially anteriorly ; the
front coxae are only slightly infuscated, and the front femora
are wholly without infuscation. In this specimen WH/LH is
.94, WF/HE is .96, and OOL/WOT is 1.35. The Bocaiuva para-
type is larger (LFW 2.9 mm) but colored like the type. In this
specimen WH/LH is .92, WF/HE is .88, and" OOL/WOT
is 1.33.
110 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
23. Rhabdepyris (Trichotepyris) amabilis Fonts
Rhahdepyris amaiilis Fouts, 1927, Proe. Ent. Soc. Washington, 29: 165-16(i
[Type: 9, MAEYLAND: Glen Echo, July (R. M. Fouts) (Coll.
Fouts)]. — Muesebeck aJid Walkley, 1951, U.S. Dept. Agri., Monogr.
2, p. 72&.
Description of female type. — Length 3.5 mm ; wings very
short, fore wing reaching slightly past the middle of the pro-
podeum, LFW 0.55 mm. Head and thorax black, faintly aeneous,
except the pronotal collar testaceous; propodeum black; abdo-
men piceous, fading to dark brown apically; palpi light brown;
mandibles and basal three antennal segments pale castaneous,
rest of antenna dull castaneous below, dark brownish above ;
tegulae testaceous ; legs bright, pale castaneous except front
coxae moderately infuscated ; wings hyaline, the veins testaceous.
Body clothed with short, golden setae. Mandibles with five
teeth, the basal three teeth subequal except the basal tooth more
rounded than the others. Clypeus with its median lobe moder-
ately prominent, rounded, with a strong median carina which
is arched in profile. Head relatively long and slender, only
1.2 X as wide as maximum width of thorax (across mesothorax) ;
WH .93 X LH ; WF .60 x WH, 1.05 x HE ; ocelli small, front
angle of ocellar triangle less than a right angle ; OOL 1.80 x
WOT. Antennal scrobes not carinate ; temples not well de-
veloped ; vertex rounded off a distance above eye tops equal to
slightly more than half HE. Front alutaceous. moderately .shin-
ing, punctures moderately strong, small, separated by 2-3 x their
own diameters. First four antennal segments in a ratio of
about 31 :7 :7 :9, segment three about as long as its apical width.
Pronotum quite long, the disc nearly 3 x the length of the
mesoscutum. the punctures small, more widely spaced than on
the front ; posterior margin paralleled by only a very faint,
non-foveolate groove. Mesoscutum short, transversely depressed,
especially laterally, the surface rather strongly alutaceous, ob-
scurely punctate ; notauli slender, diverging anteriorly ; scutel-
lar groove moderately wide, deflected backward but not enlarged
on each end. Propodeal disc about as long as wide, the median
carina complete, paralleled by two other incomplete carinae, the
surface reticulate between and beside these carinae and tending
to form two additional weak carinae close beside the median
carina ; elsewhere smooth, weakly alutaceous, with a few weak
longitudinal striae at the extreme upper basal angles. Meso-
pleurum rather irregularly pitted, the loAver fovea not defined
EVANS : REVISION OP RHABDEPYRIS 111
on its upper side. Front femora 2.3 x as long as wide; middle
tibiae not spinose above. Fore wings slender, nearly 4 x as long
as wide, with three longitudinal veins which unite apieally to
form two cells ; hind wings even more slender, reaching to
about the apex of the cells of the fore wings.
Plesiallotype. — S , VIRGINIA : Falls Church, 2 August (N.
Banks) [MCZ].
Description of male plesiallotype. — Length 3.4 mm; LFW
2.3 mm. Head and thorax piceous, abdomen dark reddish
brown, paler at base and apex ; palpi straw-colored ; mandibles
rufo-testaceous, darker basally ; antennae uniformly medium
brown except the second segment slightly paler, the scape weakly
infuscated above ; tegulae light brown ; coxae and femora dark
brown, tibiae medium brown, trochanters and tarsi light brown ;
wings hyaline, veins and stigma light brown. Clypeus with a
prominent median lobe which is somewhat rounded but with a
median tooth, which is at the end of the high, arched median
carina. WH/LH = 1.0; WF .64 x WH, 1.35 x HE; ocelli in
a small triangle, the front angle less than a right angle, OOL
1.45 X WOT. Antennal scrobes not carinate; vertex broadly
rounded off a distance above eye tops equal to about half HE.
Front somewhat shining although very strongly alutaceous, more
or less beaded, obscurely punctate. First four antennal segments
in a ratio of about 21 :6 :5 :15, segment four 2.5 x as long as
wide, segment eleven 3 x as long as thick.
Pronotal disc about 1.2 x as long as mesoscutum, measured
along midline, its posterior margin paralleled by a series of
small but well defined foveae, the disc otherwise obscurely punc-
tate, alutaceous but by no means beaded like the front. Surface
of mesoscutum about like that of the pronotum, the notauli
linear, diverging toward the front ; scutellar groove arching
backward on the sides, wider laterally than medially. Propodeal
disc 1.35 X as wide as long; disc with five carinae, the most
lateral ones rather short ; surface reticulate between the carinae,
elsewhere finely transversely striolate, the striae obsolescent pos-
teriorly; declivity with transverse striae and an incomplete
median carina ; side-pieces alutaceous, with fine longitudinal
striae toward the front. Mesopleurum alutaceous, obscurely
punctate, the upper margin of the lower fovea not well defined.
Specimens examined. — MASSACHUSETTS: 1 9, Lexing-
ton, 28 August 1955 (mesic forest, W. L. Brown) [MCZ].
MARYLAND: 1 9, Glen Echo [type, coll. Fonts]. DISTRICT
112
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
OF COLUMBIA: 1 9, \Yashington, September (J. C. Brid-
well) [USNM]. VIRGINIA: 1 9,3 S £, Falls Church, 2-11
August (N. Banks) [MCZ]. FLORIDA: 2 9 9, Gainesville,
May, October 1958, 1959 (K. W. Cooper) [USNM].
Variation. — The three males vary slightly in size (LFW
2.2-2.5 mm) and show some variation in the degree of infusea-
tion of the legs and antennae. In the two specimens other than
the plesiallotype the lateral ocelli are somewhat less far removed
from the eyes, OOL being 1.25 x WOT in both specimens. WF
varies from 1.30 to 1.35 x HE ; the propodeal disc varies from
1.2 to 1.35 X as wide as long.
The available females show so much variation that one is
tempted to place them in several subspecies. However, since
only six specimens are at hand, and since some of the variation
appears allometric (e.g. relatively longer wings in the larger
specimens) and much of it clinal (e.g. brighter colors and nar-
rower front toward the south), it would be premature to erect
subspecies at this time. Some of the variation is tabulated below
(Table IV). In spite of this variation, it should be pointed out
that the entire series is much alike with respect to form of the
mandibles and clypeus and features of the thoracic dorsum
and propodeum. The two Florida specimens are strikingly
colored, having the head ferruginous except the central part of
the front black, the thorax and propodeum ferruginous except
the mesoseutum spotted with black laterad of the notauli, and
the first two abdominal segments maculated with ferruginous
basally.
TABLE IV. VARIATION IN SIX FEMALES OF RHABDEPYRIS AMABILIS FOUTS
Locality
Length
Wing length
Color of
Color head
WH/LH
WF/HE
OOL/W(
(mm.)
(mm.)
legs
and thorax
Massachusetts
3.3
.45
brownish
black, slightly
aeneous
.93
1.13
1.75
Maryland
3.5
.55
testaceous
as above
.93
1.05
1.80
Dist. Columbia
4.7
.75
as above
as above
.94
1.05
1.65
Virginia
4.3
.60
as above
as above
.95
1.12
1.70
Florida
3.3
.50
light brown
mostly
ferruginous
.97
0.90
1.40
Florida
3.0
.40
as above
as above
.94
0.96
1.55
EVANS : REVISION OF RHABDEPYRIS 113
C. Subgenus Chlorepyris Kieffer
Chlorepyris Kieffer, 1913, Boll. Lab. Zool. Portiei, 7: 108 (type species: C.
semiviridis Kieffer 1913 [= viridissimus Kieffer 1911] ; designated by
Kieffer, 1914). —Kieffer, 1914, Das Tierreich, 41: 412-416 (key to
spp., Oriental and jSTeotropical regions). — Kurian, 1955, Agra Univ.
Jour. Res., 4: 101-105 (Oriental spp.).
Sxibgeneric characters. — Small to fairly large bethylids (up
to 10 mm) ; black, head and thorax often weakly to strongly
greenish or bluish, abdomen sometimes in part rufous; eyes
glabrous ; body clothed with fine, pale setae which are mostly
subappressed except more erect and bristling on the abdomen,
temples, and propleura ; middle tibiae of female spinose, hind
tibiae of female and tibiae of male also sometimes spinose.
Mandibles of male terminating in five small, sharp teeth, those
of female with from three to five apical teeth and also often with
a subapical tooth on the lower margin ; malar space short, about
as in Trichotepyris; antennal scrobes not margined in the known
American species ; third antennal segment of male of very
variable length, but always quite distinctly separated from the
fourth segment. Pronotum rather long, its posterior margin
not paralleled by a foveolate groove ; mesoscutum more or less
transversely depressed in the females of most species ; notauli
strong, reaching anterior margin usually only as very fine
lines; scutellar groove slender and arcuate or sometimes rather
straight and much broadened on each side, in a few species
merely forming a thin line connecting large lateral pits; pro-
podeal disc with from three to seven longitudinal carinae, its
postero-lateral angles always strongly foveolate. Claws dentate,
bifid, or trifid.
Remarks. — As mentioned in the introduction, Chlorepyris
is here redefined in a somewhat broader sense than Kieffer 's.
There are three species-groups, of which the third (the viridis-
simus group) more or less approximates Chlorepyris in Kieffer 's
sense.
KEY TO SPECIES OF SUBGENUS CHLOEEPYRIS
Females
1. Lower mesopleural fovea well defined, fully outlined or with the
upper margin indistinct on the middle fourth (Figs. 29, 30) ; man-
dibles with the preapical tooth on the lower margin small and
inconspicuous (Fig. 49) ; species of black, non-metallic coloration
{Lobatifrons group) 2
114 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Lower mesopleural fovea poorly defined, its upper margin completely
undefined for at least the middle third (Figs. 31-36) ; mandibles with
a strong, sharp tooth on the lower margin (Figs. 50-54) ; color
variable 4
2. Wings strongly suffused with yellowish; LFW at least 4.5 mm; front
broad, WF 1.25-1.50 x HE (Fig. 11); OOL more than 2.0 x WOT;
lower mesopleural fovea strongly outlined (Fig. 29)
24. htteipennis n. sp.
Wings somewhat clouded, barely if at all yellowish; LFW 3.0-3.8 mm;
OOL less than 2.0 x WOT, sometimes only slightly so ; lower meso-
pleural fovea not always fully defined above (Fig. 30) 3
3. Mandibles mostly pale ferruginous, legs also of this color except
femora sometimes darker, coxae fuscous; ocellar triangle fairly
broad, front angle only slightly less than a right angle
27. quinquelineatus Kieffer
Mandibles and legs mostly black ; front with some strong punctures on
the lower part, the punctures smaller and much more •\\adely spaced
above; ocellar triangle compact, front angle much less than a
right angle (Fig. 12) 28. nigerrimus n. sp.
4. Head well developed in front of eyes, the antennae inserted well below
bottoms of eyes (WH less than 1.05 x LH, rarely up to 1.08 x LH)
(Figs 13, 14) ; upper mesopleural fovea small, ovoid, at most about
2.5 X as long as wide (Figs. 31-33) ; claws dentate (Muscariiis
group ) 5
Head very short in front of eyes, antennae arising only slightly below
bottoms of eyes, clypeus barely protruding beyond antennal bases
(WH at least 1.07 x LH, up to 1.20 x LH (Figs. 15, 16) ; upper
mesopleural fovea elongate, 2-5 x as long as wide (Figs. 34-36) ;
head and thorax with metallic blue or green coloration (Viridissi-
mus group) 9
5. Head and thorax black, without metallic colors; scutellar groove very
slender, curved backward but barely widened at each end; front
femora strongly incrassate 6
Head and thorax weakly to strongly reflecting metallic blue or green ;
scutellar groove much widened at each end, appearing as a slender
line connecting two pits 7
6. Head and thoracic dorsum strongly shining, obscurely alutaceous;
upper margin of lower mesopleural fovea defined in front and behind
(Fig. 31) ; OOL 1.40-1.55 x WOT; LFW 3.8-4.2 mm
29. muscariiis (Westwood)
Head and thoracic dorsum (especially the latter) moderately shining,
quite distinctly alutaceous; upper margin of lower mesopleural
fovea not at all defined (Fig. 32); OOL 1.20-1.33 x WOT; LFW
3.2-3.3 mm 31. origenus Kieffer
7. Legs rufous except the coxae black; length about 4 nmi; head
metallic blue, thoracic dorsum green with bluish reflections
35. metallictis Kieffer
EVANS : REVISION OF RHABDEPYRIS 115
Legs with femora at least partially fuscous and front femora usually
with metallic reflections ; length 4.5-7.0 mm 8
8. Head and thorax with dull olive-green reflections; propodeal disc
1.25-1.36 X as wide as long; front with very small punctures which
are separated by several times their own diameters
33. vireseens n. sp.
Head and thorax bright bluish green; propodeal disc only 1.15 x as
wide as long; front with strong punctures which are separated by 2-3
X their own diameters 34. viridis (Cameron)
9. Propodeum black, without metallic reflections, contrasting to head
and thoracic dorsum, which are greenish 10
Propodeum l^luish, blue-green, or violet, either of the same color or
contrasting to the thoracic dorsum 11
10. Head and thoracic dorsum with dull olive-green reflections; propodeum
elongate, 1.25-1.28 x as wide as long; tooth of claws acute, sloping
outwards slightly; LFW 3.0-3.5 mm 36. subviridis (Kieffer)
Head and thorax intense, brilliant green; propodeum transverse, 1.38-
1.45 X as wide as long; tooth of claws somewhat blunt, sloping
outward strongly (Fig. 66) ; LFW 4.0-5.5 mm
38. viridissimus (Kieffer)
11. Legs largely ferruginous except for coxae and hind femora; upper
mesopleural fovea only about twice as long as high ; propodeum
violaceous, contrasting to the brilliant green head and thorax ....
39. fulgens (Brues)
Legs variable, but the front femora always dark and with strong
bluish reflections; upper mesopleural fovea at least 3 x as long as
high (Figs. 35, 36) ; propodeum blue or blue-green, contrasting at
most slightly with the head and thoracic dorsum, which are more
or less bluish, blue-green, or violaceous 12
12. Legs ferruginous except for the coxae and the front femora; LFW
3.5 mm; front sparsely punctate, mesoscutum with punctures only
along the notauli and parapsidal furrows; apical .4 of abdomen
ferruginous 40. tricolor n. sp.
Legs mostly dark, the hind femora black and with bluish reflections
like the front femora; LFW 4.5-4.7 mm; front and thoracic dorsum
wholly covered with small punctures; tip of abdomen weakly suf-
fused with brownish 41. violaceus n. sp.
Males
1. Third antennal segment not reduced, distinctly longer than wide, much
longer than second segment and nearly or quite as long as fourth
(Figs. 71, 72) ; lower mesopleural fovea fully outlined or the upper
margin indistinct on the middle fourth (as in Figs. 29, 30) (Lobati-
frons group) 2
Third antennal segment small, transverse, wider than long, much
shorter than fourth segment (Figs. 19-24, 73) ; lower mesopleural
fovea broadly incomplete above (Figs. 37-42) 5
116 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
2. Propodeum with five discal carinae 3
Propodeum with seven discal carinae, five parallel carinae medially
and two more oblique carinae laterad of these 4
3. Lower mesopleural fovea fully enclosed (as in Fig. 29) ; LFW 3.3-4.0
mm; front evidently alutaceous 24. luteipennis n. sp.
Lower mesopleural fovea with its upper margin indistinct on the
middle third (as in Fig. 30); LFW under 3.0 mm; front strongly
polished, at most obscurely alutaceous . . . .27. quinquelineatus Kieffer
4. Propodeum very short, transverse, 1.9 x as wide as its median length;
tip of abdomen suffused with dull ferruginous
25. lobatifrons Kieffer
Propodeum more elongate, measuring 1.6-1.7 x as wide as long; apical
third of abdomen bright ferruginous . . . .26. scptemlincatus Kieffer
5. Upper mesopleural fovea small, completely enclosed (Fig. 37) ; species
of black coloration 6
Upper mesopleural fovea elongate, open behind (Figs. 38-42) ; black
or with metallic colors 7
6. Head very broad (WH 1.12-1.20 x LH) (Fig. 19); front polished
and only very obscurely punctate, mesopleura impunctate or nearly so
29. muscarius (Westwood)
Head moderately broad (WH 1.07 x LH) (Fig. 20) ; front with large
punctures, mesopleura also punctate 30. puncticeps n. sp.
7. Third anteunal segment extremely short, about one-third the length
of the fourth segment (Fig. 23) ; head and thorax blue-green, apical
.4 of abdomen ferruginous 37. hlantoni n. sp.
Third antennal segment at least half as long as the fourth (Figs.
24, 73) ; color variable 8
8. Propodeum black ; head and thorax black or various shades of blue
or green 9
Propodeum bluish or greenish; head and thorax also metallic, some-
times contrasting slightly with the propodeum ; notauli very broad
behind 12
9. Head and thorax a very brilliant green or blue-green; claws strongly
trifid, the middle ray obliquely truncate (Fig. 67) ; second antennal
segment shorter than third segment (as in Fig. 24)
38. vindissimus (Kieffer)
Head and thorax black or weakly to moderately bh;ish or greenish ;
middle ray of claws more erect and acute (Figs. 61, 63) ; second
antennal segment longer than third segment (Fig. 21) 10
10. Propodeum short, measuring 1.35-1.46 x as wide as long, generally
with a fairly wide smooth area laterad of the lateral discal carinae;
front polished, obscurely alutaceous, with weak bluish or greenish
reflections 33. virescens n. sp.
Propodeum more elongate, 1.25-1.33 x as wide as long, generally almost
wholly covered with sculpturing ; front somewhat more evidently
alutaceous than above 11
11. Head and thorax black and without metallic colors; scutellar groove
rather weakly expanded at each end ; OOL slightlj' less than WOT
32. vesculiis n. sp.
EVANS : REVISION OF RHABDEPYRIS 117
Head and thorax more or less blue-green; scutellar groove much
widened on the sides; OOL 1.1-1.25 x WOT, the ocelli being in a
more compact triangle than above 36. suhviridis (Kieffer)
12. Head and thorax green or blue-green, apical half of the abdomen
rufous; notauli separated behind by more than half the width of
one notaulus ; propodeum with five diseal carinae
39. fulgens (Brues)
Head and thorax intensely bluish or violaceous, abdomen black;
notauli unusually wide behind, separated by only a thin line ; pro-
podeum with only three well-defined diseal carinae
41. violaceus n. sp.
TABLE V. SUMMARY OF SOME CHARACTERS OF TYPE SPECIMENS OF SPECIES OF SUBGENUS CHLOREPYRIS {??)
Species LFW WH/LH WF/HE OOLAVOT Propodeal Front Lower mesopi .
(mm) disc W/L femora fovea fully
L/W outlined
24. luteipennis 5.0 0.98 1.47 2.20 1.50 2.1 +
27. quinquelineatus 3.3 1.00 1.17 1.60 1.50 2.4 +
28. nigerrimus 3.8 1.01 1.17 1.90 1.50 2.2 +
29. muscarius 4.0 1.08 1.33 1.50 1.40 1.8
31. origenus 3.3 1.08 1.20 1.20 1.40 1.7
33. virescens 4.3 1.02 1.30 1.40 1.30 2.0
34. viridis 3.0 1.00 1.18 1.40 1.15 1.8
36. subvirldis 3.5 1.10 1.06 1.60 1.30 2.1
38. viridissimus 4.4 1.20 1.10 1.40 1.40 2.2
39. fulgens 4.0 1.16 1.23 1.50 1.40 2.0
40. tricolor 3.5 1.12 1.18 1.70 1.35 2.0
41. uiolaceus 4.5 1.17 1.20 1.60 1.35 2.4
' I fiave omitted metallicus Kieffer, since I have not seen the type or any other specimens. The specimens of muscarius
and subviridis considered are plesiallolypes rather than holotypes; the specimen of viridissimus is also a piesiallotype
of that species as well as tlie type of semiviridis Kieffer. Five species are not linown from the female sex.
TABLE VI. SUMMARY OF SOME CHARACTERS OF TYPE SPECIMENS OF SPECIES OF SUBGENUS CHLOREPYRIS (<?:?)'
Species LFW WH/LH WF/HE OOL/WOT Propodeal Antennal setj.
(mm) disc W/L three L/W
24. luteipennis 3.5 0.98 1.00 1.55 1.65 1.4
25. lobatifrons 3.6 0.97 0.94 1.50 1.90 1.6
26. septemlineatus 3.4 0.95 1.00 1.35 1.65 1.5
27. quinquelineatus 2.8 1.00 0.90 1.50 1.80 1.3
29. muscarius 3.3 1.12 1.25 1.12 1.60 0.8
30. puncticeps 3.8 1.07 1.36 1.10 1.45 0.6
32. vesculus 2.8 1.10 1.35 0.90 1.30 0.6
33. virescens 3.0 1.08 1.40 1.00 1.40 0.7
36. subviridis 2.8 1.18 1.43 1.10 1.35 0.6
37. blantoni 3.8 1.09 1.10 1.10 1.40 0.3
38. viridissimus 3.6 1.20 1.13 1.05 1.40 0.7
39. fulgens 3.9 1.20 1.24 1.00 1.40 0.7
41. violaceus 3.5 1.18 1.28 0.95 1.30 0.7
The specimens treated are either holotypes (lobatifrons, septemlineatus, muscarius, puncticeps, vesculus, subviridis,
blantoni, and viridissimus), allotypes (luteipennis, virescens), or plesiallotypes (quinquelineatus, fulgens). FTvi
species are not l<nown from the male sex.
118 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
LOBATIFRONS SPECIES-GROUP
To this group are assigned five species of black coloration in
which the scutellar groove is slender throughout, the head of
the females more or less produced in front of the eyes, the lower
mesopleural fovea well defined above except margining ridge
sometimes obsolescent for a short distance. In the males, the
third antennal segment is longer than wide and not notably
shorter than the fourth segment ; the head of the male is not
or barely wider than long, the front narrow, WF not or barely
exceeding HE.
24. Rhabdepyris (Chlorepyris) luteipennis new species
Holotype. — 9 , BRAZIL : Nova Teutonia, Santa Catarina,
March 1948 (F. Plaumann) [MCZ, No. 30,951].
Description of female type. — Length 8.5 mm ; LFW 5.0 mm.
Black ; palpi light brown ; mandibles ferruginous, infuscated
beloAV; scape black, flagellum dusky castaneous, darker above
than below ; tegulae testaceous ; coxae black, legs otherwise
fusco-ferruginous except paler apically; wings subhyaline,
strongly tinged with yellowish except apical third of fore wing
smoky, veins and stigma light amber, translucent. Mandibles
slender and rather straight, terminating in one strong tooth
basad of which are four rather indistinct, blunt teeth (Fig. 49).
Clypeus obtusely subangulate, with a median ridge which is
nearly straight in profile. WH .98 x LH; sides of head sub-
parallel behind the eyes, the vertex passing straight across a
considerable distance above eye tops. Front broad, eyes con-
vergent below ; "WF .66 x WH, 1.47 x HE ; ocelli in a compact
triangle far above eyes and close to vertex crest, OOL 2.2 x
WOT. Front strongly shining, barely alutaceous, punctures
small but sharply defined, separated by 2-4 x their own diam-
eters below, much more widely spaced above. Antennae arising
well below bottoms of eyes from beneath prominent lobes ;
first four segments in a ratio of about 23 :5 :6 :9, segment three
slightly wider than long, segment eleven about 1.5 x as long as
wide. (Fig. 11.)
Thoracic dorsum shining and obscurely alutaceous like the
front, the punctures small and widely spaced ; pronotal disc
about 2.5 x as long as mesoscutum ; notauli attenuated and
slightly diverging anteriorly ; parapsidal furrows short and
rather wide; scutellar groove moderately wnde, deflected back-
ward and slightly widened on each side. Propodeal disc 1.5
EVANS : REVISION OP RHABDEPYRIS 119
X as wide as long ; disc with five carinae between which it is
weakly transversely striate, otherwise smooth and polished;
lateral carinae paralleled by a simple groove, but the sub-
laterals not distinct ; declivity smooth except for very fine
transverse striations ; side-pieces with very delicate longitudinal
striae. Mesopleurum polished, with small, sparse punctures;
upper fovea small, ovoid; lower fovea elongate, fully outlined,
much broadened on the posterior .6 (Fig. 29). Middle tibiae
spinose above for their entire length ; claws twice-dentate, the
middle tooth short, sloping outward (Fig. 55) ; front femora
much broadened, measuring 2.1 x as long as their maximum
width.
Allotype. — S, BRAZIL: Santarem (no further data)
[USNM].
Description of male allotype. — Length 5.0 mm ; LFW 3.5 mm.
Head and thorax black ; abdomen black except segments nar-
rowly brownish apically, last two segments mostly brownish ;
palpi testaceous ; mandibles light rufo-castaneous ; antennae
wholly and uniformly light castaneous ; tegulae testaceous ; coxae
black, hind femora fuscous, legs otherwise bright rufo-castane-
ous; wings lightly tinged with yellowish brown, not darkened
apically. Clypeus as described for female. WH .98 x LH, the
general form and insertion of antennae as in female; WF .54
X WH, subequal to HE ; OOL 1.55 x WOT. Front somewhat
more evidently alutaceous and slightly more closely punctate
than in female. First four antennal segments in a ratio of about
6 :2 :3 :3, segment three about 1.4 x as long as wide, segment
eleven about 1.6 x as long as wide. (Fig. 71.)
Thoracic dorsum weakly alutaceous, the pronotum more so
than the mesonotum, punctures weak; features of mesonotum
as in female. Propodeal disc 1.65 x as wide as long, with five
carinae between which it is weakly transversely striate, else-
where smooth and polished; declivity strongly polished, barely
alutaceous; side-pieces with fine longitudinal striations. Meso-
pleurum weakly alutaceous, with small punctures, the foveae
shaped as in the female, the two foveae separated by a broad,
flat ridge. Front femora 2.1 x as long as their maximum width;
middle tibiae weakly spinose ; claws bifid, the inner ray close to
the outer ray, obliquely truncate (Fig. 56).
Paratypes. — BRAZIL : 8 ? 2 , same data as type except
various dates, September-March, 1952- '62 [MCZ, USNM, BSA,
Coll. G. R. Ferguson] ; 1 9 , Urucum, Corumba, 12-19 December
120 BULLETIN : MUSEUM OF COMPARATR^ ZOOLOGY
1919 (C. U. Exped.) [CU] ; 1 S, Santarem [USNM]. TRINI-
DAD: 1 5 , St. Augustine, 25 July 1935 (N. A. Weber)
[MCZ]. PANAMA: 1 9 , March 1923 (no further data) [MCZ] ;
1 $ , Barro Colorado Isl., Canal Zone, 14 June 1939 (J. Zetek)
[USNM]. NICARAGUA: 1 <? , San Carlos, 1933 (Wickham
Coll.) [USNTM].
Variation. — The females vary in size from 7.5 to 9.5 mm,
LFW from 4.5-5.5 mm. The Panama and Nicaragua specimens
have the legs bej^ond the coxae bright rufo-ferruginous, the tip
of the abdomen also suffused with this color, but they otherwise
show no consistent differences from the Brazilian females.
In this series WH varies from .97 to 1.00 x LH, WF from 1.25
to 1.50 X HE, OOL from 2.2 to 2.5 x WOT. The sculpturing
of the propodeum and mesopleurum shows little variation
throughout the series.
The Santarem male paratype is very similar to the allotype
in every respect. The Trinidad and Panama males have the
legs beyond the coxae and the tip of the abdomen rufo-ferru-
ginous, as in the Central American females. These two speci-
mens also have both the upper and lower mesopleural foveae
larger than in the rest of the type series, the ridge between them
narrow and round-topped. This appears to be an important
difference, and it may be that these two males go with a different,
as yet undiscovered, female ; however, in all standard measure-
ments they differ scarcely at all from the Santarem males
(although both are slightly larger, LFW 3.9-4.0 mm). The
Trinidad male is without antennae.
25. Rhabdepyris (Chlorepyris) lobatifrons Kieffer
Ehahdepyris lobatifrons Kieffer, 1910, Auu. Soe. Ent. France, 78: 297-
298 [Type: $ (not 2 as stated), BEAZIL: Para (C. F. Baker)
(Pomona College)]. —Kieffer, 1914, Das Tierreich, 41: 361.
? Ehahdepyris ohscuripennis Kieffer, 1910, Ann. Soc. Ent. France, 79: 40
[Type: $ (?), BEAZIL: Para (C. F. Baker) (location unknown)].
—Kieffer, 1914, Das Tierreich, 41: 359.
Description of male type. — Length 5 mm ; LFW 3.6 mm.
Wholly shining black except tip of abdomen suffused with dull
ferruginous ; palpi testaceous ; mandibles light castaneous, the
tips rufous; antennae wholly and uniformly bright castaneous;
tegulae testaceous ; legs bright testaceous except all coxae
strongly infuscated with yellowish brown, veins and stigma light
brown. Clypeus obtusely angulate, its median ridge very weakly
EVANS : REVISION OF RHABDEPYRIS 121
arched in profile. Head .97 x as wide as high; front rather
narrow, WF .51 x WH, .94 x HE; OOL 1.50 x WOT; front
angle of ocellar triangle slightly less than a right angle ; dis-
tance from eye tops to crest of vertex equal to roughly .5 x
HE, the vertex straight across; distance from posterior ocelli
to vertex crest about equal to width of an ocellus. Front pol-
ished, very obscurely alutaceous, punctures small although
sharply defined, separated by 1-2 x their own diameters except
more widely spaced toward the vertex. Antennae arising from
rather prominent tubercles well below bottoms of eyes ; first
four antennal segments in a ratio of about 12 :3 :7 :7, segments
three and four both 1.6 times as long as wide, segment eleven
twice as long as wide.
Thoracic dorsum polished and very obscurely punctate, like
the front, the punctures fairly strong but widely spaced, absent
on the anterior part of the mesoscutum and center of the
scutellum; pronotal disc 1.35 x median length of mesoscutum;
notauli strong, barely diverging anteriorly, represented on the
anterior half of the scutum only by hair-lines; basal scutellar
groove thin, deflected backward and considerably widened on
each side. Propodeum very short, the disc 1.9 x as wide as its
median length ; disc with seven carinae, five of them close
together, straight, and nearly complete (the median carina com-
plete), the other two spaced more widely to the sides, arching
mesad apically and stopping well short of the transverse carina ;
disc obscurely striate between the carinae, elsewhere smooth
and polished ; sublateral carinae absent ; posterior corners f oveo-
late. Mesopleurum wholly moderately alutaceous, weakly shin-
ing ; upper fovea small ; lower fovea elongate, weakly constricted
in the middle, fully outlined above and below. Middle tibiae
weakly spinose ; claws bifid.
Remarks. — The type of Kieifer's ohscuripennis is probably
lost. Kieffer did not state the sex in his original description,
but in 1914 he indicated questionably that it was a male. One
notes that the type locality, size, and color are essentially the
same in the types of lohafifrons and ohscuripennis, and no strik-
ing differences are apparent from Kieffer 's descriptions except
that the wings of the latter are "d'un brun sombre." The
wings are distinctly tinged with yellowish brown in the type
of lohatifrons, and I question that the other type is very dif-
ferent. In 1914, KiefPer keyed out ohscuripennis as having
spines on the middle tibia, lohatifrons as lacking spines, but
122 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
examination of the type of lohatifrons reveals that spines are
present. Kieffer stated that lohatifrons has 5 parallel carinae
on the propodeum ; he neglected to mention that there are two
additional, more oblique discal carinae. I suggest that lohati-
frons and ohscuripennis be regarded as probable synonyms at
least until the type of ohscuripennis is found.
26. Rhabdepyris (Chlorepyris) septemlineatus Kieffer
Ehahdcpyris septemlineatus Kieffer, 1906, Berlin. Eiit. Zeitsch., 50: 250
[Type: S, NICAEAGUA: Granada (C. F. Baker) (Pomona Coll.)]
—kieffer, 1914, Das Tierreich, 41: 361.
Description of male type. — Length 4.1 mm; LFW 3.4 mm.
Head, thorax, and basal three segments of abdomen shining
black, apical four segments of abdomen ferruginous ; mandibles
yellowish brown, darker basally and apically; antennae light
f errugiiio-castaneous, somewhat inf uscated on apical half ; tegu-
lae testaceous; coxae black, femora ferrugino-castaneous, legs
otherwise light yellowish brown ; wings subhyaline, with dark
setulae, veins and stigma brown. Clj'peus not prominent, its
apical margin obtusely subangulate, median carina not arched
in profile. WH .95 x LH ; WF .56 x WH, subequal to HE ; inner
orbits converging below; distance from eye tops to vertex crest
equal to slightly over half HE, vertex straight across; ocelli in
about a right triangle far above eye tops and close to vertex
crest ; OOL 1.35 x WOT ; hind ocelli removed from vertex crest
by less than their own diameter. Front strongly polished,
weakly alutaceous on lower sides only, punctures small but fairly
prominent, on lower front separated by about their own diam-
eters, on upper front and vertex separated by 2-4 x their own
diameters. First four antennal segments in a ratio of about
25:7:14:14, segment three about 1.5 x as long as thick, segment
eleven about 1.7 x as long as thick; pubescence moderately
coarse, subappressed, somewhat golden, erect setae short but
fairly numerous.
Pronotal disc not sharply differentiated from collar, about
1.3 X as long as mesoscutum, with smooth contours, weakly alu-
taceous and with distinct, well separated punctures. Mesoscu-
tum strongly polished, scarcely alutaceous, weakly transversely
depressed posteriorly; parapsidal furrows very short, occupying
the posterior third of the mesoscutum only, notauli weakly di-
verging anteriorly, attenuate anteriorly, occupying the posterior
half of the scutum. Scutellar disc weakly alutaceous, with a few
EVANS : REVISION OP RHABDEPYRIS 123
punctures ; basal groove narrow, of uniform width throughout
but distinctly deflected backward on each side. Propodeal disc
quadrate, 1.65 x as wide as long, lateral and transverse posterior
carinae very strong; median portion of disc with five closely
parallel carinae which are complete or nearly so, also with two
additional carinae laterad of these, extending only slightly
more than half the length of the disc ; disc otherwise polished and
with only some very M^eak sculpturing ; side-pieces weakly longi-
tudinally striolate. Mesopleurum weakly alutaceous, with a
well-defined, elongate lower fovea which is widely separated
from the small upper fovea (much as in luteipennis, Fig. 29).
Claws bifid, inner ray truncate ; middle tibiae with a few spines ;
front femora 2.1 x as long as their maximum width.
Additional specimen examined. — PANAMA : 1 $ , Barro Col-
orado IsL, Canal Zone, 11 February 1955 (C. Rettenmej^er)
[KU]._
Variation. — The Panama male is slightly larger than the
type and has the front and hind femora moderately infus-
cated, but it is otherwise very similar, the apical .4 of the
abdomen being bright ferruginous as in the type. In this speci-
ment WH is .98 x LH, WF .92 x HE, OOL 1.50 x WOT ; the
lateral pair of discal carinae extend for about .8 the length
of the propodeum.
27. Rhabdepyris (Chlorepyris) quinquelineatus Kieffer
Rhabdepyris quinquelineatus Kieffer, 1906, Berlin. Ent. Zeitschr.. 50: 249
[Type: 5, NICAEAGUA: Chinandega (C. F. Baker) (Pomona
Coll.)]. —Kieffer, 1914, Das Tierreich. 41: 360.
Description of female type. — Length 5 mm; LFW 3.3 mm.
Body shining black, without metallic hues, last two abdominal
segments weakly suffused with reddish brown ; mandibles and
antennae rather uniformly reddish brown, antennal tubercles
also suffused with this color ; tegulae testaceous ; coxae black,
legs otherwise reddish brown ; fore wing weakly clouded, more
noticeably so about the radial vein, veins and stigma brown.
Mandibles with five teeth, the basal three teeth very small
(essentially as figured for luteipennis, Fig. 49). Clypeus form-
ing a subangulate median process, its median elevation rather
weak. Head about as wade as high ; inner orbits convergent
below; WF .61 x WH, 1.17 x HE; vertex straight across a
short distance above eye tops, distance from eye tops to vertex
crest about half HE. Ocelli in a broad triangle close to vertex
124 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
crest; OOL 1.6 x WOT; posterior ocelli removed from vertex
crest by about their own diameters. Front shining, obscurely
alutaceous, with strong punctures which are separated by 1-2 x
their own diameters (much more widely spaced above). First
four antennal segments in a ratio of about 19 :5 :5 -.7, segment
three only 1.2 x as long as thick, segment eleven 1.4 x as long as
thick.
Pronotum large, with even contours, anteriorly not sharplj^
separated from collar, on the median line 1.8 x as long as meso-
scutuni; pronotum strongly shining and with large, well-sep-
arated punctures. Mesoscutum with a shallow transverse im-
pression, shining and almost impunctate ; notauli attenuate and
diverging anteriorly ; scutellar disc rather flat, basal groove
narrow, of even width throughout, its ends turned obliquely
backward. Propodeal disc 1.5 x as wide as long, its lateral
and posterior cariuae strong, also with five earinae on the median
area, all of them reaching the posterior margin or nearly so ;
disc weakly transversely striate between the earinae, on the
sides smooth and strongly polished. Mesopleurum strongly pol-
ished, with scattered small punctures, lower fovea distinct,
about as in septemlineatus. Claws bifid, inner ray truncate
(Fig. 57); front femora 2.4 x as long as wide; middle tibiae
spinose.
Phsiallotijpe. — 5 , EL SALVADOR : San Salvador, 3-4 May
1958 (0. L. Cartwright) [USNM].
Description of male plesiallotype. — Length 3.7 mm; LFW
2.8 mm. Black, apical two abdominal segments suffused with
reddish brown ; palpi testaceous ; mandibles pale castaneous, the
teeth rufous ; antennae wholly light brown, except suffused
with darker brown on upper surface ; coxae black, legs other-
wise bright ruf o-castaneous ; wings subhyaline, veins yellowish
toward base of wing. Clypeus obtusely angulate, with a low
median ridge. Head as wide as high ; eyes strongly convergent
below, WF only .49 x WH, .90 x HE ; OOL 1.50 x WOT. Front
strongly shining, barely alutaceous, with small punctures which
are close below, widely spaced above. First four antennal seg-
ments in a ratio of about 18 :6 :10 :11, segment three 1.3 x as long
as wide (Fig. 72), segment eleven twice as long as wide. Thor-
acic dorsum strongly polished, non-alutaceous, with small punc-
tures ; features as in female. Propodeum very short, the disc
1.8 x as wide as long, features as in female. Mesopleurum shin-
ing, with a few small punctures ; lower fovea elongate, well
EVANS : REVISION OF RHABDEPYRIS 125
defined except the upper margin rather indistinct toward the
middle. Claws bifid; middle tibiae with a few small spines
above ; front femora 2.2 x as long- as wide.
Additional specimens examined. — MEXICO : 1 9 , Medellin,
Veracruz (C. F. Baker no. 2154) [USNM] ; 1 $ , Progreso,
Yucatan, 23 July 1962 (H. E. Evans) [MCZ].
Variation. — In both of the Mexican specimens, the upper
margin of the lower mesopleural fovea is indistinct in the
middle. The female differs from the type also in having the
third antennal segment slightly wider than long and the femora
dark brown, contrasting to the light brown tibiae and tarsi.
The Yucatan male also has dark brown femora, but the anten-
nae are lighter than in the plesiallotype ; in this specimen the
propodeum is also 1.6 x as wide as long.
28. RiiABDEPYRis (Chlorepyris) nigerrimus new species
Holoiypc. — 9, BOLIVIA: Santa Cruz, Santiago, Novem-
ber 1959 (no collector given) [KU].
Description of female type. — Length 6.5 mm; LFW 3.8 mm.
Wholly black, shining, the apical abdominal tergite suffused
with dark brown ; palpi brown ; mandibles black, flagellum dark
brown above, light brown below ; tegulae brownish fuscous ; legs
black except apices of femora and tibae suffused with dark fer-
ruginous, tibial spines and spurs light ferruginous, tarsi castane-
ous ; wings hyaline except fore wing lightly clouded along major
veins and on apical .6, more especially' around the radial vein,
veins and stigma dark brown except veins yellowish at extreme
base of wing. Mandibles with a large apical tooth and four
small, blunt teeth in an oblique series basad of this, with a
small tooth on the lower margin (differing in no important way
from those of luteipennis. Fig. 49). Clypeus broadly angulate,
its median ridge not arched in profile. AVH 1.01 x LH ; eyes
convergent below, WF .61 x WH, 1.17 x HE ; front angle of
ocellar triangle less than a right angle, OOL 1.9 x WOT. Dis-
tance from eye tops to vertex crest about .5 x HE ; vertex nearly
straight across. Front polished, very obscurely alutaceous ; lower
part of front with a median streak and with some large, well-
spaced punctures, but the punctures on the upper half of the
front and on the vertex weaker and very widely spaced. First
four antennal segments in a ratio of about 37 :8 :8 :10, segments
three slightly wider than long, segment four slightly longer than
wide, segment eleven 1.3 x as long as wide. (Fig. 12.)
126 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Pronotal disc about 1.7 x as long as mesoscutum, strongly
polished and with small, widely spaced punctures ; mesoscutum
very weakly alutaceous, with a strong transverse depression ;
notauli complete, diverging and attenuate toward the front;
scutellar groove slender, deflected backward at each end. Pro-
podeal disc 1.5 x as wide as long, with five carinae between
which it is strongly transverse striate, the disc elsewhere smooth
and shining ; declivity alutaceous, without transverse ridges ;
side-pieces very finely longitudinally striate. Mesopleurum pol-
ished, with scattered small punctures; upper fovea small, only
slightly longer than high; lower fovea rather strongly defined
except upper margin indistinct on the middle third (Fig. 30).
Front femora 2.2 x as long as wide ; middle tibiae strongly spi-
nose; claws with the tooth short but sloping outward strongly
(Fig. 58).
Remarks. — This species is closely related to the preceding
and may only be an extreme variant of it. However, the only
known specimen is from a locality far south of the southern-
most record for quinquelineatus (Nicaragua).
MUSCARIUS SPECIES-GROUP
To this group I assign seven species, four of them without
metallic colors, the other three with the head and thorax weakly
to strongly reflecting blue or blue-green. The head of the female
is strongly developed below the eyes as in the preceding group,
the antennae arising well below the bottoms of the eyes. How-
ever, the third antennal segment of the male is very much
shorter than in that group. The claws are dentate or weakly
bifid, the scutellar groove variable and generally somewhat inter-
mediate between that of the preceding and the following species-
groups. The mandibles of the female have a strong preapical
tooth on the lower margin.
29. Rhabdepyris (Chlorepyris) muscarius (Westwood)
Epyris muscarius Westwood, 1874, Thesaurus Ent. Oxoniensis, p. 159, pi.
29, fig. 8 [Type: $, BEAZIL: Amazonia, 1861 (Bates) (HCOU)].
Rhahdepyris microstoma Kieffer, 1910, Ann. Soe. Ent. France, 78: 296-297
[Type: $ (not $ as stated), BEAZIL: Para (C. F. Baker) (Pomona
Coll.)]. New synonymy.
Rhabdepyris (Rhabdepyris) microtoma [sic] Kieffer, 1914, Das Tierreich,
41: 361-362.
Rhabdepyris (Rhabdepyris) muscarius Kieffer, 1914, ibid., p. 362.
EVANS : REVISION OF RHABDEPYRIS 127
Plesiallotype: 9 , BRAZIL: Chapada, August (Ace. no. 2966)
[CM].
Description of female plesiallotype. — Length 6.0 mm ; LFW
4.0 mm. Black except extreme tip of abdomen suffused with
brownish ; palpi light brown, mandibles rufo-testaceous, black
toward base ; scape dark brownish fuscous, ilagellum brown,
darker above than below ; tegulae testaceous ; coxae black, fe-
mora dark brown, legs otherwise light castaneous ; wings lightly
tinged with yellowish brown, veins yellowish brown, stigma dark
brown. Body setae light golden brown, short but rather con-
spicuous against the black integument. Mandibles with six
teeth, the basal three teeth small, the next tooth broad, deeply
separated from the large apical tooth, also with a sharp tooth on
the outer margin at the base of the apical tooth (Fig. 50).
Clypeus very short, angulate apically, the angle slightly greater
than a right angle. AVH 1.08 x LH; front broad, WF .66 x
WH, 1.33 X HE; ocelli in about a right triangle, OOL 1.5 x
WOT. Vertex broad, nearly straight, distance from eye tops to
vertex crest only about one-third x HE. Front shining, barely
alutaceous, punctures small, separated by 3-5 x their own diam-
eters. Antennae arising from below frontal lobes, well below
bottoms of eyes; first four antennal segments in a ratio of about
37 :9 :6 :12, segment three .7 as long as wide, segment four 1.2 x
as long as wide. (Fig. 13.)
Thoracic dorsum shining like the front, punctures small and
sparse; pronotum 1.6 x as long as mesoscutum; mesoscutum
transversely depressed on sides; notauli broad, attenuate and
diverging toward the front ; parapsidal furrows linear, extend-
ing for half the length of the mesoscutum ; scutellar furrow
very slender, slightly enlarged at each end. Propodeal disc
1.4 X as wide as long, with five discal carinae between which
it is transversely striate, elsewhere smooth except for foveolate
grooves along the lateral and posterior carinae ; declivity striate
only on the lower third ; side-pieces shining, with only very fine
sculpturing. Mesopleurum weakly alutaceous, with a few small
punctures; lower fovea broadly incomplete above (Fig. 31).
Front femora very strongly incrassate, only 1.8 x as long as
wide; middle and hind tibiae spinose ; claws dentate (Fig. 59).
Description of male type. — Length 5.5 mm ; LFW 3.3 mm.
Black ; palpi and mandibles largely testaceous ; antennae light
castaneous, both scape and flagellum sutfused with fuscous on
the upper side ; tegulae testaceous ; legs pale castaneous, except
128 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the front coxae fuscous, the other coxae and all the femora
suffused with brownish ; wings lightly, uniformly tinged with
broAvnish, the veins and stigma light brown. Body with rather
coarse, pale setae. Clypeus forming a sharp obtuse angle, the
median ridge not arched in profile. Head 1.12 x wide as high ;
front broad, WF .64 x WH, 1.25 x HE ; OOL 1.12 x WOT ; front
angle of ocellar triangle slightly less than a right angle ; ocelli
moderately large, the hind ocelli separated from the vertex
crest by about their own diameters. Vertex passing straight
across only a short distance above the eyes, distance from eye
tops to vertex crest less than WOT. Front strongly polished,
non-alutaceous except weakly so near the bottom, with minute
punctures which are separated by 3-4 x their own diameters,
except more crowded below. Antennae arising slightly below
bottoms of eyes, first four antennal segments in a ratio of
about 18 :8 :5 :9, segment three .8 x as long as wide, segment
four 1.5 X as long as wide, segment eleven about twice as long
as wide ; flagellum with rather coarse, semi-erect setulae and a
few fully erect setae. (Fig. 19.)
Thoracic dorsum polished, non-alutaceous, the punctures very
small ; maximum width of pronotal disc nearly twice its median
length, the latter 1.3 x the median length of the mesoscutum ;
notauli strong on the posterior two-thirds of the mesoscutum,
continued to the front margin only as thin lines ; basal scutellar
groove deep but thin, slightly expanded and deflected backward
on each side. Propodeal disc 1.6 x as wide as long; disc with
five strong carinae, of which the middle three are complete
and the others nearly so ; surface wealdy transversely ridged be-
tween the discal carinae, elsewhere very smooth and polished;
sublateral carina absent, but the extreme sides with a narrow
groove ; posterior angles f oveolate. Mesopleurum shining, non-
alutaceous, the upper margins of the lower fovea obsolete (Fig.
37). Middle tibiae with some fairly thick setae above, but not
really spinose ; claws bifid (Fig. 60).
Specimens examined. — BRAZIL : 1 9,1 $ , Chapada, March,
August [CM] -,2 S S , Para (C. F. Baker) [Pomona Coll., CU] ;
1 $ , Amazonia (Bates) [Type, HCOU] ; 2 9 9 , Nova Teutonia.
Santa Catarina, September, October 1957 (F. Plaumann) [Coll.
G. R. Ferguson]. PERU: 1 S , Quincemil, Dept. Cuzco, 750 m,
16 October 1962 (L. Pena) [MCZ]. COLOMBIA: 1 S, Bonda,
August [CM]. PANAMA: 2 9 9, Barro Colorado Isl., Canal
Zone, March (Bradley, Rettenmeyer) [CU, KSU] ; 1 c5 , Pacora,
EVANS : REVISION OP RHABDEPYRIS 129
Canal Zone, 13 May 1953 (F. S. Blanton) [USNM]. COSTA
RICA : 1 5 , La Fortuna, San Carlos, Alajuela Prov., 19
February 1964 (H. E. Evans) [MCZ]. MEXICO: 1 $ , Atoyac,
Veracruz, April (H. H. Smith) [BMNH].
Variation. — Although I did not compare the types of miis-
carius and microstoma direetl}^, my notes leave little doubt that
they represent the same species ; the type of microstoma is
slightly smaller (LFW 2.8 mm). The males from Chapada
and from Quincemil, Peru, are larger (LF"W 3.7 mm) but very
similar in all details. The males from Colombia, Panama, Costa
Rica, and Mexico are unusually small (LFW 2.2-2.8 mm) and
have a relatively more elongate propodeum (1.3-1.5 x as wide
as long) and a slightly broader front (WF 1.3-1.4 x HE). The
Panama specimen has the apical third of the abdomen rufous,
while the Colombia specimen has the antennae wholly testaceous.
One of the Santa Catarina females is rather small (LFW only
3.0 mm) but is colored like the plesiallotj^pe and has similar
standard measurements. The other is larger (LFAV 4.7 mm)
and has the front femora w^iolly bright ferruginous, the head
somewhat longer (WH 1.02 x LH), the front broad (WF 1.45 x
HE), the propodeal disc only 1.3 x as wide as long. The Panama
females also differ slightly from the plesiallotype. Both have
the apical third of the abdomen rufous, the head a little less
broad (WH 1.02 x LH), the front wider (WF 1.40-1.47 x HE),
the propodeum less broad (1.30-1.35 x as wide as long). How-
ever, the mandibles and all other important features agree well
with the plesiallotype. These females are comparable to the
plesiallotype in size (LFW 3.8, 4.2 mm) even though the males
from this area are very small. It is possible that I am confusing
more than one species under the name muscarius, but I do not
think so. Much of the variation parallels the variation in lutei-
pennis.
30. Rhabdepyris (Chlorepyris) puncticeps new species
Holotype. — $ , BRAZIL : Nova Teutonia, Santa Catarina,
February 1963 (F. Plaumann) [MCZ, No. 30,952].
Description of male type. — Length 5.0 mm; LFW 3.8 mm.
Body shining black except apical two abdominal segments suf-
fused with reddish brown ; palpi straw colored ; mandibles black
except apical third testaceous, teeth rufous; antennae wholly
light brown below, dark brownish fuscous above except segments
two and three wholly castaneous ; tegulae light brown ; coxae
130 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
and middle and hind femora black, hind tibiae somewhat infus-
cated, legs otherwise bright ruf o-castaneous ; wings faintly tinged
with yelloMdsh brown, apical half of fore wing slightly more
heavily clouded. Body setae rather coarse, light broAvn. Cly-
peus forming a sharp right angle, its median carina not arched
in profile. Head 1.07 x as wide as high; WF .67 x WH, 1.36 x
HE ; OOL 1.10 X WOT, the front angle of the ocellar triangle
very slightly less than a right angle. Vertex passing straight
across a short distance above eye tops. Front strongly polished,
non-alutaceous except weakly so below, wholly covered with
strong punctures which are separated by 1.5-3 x their own
diameters, except rather sparse in front of the anterior ocellus.
First four antennal segments in a ratio of about 26 :8 :5 :15, seg-
ment three nearly twice as wide as long, segment four about
1.5 X as long as wide, segment eleven 2.2 x as long as wide.
(Fig. 20.)
Thoracic dorsum wholly polished and non-alutaceous, punc-
tures small but quite strong, rather sparse on the mesoscutum
and absent from the center of the scutellar disc ; notauli strong
on the posterior two-thirds of the mesoscutum, continued to the
front margin as thin lines ; basal scutellar groove deep but thin,
slightly expanded and deflected backward at each end. Pro-
podeal disc 1.45 x as wide as long, with five strong carinae
laterad of which it is smooth and polished ; declivity with numer-
ous transverse ridges; side-pieces weakly aciculate. Mesopleurum
shining, non-alutaceous, with sparse punctures, the upper mar-
gins of the lower fovea obsolete except at the ends. Front femora
2.15 X as long as wide ; middle tibiae not spinose ; claws bifid.
Remarks. — This species is known to me from the type only.
31. Rhabdepyris (Chlorepyris) origenus Kieffer
Rhabdepyris origenus Kieffer, 1911, Ann. Soe. Sci. Bruxelles, 35: 222
[Type: $, MEXICO: Chilpancingo, Guerrero, 4600 feet, June (H. H.
Smith) (BMNH)]. — Kieffer, 1914, Das Tierreich, 41: 359.
Description of female type. — Length 5.3 mm; LFW 3.3 mm.
Black, except tip of abdomen suffused with dark brown ; palpi
light brown ; mandibles ferruginous except black at base ; an-
tennae ferruginous except under side of flagellum infuscated
for its entire length ; legs bright ferrugino-castaneous except
coxae strongly infuscated; wings subhyaline. Mandibles essen-
tially as described and figured for muscarius (Fig. 50). Clypeus
forming a rather sharp angle medially, with a strong median
ridge. WH 1.08 x LH ; WF .66 x WH, 1.20 x HE; ocelli in a
EVANS : REVISION OF RHABDEPYRIS 131
right triangle close to vertex crest, OOL 1.20 x WOT. Vertex
passing straight across a short distance above the eye tops,
distance from eye tops to vertex crest only about one-third x
HE. Front uniformly alutaceous, moderately shining, punctures
small but numerous, separated by 1.5-2.5 x their own diameters.
First four antennal segments in a ratio of about 36:10:6:11,
segment three 0.7 x as long as wide, segment eleven 1.1 x as long
as wide.
Pronotal disc about 1.5 x as long as mesoscutum, shining and
weakly alutaceous, with small punctures. Mesoscutum with the
notauli complete, linear, diverging anteriorly ; parapsidal fur-
rows also linear, extending for about half the length of the
scutum ; scutellar groove slender, slightly widened and deflected
backward at each end. Propodeal disc 1.4 x as wide as long,
with five carinae, the disc transversely striate between the
carinae, elsewhere merely weakly alutaceous ; declivity trans-
versely striate below ; side-pieces polished and with only some
very fine surface sculpturing. Mesopleurum weakly alutaceous,
weakly punctate; upper fovea small, lower fovea not at all de-
fined on the upper margin (Fig. 32). Front femora greatly
swollen, 1.7 x as long as their greatest width; claws dentate;
middle tibiae moderately spinose.
Additional specimen examined. — MEXICO: 1 9, San Bias,
Nayarit, 22 March 1962 (F. D. Parker) [UCD].
Variation. — The Nayarit female is of the same size as the
type, but differs slightly in color : the apical 2.5 segments of the
abdomen are ferruginous, the antennae somewhat lighter and
more uniformly ferruginous than in the type. Standard meas-
urements of this specimen are as folloM^s: WH 1.03 x LH,
WF 1.33 X HE, OOL 1.30 x WOT; propodeum 1.22 x as wide
as long.
32. Rhabdepyris (Chlorepyris) vesculus new species
Holotype. — $ , BRAZIL : Nova Teutonia, Santa Catarina, 14
January 1962 (F. Plaumann) [MCZ, No. 30,953].
Description of male type. — Length 4.6 mm ; LFW 2.8 mm.
Entirely black ; palpi testaceous ; mandibles testaceous except
black at extreme base, teeth rufous; antennae testaceous except
scape somewhat infuscated, especially above, flagellum slightly
darker on the upper side than below ; tegulae testaceous ; coxae
black, femora dark brownish fuscous, trochanters and hind
tibiae somewhat infuscated, legs otherwise testaceous; wings
subhyaline, veins and stigma brown. Mandibles with five teeth.
132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the basal four teeth small and sharp ; elypeiis obtusely angulate,
with a median ridge which is not arched in profile. Head rather
broad, WH 1.10 x LH ; WF .66 x WH, 1.35 x HE; ocelli
moderately large, in about a right triangle, the posterior ocelli
slightly less than their own diameters from the vertex
crest; OOL 0.90 x WOT. Vertex very broadly rounded, almost
straight ; distance from eye tops to vertex crest about one-third
X HE. Front wholly alutaceous, moderately shining, with small
punctures which are separated by 1.5-2.5 x their own diam-
eters, except sparse in front of ocellar triangle. Antennae aris-
ing well below bottoms of eyes, unusually short, first four seg-
ments in a ratio of about 15:5:4:6. segment three .6 x as long
as wide, segment four about as long as wide, segment eleven
1.4 X as long as wide ; flagellar pubescence semi-erect, pale.
(Fig. 22.)
Thoracic dorsum alutaceous and with small punctures like
the front ; pronotum about 1.6 x as long as the mesoscutum ;
notauli rather wide, much tapered and divergent anteriorly;
scutellar groove relatively short and wide, much wider on the
sides than medially. Propodeal disc 1.3 x as wide as long, with
five discal carinae and also with two more rounded ridges on
each side between the lateral discals and the laterals, the surface
rather irregularly transversely striate between the carinae and
ridges, wholly covered with sculpturing; declivity and side-
pieces both with strong horizontal ridges. Mesopleurum weakly
alutaceous, without distinct punctures ; lower fovea very broadly
open above ; upper fovea elongate, opening downward posteriorly
(Fig. 38). Front femora 2.3 x as long as wide; middle femora
not spinose; claws dentate, the tooth acute, sloping outward
somewhat (Fig. 61).
Paratypes. — BRAZIL : 5 S S , same data as type except
September, November, December, January, 1951, 1962, 1963
[MCZ, USNIM, Coll. G. R. Ferguson].
Variation. — The paratypes are very similar to the type in
color except that the antennae tend to be darker in some speci-
mens, wholly dull brownish in one. The sculpturing is very
similar except that one specimen has only one ridge between
the lateral discal and lateral carinae of the propodeum, leaving
a narrow smooth area on each side. LFW varies from 2.3 to
2.8 mm; WH/LH varies from 1.07 to 1.10; WF/HE varies from
1.28 to 1.44 x HE ; the propodeal disc varies from 1.28 to 1.33
X as wide as long. Throughout the series WOT exceeds OOL
very slightly.
EVANS : REVISION OF RHABDEPYRIS 133
33. Rhabdepyris (Chlorepyris) virescens new species
Holotypc. — 9 , BRAZIL : Nova Teutonia, Santa Catarina,
August 1963 (F. Plaumann) [MCZ, No. 30,954].
Description of female type. — Length 6.0 mm; LFW 4.3 mm.
Head and thorax black, with dull olive-green reflections ; pro-
podeum black ; abdomen shining black except the apical two
segments suffused with ruf o-testaceous ; palpi straw-colored ;
mandibles ruf o-castaneous except black at extreme base ; scape
bright rufo-castaneous, flagellum light brown below, somewhat
dusky above ; tegulae testaceous ; legs bright ferruginous except
front and hind coxae black, middle coxae and front and hind
femora partially suffused with black; wings lightly infuscated,
the fore wing with a darker cloud below the marginal cell ; veins
brown except testaceous toward base of wing. Mandibles with
a small subapical tooth on the lower margin, a large apical
tooth, and four small teeth which are close together and sep-
arated from the apical tooth by a small cutting edge (Fig. 51).
Clypeus obtusely angulate, its median ridge nearly straight in
profile. WH 1.02 x LH ; front broad, AVF .69 x WH, 1.30 x HE ;
front angle of ocellar triangle very slightly less than a right
angle, OOL 1.40 x WOT. Vertex passing straight across, dis-
tance from eye tops to vertex crest only .3 x HE. Front shining,
weakly alutaceous, with very small punctures which are sep-
arated by 3-5 X their own diameters. Antennae arising from
beloAv frontal lobes, well below bottoms of eyes; first four seg-
ments in a ratio of about 20 :5 :4 :6, segment three .8 as long as
wide, segment four 1.2 x as long as wide, segment eleven 1.3 x
as long as wide. (Fig. 14.)
Pronotum weakly alutaceous and Avith small punctures like
the front, the median length of the disc nearly twice that of
mesoscutum. Mesoscutum slightly more strongly alutaceous than
pronotum, with small punctures, weakly transversely depressed;
notauli rather Avide, diverging and attenuate anteriorly, reach-
ing anterior margin only as thin lines ; scutellar groove rather
slender, deflected sharply backward and expanded at each end.
Propodeal disc 1.3 x as wide as long, with five discal carinae
between which it is strongly transversely striate, between the
lateral discals and lateral carinae smooth and very weakly
alutaceous ; declivity with strong transverse ridges on lower half ;
side-pieces smooth except very finely alutaceous. Mesopleurum
weakly alutaceous, very weakly punctate ; upper fovea ovoid,
134 BULLETIN : MUSEUM OF COMPARATR^ ZOOLOGY
slightly longer than higli ; lower fovea broadly incomplete above
(Fig. 33). Front femora twice as long as wide; middle tibiae
spinose; claws dentate (Fig. 62).
Allotype. — $ , BRAZIL : same data as type except Decem-
ber, 1962 [MCZ].
Description of male allotype. — Length 4.5 mm ; LFW 3.0
mm. Head and thorax shining black, with a Aveak bluish green
cast ; propodeum black ; abdomen shining black, the apical two
segments suffused with brownish ; palpi straw-colored ; man-
dibles testaceous except black at base, the teeth rufous ; antennae
pale castaneous except scape fuscous above, flagellum slightly
infuscated above ; tegulae testaceous ; coxae black, femora dark
brown, legs otherwise testaceous ; wings very lightly tinged
with brownish, somewhat more evidently so around the radial
vein. Mandibles and clypeus not differing noticeably from
those of vesculus; head much wider than high, WH 1.08 x LH;
WF .68 X WH, 1.40 x HE; ocelli moderately large, in about
a right triangle, OOL and WOT subequal. Vertex broadly
rounded ; distance from eye tops to vertex crest equal to about
one-third x HE. Front strongly polished, barely alutaeeous,
the punctures minute and inconspicuous. First four antennal
segments in a ratio of about 18 :6 :5 :8, segment three .7 as long
as wide, segment four approximately as long as wide, segment
eleven about twice as long as wide ; flagellar pubescence semi-
erect, about .2 as long as width of flagellar segments. (Fig. 21.)
Pronotal disc slightly longer along midline than mesoseutum,
obscurely alutaeeous and wdth very small punctures. Mesoseu-
tum strongly polished, wholly covered with minute setigerous
punctures; notauli rather wide, diverging and strongly attenu-
ated anteriorly ; scutellar groove fairly wide, broadened and
deflected backward at each end ; center of scutellar disc polished
and impunctate. Propodeal disc 1.4 x as wide as long, with five
discal carinae between which it is irregularly transversely
ridged ; space laterad of lateral discals filled with transverse
striae which are obsolescent behind ; lateral and sublateral cari-
nas present, each subtending a somewhat f oveolate groove ;
declivity transversely ridged; side-pieces with irregular longi-
tudinal striae which tend to be coarser below. Mesopleurum
shining, obscurely alutaeeous and punctate ; upper fovea large
and continuing on down the posterior margin of the mesopleu-
rum ; lower fovea fairly well defined, but its upper margin
obsolete on the middle third (Fig. 39). Front femora 2.2 x as
EVANS : REVISION OP RHABDEPYRIS 135
long a.s wide; claws dentate, the tooth sloping outward (Fig.
63).
Paratypes. — BRAZIL : 5 9 ? , 12 $ S , same data as type
except various dates, February, j\Iarch, April, June, Septem-
ber-December, 1938, 1952, 1953, 1961, 1962, 1964 [MCZ, BMNH,
BSA, Coll. G. R. Ferguson] ; 1 c5 , Corumba, May [ANSP] ;
1 S , Santarem (no further data) [USNM]. PERU: 1 $ , Upper
Rio Paehitea, 21 July 1920 (Cornell U. Exp.) [CU]. ECUA-
DOR: 1 6 , Ongota, 8 km SE Tena, May 1963 (L. Peiia) [MCZ].
PANAMA: 1 9, Puerto Bello, 21 February 1911 (E. A.
Schwarz) [USNM] ; 1 £ , Barro Colorado IsL, Canal Zone, 18
July 1956 (C. W. & M. E. Rettenmeyer) [KSU].
Variation. — The topotypic female paratypes vary consid-
erably in size (LFW 3.3-4.0 mm) ; WH varies from 1.00 to 1.02
X LH. The amount of infuscation of the femora is somewhat
variable, but otherwise there is close agreement with the type.
The Panama female has the apical .4 of the abdomen bright
ferruginous; in this specimen LFW is 3.8 mm, WH/LII is
1.02, WF/HE 1.17. Otherwise the resemblance to the type is
close.
The males show considerable variation and I concede the
possibility that more than one species may be involved. LP"'W
varies from 2.8 to 3.7 mm. Color of the head and thorax varies
from olive -green through blue-green to a shining, metallic blue ;
the intensity of the metallic reflections also varies considerably.
In some of the topotypes, WOT exceeds OOL slightly ; in one of
them the front legs are wholly pale ferruginous, while in others
the legs are mostly blackish and the antennae darker than in
the allotype. The Panama male has the apical third of the abdo-
men bright ferruginous. This specimen and those from Peru,
Ecuador, and Santarem have the front relatively narrow (WF
1.15-1.25 X HE), but in the Corumba specimen the front is
broad (WF 1.40 x HE) ; in fact the head itself is unusually
broad (WH 1.20 x LH). The last-named specimen also has
relatively coarser sculpturing on the propodeum, especially on
the side-pieces, and in lateral view the anterior margins of the
foveae on the posterior angles can be seen to be produced up-
ward as tooth-like processes. However, tendencies in these direc-
tions can be seen in the Panama specimen; indeed, there is
much variation in details of propodeal sculpturing throughout
the entire series, although the shape of the propodeal disc is
quite constant (1.35-1.45 x as wide as long).
136 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
The variation in the color of the abdomen in this species
parallels similar variation in luteipennis and muscarius, and
the other variation is not inconsistent with the patterns in those
species.
34. Rhabdepyris (Chlorepyris) viridis (Cameron)
Epyris viridis Cameron, 1888, Biol. Centr.-Amer., Hymen. I, p. 451 [Type:
5, GUATEMALA: San Juan, Vera Paz (G. C. Champion) (BMNH)].
Ehabdepyris viridis Kieffer, 1908, Gen. Insect., 76:32.
Anisepyris viridis Kieffer, 1914, Das Tierreich, 41 : 439.
Description of female type. — Length 5.5 mm ; LFW 3.0 mm.
Head, pronotum, and mesopleura bright bluish green ; mesono-
tum bright blue; propodeum black; abdomen shining black
except each segment with a narrow, apical testaceous band
(more noticeable ventrally) and the apical 1.5 segments mostly
testaceous; palpi light brown; mandibles blackish; antennae
black, scape with green reflections; middle and hind coxae and
femora dark brownish-fuscous, with weak metallic reflections;
tibiae and tarsi medium brown; wings subhyaline, with a faint
yellowish tinge, with a strong brownish cloud just below the
stigma and radial vein. Mandibles appearing tridentate, but
probably not differing materially from those of related species.
Clypeus with an angulate median lobe. Head about as wide as
high; inner orbits subparallel below, WF .69 x WH, 1.18 x HE;
ocelli in a compact triangle, front angle much less than a right
angle ; OOL 1.4 x WOT. Vertex passing straight across a
distance above eye tops equal to about .3 x HE. Front uni-
formly but very weakly alutaceous, strongly shining, punctures
strong, separated by 2-3 x their own diameters. First four
antennal segments in a ratio of about 32 :10 :7 :10, segment three
slightly wider than long, segment eleven about as long as
wide.
Pronotal disc .57 x as long as its posterior width, 1.25 x as
long as mesoscutum ; surface of pronotum shining, like the front,
but the punctures somewhat weaker than on the front. Meso-
scutum also shining, with a few punctures on the posterior half
but otherwise smooth, not depressed; notauli complete, diverging
and becoming more slender anteriorly; scutellar groove wiaened
and deflected backward on each end. Propodeal disc only 1.15
X as wide as long, with five strong carinae, the surface trans-
versely striate between the carinae, elsewhere shining ; declivity
EVANS : REVISION OF RHABDEPYBIS 137
transversely striate, more especially below. Mesopleurum aluta-
ceous although somewhat shining; upper fovea small, lower
fovea large, broadly incomplete above (essentially as in virescens,
Fig. 33). Claws dentate; front femora 1.8 x as long as their
greatest width.
Remarks. — This distinctive species has not been recovered
since its description in 1888.
35. Rhabdepyris (Chlorepyris) metallicus Kieffer
Bhabdepyris metallicus Kieffer, 1908, Ann. Soc. Sci. Bruxelles, 32: 10
[Type: ?, NICAEAGUA: Granada (? location of type)]. —Kieffer,
1914, Das Tierreich, 41: 360.
Original description (translated and paraphrased). — ^ Length
4 mm. Head metallic blue, pro- and mesonota metallic green
with bluish reflections ; abdomen black except last two segments
red; mandibles and palpi red; scape blackish brown, antennae
otherwise red; legs red except coxae black; wings brownish.
Head subcircular, smooth and shining, with dense, fine punc-
tures. Eyes separated from vertex crest by about one-third x
HE. Third and fourth antennal segments somewhat trans-
verse, the following segments thicker, at first transverse, then
somewhat longer. Thoracic dorsum shining; pronotum punctate
like the head; mesoscutum somewhat shorter than pronotum,
almost impunctate ; notauli wide on the posterior part, divergent
and becoming very thin anteriorly. Scutellum impunctate, its
groove broad and arcuate. Propodeum hardly transverse, with
five discal carinae of which the middle three are parallel, the
lateral ones oblique.
Remarks. — I have not been able to locate the type of this
species, and I have not been able to recognize the species from
the description. I would judge it to be closely related to the
preceding and the following species. The metallic colors are
apparently much more intense than in subviridis, the antennae
and legs much lighter in color than in viridis.
ViRIDISSIMUS species-group
This group includes six brilliantly green or blue species,
all having the scutellar groove in the form of paired pits con-
nected by a slender groove {Chlorepyris in the sense of Kieffer).
The head is much wider than high in both sexes and is little
developed anteriorly, the antennae arising only slightly below
138 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the bottoms of the eyes. It is possible that metallicus Kieffer
belongs here, but since Kieffer states that the head of that species
is "fast kreisrund, " I assume that it may belong near viridis
in the preceding species-group. In the viridissimus group the
claws of the male tend to be more or less trifid ; however, there is
no sharp morphological gap between the males of these two
species-groups.
36. Rhabdepyris (Chlorepyris) subviridis (Kieffer) new com-
bination
Epyris subviridis Kieffer, 1911, Ann. Soc. Sei. Bruxelles, 35: 225-226
[Type: $, MEXICO: Teapa, Tabasco, March (H. H. Smith)
(BMNH)].
Chlorepyris subviridis Kieffer, 1913, Boll. Lab. Zool. Portici, 7 : 108.
— Kieffer, 1914, Das Tierreich, 41: 415.
Plesiallotype. — ? , PANAMA : Pacora, Canal Zone, 14 May
1953 (F. S. Blanton) [USNM].
Description of female plesiallotype. — Length 5.5 mm ; LFW
3.5 mm. Head and thorax black, with dull olive-green reflections ;
propodeum black ; abdomen shining black, apical .4 ferruginous ;
palpi straw-colored ; mandibles light castaneous, the teeth darker ;
antennae uniformly light castaneous; tegulae testaceous; coxae
black ; hind femora and outer side of front femora dark brown,
legs otherwise testaceous; wings faintly tinged with yellowish
brown, slightly darker around the radial vein, veins yellowish
toward base of wing. Mandibles with a large subapical tooth
on the lower margin, otherwise with five teeth, the basal three
teeth small and rounded (Fig. 52). Clypeus obtusely subangulate,
extending barely beyond the antennal insertions. WH 1.10 x LH ;
WF .62 X WH, 1.06 x HE; front angle of ocellar triangle much
less than a right angle, OOL 1.6 x WOT. Vertex nearly straight,
distance from eye tops to vertex crest about one-third x HE.
Front strongly shining, very weakly alutaceous, the punctures
strong, separated from one another by 1-3 x their own diam-
eters below, much more widely spaced above. Antennae arising
only slightly below bottoms of eyes, first four segments in a ratio
of about 28 :7 :5 :10, segment three about .6 as long as wide, seg-
ment four slightly longer than wide, segment eleven about as
long as wide. (Fig. 15.)
Thoracic dorsum alutaceous, not as strongly shining as the
front, the punctures small and well spaced; pronotum about 1.25
EVANS : REVISION OF RHABDEPYRIS 139
X as long as mesoscutum along midline. Notauli elongate, attenu-
ate and diverging anteriorly; scutellar groove very narrow
medially, connecting two oblique, elliptical pits. Propodeal disc
1.30 X as wide as long, with five discal carinae, smooth and
polished between the lateral discals and the laterals ; declivity
polished, with some transverse ridges below ; side-pieces shining
and with very fine surface sculpturing. MesoiDleurum alutaceous
and with numerous small punctures ; upper fovea small, slender ;
lower fovea broadly incomplete above (Fig. 34). Front femora
2.1 X as long as wide; middle tibiae spinose; claws dentate, the
tooth fairly long and sloping outward somewhat.
Description of male type. — Length 5.0 mm; LFW 2.8 mm.
Head and thorax black, with moderately intense dark blue-green
reflections ; propodeum black ; abdomen shining black except
apical two segments suffused with brown; palpi and mandibles
straw-colored, the mandibular teeth rufous ; antennae light cas-
taneous, all except the apical segment somewhat infuscated
above, scape especially so ; tegulae testaceous ; legs bright ruf o-
eastaneous except all coxae blackish ; wings subhyaline, veins
and stigma brown. Mandibles with the usual five teeth ; clypeus
obtusely svibangulate, with a low median ridge. WH 1.18 x LH ;
WF .70 x AVH, 1.43 x HE; front angle of ocellar triangle
slightly less than a right angle, OOL 1.10 x WOT. Vertex
broadly rounded off a short distance above the eye tops. Front
strongly shining, weakly alutaceous, punctures small but sharply
defined, on lower part separated by 2-3 x their own diameters,
on upper part by 4-6 x their own diameters. First four antennal
segments in a ratio of about 20:10:6:11, segment three .6 as long
as wide, segment four about as long as thick, segment eleven
2.5 x as long as thick ; flagellar pubescence pale and erect, longest
setulae of segment eleven about half as long as width of seg-
ment (Fig. 73).
Thoracic dorsum shining, weakly alutaceous, the punctures
weak ; pronotal disc slightly longer than mesoscutum ; notauli
very wide behind, much narrowed anteriorly, reaching anterior
margin of scutum as thin lines; scutellar groove wider medially
than in the female, widened and deflected backward at each
end. Propodeal disc 1.35 x as wide as long, with five discal
carinae, only the median one intercepting the transverse carina;
lateral and sublateral carinae strong, disc also with a round-
topped ridge between the sublaterals and the lateral discals and
mostly covered with transverse striae ; declivity transversely
140 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
ridged below ; side-pieces with strong longitudinal ridges, more
especially below. Mesopleurum shining, weakly alutaceous; up-
per fovea open behind, as in the preceding species ; lower fovea
broadly open above. Claws trifid (Fig. 65).
Other specimens examined. — HONDURAS: 1 9, "Banana
debris, 3-25-40, lot no. 40-6469" (presumably intercepted at
quarantine) [USNM]. COSTA RICA: 1 S, Turrialba, 17
June 1949 (K. AV. Cooper) [USNM]. PANAMA: 2 $S,
Pacora, Canal Zone, same data as plesiallotype except 13 May
1953 and July 1953 [USNM].
Variation. — The Honduras female is strikingly like the female
from Panama; WF is 1.13 x HE; OOL is 1.53 x WOT; the
upper mesopleural fovea is even more elongate. The males are
all slightly larger than the type (LFW 3.2-3.3 mm) ; the Pacora
specimens have paler antennae, only very slightly infuscated
above, and these specimens also have the hind femora infus-
cated. In the males, WII varies from 1.13 to 1.18 x WH, WF
from 1.33 to 1.38 x HE, OOL from 1.13 to 1.22 x WOT. In the
Pacora males the antennae are more elongate than in the
Turrialba and Teapa males, segment four being distinctly longer
than wide, segment eleven more than 3 x as long as wide.
37. Rhabdepyris (Ciilorepyris) blantoni new species
Holotype. — $ , PANAMA : Pacora, Canal Zone, 13 May
1953 (F. S. Blanton) [USNM, No. 67,541].
Description of male type. — Length 5.5 mm ; LFW 3.8 mm.
Head and thorax black, shining, with moderately strong bluish
green reflections ; propodeum black, with faint dark blue reflec-
tions in certain lights ; abdomen shining black, the apical .4
bright ferruginous; palpi and mandibles testaceous, the latter
dark at tips; antennae bright rufo-testaceous except the scape
strongly infuscated ; tegulae testaceous ; coxae black, femora
medium brown, legs otherwise testaceous; wings lightly tinged
with brownish, veins and stigma brown. Mandibles with the
usual five teeth; clypeus short, obtusely angulate. WH 1.09 x
LH ; WF .62 X WH, 1.10 x HE ; ocelli in about a right triangle,
OOL 1.1 X WOT. Front shining, weakly alutaceous except not
at all so on the vertex, with sharply defined punctures which
are separated by only 1-2 x their own diameters. First four
antennal segments in a ratio of about 28 :7 :4 :11, segment three
unusually short, only about .3 as long as wide, segment four
EVANS : REVISION OF RHABDEPYRIS 141
very slightly longer than wide, segment eleven about 1.5 x as
long as wide. (Fig. 23.)
Thoracic dorsum polished, non-alutaceous, wholly covered with
small punctures; notauli moderately wide behind, tapering and
diverging anteriorly; scutellum with a pair of large, strongly
oblique pits which are connected by a slender groove. Pro-
podeal disc 1.4 x as wide as long, with five carinae ; lateral and
sublateral carinae running closely parallel, the latter paralleled
by a broad, longitudinal groove; space between this groove
and the lateral discal carinae shining and with only some very
weak surface sculpturing ; declivity with transverse ridges below ;
side-pieces with moderately coarse longitudinal striae. Meso-
pleurum polished, punctate ; upper fovea elongate, open behind ;
lower fovea large, its upper margin obsolete (Fig. 40). Front
femora 2.3 x as long as wide ; middle tibiae weakly spinose ;
claws trifid, the middle ray truncate (Fig. 64).
Paratypes. — PANAMA: 5 $ $, same data as type [USNM,
MCZ].
Variation. — Little variation is evident in this series. There
is some slight variation in the hue of the metallic colors of the
head and thorax and in the degree of inf uscation of the femora ;
in some specimens the bluish reflections of the propodeum are
barely evident. In the paratypes, LFW varies from 3.1 to 3.5,
WH from 1.06 to 1.09 x LH, WF from 1.1 to 1.2 x HE, OOL
from 1.0 to 1.1 X WOT.
Remarks. — It is possible that these specimens represent the
male sex of metallicus Kieffer or tricolor n.sp., but there are
enough doubts in my mind to justify recognizing them as a
distinct species for the present. The very short third antennal
segment is unique in this subgenus.
38. Rhabdepyris (Chlorepyris) viridissimus (Kieffer) new
combination
Epyris viridis Kieffer, 1911, Ann. Soc. Sci. Bruxelles, 35: 223-225 [Type:
$, MEXICO: Teapa, Tabasco, March (H. H. Smith) (BMNH)]. Pre-
occupied by Cameron, 1888.
Epyris viridissimus Kieffer, 1911, ibid., p. 225 [Type: S, MEXICO:
Teapa, Tabasco, March (H. H. Smith) (BMNH)].
Chlorepyris semiviridis Kieffer, 1913, Boll. Lab. Zool. Portici, 7: 108 (new
name for viridis Kieffer, not Cameron). — Kieffer, 1914, Das Tier-
reich, 41 : 414. New synonymy.
Chlorepyris viridissimus Kieffer, 1914, ihid., p. 415.
142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Description of female type of semiviridis Kieffer. — Length
7.8 mm; LFW 4.4 mm. Head and thorax brilliant green; pro-
podeum black; basal 3.5 segments of abdomen shining black, re-
mainder of abdomen bright rnfo-castaneous ; palpi light brown ;
mandibles ferruginous; antennae bright rufo-castaneous, slightly
paler apically than basally; tegulae with greenish reflections;
legs bright rufo-castaneous except all coxae suffused with black ;
fore wing lightly tinged with yellowish, basal third and apical
.4 lightly infuscated, leaving a subhyaline area below the stigma.
Mandibles with five apical teeth and a strong subapical tooth on
the lower margin (Fig. 53). Clypeus short, obtusely angulate,
its median ridge slightly concave in profile. WII 1.2 x LH; WF
.61 x WH, 1.10 X HE ; ocelli in a small triangle, front angle less
than a right angle, OOL 1.4 x WOT. Front polished, obscurely
alutaceous, punctures strongly defined, separated by 2-4 x their
own diameters. First four antennal segments in a ratio of about
22 :6 :5 :7, segment three about .7 x as long as wide, segments
four and eleven each about as long as wide.
Thoracic dorsum shining, obscurely alutaceous, and with well
spaced strong punctures like the front ; notauli moderately wide
behind, tapering and diverging anteriorly ; scutellar groove rela-
tively narrow, deflected backward and slightly widened at each
end. Propodeal disc 1.4 x as wide as long, the five discal carinae
all reaching the transverse carina; lateral and posterior carinae
margined within by broad, subfoveolate grooves; surface trans-
versely ribbed between discal carinae, elsewhere with weak sculp-
turing. Mesopleurum polished, weakly alutaceous, with small,
well-spaced punctures ; upper fovea about 3 x as long as high ;
lower fovea large, broadly open above. Front femora 2.2 x
as long as wide ; middle tibiae strongly spinose ; claws trifid
(Fig. 66).
Description of male type. — Length 6 mm; LFW 3.6 mm.
Head and thorax brilliant green ; propodeum black ; basal
4.5 segments of abdomen shining black, remainder bright rufo-
castaneous ; palpi straw-colored ; mandibles rufo-testaeeous, the
teeth darker ; clypeus brownish ; antennae with the scape fuscous,
the flagellum pale rufo-castaneous except basal few segments
weakly infuscated on upper side; tegulae brown, with blue-
green reflections; legs bright rufo-castaneous except all coxae
blackish ; wings very lightly infuscated, more especially so
around radial vein of fore wing. Mandibles 5-toothed; clypeus
verv short, obtusely angulate. WH 1.20 x LH ; WF .61 x WH,
EVANS : REVISION OF RHABDEPYRIS 143
1.13 X HE; front angle of oeellar triangle very slightly less
than a right angle, OOL 1.05 x WOT. Front shining, obscurely
alutaceous, the punctures strong, somewhat more closely spaced
than in the female, separated by 1-3 x their own diameters.
First four antennal segments in a ratio of about 28 :6 :8 :13,
segment three about .7 as long as thick, segment four about as
long as thick, segment eleven about twice as long as thick.
Thoracic dorsum weakly alutaceous, wholly covered with rather
large punctures ; notauli Avide behind, separated by about their
own widths, diverging and attenuate anteriorly ; scutellar groove
narrow, connecting two large, oblique pits. Propodeal disc 1.4 x
as wide as long, with five diseal carinae ; lateral and sublateral
carinae closely parallel, the latter paralleled by a shallow groove
along its mesal margin ; postero-lateral foveae very strong ; side-
pieces with 8-10 coarse, irregular ridges. Mesopleurum alu-
taceous, with shallow punctures ; upper fovea elongate, open
behind (Fig. 41) ; lower fovea not well defined above. Front
femora 2.4 x as long as wide; claws trifid (Fig. 67).
Other specimens examined. — MEXICO : 1 9 , Mazatlan, Sina-
loa, 20 July 1959 (H. E. Evans) [MCZ] ; 1 9 , Cordoba, Vera-
cruz, 20 January 1908 (F. Knab) [USNM] ; 3 $ i , Teapa,
Tabasco (same data as type) [BMNH]. GUATEMALA: 1 9,
"Guatemalan forest," 3 March 1909 (F. Knab) [USNM].
Variation. — The females vary considerably in size (LFW
4.0-5.3 mm) but differ in color only in having the green colora-
tion somewhat blue-green in the Cordoba specimen. This same
specimen also has the punctures of the head and thorax some-
what weaker and more widely spaced, also the scutellar groove
very narrow indeed, connecting two large, oblique pits, more as
in the male. Since the other two females show some variation in
the scutellar groove, I doubt if the Cordoba specimen is more
than an extreme variant in this regard. The Mazatlan specimen
is the largest, being fully 9 mm long, and has the thoracic dor-
sum and pleura much more evidently alutaceous than do the
other females. In the females, WH varies from 1.3 to 1.20 x LH,
WF from 1.10 to 1.25 x HE, OOL from 1.4 to 1.6 x WOT.
The three topotypic males resemble the type closely, but some
are more blue-green than green, and the head tends to be
slightly less broad in some (WH 1.15-1.20 x LH; WF 1.10-1.20
X HE).
144 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
39. Rhabdepyris (Chlorepyris) fulgens (Brues) new combi-
nation
Epyris fulgens Brues, 1907, Bull. Wise. Nat. Hist. Soc, 5: 99-100 [Type:
9, TEXAS: Brownsville, 4 August (C. Sehaefeer) (USNM, No.
42,702)].
Chlorepyris fvlgens Kieffer, 1914, Das Tierreieh, 41 : 414. — Muesebeck and
Walkley, 1951, U.S. Dept. Agr. Monogr. 2, p. 730.
Description of female type. — Length 7.0 mm; LFW 4.0 mm.
Head and thoracic dorsum green, with bluish green reflections
in certain lights ; mesopleurum aeneous, grading into bluish
below; propodeum violaceous; basal three abdominal segments
and basal part of fourth tergite black, remainder of abdomen
rufo-testaceous ; palpi testaceous ; mandibles testaceous, the teeth
rufous ; antennae pale ferruginous except somewhat dusky on
outer surface, segments two and three wholly dusky ; tegulae
with greenish reflections ; legs bright rufo-testaceous except front
coxae black, hind coxae Aveakly infuscated basally; wings sub-
hyaline, veins and stigma light brown. Mandibles as in the
preceding species and as sho^vn in Figure 54 ; clypeus broadly
angulate, the median ridge concave in profile. WH 1.16 x LH;
WF .63 X WH, 1.23 x HE ; front angle of oeellar triangle slightly
less than a right angle, OOL 1.5 x WOT. Front strongly pol-
ished, obscurely alutaceous below, with a linear median impres-
sion opposite the lower third of the eyes ; punctures of front
strong, separated by 2-4 x their own diameters. First four
antennal segments in a ratio of about 18:5:4:6, segment three
considerably wider than long, segment four slightly longer than
wide (Fig. 74).
Thoracic dorsum shining, obscurely alutaceous; punctures of
pronotum strong, separated by 2-4 x their own diameters, those
on the mesoscutum occurring mostly along the notauli ; notauli
linear, only slightly widened behind, complete ; scutellar groove
narrow, much widened on each side. Propodeal disc 1.4 x as
wide as long, with five discal carinae between which it is trans-
versely ribbed, the space between the lateral discals and the
laterals smooth and merely ver}^ finely striolate ; side-pieces
finely striolate. Mesopleurum weakly alutaceous, with small
punctures ; upper fovea about twice as long as high ; lower fovea
broadly incomplete above. Front femora twice as long as wide ;
middle tibiae strongly spinose, hind tibia more weakly so;
claws trifid (Fig. 68).
EVANS : REVISION OF RHABDEPYBIS 145
Plcsiallotijpe.— $, HONDURAS: Tegucigalpa (F. J. Dyer)
[USNM].
Description of male plesiallotypc. — Length 5.5 mm; LFW
3.9 mm. Head and thorax brilliant green, with the thorax with
some bluish tints ; propodeum bluish green, somewhat less bril-
liant than the head and thorax, especially on the side-pieces;
abdomen black, apical .4 rufo-testaceous ; palpi and mandibles
testaceous, the latter with rufous tips; scape fuscous, fiagellum
rufo-testaceous except basal segments moderately infuscated on
upper surface ; tegulae brown, with green reflections ; legs bright
rufo-testaceous except all coxae infuscated ; wings tinged with
brownish, veins and stigma brown. Clypeus obtusely angulate,
its median carina concave in profile. "WH 1.20 x LH; WF .63
X AYH, 1.24 x IIE ; front angle of ocellar triangle very slightly
exceeding a right angle, OOL and WOT subequal. Vertex
straight across in center, broadly rounded on the sides. Front
polished, obscurely alutaceous below but non-alutaceous above,
punctures strong, separated by 1-3 x their own diameters. First
four antennal segments in a ratio of about 25 :7 :8 :13, segment
three .7 as long as wide, segment four very slightly longer than
wide, segment eleven 1.8 x as long as wide. (Fig. 24.)
Thoracic dorsum shining, wholly covered with fairly strong
punctures ; notauli wide behind, separated by slightly less than
their own widths, strongly diverging and attenuate anteriorly ;
scutellar groove strong, much widened at each end. Propodeal
disc 1.4 x as wide as long, with five carinae between which it is
strongly transversely ridged; lateral and sublateral carinae
closely parallel, the sublaterals subtending a broad groove which
occupies nearly half the space between these carinae and the
lateral discals ; postero-lateral foveae very large ; side-pieces with
only about six large, longitudinal ridges. Mesopleurum with the
upper fovea large, open behind; lower fovea large, well defined,
its upper margin partially incomplete (Fig. 42). Front femora
2.4 X as long as wide ; claws trifid, about as in viridissimus
(Fig. 67).
Remarks. — This species is known to me only from the two
specimens described above.
40. Rhabdepyris (Chlorepyris) tricolor new species
Holotype. — 9, ECUADOR (label reads: on roses ex Ecua-
dor, VII-29-1962; F. J. Formichella, lot 62-26784) (presum-
ably intercepted at quarantine) [USNM, No. 67,542].
146 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Description of female type. — Length 6.0 mm; LFW 3.5 mm.
Head, thorax, and propodeum brilliant dark blue-green, outer
side of front femora also of this color ; abdomen black except
apical .4 bright ferruginous; palpi and mandibles testaceous
except the latter darker at extreme base and apex; antennae
fuscous except segments 4-12 testaceous beneath, the apical
segment wholly testaceous; coxae black, legs otherwise bright
ferruginous except outer side of front femora dark like the
body; wings tinged with brown, the base of the fore wing, in-
cluding the veins, somewhat yellowish. Mandibles essentially
as figured for the preceding species (Fig. 54). Clypeus obtusely
angulate, barely protruding beyond the strong antennal lobes.
WH 1.12 X LH • WF .64 x WH, 1.18 x HE ; front angle of ocellar
triangle less than a right angle, OOL 1.7 x WOT. Vertex
straight, distance from eye tops to vertex crest equal to about
one-third x HE. Front polished, very obscurely alutaceous, with
strong punctures which are separated, in the center of the front,
by 2-4 X their own diameters (more crowded beloAv, le.ss so
above). Antennae arising only slightly below bottoms of eyes,
first four segments in a ratio of about 30 :7 :6 :9, segment three
considerably wider than long, segment four about as long as
wide, segments 5-12 very slightly wider than long. (Fig. 16.)
Thoracic dorsum shining but slightly more evidently alu-
taceous than the front ; pronotum with numerous fairly strong
punctures, slightly longer than mesoscutum. ]\Iesoscutum with
a few punctures along the parapsidal furrows and notauli, the
latter complete but reduced to thin lines on the anterior half;
scutellar groove a mere line connecting two oblique, elliptical
pits. Propodeal disc 1.35 x as wide as long, with five discal
carinae between which it is transversely striate and laterad of
which there is weak surface sculpturing ; sublateral carinae not
distinct, but lateral carinae paralleled by a broad, transversely
striate groove ; declivity with transverse ridges below ; side-pieces
shining, with weak longitudinal aciculations. Mesopleurum
weakly alutaceous, weakly punctate ; upper fovea extending most
of the length of the mesopleurum, nearly 5 x as long as wide ;
lower fovea large, its upper margin mostly indistinct (Fig. 35).
Front femora moderately incrassate, measuring 2.0 x as long as
wide; middle and hind tibiae strongly spinose; claws trifid (as
figured for fulgens, Fig. 68).
Bemarks. — This species is known only from the type. It
EVANS : REVISION OF RHABDEPYRIS 147
seems definitely to belong to this complex of Bhahdepyris, al-
though the scutellar pits differ hardly at all from those of some
species of Epyris.
41. Rhabdepyris (Chlorepyris) violaceus new species
Holotype. — 9 , BRAZIL : Nova Teutonia, Santa Catarina, 30
August 1938 (F. Plaumann) [BMNH].
Descriptio7i of female type. — Length 7.5 mm ; LFW 4.5 mm.
Head and thorax, including scape, tegulae, and outer sides of
femora, brilliant blue-violet, with greenish tints in certain lights ;
propodeum dark blue, contrasting slightly to head and thorax;
abdomen black except apical tergite brownish ; mandibles black
except apical fourth ruf o-testaceous ; antennae fuscous except
front tibiae and all the tarsi light brown; wings lightly tinged
with brown, veins and stigma brown. Mandibles with five apical
teeth and a large subapical tooth on the lower margin. Clypeus
obtusely subangulate, barely produced beyond antennal inser-
tions. WH 1.17 X LH; WF .65 x WH, 1.20 x HE; front angle
of ocellar triangle much less than a right angle, OOL 1.60 x
WOT. Vertex broad, straight; distance from eye tops to vertex
crest equal to about .3 x HE. Front shining, weakly alutaceous,
with large punctures which are separated by only 1-2 x their
own diameters (more widely spaced at level of ocelli). First
four antennal segments in a ratio of about 40 :9 :8 :12, segment
three very slightly wider than long, segments four and eleven
both approximately as wide as long.
Thoracic dorsum weakly alutaceous and punctate like the
front ; notauli complete, rather slender and only slightly attenu-
ate anteriorly; scutellar grooves very thin, connecting a pair of
small lateral pits. Propodeal disc 1.35 x as wide as long, the five
discal carinae parallel and complete, the surface weakly trans-
versely ridged between the carinae, elsewhere with only very
weak surface sculpturing ; side-pieces shining, with extremely
fine longitudinal striations. Upper mesopleural fovea about 3 x
as long as high, lower fovea broadly incomplete above (Fig. 36).
Front femora only slightly incrassate, measuring 2.4 x as long
as wide; claws trifid, middle ray truncate (Fig. 69).
Allotype. — S , BRAZIL : Rio Caraguata, Matto Grosso,
May 1953 (F. Plaumann) [MCZ].
Description of male allotype. — Length 6.0 mm ; LFW 3.5 mm.
Coloration as described for female, except with slightly stronger
148 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
greenish reflections, even on the propodeum, which contrasts
very little with the thorax; abdomen entirely black; mandibles
brownish, paler apically; antennae fuscous except flagellar seg-
ments testaceous beneath ; legs and wings colored as in female.
Clypeus broadly subangulate. WH 1.18 x LH ; WF .64 x WH,
1.28 x HE ; ocellar triangle rather broad, WOT exceeding OOL
slightly. Front polished, non-alutaceous, punctures strong, sep-
arated by 1.5-3 X their own diameters. First four antennal seg-
ments in a ratio of about 20 :6 :8 :11, segment three considerably
wider than long, segment four slightly longer than wide, seg-
ment eleven about twice as long as wide.
Thoracic dorsum slightly alutaceous, with strong punctures
like the front ; notauli very wide behind, separated by only a
thin carina, diverging and attenuate anteriorly ; parapsidal fur-
rows strong and complete, somewhat sinuate ; scutellar disc
rather flat, the basal groove rather short, expanded on each side
to form oblique pits. Propodeal disc 1.3 x as wide as long, some-
what narrowed behind; disc with only three carinae, between
which it is transversely ridged, sublateral carinae strong, sub-
tending a broad groove which occupies about half the space be-
tween the sublaterals and the lateral discals; this groove, like
that between the sublateral and lateral carinae, is polished and
without sculpturing; side-pieces with only six strong, longi-
tudinal ridges. Mesopleurum wnth the upper fovea very large,
open behind. Claws trifid (Fig. 70).
Paratypes. — 2 5 9, BRAZIL : same data as type except
dated 10 September 1938 and 21 December 1937 [BMNH, MCZ].
Variation. — In both paratypes the propodeum has some
greenish reflections and therefore does not contrast M-ith the head
and thorax as much as in the type. Otherwise these females are
very similar to the type in color, size, and standard measure-
ments. In both specimens the ocelli are slightly less far removed
from the eye margins, OOL being 1.43 and 1.50 x WOT.
Remarks. — This is one of the most brilliantly colored of all
bethylid wasps and doubtless the most highly evolved member
of this subgenus.
ALPHABETICAL LIST OF ABBEEVIATIONS USED IN TEXT
Structures
HE: height of eye (maximum, lateral view)
LFW : length of fore wing
EVANS : REVISION OF RHABDEPYRIS 149
LH : length of head (apical margin of clypeus to median vertex crest)
OOL: oeello-ocular line (minimum distance from eye to lateral ocellus)
WF: width of front (measured at its minimum point)
WH: width of head (maximum, including eyes)
WOT: width of ocellar triangle (including lateral ocelli)
Institutions
AMNH : American Museum of Natural History, New York
ANSP: Academy of Xatural Sciences of Philadelphia
BMNH: British Museum (Natural History), London
BSA : Brasil: Secretaria da Agricultura, Sao Paulo
CAS : California Academy of Sciences, San Francisco
CIS : California Insect Survey, Berkeley
CM: Carnegie Museum, Pittsburgh, Pa.
CU : Cornell University, Ithaca, N.Y.
ENAC : Eseuela Nacional de Agricultura, Chapingo, Mexico
HCOU : Hope Collections, Oxford University, England
INHS : Illinois Natural History Survey, Urbana
KU: Kansas University, Lawrence
KSU : Kansas State University, Manhattan
MCZ : Museum of Comparative Zoology, Cambridge, Mass.
UA: University of Arizona, Tucson
UCD : University of California, Davis
USNM : United States National Museum, Washington
LIST OF SPECIES OF REABDEPYE18
OCCURRING IN THE AMERICAS
Subgenus Bhabdepyris Kieffer, 1904
1. mellipes Evans, n. sp. ( 2 , Florida)
2. huachucae Evans, n. sp. (9, Arizona)
3. muesebecJci Evans, n. sp. ( 5 , S , Mexico to Bolivia)
4. gracilis Evans, n. sp. (9 , S , California ; Durango, Mexico)
5. minutulus Evans, n. sp. ( ? , Peru)
6. nigriseapus Evans, n. sp. ( 9 , Argentina)
Subgenus Tricliotepyris Kieffer, 1906
Nigropilosus species-group
7. nigropilosus Evans, n. sp. ( 9 , Panama, Brazil)
Megacephalus species-group
8. megacephalus (Ashmead), 1893 (9,5, California to Texas)
9. werneri Evans, n. sp. (9, $ , Arizona)
10. apaclie Evans, n. sp. ( 9 , S , Arizona; Sonora, Mexico)
11. texanus Evans, n. sp. (9, $, Texas and Arizona to Morelos, Mex-
ico)
12. mexicanus Evans, n. sp. ( 9 , i , Morelos and Chiapas, Mexico)
13. fortunatus Evans, n. sp. ( 9 , Costa Rica)
150 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
14. lupus Evans, n. sp. ( $, Morelos, Mexico)
15. carolinianus Evans, n. sp. ( $ , South Carolina, Florida)
16. subaeneus Kieffer, 1906 ( 9 , Nicaragua)
17. angusticeps Evans, n. sp. ( $ , Arizona)
18. olivaceus Evans, n. sp. ($ , Panama, Colombia)
19. plaumanni Evans, n. sp. ( 5 , $ , Brazil)
Puldhripennis species-group
20. pulcliripennis Evans, n. sp. ( $ , Costa Eica)
21. iridescens Evans, n. sp. ( 9 , Morelos, Mexico)
22. cupreolus Evans, n. sp. ( $ , Brazil)
23. amahilis Pouts, 1927 ( 9 , <? , Massachusetts to Florida)
Subgenus Chlorepyris Kieffer, 1913
Loiatifrons species-group
24. luteipennis Evans, n. sp. ( $ , S , Brazil to Nicaragua)
25. lohatifrons Kieffer, 1910 ( $ , Brazil)
Synonym: ohscuripennis Kieffer, 1910
26. septemUneatus Kieffer, 1906 ( $ , Nicaragua, Panama)
27. quinquelineatus Kieffer, 1906 (9,5, Mexico to Nicaragua)
28. nigerrimus Evans, n. sp. ( 5 , Bolivia)
Musoarius species-group
29. musoarius (Westwood), 1874 (9,5, Mexico to Peru and Brazil)
Synonym: microstoma Kieffer, 1910
30. puncticeps Evans, n. sp. ( $ , Brazil)
31. origenus Kieffer, 1911 ( 9 , Mexico)
32. vesculus Evans, n. sp. ( $ , Brazil)
33. virescens Evans, n. sp. ( 9 , S , Panama to Peru and Brazil)
34. viridis (Cameron), 1888 ( 9, Guatemala)
35. metallicus Kieffer, 1908 ( 9 , Nicaragua)
Viridissimus species-group
36. suhviridis (Kieffer), 1911 (9,5, Mexico to Panama)
37. hlantoni Evans, n. sp. {$ , Panama)
38. viridissimus (Kieffer), 1911 (9,5, Mexico, Guatemala)
Synonjan : semiviridis Kieffer, 1913
39. fulgens (Brues), 1907 (9,5, Texas to Honduras)
40. tricolor Evans, n. sp. ( 9 , Ecuador)
41. violaceus Evans, n. sp. ( 9 , 5 , Brazil)
(Received May 28, 1964.)
Valid names
main references
amabilis, 110
angusticeps, 102
apache, 91
hlantoni, 140
caroUnianus, 100
cupreolus, 108
fortunatus, 97
fulgens, 144
gracilis, 76
huachucae, 73
iridescens, 107
lobatifrons, 120
lupus, 99
luteipennis, 118
megacephalvs, 85
melllpes, 72
metallious, 137
mexicanus, 96
microstoma, 126
minutulus, 77
muesebecki, 74
muscarius, 126
myrmecophilus, 70
INDEX
are printed in italics. Page numbers refer to
nigerrimus, 125
nigriscapns, 78
nigropilosus, 84
obscuripennis, 120
olivaceus, 103
origemis, 130
pallidipennis, 79
plaumanni, 104
pulchripennis, 106
puncticeps, 129
quinquelineatus, 123
semiviridis, 141
septemlineatus, 122
subaeneus, 101
subviridis, 138
texanus, 93
tricolor, 145
vesculus, 131
violaceus, 147
virescens, 133
viridis, 136
viridis, 141
viridissimus, 141
werneri, 89
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 1
Heads of female RJiaidepyris
Fig. 1. R. mellipes n. sp., type
Fig. 2. E nigriscapus n. sp., type
Fig. 3. B. nigropilosus n. sp., type
Fig. 4. R. werneri n. sp., type
Fig. 5, R. apache n. sp., type
Fig. 6. R. texanus n. sp., type
EVANS : REVISION OF RIIABDEPYRIS
I, mellipes
2 nigriscapus
3. nigropilosus
4. wernen
5 opache
6. texanus
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 2
Heads of female Bhabdepyris
Fig. 7. B. cnrolinianus n. sp., type
Fig. 8. B. angu-sticeps n. sp., type
Fig. 9. B. fortimatus, n. sp., type
Fig. 10. B. iridescens n. sp., type
Fig. 11. B. luteipennis n. sp., type
Fig. 12. B. nigerrimus n. sp., type
EVANS : REVISION OF RHABDEPYRIS
1. carolinionus
8. angusticeps
9. fortunatus
10. iridescens
II luteipennis
\Z. nigernmus
BULLETIN : MUSEUM OP COMPARATR^ ZOOLOGY
PLATE 3
Heads of Ehabdepyris (females and males)
Fig. 13. jB. muscarius (Westwood), plesiallotype female
Fig. 14. R. virescens n. sp., type female
Fig. 15. E. suiviridis (Kieffer), plesiallotype female
Fig. 16. B. tricolor n. sp., type female
Fig. 17. i?. werneri n. sp., allotype male
Fig. 18. R. texanus n. sp., allotype male
EVANS : REVISION OF RHABDEPYRIS
13 muscarius
14. virescens
15 subviridis
16. tricolor
^ly
17 werneri
18. lexonus
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 4
Heads of male BJiabdepyris
Fig. 19. B. musoarius (Westwood), specimen from Chapada, Brazil
Tig. 20. E. puncticeps u. sp., type
Fig. 21. R. virescens n. sp., allotype
Fig. 22. B. vesculus n, sp., type
Fig. 23. B. blantoni n. sp., type
Fig. 24. B. fulgens (Brues), plesiallotype
EVANS : REVISION OF RHABDEPYRIS
19 muscarius
20. puncticeps
21. virescens
22. vescufus
-z>
23. blantoni
24. fulgens"
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 5
Mesopleuia of female lihabdepyris
Fig. 25. B. werneri n. sp., type
Fig. 26. E. apaohe n. sp., type
Fig. 27. E. texanus n. sp., type
Fig. 28. R. carolinianus n. sp., type
Fig. 29. E. luteipennis n. sp., type
Fig. 30. R. nigerrimus n. sp., type
Fig. 31. E. musoarius (Westwood), plesiallotype
Fig. 32. E, origcnus Kieffer, type
Fig. 33. E. virescens n. sp., type
Fig. 34. E. subviridis (Kieffer), plesiallotype
Fig. 35. E. tricolor n. sp., type
Fig. 36. E. violaceus n. sp., type
EVANS : REVISION OF RIIABDEPYRIS
25 werneri
26 opache
27. texanus
28. carolinianus
29. lufeipennis
30. nigerrimus
31. muscorius
32. origenus
33. virescens
34. subviridis
35. tricolor
36. violoceus
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 6
Mesopleura of male Khabdepyris
Tig. 37. E. muscarius (Westwood), specimen from Chapada, Brazil
Fig. 38. B. vesculus n. sp., type
Fig. 39. E. virescens n, sp., allotype
Fig. 40. B. blantoni n. sp., type
Fig. 41. B. viridissimus (Kieffer), type
Fig. 42. B. fulgens (Brues), plesiallotype
Mandibles of female BJiabdepyris
Fig. 43. B. megacephalus (Aslimead), type
Fig. 44. B. iverneri n. sp., type
Fig. 45. B. fortunatus n. sp., type
Fig. 46. B. oarolinianus n. sp., type
Fig. 47. B. subaeneus Kieffer, type
Fig. 48. B. angusticeps n. sp., type
Fig. 49. B. luteipennis n. sp., type
Fig. 50. B. muscarius (Westwood), plesiallotype
Fig. 51. B. virescens n, sp., type
Fig. 52. B. subviridis (Kieffer), plesiallotype
Fig. 53. B. viridissimus (Kieffer), plesiallotype
Fig. 54. B. fulgens (Brues), type
EVANS : REVISION OF RIIABDEPYRIS
37 musconus
38 vesculus
39 virescens
40 blantoni
41 \/iridissimus
42 fulgens
43 megacephalus
44, wernen
45. fortunatus
46. carolinianus
47 subaeneus
48. angusticeps 49. luteipennis 50. muscorius
51. virescens
5Z. subviridls
53. viridissimus
54. fulgens
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 7
Claws of posterior tarsi of Ehabdepyris
Fig. 55. E. luteipennis n. sp., type female
Fig. 56. E. luteipennis u. sp., allotype male
Fig. 57. E. quinquelineatus Kieffer, type female
Fig. 58. E. nigerrimus n. sp., type female
Fig. 59. E. musearius (Westwoocl), plesiallotypc female
Fig. 60. E. musearius (Westwood), type male
Fig. 61. E. vesoulus n. sp., type male
Fig. 62. E. virescens n. sp., type female
Fig. 63. E. virescens n. sp., allotype male
Fig. 6-1. E. hlantoni n. sp., type male
Fig. 63. E. suiviridis (Kieffer), type male
Fig. 66. E. viridissimus (Kieffer), plesiallotype female
Fig. 67. E. viridissimus (Kieffer), tj-pe male
Fig. 68. E. fulgens (Brues), type female
Fig. 69. E. violaoeus n. sp., type female
Fig. 70. E. violaoeus n. sp., allotype male
Basal segments of antennae
Fig. 71. E. luteipennis n. sp., allotype male
Fig. 72. E. quinquelineatus Kieffer, plesiallotype male
Fig. 73. E. subviridis (Kieffer), type male
Fig. 74. E. fulgens (Brues), type female
EVANS : REVISION OP RHABDEPYRIS
cr:^ cz:^ C3^ c:::^^
55. luteipennis ? 56, luteipennis ^ 57 quinquelineatus ? 58. nigerrimus *
C^/
/
59. mjscarius ? 60. muscarius S 61. vesculus s
62. virescens s
63. virescens s 64. blantoni i 65. subviridis J 66. viridissimus ?
^^
C2^ CZ^ C2^
67. viridissimus cJ" 68. fulgens ¥ 69. violaceus ? 70. violaceus i
71 luteipennis cT
XDZI]
72. quinquelineatus d"
73. subviridis i
74. fulgens ?
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, Xo. 3
NELDASAUBU8 W RIGHT AE,
A NEW RHACHITOMOUS LABYRINTHODONT
FROM THE TEXAS LOWER PERMIAN
By John Newland Chase
Ohio Wesleyan University, Delaware, Ohio
With Five Plates
CAMBRIDGE, MASS., U.S.A,
PRINTED FOR THE MUSEUM
June 25, 1965
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MUSEUM OF COMPARATIVE ZOOLOGY
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Bulletin of the Museum of Comparative Zoology
HAKVAED UNIVEESITY
Vol. 133, No. 3
NELDA8AUBUS W BIGHT AE,
A NEW RHACHITOMOUS LABYRINTHODONT
FROM THE TEXAS LOWER PERMIAN
By John Newland Chase
Ohio Wesleyan University, Delaware, Ohio
With Five Plates
CAMBRIDGE, MASS., U.S.A.
FEINTED FOR THE MUSEUM
June, 1965
Bull. Mus. Comp. Zool., Harvard Univ., 133(3) : 153-225, June, 1965
No. 3. Neldasaurus wrightae, A New Rhachitomous
Labyrinthodont from the Texas Lower Permian^
By John Newland Chase
Department of Zoology,
Ohio Wesleyan University, Delaware, Ohio
TABLE OF CONTENTS
Introduction 156
Acknowledgements 157
Systematic Description 158
Generic and Specific Diagnoses 158
Descriptive List of Materials 159
Anatomical Description of Genus and Species 160
General Skull Morphology 160
Bones of the Dermal Skull Roof 163
The Sensory Canal System 168
Palate and Palatoquadrate 169
Parasphenoid and Braincase 176
Occiput 179
Lower Jaw 182
Vertebral Column 188
Ribs 193
Appendicular Skeleton 196
Scales 203
Other Occurrences of Neldasaurus 205
Discussion 206
Comparison of Neldasaurus with other Genera of
Trimerorhaehoids 206
Interrelationships of Trimerorhachoid Genera 215
Trimerorhachoid Origins and Relationships 220
Literature Cited 222
1 This paper is part of a thesis submittefl to the Department of Biology of
Harvard University as partial fulfillment of the requirements for the degree of
Doctor of Philosophy, August, 1962.
156 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
INTRODUCTION
Since their discovery in 1875, the Lower Permian redbeds of
north central Texas have been an important source of fossil
vertebrates. Until recent decades, however, the fauna of this
region (mainly described by Cope, Case, and Williston) was de-
rived from a very limited series of horizons, which include the
upper portions of the Wichita Group and the lowest part of the
overlying Clear Fork. Collections made in the uppermost beds
of the Clear Fork by Olson (1948, 1951 a-c, 1955) and in the
Lower "Wichita formations by Romer and associates (Romer,
1935, 1947, 1958) have extended our knowledge and given us
some concept of the faunal assemblages that preceded and fol-
lowed those of the typical Lower Permian faunas. This paper is
concerned with certain amphibian remains recovered from the
lower levels of the Wichita.
A number of expeditions from the Museum of Comparative
Zoology at Harvard College have been made to the Texas red-
beds from 1934 onward. In most cases these had the special ob-
jective of obtaining remains from the lower horizons of the
Wichita Group — the Putnam, Moran and Pueblo formations in
descending order. The finds have included numerous identifiable
amphibian specimens (mainly from the Putnam and Moran)
which, for the most part, pertain to genera already well known
from the higher, "classic," Wichita formations, the Admiral and
the Belle Plains. These include such forms as T rimer orhachis,
Eryops, Broiliellus, Tersomius, Parioxys and Acheloma. How-
ever, the lower formations showed indication of a somewhat dif-
ferent and more primitive faunal assemblage, particularly in the
occurrence of the primitive rhachitome, Edops (Romer and
Witter, 1942).
A further indication of the faunal difference between the
upper, typical, Wichita and the lower Wichita formations was
the recovery in the earlier trips to these lower beds of remains of
an obviously new rhachitome too fragmentary to permit ade-
quate description. Vertebral and limb materials showed some
similarity to T rimer orhachis, a form not identified with cer-
tainty below the Putnam (Olson, 1955), but such skull remains
as were recovered indicated a narrow, attenuated snout quite
different from that of Trimerorhachis.
A collecting trip from the Museum of Comparative Zoology in
1954 under the direction of Dr. Alfred S. Romer resulted in the
CHASE : NELDASAURUS WRIGHTAE 157
recovery of a nearly complete skull of this problematical form,
with attached lower jaws and some postcranial material in a
common block. The certain association of the skull with post-
cranial materials made possible for the first time a comprehen-
sive description of this new form and the assignment of the
specimens in previous collections to a new genus.
The fossil material described in this paper is assigned to a new
genus and species of rhachitomous labyrinthodont for which I
propose the name Neldasaurus wrightae. Both generic and speci-
fic designations are in honor of Miss Nelda E. Wright, Research
Assistant and Editor of Publications of the Museum of Com-
parative Zoology, who discovered the holotype specimen.
ACKNOWLEDGEMENTS
My appreciation is due to several persons whose aid I re-
received during the course of this study. I should like first to
express my sincere thanks to Dr. Alfred S. Romer, under whose
inspiring direction this work was undertaken and whose con-
tinued interest and advice have been of invaluable assistance. I
am indebted to Dr. Donald Baird of Princeton for many helpful
suggestions and permission to use his information on the Linton
fauna ; to Dr. Nicholas Hotton III of the United States National
Museum for information on Acroplous vorax; to Dr. Edwin H.
Colbert of the American Museum of Natural History for access
to specimens of Trinierorhachis ; to Dr. Craig C. Black of the
Carnegie Museum for the loan of specimens ; to Dr. Robert
Carroll then at the Museum of Comparative Zoology for informa-
tion pertaining to early edopsoids ; to Dr. Carl Cans of the Uni-
versity of Buffalo and Mr. Frank White of the Biological Labo-
ratories at Harvard for assistance in taking photographs of the
specimens. Thanks are also due to Mr. Arnold Lewis of the
Museum of Comparative Zoology at Harvard, who gave freely
of his time and knowledge in introducing me to the techniques
of fossil preparation. Thanks are due also to Dr. Ernest E.
Williams and Professor Bryan Patterson of the Museum of
Comparative Zoology for critical reading of the manuscript. I
am indebted to Miss Nelda Wright for many valuable sugges-
tions during the preparation of this paper.
A year of this study was made possible by a Danforth Teacher
Study Grant, for which I am very grateful. Preparation of the
manuscript was aided by a grant from the Shell Assist Fund.
158 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
SYSTEMATIC DESCRIPTION^
Class AMPHIBIA
Order TEMNOSPONDYLI
Suborder EHACHITOMI
Superfamily TRIMERORHACHOIDEA
Family TRIMERORHACHIDAE
Neldasaukus, gen. n.
Type species. Neldasaurus wrightae sp.n.
Generic Diagnosis. — A low-skulled labyrinthodont amphibian
which closely resembles T rimer orhachis in size, many skull pro-
portions, body and limbs, dermal roof pattern, and palate.
Neldasaurus differs from T rimer orhachis in the following re-
spects : orbits midway between snout and occiput ; snout nar-
row ; lacrimal elongate but not reaching the external naris ;
nasals, prefrontals and frontals elongate ; external nares close
together, their long axes more or less parallel ; jugal entering
the orbital border; prevomers elongate and the choanae broadly
separated from the anterior border of the interpterygoid vacui-
ties ; larger number of teeth than in T rimer orhachis in the lower
jaw and the marginal series of the upper jaw; a tusk pair on
the ectopterygoid ; a foramen in the lower jaw for the re-
ception of vomerine tusks ; opisthotic and prootic unossified ;
pleurocentra as high as the intercentra; clavicles meeting in
front of the interclavicle.
Neldasaurus wrightae sp. n.
Holotype. — MCZ 2200, including a nearly complete skull and
attached lower jaws, central and neural arch vertebral elements,
a partial pectoral girdle, limb bones and remnants of dermal
armor.
Type Locality and Horizon. — Terrapin School, Section A-
1266, BBB + CRR Survey, Archer County, Texas ; Moran For-
mation, "Wichita Group, Permian. Collected by A. S. Romer
party, 1954.
Diagnosis. — As for genus.
Referred Specimens. — UCZ Nos. 1371, 1381, 1438, 1463,
2404, 2406, 2407, 2516, 2518, 2519.
1 The classification used here follows Homer (1947).
CHASE : NELDASAURUS WRIGHTAE 159
DESCRIPTIVE LIST OF MATERIALS
The material consists of the holotype and the following speci-
mens, all from the Wichita Group of the Texas Lower Permian.
MCZ 1371. A partial skull and left lower jaw, dorsal verte-
brae — some in articulated series, ribs, a partial clavicle and
iuterclavicle, a nearly complete forelimb, dermal scales and
armor. There is also what appears to be a pelycosaur foot bone.
Putnam Formation, Table Branch, South Fork of the Little
Wichita River, G. AV. Stell Survey, Section A-382, 10 miles west
of Anarene in Archer County, Collected by R. V. Witter party,
1936.
MCZ 1381. Fragmentary remains of several small individuals.
Same data as for MCZ 1371.
MCZ 1438. Poorly preserved remains of skulls and postcra-
nial materials of at least two individuals of little use for pur-
poses of description. Putnam Formation, 1 mile west of Archer
City in the Archer City bone bed. Section 151, American Tribune
New Colony Subdivision, southwest part Archer County. Col-
lected by R. V. Witter party, 1936.
MCZ 1463. Fragmentary cranial and postcranial materials in
a limey nodule. Section 16, Falls County School Land, Archer
County, about 7 miles southwest of Anarene. Collected by R. V.
Witter party, 1936. The formation is either Putnam or Moran.
MCZ 2404. Fragmentary portions of lower jaw and postcra-
nial materials. Moran formation at Terrapin School, BBB -f
CRR Survey, A-1266 in southern Archer County. Collected by
A. S. Romer party, 1950.
MCZ 2406. A large number of discrete skull and postcranial
materials, mostly fragmentary, including the proximal ends of
a humerus and a femur. There is also a Trimerorliachis-like
intercentrum, a complete ring central element and scraps of a
larger labyrinthodont. Moran Formation near Padgett in the
northeast part of William Tryndale Survey, Young County.
Collected by A. S. Romer, 1942.
MCZ 2407. Fragmentary vertebral materials, the distal ends
of a radius and a femur and portions of a pterygoid. Collection
site as for MCZ 2406. Collected by R. V. Witter, 1942.
MCZ 2516. Separate and disarticulated elements, mostly in-
complete ; including parts of a lower jaw, a right femur, an
ilium, and vertebrae. Moran Formation, %^ mile north and 1
mile west of Padgett Schoolhouse, Young County. Collected by
R. V. Witter party, 1936.
160 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
MCZ 2518. Fragmentary skull and jaw materials, numerous
partial vertebral elements, ribs, pectoral plates and two ilia,
probably all pertaining to one individual. Moran Formation in
Padgett, the northwest part of the William Tryndale Survey,
MCZ Field No. 9, Young County. Collected by A. S. Eomer,
1951.
MCZ 2519. A partial intercentrum. Moran Formation west
of Cottonwood Creek, about Section 48, County School Land,
Archer County. Collected by A. S. Romer, 1954.
ANATOMICAL DESCRIPTION OF GENUS AND SPECIES
General Skull Morphology
Of the three principal skull specimens, the holotype, MCZ
2200 (Pis. 1 and 2), is the most complete. The figures and de-
scription are primarily based on this specimen, but supplemen-
tary information was obtained from MCZ 1371 (PI. 3), MCZ
1438 (PL 4), and also the fragments included in MCZ 2518.
The holotype skull is described first.
The dermal roof, although fairly complete, is displaced to the
left in relation to the palate and has been subjected to post-
mortem crushing so that the skull appears to have retained its
original depth only in the right suborbital region. Horizontal
pressures have caused some overlapping of the medial boundaries
of the central dermal bones.
Intrusion of the articular portions of the lower jaws from be-
low partially obscures the relationships of the quadrate region.
The anterior extremity of the snout is incomplete in all of the
specimens, but the portions that remain permit a reasonable res-
toration. The right lower jaw of the holotype has retained a
fairly normal position throughout much of its length so that
most of the lateral skull margin is visible. However, the posterior
quarter of the ventrolateral border of the skull roof has been
damaged and details of structure are hard to trace.
Post-mortem flattening of the skull and damage to the skull
margins introduce some uncertainty in the determination of
cranial dimensions. However, the figures obtained from the re-
constructed skull roof, the palate and the lower jaw, give the
general dimensions, as set forth in Table 1, with a fair degree of
accuracy. Determination of the depth of the skull was based
primarily on the transverse dimensions of the palate and the
resultant restoration of the skull roof to fit these dimensions.
CHASE : NELDASAURUS WBIGHTAE 161
The skull (Figs. 1, 2, 3 and 5) is of moderate size. The snout
is narrow but bluntly rounded anteriorly, the skull increasing
in breadth toward the back, having its greatest width in the oc-
cipital region. Allowing for a slight backward projection of the
condyle from the occipital border, the suspensorium is about 1.5
cm behind the level of the condyle. However, the otic notch is
not as highly developed as in Trimerorhachis, and the contour of
the central posterior rim of the skull table is less concave than
it is in that animal.
The occiput slopes downward and backward at an angle of
approximately 23° from the vertical in the preserved skull.
But, though some degree of slope may be normal, the occiput
may have been more nearly vertical in life.
The depth of the skull midway between the orbits and the
end of the snout is approximately 13 mm, and the height of the
cheek below the orbit is 14 mm. The height of the occiput is
about 39 mm. Thus, the depth of the skull increases from front
to back, the increase being considerably more pronounced in the
postorbital region, the snout being generally flattened.
Determination of the original skull roof topography is difficult
because of post-mortem distortion. The evidence seems to indi-
cate a more or less flat or slightly concave surface in the central
region of the skull table bounded by a low ridge on each side
which runs from the tabular to the outer posterior corner of the
orbit. The outer face of the ridge slopes gently downward, be-
coming confluent with the cheek, and the inner face dips down
to the central region of the skull table.
The mid-orbit to snout length is approximately 49.8 per cent
of the midline skull length. The dorsally directed orbits are
oval in shape and somewhat smaller in relation to overall skull
size than those of Trimerorhachis'^ , though, as in Trimerorhachis,
the interorbital space is narrow. The inner border of the orbit
is slightly elevated above the surface of the skull roof, the
smooth, medial margin having a depth of 3 mm. The rest of the
margin, which shows no appreciable elevation, is not as thick.
A small parietal foramen is present in the midline between the
parietals, 2.9 cm behind the posterior orbital border. The dis-
tance from the orbit to the parietal foramen is equal to 38 per
cent of the skull table length, a shorter distance than that seen in
Trimerorhachis, where the comparable figure is 50 per cent.
1 This comparison is based on the ratio of skull width X skull length/orbit
width X orbit length.
162
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
In agreement with the dimensions of the rostrum, the external
nares are not as widely separated as they are in other tri-
merorhachoids. The medial margin of the naris is slightly ele-
vated above the surface of the internarial region, and the opening
is somewhat constricted in its posterior third. The lateral border
of the naris is separated from the skull margin by a dorsal ex-
posure of the premaxilla and the maxilla of approximately
5.5 mm.
Skull MCZ 1371 (PI. 3) exhibits overall dimensions that are
closely comparable to those of the holotype, and the dimensions
given in Table 1 can be accepted with a fair degree of certainty.
Extreme flattening makes the determination of skull height dif-
ficult, but it appears to agree closely with that of MCZ 2200.
The preserved portions of skull MCZ 1438 (PI. 4) appear to
represent a somewhat larger animal than the preceding forms.
Overall length cannot be determined, but the dimensions of the
preorbital region suggest proportions similar to those of MCZ
2200 and 1371.
The dermal roof bones of MCZ 2518 are disarticulated and
incomplete so that comparison of general dimensions of this
specimen and the others is impossible.
TABLE I
Measurements
(in mm) of Dermal Skull Eoof
MCZ
MCZ MCZ
No.
No. No.
Length
2200
1371 1438
Snout to
end of skull
table
158
— —
Snout to
quadrate
173
174 —
Snout to
mid-orbit
78
80 86
Orbit
20
20 (est.) —
External
naris
13
— 14
Width
Snout behind naris
40
42 46
Orbital region
70
74 —
Quadrate
level
120
— —
Orbit
16
15 (est.) —
External
naris
8
— 8
Internarial
10
— 12
K eight
Mid-rostral
13
— 18 (est.)
Suborbital
14
14 (est.) 15 (est.)
Occiput
39
— —
chase : neldasaurus wrightae 163
Bones of The Dermal Skull Roof
(Figures 1 and 2)
Some sutures on one or another of the skull specimens, MCZ
2200, 1371, 1438, 2418, were readily traced, but the location
of others, which were not recognizable per se, was based on
the sculpture patterns of individual bones. Centers of ossifica-
tion were determined by considering their relationship to the
sensory canal system, which is typically associated with the cen-
ter of growth in each bone (Bystrow, 1935; Westoll, 1943; Par-
rington, 1949). I believe that the general pattern of the skull
roof of MCZ 2200, as figured, is reasonably accurate.
The dermal roof pattern closely resembles that of Trimerorha-
chis. Differences in the shape of individual bones in the two
animals are obviously due in large measure to the differences in
skull proportions mentioned above.
The dermal bones of the skull roof are treated here as com-
prising four groups: (1) the paired elements of the dorsal mid-
line, (2) the circumorbital bones, (3) the bones of the temporal
region that form the lateral margin of the skull table, and (4)
the marginal tooth-bearing bones and the cheek bones.
(1) The nasals are long, roughly rectangular bones which ex-
tend more than half the distance from the naris to the
orbit, their shape sharply contrasted to the squarish nasals in
T rimer orhachis.
The elongate frontals, with over half their length extending
into the preorbital region are also in contrast to conditions in
T rimer orhachis, in which form the nasals make a much greater
contribution to the midfacial area than do the frontals.
The parietals form most of the central portion of the skull
table, extending from the posterior border of the frontals to the
postparietal suture. A parietal opening of modest size is located
centrally between them.
The roughly pentagonal postparietals are relatively small
bones. Posteriorly, they turn down over the rim of the skull
table in an occipital exposure. A stout occipital flange of the
postparietal, which rests on a dorsal extension of the exoccipital,
bounds the supraoccipital region laterally and forms the medial
rim of the posttemporal fenestra (Fig. 5). The tabular does not
appear to take part in the formation of the descending flange.
(2) The circumorbital series, including the prefrontal, post-
frontal, postorbital, jugal, and lacrimal, is complete.
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BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PMX
Fig. 1. Dorsal view of skull roof of Neldasauriis wrightae, n. gen., n.
sp., restored, X -75. Portions of the sensory canal grooves and some sutures
on the right have been restored to agree with conditions on the left.
Abbreviations: EO, exoccipital; F, frontal; IT, intertemporal; J, jugal;
L, lacrimal; MX, maxilla; N, nasal; P, parietal; PF, postfrontal; PMX,
premaxilla; PO, postorbital; PP, postparietal; PEF, prefrontal; QJ, quad-
ratojugal; SQ, squamosal; ST, supratemporal; T, tabular.
CHASE : NELDASAURUS WRIGHT AE 165
The prefrontal is a narrow bone which forms the anteromedial
margin of the orbit, ending halfway back on the medial rim.
The postfrontal is narrow anteriorly where it joins the pre-
frontal to complete the medial rim of the orbit. Behind the
orbit it becomes somewhat wider.
The jugal is a large bone approximately 7 cm long, and the
longest skull roof bone, aside from the maxilla. Unfortunately,
the suborbital region in all of the specimens is poorly preserved
or damaged, but a break in the surface contour of the right
cheek below the orbit in MCZ 2200 and the pattern of the sculp-
ture in this area suggest that there was a lacrimal-jugal suture
about halfway back on the lateral rim of the orbit. If this inter-
pretation is correct, the jugal forms at least a small part of the
orbital border.
The postorbital forms most of the posterior orbital rim, meet-
ing the jugal laterally on the outer margin of the orbit, though
the exact position of the suture is not known.
The lacrimal is elongate. It forms much of the anterolateral
rim of the orbit and, though it fails to reach the naris, it extends
four-fifths of the distance from the orbit to the naris in the
holotype skull.
(3) Primitively, the lateral margin of the skull table includes
an intertemporal bone as well as supratemporal and tabular.
The posterior limits of the intertemporal, a relatively small bone
here, were difficult to trace but the anterior and medial bounda-
ries were well defined on MCZ 2200 and MCZ 2518.
The tabular is the smallest bone in the skull table. Laterally
it has a short contact with the squamosal, excluding the supra-
temporal from the margin of the otic notch, a feature which
tends to accompany modest development of the latter (Homer,
1947, p. 24). Medially, where it meets the postparietal, the
tabular forms most of the dorsal margin of the posttemporal
fossa. Midway in its posterior border the tabular has a ventral
projection, near the inner edge of the otic notch, which is the
posterior end of a low ridge that continues anteriorly under the
skull roof for a distance of 7 mm. The depth of this ridge, or
flange, is approximately 3 mm.i
Immediately anterior to its posterior border the ventral sur-
face of the tabular is excavated so that a depression is formed.
1 Case (193.5) mentions a "low riisoso line" rniinine: forward and Inward on the
under surface of the tabular from the inner edge of the otic notch in his Trimero-
rhachis specimen 16002, which may be comparable to the structure described here.
166
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
bounded medially by the "flange"
described above and posteriorly by
the posterior rim of the tabular. Lat-
erally this fossa, which is about 9 mm
wide, is bounded by a ventrally-
directed process from the posterome-
dial edge of the squamosal. Little of
the underside of the skull roof of this
region can be seen and its precise
character, partially visible only in the
holotype, is difficult to interpret be-
cause of its imperfect condition.
(4) The boundaries of the marginal
tooth-bearing bones, the premaxilla
and maxilla, were hard to find in the
rostral region. Their outlines as fi-
nally determined are a composite of
the patterns that could be seen on
parts of four specimens, MCZ 2200,
1371, 1438, and 2518. The premax-
illa, seen best in MCZ 1438, appar-
ently had a small dorsal exposure an-
teriorly, separating the anterior rim
of the external naris from the rostral
border by a distance of approximately
3 mm. The union of the premaxilla
with the nasal posteriorly appears to
have been close to the anterior narial
rim on the medial side. Medially, the
premaxilla joins its fellow of the op-
posite side, and posteriorly, lateral to
the naris, it joins the maxilla near the
middle of the lateral rim of the naris.
Fig. 2. Skull of Neldasaurus wrightae, in lateral aspect. Eestored, X -75.
Abbreviation: Q, quadrate. Other abbreviations as in Figure 1.
CHASE: NELDASAURUS WRIGIITAE 367
The maxilla extends along the lateral margin of the skull for
about two-thirds of the skull length, its dorsal exposure dwin-
dling rapidly towards the rear. Posteriorly, it comes close to the
quadratojugal. Conditions here are uncertain, but the jugal
appears to enter the skull margin for a short distance between
the maxilla and quadratojugal.
The postorbital cheek region is formed in typical fashion by
the squamosal and quadratojugal. Posteriorly the squamosal has
a contact with the anterior portion of the quadrate and forms
part of the outer margin of the otic notch. On the inner posterior
margin of the cheek it has a ventral and anteriorly directed proc-
ess, about 6 mm deep, which joins the quadrate ramus of the
pterygoid on its lateral edge.
The quadratojugal forms the posterolateral margin of the
skull below the squamosal. Conditions in the left quadrate region
of MCZ 2200, as near as can be determined, are like those in
MCZ 1371, where it is better preserved. The posterior edge of
the quadratojugal, forming a shallow, concave rim, is separated
from the lateral margin by a sharp angle. The inner posterior
end of the bone forms a medial projection posterior to the
sutural surface for the squamosal. The posterior rim of this
medial process probably formed part of the border of the
quadrate foramen. The sutural face on the quadratojugal for
articulation with the squamosal, therefore, does not reach the
anterior edge of the foramen, and the squamosal does not reach
its border. The remainder of the border of the quadrate foramen
was apparently formed by the quadrate. Conditions here ap-
pear to be comparable to those seen in T rimer orhachis (Case,
1935, fig. 4; Williston, 1915, fig. 3).
The pattern of the dermal roof bones of MCZ 1371 (PL 3) ap-
pears to be identical to that of the holotype. This specimen
offers further evidence of the relations of the lacrimal, which is
here prevented from reaching the posterior narial border by a
union of the maxilla and the nasal. Also, although the situation
is not entirely clear, the lacrimal appears to have a stout union
with the jugal below the orbit so that the jugal forms more than
half of the lateral orbital border.
The skull roof of MCZ 1438 (PI. 4) is incomplete, but it ap-
pears to follow the general pattern seen in MCZ 2200 and 1371.
In contrast to its form in the other skulls, the lacrimal here
reaches to, or nearly to, the external naris. The only other
apparent difference is in the slightly larger dimensions of the
168 BULLETIN : MUSEUM OF COMPARATn^E ZOOLOGY
individual bones, which agree with the general porportions of
the skull.
As stated earlier, the skull roof of MCZ 2518 consists of a
number of isolated fragments. In so far as it can be traced, the
dermal roof pattern is like that of the holotype.
The sculpture of the dermal roof bones consists of ridges and
pits, or grooves, radiating from a center of ossification in each
bone. Notable differences between the sculpture pattern in
Neldasaurus and T rimer orhachis are obviously correlated with
differences in the shape of individual bones in the two animals.
As any figure shows (Bystrow, 1938, fig. 11), there is no linear
sculpture in Trimer orhachis, whereas in Neldasaurus there is a
suggestion of linear sculpture on adjacent regions of the frontals
and nasals and on the frontals and parietals. The sculpture also
tends to become linear in portions of other long bones in Nelda-
saurus such as the lacrimal and jugal.
The Sensory Canal System
The channels of a sensory canal system are well defined on the
surface of the dermal roof bones in Neldasaurus (Figs. 1 and
2). In all the specimens the pattern of these channels is re-
markably constant, and, except for the anterior rostral portions
of the supraorbital and infraorbital channels, the pattern is
clear.
The pattern of the sensory canal system generally resembles
that of Trimer orhachis (Case, 1935, fig. 5 A) but differs in some
particulars. In Neldasaurus the supraorbital groove crosses the
postfrontal, prefrontal, and nasal but does not enter the lacrimal
as in Trimer orhachis. The anterior portion of the infraorbital
groove in Neldasaurus runs from the jugal along the suture be-
tween the maxilla and lacrimal, then continues for some distance
on the lacrimal. The anterior portion of the infraorbital groove
in Trimerorhachis is confined to the maxilla except for a short
loop onto the lacrimal.
The position of the temporal groove in Neldasaurus is esssen-
tially as in Trimerorhachis, though it is slightly more medial in
Neldasaurus. A jugal sulcus of the sensory canal system,
separated by a short gap from the infraorbital groove on the
jugal, crosses the jugal-squamosal suture near its midpoint and
has the shape of a medially convex curve on the central region
of the squamosal, swinging laterally near the posterior margin
CHASE : NELDASAUEUS WRIGHTAE 169
to end at the posterolateral corner of the bone. It does not ap-
pear to have passed across the quadratojugal, but over the pos-
terior border of the squamosal, just behind the end of the qua-
dratojugal, to its connection with the lower jaw system. This
groove is unknown in T rimer orhachis.
The courses traced by the sensory canal grooves of MCZ 1371,
1438 and 2518, where they can be seen, are the same as those on
the holotype.
Palate and Palatoquadrate
(Figure 3 and Plate 2)
In all of the skull specimens the lower jaws lay in more or
less normal closed position against the ventral surface of the
skull, thus obscuring the lateral palatal surface. Fortunately,
it was possible to expose this region by removing the right lower
jaw ramus and 2.5 cm of the anterior end of the left jaw ramus
from the ventral surface of the holotype skull. The description
and reconstruction of the palate and the ventral skull surface, as
shown in Figure 3, were mainly based on this specimen. Restored
portions are hatched in the figure.
The general form and proportions of the central and posterior
portions of the palate in Neldasaurus, including the presence of
large interpterygoid vacuities, resemble conditions in Trimero-
rhachis. However, the region of the palate in front of the inter-
pterygoid vacuities in Neldasaurus, reflecting the proportions of
the elongate snout, is more extensive.
The anterior end of the palate is incomplete, but the evidence
suggests that the premaxilla had a moderate palatal exposure.
The ventral exposure of the maxilla, throughout most of its
length, represents little more than the width of the tooth row.
The maxilla accounts for nearly two-thirds of the lateral mar-
gin ; its posterior end briefly enters the anterior portion of the
lateral rim of the subtemporal fossa. The jugal apparently
enters for a short distance into the ventral margin of the skull
posterior to the maxilla, but most of the lateral border of the
fossa is formed by the quadratojugal.
Determination of the extent of the palatal exposure of the
maxilla adjacent to the internal naris was difficult, and some un-
certainty exists. Since the palatine appears to form most if not
all of the lateral border of the choana, the maxilla could enter
170
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 3. Reconstruction of skull of Neldasaurus wrightae in palatal as-
pect, X -75. Eestored areas are hatched. Abbreviations: APF, anterior
palatal fenestra; BO, basioccipital; CH, choana; EC, ectopterygoid; EO,
exoccipital; J, jugal; MX, maxilla; PAL, palatine; PMX, premaxilla; PS,
parasphenoid ; PT, pterygoid; Q, quadrate; QJ, quadratojugal ; SE, sphen-
ethmoid ; VO, vomer.
CHASE : NELDASAURUS WRIGHTAE 171
its border (if at all) only at the outer anterior corner. Exclu-
sion of the maxilla from the lateral rim of the choana by a union
of the vomer and the palatine, which appears to be the case
here, is to my knowledge reported in only one other form,
Eohrachyops townendi (Watson, 1956). The suture between the
maxilla and premaxilla can not be seen in ventral aspect. Never-
theless, on the basis of its location on the lateral skull margin, it
probably lies close to the level of the anterior member of the
vomerine tusk pair.
The marginal teeth are small, crowded and numerous. By
counting the teeth in areas where they can be seen, it was pos-
sible to estimate a total number of 93 maxillary teeth. The an-
terior teeth are less than 1 mm in diameter, and in the anterior
portion of the maxilla there are slightly less than 10 teeth per
cm, while towards the posterior end, where the teeth are still
smaller, there are approximately 11 per cm. The length of the
tooth-bearing margin of the premaxilla, as reconstructed, is
approximately 2.5 cm. The narrow palatal exposure of this bone
suggests small tooth size in this region also and, allowing for
some decrease in the number of teeth per unit area, the pre-
maxilla should have held at least 15 teeth. In some regions of
the maxilla it was noted that teeth alternated with replacement
sockets, though no consistent pattern could be seen.
Although the anterior portion of the palate in MCZ 2200 is
damaged, there is evidence of the presence of two sets of open-
ings, the anterior palatine fenestrae and the choanae. Immedi-
ately anterior, and slightly medial, to the anterior vomerine tusk
on each side there is an opening in the palate. In the matrix
deposited in this region, on both sides, there are remnants of the
upper ends of symphysial tusks belonging to the lower jaw. It
appears that anterior palatal fenestrae were present on the
suture between the premaxillae and the vomers in Neldasaurus.
The matrix-filled space containing the upper end of the tusk is
confluent with a gap in the anterior part of the floor of the ex-
ternal naris in MCZ 2200. The condition of this region in MCZ
1371 and 1438 is not clear but appears to be similar in the former
at least. A comparable situation exists in Eupelor {Buettneria) .
The choanae are located halfway between the anterior end of
the interpterygoid vacuities and the end of the snout. They are
suboval in outline, with a length of approximately 1.5 cm and a
greatest width of .6 cm. Anteriorly, they are constricted by a
lateral curvature of the medial border. The lateral margin of
172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the choana is formed in the main, and perhaps entirely, b}^ the
palatine, the anterior and medial margins by the vomer. Al-
though the palate was crushed against the ventral surface of the
skull roof in this region of the type specimen, it is apparent that
the vomer formed a stout rim on the medial and anterior borders
of the opening. The anterior ends of the choanae are 1 cm behind
the level of the posterior rims of the external nares. Therefore,
to reach the internal naris from the outside, air had to pass
backward through a short passageway between the skull roof
and the palate — a passageway surely enclosed in a cartilaginous
nasal capsule.
The central palatal surface anterior to the interpterygoid
vacuities is formed by the vomers. These bones, conforming to
the contours of the snout, are approximately four times as long
as they are wide. Anteriorly, the vomer is bounded by the pre-
maxilla, and forms the posterior rim of the anterior palatal
fenestra ; laterally, it joins the maxilla and the palatine, and
forms the medial and posterior boundaries of the choana. The
vomer meets the palatine midway on the posterior rim of the
choana. The suture between the two bones appears to run diago-
nally backward and outward, medial to the palatine tooth-row.
Posteriorly, the vomer forms the anterior margin of the inter-
pterygoid vacuity, but the two vomers are separated in the mid-
line, posteriorly, by the wedge-shaped anterior end of the cultri-
form process of the parasphenoid. At the point where the vomers
meet medially the end of the cultriform process appears to pass
dorsal to them.
The posterolateral sutural relations of the vomers are obscured
by numerous cracks and the poor preservation of the palatal
surface. However, it appears that the vomer formed the antero-
lateral border of the interpterygoid vacuity, retaining a primi-
tive contact with the pterygoid medially near the anterior end of
that opening.
The palatine is bounded laterally by the maxilla and medially
by the vomer. Posteriorly, the palatine meets the ectopterygoid.
This suture was hard to find but appears to run diagonally for-
ward across the palatal toothrow immediately in front of an en-
larged tooth pair and 2.5 cm behind a pair of palatine tusks. If
this is the correct position of the suture, the ectopterygoid bears
a pair of tusks at its anterior end. It is of interest to note that in
T rimer orhachis, as described by Case (1915), the ectopterygoid
is a short bone, and the palatine supports not only an anterior
CHASE : NELDASAURUS WRIGHTAE 173
fang pair but a second pair of enlarged teeth as well. No en-
larged tusks occur on the ectopterygoid in that form. A review
of the literature failed to uncover any different description of
the palatine-ectopterygoid suture in Trimerorhachis. Thus, if the
present interpretation is correct, Neldasaurus is more primitive
than TrimerorhacJiis in this respect.
Assuming that the position of the suture is correct, the
ectopterygoid is a long bone, about equal in length to the pala-
tine. ]\Iedially, it is bounded by the palatal ramus of the ptery-
goid and laterally by the maxilla. Posteriorly, where it enters
the anterior border of the subtemporal fossa, it is very narrow.
The palatine and ectopterygoid are thickened under the pala-
tal tooth row, and join the maxilla in a firm articulation on the
medial surface of that bone. The palatine is stouter than the
ectopterygoid. The union of these lateral palatal bones with the
maxilla does not reach the ventral edge of the latter with the
result that, in ventral aspect, there is a shallow longitudinal
groove along the lateral edge of the palate just inside the skull
margin.
In primitive fashion, the vomer, the palatine and, apparently,
the ectopterygoid each bears a pair of tusks. Those of the palatine
are the largest and those of the ectopterygoid are the smallest.
It was not possible to determine whether or not any small teeth
were present on the vomers between the choanae. However, both
the palatine and ectopterygoid tusks are accompanied by a
single row of small teeth.
On the left side of the palate in the type specimen the vo-
merine tusk pair is represented by an empty socket with a tusk
behind it. On the right the conditions are reversed. The vo-
merine tusk is .5 cm in diameter near the base and is some-
what over 1 cm long.
The right palatine bone also supports a tusk pair at its an-
terior end, the posterior tusk represented by an empty socket.
The palatine tusk preserved is larger than that of the vomer,
with a diameter at the base of .6 cm and an approximate length
of 1.2 cm. Behind the tusk pair on the palatine is a close-set
row of 8 small teeth. The more anterior ones are oval in out-
line, their long axes perpendicular to the tooth row. The pos-
terior ones are round in section. Larger than the maxillary
teeth, the palatine teeth average 2 mm in diameter. The third
pair of tusks is located at the anterior end of the ectopterygoid.
The anterior member of the pair has a basal diameter of 3 mm,
174
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
and the posterior a diameter of 4 mm. Behind these, extending
nearly to the end of the ectopterygoid, is a row of 21 teeth, de-
creasing slightly in size towards the posterior end of the row.
Their diameter at the base is about 1 mm.
The pterygoid of the type specimen, though damaged, is
nearly complete. There are also portions of pterygoid bones in
MCZ 2406 and 2407.
The pterygoid consists of a horizontal palatal ramus, a stout
central body and a vertical quadrate ramus. The curved, me-
dial margin of the palatal ramus forms the lateral and postero-
lateral borders of the interpterygoid vacuity. The bone in gen-
eral is fairly stout but is thin at its medial edge. The palatal
ramus has a narrow anterior process, bounded laterally by the
ectopterygoid, which articulates with the vomer anteriorly. The
posterior portion of the palatal ramus is expanded horizontally.
The outer edge of the pterygoid is here thickened and turned
ventrally to form a flange about 1 mm deep which forms the
anterior portion of the medial border of the subtemporal fossa;
the remainder of the border is supplied by the central body and
the quadrate ramus of the pterygoid.
The basal articulation was apparently movable (Fig. 4, A).
The thickened, horizontal body of the pterygoid opposite the
..j^yOy-
B
Fig. 4. Neldasaurus wrightae. A, reconstruction of the basal articular
region of the skull of MCZ 2200 in ventral view, X -75. (Evidence for the
presence of the teeth shown on the right pterygoid based on MCZ 2406 and
2407). B, projection of a portion of the medial surface of the left palato-
quadrate of MCZ 2200, X -75. Abbreviations: BO, basioccipital ; CE, conical
recess; EO, exoecipital; ET, "excavatio tympanica"; PS, parasphenoid ;
PT, pterygoid; RC, pocket for rectus capitus muscle ?
CHASE : NELDASAURUS WRIGHTAE 175
basal articular region of the basisphenoid has an internal process
that presumably articulated with the basipterygoid process of
the braincase. The internal process, seen best on the right in
MCZ 2200, curves dorsally from the level of the central body
of the pterygoid so that its tip is nearly .5 cm above the lower
surface of the bone. An ascending process rises from the medial
edge of the pterygoid just behind the tip of the internal process.
Its height above the internal process is about .5 cm, but it rises
rapidly posterior to this point to a height of 1.7 — 1.8 cm opposite
the basipterygoid process of the braincase. The concave pos-
terior face of the internal process of the pterygoid forms the
anterior and ventral walls of a socket or "conical recess" for
the reception of the basipterygoid process of the braincase.
The back wall of the "conical recess" is formed by an essen-
tially vertical ridge that rises from the dorsal surface of the
pterygoid, just inside the medial edge of the bone, about 1 cm
behind the tip of the internal process. This leaves a very limited
socket for the reception of the basipterygoid process. Behind the
ridge there is a vertical groove, about 5 mm wide, which separates
the ridge from the anterior end of the vertical quadrate ramus
(Fig. 4, B). The latter, rising above a small medial projection
of the pterygoid, forms a second ridge posterior to the groove.
The posterior face of the ridge thus produced at the root of the
quadrate ramus, and the medial face of the quadrate ramus ad-
jacent to it, share a concavity just above the ventral surface of
the pterygoid. It is obvious that this deeply excavated area rep-
resents the "excavatio tympanica" of Bystrow and Efremov
(1940). It appears that Neldasaurus is unusual in that the pos-
terior wall of the "conical recess" is separated by a groove from
the ascending ridge at the base of the quadrate ramus which
forms the anterior wall of the tympanic excavation.
The quadrate ramus of the pterygoid is approximately 4 cm
long and curves outward slightly as it proceeds from its root to
an attachment with the quadrate at the outer posterior corner
of the skull. Just behind the root, the quadrate ramus, as noted,
becomes more or less vertical in the skull, apparently ascending
dorsally and somewhat medially. Near the level of, or just an-
terior to, the otic notch the vertical flange of the quadrate ramus
overlaps on the medial side a vertically descending flange of the
squamosal. The posteromedial face of the quadrate ramus ap-
pears to have had an irregular surface, marked by longitudinally
oriented ridges and grooves. There is a prominent, more or
176 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
less horizontal, elongate groove in the quadrate ramus, just be-
low its connection with the descending flange of the squamosal.
Although post-mortem flattening of this region has introduced
some uncertainty in dorso ventral relationships, it seems highly
probable that this groove is homologous with the groove in the
posteromedial face of the quadrate ramus of T rimer orhachis,
which apparently serves as a floor for the tympanic cavity
(Watson, 1956).
Little can be seen of the quadrate bone. On the medial side of
the left cheek in the holotype, however, an anterior extension of
the quadrate, which diminishes in size anteriorly, meets the
squamosal dorsally and the quadrate ramus of the pterygoid
ventromedially.
There is no evidence of an ossified epipterygoid in Nelda-
saurus. A review of the Trimerorhachis materials in the collec-
tion of the Museum of Comparative Zoology confirmed earlier
reports that there is no ossified epipterj-goid in that form either.
The obstinate character of the matrix left no indication of
palatal teeth in areas other than those described in MCZ 2200,
but the fragments of the pterygoid bones in MCZ 2406 and
2407 show that the horizontal flange and part of the palatal
ramus are covered by a densely-packed shagreen of small teeth
except at the medial edge of the bone.
Parasphenoid and Braincase
The braincase is fairly complete in ventral aspect in the holo-
type specimen. The dorsal surface is crushed against the under-
side of the skull roof and little can be seen of the lateral surface.
The only other braincase material consists of a badly weathered
partial parasphenoid with MCZ 1438 and a small portion of
the left side of an isolated parasphenoid in MCZ 2518. The de-
scription of the parasphenoid and the braincase is based on the
holotype.
The proportions of the braincase are remarkably similar to
those of TrimerorJiachis (Case, 1935; Watson, 1956). The sphe-
nethmoid region is narrow, but behind its articulation with the
pterygoids the braincase is expanded and more or less flattened.
As is usual, the ventral surface of the braincase is sheathed by
the parasphenoid.
The parasphenoid narrows gradually in front of the basi-
pterj^goid process of the braincase, giving rise to an anteriorly
CHASE : NELDASAURUS WRIGHTAE 177
directed cultriform process, which forms the medial border of
each interpterygoid vacuity. The cultriform process is about 6.6
cm long, fairly broad at each end, but narrow in the central
region. The anterior end of the process is wider than in Tri-
merorhachis, but it does not project as far forward into the
anterior palatal surface as it does in such forms as Sanrerpeton
or the Triassic metoposaurs.
The anterior and posterior portions of the ventral surface of
the cultriform process are nearly flat; the central portion is
ventrally convex in cross section, the lateral edge being approxi-
mately 2 mm above the level of the ventral surface. The surface
of the cultriform process is marked by fine longitudinal ridges
and grooves, which also appear to some extent on the main body
of the parasphenoid. There is no evidence of teeth on the ventral
surface of the parasphenoid, though the rigorous preparation
required to remove the matrix could have obscured them.
The main body of the parasphenoid is broad and essentially
flat in ventral view (Fig. 4, A). Behind the root of the cultri-
form process the parasphenoid expands laterally, the central
region between the laterally expanded ''wings" being moder-
ately concave. Laterally, the parasphenoid sheaths the anterior,
ventral and posterolateral faces of the basipterygoid process.
The "core" of the process, ossified as part of the basisphenoid
in many labyrinthodonts, was here apparently cartilaginous.
It would have been exposed at the tip of the process above the
parasphenoid, where it could have entered the "conical recess"
of the pterygoid to meet the cartilaginous epipterygoid. The
anterior margin of the parasphenoid covering the basipterygoid
process is directed diagonally outward and backward in direct
apposition to the posterior margin of the internal process of
the pterygoid. As in Trimcrorhachis, but in no other Permian
temnospondyl, the articular process of the parasphenoid is set
off from the main body of the bone bj^ a groove which runs di-
agonally outw^ard and backward, widening and increasing in
depth distally. The anteroventral rim of the articular process of
the parasphenoid issues from the side of the braincase on the
same level as the ventral surface of the main body of the bone.
Behind the basipterygoid processes, the parasphenoid expands
laterally to a width of approximately 5 cm. The margin of the
bone then curves sharply inward, matching posteriorly the width
of the basioccipital which is 2.2 cm across the base. The pos-
terior border of the braincase has been damaged, but it appears
178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
that the parasphenoid reaches, or nearly reaches, the posterior
ventral rim of the basioccipital, although it is extremely thin at
this point.
A lateral extension of the parasphenoid turns upward on
either side around the basioccipital, gaining a narrow exposure
on the lateral face of the braincase at the level of the condyle.
Anteriorly this lateral exposure increases, attaining a height of
nearly 5 mm ventral to the fenestra ovalis, the ventral rim of
which was apparently formed by the parasphenoid. The dorsal
and lateral margins are incomplete. In T rimer orliachis the
lateral concave face of the parasphenoid behind the basiptery-
goid process contains a deep pocket ventral to the fenestra
ovalis, which Watson suggests could have received a rectus
capitus muscle. Although this region in Nelclasaurus is not com-
plete, the details that can be seen suggest that a similar, though
less pronounced, fossa for muscle insertion was present.
Careful scrutiny failed to disclose the presence of any open-
ings for the palatine or internal carotid arteries in the ventral
or lateral surfaces of the parasphenoid. However, these openings
are often small and are not consistently demonstrable even in a
well known form like Trimerorhachis. Failure to find them in
Neldasaurus can probably be attributed to their small size or to
the rigorous treatment necessarily employed in removing the
matrix from the surface of the bone.
Some features of the dorsal surface of the parasphenoid can
be determined from exposed portions of the lateral edges, and
from the face of a break which runs diagonally across the main
body of the braincase. The outer end of a broad groove in the
dorsal surface of the parasphenoid can be seen on the lateral
surface of the braincase above the laterally expanded "wing."
The groove is 6 mm wide at its lateral end and is directed an-
teromedially. It appears to be comparable to a similar groove
on the upper surface of the parasphenoid in Trimerorhachis
and, as Watson (1956) suggests, probably sheathed the lower
medial end of the prootic.
Immediately anterior to the basipterygoid process, the lateral
edge of the parasphenoid is thin, but both faces of a break in
this region exhibit a thickened mass of bone. Although the form
of this thickened mass is partially obscured by matrix, it appears
to be part of a strongly developed transverse ridge of bone on
the upper surface of the parasphenoid between the roots of the
basipterygoid processes. The ridge could occupy the position to
CHASE : NELDASAURUS WRIGHTAE
179
be expected of the posterior rim of a depression for the pituitary.
Reduction of ossification in the braincase is well advanced.
The basisphenoid and otic capsule were apparently cartilaginous,
since neither is preserved in the holotype. There is, however,
evidence of an ossified sphenethmoid. A flat bone about 1 cm
high appears to occupy the space between the cultriform process
and the skull roof from the anterior end of the interpterygoid
vacuity back to a point not far in front of the level of the parie-
tal foramen, a distance of about 4 cm. This bony sheet, bounding
laterally the space above the cultriform process, undoubtedly
represents an ossified sphenethmoid of the right side. There is no
evidence that there was any lateral expansion of the bone like
that seen in Edops and Eryops. Unfortunately, the stapes is not
preserved in any specimen.
Occiput
(Figure 5)
The occipital region of the holotype, though damaged, is
nearly complete and can be restored. An isolated partial condyle
among the materials associated with MCZ 1371 also offers some
information, but the description is based mainly on the holotype.
The occipital region of MCZ 2200 has been subjected to post-
mortem flattening, resulting in some dislocation of its parts. The
dorsal portions of the exoccipitals are missing, and only the left
side of the condyle is reasonably intact.
The incomplete nature of the exoccipitals hindered an attempt
to get an accurate picture of the proportions of the occiput.
Fig. 5. Neldasaurus wrightae (MCZ 2200 and MCZ 1371). Eeconstruc-
tion of the skull in occipital aspect, X -75 approx. Abbreviations : BO, basi-
oeeipital; FM, foramen magnum; POP, paraoccipital process; PS, para-
sphenoid; PT, pterygoid; PTF, posttemporal fossa. Other abbreviations
as in Figure 1. Eestored features are in dotted lines.
180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
However, on the basis of the general similarity of the structure
and proportions of the occipital region of Neldasaurus and
T rimer orhachis, the height of the dorsal portions of the exocci-
pitals of Neldasaurus was estimated by comparison with Wat-
son's figures for T rimer oj'hachis. Accordingly, the height of the
occiput of Neldasaurus was approximately 3.9 cm. As noted
earlier, the backward slope of the occiput was probably not
pronounced in life.
The occipital condyle is a single, subcircular structure with a
concave posterior face. Its lower division is formed by the basi-
oecipital, best seen in MCZ 1371. The basioccipital is wedge-
shaped in lateral view and extends forward ventrally into the
floor of the brainease for a distance of approximately 1.5 cm.
The dorsal surface of this anterior extension is unfinished. As
said earlier, the parasphenoid covers the ventral surface of the
basioccipital to, or nearly to, its posterior border. As in Tri-
merorhachis, the ventral surface of the basioccipital has longi-
tudinal ridges which were in contact with the upper surface of
the parasphenoid.
The exoccipitals, resting directly on the basioccipital, form
the dorsolateral portions of the condjde and complete its concave
posterior face. The position of the suture between the basiocci-
pital and exoccipital can not be determined with certainty but
appears to be about half-way between the base and the top of
the condyle. Dorsally, the condylar portions of the exoccipitals
are well separated, but they approach each other in the midline
ventrall}". Above the center of the condyle, which contains a
notochordal pit, the exoccipitals are only separated by a narrow
groove. Above the condyle the exoccipital had a dorsal process,
represented in the specimens by its basal portion only, which
formed the lateral wall of the foramen magnum and articulated
dorsally with the inferior surface of the descending flange of the
postparietal. Anteriorly, a projection of the basal portion of
the exoccipital of each side extends forward in the floor of the
brainease above the anterior extension of the basioccipital, but
details are obscure. Probably, as in T rimer orhachis, it provided
a floor for the medullary region of the brainease. These an-
terior extensions of the exoccipitals do not quite meet in the
midline. The structure of this region, in so far as it can be
determined, corresponds generally to that described for Tri-
mer orhachis by Watson.
CHASE : NELDASAURUS WRIGIITAE 181
Anterior and somewhat lateral to the base of the dorsal exten-
sion of the exoccipital, there is a small process of the exoccipital
which forms the anterior wall of the vagal foramen. The pos-
terior wall of the foramen is formed by the base of the ascending
process of the bone. Just posterior to the vagal foramen the
lateral wall of the exoccipital is pierced by a singe foramen for
nerve XII. The inner opening of this foramen is in the medial
surface of the bone, anterior and slightly ventral to its lateral
exit, as seen in MCZ 1371.
The ventral surface of the condyle is broad and more or less
flat. The width of the condyle in MCZ 2200 is 2.2. em and its
height is 1.7 cm, while the isolated condyle of MCZ 1371, with
a width of 2.3 cm and a height of 1.3 cm, is even lower. These
proportions seem to reflect a general trend towards flattening in
Neldasaurus. The lateral sides of the condyle rise abruptly
from the more or less flattened base. In lateral aspect they are
considerably narrower than the sides of the condyle in Trimero-
rhachis. Taken together, these characteristics of the condyle
resemble the proportions of the intercentra of Neldasaurus, in
which slender, upright lateral processes arise from the edges of
a broad ventral base.
The occipital exposure of the postparietal has a slight back-
ward slope, which is interpreted as being normal. There is no
supraoccipital ossification. The ventral articular surface of the
postparietal flange slants laterally and ventrally at an angle of
45 degrees to the horizontal, ending laterally in a blunt point.
The medial portion of this flange presumably articulated with
the upper end of the exoccipital; the laterally projecting outer
corner may have rested on the base of a cartilaginous paroccipi-
tal bar. There is some evidence that a cartilaginous paroccipital
bar extended between the exoccipital and the tabular. The upper
end of the bar could have been received in the fossa in the ven-
tral surface of the tabular already described. The lower end of
the bar presumably met the anterior surface of the exoccipital
ventromedially. There is no indication that the tabular or the
exoccipital invaded the paroccipital bar, the latter thus forming
the whole of the lower boundary of the posttemporal fossa. The
remainder of the fossa, as in T rimer orkacliis, is formed by the
postparietal and the tabular.
182 bulletin : museum of comparative zoology
Lower Jaw
(Figure 6)
All of the lower jaw specimens have suffered some post-mortem
damage and distortion, especially in the articular region. The
most complete jaws, MCZ 2200, 1371 and the anterior por-
tions of 1438, are crushed against the ventral surface of the skull
so that much of their dorsal and medial surfaces is obscured.
Fragments of lower jaws are also present in MCZ 2516, 1381,
1463, 2406, 2404, and 2518.
The right jaw of the holotype specimen is well preserved
throughout its prearticular extent, the articular region being
hidden. The left jaw is nearly complete from front to back in
ventral aspect, the articular region, though twisted, retaining
its connection with the prearticular ramus. The infradentary
portions of both jaws have been flattened against the palate, los-
ing their original relation to the vertical plane of the dentary.
The left lower jaw of MCZ 1371 (Fig. 6, A) has undergone
some distortion and damage, but the preservation of the surface
permits a more accurate tracing of the suture pattern than either
jaw in MCZ 2200. The anterior tip is missing, but the contours
of the preserved portion permit an accurate restoration. Also,
though it is incomplete, the articular region is more readily vis-
ible in this specimen.
The reconstruction of the lower jaw shown in Figure 6, B is
a composite, based primarily on MCZ 2200 and 1371. The rela-
tion of the articular region to the ventral rim of the jaw is
based on this portion of a small jaw in MCZ 1381, but this
relationship is, admittedly, subject to some uncertainty in the
larger specimens where the articular region has shared in the
general flattening of the ramus.
Aside from a few details, the lower jaw of Neldasaurus closely
resembles the lower jaw of TrimerorJiachis. Viewed from below,
the anterior end of the jaw is bluntly rounded, the end of each
ramus swinging abruptly towards the symphysis from its lateral
border, thus paralleling the blunt contours of the snout. The
jaw is very shallow anteriorly, its strongly curved outer surface
directed as much ventrally as it is laterally. Posteriorly, it be-
comes progressively deeper, and the convexity of the lateral
surface, though continued, is less pronounced. The ventral rim
of the jaw was apparently medial to the plane of the dorsal rim
CHASE : NELDASAURUS WRIGHTAE
183
in life. The medial surface, forming a sharp angle with the lat-
eral surface ventrally, is essentially vertical. Posteriorly, the
ventral margin of the jaw curves sharply upward to the articu-
lar region. There is a modest retroarticular process which ex-
tends for a distance of 6 mm behind the posterior rim of the
glenoid fossa on the medial side of the jaw. Two cm behind the
anterior end of the jaw the dorsal edge of the dentary rises to a
point 3 mm above its anterior level; 3 cm behind this point a
second, though less abrupt, increase in height occurs. From
here, the dorsal margin of the jaw rises gradually to the articu-
lar region.
CHT
SANG
AN 6
SANG
S MAND
SACC
Fig. 6. Lower jaw of Neldasaurus wrightae X -50. A, ventral view of
left ramus of MCZ 1371. B, external view of left ramus restored (based
mostly on MCZ 2200 and MCZ 1371). Abbreviations: AMF, anterior Meckelian
fenestra; ANG, angular; CHT, foramen for chorda tympani; D, dentary;
PA, preartieular ; PMF, posterior Meckelian fenestra; SACC, accessory
sulcus; SANG, surangular; SD, dentary sulcus; SMAND, mandibular sul-
cus; SP, splenial; SPP, posterior splenial; VT, fenestra for reception of
vomerine tusk.
The bones of the outer surface of the jaw are sculptured. The
medial surface and the ventral rim are smooth.
Two narrow, elongate Meckelian fossae are present just above
the ventral margin on the inner surface of the jaw. There is a
foramen for the chorda tympani just under the edge of the
glenoid fossa at the root of the retroarticular process on the
medial surface of the jaw.
184 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
In the discussion of the palatal dentition, a fenestra for the
reception of a vomerine tusk in the lower jaw was mentioned.
In both lower jaws of MCZ 2200 and 1438 and the single jaw
of 1371 there is a hole that penetrates the jaw from top to bot-
tom to accommodate the tip of a vomerine tusk. That this is
not the result of post-mortem flattening of the skull is shown
by the absence of breakage of the surface bones and the finished
rim of the opening. The opening of the fenestra in the ventral
surface of the jaw is suboval in outline, the long dimension be-
ing directed anteromedially. It is approximately 4 mm wide
and 5 mm long, and is located on the suture between the dentary
and the splenial at, or slightly behind, the level of the posterior
limit of the symphysis. In MCZ 2200 a second fenestra is present
in the anterior end of the postsplenial for the accommodation
of a palatine tusk. This second fenestra, however, is not evident
in MCZ 1371 or 1438.
Colosteus scutellatus and Ei'petosauriis rodiafus (Romer,
1930) have a groove in the outer surface of the lower jaw for
the reception of a large premaxillary fang. Some Crocodylidae
have developed fossae which penetrate the upper surface of the
anterior end of the snout for the lower jaw fangs and, of course,
anterior palatal openings are present in a large number of tem-
nospondylous labyrinthodonts. But perforation of the lower
jaw for the accommodation of an upper jaw tusk appears to
be unique in Neldasaurus.
The absence of the posterior fenestra in two of the skulls may
reflect individual variation in the depth of the jaw in relation
to the length of the tusks, or perhaps in older animals large
tusks may gradually wear a smooth opening through the lower
jaw.
The symphysis is formed by the dentary anteriorly, and the
splenial (presplenial) forms the posterior half. Ventrally, it is
approximately 1.5 cm long; dorsally it is 2.2 cm long. A medial
expansion of the splenial near its anterior end meets a similar
expansion from the opposite side, thus lengthening the extent
of the symphysis dorsally. The posterior border of the medial
process so formed lies 1 cm above the ventral rim of the jaw.
A somewhat similar condition appears to obtain in Trimero-
rhachis, though developed to a lesser degree.
The dentary occupies most of the upper portion of the lateral
surface of the jaw ramus and, as already shown, forms most of
CHASE : NELDASAURUS WRIGHTAE 185
the anterior end of the jaw. Except for some uncertainty con-
cerning the posterior extent of the dentary, its relation to the
other dermal elements is similar to that of other rhachitomes.
It is bounded ventrally by the splenial, postsplenial and angu-
lar from front to back, and overlaps each of these elements to a
variable extent. The dentary narrows abruptly at the back, its
ventral edge ascending diagonally upward and backward. Al-
though its relation to the surangular is not clear, an anterior
process of the surangular appears to be interjected between the
upturned posterior end of the dentary and the dorsal border of
the angular. Much of the dentary is sculptured, but the dorsal
portion along the tooth row is quite smooth except for the pres-
ence of fine, longitudinal striations.
The splenial, as stated, forms the posterior portion of the sym-
physis, its anterior end meeting the dentary in a serrate suture.
On the outer surface of the jaw it has a long diagonal external
suture with the postsplenial, which slants downward and back-
ward, and internally reaches the anterior Meckelian fenestra,
forming the anterior rim of that opening.
The postsplenial extends posteriorly to the posterior Meckelian
fenestra, forming the anterior third of the border of that open-
ing. Posteriorly, it has a ventrally directed, diagonal suture
with the angular on the lateral surface of the jaw. The post-
splenial turns over the ventral rim of the jaw, appearing at the
level of the posterior Meckelian fenestra to have a height well
over 1 cm on the medial surface.
The large angular forms the posteroventral rim and most of
the lateral surface of the jaw below and anterior to the articu-
lation. Posteriorly and dorsally the angular is bounded by the
surangular, anteriorly and dorsally by the dentary. It has a
rather long anterior extension, confined to the lateral face of
the ramus, which projects between the postsplenial and the den-
tary in a gradually narrowed process. Ventrally, a suture be-
tween the angular and the prearticular is located on, or just
medial to, the ventral rim of the jaw.
The surangular is not complete in any of the specimens.
However, on the medial surface of the articular region of the
jaw of MCZ 1371 a rather large crack slants upward and back-
ward from the end of the angular to the lip of the glenoid fossa.
In the absence of other evidence, and because of its position,
this "crack" is tentatively identified as the suture between the
surangular and the prearticular. Thus, the surangular would
186 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
form the posterior border of the glenoid fossa. Immediately
posterior to the "suture," and, as noted previously, just under
the edge of the glenoid fossa, is a foramen for the chorda tym-
pani. Its dorsal position and the fact that it is nearly, if not
completely, surrounded by the surangular contrasts to the sit-
uation in T rimer orhachis. In that form, the surangular, angular
and prearticular all contribute to the borders of the foramen,
according to Case's figures, and its position is more ventral. The
foramen for the chorda tympani in MCZ 2200 has a lower posi-
tion than that observed in MCZ 1371, suggesting that its loca-
tion is subject to individual variation.
The limits of the articular bone can not be determined and a
dorsal view of the glenoid fossa is seen only in MCZ 1371. Un-
fortunately, it has been damaged. The fossa consists of a con-
cave depression passing diagonally inward from the outer pos-
terior corner of the jaw. A slight ridge separates an outer ar-
ticular facet from a longer facet on the inner side of the fossa.
The full extent of the prearticular can not be determined, nor
can its relation with the coronoids be seen in surface view. In
typical fashion it forms the medial wall of the adductor fossa
above the angular and postsplenial. Its anterior end extends for
some distance in front of the posterior Meckelian fenestra. The
medial edge of the adductor fossa is, as typically, lower than the
lateral edge. The fossa appears to end anteriorly at the level of
the posterior rim of the posterior Meckelian fenestra, having a
length of approximately 4 cm.
Unfortunately, the coronoid series is not visible in any of the
specimens. The only source of information is that supplied by
cross sections of the right lower jaw of MCZ 2200 (Fig. 7).
A section through the anterior coronoid, Section J^, shows three
small teeth in a transverse series. Presumably these teeth rep-
resent the condition continued throughout the rest of the bone.
Traces of similar small teeth were also found in sections showing
the other coronoid elements in Neldasaurus.
Section Ji was made approximately 3.5 cm behind the tip of
the jaw and 2.3 cm in front of the anterior end of the anterior
Meckelian foramen. It should, therefore, pass through the den-
tary, the splenial and the anterior coronoid. As can be seen in
the figure, some distortion is evident, but the three elements can
be identified.
A comparison of this section with a section taken through the
anterior end of the jaw of T rimer orhachis (Broom, 1913, fig.
CHASE : NELDASAUBUS WRIGHTAE
187
9D) demonstrates a basic similarity in structure, though the jaw
of Neldasaurus is relatively more narrow.
Section J2 was made approximately 1 cm in front of the pos-
terior Meckelian foramen. This section shows very clearly a
medial extension of the dentary below the tooth row and above
the Meckelian space. The lower portion of the jaw has been
folded medially to the vertical plane of the dentary. If this
were drawn back to a reasonably normal position, the height of
the jaw at this level would be approximately 1.8 cm and the
ventral rim would lie a considerable distance medial to the lat-
eral surface of the upper border of the dentary.
PAL?
ANG SPP
J2
Fig. 7. Transverse sections of the right lower jaw and lateral palatal
border of MCZ 2200, X 1- Ji, Section 3.5 cm behind the anterior end of
the ramus; J2, Section 1 cm in front of the posterior Meckelian foramen.
Abbreviations: Ci, anterior coronoid; lAC, inferior alveolar canal; MSP,
Meckelian space; PD, palatal debris; V?, vomer. Other abbreviations as in
previous figures.
The sensory canal system of the lower jaw appears to be rep-
resented by three lateral line grooves — a mandibular, a den-
tary, and an accessory. The mandibular sulcus originates on the
posterior margin of the jaw, lateral to the retroartieular process.
Its course is readily traced in MCZ 1371 and 2200, where it pro-
ceeds along the ventrolateral edge of the jaw, just above the
ventral rim, to the posterior border of the symphysis.
The dentary sensory groove arises from the mandibular groove
below the articular region, and runs forward along the upper
margin of the angular. Anterior to the midpoint of the angu-
lar it can not be traced with certainty.
There is some evidence in MCZ 1463 and in the small jaw of
1381 of the presence of an accessory sulcus, dorsal to the dentary
sulcus, at the posterior end of the jaw. Due to poor preservation
it can not be positively identified in either 1371 or 2200.
188 BULLETIN : MUSEUM OE COMPARATIVE ZOOLOGY
By piecing together the information that can be obtained
from MCZ 2200 and 1371, it is possible to give a fairly complete
account of the lower jaw dentition. Neldasaurus possesses a pair
of prominent symphysial tusks on either ramus, each with a
diameter at the base of about 5 mm and a height of at least 1 cm.
The teeth of the dentary series in MCZ 2200 and 1371 agree
generally in size and number in comparable sections of the jaws
of the two specimens, and it appears that the lower jaw of Nel-
dasaurus contained 60 or so teeth. The teeth are more crowded
than they are in Trimerorhachis (where there are about 48
teeth in the lower jaw), a feature which reflects the high tooth
count of the maxillary series already described.
An attempt to recognize a consistent pattern of tooth replace-
ment was unsuccessful.
There is a gradual reduction in tooth size from front to back,
but with some suggestion of regional enlargement not far be-
hind the anterior end of the dentary in MCZ 2200. The teeth in
MCZ 1371, though all have lost their tips, are better preserved
than in either jaw of MCZ 2200. Again, there is regional en-
largement of the teeth somewhat behind the anterior end of the
tooth row. Near the anterior end of the dentary the teeth have
a diameter at the base of 1.75 mm. The enlarged teeth that fol-
low have a diameter of 2.5-3.0 mm. In the lower jaw^ tooth series
here described, teeth 18-22 are enlarged, 21 and 22 being the
largest. Comparison of the position of this series with the palatal
dentition shows that these teeth would be located between the
vomerine and palatine tusks, though closer to the latter. Be-
hind this series a second, though less pronounced, enlargement
occurs in teeth 33-38.
Vertebral Column
(Figures 8 and 9)
In addition to numerous isolated elements, mostly centra and
intercentra, three specimens show associated vertebral material
in more or less continuous series.
The vertebral column of MCZ 2200 is distorted and much of
the detail is obscure. However, it is possible to trace a reason-
ably continuous presacral column in which there were approxi-
mately 34 vertebrae. MCZ 2518 includes a number of blocks con-
taining presacral vertebrae and several isolated elements, in-
cluding 33 intercentra. Twenty-five vertebrae can be accounted
CHASE : NELDASAURUS WRIGHTAE
189
for in MCZ 1371. It seems probable, therefore, considering the
possibility of loss of some elements, that Neldasaurus had ap-
proximately 34 presacral vertebrae. The estimated number for
Trimcrorhachis is 31 (Williston, 1915; Case, 1935).
Unfortunately, no atlas vertebra was found and there are only
three elements possibly belonging to the caudal series.
On the basis of the relative positions of the vertebral elements
in blocks, variations in size of the elements, and variations in
the height of the rib facets on centra and intercentra, the pre-
sacral vertebral columns of MCZ 2200, 1371 and 2518 were re-
constructed. ^ The similarity of the vertebrae of all the Nelda-
saurus specimens makes a detailed description of each specimen
unnecessary.
B
'•<irr:i
Fig. 8. Partial reconstruction of presacral vertebral column of Nelda-
saurus wrightae, X -5. A, vertebrae 2-5; B, vertebrae 11-16; C, vertebrae
19-26.
lA detailed description of the reconstruction of the vertebral column for each
specimen is given in the thesis on which this paper is based, on file at Harvard
University.
190 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
The Neldasaurus vertebrae conform closely to the typical
rhachitomous pattern. Each vertebra consists of a neurocentrum,
a single intercentrum open at the top, and paired lateral pleuro-
eentra. The latter are well developed and have a height approxi-
mately equal to that of the intercentra. Figure 9, A and B,
shows three vertebrae from the mid-dorsal region and two verte-
brae from the anterior dorsal region of MCZ 1371.
There is little regional difference in the size of the intercentra
and pleurocentra of the presacral vertebral column. Those of
the mid-dorsal region are slightly larger than the elements of
the anterior and posterior regions. The average height of the
first ten intercentra in MCZ 1371 is 1.9 cm, and the average
length of the base is 1.2 cm. In the mid-dorsal region these fig-
ures are 2.2 cm and 1.5 cm, respectively. Elements near the
posterior end of the series are intermediate in size between the
anterior and mid-dorsal elements.
The intercentrum has an expanded, somewhat thickened base,
from the lateral edges of which tapering, dorsally-directed proc-
esses curve upward around the notochordal space, giving a tall
wedge-shaped appearance in side view. The ascending process
ends in a blunt point. In none of the specimens do the inter-
centra form complete rings, though the dorsal tips of some ap-
proach each other quite closely. The ascending process is marked
off from the base, on the lateral face of the intercentrum, by a
usually prominent longitudinal ridge. Above the ridge the lat-
eral face of the ascending process is concave. In ventral aspect
the base of the intercentrum is widest posteriorly. Both anterior
and posterior edges of the base are j^rotuberant, having a medial
swelling which is usually more developed anteriorly. The an-
terior and posterior edges of the base turn down so that the
central region is modestly concave in lateral view. However, this
concavity is not uniform, and in most intercentra a low, median,
rounded ridge interrupts the depression.
There is an articular facet for the capitulum of the rib on the
posterior border of the ascending process. In some instances
this is a strong laterally directed projection, in others it is less
pronounced. The height of the facet above the base of the inter-
centrum becomes progressively greater from front to back in
the vertebral column. In some anterior intercentra the facet is
at the very base of the lateral face, while in the posterior ones
it is at the tip of the ascending process. The intercentra are
stouter and better ossified than those of T rimer or Jiachis, so that
the notochordal canal is relatively smaller than in that animal.
CHASE : NELDASAURUS WRIGHTAE
191
However, the base of the intercentrum is not as highly ossified
as it is in Eryops. The ventral face and the ascending process
are sheathed by dense periosteal bone, but on the lateral face of
the latter the finished bone does not quite reach the edge or the
tip of the process. The ventral and lateral faces of some inter-
centra are marked to varying degrees by small pits, while others
have a smooth surface. The occurrence and development of pits
is highly variable in intercentra of comparable size, even from
the same individual. Prominent longitudinal ridges are not a
common feature of the outer surface of the base and ascending
process of the intercentra of Neldasaurus as they are in Trimero-
rhachis.
B D
Fig. 9. Vertebral elements of Neldasaurus wrightae, MCZ 1371. A, mid-
dorsal vertebrae, left lateral aspect. B, anterior dorsal vertebrae, left
lateral aspect. C and D, ventral and anterior view of an intercentrum.
E, lateral and anterior view of a left pleurocentrum. All X 1-
192 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Three partial intercentra, possibly referable to the caudal
series, are present among the specimens. Two are associated with
MCZ 2200 ; the other is an isolated element associated with MCZ
2518. The base is not flattened but strongly curved. Dense
perichondral bone is confined to the ventral surface and one
edge of the ascending process, the top of which is missing. The
intercentrum base is thickened dorsally by endochondral bone
which constricts the notochordal space. There is no evidence of
haemal arches. In the absence of other information these ele-
ments are presumably intercentra of proximal caudals.
The pleurocentra, like the intercentra, are very uniform in
structure. They are prominent elements, and, as noted, have a
height approximately equal to that of the intercentra. They have
a blunt, oblong, dorsal head, about twice as long as it is wide.
The ventral end is pointed. Their shape is essentially that of a
slender wedge with the tip down, and they are unusual in reach-
ing nearly to the bottom of the intercentra. In these features
they are in sharp contrast to the small diamond-shaped pleuro-
centra of T rimer orhachis or the blocky wedges of Eryops. The
lateral face of the pleurocentrum is covered by a layer of dense
bone that curves outward along the edges so that the surface is
concave. There are modest transverse ridges within the vertical
groove so formed, and small pits which, like those of the inter-
centra, are variable as to number and arrangement. The finished
bone does not reach the anterior or dorsal border of the pleuro-
centrum. In some, the anterior, lateral edge of the bone is re-
flected to produce a small facet, which presumably shared with
the intercentrum in supporting the capitulum of the rib. In
anterior aspect the pleurocentra are strongly curved to fit the
notochordal canal.
The neural arches are low and the presence of a narrow strip
of unfinished bone surface between the lateral halves of the arch
suggests that the arches were formed as separate lateral ossifica-
tions. There are well developed anterior and posterior zygapo-
physes, but there is no discrete transverse process. The pedicle
of the arch is modestly developed and is more truncate than it is
in such a form as Eryops. The tuberculum of the rib presumably
articulated with the ventrolateral area of the base of the arch
pedicle.
The articular faces of the zygapophyses are roughened, and
that of the anterior zygapophysis slopes inward for the reception
of the posteriorly and ventrally directed face of the posterior
CHASE : NELDASAURUS WRIGHTAE 193
zygapophysis of the preceding arch. The neural spines are low
and only moderately expanded from front to back.
The neural arch elements show greater regional variation than
the central elements. Thus, the neural arches of the "cervical"
region (Fig. 9, B) have low neural spines, the fourth neural
arch from the front as preserved in MCZ 1371 being 2 cm high,
the arch pedicle accounting for 50 per cent of the total height.
The pedicle descends for a short distance below the zygapophysis,
and is curved sharply outward in a strong lateral projection • —
an incompletely ossified transverse process. The neural spines of
this region have a slight backward tilt. The neural elements of
the mid-dorsal region are taller, the additional height resulting
from elongation of the neural spine (Fig. 9, A). The height of
neural element 12 is 2.4 cm; the height of the arch above the
pedicle is 1.8 cm, 75 per cent of the total height. The pedicle
is truncated and the anterior zygapophyses are only slightly
above its base. The neural arch of this region does not have a
laterally directed pedicle as does that of the anterior region.
The posterior zygapophyses are well separated from the anterior
processes on each arch as a result of the strong backward slant
of the mid-dorsal spines, which have a sharper backward tilt
than those of the anterior region.
Ribs
(Figure 10)
There are many ribs preserved in the specimens but most are
fragmentary and only a few, confined to MCZ 1371, have re-
tained their original association with the vertebrae.
With one possible exception, the ribs preserved appear to be-
long to the dorsal series. A fragment from MCZ 2200 seems to
be the distal portion of a rib with a "finished" end, which may
belong to the "cervical" region. The proximal end is missing.
The portion of the shaft preserved is 2.2 cm long, slightly curved
and oval in cross section. The shaft is uniformly slender, with a
diameter of only 0.2 cm, in contrast to the heavier dorsal ribs
described below.
There is some regional variation in the dorsal ribs and also
variation between specimens, but a common pattern can be
traced. The shafts of the dorsal ribs are only slightly curved, so
that the distal ends of the ribs of opposite sides are well sepa-
rated ventrally. A rough calculation of the distance spanned by
194
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
^:r3
Fig. 10. iiibs of Neldasaurus wrightae. A and B, partial anterior ribs.
C, partial rib from the mid-dorsal region. D, partial rib from the posterior
presacral region. Other ribs as numbered. All from MCZ 1371. All X -75,
approx.
opposing ribs articulating with the vertebral column gives a fig-
ure of 11.5 cm. If articulated, the ribs would appear to run
essentially straight out, suggesting a flat back (and belly) in
Neldasaurus. As the longest ribs preserved are incomplete dis-
tally, and since no provision was made for a possible cartila-
ginous cap on the rib head, this figure is reasonable when com-
pared to the width of the pectoral girdle, which is 13.6 cm.
Though most of the ribs taper distally, a few appear to have had
a modest expansion of the distal portion of the shaft. Proximally
the shafts are more or less triangular in section ; distally they are
CHASE : NELDASAURUS WRIGHTAE 195
elliptical, the long axis of the ellipse oriented dorsoventrally.
The rib heads are expanded, though single, and some distinction
can be made between confluent capitular and tubercular articular
areas.
As noted, there is variation between specimens, so the rib
series of MCZ 2200 and MCZ 1371 are described separately. The
ten largest ribs preserved in MCZ 2200 probably belong to the
anterior region of the dorsal series, and are all of comparable
size. Of these ribs, the longest as preserved is 4.5 cm and is repre-
sentative (Fig. 10, B). The head, which was presumably con-
tinued in cartilage, is broad, measuring 1.0 cm in the long axis.
Proximally the shaft is stout, having a width of .6 cm on the
"capitular" (medial) side and tapering distally to a diameter
of .4 cm at the end as preserved. There is no clear division of
capitulum and tuberculum on the anterior face of the rib head
but the posterior face contains a shallow V-shaped depression
in the central proximal region separating a rounded capitular
area from a narrow tubercular area. The lateral edge of the
shaft bears a low, narrow keel. Behind the keel, along the pos-
terodorsal surface of the rib, a groove runs the length of the
shaft.
There are parts of 12 slightly smaller ribs, mostly fragments
of the shaft region, presumed to belong to the mid-dorsal series.
The longest and most complete of these is shown in Figure 10, C.
The rib is incomplete distally but is 4.2 cm long. It is less mas-
sive than the anterior ribs; the shaft is more slender and the
groove along the posterodorsal border of the shaft is less pro-
nounced. Unlike the anterior ribs, the shaft is straight. There
is almost no distinction between capitular and tubercular areas,
though the head is flattened.
Posterior dorsal ribs are represented by a single incomplete
rib and fragments of two others. The former consists of the head
and 2.5 cm of the proximal portion of the shaft (Fig. 10, D).
The shaft is slender, the lateral margin forming a thin keel. It
does not appear to be grooved. The medial surface is not visible.
The head, which is .7 cm long, is thickened and is expanded
medially in the central proximal region, so that it forms a nearly
oval articular area in end view, and is more sharply marked off
from the shaft than the heads of the anterior ribs. There is no
separation of capitular and tubercular regions.
The ribs of MCZ 1371 (Fig. 10, E) are similar to those of
MCZ 2200, but none are as massive as the anterior dorsal ribs of
196 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the latter. Regional differentiation is less marked in this speci-
men. The longest rib as preserved, associated with the 5th verte-
bra of the anterior portion of the vertebral column, is 4.4 cm
long. The shaft tapers distally and is slightly curved. The head
is flattened, being .8 cm wide, but there is little differentiation
between capitular and tubercular areas. The depression noted
in the rib heads of MCZ 2200 is lacking. There is a keel along
the lateral edge in the proximal half of the rib shaft.
The ribs associated with the mid-dorsal vertebrae in this speci-
men differ from the anterior ribs in that the heads, though broad,
are less flattened. There is some indication of a longitudinal
groove like that seen in the ribs of the holotype specimen. The
ribs associated with the more posterior series of vertebrae here
resemble the mid-dorsal ribs. They are all incomplete distally,
the longest being 3.0 cm long.
Thus, the ribs of Neldasaurus show some decrease in size from
front to back. The heads of the anterior and mid-dorsal ribs are
more or less flattened, and the posterior ribs have an expanded
head, oval in end view. In some (MCZ 2200), distinction between
capitular and tubercular areas is strongly pronounced in the
anterior ribs, modest or lacking in the mid-dorsal ribs. A lateral
keel appears to be a common feature of the rib shaft. The rib
shafts show little curvature.
Appendicular Skeleton
Shoulder Girdle. Parts of the dermal pectoral girdle can be
seen in three specimens. MCZ 2200 shows the ventral surface
of the clavicles and part of the interclavicle on the underside of
the block behind the skull (Plate 2). The anterior end of the
right clavicle was broken off but remained in the matrix. The
clavicles are pushed together along the mid-line, so that there is
little ventral exposure of the interclavicle. Part of the lateral
edge of the left clavicle can be seen in dorsal view on the upper
side of the block. MCZ 1371 includes the posterior half of a
clavicle and a small portion of the interclavicle. MCZ 2518 in-
cludes a number of sculptured fragments of the clavicular girdle.
The dermal pectoral girdle of Neldasaurus has been restored
in ventral aspect in Figure 11. The clavicles are based mainly
on MCZ 2200. The posterior portion of the interclavicle is drawn
from MCZ 1371. Although the latter animal was somewhat
larger than the holotype, the structure and proportions of the
girdle are apparently identical.
CHASE : NELDASAURUS WRIGHTAE
197
The ventral dermal girdle of Ncldasaurus forms a broad thora-
cic shield whose anterior end appears to have extended forward
under the braincase to the level of the posterior end of the basi-
pterygoid process of the paraspheniod.
B
Fig. 11. Pectoral girdle of Neldasaurus wrightae, MCZ 2200. A, left
scapulocoracoid In medial aspect, X -75. B, restoration of the dermal pec-
toral girdle, X -375.
The greatest width of the dermal clavicular girdle nearly
equals the width of the skull. The clavicle has a roughly tri-
angular outline. Its medial border is irregular but essentially
straight; the posterior border is more or less transverse but
slants diagonally forward and outward in normal position. The
width of the clavicle across the base is 4.4 cm; its length is
approximately 8.8 cm. A similar ratio obtains in MCZ 1371.
There is a modest medial projection of the posteromedial corner
of the bone. The lateral border curves inward anteriorly, but
this curvature is not as pronounced as it is in such trimerorhachids
as T rimer orliachis or Acroplous (Hotton, 1959). The outline in
general closely resembles that of Buettneria (Case, 1922).
The lateral edge of the clavicle is curved upward throughout
most of its length. About 3 cm anterior to the posterolateral
corner of the bone, the upturned lateral edge thickens and rises,
forming a stout ridge, the base of the scapular process, which
increases in height posteriorly. The dorsal tip of this process is
missing.
Although the interclavicle is obscured ventrally in MCZ 2200,
its dorsal surface in that specimen and the portion preserved in
MCZ 1371 show that it is a sizable element with a length some-
what greater than that of the clavicles. Its posterior end is
198 BULLETIN : MUSEUM OF COMPABATWE ZOOLOGY
bluntly truncated and extends only a short distance beyond the
posterior borders of the clavicles. Anteriorly its ventral exposure
is reduced by the converging medial edges of the clavicles, whose
anterior ends seem to meet in front of it. An unsculptured, but
striated flange of bone projects from the anterolateral edge of
the interclavicle to provide a surface for the reception of the
ventrally overlapping edge of the clavicle.
The ventral surfaces of the clavicles and the interclavicle have a
sculpture consisting of prominent ridges and grooves radiating
from centers in the respective bones. The center of the sculpture
in the clavicle is near the posterolateral corner, that in the inter-
clavicle is in the midline about a quarter of the distance forward
from the posterior end. The sculpture is reticulate near the
center of ossification, but rapidly becomes linear as it radiates
from the center, particularly in the clavicle. The linear aspect
of the sculpture is strongly developed, and anastomoses between
adjacent ridges are less evident than in the thoracic girdle of
Trinierorhachis. No cleithrum has been found.
The only trace of the endochondral shoulder girdle is a dam-
aged left scapulocoracoid in MCZ 2200 (Fig. 11, A). The an-
terior margin is incomplete and a large part of the central region
is missing. The general shape of the remaining portion is re-
markably similar to the scapulocoracoid of Trimerorliachis
(Williston, 1915, fig. 5, B, C, D ; Case, 1935, figs. 15 and 16),
and the missing portions have been restored by reference to
that form as well as by continuation of the contours suggested
by the broken edges of the bone.
The scapulocoracoid of Neldasaurus consists of an expanded
blade above the articular surface of the glenoid region. In
lateral view the anterior border as restored has a convex outline
and the outline of the posterior border is strongly concave dor-
sally, less so ventrally. The bone as preserved is 3 cm high and
its greatest anteroposterior extent was presumably about 2.5 cm.
Thus, correlated with the flat body of Neldasaurus, it is propor-
tionately shorter than the scapulocoracoid of a form like Eryops.
The dorsal and ventral edges are unfinished and were presum-
ably continued by cartilage. The thickness of the edges of the
bone as preserved suggests that the anterior edge was thinner
than the dorsal and posterior margins.
The posterior margin is thickened and rounded, the lower
two-thirds being confluent with a modest vertical ridge on the
medial side of the bone above the articular surface of the glenoid
CHASE : NELDASAURUS WRIGHTAE
199
region. Anterior to this ridge the posterior portion of the scapu-
lar blade is pierced by an oval supraglenoid foramen about one-
third of the way above the base.
Although the general form of the scapulocoracoid of Nelda-
saurus is strikingly similar to that of T rimer orhachis there are
marked differences in proportion. Where the scapulocoracoid of
T rimer orhachis is short and stocky, in Neldasaurus it is lightly
constructed, taller and thinner ; the articular area of the base is
only half as large as that area in Trimerorhachis. The vertical
ridge above the articular surface in Trimerorhachis is separated
from the thickened posterior margin of the bone, whereas in
Neldasaurus it is confluent with the posterior margin.
Anterior Limb (Figure 12). A partial left forelimb and
manus are preserved in MCZ 1371 (PL 5). The manus as pre-
served has four metacarpals, with two phalanges articulated
with the third and one with the fourth. The carpus was not
preserved and was presumably unossified.
There is an incomplete, disarticulated right forelimb in MCZ
2200.
Fig. 12. Neldasaurus wrightae. Left anterior limb restored from MCZ
2200 and MCZ 1371, X -75. Eestored areas are hatched.
200 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Partial humeri associated with MCZ 2516, 2518, and 2406
show some surfaces not visible in the specimens described above.
The reconstruction of the left forelimb of Neldasaurus is based
mainly on MCZ 1371 with the proximal end of the ulna added
from MCZ 2200. As can be seen from Plate 5, some re-orientation
is necessary to put the elements in good articulation.
The forelimb is small for an animal the size of Neldasaurus.
The humerus, which is relatively long, is of the type seen in
T rimer orhachis. It is about 4.5 cm long and shows some develop-
ment of a shaft region. The width of the distal end is 2.3 cm ;
the proximal end is only 1.2 cm wide. The ends are "twisted"
so that the planes of their surfaces form an angle of 27 degrees.
There is a well developed deltoid crest on the radial edge of the
humerus, just above the middle of the bone, from which an
unfinished area along the radial border reaches nearly to the
proximal articular surface. There is a small process .5 cm below
the top of the humerus on the lateral face as seen in MCZ 2518.
In the normally oriented limb its position is near the posterior
edge and it probably represents the processus latissmus dorsi.
The deltoid crest and this process are connected by a curved
band of short, radial striations, presumably supplying part of
the deltopectoral insertion. The entepicondyle is strongly de-
veloped. The supinator process is continuous with a ridge which
projects beyond the ectepicondyle on the distal radial margin
of the bone.
The radius is a short, stout bone, about 2.2 cm long, and
expanded at both ends. In end view the proximal end is nearly
circular, the distal end more like a flattened oval. The bone is
not evenly rounded, and what appears to be the medial surface
is rather sharply separated from a lateral-anterior surface by a
pronounced longitudinal ridge. The side of the shaft opposed to
the ulna is concave.
The ulna is slightly longer than the radius, as would be ex-
pected, though the exact length cannot be determined. The shaft
is curved and very narrow. The proximal end is strongly ex-
panded. The distal end is only slightly expanded. A ridge,
similar to that seen in the ulna of Triynerorhachis, occurs on
the inner posterior border of the proximal end of the bone.
The articular ends of the bones described above are unfinished
and excavated, indicating that they were originally capped by
cartilage.
The manus was preserved in lateral aspect. The limb bones
on the slab are seen from the medial aspect and must be turned
CHASE : NELDASAURUS WRIGHTAE 201
over to agree with the position of the manus. This move correctly
places the radius above the first metacarpal.
There are four digits in the manus. The phalangeal count as
restored is 2.3.3.2. The metacarpals are relatively long, slender
bones and have a flattened-oval outline in cross section. The two
central metacarpals are the longest. The first metacarpal, on the
radial side of the manus, is the shortest, and is only a little more
than half as long as the second and third. The fourth metacarpal
is smaller than the middle two but larger than the first. The
phalanges as preserved are slender bones, with the articular
swelling at the distal end less pronounced than that at the
proximal end. In both metacarpals and phalanges, as is fairly
common, most distal articular surfaces are convex, whereas the
proximal articulations are flattened.
The pectoral limb of Neldasaiirus is strikingly similar to that
of T rimer orhacliis. It would be difficult, if not impossible, to dis-
tinguish between individual bones of the two forms. As can be
seen in Case's reconstruction (Case, 1935, fig. 26, p. 267), the
restored manus of Neldasaurus matches that of Trimerorhachis.'^
Pelvic Girdle (Figure 13). MCZ 2518 contains the only pelvic
girdle material. There are two ilia with this specimen. The
left ilium is seen from the medial side. The lower part of the
right ilium is separated from the blade, which is in the same
block, and can be seen both medially and laterally. There are
no remains of pubis or ischium.
The height of the ilium is approximately 3.3 cm; the expanded
base is 2 cm across. The iliac blade, slender but expanded dis-
tally, is directed dorsally. The tip of the blade is 1.2 cm long,
but its width in the middle of the shaft is only .5 cm. The slender
proportions of the ilium recall the slight construction of the
scapulocoracoid. The general form is similar to that of Trimero-
rhachis, but the ilium is relatively taller and more slender than
in that animal. The lateral surface of the base is somewhat con-
cave. The central portion of the base is strongly produced medi-
ally and somewhat A^entrally. The upper portion of the acetabu-
lum is located above the edge of the base in the center of the
lateral concave face. The unfinished articular surface is bounded
by an upraised ridge dorsally. The ventral surface of the ilium
shows an unfinished, excavated area that was formerly continued
in cartilage. There is no indication of an attachment area for a
1 Case has reversed the position of the radius and ulna in his figures.
202
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
sacral rib on the inner side of tlie iliac blade. The top of the iliac
blade is unfinished and was presumably continued in cartilage.
Posterior Limh (Figure 13). Pelvic limb material is fragmen-
tary and poorh^ represented. Portions of femora include the
damaged proximal and distal ends of a right femur from MCZ
2406, an incomplete shaft from MCZ 2518, a damaged distal
end of a femur from MCZ 2407, and the proximal end of a small
femur and the distal end of another in scrap associated with
MCZ 2404. The last named elements are about half the size of
those from MCZ 2406 but the structure is the same. The femur
is of the peculiar Trimerorhachis type, with a cylindrical shaft
and expanded ends. There is a shallow depression on the ventral
proximal surface, bounded anteriorly by a projecting trochanter.
The outer edge of the latter curves inward toward the shaft
distally. The posterior margin of the depression opposite the
trochanter is not strongly developed, so that the Y-shaped pat-
tern of ventral ridges common in this area (Romer, 1947) is
lacking. The adductor crest is represented by a modest, narrow
ridge. Distally, this ridge ends in the center of the ventral sur-
face, a short distance above the end of the bone. Below it.
I
Fig. 13. Pelvic girdle and limb elements of Neldasaurus. A, right ilium,
outer side. B, left ilium, inner side. C, right ilium, distal surface. A-C,
MCZ 2518, X -75 approx. D and E, ventral and proximal views of proximal
end of femur. F and G, ventral and distal vievrs of distal end of femur.
D-G, MCZ 2406, X 1-5 approx. H and I, inner and proximal view of
proximal end of a right tibia of MCZ 2404, X -75 approx.
CHASE : NELDASAURUS WRIGHTAE 203
centrally, is a popliteal space. In MCZ 2407 this is a smooth
depression, but in MCZ 2406 it is a rugose, unfinished surface,
bounded dorsally by a low upraised ridge. The distal end of the
femur has a double condylar area for the head of the tibia and a
smaller, lateral condyle for the fibula.
The lower limb bones are represented by the proximal portion
of a right tibia and the distal end of a fibula from MCZ 2404,
and a similar fragment of fibula fi-om MCZ 2407. The proximal
end of the tibia is massive and there is a well developed cnemial
crest. The bone tapers rapidly to a slender shaft below the head.
The width of the proximal end is 1.7 cm, while the shaft, which is
oval in cross section, is only 5 mm across (Fig. 13, H and I).
The fragment assigned to the fibula shows a sub-oval shaft,
below which the distal end is considerably expanded, though
flattened. Identification of this bone is uncertain.
As far as known, the posterior limb bones of Neldasaurus are
indistinguishable from those of T rimer orliachis, with the pos-
sible exception of the distal end of the femur of MCZ 2406.
Review of the T rimer orliachis material in the collection of the
Museum of Comparative Zoology failed to uncover any femora
with the peculiar popliteal fossa of that bone.
Assignment of these materials to Neldasaurus is not as certain
as was the case with the pectoral limb. A single intercentrum of
T rimer orhachis-tyi)e is present in MCZ 2404 and another in
MCZ 2406. MCZ 2516 contains two small Trimerorhachis-like
intercentra and a pleurocentrum. However, all of these speci-
mens are from the Moran formation, from which there has
been no definite evidence of the occurrence of T rimer orliachis
(Olson, 1955). Most of the materials included in the specimens
listed here are N eldasaurus-\\ke, and the limb materials described
compare favorably in size with the other Neldasaiirus elements
in these specimens. For this reason, and because the few
Trimerorhachis-like vertebral elements do not seem sufficient
evidence on which to establish the presence of that animal in the
Moran, reference of the posterior limb elements to Neldasaurus
seems reasonable.
Scales
Remnants of dermal armor consisting of thin, imbricated bony
scales are preserved in a number of places in the specimens.
Isolated patches of integumental material are present in MCZ
2200 and MCZ 2518, and in MCZ 1371 considerable areas are
204
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
preserved. The pattern of scale distribution is obscure, but the
occurrence of scale material in association with much of the
skeleton suggests that scales covered most of the dorsal surface
of the body.
Scale structure and arrangement is best seen in MCZ 1371
(Fig. 14). In most cases the dermal covering as preserved con-
sists of groups of small, parallel bony rods, the longest about
1 cm long, covering all parts of the skeleton with the exception
of the skull and dermal shoulder girdle. Isolated patches of rods
were found on the posterior portions of the clavicles and inter-
clavicle but their separation from the main mass of integumental
material suggests post-mortem displacement. In a number of
areas the integument appears to have contained as many as seven
layers of bony rods, one above the other. This condition is sim-
ilar to that of the layers of ''bony fibrillae" described by Willis-
ton (1916) in Trimerorhachis.
Fig. 14. Dermal scales of Neldasaurus wrightae, MCZ 1371, X 5.
More or less complete scales are seen in only a few places.
The scales are elongate and rounded at both ends. As near as
can be determined, they are 10-12 mm long and 4-5 mm mde,
CHASE : NELDASAURUS WBIGHTAE 205
and, as shown by their layered condition, overlapped to a con-
siderable degree. However, they are very thin, and even in
areas where they are in several layers they easily conform to
the contours of the underlying skeleton. Colbert (1955) has
shown that the "bony fibrillae" described by Williston were in
fact remnants of elongate bony scales in Trimerorhachis. Com-
parison of the scales of Neldasaurus with the scales of Trimero-
rhachis from the specimens on which Colbert based his paper
shows the squamation of the two forms to be alike. As in
TrimerorhacTiis , some scales have a superficial layer of fine longi-
tudinal striations, below which is a layer of bony rods of variable
width arranged in concentric rings. The presence of the super-
ficial layer, however, is rare in Neldasaurus specimens.
OTHER OCCURRENCES OF NELDASAURUS
Fragmentary remains of small rhachitomous amphibians have
been recovered from several Dunkard localities in West Virginia
(Romer, 1952). Early collections of some of these were referred
to Trimerorhachis (Whipple and Case, 1930; Tilton, 1926).
Romer (1952), noting the absence of proof of the existence of
Trimerorhachis in these beds, described a series of vertebrae
from these collections, comparing them with ' ' a rhachitome from
the lower Wichita formations (Putnam, Moran) of Texas" in
reference to the then undescribed Neldasaurus. Comparison of
an isolated intereentrum from locality 28 in the Greene Group
and an articulated series of three vertebrae from locality 4 of
the Washington Group of West Virginia (Moran, 1952; Romer,
1952) (Carnegie Museum nos. 8568 and 8569) has been made
with comparable elements for Neldasaurus. These elements,
(see Romer, 1952, fig. 2, p. 65 and pi. 2, figs. 3 and 4) are
remarkably similar to those of Neldasaurus, particularly in the
height of the pleurocentra and narrow dimensions of the ascend-
ing processes of the intercentra.
Hotton (1959) compared vertebral elements of Acroplous with
those described by Romer, along with some other material from
the Dunkard region, finding an isolated parasphenoid and the
vertebral elements to bear a close resemblance to those of
Acroplous.
Correlation of the continental deposits of the Dunkard with
Texas beds is somewhat uncertain, but it is generally assumed
that the Washington and Greene Groups of the Dunkard are
equivalent in age to the Wichita in Texas (Romer, 1958) and
206 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
represent the base of the Lower Permian in that region. Further,
Romer (1952) has demonstrated a general similarity in the
faunal assemblages of the Dunkard and the Lower Permian
redbeds of Texas.
On the basis of the resemblance of the circumehordal elements
of Neldasaiirus and the Dunkard specimens, and the approximate
correlation of the Texas and West Virginia beds, there is a strong
likelihood that Ncldasaurus (and/or Acroplous) or some closely-
related form was present in the Dunkard.
DISCUSSION
CoMPAEisoN OF Neldttsaurus with other Genera
OF Trimeroriiachoids
The description of this new genus, obviously comparable in
many ways to Trimerorhachis, suggests the advisability of a
general review of the trimerorhachoids — the membership of this
group of labyrinthodonts, their structural patterns, classifica-
tion and relationships. To this end, the first portion of the dis-
cussion consists of a morphological comparison of Ncldasaurus
with the other trimerorhachoid genera.
Animals which have been suggested by various authors as
belonging to the trimerorhachoid complex include, Trimero-
rhachis, Saurerpeton, Eohrachyops, Acroplous, Dvinosaurus,
Chalcosaurus, Slaugenhopia, Eugyrinus and other "peliontids,"
and Dawsonia. Although the entire group is reviewed, detailed
comparisons are confined to those genera about whose structure
we have sufficient knowledge.
Before proceeding with individual comparisons, we may note
that the elongate snout of Neldasaurus is sharply contrasted to
the abbreviated snout of other members of the group. Other
features apparently related to snout proportions in which
Neldasaurus differs from typical trimerorhachoids include elon-
gation of the dermal bones of the face and palate, failure of the
lacrimal to reach the naris, and broad separation of the ehoanae
from the interpterygoid vacuities.
Neldasaurus and Trimerorhachis (Fig. 15). It is obvious
from the description given that Neldasaurus, apart from facial
length, has the combination of primitive and advanced characters
seen in the Trimerorhachoidea, as this term is used by Romer.
Further, except in this one feature, Neldasaurus is obviously
closely related to Trimerorhachis and will fit into the family
CHASE : NELDASAURUS WRIGHTAE
207
Trimerorliacliidae (Romer, 1947, p. 312). T rimer orhacJiis is a
well known form from the Texas redbeds whose anatomy has
been described by Cope (1878), Broom (1913), Williston (1915,
1916), Case (1911, 1935), and Watson (1956).
B
Fig. 15. Trimerorhachid skulls reduced to the same width. Dorsal and
palatal views of A-B, N eldasaurus ; C-D T rimer orhachis after Case; E-F,
Saurerpeton after Romer and Watson.
208 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
In addition to the resemblances between Neldasaurus and
T rimer orhachis related to general level of organization, the two
show numerous close resemblances of a special nature. The skulls
are alike in such features as the general pattern of dermal bones,
narrow interorbital width, prominent sensory canal grooves,
relation of the anterior palatal bones to the interpterygoid vacu-
ities, symphysial tusks and anterior palatal fenestrae, and the
remarkably similar basicranial articulation. The lower jaws are
similar, each having a modest retroarticular process and neither
showing a coronoid process. The postcranial skeletons of
Neldasaurus and T rimer orhachis are much alike and the limb
bones, as noted, are nearlj^ identical.
In spite of their close resemblance, there are a number of
significant differences between Neldasaurus and T rimer orhachis
other than those related to snout proportions. In some respects
Neldasaurus seems to be somewhat more primitive than Tri-
merorhachis. Here, in contrast to Trimerorhachis, a low ridge
bounds the skull table laterally, the skull is deeper posteriorly,
the jugal enters the orbital border, and a tusk pair persists on
the ectopterygoid.
Neldasaurus and Trimerorhachis also differ in several special
characters. Prootic, opisthotic and basisphenoid elements, as far
as known, were persistently cartilaginous in Neldasaurus, where-
as in Trimerorhachis they were ossified. The pattern of sensory
canal grooves in Neldasaurus differs from that of Trimerorhachis
in failure of the supraorbital groove to enter the lacrimal and
in the presence of a jugal sulcus, unknown in Trimerorhachis.
The lateral border of the choana in Neldasaurus is formed by an
anterior extension of the palatine. In Trimerorhachis it is
formed in more "normal" fashion by the maxilla. Separation
of the posterior wall of the "conical recess" from the anterior
wall of the "excavatio tympanica" by a groove in the medial
surface of the pterygoid in Neldasaurus is not shown by Tri-
merorhachis. Neldasaurus, with a remarkably high tooth count,
has twice as many marginal upper jaw teeth as Trimerorhachis
— about 108 as contrasted to 50. The dentary also bears more
teeth in N eldasaurus than in Trimerorhachis ■ — the approximate
count being 60 and 43 respectively. This may in part be related
to snout elongation in Neldasaurus, but tooth number is not nec-
essarily correlated with snout length in labj'rinthodonts (Chase,
1963).
A notable difference in the proportions of pleurocentra and
intercentra is seen in the vertebral columns of Neldasaurus and
Trimerorhachis.
CHASE : NELDASAURUS WRIGHTAE 209
Saurerpeton (Fig. 15). This small trimerorhachoid, known
mainly from the Upper Pennsylvanian of Linton, Ohio, has been
suggested as being closely related to T rimer or had lis (Romer,
1947). Romer further believed that "Pelion lyelli," also from
Linton, was a trimerorhachoid and erected a family Peliontidae
within the Trimerorhachoidea for its reception. However, recent
studies by Baird and Carroll (Carroll, 1964) show that the type of
"Pelion" is actually a primitive dissorophid of the genus Aniphi-
bamus, and that the characters which Romer gave to the ' ' Pelion-
tidae" actually pertain to Saurerpeton. All other Linton speci-
mens with an intertemporal previously described as "Pelion/'
and "Branchiosaurauus" (Romer, 1930) as well, apparently
belong to Saurerpeton.
Saurerpeton shows definite trimerorhachoid characters al-
though these are generally developed to a lesser degree than in
Neldasaurus and Trimerorhachis. It also resembles them in
such special characters as sensory canal grooves on at least some
of the dermal roof bones, and anterior palatal fenestrae for the
reception of symphysial tusks. These openings are clearly shown
in Romer 's "Pelion" (1930), though Steen (1931) and Watson
(1956) reconstruct "Pelion" without anterior palatal fenestrae.
It appears from Steen 's figure 15 of "Pelion lyelli" Wyman
that the prevomers are displaced from right to left so that, as
Romer (1947) suggests, she has interpreted the anterior palatal
fenestrae as the choanae. In this she is followed by Watson. In
some Saurerpeton specimens (Romer, 1930, fig. 6) it appears
that the palatine nearly reaches the prevomer lateral to the
choana, a feature suggestive of the pattern in Neldasaurus.
Saurerpeton resembles Trimerorhachis more than Neldasaurus
in such features as the proportions of the antorbital region and in
the small ectopterygoid in the palate.
In the retention of three palatal tusk pairs, Saurerpeton re-
sembles Neldasaurus rather than Trimerorhachis.
In several significant features Saurerpeton differs from Nelda-
saurus and Trimerorhachis, and in part, in keeping with its
greater age, it is seemingly more primitive. In Saurerpeton the
otic notch, though reduced, is more highly developed than in
Neldasaurus and Trimerorhachis. The orbits in Saurerpeton are
widely spaced and the jugal forms most of the lateral orbital
border. The skull in Saurerpeton appears to be deeper than in
Neldasaurus and Trimerorhachis. Watson (1956) estimates a
width to height skull ratio in "Pelion" of 1:1. The ratio in
Neldasaurus is 3:1 and in Trimerorhachis (Watson, 1956) it is
210 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
4.4 :1. Saurerpeton has fewer marginal upper jaw teeth (about
38) than Neldasaurus and T rimer orhachis and also lacks the
large number of small palatal teeth seen in them.
Saurerpeton differs from Neldasaurus and T rimer orhachis in
several special characters as follows :
(1) The occiput projects posteriorly beyond the skull table.
(2) The quadrate condyle appears to have been somewhat
anterior to the occipital condyle and well below the level
of the floor of the braincase.
(3) The tabulars are remarkably small.
(4) The palatine enters the border of the interpterygoid
vacuity between the vomer and the pterygoid.
(5) The maxilla is truncated posteriorly and does not reach
the subtemporal fossa.
(6) The quadrate ramus of the pterygoid is short and has a
vertical, ventrally directed tlange laterally.
(7) There is a coronoid process in the lower jaw.
The very long skull table in Saurerpeton is a remarkable
feature (Romer, 1947), and both supratemporal and squamosal
share in the elongation to an extent not seen in Neldasaurus and
T rimer orhachis.
EoBRACHYOPS (Fig. 16). This genus has been described by
Watson (1956) on the basis of a specimen from the Lower Perm-
ian of Texas. There is no preserved record concerning the col-
lection of the specimen, but the matrix strongly suggests the
Clear Fork and, since no materials of that group from higher
levels were collected before Olson's recent work, it is highly
probable that Eohrachyops comes from the Arroyo Formation.
Watson's specimen lacked the anterior end of both palate and
lower jaw. Eohrachyops resembles other trimerorhachoids in its
general level of organization and can readily be admitted to the
Trimerorhachoidea by accepting Ilotton's modification of
Romer 's definition of the superfamily to include the phrase,
"otic notch often poorly developed or lacking" (Hotton, 1959).
Eobrachyops differs from Neldasaurus, Trimer orhachis and
Saurerpeton in some special characters. It apparently had an
ossified supraoccipital ; the lacrimal is confined to the orbital
region, its usual area being provided for by an enlarged septo-
maxilla; the prefrontal reaches from orbit to naris; sensory
canal grooves are essentially confined to circumorbital rings.
In some ways Eohrachyops appears to be advanced beyond the
level of the other animals so far considered. The exoccipitals
CHASE : NELDASAURUS WRIGHTAE
211
are expanded, enclosing the vagal foramen and meeting the
tabulars, thus sheathing posteriorly the cartilaginous paroccip-
ital. Sphenethmoid and quadrate, as well as the prootic elements,
are unossitied. The interpterygoid vacuities are larger than in
B
Fig. 16. Trimerorhachid skulls reduced to the same width. Dorsal and
palatal views of A-B, N eldasaiirus ; C-D, Eobrachyops after Watson; E-F,
Acroplous after Hotton.
212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the previously discussed trimerorhaehoids, being separated from
the ehoauae only by a narrow strip of bone. The tabulars are
reduced to narrow strips. There is no otic notch.
Eobrachyops resembles Neldasaurus in two important skull
features not shared by other trimerorhaehoids. The elongate
ectopterygoid and the unusual palatine-vomer union lateral to
the choana suggest a real relationship between them.
On the other hand, Eobrachyops resembles Saurerpeton rather
than Neldasaurus and T rimer orliachis in the seven characters
listed above (p. 210). Further, we can add to this list of re-
semblances: (8) Jugal contact with the orbital border broad.
(9) Skull deep posteriorly. Also, Eobrachyops, like Saurerpeton,
has widely-spaced orbits.
The proportions of intercentra and pleurocentra in Eobrachy-
ops are like those of Neldasaurus and are contrasted to those of
Trimerorhachis. Conditions in Saurerpeton are uncertain.
AcROPLOUs (Fig. 16). This genus, recently described by
Hotton (1959), comes from the Speiser Formation, Council
Grove Group of the Wolf camp series of Kansas. According to
correlation tables of Permian formations in North America
(Dunbar et al., 1960), this level is approximately equivalent to
that of the Putnam Formation of the Wichita in Texas. Hence,
Acroplous was a contemporary of Neldasaurus and Trimero-
rhachis. Acroplous is a highly specialized form with an extremely
short face, in which the dermal bones are crowded by large,
dorsally situated nares. However, Acroplous has all of the im-
portant trimerorhachoid characters.
In some characters Acroplous resembles one or another of the
animals previously considered. The slit-like otic notch in
Acroplous suggests a primitive condition and most nearly re-
sembles that of Saurerpeton. The small ectopterygoid is like
that seen in Saurerpeton and Trimerorhachis. The lack of an
ossified quadrate and sphenethmoid in Acroplous, as well as
the presence of an ossified supraoccipital, recalls conditions in
Eobrachyops. Also, as in Eobrachyops, the tabulars are strip-
like, and the sensory canal grooves are confined to the orbit
region.
Most significant is the fact that Acroplous resembles Eobrachy-
ops and Saurerpeton in the nine important characters listed
above, which distinguish the last two animals from Neldasaurus
and Trimerorhachis.
The proportions of the intercentra and pleurocentra of
Acroplous resemble those of Neldasaurus and Eobrachyops.
CHASE : NELDASAURUS WRIGHTAE 213
DviNOSAURUS. This perennibraucliiate labyrinthodont of the
Russian Upper Permian has been described by Amalitsky (1924),
Sushldn (1936), and Bystrow (1938). Dvinosaurus has a short
face, anteriorly placed orbits, a very short skull, and lacks an
intertemporal element. Some of these characters may be asso-
ciated with "larval tendencies."
Dvinosaurus shows resemblance to the earlier trimerorhachoids
in almost every feature. The only notable exceptions are the
apparent lack of the intertemporal and — most important — the
shortness of the skull as a whole. Bystrow suggests a probably
compound nature of the postorbital — the posteromedial portion
of the bone representing a fused intertemporal moiety. The
absence of a discrete intertemporal is, therefore, no bar to
trimerorhachoid relationship for Dvinosaurus. Further, though
the skull is short, even here the proportions of pre- and post-
orbital segments are those of trimerorhachoids, and the general
shortness can be attributed to neoteny, since it is known in sev-
eral cases that the skull of larval labyi-inthodonts is proportion-
ately much shorter than in the adult.
Dvinosaurus is set apart from other trimerorhachoids by cer-
tain special characters, such as enclosure of the quadrate foramen
by the quadratojugal, a greatly reduced palatine, a somewhat
specialized palatal dentition, and a long-stemmed inter clavicle.
Further, it shows several advanced characters, as might be ex-
pected in a late-surviving trimerorhachoid. In two skull features
— an essentially double occipital condyle and a tabular process
which forms the outer end of the paroccipital bar — Dvinosaurus
is advanced over all other trimerorhachoids. The exoccipitals in
Dvinosaurus are expanded to surround the vagal foramen, a
feature in which Dvinosaurus is more advanced than other mem-
bers of this group with the exception of Eohrachyops. The de-
gree of ossification in Dvinosaurus, in which the otic elements
and the sphenethmoid remain cartilaginous while a small ossi-
fied supraoccipital occurs in at least some specimens, recalls
conditions in Acroplous and Eohrachyops. Dvinosaurus also has
an unossified gap between the posterior end of the quadrate
ramus of the pterygoid and the squamosal ( Bystrow 's ' ' pterygo-
squamosal fissure"). A similar condition obtains in Eohrachyops
but in no other trimerorhachoid.
In several characters Dvinosaurus seems to resemble Saurerp-
eton, Acroplous and Eohrachyops rather than Neldasaurus and
Trimerorhachis, although in Dvinosaurus these characters are
214 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
generally developed to a lesser degree. In Dvinosaurus the occi-
put projects behind the skull table, the quadrate condyle is
about even with or slightly in advance of the level of the occipital
condyle and lies below the level of the ventral surface of the
parasphenoid, the pterygoid has a ventral flange laterally, the
jugal has a broad contact with the orbital border, the orbits are
wide-spaced, and there is a coronoid process in the lower jaw.
Although the skull of Dvinosaurus is deepened posteriorly, it
is considerably flatter than in Saurerpeton, Acroplous or
Eohrachyops. Furthermore, Dvinosaurus resembles Neldasaurus
and T rimer orhachis and is in contrast to the other three animals
in several important characters. In Dvinosaurus the otic notch
is reduced and is essentially comparable to the otic notch in
Neldasaurus and T rimer orhachis. The tabular is small, though
it is not reduced to a narrow strip. The pattern of the sensory
canal grooves on the dermal roof bones of Dvinosaurus closely
resembles that of T rimer orhachis and is nearly identical to that
of Neldasaurus.
In Dvinosaurus, as in Neldasaurus and T rimer orhachis, there
is a union of the vomer and the palatine in the palate, and the
maxilla is not truncated posteriorly but reaches the anterior end
of the subtemporal fossa.
The vertebral elements of Dvinosaurus are most like those of
T rimer orhachis.
Chalcosaurus. This poorly known labyrinthodont was tenta-
tively assigned to the Trimerorhachidae by Komer (1947). The
original specimen has been lost, but Efremov redescribed the
genus on the basis of the literature and published a figure (1940,
pi. II, fig. 2). On the basis of the description, assignment of
Chalcosaurus to the trimerorhachid group seems reasonable, but
knowledge of this animal is too scanty to permit definitive
comparisons.
Slaugenhopia. This genus, recently described by Olson
(1962), is based on partial skull material from the San Angelo
of Texas and a partial lower jaw from the same horizon (Olson
and Beerbower, 1953). The incomplete nature of the remains
precludes definitive discussion and we here provisionally accept
Olson's assignment of Slaugenhopia to the Trimerorhachidae.
EuGYRiNUS. This small, short-faced genus from the Pennsyl-
vanian Coal Measures in Lancashire has been described by
Woodward (1891), and by Watson (1921, 1940), and its phylo-
genetic position has been considered by Romer (1947). Romer
assigned it to the "Peliontidae, " a group now known to be
CHASE : NELDASAURUS WRIGHTAE 215
unreal, and considered it a young, though probably post-larval
member of the primitive rhachitome group. Further, AVatson
(1940) has compared Eugyrinus with Dendrerpeton. More re-
cent unpublished studies by Carroll indicate that Eugyrinus is
more closely allied to the edopsoids than to the trimerorhachoids,
and should probably be removed from the trimerorhachoid com-
plex and placed near the base of the edopsoid group. Hence,
Eugyrinus is not here considered as a trimerorhachoid.
Other "peliontids." Included tentatively in Romer's fam-
ily Peliontidae with "Pelion" and Eugyrinus were Erpcto-
cephalus and various Mazon Creek "larvae."
The position of Erpetocephalus is problematical. Aside from
traces of shoulder girdle, only the dorsal surface of the head is
known (Romer, 1947). According to Baird (in litteris), "Erpe-
tocephalus is a labyrinthodont of uncertain position ; Romer
(1945) very plausibly classified it as a dendrerpetontid edop-
soid." There seems to be no strong argument for assigning it to
the trimerorhachoid complex.
Baird 's studies have further shown that the Mazon Creek
larvae are either indeterminate or that they are not trimero-
rhachoids.
Dawsoxia polydens was based on a number of fragmentary
remains of Carboniferous age (Fritsch, 1901). Romer (1945)
reviewed the material, finding identification uncertain. Figures
(Fritsch, 1901, vol. 1, figs. 42, 43) and copper casts of the orig-
inal specimens support the assumption that we are here dealing
with a primitive temnospondyl of indeterminate affinities.
Interrelationships of Trimerorhachoid Genera
The genera here considered trimerorhachoids, in spite of many
variations in structure, share a large number of characters re-
lated to their general level of organization. The resemblances
that in combination suggest the Trimerorhachoidea to have been
a natural group are :
(1) Presence of an intertemporal element- — the compound
nature of the postorbital in Dvinosaurus justifies its in-
clusion here.
(2) Single occipital condyle — the condyle tending towards
a double condition in Dvinosaurus ; conditions in Acrop-
lous are incompletely known.
(3) A movable basal articulation.
216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
(4) Tendency towards flattening of the skull, at least in the
antorbital region.
(5) Otic notch small, poorly developed or lacking.
(6) Postorbital segment of the skull expanded.
(7) Usually short-faced — Neldasaurus is the only exception.
(8) Quadrate close to the level of the occipital condyle.
(9) Presence of sensory canal grooves on the dermal roof
bones, often well developed.
(10) Interpterygoid vacuities enlarged.
(11) Broad cultriform process of the parasphenoid.
(12) Anterior palatal feuestrae and symphysial tusks, where
known.
(13) A modest retroarticular process in the lower jaw.
(14) Ventral dermal shoulder girdle expanded.
(15) Body flattened.
(16) Limbs small.
Phylogenetic relationships and classification of the trimero-
rhachoids have been recently treated, principally by Komer
(1947), Watson (1956), and Hotton (1959), with varying re-
sults. The description of Neldasaurus and the discussion of
trimerorhachoids in the last section provide an opportunity to
review relationships within the group.
Neldasaurus resembles T rimer orhachis more closely than any
other trimerorhachoid. However, these animals differ in special
characters, some of advanced nature, which iiidicate that the re-
lationship between them is probably not an ancestor-descendant
one. Their typically trimerorhachoid features and their appear-
ance near the base of the Permian suggest that they probably
arose from a trimerorhachoid ancestor in the Pennsylvanian.
Saurerpeton has been suggested as being closely related to the
ancestry of T rimer orliachis by Romer (1947) and Hotton (1959).
Several primitive features in Saurerpeton anticipate conditions
in Neldasaurus and T rimer orhachis. However, some characters,
e.g. the anterior position of the quadrate condyle, and failure of
the pterygoids to meet the vomers, suggest that Saurerpeton has
already advanced beyond the evolutionary stage represented by
Neldasaurus and T rimer orhachis. We are thus led to the con-
clusion that Saurerpeton, though close to the ancestry of Nelda-
saurus and Trimerorhachis, is not directly ancestral to them.
Eohrachyops, though sharing some special characters with
Neldasaurus and Trimerorhachis, shares a long list of important
characters with Saurerpeton which are in contrast to conditions
CHASE : NELDASAURUS WRTfiTITAE 217
ill Xcldasdunis and Trimei'orliachis. The combination of primi-
tive and special characters shown by Saurerpeton would logically
place it close to the ancestry of Eohrachyops.
Aero pious has been shown to resemble Saurerpeton and
Eohrachyops in characters that distinguish them from Nelcla-
saurus and Trimcrorliachis. Ilotton (1959J recognized the sim-
ilarity of Acroplous to Eohrachyops, but placed it closer to
Trimcrorhachis mainh' on the basis of three characters. These
were :
(1) The presence of narrow midline elements in Acroplous
and T^im erorh ach is.
(2) Symphysial tusks and anterior palatal fenestrae in
Acroplous and Trimerorhachis.
(3) The "normal" pattern of palatal bones adjacent to the
clioana in Acroplous and Trimerorhachis, as opposed to
the "peculiar" situation in Eohrachyops, where the pal-
atine provides the lateral boundary of the choana.
A review of these dilferences suggests that they may not sup-
port closer relationshii? of Acroplous to Trimerorhachis than to
Eohrachyops.
(1) The narrow midline elements in Trimerorhachis, as well
as in Neldasaurus, appear to result from a medial shift in
the position of the orbits. In primitive temnospondyls the
orbits are widely separated, the postorbital is confined
to the posterior border of the orbit and the jugal forms
much of the lateral border. With a medial shift in posi-
tion, the orbit would, in effect, "move away" from the
jugal, the postorl)ital would have a more lateral position
relative to the orbit and by an anterior extension could
replace (cf. Trimerorhachis) or nearly replace (cf. Nelda-
saurus) the jugal on the outer orbital border; the mid-
line elements would at the same time be constricted.
In Acroplous the strong jugal contact with the lateral
border of the orbit and the position of the postorbital
immediately behind the orbit suggest that the narrow
midline elements here are the result of absolute increase
in size of the orbits rather than a shift in position.
(2) The anterior end of the palate and the lower jaw in
Eohrachyops are not known, and hence we do not know
whether symphysial tusks and anterior palatal fenestrae
were present or not in Eohrachyops.
218 BULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY
(3) The uniusual union of the palatine with the vomer on
the lateral border of tlie choana in Eobrachnops is a real
differenee between Eohrachyops and Acruplous. IIow-
evei', the occurrence of a similar pattern in ycldas(iun(.s.
as noted above, a form obviously close to Triinerorhachis,
suggests that the "normal" pattern may have been modi-
fied in parallel fashion in Edhyachijops and Nddasaurus.
Although in many respects, as llotton noted, Acrophnis aj)-
pears to be a morphological intermediate between Saiircrpeton
and Eohrachyops, special characters in Acroplous suggest that
it probably represents a specialized side branch of the line lead-
ing from Saitrcrpcton to Eobrachyops.
The aberrant structure of DL'i)iosaiinis appears to remove it
from close relationship to any trimerorhaehoid. Romer (1947)
and liotton (1959) suggested relationship of Dvinosaurus to
Trimerorluichis and Saiircrpdon \ Watson (1956) placed it in
the line stennning from Eohrachyops to the Triassic brachyopids.
In spite of resemblances of Acroplous and Eohrachyops to
Dvinosaurus, these animals are too specialized to be reasonably'
considered ancestral to Dvinosaurus. Further, Dvinosaurus
shares witli Xddusaurus and Triiin r(nh(ivl}is a significant num-
ber of those characters which su])])()rt separation of Xcldasaurus
and Triinerorhachis from Acroplous and Eohrachyops. Hence,
we arrive at the conclusion that, as suggested by Iiomer and
llotton, Dvinosaurus is related to TriinerorJiacliis and miglil
reasonably be considered a terminal member of the Triinero-
rhachis-Neldasaurus group.
Review of the Trimerorhachoidea supports the assumption that
they form a natural group. The superfamily can be defined b\
the characters listed on pages 215-216. The pliylogenetic conclu-
sions reached above suggest the existence of three subgroups with-
in the Trimerorhachoidea. The families Trimerorhaehidae and
Dvinosauridae of Romer can be retained, the first including
T rimer orhachis, Neldasaurus, Chalcosaurus and Slaugenhopia,
the second containing Drinosaurus. I propose the erection of a
new family Saurerpetontidae, with Saurcrpcton as the type
genus, for the reception of Saurerpeton, Acroplous and
Eohrachyops.
CHASE : NELDASAURUS WRIGHTAE 219
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220 bulletix : museum of comparative zoology
Trimerorhachoid Origins and Relationships
The origin of the trimerorhaehoids is problematical. Some
characters of this group are advanced and of a sort typically
met with in Triassic labyrinthodonts. On the other hand, a
number of primitive features in the trimerorhaehoids, notably
single condyle, movable basal articulation, and retention of inter-
temporal, are not found in labyrinthodonts above the edopsoid
level and hence suggest an origin of the trimerorhaehoids from
an early member of the edopsoid group.
Among the earliest edopsoids are such Pennsylvanian forms
as Gaudrya from Nyfany, the dendrerpetontids from Joggins,
and the English edopsoid-like labyrinthodont, Eugyrinns. A
trend in dermal roof pattern in edo]isoids, characterized by
withdrawal of the lacrimal from the anterior orbital border, is
already well established in (laudrya and the dendrerpetontids.
which would bar them from being directly ancestral to the
trimerorhaehoids.
Eugyrinus shows a number of characters that might be ex-
pected in a trimerorhachoid ancestor, such as: (a) short skull.
(b) broad snout, (c) short face, (d) small otic notch, (e) skull
table moderately expanded, (f) apparently single condyle, (g)
parietal foramen close to orbits, (h) primitive dermal roof pat-
tern, including intertemporal, (i) traces of lateral line grooves
on dermal bones of skull roof, (j) open palate, but with ptery-
goids still extending anteriorly to nearly meet cultriform jiroe-
esses on the anteromedial edges of the interpterygoid vacuities,
(k) apparently movable basal articulation, (1) lower jaw with a
modest retroarticular process and coroiioid jirocess, the last
named structure occurring in many, though not all trimero-
rhaehoids.
Only in one specific point is Eugyri)ius too specialized to be a
trimerorhachoid ancestor — the fact that, unlike conditions in
any trimerorhachoid, the quadratojugal enters into the jaw
articulation. However, an edopsoid resembling Eugyrinus in all
features except this one specalization Avould be a reasonable
trimerorhachoid ancestor.
The possible relationship of trimerorhaehoids to the Triassic
brachyopids has been treated by Watson, Nilsson and Romer.
Watson (1956) proposes Eohrachyops and Dvinosaurus (1919.
1956) as ancestors of the Triassic brachyopids; Dvinoscn())is,
according to that author, represents a more or less intermediate
staue in a line stemming from an ancestral form close to
CHASE : NELDASAimrS WRIGIITAE 221
Eobravhyops in the Lower Permian. Nilsson (1937) agrees in
general that Dvinosaurus. tliough not directly ancestral, may be
close to the ancestry of brachyopids. Romer (1947) denied
relationship of Dvinosaurus to the brachyopids, considering it
closer to Saurerpeton and Trivicrorhachis. and suggested the
metoposaurs as brachyopid ancestors.
In its parabolic skull outline and anteriorly placed orbits,
Dvinosaurus superficially resembles the brachyopids. However,
the pattern of dermal bones in the skull roof of Dvinosaurus,
including reduction of the postparietal and the topography of
the postorbital region, is in contrast to conditions in the brachy-
opids where the postparietals are not notably reduced and the
postfrontal-supratemporal contact, as Romer points out, is more
reasonably derived from the more "normal" pattern of typical
rhachitomes. Further, the lacrimal contact with the orbit and
naris in Dvinosaurus differs from the brachyopid condition, in
which the lacrimal is typically reduced and withdrawn from the
orbit border. The skull table of Dvinosaurus is shorter than in
the brachyopids. AVatson "writes off" such features as a mov-
able basal articulation and tlic greater anterior extent of the
pterygoids in Dvinosaurus as merely the retention of primitive
characters. However, the retention of such a remarkably primi-
tive feature as a movable basal articulation at such a late
date is a condition one would hardly expect if we were, in
Dvinosaurus, confronted by an immediate brachyopid ancestor.
Other palatal structures of Drinosaitrus are also features
hardly to be expected in an innnediate ancestor of the brachy-
opids. In Dvinosaurus, the vomers are expanded to meet the
ptei-ygoids on the lateral border of the interpterygoid vacuities,
so that the palatine, which in Dvinosaurus is a remarka])ly small
bone, takes no part in the border of the interpterygoid vacuity,
whereas in brachyopids the palatine forms the anterior border
of that opening. Perhaps significant is the presence of anterior
palatal fenestrae in Dvinosaio-iis — none are recorded in the
brachyopids.
Some differences between Dvinosaurus and the brachyopids
may reflect a more primitive level of organization in Dvinosaurus
and its obvious neoteny. However, the number of morphological
differences that exist between Dvinosaurus and the brachyopids
rule against a close relationship.
Eohrachijops is more like the bracliyopids than is Dvinosaurus
in such features as: less broadly rounded snout; nares, though
222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
widely separated, closer together than in Dvinosaurus; orbits of
more "normal" proportions; very shallow cheek below orbit;
pattern of postorbital region ; lacrimal, though still in contact
with orbit, greatly reduced; longer skull table than in Dvino-
saurus: entrance of palatine into the margin of interpterygoid
vacuity.
On the other hand, such specialized characters in Eohrachyops
as tiny tabulars, great reduction of sensory canal grooves, ab-
breviated maxilla, peculiar pattern of palatal bones adjacent to
choanae, and position of quadrate articulation further forward
than in some brachyopids, tend to remove it from a directly an-
cestral position.
In i)hu"ing Eohrachyops close to the ancestry of the brachy-
opids, Watson listed five characters in which Eohrachyops and
Dvinosaurus resemble the brachyopids and "differ from all other
labyrinthodonts" (1956, p. 365). This, which would appear to
be a clinching argument, loses strength on closer examination.
Two of the five characters, it is true, are found only in Eohrachy-
ops, I)ri)iosau)-us and brachyopids. These are: (1) a distinct
space, formerly occupied by a cartilaginous ridge on the posterior
surface of the quadrate, separates the hinder border of the ptery-
goid from those of the squamosal and ({uadratojugal ; (2) the
outer surface of the squamosal and (juadratojugal passes round
onto the posterior face of the quadrate and there forms a later-
ally concave, nearly vertical surface.
However, the other three charact<'rs cited by Watson are not
exclusive to Eohrachyops, Dvinosaurus and the brachyopids.
but are present in other trimerorhachoids (Sauro'peton, Acrop-
Jous), and two of the three — projecting occiput (m?toposaurs.
plagiosaurs) and ventral position of the quadrate condyle (edop-
soids, eryopsoids) — may also be found in other labyrinthodont
groups.
If Eohrachyops is a Permian representative of the brachyopids,
special cliaracters and typically advanced features in Eohrachij-
ops would certainly debar Dvinosaurus from a position inter-
mediate between Eohrachyops and later brachyopid-;.
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iy:i-±. On the Dvinosainidac, a family of labyrinthodonts from the
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CHASE : NELDASAURUS WBIGHTAE 223
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1943. The origin of tetrapods. Biol. Rev., 18: 78-98.
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BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
^-^■■P^' ■'
PLATE 1
Xeldasaurus icrightae, n. gen., ii. sp. Holotype skull in dorsal view.
MCZ 2200, Block A, X -66 approx.
CTIASE : XELDASAT'RfS WRTGTITAE
PLATE 2
Xrhld.sauras lorightae, n. gen., ii. sp. Ilolotype skull in palatal view.
MCZ 2200, Block A, X -66 approx.
BULLETIX : MUSEUM OF COMPARATIVE ZOOLOGY
PLATE 3
N eld u. sail ni.s icriglitue, slab showing dorsal vertebrae in series and a par-
tially articulated left forelimb. MCZ 1371. Block A, X 1 approx.
Bulletin of the Museum of Comparative Zoology
HARVAKD UNIVEKSITY
Vol. 133, No. 4
THE GENERA OP THE CHILOCORINI
(COLEOPTERA, COCCINELLIDAE)
By
Edward A. Chapin
Museum of Comparative Zoology
CAMBEIDGE, MASS., U.S.A.
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Bulletin of the Museum of Comparative Zoology
HAEVAED UNIVEESITY
Vol. 133, No. 4
THE GENERA OP THE CHILOCORINI
(COLEOPTERA, COCCINELLIDAE)
By
Edward A. Chapin
Museum of Comparative Zoology
CAMBEIDGE, MASS., U.S.A.
FEINTED FOE THE MUSEUM
September, 1965
Bull. -Mils. Coinp. Zool., Hiuvnnl Univ., 133 (4): 227-271, Sept. 1965
Xo. 4. — The Genera of the Chilocorini {Coleopi< va, Coccinellidae)
By
Edward A. Chapin
INTRODUCTION
It is now ninety-one years since the latest attempt tu evaluate
the described genera of Coccinellidae of the world was made by
G. R. Crotch (A Revision of the coleopterous family Coccinel-
lidae, London, 311 pages, 1874). Since 1874, there has appeared
a bibliographic catalog of the species of this family by R. Kors-
chefsky (Coleopterorum Catalogus, Junk-Schenkling, Partes 118,
120, Coccinellidae, 659 pages, 1931-1932). A critical review of
the genera and species of this family is long overdue but has
now become virtually impossible for any one worker to ac-
complish. So it seems best to select some small, well circum-
scribed group within the famil.y as a starting point. I have
chosen the Chilocorini, a well known and easily recognized tribe.
This tribe is a very compact group of genera distributed
generally throughout the world. In the Korchefsky catalog
twenty-four generic or subgeneric names are listed. One of
these, Clanis, is a preoccupied name that has already been re-
placed. A second, Notolipernes, had been placed as a s;snionym
of Telsimia and removed from this tribe at the time that the
catalog was going through press. Corystes Mulsant was de-
scribed in the Hyperaspini, and its placement has been under
discussion since Chapuis {in Lacordaire, 1876, Genera Coleopt.,
12:244, 249) referred it to the Chilocorini. The genus is here
returned to the Hyperaspini where it apparently belongs. I have
found no explanation for the inclusion of Elpis Mulsant in the
Chilocorini and, after an examination of the type species, E.
dolens Mulsant, have returned it to the Coccinellini to a position
near Menochilus Timberlake. There remain twenty names that
are properly included in this tribe. Of these, I have been able
to examine specimens, usually of the type species, of seventeen.
In the course of my work I have found it necessary to propose
two new genera. As far as possible the characterizations of the
genera have been taken from the respective tjqDe species.
230 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
I wish to express my thanks to Dr. P. J. Darlington, Jr. for
encouragement and assistance in securing necessary material for
study, to Dr. R. D. Pope for specimens of Elpis dolens Mulsant,
Endochilus plagiatus Sicard, and Endochilus styx Sicard, to Mr.
E. B. Britton and Mr. G. F. Gross for specimens of three species
of Orcus. My thanks are also due to Mrs. Morna MacLeod for
preparing the illustrations from my tracings and to Mrs. Neil
Barth for typing the manuscript.
Specimens for study have been dissected, the pertinent parts
cleared of muscular tissue in cold KOH 10 per cent solution,
and passed through the alcohols and oil of wintergreen (methyl
salicylate) into Canada balsam. The drawings were made by
tracing the outlines projected onto paper at various magnifica-
tions through a Bioscope projection apparatus. There follows
a discussion of the parts studied.
STRUCTURE AND TERMINOLOGY
Antenna. The number of segments varies from ten to seven.
The basal segment may be more or less cylindrical or it may be
produced laterally at its apex in a rounded or subconical lobe
(HaJmus, Priasus, Orcus). The apical segment may be very
short and embedded in the penultimate segment or it may be as
long as or longer than the penultimate segment {Anisorcus,
Orcus, Chilocorus, Egius, Priasus).
Mandibles. In place of the supplementary tooth (absent in all
Chilocorini), there is a short knife-like ridge on the ventral
aspect running backward a short distance from the apex. In
general the mandibles follow a pattern throughout the Chilo-
corini. In three cases {Axion, Egius, Priasus) the apices are
obtusely triangular, without the hooked tip of the others. In
Halmus, the right, and to a lesser extent the left, mandible has
a more or less broad triangular internal tooth. Rudiment of this
tooth can be seen on the right mandible of Anisorcus.
Maxilla. 1 have not attempted to describe the differences
which exist in the maxilla proper, but have confined my atten-
tion to the palpus. The terminal segment, described as securi-
form for the family as a whole, varies from that shape in several
instances. In Anisorcus, Egius, and Phaenochilus the segment
is cylindro-acuminate while in Chilocorus and Priasus the sides
are almost parallel. In Halmus the segment is barrel-shaped Avitli
oblique apex.
CHAPIN : GENERA OF THE CHILOCORINI 231
Liijulu. Ill general this sclerite is soft, quadrate, with an-
terior angk^s rounded. It bears the labial palpi which show
some variation. The terminal segment may be shorter than the
preceding, or longer, as in Harpasus and Axion. It may be
slender, cylindro-acuminate as in E.iochomus or short, stout, and
conical as in Halmus.
The number of visible abdominal sternites has been considered
as one of the prime characters in coccinellid taxonomy. How-
ever, in the Chilocorini, most of the genera have six visible
sternites in the male and five in the female. This is because the
fifth in the male is truncate or broadly emarginate, thereby dis-
closing the sixth sternite. In the genus Orcus, the fifth is as
broadly rounded in the male as in the female, thereby con-
cealing the sixth in both sexes. On the other hand, in the genera
Halmus, Anisorcus, and Egiius, the fifth is truncate or emargi-
nate in the female as in the male, thereby disclosing the sixth
in both sexes. Except for these slight difi'erences, the fifth
sternite is unmodified. The first visible sternite carries the
metacoxal arcs — thin ridges arising at the sides of the inter-
coxal process of the sternite and extending out. In the Chiloco-
rini we find the same types that have been long used in the
classifications of the Scymnini. Thus, in Exochomus, Brunms
(male), and Brumoidcs the arcs are complete or closed, as in
Fullus. In Axiom, Brumus (female), Cladis {Clanis renamed),
Curinus, Harpasus, and Zagrcus, the ares are incomplete but
directed forward at their extremities as in Scymnus. These
genera are characterized by the presence of tibial spurs. The
arcs are incomplete and directed outward at their extremities or
are fused with the posterior margin of the sternite in those
genera without tibial spurs. Afiisorcus, Orcus, and Priasus have
arcs directed outward, as in Nephus, while Chilocorus, Egius,
and Halmus have arcs fused with the margin of the sternite as
in Diomus.
Legs. The femora are unmodified throughout the group, ex-
cept for a shallow groove for the reception of the tibiae. The
tibiae show some modifications. In Brumus and Brumoidcs, the
legs are unusually slender, and in Brumoidcs especially, the
tibiae are many times longer than broad. Three genera, Chilo-
corus, Egius, and Phacnochilus have developed a stout, tri-
angular tooth on the outer margin of the basal half of the tibia.
232 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
Claws. Ill general the form of the tarsal claws is characteristic
of the genus. In this tribe there are three genera with com-
pletely simple claws {Brunius, Brumoides, and Priasus) ; claws
with a very small triangular tooth basally or slightly postmedian
are present in Endochilus and Parapriasus (in part) ; and claws
with a heavy triangular or subquadrate tooth are in the other
genera. Usually the tA'pe of claw can be determined from a dry
specimen but it may be necessary to mount a claw in balsam
in order to perceive a minute tooth as in Endochilus or in some
species of Parapriasus. In the figures which accompany this
paper, all illustrations of claws are drawn to the same scale.
Male genitalia. The median lobe, except in one genus, is tri-
angular in dorsal view. It may be perceptilily asymmetrical as
in Axion, Chilocorus, Curinus, Egius and Exochomus. It may he
less than twice as long as broad, in Brumoides and Claelis, or as
much as seven times longer than broad as in Egius and Exo-
cJwniiis. In Egius, the lobe is longer than the parameres, in
other genera equal {Chilocorus) or shorter. The exception is the
genus Halmus, in Avhich case the lobe is parallel sided, longer
than the parameres, the tip upturned and in side view concave
in the basal two-thirds and again in the apical third. The
parameres are slender throughout as in Halmus and Priasus or
are moderately broad and spatulate and may be slightly or
strongly constricted in basal half. In the genus Anisorcus they
are very broad and concave, without any basal constriction.
The trahes is simple, slightly curved, and slightly thickened at
its free extremity. It is unusually thin in Axion. The sipho
does not vary much from genus to genus. Moderately stout and
curved, with the siphonal capsule more or less similar in all
genera but Anisorcus, and the apex variously modified, the apex
has more value for specific than for generic identifications. In
the illustrations, the aedeagus and sipho are always drawn to the
same scale.
Female gcmtalia. The recepiaculum seminis is of a type not
to be found, to the best of my knowledge, elsewhere in the
Coccinellidae. The body is stout, very much so in the l)asal part,
the ramus is entirely absent, the nodulus usually absent but
represented in some genera {Egius, Zagreus) by a small button-
like dome, and the cornu small, rounded, and bent at right
angles to the body. In four genera (Anisoreus, Chilocorus,
Phaenochilus, and Egius) most of the cornu is replaced by an
appendix, definitely separate from the body. In the genus
CHAPIN: GENERA OF THE CHILOCORINI 233
Halmus the body is more slender and appears nearly coceinel-
line in its form. The accessory gland is moderate to very long,
sometimes, as in Axion, several times as long as the recepta-
culum. The sperm duct is extraordinary as among Coccinel-
lidae in my experience. The duct consists of two parts : one
of small diameter arises at the head of the bursa copulatrix, at
the base of the infundibulum, and connects with the second part
of greater diameter, which leads to the receptaculum. These
parts are roughly equal in length. Two genera deviate from
the norm. In Anisorcus, the first part seems to be an ex-
tension of the bursa ; it is of greater diameter than usual but
is very thin-walled and definitely shorter than the second part.
In Halmus the first part is longer than the second and almost
as great in diameter. The infundibulum is present in all but
four genera and is very variable in shape and size. In Egius,
Phaenochilus, and Chilocorus it is replaced by a fleshy pro-
tuberance, unsclerotized, on the head of the bursa; in Anisor-
cus it is completely lacking. The bursa copulatrix in all genera
is a simple thin-walled sac with no modifications, such as are
seen in certain species of Hippodamia. The hemisternites (di-
vided eighth sternite) are usually subtriangular, more or less
elongate, each with a small button-like stylus at apex. In the
genera Endochilus, Anisorcus, Egius, Phaenochilus, and Chi-
locorus, the styli are absent while in Halmus they are cylindri-
cal and longer than wide.
In the key which follows it will be noticed that half of
the genera studied lack tibial spurs, all but two of which
{Chilocorus and Egius) are restricted to the Old World, while
the other lialf of the genera possess tibial spurs and all but
two {Brumus and Brumoides) are confined to the New World.
Of these four exceptions, one of each pair is nearly world-
wide, the other closely related to it. This suggests that Bru-
moides developed first in the Ncav World, then spread into most
of the Old World, with Brumus as an Old World offshoot, and
that Chilocorus was originally an Old World genus which
invaded the New World, with Egius as its New World segre-
gate.
234 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY
SYSTEMATIC SECTION
Tribe CHILOCORINI Costa
Costa, 1849, Fauna Eegno Napoli, 1:9.
Coccinellidae of small to large size, 2.0-8.0 mm, form oval
to nearly circular, moderately to strongly convex, upper sur-
face glabrous or pubescent. Antenna of less than eleven seg-
ments, short, terminal segments forming a fulsiform club, its
origin concealed beneath the genal extension of the clypeus
which enters or passes immediately beneath the eye. Mandible
without supplementary apical or subapical tooth but with a
sharp ridge which runs back a short distance from apex on
the ventral face. Prosternal lobe without carinae. Abdomen
usually with six visible sternites in male, five in female, some-
times five in both sexes or six in both sexes. Legs usually
normal, occasionallj' with modified tibiae, tibiae with or with-
out terminal spurs, tarsi four-segmented, tarsal claws simple,
or swollen at base, or with basal tooth. Male genitalia : median
lobe of aedeagus usually elongate triangular, compressed, often
asymmetrical near apex, parameres shorter or longer than the
median lobe, occasionally modified at apices. Female genitalia :
receptaculum seminis very stout, somewhat bent, ramus absent,
nodulus occasionally developed but never prominent, cornu a
small terminal portion, sometimes with an appendix, accessory
gland usually very large. Sperm duct very long, of two parts.
The part leading into the receptaculum, usually half the total
length, is a moderately coarse tube, the part leading from the
bursa a thinner, usually much thinner, tube. Infundibulum
present or absent; if present the sperm duct arises at its base.
Bursa copulatrix a large, simple sac. Hemisternites usually
elongate, sometimes short, subtriangular, rounded at apices, styli
small, usually button-shaped with two to four long setae, some-
times absent.
KEY TO GENEBA OF TEIBE
1. Tibial spurs present on legs II and III, metacoxal ares as in
Scymnus or Piilliis, antenna 10, 9, or 8-segniented 2
Tibial spurs absent on all legs, metacoxal arcs as in Xcphus or
Diomus, antenna 9, 8, or 7-segmented 9
2. Tarsal claw simple, sometimes slightly thickened at base 3
Tarsal claw with tooth in basal half, which is sometimes quadrate,
sometimes acute, often very small 4
3. Antenna 10-segmented, Old World Brumus Mulsant
Antenna 8-segmeuted, world-wide Brumoides n. gen.
4. Antenna 10-scgmented 5
CHAPIN : GENERA OF THE CHILOCORINI 235
Antenna 9-segmented Hnrpasus Mulsant
Antenna 8-segmented Zagreus Mulsant
5. Elytra! epipleura f oveolate for reception of tips of femora 6
Elytral epipleura not foveolate 8
6. Elytra without retlexed margins, with marginal bead, size small
to moderate, up to 6 mm 7
Elytral margins feebly but distinctly retlexed, with or without
marginal bead, size very large, up to 8 mm . . Axion Mulsant
7. Size larger, up to 6 mm, elytra blue, parameres not appendieulate
Curinus Mulsant
Size smaller, up to 3.75 mm, elytra black or bluisli l>lack, with
red basal spot, parameres appendieulate Arawana Leng
8. Metacoxal ares complete or virtually so . Exocliomus Eedtenbacher
Metacoxal arcs widely incomplete Cladis Mulsant
9. Eeceptaculum seminis without apical appendix, infundibulum
usually present (absent in Endochilus) 10
Eeceptaculum seminis with an apical appendix, infundibulum
absent 14
10. Antenna 9-segmented 11
Antenna with less than 9 segments 12
11. Tarsal claw without basal tooth but somewhat swollen at base,
elytral margin feebly retlexed in apical half Priasus Mulsant
Tarsal claw with sharp quadrate basal tooth, elytral margin
strongly retlexed throughout Parapriasus n. gen.
12. Upper surface wholly or in large part pubescent, infundibulum
absent Endochilus Weise
Upper surface glabrous, anterior angles of pronotum sometimes
sparsely hairy, infundibulum present 13
13. Antenna 7-segmented Hahnus Mulsant
Antenna 8-segmented Orciis ]\lulsant
14. Antenna 7-segmented, all tibiae without tooth on outer margin
in basal half Anisorcus Crotch
Antenna 8-segmented, tibiae II and III, sometimes I, with tri-
angular tooth on outer margin at extremity of tarsal groove 15
15. Terminal segment of maxillary palp one and one-half times as
long as wide, its apex strongly oblique, elytral margin feebly
retlexed Chilocorus Leach
Terminal segment of maxillary palp two or three times as long
as wide, apex acuminate 16
16. Elytra strongly alutaceous, the lateral margins not retlexed,
terminal segment of maxillary palp about twice as long as
wide, cylindro-acuminate, tooth on tarsal claw weak, Xew
World Egius Mulsant
Elytra not alutaceous, the lateral margins strongly retlexed, ter-
minal segment of maxillary palp three times as long as wide,
gradually narrowed to a blunt point, tooth on tarsal claw
very strong, nearly as long as apical portion of claw. Old
World Phaenochihis Weise
236
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
BrUMUS Mulsant
Brumus Mulsant, 1850, Species Trimeres Seeuripalpes, p. 492; Crotch, 1874,
Eevision of the Coceinellidae, p. 195; Weise, 1885, Best. -Tab. Europ.
Coleopt., Coeciiielliden, II, ed. 2, p. 5.
Type species. Cocoinella octosignata Gebler, through synonymy with C.
desertorum Gebler, by subsequent designation of Crotch, 1874.
Chilocorini with form broadly oval, moderately convex, upper
surface glabrous. Antenna ten-segmented ; first segment stout,
less than twice as long as wide, constricted at base ; second
almost as wide as first, slightly longer than wide ; third to fifth
mutually similar in form, the fifth shorter ; sixth and seventh
mutually similar, each wider than long ; eighth and ninth
large, eighth almost as long and ninth longer than sixth and
seventh together ; tenth small, more or less embedded in the
oblique apex of the ninth. Mandibles stout, somewhat angulate
on the outer margin. Terminal segment of maxillary palp
rather elongate, apex strongly oblique, lacinia with a row of
three shorter spines along the edge of the row of long setae.
Terminal segment of labial palp cylindro-acuminate, rounded
Fig. 1. Brumus octosignatus (Gebler). In this figure and all subsequent
figures the lettering is as follows : A — antenna, B — mandibles, C —
maxilla, B — ligula, E — legs, F — claw, G — first abdominal sternite,
S — aedeagus, I — sipho, J — sclerotized portions of female genitalia.
CHAPIN : GENERA OF THE CHILOCORINI 237
at apex. Prosteriial lobe moderately wide, truncate at apex. Elytra
with lateral margins very slightly reflexed, finely beaded, epi-
pleura not foveolate for reception of the femoral apices. Abdo-
men with six visible sternites in male, five in female. Meta-
coxal arcs almost complete in male, widely incomplete in female.
Legs slender, femur of leg III a little stouter than the others,
tibial spurs present, tarsal claws unusually long and slender,
without basal tooth or swelling. Male genitalia : median lobe
lanceolate, slightly wider at middle of length, four and one-half
times as long as greatest width ; parameres moderately wide
at base and apex, constricted near middle of length, slightly
longer than the median lobe ; trabes rather stout, as long as the
main parts of the aedeagus ; sipho rather long and slender, of
nearly uniform diameter through most of its length, apex
blunt, with a series of oblique lines or grooves shortly before
apex. Female genitalia : receptaculum seminis stout, cornu
rather small, bent and rounded ; accessory gland large ; sperm
duct shorter than usual, the parts equal in length, the slender
part very slender; infundibulum present, rather longer than
usual ; hemisternites short and blunt, triangular, apices rounded,
styli apical, button-shaped, with two or three long setae each.
This genus was established by Mulsant to include two species,
Coccinella desertoriim Gebler (=C. octosignata Gebler) and
Coccinella sutiiraJis Fabricius. I have removed the second
species to serve as the type of a new genus, Brumoides. Brumus
octosignatus (Gebler) is the only species of Brumus known to
me.
Brumoides new genus
Brumiis, — Leng, 1908, Journ. New York Ent. Soc, 16:34; Casey, 1908,
Can. Ent., 40:409.
Type species. Coccinella suturalis Fabricius, by present designation.
Chilocorini with form oval, moderately convex, upper surface
glabrous. Antenna eight -segmented ; first segment short and
stout, slightly bent; second, as wide as first at base, gradually
tapering to half that width at apex; third, fourth, and fifth
similar and nearly equal, all slightly longer than wide ; sixth
one and one-half times longer than wide, of nearly the same
diameter as fifth ; seventh about equal in length to sixth, apex
obliquely truncate ; eighth small, partly embedded in apex of
seventh. Mandibles not heavily developed, outer margin strongly
convex in apical two-thirds, concave in basal third. Terminal
238
BULLETIN: MUSEUM OP^ COMPARATIVE ZOOLOGY
segment of maxillary palp securiform with apex strongly oblique,
lacinia with row of three short, stout spines along the edge of
the row of longer setae. Labial palp rather long and slender,
the terminal segment cylindro-acuminate, the apex rounded.
Prosternal lobe narrow, truncate. Elytral margin very finely
reflexed, with bead, epipleura not foveolate for the reception of
the femoral apices. Abdomen with six visible sternites in male,
five in female. Metacoxal ares complete. Legs slender, femora
not noticeably inflated, tibiae slender, tibial spurs present, tarsi
rather long, tarsal claws moderately long, slightl^y thickened
at base but Avithout angular basal tooth. Male genitalia : median
lobe triangular, less than twice as long as wide at base ; para-
meres thin, spatula-shaped, hardly at all constricted in basal
half, nearly twice as long as median lobe ; trabes rather slender,
equal in length to the main parts of the aedeagus; sipho mod-
erately slender, of nearly even diameter in basal half, tapering
slightly in apical half, apex twisted and terminating in a short
finger-like process. Female genitalia : receptaculum seminis
much as in Brumus; sperm duct moderately long, the thin
portion not much more slender than the thick portion ; infundi-
bulum present, small ; hemisternites moderately long, parallel-
sided to just before apices where they taper to blunt points.
Fig. 2. Bnnnoifles suturalis (Fabricius).
CHAPIN : GENERA OP THE CHILOCORINI 239
styli small, button-shap(Ml. willi two or three long setae from
each.
The following species have been studied and belong to Brii-
moides -. Coccmella suturalis Fabricius, Exochonius foudrasii
Mulsant, E. hoegei Gorham, E. dcscrtorum Casej^ E. j)arvi-
collis Casey, E. histrio Fall, Exochonius {Brumus) davisi Leng,
E. (B.) ncvadensis Leng, Brumus septentrionis Weise.
HaePASUS Mulsant
Orcus {Harpasus) Mulsant, 1850, Species Trimeres Seeuripalpes, p. 473.
Harpasus, — Crotch, 1874, Revision of the Coccinellidae, p. 190 (as synonym
of Curinus Mulsant).
Type species. Orcus {Ear2)a.svs) pallidilahris Mulsant, by subsequent
designation of Crotch 1874.
The ensuing description has been taken from specimens of
Orcus (Harpasus) zonatus Mulsant, one of the originally in-
cluded species.
Chilocorini with form very broadly oval, nearly circular,
moderately convex, upper surface glabrous. Antenna nine-
segmented; first segment short and stout, narrowed at base;
second as long but not quite as wide as first ; third about as
long as width at apex which is twice width at base ; fourth to
sixth mutually similar in shape, the fourth half as long as third
and slightly wider ; fifth and sixth each a little longer and
wider than the preceding segment ; seventh almost as long as
fifth and sixth combined and still wider; eighth more than twice
as long as seventh, sides nearly parallel ; ninth small, conical.
Mandibles with outer margin feebly concave in basal half, then
sharply convex to apex. Terminal segment of maxillary palp
elongate, the sides nearly parallel, the apex strongly oblique,
lacinia with row of four short, stout spines along the edge of
the row of longer setae. Terminal segment of labial palp
cylindro-acuminate, rounded at apex. Prosternal lobe rather
broad, its lateral margins strongly grooved, the groove then
following the margin of the coxal cavity. Elytral margin nar-
rowly reflexed, finely beaded, epipleura foveolate for the recep-
tion of the femoral apices. Abdomen with six visible sternites
in male, five in female. Metacoxal arcs widely incomplete. Legs
slender, the femora not much enlarged, tibiae simple, tarsi
normal, tarsal claws bent, basal tooth large, quadrate. Male
genitalia : median lobe lanceolate, greatest width in basal third,
four times as long as wide ; parameres spoon-shaped, strongly
240
BULLETIN: MUSEUM OF COMPAEATIVE ZOOLOGY
Fig. 3. Harpasus zonatus Mulsant.
constricted in basal half, longer than the median lobe by its
greatest width ; trabes rather stout, a little longer than the
median lobe; sipho stout, of uniform diameter throughout most
of its length, apex slightly twisted. Female genitalia : recepta-
eulum seminis stout, cornu small; just before the cornu the
body is girdled by a slightly prominent band ; sperm duct long,
the thick portion relatively not as thick as usual, the thin por-
tion extremely thin ; infundibulum present ; hemisternites tri-
angular, rounded at the apices, styli small, button-shaped, with
two or three long setae each.
Harpasus zonatus Mulsant is the only species belonging to
this genus that I have been able to study. From descriptions,
it appears to be quite distinct from the type species, H. palli-
dilahris Mulsant and an examination of that species may show
that the two are not congeneric.
ZaGREUS Mulsant
ExocJiomiis (Zagreus) Mulsaut, 1850, Species Trimeres Securipalpes, p. 488.
Type species. Exochomus (Zagreus) bimaculos^^■s Mulsant, by present
designation.
CHAPIN : GENERA OF THE CHILOCORINI
241
Chiloeorini with form very broadly oval to nearly circular,
moderately convex, upper surface glabrous. Antenna eight-
segmented ; first segment stout, slightly bent, half again as long
as wide ; second as stout at base as first, tapering to apex, as long
as wide; third and fourth nearly equal in length, each wider
at apex than at base, fourth wider than third; fifth as long as
fourth but wider; sixth as long as wide; seventh about as long
as first, with apex strongly oblique ; eighth short, conical, em-
bedded in seventh. Mandible moderately stout, its outer margin
broadly curved. Terminal segment of maxillary palp subsecuri-
form with apex strongly obliciue, lacinia with oblique row of
seven slender spines on outer face. Terminal segment of labial
palp cylindro-acuminate, a little more than twice as long as
wide, truncate at apex. Prosternal lobe narrow. Abdomen with
six visible sternites in male (in a related species), five in female.
Fig. 4. Zagreus himaculosus Mulsant.
Metacoxal arcs incomplete, ending about half way to base of
segment, posterior margin of fifth sternite broadly rounded in
female. Legs with tibial spurs, femora moderately stout, tibiae
slender, shallowly excavate near apices, the margin of excavation
242 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
edged with a row of slender bristles. Tarsal claws wdth stout,
subquadrate basal tooth on each. Male genitalia not studied.
Female genitalia : receptaculum seminis with nodulus distinct
from body, but not prominent, eornu curved, rounded at apex,
without appendix ; accessory gland about twice as long as
receptaculum ; sperm duct long, the thick portion much thicker
than the very thin portion ; infundibulum present ; hemister-
nites short and stout, triangular, about twice as long as wade
at base, apices broadly rounded, styli short, button-shaped, with
long setae.
The following species have been determined as belonging to
the genus Zagreus. Exochomus (Zagreus) himaculosus Mulsant,
E. (Z.) cinctipennis ]\Iulsant, Exocliomus jordani Mulsant, E.
adelae Crotch, E. histillafus Weise, E. decempunctatus Weise,
E. guttatus Weise, E. suhcoendeus AVeise, and E. ritchiei Si-
card.
AxiON Mulsant
Exocliomus {Axion) Mulsant, 1850, Species Trimeres Securipalpes, p. 477.
Axion, — Crotch, 1874, Eevison of the Coccinellidae, p. 191; Gorham, 1892,
Biol. Centr.-Amer., Ins., Coleop., 7:176; Casey, 1899, Journ. New York
Ent. Soc, 7:105; Leng, 1908, Journ. New York Ent. Soc, 16:34.
Type apecies. Coccinella tripustulata DeGeer, by subsequent designation of
Crotch, 1874.
Chilocorini with form subcircular, size large to very large,
strongly convex, upper surface glabrous. Antenna ten-seg-
mented; first segment almost straight, twice as long as narrow
median portion ; second barrel-shaped, very little longer than
wide ; third through sixth similar in shape, narrow at base and
widening from base to apex ; third as long as wide, fourth and
fifth each shorter than the preceding segment, sixth slightly
.longer than fifth ; seventh through tenth forming a compact
oval club, seventh and eighth equal in length, ninth longer
with the apex oblique, tenth small, deeply embedded in ninth,
conical. Mandible very stout, subangulate near middle of outer
margin. Terminal segment of maxillary palp wider toward apex
than at base, apex strongly oblique, lacinia with an irregular row
of seven or eight stout spines on the outer side. Terminal
segment of labial palp cylindro-acuminate, two and one-half
times longer than wide. Prosternal lobe narrow, truncate at
apex. Pronotum without fine marginal line across base. Elytra
with lateral margins slightly reflexed, epipleura with very small
foveae for reception of the femoral apices. Abdomen M-ith six
CHAPIN: GENERA OF THE CHILOCORINI
243
Fig. 5. Axion tripustulatum (DeGeer).
visible sternites in male, five in female, fifth sternite of male
broadly emarginate. Metacoxal arcs broadly incomplete. Femora
not noticeably stout, tibiae moderate, feebly excavated at apices
for reception of first segments of tarsi, tarsal claw with very
strong quadrate, plate-like tooth in basal half. Male genitalia:
median lobe long, slender, slightly asymmetrical in apical third ;
parameres noticeably longer than median lobe, paddle-shaped
and constricted at basal third; trabes very slender, about as
long as a paramere; sipho slender, siphonal capsule well de-
veloped, apex of sipho blunt, slightly twisted. Female geni-
talia: receptaculum seminis much as in Curinus; sperm duct
proportionately shorter than in Curinus and differing in that
the thick portion is longer than the thin portion ; inf undibulum
present, inverted Y-shaped; hemisternites parallel nearly to
apices, then rounded triangular, styli button-shaped, each with
three or four long setae.
This purely North American genus is in need of revision.
Eight trivial names have been associated with the generic name
^Axion ; four of these were considered as valid specific names in
the world catalog.
244
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
CUEINUS Mulsant
Orciis (Ctirinus) Mulsant, 1850, Species Trinieres Securipalpes, p. 472.
Curinus, — Crotch, 1874, Eevision of the Coccinellidae, p. 190 ; Gorham,
1892, Biol. Centr.-Anier., Ins., Coleop., 7:176; Koischefsky, 1932,
Coleopt. Catalogus (Jimk), pars 120, p. 252.
Type species. Orcus (Curinus) coeruleus Mulsant, monobasic.
Chilocorini with form nearly circular, strongly convex, upper
surface glabrous. Antenna ten-segmented ; first segment stout,
slightly curved, slightly longer than wide ; second barrel-shaped,
one and one-half times longer than wide and nearly as wide as
the first; third half as wide at base as second, nearly twice as
wide at apex as at base ; fourth nearly equilateral but slightly
wider apically; fifth through tenth forming a compact fusiform
club ; fifth slightly wider than long ; sixth and seventh almost
equal in length, each wider at apex than at base; eighth longer
and wider than seventh, maximum width of club at apex of
eighth; ninth about twice as long as eighth, sides gradually
tapering to apex which is oblique ; tenth short and conical,
Fig. 6. Curinus coeruleus Mulsant.
CHAPIN : GENERA OF THE CHILOCORINI 245
slightly Avider than long. Mandible stout, subangulate at mid-
dle of outer margin. Terminal segment of maxillary palp with
lateral margins subparallel and apical margin strongly oblique,
laciuia with irregular row of five stout spines on outer face.
Terminal segment of labial palj) elongate cylindrical but with
outer margin slightly curved, about three times longer than
wide. Prosternal lobe flat, moderately broad. Elytral margins
slightly reflexed, with fine marginal bead, epipleura foveolate
for the reception of the femoral apices. Abdomen with six visible
sternites in male, five in female, apical sternite of male emar-
ginate or notched, of female simple and rounded. Metacoxal
arcs on first sternite incomplete, failing to reach base of ster-
nite. Legs with femora moderately stout, tibiae near apices
shallowly excavate, the margins of the excavation edged with
rows of slender bristles, tarsal claws stout with subquadrate
basal tooth. ]\Iale genitalia : median lobe slightly asymmetrical ;
parameres longer than the median lobe, constricted near basal
third ; trabes slender, about as long as paramere ; sipho mod-
erately stout, of even diameter nearly to apex, where the tube
undergoes a quarter turn, the apex truncate. Female genitalia :
receptaculum seminis more or less globular, basal portion with-
out well-defined nodulus or ramus, cornu short, small, and bent ;
accessory gland long and slender ; sperm duct very long, the
thin part about twice as long as the thick part ; infundibulum
present, inverted Y-shaped; hemisternites elongate triangular,
styli button-shaped, with three or four long setae from each.
With the removal of Curinus peleus (Mulsant) to Exochomus,
and the reestablishment of Harpasus, with the transfer of its
species from Curinus to it, Curinus is left with the type species
and one very doubtfully included species from Chile, Curinus
ruizi Brethes. Brethes figures the antenna as having eleven
segments, which if true, makes that species unique in the tribe.
I think it more likely that there is an error in the drawing and
that the species belongs in Harpasus.
ArAWAXA Leng
Exochomus {Arawaiia) Leng, 1908, Journ. New York Ent. Soc, 16:34, 38;
Casey, 1908, Can. Ent., 40:409.
Type species. Exochomus arisoniciis Casey, by original designation of Leng.
Chilocorini with form broadly oval, strongly convex, upper
surface glabrous. Antenna ten-segmented; first segment short
and stout, bent at almost a right angle; second equally stout,
246
BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
as long as wide ; third strongh' wedge-shaped, much narrower
at base than at apex, about as long as width at apex ; fourth to
seventh similar in shape ; fourth and fifth nearly equal, sixth
and seventh each slightly wider and longer than the preceding
segment ; eighth one-half longer but hardly wider than seventh,
ninth twice as long as eighth and as wide, its apex strongly
oblique; tenth small, embedded in apex of ninth. Mandibles
stout, somewhat angulate at middle of outer margin. Terminal
segment of maxillary palp twice as long as wide, apex strongly
oblique, lacinia with irregular row of six stout spines along the
edge of a row of long setae. Terminal segment of labial palp
slender cylindro-acuminate, rounded at apex. Prosternal lobe
moderately broad, truncate. Elytral margin not reflexed, very
finely beaded, epipleura foveolate for the reception of the
femoral apices. Abdomen Avith five visible sternites in both
sexes. Metacoxal arcs nearly complete. Legs moderately stout,
femur of leg I a little stouter than the others, tibia I with the
outer margin expanded into a thin keel, tibiae II and III with
spurs, tarsal claws stout, strongly hooked, with triangular tooth
in basal half. Male genitalia : median lobe lanceolate, four
times longer than its greatest width at basal third ; parameres
noticeably longer than median lobe, strongly constricted near
Fig. 7. Araicana arizonica (Casey).
CHAPIN : GENERA OF THE CHILOCORINI 247
base, each with a finger-like process at its apex; trabes slender,
one and one-half times as long as the median lobe ; sipho long
in proportion to the aedeagus, of uniform diameter throughout
most of its length, siphonal capsule small, apical portion of
sipho slightly expanded and twisted. Female genitalia : reeep-
taculum seminis much as in the genus Curinus; sperm duct
with the thicker portion considerably longer than the extremely
thin portion ; inf undibulum present, inverted Y-shaped ; hemi-
sternites about twice as long as wide, generally tapering to the
blunt apices, styli apical, button-shaped, each with two or three
long setae.
In addition to the type species, I am including two other
species in the genus Arawmia, Exochomus scapularis Gorham
from Central America and Exochomus cuhensis Dimmock from
Cuba, this last with some hesitation. E. cuhensis is included
because, while the fore tibiae do not have the thin keel but are
simply broadly curved on their outer margin, the male genitalia
are quite similar to those of the other two species and do have
the finger-like processes at the apices of the parameres. I con-
sider the structure of the genitalia of more importance than
that of the fore tibiae. All three species are similar in general
appearance.
Exochomus Redtenbache
Exochomus Redtenbacher, 1843, Tentauieii dispositiouis Coleopterorum
pseudotrimerorum, p. 11 ; Eeprint of aljove in Gerniar, 1844, Zeitseh.
f. Ent., .5:121; Mulsaiit, 1850, Species Trimeres Securipalpes, p. 476;
Crotch, 1873, Trans. Amer. Ent. Soc, 4:376; 1874, Revision of the
Coecinellidae, p. 192; Barovsky, 1922, Ann. Mus. Zool. Russie, 23:293,
figs, la, lb, Ic; Korschefsky, 1932, Coleopt. Catalogus (Junk), Coe-
cinellidae, Pars 120, p. 252.
In 1922, Barovsky proposed a division of ExocJiomus into
three subgenera, based on the structure of the tarsal claws, as
follows :
I. Unguiculi dente valido basali instructi
Subg. Exochomus, s.str.
II. Unguiculi dente indisiincto basali instructi
Subg. Parexochomus new
III. Unguiculi dente subacutangulo post medium instructi . .
Subg. Anexochomus new
'o
248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
ExocHOMUs (ExocHOMus) Redteiibacher
ExocJiomus (Exochomus) — Barovsky, 1922, Ann. Mus. Zool. Eussie, 23:291,
fig. la.
Type species. Coecinella 4-pustulata Limie (quadrip-ustulata auct.), sub-
sequent designation of Korschefsky, 1932. Two previous type designa-
tions for this genus had been made, both by Crotch and both invalid,
since the species designated were not among those originally included.
Chilocorini with form broadly oval to almost circular, mod-
erately convex, upper surface glabrous or pubescent. Antenna
ten-segmented ; first segment slightly curved, a little less than
twice as long as wide ; second barrel-shaped, as long as wide and
same diameter as first ; third obconical, half as long and half
as wide as second; fourth, fifth, and sixth nearly equal, each
slightly wider apically than at base; seventh similar in shape
to sixth but about half as long as wide, and forming with the
eighth, ninth, and tenth a slender fusiform club ; eighth almost
as long as wide, at base equal in width to apex of seventh,
greatest width of club at apex of eighth ; ninth as long as
eighth, conical with apex strongly oblique; tenth small, conical
deeply embedded in ninth. Mandible heavy, angulate at middle
of outer face. Terminal segment of maxillary palp subsecuri-
form with apex strongly oblique, lacinia with row of five long,
slender (not short, stout) spines on outer face. Terminal seg-
ment of labial palp cylindrical, twice as long as wide. Pro-
sternal lobe narrow, truncate at apex, the anterior coxae almost
contiguous. Pronotum very finely margined across base, lateral
margins slightly reflexed. Elytral margin strongly beaded,
epipleura not foveolate for the reception of the femoral apices.
Abdomen with six visible sternites in male, five in female. Meta-
eoxal arcs complete or virtually so. Legs with moderately stout
femora, tibiae slender, tarsal claws strong, with subquadrate
basal tooth on each. Male genitalia : median lobe long, slender,
parallel in basal two-thirds, slightly asymmetrical in apical
third; parameres about one-eighth longer than the median lobe,
slender, somewhat constricted in basal half, spoon-shaped in
apical half; trabes slender, almost as long as main body of the
aedeagus; sipho moderately slender, of even diameter through-
out most of its length, siphonal capsule rather heavy, apex of
sipho twisted. Female genitalia: receptaculum seminis a stout
body without clearly defined nodulus or ramus, cornu more
slender than body, stronglj- curved; accessory gland slightly
longer than receptaculum ; sperm duct long, the parts about
CHAPIN : GENERA OF THE CHILOCORINI
249
Fig. 8. Exochomus quadripustulatus (Linne).
equal in length, the thick part noticeably thicker than the thin
part ; infundibulum present, tapering slightly from a broad
Imse and slightly curved; hemisternites elongate, subparallel,
about three times as long as wide at base, apices acutely
rounded, styli button-shaped, each with two or three long setae.
The following species have been verified as belonging to
Exochomus sensii stricto: Coccinella quadripustulata Linne, C.
aethiops Bland, C. flavipcs Thunberg, C. marginipennis Leconte,
C. nigripcnnis Erichson, Orcus pelews Mulsant, Exochomus
childreni Mulsant, E. calif or nicus Casey, E. contristatus Mul-
sant, E. jamaicensis Sicard, E. latiusculus Casey, E. lituratus
Gorham, E. metaUicus Korchefsky, E. mormonicus Casey, and
E. uropygidialis Mulsant.
Eighteen species, originally described in Exochomus, have
been removed from this genus. Two are here placed in Arawmia,
seven in Zagreus, and seven in Brumoides. Two species, Exo-
chomus tricoJoratus Gorham and E. championi Gorham are re-
moved from the Chilocorini and should be placed in the Coc-
cinellini near Cycloneda.
250 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
ExocHOMus (Parexochomus) Barovsky
Exocliomus {Parexochomus) Barovsky, 1922, Ann. Mns. Zool. Eussie, 23:292,
fig. lb.
Type species. Exochomus puhcscens Kiister, by present designation.
I have been unable to examine any specimens conforming to
Barovsky 's description. Barovsky assigns the following palae-
arctic species to this subgenus : E. (P.) puhcscens Kiister, E. (P.)
anchorifcr Allard, E. (P.) semenovi Weise, and E. (P.) kirgi-
zorum Barovsky.
Exochomus (Anexochomus) Barovsky
Exochomus (Anexocliomus) Barovsky, 1922, Ann. Mus. Zool. Eussie, 23:292,
fig. Ic.
Type species. Exocliomus undul-atus Weise, by present designation.
A single female specimen from Pusan, Korea, in the collec-
tion of the Museum of Comparative Zoology appears to be a
color variant of E. (A.) mongol Barovsky. It satisfies the orig-
inal description in every respect except that the subapical spot
on the elytron is missing. Its tarsal claws conform to figure Ic
of the original description. A study of the gross anatomy shows
no character of importance that would warrant a change of
status for Anexochomus. Barovsky assigns the following species
to this subgenus : E. (A.) un(lulatusWehe,E. (A.) kiritshenkoi
Barovsky, and E. (A.) mongol Barovsky, all from the palaeare-
tic region.
-e^
ClAJ)IS Mulsant
Exochomus (Clanis) Mulsant, 1850, Species Trimeres Securipalpes, p. 479
(name preoccupied).
Exochomus (Cladis) Mulsant, 1850, idem, Appendix, p. 1033 (new name
for Clanis Mulsant not Hiibuer 1819).
Cladis, — Crotch, 1874, Eevision of the Coccinellidae, p. 192; Weise, 1883,
Wien. Ent. Zeitseh., 2:67; Gorham, 1894, Biol. Centr.-Amer., Ins.,
Coleopt., 7:179.
Type species. Coccinella nitidula Fabricius, through synonymy with Ex-
ochomus (Clanis) uva Mulsant, by subsequent designation of Crotch,
1874.
Chilocorini with form broadly oval, moderately convex, upper
surface glabrous. Antenna ten-segmented ; first segment slightly
bent, about one and one-half times longer than wide; second
equal in width to first, almost equilateral; third as long as sec-
ond but only half as wide ; fourth nearly as wide as third.
CHAPIN : GENERA OF THE CHILOCORINI
251
equilateral; fifth to eighth rather similar, each wider and
shorter than the preceding ; ninth large, equal to the seventh
and eighth conihined; tenth small, more or less embedded in
the oblique apex of the ninth. Mandible with the outer margin
moderately curved, with no unusual modifications. Terminal
segment of maxillary palp broad and short, apex stronglj-
oblique, lacinia with two or three stout spines along the edge
of the row of long setae. Terminal segment of labial palp
cylindro-acuminate, rounded at apex. Prosternal lobe narrow,
truncate at apex. Elytra with lateral margin not reflexed, epi-
pleura not foveolate for reception of the femoral apices. Abdo-
men with six visible sternites in male, five in female. Metacoxal
Fig. 9. CJadis nitidula (Fabrit-ius).
ares widely incomplete. Legs with femora not unusually stout,
tibiae simple, rather slender, shallowly excavate near apices,
tarsal claws strongly curved, each with a large, quadrate basal
tooth. Male genitalia : median lobe acutely triangular, two
and one-half times longer than wide at base; parameres very
252 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
broad in apical half, strongly constricted in basal half, notice-
ably longer than the median lobe ; trabes moderately stont,
curved, about as long as a paramere from basal attachment to
apex ; sipho moderately stout, of nearly uniform diameter
throughout, apex somewhat twisted, blunt. Female genitalia :
receptaculum seminis much as in Curinus or ExocJwmus, acces-
sor}^ gland moderate in size ; sperm duct moderately long, the
thick portion slightly longer than the thin portion; infundi-
bulum present, inverted Y-shaped; hemisternites moderately
heavy, elongate triangular, apices rounded, styli minute, button-
shaped, each with two or three long setae.
Mulsant segregated two supposedly new species under the
name Clanis, which name had been previously used by Hiibner.
At present it is considered that Mulsant 's "species" are not
distinct and are synonyms of the much earlier Fabrician species,
which occurs on various of the West Indian islands. It has
also been reported from Buenos Aires.
PrIASUS :\lulsant
Orcus (PriasHs) Mulsant, 18.50, Species Trimeres Securipalpes, j). 4(37;
Crotch, 1874, Eevision of the Coceinellidae, p. 188.
Type species. Coccinclla bilunulata Boisduval, by suljscqueiit designation
of Crotch, 1874.
Chilocorini with form nearly circular, strongly convex, upper
surface glabrous. Antenna nine-segmented ; first segment slightly
bent, strongly constricted at middle, apically produced laterally
in a small, rounded lobe ; second stout, barrel-shaped, nearly
equilateral ; third, fourth, and fifth trapezoidal, decreasing in
length and slightly increasing in width at apices; sixth longer
than fifth and slightly wider ; seventh and eighth each longer
than the preceding, the eighth slightly produced laterally at
apex ; ninth longer than eighth, subcylindrical, apex slightly
oblique. Mandible stout, the outer margin not strongly curved,
apex unusually blunt. Terminal segment of maxillary palp
with outer margin convex, inner margin concave, the sides
nearly parallel, apex oblique ; lacinia with regular row of four
stout spines on outer face. Terminal segment of labial palp
cylindro-acuminate, truncate at apex. Prosternal lobe rather
narrow, slightly dilated behind, apex rounded. Pronotum with
a strong marginal line. Elytra with lateral margins only
slightly reflexed, epipleura not foveolate for the reception of
the femoral apices. Abdomen with six visible sternites in male,
CHAPIN : GENERA OF THE CHILOCORINI
253
five in female; metaeoxal arcs incomplete, running almost into
the posterior margin of the sternite in female, curving toward
anterior margin for a short distance in male. Legs stout, an-
terior femora about twice as long as wide, tibiae simple, slightly
expanded apically, tarsal claws short and stout, swollen at
base, without basal tooth. Male genitalia : median lobe elon-
gate triangular ; parameres straight, parallel-sided, rounded at
apices, noticeably longer than median lobe ; trabes as long as a
paramere with its basal attachment; sipho slender, of even
diameter except near the siphonal capsule where it is somewhat
thicker, apex twisted and with a short, sharp apical process.
Fig. 10. Priasus hihniuJatus (Boisduval).
Female genitalia : receptaculum seminis stout bulbous, nodulus
hardly differentiated, ramus absent, cornu short, stout, and
rounded ; accessory gland about four times longer than the
receptaculum; sperm duct rather short, the thick portion equal
in length to the thin portion and about three-fourths as long
as the accessory gland; infundibulum present; hemisternites
elongate, the sides almost parallel to near apices Avhich are
abruptly narrowed and rounded, styli button-shaped, hardly
prominent, each with two or three long setae.
254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Mulsant placed three species, all Australian, under this name.
I have been able to study all three and have found that, Avhile
the type species has toothless claAvs, the other two have claws
which are equipped with a basal tooth on each. In one case, Coc-
cinella aiistralasiac Boisduval, the tooth is large and conspicu-
ous, in the other, C. nummularis Boisduval, the tooth is much
smaller and is somewhat transparent and difficult to see. These
two species have been referred to the following genus.
ParAPEIASUS new genus
Type species. Coccinella australasiae Boisduval, by present designation.
Chilocorini with form nearly circular, strongly convex, upper
surface glabrous. Antenna nine-segmented; first segment a
little longer than second, bent, constricted at basal third,
apically widened : second barrel-shaped, wider at base than at
apex, nearly equilateral ; third to seventh similar in shape, third
longer than wide, fourth shorter than third, fifth shorter than
fourth or sixth, seventh as long as sixth but slightly wider;
eighth longer than wide ; ninth longer than eighth and tapering
to a blunt point. Mandible stout, apex short and acute, outer
margin subangulate at middle of length. Terminal segment of
maxillary palp subsecurif orm, apex oblique ; lacinia with a
patch of three or four short and stout bristles, in addition to
the row of long setae. Terminal segment of labial palp cylindro-
acuminate, rounded at apex. Prosternal lobe moderate, convex,
margined laterally, the bead prolonged part way around the
coxal cavity. Elytral margin rather strongly reflexed, without
marginal bead ; epipleura not foveolate for the reception of
the femoral apices. Abdomen with six visible sternites in male,
five in female, the fifth sternite broadly truncate in male,
broadly rounded in female. Metacoxal arcs broadly open, the
line running from the intercoxal process out and back almost
to the posterior margin of the sternite, then curving forward
for a short distance. Legs moderately stout, tibiae slightly
widened toward apices, tibia I with a row of small denticles
along outer margin, tarsal claw stout, with a subquadrate, trans-
lucent tooth in basal half. Male genitalia : median lobe about
as long as the parameres, slender, elongate triangular but with
the lateral margins nearly parallel in basal two-thirds, apical
third tapering to a slender blunt apex ; parameres slender, not
noticeably constricted, apices broadly rounded ; trabes slender,
slightly curved, a little shorter than the median lobe ; sipho
CHAPIN: GENERA OF THE CHILOCORINI
255
Fig. 11. Parapriasus australasiae (Boisduval).
rather similar to that of Priasus. Female genitalia : receptacn-
lum seminis similar in shape to that of Priasus but less stout,
especially the cornu; accessory gland about twice as long as
receptaculum ; sperm duct with thick portion a little longer
than the very thin portion ; infundibulum a simple, bar-like
structure; hemisternites long, slender, and tapering to blunt
apices, styli terminal, button shaped, each giving rise to two or
three long setae.
Two species have been seen which belong to this genus, Coc-
cinella australasiae Boisduval and C. nummularis Boisduval,
both from Australia. In the latter the tooth on the tarsal claw-
is less well developed and less easily seen.
EnDOCHILUS Weise
EndoChilus Weise, 1898, Deutsch. But. Zeitsch., p. 119, pi. 1, fig. 10, 1910,
Verb. Naturf. Ver. Briinn, 48:52; Sicard, 1929, Ann. Mag. Nat. Hist.,
(10) 4:518.
256 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Type species. Endochilus cavifrons Weise, by subsequent designation of
Korschefsky, 1932.
As the type species is not available to me, I have prepared
the generic description and illustrations from specimens of E.
plagiafiis Sicard.
Chilocorini with form nearly circular, not strongly convex,
margins broadly but not abruptly reflexed, upper surface wholly
or partly set wdth fine pubescence. Antenna eight-segmented ;
first segment constricted at middle, twice as long as width at
constriction ; second elongate globose, slightly shorter and wider
than first; third, fourth, and fifth wedge-shaped, each slightly
shorter and wider than the preceding, third and fourth together
about as long as the second ; sixth longer and wider than fourth
and fifth together ; seventh slightly longer than sixth but hardly
wider, the margins nearly parallel ; eighth conical, as long as
sixth, its apex rounded. Mandible with outer margin evenly
curved. Maxillary palp very stout, the terminal segment
elongate-conical, four times as long as the penultimate segment ;
lacinia with a row of four short, stout spines on outer face.
Labial palp stout, the terminal segment conical, a little longer
than the penultimate segment. Prosternal lobe broad, flat, not
margined. Elytral margin subexplanate, slightly reflexed, with-
out bead ; epipleura weakly f oveolate for the reception of the
femoral apices. Abdomen with five visible sternites in either
sex, in both the fifth sternite is broadly rounded. Metacoxal
arcs run from the sides of the rather broad intercoxal process
almost straight toward the posterior angles of the sternite, where
the lines are slightly recurved. Legs slender, femur I a little
stouter than the others, tibiae simple, tarsal claws gently and
evenly curved, each with a small, obtuse, triangular tooth at
base. Male genitalia : median lobe lanceolate, its greatest width
at middle of length, apex acuminate ; parameres slightly longer
than the median lobe, their bases concave and partly embracing
the lobe ; trabes slender, about as long as median lobe, gradu-
ally broadened from apical third to apex; sipho long and slen-
der, of nearly uniform diameter throughout most of its length,
siphonal capsule small, apical portion slightly enlarged just back
of the acuminate apex. Female genitalia : receptaculum seminis
bent sausage-shaped, strongly constricted at basal fourth, nodu-
lus not prominent, cornn stout, without apical appendix, acces-
sory gland small, globular ; sperm duct short, the thick portion
shorter than the length of the receptaculum, the thin portion
CHAPIN : GENERA OF THE CHILOCORINI
257
Fig. 12. Endochilus -plagiatus Sieard.
shorter than the thick portion ; infnndibulum absent ; hemister-
nites elongate, sides nearly parallel, about five times longer than
average width, apices rounded, styli absent, represented by two
long, apically placed, setae.
A genus of a few, below average size, species, mostly from
West Africa south of the Gulf of Guinea. The genus is im-
mediately distinguished from other chilocorine genera by the
peculiar conformation of the head. The genal lobes and the
elypeus form the transverse frontal margin of the head, with a
forward extension below the eyes equal to half the long diam-
eter of the eye. This margin passes below the eye and joins
the temporal areas of the head. Thus there is no invasion of the
eye itself.
HaLMUS Mulsant
Orcus (Ealmus) Mulsant, 1850, Species Trimeres Securipalpes, p. 471.
Ealmus, — Weise, 1923, Arkiv Zool., 15(12): 134.
Type species. Coccinella chalyhea Boisduval, the genus being originally
monobasic.
Chilocorini with form nearly circular, moderately convex,
upper surface glabrous. Antenna seven-segmented; first seg-
ment bent, constricted in basal half, apically produced laterally
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in a conical lobe ; second nearly equilateral, barrel-shaped ; third
slender, twice as long as wide, the apex oblique ; fourth and fifth
similar in shape and proportions, the fifth half again as large
as the fourth ; sixth as wide at base as fifth, beyond middle it
tapers to half its basal diameter ; seventh conical, slightly longer
than its diameter at base. Mandibles rather delicate, the outer
margins strongly curved, apices acute ; just distad of the median
notch the inner margin is cut away so that it appears to be
tridentate. Terminal segment of maxillary palp broadly oval,
the apical margin oblique ; lacinia with an irregular row of
six or seven stout spines on outer face. Terminal segment of
labial palp short, stout, and conical. Prosternal lobe moderately
broad, truncate at apex. Pronotum without basal marginal line.
Elytra with lateral margins reflexed, epipleura not foveolate
for the reception of the femoral apices. Abdomen similar in
both sexes, with six visible sternites, the fifth sternite in both
sexes truncate. Metacoxal arcs incomplete, running parallel and
close to the posterior margin of the sternite. Legs with femora
not noticeably stout, tibiae simple, tarsal claws strongly curved,
with broad, quadrate, plate-like tooth in basal half. Male
genitalia : median lobe long, narrow, parallel-sided nearly to
apex where it is rounded, as seen in dorso ventral ^dew ; in
Fig. 1.3. Eahnus chalybetis (Boisduval).
CHAPIN : GENERA OF THE CHILOCORINI 259
lateral view it is somewhat slipper-shaped, the apex or "toe"
turned up and with a triangular projection in the dorsal margin
at apical third; parameres very slender, slightly shorter than the
median lobe ; trabes a little longer than body of aedeagus ; sipho
slender, of even diameter throughout, apex somewhat twisted
and with a short process. Female genitalia : receptaculum
seminis curved, sausage-shaped, nodulus and ramus not defined,
cornu separated from the main body by a slight constriction and
broadly rounded apically, accessory- gland longer than the
receptaculum ; sperm duct comparatively short, the thick sec-
tion shorter than the thin section but of almost the same diameter
except at the junction point ; infundibulum present ; hemister-
uites short, and broadly triangular, styli cylindrical, nearly
twice as long as the diameter at base, each with two long setae.
Halnius chalybeus is a median sized species, 3.0-3.7 mm long,
clear blue to blackish-blue above except that in the male the head
and flanks of the pronotum are yellowish-white. This is one of
the few genera that shows a sexual difference in coloration. The
species is further distinguished by the unusual form of the
genitalia of both sexes.
Weise was apparently the first to raise Mulsant's subgenus
Halmus to full generic status. He did this because of the un-
usual shape of the genae. Here the genal lobe, which extends
out over the antennal insertion and on into the eye, is abruptly
narrowed at the inner margin of the eye and is continued into
the eye as a fine line. This character, apparently unique among
the Chilocorini, when taken together with the other character-
istics which deviate from the norm, indicates that the genus
stands in a rather isolated position in the tribe.
For the present, I am referring Orcus ovalis Blackburn to this
genus. It agrees with Hahnus chalyheus in its antennae, mandi-
bles, ligula with labial palps, legs, metacoxal arcs, visible sixth
sternite in the female, and receptaculum seminis. It differs in
the broadly securiform maxillary palps, the male genitalia
(which are simple as in most Chilocorini), and most importantly
in the form of the clypeus and genal lobes. The clypeus is not
more or less deeply emarginate without a marginal bead, as in
most other Chilocorini, but is straight across the front, without
a trace of an emargination, and with a strong marginal bead.
The genal lobes are as in most other Chilocorini, not suddenly
contracted at the inner margin of the eye. "When more of the
Australasian "Orcii.^" are thoroughly studied, it may seem
best to establish 0. ovalis as the type of a distinct genus.
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OrCUS Mulsant
Orcus Mulsaut, 1850, Species Trimeres Seeuripalpes, p. 465; Crotch, 1874,
Eevision of the Coccinelliclae, p. 188 ; Weise, 1923, Arkiv Zool.,
15(12) :133.
Type species. Orcus jantliinus Mulsant, by subsequent designation of
Crotch, 1874.
Cliilocorini with form nearly circular, moderately convex,
upper surface glabrous. Antenna eight-segmented ; first seg-
ment slightly curved, less than twice as long as wide, constricted
at basal third, apex with a small, rounded lateral lobe ; second
barrel-shaped, a little longer than wide; third obconical, much
narrower at base than apex of second, less than twice as long
as wide ; fourth through seventh mutually similar in shape but
becoming progressively longer and wider ; seventh somewhat
oblique at apex ; eighth nearly twice as long as wide, rounded
acuminate. ]\Iandible heavy, subangulate at middle of outer
margin. Terminal segment of maxillary palp stout, subsecuri-
form, the apex strongly oblique; lacinia with three or four
short, stout spines on outer face. Terminal segment of labial
palp cylindrical, slightly tapering, twice as long as wide at
Fig. 14. Orcus janthinus Mulsant.
CHAPIN: GENERA OF THE CHILOCORINI 261
base. Prosternal lobe moderately broad, truncate at apex. Pro-
notuni finely margined across median third of base. Elytra
with lateral margins slightly reflexed, epipleura weakly foveo-
late for the reception of the femoral apices. Abdomen similar
in both sexes, each with five visible sternites, the fifth evenly
rounded. Metacoxal arcs broadly open, the line running par-
allel to the posterior margin of the sternite, nearly to the lateral
margin. Legs w-ith femora moderately stout ; tibiae excavate in
apical half or more for the reception of the tarsi ; tarsal claws
strongly curved, each with a large quadrate, plate-like tooth in
basal half. Male genitalia : median lobe long, lanceolate, not
visibly asymmetrical apically ; parameres slender, straight, not
broadened in apical half, slightly longer than median lobe;
trabes longer than median lobe ; sipho rather slender, terminat-
ing in a slender, acute spine. Female genitalia : receptaculum
seminis a stout body with nodulus indicated by a slight prom-
inence, ramus not defined, cornu strongly curved and broadly
rounded at apex, accessory gland much longer than recepta-
culum; sperm duct with parts nearly equal in length, the thick
section about three times the diameter of the thin section; in-
fundibulum present; hemisternites slender, three times longer
than wide at base, styli button-shaped, each with two or three
long setae.
Orcus janthinus is a large species, 5.0-5.7 mm, uniformly
steel-blue above with the exception of the reddish-yellow labrum.
Beneath, the meso- and metathoraces and legs are castaneous,
the antennae and abdomen reddish-yellow. The series studied
is from Bogor, Java, taken in February, 1954, by Mr. 0. D.
Deputy. I have seen no other species of this genus.
AnISORCUS Crotch
Anisorcus Crotch, 1874, Eevision of the Coccinellidae, p. 190; Weise, 1902,
Termes. Fiizetek, 25:508.
Type species. Anisorcus fryi Crotch, liy original designation.
Chilocorini with form broadly oval, moderately convex, upper
surface glabrous. Antenna seven-segmented ; first segment short,
bent, hardly constricted; second barrel-shaped, of same diam-
eter and as long as the first; third obconical, shorter than and
much narrower than second; fourth and fifth mutually similar
in shape, the fifth slightly wider and longer than the fourth
but shorter than the third; sixth similar in shape to the second
and slightly longer; seventh nearly twice as long but not quite
262
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as wide as sixth, cyliiidro-aeuminate, the apex rounded. Mandi-
bles moderately stout, the inner margin of the right mandible
with a small tooth just in front of the notch, outer margins
broadly rounded, apices acute. Terminal segment of maxillary
palp cylindrical, tapering slightly toward apex which is oblique
but not strongly so; lacinia with a patch of nine or ten stout
spines on outer side. Terminal segment of labial palp cylindro-
acuminate, twice as long as wide at base. Prosternal lobe mod-
erately broad, truncate at apex. Pronotum without fine marginal
line across base. Elytra with lateral margins slightly reflexed,
very finely beaded ; epipleura shallowly foveolate for the recep-
tion of the femoral apices. Abdomen alike in both sexes, each
with six visible sternites, the fifth sternite in each is truncate.
Metacoxal arcs nearly straight from intercoxal process nearly
to posterior margin of sternite where they turn abruptly and
follow margin nearly to lateral margin of sternite. Legs with
femora not noticeably stout ; the tibiae rather broad and short ;
tarsal claws each with a strong tooth in basal half. Male geni-
talia : median lobe elongate triangular, not noticeably asym-
metrical apically ; parameres very broad, not constricted in
Fig. 15. Anisorcus fryi Crotch.
CHAPIN : GENERA OF THE CHILOCORINI 263
basal half, one-third longer than median lobe ; trabes slender,
as long as main portion of the aedeagns ; siplio rather slender,
the siphonal capsule poorly developed, apex of sipho slightly
recurved and covered by a membraneous hood. Female genitalia :
receptaculum seminis with a very stout body, nodulus and ramus
not defined, cornu very short and ends in a flat, rounded ap-
pendix, accessory gland only a little longer than receptaculum ;
sperm duct in two parts, the thick section is of even diameter
throughout and only slightly thicker than the thin section at
or near the junction of the two sections ; the thin section, nor-
mally thin throughout its length, remains of small diameter for
but a short distance, then gradually enlarges its diameter until
it joins the bursa copulatrix; infundibulum absent; hemister-
nites very elongate triangular, rounded at tips, styli apparently
absent.
Anisorcus fryi is a species of below medium size, 3.0-3.2 mm.
piceous black above except for head, pronotum, and narrow
lateral margins of the elytra which are reddish-yellow. The
under parts are uniformly reddish-yellow. The structure of the
female genitalia apparently indicates a relationship with Cliilo-
corus and Phaenochilus. The series before me comes from Suva,
Fiji Is., taken in March 1960 by Mr. N. L. li. Krauss. Two
other species were included in this genus at the time of descrip-
tion ; I have seen neither of them.
ChILOCORUS Leach
Chilocoriis Leach in Brewster, 1815, Edinburgh Encyclopedia, 9:116; Eed-
tenbacher, 1843, Tcntamen dispositionis Coleopterorum pseudotrimer-
orum, p. 11; Eeprint of above in Germar, 1844, Zeitsch. f. Ent., 5:118;
Mulsant, 1850, Species Trimeres Seeuripalpes, p. 452; Crotch, 1874,
Eevision of the Coccinellidae, p. 183; Gorham, 1892, Biol. Centr.-Amer.
Ins., Coleopt., 7:175.
Type species. Cocci7iella cacti Linne, monobasic.
Chilocorini wath form broadly oval, strongly convex, upper
surface glabrous except for a few hairs on the flanks of the
pronotum. Antenna eight-segmented ; first segment elongate,
subcylindrical, slightly bent, more than twice as long as wdde
at widest part ; second nearly equilateral, dome-shaped, greatest
width near base, rounded in apical third; third wedge-shaped,
apex twice as wide as base and wider than length ; fourth nearly
quadrate, one-third wider than long ; fifth through eighth form-
ing a fusiform club, fifth and sixth each equal in length to fourth.
264
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each wider at apex than at base, seventh slightly longer and
wider than sixth, eighth conical, half again as long as seventh
but slightly narrower, slightly excavate on one side near apex.
Mandible stout, apex slender, outer margin subangulate near
middle. Terminal segment of maxillary palp with lateral mar-
gins nearly parallel, apical margin strongly oblique, lacinia with
a patch of eight stout spines on outer face. Terminal segment
of labial palp elongate subconical, slightly more than twice
as long as wide at base. Prosternal lobe flat, moderately wide.
Fig. 16. Chilocorus cacti (Linne).
Elytral margins not reflexed, finely beaded; epipleura shal-
lowly foveolate for the reception of the femoral apices. Abdo-
ment with six visible sternites in male, five in female ; metacoxal
arcs in form of a quarter circle, merging with the posterior
margin of the sternite ; margin of fifth sternite transverse,
straight, of sixth slightly emarginate in male, margin of fifth
evenly rounded in female. Legs with stout femora, tibiae with
a triangular tooth on each at basal third, tarsal claws with
small, quadrate tooth on each at base. Male genitalia : median
CHAPIN : GENERA OF THE CHILOCORINI 265
lobe very slightly asymmetrical, elongate triangular with sides
gently curved; parameres hardly longer than median lobe, mod-
erately stout ; trabes slender, longer than the main parts of
the aedeagus ; sipho moderately stout, of even diameter through-
out most of its length, near apex it is twisted through a
quarter turn, apex truncate. Female genitalia : reeeptaculum
seminis a stout body without dilferentiation into nodulus and
ramus, eornu very short and bent, with a falciform appendix
at apex, accessor}^ gland very long and slender ; sperm duct
long, in the usual two sections, the thick section only slightly
thicker than the thin section ; infundibulum absent, replaced
by a fleshy, unpigmented protuberance ; hemisternites rather
broadly triangular.
Chilocorus Leach is very closely related to Egius Mulsant, a
monobasic West Indian genus and also to Phacnochilns AVeise
of the Malayan region. The absence of tibial spurs and the
worldwide distribution of the species of Chilocorus suggests that
the Old World is the original home of the genus. Chilocorus is a
large genus, with many species in the Palaearctic and Ethiopian
regions, fewer in the Indo-malayan, still fewer in the Nearctic,
Neotropical and Australian regions. A thorough study of the
species would probably result in the splitting of the genus.
Egius Mulsant
Egius Mulsant, 1850, Species Trimeres Securipalpes, p. 464.
Chilocorus, — Crotch, 1874, Eevision of the Coccinellidae, p. 188 (part).
Type species. Egius platycepJialus Mulsant, monobasic.
Chilocorini with form nearly circular, flattened convex, upper
surface glabrous. Antenna eight-segmented; first segment short
and stout, nearly equilateral; second longer than first but of
equal width, slightly narrower at apex than at base ; third less
than half as long as second, twice as wide at apex than at base ;
fourth similar in shape to third but longer and wider ; fifth a little
shorter but about as wide as fourth; sixth similar in shape to
fifth but a little wider ; seventh longer than wide, a little longer
than sixth; eighth longer than seventh, tapering to a rounded
apex. Mandibles stout, apices rather blunt, outer margin feebly
concave in basal half, nearly straight in apical half. Terminal
segment of maxillary palp long and narrow, tapering, apex
slightly oblique ; lacinia with a row of six or seven short, stout
spines parallel to the row of longer setae. Terminal segment of
labial palp elongate conical, rounded at apex. Prosternal lobe
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moderate in width, rounded at apex, the margins with a strong;
bead. Elytral margin not reflexed, with a fine marginal bead
in basal third, epipleura not foveolate for the reception of the
femoral apices. Abdomen with six visible sternites in both sexes ;
metacoxal ares joining the posterior margin of the sternite near
the lateral margin. Legs rather stout; femora not inflated; tibiae
with a triangular tooth on outer margin at middle of length ;
tarsi broad, tarsal claws each with a feeble basal tooth. Male
genitalia : median lobe elongate triangular, slightly asymmetrical
Fig. 17. Egius plaiyceplialus Mulsant.
near apex, side margins nearly parallel in basal half ; parameres
flattened, not constricted in basal half, slightly shorter than
median lobe ; trabes moderately stout, equal in length to median
lobe ; sipho rather slender, the apex twisted. Female genitalia :
reeeptaculum seminis and accessory gland similar to those parts
in Chilocorus; sperm duct with slender section very thin, total
length of duct proportionately longer than in Chilocorus; in-
fundibulum absent, replaced by a fleshy protuberance on the
CHAPIN : GENERA OF THE CHILOCORINI
267
bursa copulatrix ; hemisternites short, rather broad, somewhat
ereseent-shaped, styli apparently absent.
The type and only species of this genus is found on Cuba.
While anatomically very close to Chilocorus, its superficial ap-
pearance is very different from that of any of the recorded
species of Chilocorus. The very strongly alutaceous surface of
the head and elytra, and the almost iridescent metallic greenish-
purple color of the elytra set it apart immediately.
PhAEXOCHILUS Weise
Phaenochilus Weise, 1895, Ann. Soe. Ent. Belgique, 39:135; 1913, Philip-
pine Journ. Sci., 8, D:241.
Type species. Phaenochilus punctifrons Weise, by subsequent designation
of Korschefsky, 1932.
Chilocorini with form nearly circular, strongly convex, upper
surface glabrous. Antenna eight segmented ; first segment short,
stout, curved; second longer and thicker than first, much nar-
rower at apex than at base ; third nearly twice as long as wide at
base, apex half again as wide as base ; fourth to sixth mutually
similar in shape, fourth and fifth nearly equal, seventh slightly
Fig. 18. Phaenochilus punctifrons Weise.
268 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
longer and wider than fifth ; eighth twice as long as sixth, taper-
ing to a blunt point. Mandible somewhat longer than broad, outer
margin not strongly curved, apex rather elongate and slender.
Maxillary palp long and slender, the terminal segment very long,
about three times longer than wide, tapering to a blunt point ;
lacinia wdth a patch of about seven stout spines on outer face.
Terminal segment of labial palp slender acuminate, rounded at
apex. Prosternal lobe moderately wide, convex, rounded at apex,
the lateral margins beaded, the bead continued partly around
the anterior coxal cavities. Margins of elytra strongly reflexed,
with extremely fine marginal bead ; epipleura shallowly f oveolate
for the reception of the femoral apices. Abdomen with six visible
sternites in male, the fifth sternite broadlj' truncate, five in
female, the fifth broadly rounded. Metacoxal arcs run outward
and back nearly to the posterior margin of the sternite, thence
following the margin nearlj^ to the lateral margin of the abdo-
men. Legs rather short ; femora moderately stout ; tibiae slender,
those of legs II and III with triangular tooth at the top of the
tarsal groove, that on leg III stronger than that on leg II ;
terminal segment of tarsus stout, claws stout, the apical half at
nearly a right angle to the basal half, the basal tooth elongate,
parallel to and nearly as long as the apical half of the claw.
Male genitalia : median lobe equal in length to the parameres,
widest at apical third, side margins straight in basal two-thirds,
apical third triangular, apex acutely pointed ; parameres paddle-
shaped, not noticeably constricted basally, apices bluntly
rounded ; trabes slender, longer than the median lobe ; sipho
rather slender, of even diameter in first four-fifths, apical fifth
much more slender and ending in a harpoon-shaped j)oint.
Female genitalia : receptaculum seminis very stout, much like
that of Chilocorus, appendix of cornu strongly developed, ac-
cessory gland somewhat longer than long diameter of the re-
ceptaculum; .sperm duct with thick portion more than twice as
long as the very thin portion ; infundibulum absent, replaced by
a fleshy tab at apex of bursa copulatrix ; hemisternites moder-
ately long, triangular, styli not prominent, appearing as pits
near apices of these sclerites, each pit furnished with two or
three long setae.
This small genus of three species, P. punctifrons Weise, P.
ruficollis Weise, and P. monostigma Weise, is at home in the
Indomalayan region from Java to Mindanao in the Philippines.
Specimens of all three species are in my hands. The genus may
CHAPIN : GENERA OK THE CHILOCORINI 269
be immediately recognized by the long, slender maxillary palps
and the broadly reflexed elytral margins.
TriCHORCUS Blackburn
Trichorcus Blackburn, 1892, Trans. Proe. Koy. Soe. South Australia, 15:73.
Type species. Trichorcus cinctus Blackburn, monobasic.
The original diagnosis is quoted in full below.
"Trichorcus (gen. nov. Coccinellidarum) Ab Oreo differt
corpore baud metallico, subopaco, dense pubescenti.
I can find no structural character to distinguish the genus
from Orcus, but it seems scarcely possible to place in that genus
a non-metallic subopaque species densely clothed with pubes-
cence."
From the description of the type species little of importance
can be learned except that the individuals of this species are a
trifle wider than long. It will be necessary to study specimens
of this species in order to disclose its relationships.
CHIL0C0RU8 (TrICHOCORUS) Sicard
Chilocorus {Trichocorns) Sicard, 1921, Bull. Soc. Portugaise Sci. Nat.,
8:213; Mader, 1954, Explor. Pare Nat. Albert, Mission de Witte, Lief.
80, Coccinellidae, pars III, p. SO.
Type species. Chilocorus pilosus Sicard, by present designation.
It is not possible to place this taxon in its proper position in
the tribe from the information contained in the above references.
Two species were included in the original description and these
ditfer in the formation of the anterior tibiae. The type species
is said to have angulate but not spinose tibiae, a very small
scutellum, upper surface clothed with somewhat long, gray
pubescence, elytral margins not at all reflexed, basal margin of
pronotum sinuate either side of the scutellum, and other charac-
ters mostly relating to color.
Unfortunately, through a misreading of the original descrip-
tion, Dr. Mader has introduced a false character into his key.
Sicard states that the pubescence is "dirigee en avant" on the
pronotum only, on the elytra it is "diriges {sic!) en arriere."
CORYSTES Mulsant
Cnrystcs Mulsant, 1850, Species Trimeres Securipalpes, p. 506; Crotch, 187-1,
Kevision of the Coccinellidae, p. 208; Chapuis, in Lacordaire, 1876,
Gen. Coleopt., 12:24-4, 249; Gorham, 1894, Biol. Centr.-Amer., Ins..
270 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Coleopt., 7:182; Weise, 1899, Deutseh. Ent. Zeitscli., p. 269, 1903, ibid.,
p. 208; 1904, ibid., p. 358; 1927, Arkiv f. Zool., 18A(4), no. 34,
pp. 12-13.
Type species. Corystes hypocrita Mulsant, monobasic.
Mulsant, 1850, placed his new genus Corystes in the third
branch, Thalassaires, of his liyperaspiens. He took note of the
fact that the structure of the anterior part of the head was dif-
ferent from that of his Chilocoriens in that in Corystes there is a
break in continuity between the clypeus and genal lobes and no
break in other Chilocoriens.
Crotch, 1874, agrees wdth Mulsant in placing Corystes near
Thalassa but brings these genera to the beginning of his Hyper-
aspides, directly following his group, Chilocorides.
Chapuis, 1876, removed the genus from his Hyperaspites and
placed it as a transitional form in the Chilocorites. However, he
noted, as Mulsant had before, that the structure of the head is
not typically chilocorine and that otherwise the characters of
the genus are largely hyperaspine.
Gorham, 1894, added little to the clarification of the problem.
He accepted Chapuis' findings and continued Corystes in the
Chilocorini.
Weise, 1899, working with specimens which he believed at the
time were Corystes hypocrita, returned the genus to the Hyper-
aspini. In 1903, he added a second species to the genus, noting
that the structure of the head of the new species was somewhat
different from that of the original species. In 1904, Weise again
asserts that the genus is undoubtedly hyperaspine.
In 1927, in a posthumous paper, Weise reverses himself on
the ground that the species dealt with in 1899, 1903, and 1904,
were misidentified. In this paper, he sets up a new genus, Dia-
zonema, type I), fallax Weise {Corystes hypocrita Weise, not
Mulsant, renamed), in the Hyperaspini and redescribes Corystes
hypocrita Mulsant from specimens from San Fermin, Bolivia.
He now places Corystes in the Chilocorini but notes the difference
between it and the rest of the Chilocorini in the structure of the
clypeus and genal lobes.
I have before me a short series of a species from Colombia
which I am convinced represents an undescribed species of
Corystes. There is no question in mj- mind that the species does
belong in the Hyperaspini near Thalassa. The antenna with
the antepenultimate segment very long, the mandible with a
second tooth very near the apex, the male genitalia with a de-
pressed, strongly asymmetrical median lobe, and otlier characters
CHAPIN : GENERA OF THE CHILOCOBINI 271
are all foreign to the Chilocorini but common to all Hyperaspini
known to me.
ElPIS Mulsant
Elpis Mulsant, 1850, Species Triineres Securipalpes, p. 449; 1866, Mono-
graphie ties Coecinellides, pp. 283, 291; Crotch, 1874, Revision of the
Coceinellidae, p. 181; Chapuis in Lacordaire, 1876, Gen. Coleopt.,
12:191, 193; Sicard, 1909, Ann. Soc. Ent. France, 78:64, 82; Kors-
ehefsky, 1932, Coleopt. Catalogus (Junk), pars 120, Coceinellidae, p.
248; Mader, 1941, Explor. Pare Nat. Albert, Mission de Witte, fasc.
34, p. 185; 1954, op. cit., fasc. 80, pp. 117, 118.
Type species. Elpis dolens Mulsant, monobasic.
The above are all the references to Elpis that I have been able
to find in the literature. All of the authors, except Korschefsky,
agree in placing this genus in the Coccinellini, near Menochilus.
I have examined a specimen of the type species and it is obvious
that the genus does belong in the Coccinellini. Korschefsky gave
no explanation for placing the genus in the Chilocorini.
(Eeeeived 8 December 1964.)
Bulletin of the Museum of Comparative Zoology
HAEVARD UNIVERSITY
Vol. 133, No. 5
COMMENTS ON SOME RECENT CHANGES IN THE
CLASSIFICATION OF THE CIIDAB (COLEOPTERA)
By
John F. Lawrence
CAMBRIDGE, MASS., U.S.A.
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Bulletin of the Museum of Comparative Zoology
HAEVARD UNIVERSITY
Vol. 133, No. 5
COMMENTS ON SOME RECENT CHANGES IN THE
CLASSIFICATION OF THE CIIDAE (COLEOPTERA)
By
John F. Lawrence
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
October, 1965
Bull. Mus. Comp. Zool., Harvard Univ., 133(5) : 273-293, October, 1965.
No. 5 — Comments on Some Recent Changes in the Classification
of the Ciidae (Coleoptera)
By
John F. Lawrence
Musum of Comparative Zoology, Harvard University, Cambridge, Mass.
The family Ciidae includes at present 40 genera and about
550 species. The group has been badly neglected in the past,
and although a few excellent local monographs have appeared,
the last treatment of the world fauna was published in 1848.
The European fauna has probably been more extensively studied
than that of any other area, but most of the workers have not
been specialists and have tended to be provincial in their ap-
proach, completely ignoring, for instance, the contributions to
the Holarctic fauna made by Japanese and Americans. Lohse
(1964) produced a short paper in which some of the European
generic concepts were reexamined and 3 new generic group names
proposed. This recent contribution is certainly welcome, since
it represents the first step in clarifying the relationships within
this difficult group. In the present discussion, I will review
Lohse 's proposals and reevaluate his concepts in the light of my
own work on the North American Ciidae. Since the histories of
the various European generic names have been discussed pre-
viously, it may seem redundant to include them here ; I think this
is justified, however, considering that even Lohse overlooked
or misinterpreted some of these historical facts.
1. ENTYPVS, RHOPALODONTUS, AND 8ULCACIS
Although the genera Entypus and Rhopalodontus represent
distinct taxa whose species are only distantly related, the nomen-
clatural histories of the two names are so interwoven that it
seems practical to discuss them together. Ropalodontus Mellie
(justifiably emended to Rhopalodontus by Gaubil, 1849) was
first proposed in 1847 for the species Cis perfomtus Gyllenhal,
the main diagnostic character being the expanded, rounded, and
externally spinulose protibial apices. In the same year, Redten-
bacher described Entypus, which included only the species Cis
affinis Gyllenhal (misidentified as Apate fronticornis Panzer),
characterized by having 9-segmented antennae, with the 3rd seg-
ment longer than the following 3 together, and spinulose protibial
276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
apices. In the following year, Mellie included C. affinis in his
genus Ennearthron, which had been described some months prior
to the publication of Redtenbacher's work, and placed Entypus
in synonymy. Mellie included the true A. fronticornis in the
genus Cis, primarily because of its 10-segmented antennae.
Thomson (1863) considered Ehopalodontus and Orophius Red-
tenbacher (=Octotemnus Mellie) to be so different from other
members of the family that he included them in a separate tribe,
the Orophiina. Being a careful worker, he found several other
characters which consistently distinguished the group, such as
the conical, projecting, and subcontiguous procoxae, short pro-
sternum, deep antennal sulci, and subequal 3rd and 4th antennal
segments. In the same paper, Thomson noted the close similarity
between Ennearthron affine (Gyllenhal) and Cis fronticornis
(Panzer), and in spite of the difference in antennal segmenta-
tion, he placed them both in Entypus. Mellie 's name Ennear-
thron could not be applied, because a very different species, Cis
cornutns Gyllenhal, had been designated as its type by Demarest
(1860). Kiesenwetter (1877) included Entypus as a distinct
subgenus of Cis and added to it a third species, Ennearthron
ivagai Wankowicz, which had been described as having spinulose
protibial apices. He considered the Orophiina to be a separate
family, the Orophyidae, to which he added Xylographus Mellie.
A more extreme view was taken by Seidlitz (1872), who placed
these genera in a tribe of the family Anobiidae.
In spite of the efforts of Thomson and Kiesenwetter, most
European entomologists chose to ignore these two concepts and
to rearrange the above species according to more superficial char-
acters, such as the number and relative lengths of antennal seg-
ments. Thus, Jacquelin Du Val (1861) placed C fronticornis in
the genus Rhopalodontus in spite of its totally different pro-
coxal structure, and returned affinis to Ennearthron, because of
its 9-segmented antennae. In 1915, Peyerimhoff placed Cis hi-
cornis Mellie in Rhopalodontus on the basis of antennal and pro-
tibial characters only, and Nobuchi (1960a) described 2 more
species, R. japonicus and R. tokunagai, which also appear to be
in the fronticornis group. Until quite recently, then, the genus
Rhopalodontus contained two distinct and distantly related
groups of species : those resembling Ennearthron affine and be-
longing properly to Entypus of Redtenbacher and Thomson, and
those resembling R. perforatus.
The situation was clarified by Lohse (1964), who had restudied
the European species carefully enough to recognize the true
LAWRENCE: CLASSIFICATION OF CIIDAE 277
relationships. By reviving the concept of Redtenbacher and
Tliomson and uniting E. affinc, E. fronticornis, and R. hicornis
into a single genus, he not only called attention to the apparent
affinities of these three species, but also eliminated discordant
elements from both Ennearthron and Rhopalodontus. In addi-
tion to the above, Lohse added Cis bidentulus Rosenhauer to the
group. Because of the variation in antennal segments, he pro-
posed the subgenus Entypocis (with C. bidentulus as its type)
for those species with 10-segmented antennae, leaving Entypus
affinis the only member of the nominate subgenus. Ennearthron
wagai Wankowicz (1869) (mis-cited in Lohse as E. wagai
Wanka) was placed in a new genus, Wagaicis, because of its very
narrow and laminate prosternal process.
I agree with Lohse 's revival of Entypus and his inclusion of
the 4 European species. In my own studies of North American
Ciidae, I have encountered two more species, Cis curtidus Casey
and Sulcacis lengi Dury, which also belong to this group. I have
compared these with specimens determined as and fitting the
descriptions of E. fronticornis, E. affinis, and E. bidentulus. C.
curtulus is very similar to E. bidentulus, w^hile S. lengi is unique
in having 9-segmented antennae, a somewhat shortened proster-
num, and no sexual modifications on the head of the male. Since
lengi is here selected as the type of Stdcacis Dury (1917), this
genus becomes a junior synonym of Entypus. I would tentatively
add R. japonicus and R. tokunagai, both from northern -Japan,
to this genus, on the basis of descriptions and illustrations. I do
not think the proposed division into subgenera is necessary,
especially if it is based on the number of antennal segments.
There is one final matter that appears to have been overlooked
by all of the w^orkers in the Ciidae. This is the fact that the name
Entypus was not available when used by Redtenbacher, having
already been applied to a pompilid wasp (Hjanenoptera) by
Dahlbom (1843). Entypus is a primary junior homonym and
must be replaced. Of the two remaining available names which
have been applied to the group, Sulcacis Dury is the oldest and
is here considered as a replacement name. Although Sulcacis
lengi Dury is the most atypical member of the group, it becomes
the type species because of the priority of the generic name.
I do not think that S. lengi is distinct enough to be placed in a
separate genus ; if this species were to be removed from the group,
however, then Lohse 's Entypocis is the next oldest available
generic name and E. bidentulus would be considered the type.
278 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
A synonymy and redefinition of Sulcacis is given below.
SULCACIS Dury
Sulcacis Dury, 1917, Jour. Cincinnati Soc. Nat. Hist., 22(2) :20; Leng,
1920:247. Type, by present designation, Sulcacis lengi Dury, 1917:21.
Eniypus Eedtenbaelier, 1847, Faun. Austr., (3):350 {non Dahlbom,
1843:35); Thomson, 1863:193; Sahlberg, 1926:78; Lohse, 1964:118.
Type, by monotypy, Cis affinis Gyllenhal, 1827:628 (misidentified as
Apate froniicornis Panzer).
Cis {Eniypus), Kiesenwetter, 1877:190; Eeitter, 1878:21.
Entypus (Eniypocis) Lohse, 1964, Ent. Blatter, 60(2) :121. Type, by orig-
inal designation, Cis hideyitiilus Eosenhauer, 1847:58.
Cis (in part), Gyllenhal, 1827:624; Mellie, 1847:109; Mellie, 1848:236
Lacordaire, 1857:551; Jacquelin Du Val, 1861:237; Seidlitz, 1872:44
Abeille de Perrin, 1874:19; Eeitter, 1878:27; Seidlitz, 1891:282
Casey, 1898:78; Sehilsky, 1900 :37E; Eeitter, 1902:47; Dalla Torre,
1911:5; Dury, 1917:5.
Ennearthron (in part), Mellie, 1848:360; Lacordaire, 1857:552; Jacquelin
Du Val, 1861:238; Abeille de Perrin, 1874:80; Eeitter, 1878:30;
Sehilsky, 1900 :37B; Eeitter, 1902:59; Dalla Torre, 1911:23; Nobuchi,
1960a: 41.
Ehopalodontus (in part), Jacquelin Du Val, 1861:238; Abeille de Perrin,
1874:76; Eeitter, 1878:30; Sehilsky, 1900 :37D; Eeitter, 1902:57;
Dalla Torre, 1911:21; Peyerimhoff, 1915:26; Nobuchi, 1960a: 39.
Form oblong to elongate, strongly convex and cylindrical ; ves-
titure short, usually dual, consisting of both erect and inclined
bristles. Head moderately declined, partly covered by pronotum ;
frontoclypeal ridge of male usually with 2 small teeth or tuber-
cles ; antennal fossa relatively deep. Antenna 9- or 10-segmented,
with 3-segmented club; segments III and IV subequal, or III
longer than next 2 combined ; maxillary palp narrow and
elongate. Pronotum somewhat constricted anteriorly, narrowly
margined laterally; anterior edge usually simple in both sexes.
Elytra with relatively coarse, indistinctly dual, uniform or
seriate punctation. Prosternum slightly tumid, concave laterally,
slightly shorter than iutercoxal process which is narrow but not
laminate. Protibia expanded at apex, outer apical angle forming
a rounded process bearing several small spines. Metasternum
convex, the suture short. Meso- and metatibiae slightly expanded
and spinulose at apex. Sternite III of male with median pubes-
cent fovea.
A small genus with species throughout the Holarctic region.
Intermediate in its characters between Cis and Eridoulus, on the
one hand, and Ceracis, Wacjaicis, and Malacocis on the other, dif-
fering from the former by the spinulose protibial apices and from
LAWRENCE : CLASSIFICATION OF CIIDAE 279
the latter by the broader intercoxal process. The species which
have been studied biolo<i'ically occur primarily on fungi of the
Folyporns versicolor group (Paviour-Smith, 1960).
2. HADRAULE, ERIDAULU8, AND XE8T0CI8
Two of Thomson's genera, Iladraulc and Eridaulus, have been
involved in considerable taxonomic confusion over the past 100
years. Lohse (1964), in his analysis of the problem, has referred
to it as a " nomenklatorischen Tragikomodie," an opinion with
which I am inclined to agree.
Hadraide was originally proposed for the species Cis elongatu-
lus Gyllenhal, characterized by the somewhat flattened body form,
subquadrate prothorax, well-separated procoxae, and regularly
striato-punctate elytra. Seidlitz (1872), Kiesenwetter (1877),
and Reitter (1878) considered it to be a subgenus of Cis. Schil-
sky (1900) first expanded the subgenus to include certain other
species with a more or less parallel form, subquadrate prothorax,
and elytra with rows of setae alternating with rows of large
punctures. In addition to C. elongatulus (misspelled as elonga-
tus in text, p. 59), he placed Cis setifer Reitter and Cis striatulus
Mellie within Hadraule. Reitter (1902) continued the trend by
including the following species as well : C. seriatopilosus Mot-
schulsky, C. heiroglyphicus Reitter, C. hifasciatus Reitter, C.
comptus Gyllenhal, C. coriaccus Baudi, and C. pumilio Baudi.
Jacobson (1915) doubtfully placed Cis fuscipcs Mellie, described
from North America and Madeira, in the same group. Roubal
(1936), having examined the type of C. dongatulus and dis-
covered that it had 9-segmented, rather than 10-segmented, an-
tennae, transferred the subgenus (including only the type
species) to the genus Enncarthron Mellie. Through a misunder-
standing of the rules of nomenclature, Roubal replaced Had^raule
with a new name, Knahlia. Lohse (1964) restored Hadraule
and suggested that it be used in the original sense, as a separate
genus including only the type species, Hadraule elorigafula.
Lohse further stated that Cis comptus and its relatives should
be placed in the subgenus Eridaulus. This is entirely wrong,
being based on an erroneous type designation, as will be ex-
plained below. The rest of the species mentioned above do seem
to form a natural group, to which the American species Cis falli
Blatchley, Cis striolatus Casey, and Cis versicolor Casey might be
added. H. elongatula shows a marked similarity to members of
the C. comptus group and also to MapJioca hlaisdelli Casey, the
280 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
latter differing mainly in the number of antennal club segments.
The genus Eridaulus Thomson (1863) originally included two
species, Anohium nitidum Fabricius and Cis jacquemarti Mellie,
which were characterized by the 10-segmented antennae, reflexed
"gular margin," elytra with larger and smaller punctures (dual
punctation), short, carinate prosternum, and produced and den-
tate protibial apices. Seidlitz (1872), Sahlberg (1926), and a
few others retained it as a genus, but most later authors reduced
it to a subgenus of Cis. Seidlitz added Cis glabra f us Mellie and
C. lineatocrihratus Mellie, and Schilsky (1900) included C.
quadridens Mellie as well. Roubal added one more European
species, Cis hitu'berculosns Roubal, 1937 {=Cis hituherculatus
Roubal, 1912, non Gorham), and in recent years Chujo (1940),
Miyatake (1954), Nakane and Nobuchi (1955), and Nobuchi
(1955, 1960b) have described several others from Japan. The
only worker to designate a type for Eridauhis was Abeille de
Perrin (1874), who selected Cis comptus Gyllenhal. This species,
however, was not originally included in the genus, and is there-
fore unavailable as a type. In the same work, Abeille de Perrin
designated C. nitidus as the type of Entypus, making a similar
error. The name Eridaulus, then, is still available for the Cis
nitidus group, contrary to the statement of Lohse (p. 119), and
the type must still be designated. Since I have not examined the
type specimens of either C. nitidus or C. jacquemarti, I hesitate
to select one of them as the type at the present time.
In the past several years, I have examined specimens of Ciidae
from various parts of the world, and it has become apparent to
me that a number of other species described in Cis and the major-
ity of those included in Xestocis Casey should be placed in
Eridaulus. Casey (1898) proposed the genus Xestocis for 5
North American species with a carinate prosternum. Dury
(1917), Brethes (1922), and Hatch (1962) added 14 more
species, and most of those examined fall within this group.
Since a type has not been selected for Xestocis, I here designate
X. levettei Casey, the best known of the American species. Miya-
take (1954) noted the similarity of X. levettei to Cis (Eridaulus)
konoi Chujo, and I have noticed the resemblance of the former to
C. nitidus, with respect to dual elytral punctation, carinate
prosternum, dentate protibial apices, secondary sexual characters,
male genitalia, and larval urogomphi. I think there is little
doubt that Xestocis should be considered a junior synonym of
LAWRENCE: CLASSIFICATION OF CIIDAE 281
Eridaulus. Since a number of species synonymies and new gener-
ic combinations are involved, I will not present here a complete
list of American forms to be included in Eridaulus, but will
publish this separately in a forthcoming revision of the New
World species. In addition to the American forms, several species
of Cis from the Pacific region, such as Cis i^^^'Cificus Sharp
(Hawaii) and C. agariconae Zimmerman (Micronesia) should be
placed in the genus.
Although Eridaulus appears to be quite close to the genus Cis,
I think it should be given generic rank for several reasons.
According to Mayr, Linsley, and Usinger (1953), the size of the
gap between two genera should be inversely proportional to the
sizes of the groups involved. With its present constitution, the
genus Eridaulus is large enough, both in numbers of species and
in extent of distribution, to warrant generic distinction. There
are about 30 named species, and several more apparently unde-
scribed, extending throughout the Holarctic region and as far
south as New Zealand in the Pacific area. Furthermore, within
the genus, several distinct subgroups can be recognized, and
certain independent character trends can be followed. Finally,
the group displays a biological unity, most of the species being
associated with the larger, woody fruiting bodies of fungi such
as Fomes and Ganoderma. This habit, which has been inde-
pendently evolved in several unrelated genera, such as Ceracis
and Xylographus, may be associated with structural modifica-
tions, such as the prosternal carination and convex form of the
adult and the increased molar area in the larval mandible.
Paviour-Smith (1960) noted that the "headquarters" of E. niti-
dus was the fungus Ganoderma applayiatum, although the beetle
was recorded from several other fungi, and this is quite similar
to the host range and headquarters of the American E. levettci.
Saalas (1923) reported E. quadridens, E. Uneatocribratus, and
E. jacquemarti from Fomes pinicola, and the Japanese species
E. rufocastaneus and E. nikkoensis have been taken on the same
fungus. E. hiarmatus (Mannerheim) and several Eridaulus irom.
western North America occur on Fomes piuicola, F. annosus
and related fungi. In Hawaii, where most of the Ciidae are col-
lected by beating dead vines and branches, E. pacificus is the only
species which has been reported feeding on the woody bracket
fungi. It appears to me, then, that the members of this group
represent a morphologically and biologically distinct genus, to
which the name Eridaulus should be applied. Synonymies of
282 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Iladraule and Eridaulus and a redefinition of the latter genus are
eiven below.
fc>
HaDRAULE Thomson
Hadraule Thomson, 1863, Skand. Col., 5:182; Sahlberg, 1926:71; Lohse,
1964:119. Type, by monotypy, Cis elongaiulus Gyllenhal, 1827:627.
Cis {Hadraule), Seidlitz, 1872:44; Kiesenwetter, 1877:188; Reitter,
1878:24; Seidlitz, 1891:281; Sehilsky, ]900:37F (in part); Reitter,
1902:47 (in part); Dalla Torre, 1911:5; Jacobson, 1915:953 (in part).
Tityocis Peyerimhoff, 1918, Bull. Soc. Ent. France, 1918:141. Type, by
monotypy, Pityocis coarctatus Peyerimhoff, 1918:142 ( = Cis elongatulus
Gyllenhal). [See Peyerimhoff, 1933.]
Ennearthron {Knahlia) Roubal, 1936, Festsehr. 60 Geburtst. Embrik Strand,
1:53. Type, by monotypy, Cis elongatulus Gyllenhal, 1827:627.
Hadraida Leng, 1920:246, incorrect subsequent spelling.
Eridaulus Thomson
Eridaulus Thomson, 1863, Skand. Col., 5:191; Seidlitz, 1872:45; Sahlberg,
1926:79. Originally included species: Anohium nitidum Fabricius,
1792:238 and Cis jacquemarti Mellie, 1848:328. Type not designated.
Cis {Eridaulus), Kiesenwetter, 1877:191; Reitter, 1878:21, 24; Seidlitz,
1891:281; Sehilsky, 1900 :37F; Reitter, 1902:48; Dalla Torre, 1911:5;
Roubal, 1912:29; Roubal, 1937:39; Chujo, 1940:132; Miyatake,
1954:49; Nakane and Nobuchi, 1955:49; Nobuchi, 1955:56; Nobuchi,
1960b :65.
Xestocis Casey, 1898, Jour. New York Ent. Soc, 6(2) :85; Dalla Torre,
1911:20; Dury, 1917:15 (in part); Leng, 1920:247; Hatch, 1962:231.
Type, by present designation, Xestocis Icvettei Casey, 1898:85. NEW
SYNONYMY.
Anohium, Ptinus, Cis auctt.
Form oblong, strongly convex ; vestiture of short, fine hairs,
longer recumbent hairs, or short, stout setae. Head moderately
declined, partly covered by pronotum ; f rontoclj'peal ridge of
male produced on each side forming 2 flattened plates which
are rounded to triangular ; genal ridge strongly elevated and
carinate, forming relatively deep antennal fossa. Antennae 10-
segmented, with a 3-segmented club. Pronotum strongly convex,
variously margined laterally, anterior angles rounded to pro-
duced and acute ; anterior edge in male simple or produced and
cmarginate. Elytra with dual punctation, consisting of larger
nude macropunctures, and smaller setiferous micropunctures,
the punctures uniform or seriate. Presternum relatively short,
strongly tumid, concave laterally and carinate mesially ; inter-
coxal process relatively narrow but not laminate, subacute at
LAWRENCE : CLASSIFICATION OF CIIDAE 283
apex. Protibia expanded at apex, outer apical angle produced
and dentate. Sternite III of male with median pubescent fovea
or patch.
A large genus with a primarily Holarctic distribution, but
extending south at least in the Pacific region. Closely related to
Cis and Strigocis, but distinguished by the strongly carinate pro-
sternum, dual elytral punctation, and produced and dentate pro-
tibial apices. The genus differs from Hadraule, Orthocis, and
Dulichocis in general body form and by the dentate protibial
apices, as well as by the carinate prosternum. Most of the
species in this genus feed on the larger woody fungi, such as
Fomes and Ganoderma.
3. ORTHOCIS, MELLIEICIS, AND DOLICHOCIS
Casey (1898) proposed the genus Orthocis for the two Ameri-
can species, Cis puncfatus Mellie and Orthocis aterrima Casey,
which were distinguished from the species of Cis by the "...
more parallel form of the body . . . glabrous surface, margined
elytral suture, and . . . simple apex of the anterior tibiae."
Dury (1917) described a third species, 0. longida, from the east-
ern U.S. and Kraus (1908) added two species, 0. huesanus and
0. pidcher, from Florida. Another species, 0. platcnsis Brethes
(1922), was described from Argentina.
In the European literature, it has long been recognized that
Cis alni Gyllenhal, and several related forms, such as C. perrisi
Abeille de Perrin and C. coluhcr Abeille de Perrin, form a
distinct group, characterized by the elongate body form, short
pubescence, simple protibial apices, and lack of distinct sexual
modifications on the head or prothorax of the male. Lohse
(1964) proposed for these species the subgenus Mcllieicis and
selected Cis alni as its type. Although the exact limits of the
subgenus were not given in Lohse 's paper, it was indicated that
all species of Cis with rounded or truncate protibial apices should
be included. Having studied the European literature and ex-
amined specimens in the American collections, I have come to
the conclusion that both Cis punctatns Mellie sensu Casey and
Orthocis aterrima Casey are very closely related to the Palearctic
species in the Cis alni group. The name Orthocis, then, should
apply to this group of species, and McUicicis should be considered
a junior synonym.
In 1908, Kraus described 3 species of Ennearthron, E. annida-
tum, E. transversatum, and E. pallidum, from the southeastern
284 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
United States and Cuba, which differ from the species of Orthocis
only in the possession of 9-segmented antennae. I think it is only
sensible to expand the definition of Orthocis to include these
species as well, instead of leaving them in a genus which appears
to be a conglomerate of unrelated forms. If the limits of the group
are thus expanded, it becomes necessary to consider several other
species which might be included. In 1919, Dury described the
genus Dolichocis with the single species D. manitoha Dury, the
main characteristics being the 9-segmented antennae, narrow,
cylindrical form, vestiture of short bristles, and rounded pro-
tibial apices. The species differs from the American species
of Orthocis, not only in its antennal segmentation, but by its
stouter pubescence, anteriorly constricted prothorax, and the
presence of 2 clypeal tubercles and a pubescent fovea on the
head of the male. Hatch (1962) described a second species, D.
incUstinctus, from western North America, which is very similar
to and obviously congeneric with E7inearthron laricinmn (Mel-
lie) of the European fauna. Other Palearctic species which might
be in the same group are Ennearthron yuasai Chujo, E. pruino-
siilum (Perris), and E. poriae Nakane and Nobuchi, the last two
of which resemble Orthocis more than Dolichocis in the more
quadrate prothorax with broader lateral margins and in the
absence of sexual modifications on the head of the male. The
problem is made more complex by the presence of 3 Palearctic
species, Cis festivus (Panzer), Cis pygmaeus (Marsham),
and Cis rhododactylus (Marsham), and one North American
species, Cis angustus Hatch, which share certain character-
istics with both Cis and Orthocis. Nyholni (1953) noted
that the first three all have truncate or rounded protibial
apices, and I have observed the same character in C. angustus.
Yet all of these have sexual ornaments on the head of the
male, and they differ in general appearance from typical Ortho-
cis. The setting of generic limits, if this is at all justified,
presents a difficult problem, which can be solved only after a
more detailed investigation of all species involved. In the pres-
ent discussion, I will make a few suggestions based on a rather
superficial study of a large number of forms from various
parts of the world and a more intensive study of American
representatives.
In the North American fauna, both Orthocis and Dolichocis
seem to be well defined both morphologically and biologically.
All of the species of Orthocis are elongate and parallel, with a
LAWRENCE: CLASSIFICATION OF CIIDAE 285
vestiture of very short and fine hairs, a fairly smooth and
shining surface, margined elytral suture, flat or slightly tumid
prosternum, rounded protibial apices, and an absence of sexual
ornaments on the head of the male. There is a tendency in some
species (0. longulus Dury) towards an extreme attenuation of
the body, and several subtropical forms are bicolored and may
have 9-segmented antennae. Field observations indicate that
at least some species occur under bark, in decaying branches and
vines, and apparently not in the tougher fruiting bodies of most
Polyporaceae. The genus is more common in the southern part
of the continent, and a number of unnamed species have been
seen from Central and South America. The two species of Boli-
chocis, on the other hand, dift'er in having the prothorax con-
stricted anteriorly, the vestiture consisting of short, stouter
bristles, and the head modified in the male. Both species occur
in the northern part of the continent, and, like most Eridaulus,
they occur mainly on the fruiting bodies of woody fungi.
In the European fauna, generic distinctions are not so easily
made. Cis coluber Abeille de Perrin, C. reflexicollis Abeille de
Perrin, and C. juglandis Reitter appear to be closely related to
0. alni, but they all have a stouter pubescence. Members of the
Cis festivus group differ both in vestiture and in the presence
of clypeal tubercles in the male. Of the species with 9-segmented
antennae, E. laricinum undoubtedly belongs to Dolichocis, but E.
priiinosulum appears to be closer to Orthocis. One character
which has been overlooked by most European workers is the
margined elytral suture in species of Orthocis. At the apex of
each elytron, there is a distinct raised margin which curves
laterad, leaving a narrow flat area between it and the edge of
the suture. I have observed this in 0. alni, as well as in the
American species. This character, in combination with others
mentioned above, might serve to distinguish consistently the
species of Orthocis from those of Cis and Dolichocis. Several
other features which I have observed only in the North American
Orthocis are the elongate prementum, elongate maxillary stipes
with a flattened laterally placed laeinia, and the deeply emar-
ginate, mesially unpigmented, 8th sternite of the male. An in-
vestigation of these and other more cryptic characters in the
European species may shed some light on their true relation-
ships.
Orthocis, like Eridaulus, is so similar to Cis that the question
arises whether a generic distinction is justified or not. I think
286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
that the same criteria can be used here as were applied in the
case of Eridaulus. The size of the group is quite large, especially
in view of the number of unnamed species which I have seen,
and the distribution is probably cosmopolitan, with the majority
of species occurring in tropical and subtropical regions. Several
unique trends occur within the genus, and it appears that the
group may have given rise to some of the peculiar endemic forms
on the Hawaiian Islands and in other Pacific areas. The latter
resemble GrfJiocis in the minute vestiture, flattened prosternum,
simple protibial apices, and lack of male ornaments, but they
vary considerably in body form, sculpture, and coloration. The
trend towards extreme attenuation has already been mentioned.
The biological characteristics of the genus, although briefly
mentioned above, deserve further comment. Most of the species
of Ciidae feed on the sporophores of wood-rotting fungi, espe-
cially the more fibrous or woody fruiting bodies of the Poly-
poraeeae and Hydnaceae. These species have evolved various
morphological and physiological adaptations to enable them to
cope with a substrate which not only presents a physical barrier
to feeding but usually contains little moisture and a high per-
centage of pure chitin, unavailable to insects not possessing a
symbiotic intestinal flora. In addition, there has been a tendency
for various species to specialize in certain groups of host fungi
(Paviour-Smith, 1960). The species of Ortliocis, whose habits
have been recorded, appear to be general fungus feeders, oc-
curring under bark, in dead branches and vines, in more humid
situations, and in association with mycelia and fruiting bodies
of a wide variety of wood-rotting fungi. The fungus sporophores
are usually of a softer and more ephemeral type and their loca-
tions permit a certain amount of bacterial and fungal decompo-
sition, so that the resulting substrate is more easily utilized by
the beetles. Beniek (1952) has reported Cis alni from such
diverse fungi as Exidia glandulosa (Tremellaceae), Stereum
rugosuni (Thelephoraceae), and Auricularia auricula-judae
(Auriculariaeeae), while Lucas (1849) found the same species
(as C. puncUdatus Lucas, non Gyllenhal) on Schizophyllum
commune (Agaricaecae). Blatchley (1910, 1923) noted that
0. punctatus was taken by sifting debris from an oak log and
that 0. pulchcr was found by beating dead branches of oak.
Kraus (1908) bred Ennearthron fransvcrsatum from decaying
rattan vines. Perris (1877) described the larva of Cis coluber
from dead branches of chestnut and oak in which the fungus
LAWRENCE: CLASSIFICATION OF CIIDAE 287
Thclcphora was oTowing; he also noted that the beetle could not
be found on branches still remaining on the tree. In the same
paper, he noted that the habits of C. alni and C. reflexicollis
Abeille de Perrin were similar, and that C. ohlongus Mellie and
C. pruinosulus were taken on fungus-infested elm branches. Zim-
merman (1938, 1942) and Swezey (1954) have recorded a num-
ber of Pacific island species from dead vines and branches, and
those which I have examined seem to fall within this same group.
One fact that makes the study of the particular group more
difficult is that most of the shared characters are both simple
and apparently primitive. Most of the special modifications
which are characteristic of the family as a whole, such as the
development of protibial structures for boring (expanded apices,
teeth, combs, spines), various prosternal modifications (shorten-
ing, carination, reduction of intercoxal process and increased
coxal size), presence of sexual ornaments on the head and pro-
notum of the male, and the development of more complex puncta-
tion and vestiture, are all absent in the species of OrtJiocis. In
addition, the generalized fungus-feeding habit may be considered
primitive. Members of closely related families, such as the
Lathridiidae and Corylophidae, share this habit with the species
of Orthocis, and Crowson (1955) thinks that this may have been
characteristic of the ancestral Cucujoidea. If these characters
are primitive or plesiomorphic, they are, according to Hennig
(1965), less reliable than derivative or apomorphic ones in deter-
mining relationships. If, on the other hand, Orthocis represents
a collection of two or more convergent groups, which have
secondarily developed this type of feeding habit and thus have
lost, through disuse, the structural modifications associated with
boring, this convergence would probably be difficult to detect
because of the simplicity of the resulting characters. I favor the
hypothesis that these characters are primitive, rather than
derivative, and that the feeding habits probably represent those
of the common ancestor of the group. In certain other species,
such as Hadraiile elongaiula and Maphoca hlaisdelli, which have
similar biologies, this simplified condition also occurs, but other
characters indicate that both of these are derivative forms,
which may have evolved from a species in the Cis comptus group.
I think that most of the species discussed above should be
placed in a distinct genus, to which the name Orilwcis applies.
The exact limits of the genus remain uncertain, especially with
reference to the inclusion of Ch festivns and its relatives, and
288 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
it is hoped that Dr. Lohse will shed some light on the relation-
ships of these European species. I would definitely exclude Cis
angustus Hatch, which does have a truncate protibial apex, but
which occurs on Fomes pinicola, has clypeal tubercles in the male,
and lacks a margin on the elytral suture.
The species of Dolichocis, although they share certain char-
acters with Orthocis, should remain in a separate genus, differing
by the absence of the margin on the elytral suture, the anteriorly
constricted prothorax, cylindrical form, and vestiture of short,
stout setae. The genus should include the species laricinus (Mel-
lie), yuasai (Chujo), mayiitoha Dury and indistinctus Hatch.
All four species occur on the larger, woody fruiting bodies of
fungi, such as Fomcs pinicola and F. officinalis. Synonymies and
redefinitions of Orthocis and DolicJiocis are given below.
Orthocis Casey
Orthocis Casey, 1898, Jour. New York Eiit. Soc, 6(2) :84; Kraus, 1908:77;
Dalla Torre, 1911:20; Dury, 1917:13; Leng, 1920:247; Brethes,
1922:302. Type, by present designation, Orthocis aterrima Casey,
1898:84.
Cis (Mellieicis) Lohse, 1964, Ent. Blatter, 60(2) :122. Type, by original
designation, Cis alni Gyllenhal, 1813:386. NEW SYNONYMY.
Enncarthron, Kraus, 1908:78.
Cis (in part), auctt.
Form elongate and somewhat depressed to narrowly elongate
and cylindrical; vestiture of very short fine hairs or stouter
bristles. Head slightly to moderately declined, only slightly cov-
ered by pronotum; frontoclypeal ridge without distinct sexual
modifications in male ; antennal fossa shallow. Antenna 9- or 10-
segmented, with 3- segmented club, segments III and IV usually
elongate ; maxillary stipes elongate, laeinia lateral, palp rela-
tively stout ; prementum elongate. Pronotum subquadrate, sides
narrowly margined to broadly margined and explanate, anterior
angles truncate to slightly produced and rounded ; anterior edge
simple in both sexes. Elytra usually parallel sided; punctation
single and relatively uniformly distributed; suture margined
posteriorly, the margin curved laterad just before apex. Pro-
sternum flat to slightly tumid, longer than intercoxal process
which is fairly broad ; procoxae subtransverse, narrowly open
behind. Protibia only slightly expanded at apex, outer apical
angle truncate or rounded. Metasternum slightly convex, the
suture moderately long. Sternite III of male with a median
pubescent fovea or patch.
LAWRENCE: CLASSIFICATION OF CIIDAE 289
A large, widespread g:emis, but mainly tropicopolitan. Closely
related to Cis and Dolichocis, from which it differs by the
rounded protibial apices, margined elytral suture, and lack of
distinct sexual ornaments in the male. The species appear to be
general fungus feeders, occurring often on dead vines and
branches penetrated by fungus mycelia.
Dolichocis Dury
Dolichocis Dury, 1919, Canad. Ent., 51:158; Hatch, 1962:234. Type, by
monotypy, Dolichocis manitoha Dury, 1919:158.
Cis (in part), Mellie, 1848:236; Lacordaire, 1857:551; Jaequelin Du Val,
1861:237; Seidlitz, 1872:44; Kiesenwetter, 1877:178.
Ennearthron (in part), Abeille de Perrin, 1874:80; Eeitter, 1878:30; Seid-
litz, 1891:285; Schilsky, 1900 :37B; Eeitter, 1902:59; Dalla Torre,
1911:23; Chujo, 1941:85.
Form elongate, cylindrical; vestiture of short, stout, suberect
bristles or squamae. Head moderately declined, partly covered
by pronotum ; frontoclypeal ridge of male bituberculate, vertex
simple or foveate ; antennal fossa shallow. Antenna 9-segmented,
with 3-segmented club ; maxillary stipes subquadrate, lacinia
subterminal, palp relatively stout. Pronotum almost as long as
wide, somewhat constricted anteriorly, sides narrowly margined,
anterior angles not or barely produced ; anterior edge simple in
both sexes. Elytra elongate and subparallel ; punctation single
and fairly uniformly distributed. Prosternum slightly tumid,
slightly longer than intercoxal process which is fairly broad and
blunt at apex ; procoxae subtransverse, narrowly open behind.
Protibia only slightly expanded at apex, outer apical angle
truncate or rounded. Metasternum slightly convex, suture mod-
erately long. Sternite III of male with a median pubescent
fovea.
A small genus restricted to the Holarctic region. Closely re-
lated to Cis and Orthocis, from which it differs in the rounded
protibial apices, stout vestiture, constricted prothorax, and lack
of a margin along the elytral suture. The 4 species all occur on
Fomes pinicola and related fungi.
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November, 1005
Bull. Mus. Comp. Zool., Harvard Univ., 133(6): 295-335, November, 1965.
No. 6 — The Fossil Elephant Shrews (Family Macroscelididae)
By Bryan Patterson
INTRODUCTION
The history of this peculiar group of mammals was osteusibly a
blank until 1937, when Broom described an extinct species of
Elephanfulus from the Pleistocene of South Africa. Four extinct
genera, however, had been described earlier, from 1910 on, but,
having been placed incorrectly in other groups of mammals, had
gone unrecognized. One of these, Palacolhenfoides Stromer, has
been detected as a macroscelidid by Butler and Hopwood (1957);
the other three, Metoldobofes Schlosser, Myohyrax Andrews and
Protypotheroides Stromer are here placed in the family for the first
time. Palaeothentoides was originally described as a marsupial,
Metoldobofes as a mixodectid insectivore, and Myohyrax and
Protypotheroides as hyracoids. So extraordinary a situation is, I
believe, without parallel in mammalian paleontology.
This paper came into being in a rather roundabout way. Crea-
tures with names such as Palaeothentoides and Protypotheroides
have a certain fascination for anyone with a taste for South
American fossil mammals. Being in Europe during the spring of
1957, I took the opportunity of visiting Rlunich and examining
material of both forms. The collection included an important
undescribed specimen of Palaeothentoides africanus, and this I was
\Try kindly permitted to borrow for further study. Shortly after
completing the description of it, and with the details of macro-
scelidid dental structure fresh in mind, I had occasion to consult
Schlosser 's memoir (1911) on Fayum mammals during a discussion
of the dental formula of Parapithecus. There, staring up from the
plate, was Metoldobotes, an ob\-ious macroscelidid. (Discussions of
Parapithecus are perennial ; it is gratifying to be able to report this
useful by-product of one of them.) On examining Protypotheroides
beetzi in Munich I had noted a decided resemblance to Palaeo-
thentoides, but at the time had considered it to be an interesting
example of convergence, one about on a par with the resemblance
to the interatherid typotheres. It was only after consulting Whit-
worth's study (1954) of the Miocene hyracoids of Kenya in quite
another connection that I began to suspect the Myohyracinae
really were macroscelidids, a suspicion that hardened to conviction
upon examination of Stromer's figures (1926) of postcranial
remains from the Miocene of Southwest Africa. Here was a group
298 bulletin: museum of comparative zoology
of elephant shrews that had masqueraded as ungulates for half a
century. Broom's (1948) Mylomygale spier si from the Pleistocene,
a form with hypsodont, rodent-like cheek teeth, revealed the
former existence of yet another phylum of the family. It became
clear that the surviving forms gi\'e little hint of a rather remark-
able radiation that went on within the African continent through-
out much of the Cenozoic. There is nothing remotely primate-like
about the extinct phyla so far known. The fossil record in fact very
definitely supports those who ha\'e been unable to accept the
hypothesis of a close relationship between the elephant shrews and
the trecshrews. As LcGros Clark has well put it (1959, pp. 318-
319n): "In retrospect it is difficult to understand this taxonomic
association . . .the differences . . . are so marked as to make it
clear that they are really quite divergent types."
This study has been aided by National Science Foundation
Grants G-3120 and GP 1188, which made it possible for me to
examine specimens in Europe and in Kenya. For access to material
I am indebted to the authorities of the British Museum (Natural
History) and of The American Museum of Natural History, to
Dr. Richai'd Dehm and to Dr. L. S. B. Leakey. Miss Margo
Hayes has assembled the final manuscript and assisted in checking
localities and references. The photographs are by Mr. Frank
White and the drawings by Mrs. Dorothy Marsh. Figures of
fossil macroscelidids are widely scattered in the literature and a
number of them are in rather rare publications. Those pertinent
to the work are accordingly redrawn here. Publication has been
aided by National Science Foundation Grant GB-500.
TAXONOMY AND MORPHOLOGY
MACROSCELIDIDAE
IMACROSCELIDINAE
MetOLDOBOTES Schlosser
Metoldobotes Schlosser, 1910, p. 507; Matthew, 1910, p. 702.
Metolbodotes Schlosser, 1911, p. 70; Matthew, 1915, p. 467.
Type species: M. stromeri Schlosser, 1910.
Distribution: Early Oligocene, north Africa.
Emended diagnosis: I3 (?), C, P4, M2. I3 not bifid, with pro-
nounced vertical groove on lingual side; C bluntly pointed, three
sided; Pi_3 short relative to P4 — M2; Pi single rooted, roots of P2
very closely appressed, P3 with small anterior and posterior cusps,
without posterior accessory cusp; P4 and lower M with crista
PATTERSON : FOSSIL ELEPHANT SHREWS 299
obliqua running to posterior face of trigonid, talonid of M3 short
relative to trigonid. Horizontal ramus of mandible relatively short,
deep; ascending ramus steep; symphysis long, extending to P3.
]\Ietoldobotes stromeri Schlosser
(Fig. 1 c, d)
Metoldobotes slronicri Schlosser, 1010, p. 507; Matthew, 1910, p. 702.
Metolbodotes stromeri Schlosser, 1911, pp. 70-72, 147, 157, 163, 164; pi. 9, fig. 5.
Type: An incomplete right horizontal ramus in the Stuttgart
collections, with I3, C, P3 — M2, alveoli for I2, Pi, roots of P2.
Hyyodigm: Type only.
Horizon and locality: Fluviomarine series, early Oligocene;
Fayum, Egypt.
Diagnosis: As for the genus. The fragmentary type specimen
appears to represent an animal somewhat larger than Rhynchocyon
peter si and Protypotheroides beetzi, and hence the largest known
member of the family.
Discussion: Metoldobotes, with the exception of brief comments
by Matthew and passing mentions in various editions of Zittel,
has remained essentially unnoticed in the literature since its
description. Schlosser tentatively assigned it to the JN'Iixodectidae,
a reference which, as Matthew (1915, p. 467) stressed, had nothing
to recommend it in the way of positive resemblances between the
Fayum form and any mixodectid.
The type ramus, incomplete anteriorly, preserves, in series, an
anterior alveolus, an incisor, a partially erupted, conical tooth, the
ah-eolus of a single-rooted tooth, the very closely appressed roots of
a double-rooted tooth and the last four cheek teeth. Schlosser
interpreted this array as Ii,2,3, C, P3,4, Mi,2,3, but he made no
comparisons with any macroscelidid. Inspection of his figures at
once reveals an impressive number of resemblances to the various
members of this family, and suggests that the dental formula is in
reaUty 1(0,2,3, C, Pi,2,3,4, Afi,2, as in Rhynchocyon, Petrodromus,
Elephantulus, Macroscelides, and Mylomygale (an alveolus for Ii in
this interpretation is lacking, but the specimen is incomplete
anteriorly). In the ensuing remarks the teeth will be so designated.
The crown of 1 3 is described by Schlosser as being about half the
height of the root; this is true of the incisors of macroscelidines.
A grooN'e is present on the lingual faces of the incisors of members
of this subfamily, although in no case is it as pronounced as shown
in Schlosser 's figure. The canine in Rhynchocyon is sometim3 5
(e.g. MCZ 38782), although by no means invariably, not fully
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erupted until after the posterior cheek teeth have come into wear.
Pi is single-rooted in Macroscelides, Mylomygale, and in some
species of Elephant iilus^. The roots of P2 are closely appressed in
Macroscelides. P3 of Mefoldohotes resembles that of all members of
a
Figure^ 1. r, d, Meioldobotes slronicri, ciowii view of (ifiititioii ;inil lateral
view of iiKUitliWle, c X3, d x|; redrawn from Sclilosser. «, I'etrodroiiius
nigriseia, MCZ 22434, 6, Rhi/nchoei/on petersi, MCZ 22573, ert)wn views of
P3 - Ml, not to scale.
' Among the Elephantulus material in the Museum of Comparative Zoology collections I
have found Pi to be single rooted in E. o-ularis (29 specimens), ruftsceris (•'>), and pulchtr (ti);
variable in rupestris (3 single and 3 double); and double rooted in intufi (5). and ;i/sopes (2).
PATTERSON: FOSSIL ELEPHANT SHREWS 301
the family in its proportions relative to P4 — M2 and in its posses-
sion of low anterior and posterior cusps, and that of the macrosceH-
dines in the absence of an accessory cusp on the posterior slope of
the protoconid. The large P4 has a wide molariform talonid and a
narrower, elongate trigonid. The tooth is typically macroscelidid
in these features, and also as regards size relative to the molars and
the structure and arrangement of the cusps and crests. The
molars of Metoldobofes resemble those of the living macroscelidids
in the nearly equal heights of the trigonids and talonids, the
metaconids and entoconids higher than the protoconids and
hypoconid, the absence of labial and lingual cingula and the struc-
ture and positions of cusps and crests generally. The paraconid is
median, as in the macrosceUdines. The crista obhqua runs to
the center of the posterior face of the trigonid, as in the Miocene
Rhynchocyon clarki (Butler and Hopwood, 1957, p. 10), rather than
to the metaconid, as is the case in living forms. The relative
lengths of Mi and M2 are approximately as in Rhynchocyon, and
the talonid of AI2 appears to be about as small and short relative
to the trigonid as in R. cirnei (e.g. MCZ 43735). In agreement with
Rhynchocyon and the macroscelidines, the mental foramina are
beneath Pi_2 and P4; the ascending ramus, to judge from Schlosser's
figure, arises abruptly well behind the last molar. The masseteric
fossa is shallow and, as in macroscelidines, extends down to the
level of the tooth row. The horizontal ramus is deeper than in
either the Rhynchocyoninae or the Macroscelidinae, shallower
than in the hypsodont Myohyracinae. That part of the tooth row
anterior to P4 is somewhat shorter relative to the length of the
series as a whole than in living members of either of the first two
subfamilies, but is approximately comparable to Myohyrax and
Mylomygale in this respect. The symphysis, fide Schlosser, extends
to P3 (his P4) and is hence longer than in all other known members
of the family, in which it terminates beneath C or Pi.
On the evidence available it is difficult to assign Metoldobotes to
subfamily with any confidence. The Fayum form does not, even
incipiently, display any of the specializations of myohyracines or
mylomygalines. It does resemble both the Rhynchocyoninae and
the Macroscelidinae, agreeing with one or the other now in this
character, now in that, and differing from both of them in the long
symphysis. Resemblances to Rhynchocyon — and differences from
the macroscelidines — are the small size of the talonid of J\l2 rela-
tive to the trigonid and the small size of M2 as a whole relative to
Ml. M. stromeri resembles the Miocene R. clarki in that the crista
obliqua runs only to the posterior face of the trigonid and not to the
302 bulletin: museum of comparative zoology
apex of the metaconid, but this is interpretable simply as a primi-
tive character possessed in common. Resemblances to the ]\Iacro-
sceUdinae, or at least to some of them, are the lack of a posterior
accessory cusp on P3, the small single-rooted Pi, the closely
appressed roots of P2, the median position of the paraconid in the
molars, the lingual groove of the incisor, and the steeply rising
ascending ramus. In sum, the characters suggest relationship with
the macroscelidines rather than with the rhynchocyonines, and I
very tentatively place Metoldobotes in the Macroscelidinae.
PalAEOTHENTOIDES Stromer
Palaeothentoides Stromer, 1932, p. 185.
Type species: P. africanus Stromer, 1932.
Distribution: Early Pleistocene?, southwest Africa.
Emended diagnosis: Lower postcanine formula P4, M3; Pi two-
rooted, not incisiform; P2_3 with anterior cusps little separated from
protoconids, P3 without metaconid and entoconid rudiments; P4
narrow, metaconid decidedly posterointernal to protoconid,
reentrant valley between metaconid and entoconid nearly filled by
swelling on crest running anteroexternally from entoconid, para-
conid crest high, anterointernal sAvelling partially obhterating cleft
between anterior crest and metaconid; Mi_2 with very slight,
shallow clefts between paraconids and metaconids, sides of deep
reentrants between metaconids and entoconids parallel, not
ventrally converging; horizontal ramus of nearly even depth
beneath cheek teeth, shghtly downcurving anteriorly.
Palaeothentoides africanus Stromer
(Fig. 2; PL 1)
Palaeothentoides africanus Stromer, 1932, pp. 178-185, figs. la-2b; Butler and
Hopwood, 1957, p. 11.
Type: Munchen No. 1931. VII. la, left ramus with P3 - M2,
alveoli for Pi_2, M3.
Hypodigm: Type, and Munchen Nos. 1931. VII. lb, fragment of
left ramus with M2-3 (now lost), and 1932. I. 501, left ramus with
Pi - M3.
Horizon: The "intermediate terrace" of Wagner and Merensky
(1929, p. 29, fig. 5); age uncertain, possibly early Pleistocene.
Stromer (1931, p. 41; 1932, p. 185) considered this to be ''wohl
Mittelpliocan," which would now, with the transfer of the Villa-
franchian to the Pleistocene, be regarded as late Phocene. The
two other forms definitely identified by Stromer from this level,
PATTERSON : FOSSIL ELEPHANT SHREWS 303
Enhydriodon and Hijaena (especially the latter), are not incon-
sistent with a Pleistocene age.
Locality: Klein Zee (or Kleinsee), near the mouth of the Buffels
River on the coast of Little Namaqualand, some 25 miles SSE of
Port Nolloth, Union of South Africa.
Diagnosis: As for the genus. Comparable in size to the smaller
hving macroscelidines.
Description: The first premolar is a long, narrow, double-rooted
tooth with a simple crown consisting of a procumbent protoconid,
which extends forward beyond the anterior root, connected by a
crest to a smaller posterior cusp. The lingual face of the tooth is
very slightly convex, the labial vertically grooved between the
cusps. P2 bears a small cusp on the anterior slope of the erect
protoconid; labial and lingual grooves are present anterior to the
protoconid, and the labial groove between protoconid and posterior
cusp is much larger and deeper than in Pi. All these features are
accentuated in P3 : the anterior cusp is larger, the grooves deeper,
and the posterior cusp larger and wider; there is no metaconid or
entoconid rudiment. These three teeth progressively increase in
length and height. P1-3 of Palaeothentoides are very similar to the
corresponding teeth of Nasilio, differing from those of the other
living genera in various particulars. Thus, in Macroscelides, Pi is
single-rooted and similar in structure to I2 — C, while P2-3 are
higher crowned relative to length ; in Elephantulus, Pi has a higher
protoconid, P2-3 have the anterior cusps well separated from the
protoconid, P3 has metaconid and entoconid rudiments, and a
rudiment of the metaconid is occasionally seen on P2; in Petro-
dromus and Rhynchocyon, all three teeth are higher, more piercing
(especially the caniniform Pi of the latter), anterior cusps are
either lacking entirely (Rhynchocyon) or rudimentary {Petro-
dromus), while the posterior cusps are very small in both and, in
Rhynchocyon, confined to P3.
P4, the longest of the cheek teeth, is submolariform, the talonid
completely as in Mi_2, the narrow trigonid not. The entoconid
is fully as large as in the molars and, as in them, higher than the
hypoconid. In the trigonid, the anterior cusp is set off labially
from the protoconid by a vertical groove wider and shallower than
the corresponding ones on the anterior premolars. The groove be-
tween these two cusps on the lingual side is shallow and partially
filled by a buttress on the side of the ridge connecting them. The
metaconid is well developed, nearly as high as the protoconid and
decidedly posterointernal to it in position ; the short crest between
the cusps bears a small, shallow groove on its lingual side. The
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crista obliqua runs to the apex of the meta-
conid. The labial reentrant between trigonid and
talonid is fully as large and deep as in Mi_2, but
the lingual is to a great extent filled by a vertical
swelhng on the side of the ridge running from the
entoconid to the crista obliqua. The sides of this
lingual reentrant are nearly parallel for most of
their heights and converge to form a U only near
the base of the enamel. The tooth continues the
progressive increase in crown height seen in Pi_3.
Although thoroughly macroscelidid in structure,
P4 is the most distinctive tooth of the series.
None of the hving forms has the labial reentrant
between trigonid and talonid nearly filled by a
swelhng, and in none is the metaconid so far
posterointernal to the protoconid. In all except
Macroscelides the anterior crest and the meta-
conid are widely separated by a deep groove and
the sides of the lingual reentrant converge toward
the base, forming a V.
Mi_2 consist essentially of two triangular pris-
matic columns connected by the narrow isthmus
formed by the crista obhqua. The protoconids,
hypoconids and paraconids are angulate, the
metaconids and entoconids more rounded. The
paraconids and metaconids are separated by very
shallow vertical grooves. The lingual and labial
reentrants are large, deep, and extend nearly to
the base of the enamel; their sides are parallel
and U-shaped below. The trigonids are wider
and larger than the talonids, particularly on Mi.
The metaconids and entoconids are higher than
the protoconids and hypoconids, and there is a
shght indication of a hypoconuhd. This cusp may
have been larger on the unworn crown, if we may
judge from Macroscelides in which it is very
prominent on unerupted molars but rapidly be-
comes worn away. On the hngual sides of the
crests running forward from the entoconids are
faint vertical swelhngs corresponding to the
Figure 2. Palaeothentoides africanus, dorsal view of mandible, Munchen
No. 1932. I. 501. X8.
PATTERSON : FOSSIL ELEPHANT SHREWS 305
prominent one in P4. These molars are the highest crowned of the
cheek tooth series. M3 is vestigial and much lower crowned than
M2. It is composed of the trigonid only, on which the small
metaconid is the highest element and the protoconid is sub-
ordinated in the paraconid crest; metaconid and paraconid are
separated lingually by a shallow depression, and there is a slight
vertical ridge on the posterior face, the last vestige of the talonid.
The enamel is continuous on all cheek teeth ; it is thick on P4 — M2
although thinning at the paraconids in the molars. M3 is indis-
tinguishable from that of Nasilio. Mi_2, on the contrary, are quite
different from those of this form and very close indeed to those of
Macroscelides, differing only in their slightly lower crowns and, at a
corresponding stage of wear, in the presence of enamel around the
paraconids. In the other living forms, Mi_2 are somewhat lower
crowned, have wide lingual grooves between the paraconids and
metaconids, and V-shaped sides to the lingualr eentrants separating
trigonids and talonids.
Seen from above, the ramus curves very slightly inward from
P2 forward and gently outward from M3 backward. There is no
trace of the symphysis on the part preserved, indicating that, as in
all living macroscelidids except Petrodromus, this did not extend
posteriorly beyond the level of the canines. The inner face, as
noted by Stromer, is nearly flat, the outer swells out gently oppo-
site the molars. As in other forms, there is a posterior mental
foramen beneath P4 and an anterior beneath Pi. The height of the
horizontal ramus remains rather constant beneath the cheek teeth,
decreasing less anteriorly than in the living forms. The ventral
border is gently convex beneath the molars and P4, and shows a
more marked tendency to turn down beneath Pi_2 than in any
other form. As is usual in the group, the ascending ramus begins
to rise well behind the last molar and the masseteric fossa is
shallow and poorly defined.
Discussion: Palaeothentoides is unquestionably a valid genus. It
resembles Nasilio and Macroscelides, combining characters of both,
and can be referred with assurance to the Alacroscelidinae. There
is no need to belabor the fact that this form is no marsupial. A
resemblance does exist, particularly in the trigonid, between P4 of
Palaeothentoides and ^Ii of Palaeothentes, but this is far from exact.
The labial and lingual reentrants separating trigonid and talonid
that are large and deep in the African form are, for example,
shallow in the South American one, and in any event the teeth
Stromer compared are not homologous.
P4
Ml
Mo
Ma
2.8
2.4
2.1
0.8
1.4
1.7
1.5
0.8
306 bulletin: museum of comparative zoology
Stromer himself realized almost at once that Palaeothentoides
was not a marsupial. A separate of his paper in my possession
bears a tj^pewritten shp reading: "Berichfigung: Durch einen neuen
Fund ist erwiesen, dass Palaeothentoides 4 Pin and 3 M hat, also
sicher kein Didelphier ist." So far as I am aware, however, he
never published this retraction. The "new find" referred to is of
course No. 1932. I. 501, here described and figured for the first
time.
Measxirements, in mm, of Mi'mchen No. 1932. I. 501.
Pi P2 P3
Length 1.4 2.0 2.3
Width 0.6 0.7 0.9
Length Pi - M3, 14.0; Pi_4, 8.6; Mi_3, 5.6.
Height of ramus beneath Mi, external, 2.9.
ElephantuLUS Thomas and Schwann
Elephantulus Thomas and Schwann, 1906, p. 577.
Elephayitomys Broom, 1937, p. 758.
In the course of describing the Pleistocene E. langi, Broom
observed that certain species of the genus, langi among them, have
a molariform P-. Believing that E. rupestris, the type species,
lacked this character, he proposed Elephantomys, with E. langi as
type, for the reception of those species possessing it. Shortly
thereafter, specimens from what he supposed to be the type
locality of E. rupestris having come to hand, he concluded that
this species did after all have a molariform P-, a fact which in his
opinion effectively suppressed Elephantomys. He did not go on to
erect a new genus for those forms with a non-molariform P'.
Elephantomys was subsequently revi^'ed by Ellerman, Morrison-
Scott and Hayman (1953, p. 8). Stating that P- of the type of
E. rupestris was non-molariform, they recognized Elephantomys as
a subgenus, distinguishing it on the basis of the molariform P- and
the possession of less flattened bullae, in which the lateral (i.e.
tympanic) portion is higher relative to the median (i.e. ento-
tympanic) than in Elephantulus^ . These distinctions are not valid.
As regards P-, what is involved is a forward extension of the
molarisation field. In Rhynchocyon and Petrodromus this extends
hardly or not at all beyond P^; in Nasilio and Macroscelides P^-s
have been incorporated to the degree that two lingual cusps are
1 They also reduced Nasilio to the rank of a subgenus of Elephantulus. This does not seem
justified. In addition to possessing Ms, Nasilio has higher-crowned posterior cheeli teeth and
different hind limb proportions (Evans, 1942).
PATTERSON: FOSSIL ELEPHANT SHREWS 307
present on them. Elephantulus is in a state of flux. Among the
material available to me, P^ is molariform in intufi (5 specimens)
and fuscipes (2), non-molariform in rupestris (5) and pulcher (7).
Although predominantly non-molariform in ocularis (29) and
rufcsccns (6), it is nevertheless \'ariable in these species, even
within what are surely local populations. Thus, in rufescens from
Mt. IMbololo, Kenya, of two specimens collected on the same day,
one (MCZ 31800) has two lingual cusps on P^, another (MCZ
31802) one. Within ocularis, a small series from Unyanganyi,
Tanganyika, Tanzania, includes two specimens (MCZ 25660 and
25683) with two lingual cusps on this tooth and three with one; a
specimen from Dodoma, Tanganyika (MCZ 22841), has two
hngual cusps on the left side and one on the right. Within the
"non-molariform" species, P''' as well as P- is variable in respect of
lingual cusp development. A distinction based on bulla structure
cuts across one based on premolar structure. Thus the "non-
molariform" pulcher has a "flattened" bulla, and the predomi-
nantly "non-molariform" rufescens and ocularis have "less
flattened" ones. Elephantomys does not merit recognition.
A far reaching proposal for a division of Elephantulus, and
indeed of the whole subfamily, has been advocated by Van der
Horst, who, with co-workers, devoted many years to study of the
embryology of the genus, with particular reference to E. myurus
jamesoni (I employ Van der Horst's names in this paragraph). In
the course of his work there emerged the remarkable facts that in
this form approximately sixty eggs are liberated and approximately
sixty corpora lutea develop in each ovary, only one of which be-
comes implanted, the Graafian follicle is remarkably small, and
fat globules are lacking in the ova. Macroscelides prohoscideus was
found to agree in all these particulars, and E. capensis in all save
for the presence of a few fat globules. E. intufi and E. rupestris
stand in striking contrast. In these species only two eggs per ovary
are liberated, the Graafian follicle is of normal type, and fat
globules are present. Petrodromus tetradactylus is in agreement with
them except for the apparent absence of fat globules. On the basis
of all this. Van der Horst has suggested (e.g. 1944) that there are
only two genera of macroscelidines and that the division passes
through the genus Elephantulus of current usage. He has not
spelled out what would result were his suggestion to be adopted,
but this can be simply put. We would have two genera: Macro-
scelides, with prohoscideus, myurus and capensis, and Petrodromus,
with tetradactylus, rupestris and intufi; all other species would have
to remain in limbo until comparable investigations had been
308 bulletin: museum of comparative zoology
carried out on them. If these characters were indeed the touch-
stone of macroscehdine systematics such a situation would be
acceptable, but there is no real evidence that they are. The genera
recognized by mammalogists over the years are clear-cut taxa,
distinguishable by different combinations of characters. The
sporadic occurrence of the curious ovarian characters suggests that
these were independently acquired, or, alternatively, perhaps lost,
at various times within the group.
This possibility is reenforced if the classification, other than
subgeneric, of the South African species and subspecies of Ele-
phantuhis proposed by Ellerman, Morrison-Scott and Hayman is
correct. Van der Horst's myurus jamesoni is their rupestris
jamesoni, his capensis is their rupestris capensis, his intufi and
rupestris may be their rupestris and intufi. His division of the
subfamily would thus run between subspecies of rupestris in their
arrangement. The genetic basis of the ovarian peculiarities may
be of a rather simple sort.
Elephantulus langi (Broom)
Elephaniomys langi Broom, 1937, pp. 758-760, fig. 5.
Elephantulus langi Broom, 1938, p. 251; 1948, p. 5.
Horizon: Pleistocene.
Locality: Cave deposit at Schurveberg, 15 miles west of Pretoria,
Transvaal, Union of South Africa.
E. langi is evidently represented by rather rich material from the
Schurveberg cave deposit, and Broom's description is of the most
preliminary sort. The relationship between langi and living forms
remains to be determined.
Elephantulus antiquus Broom
Elephantulus antiquus Broom, 1948, pp. 5-6, fig. 3.
Horizon: Earlier Pleistocene.
Locality: Bolt's worldngs, Sterkfontein, Transvaal, Union of
South Africa.
E. antiquus is evidently distinct from E. langi — it has, e.g.,
a non-molariform P^ — but little more can be said. As in the case
of E. langi, there is fairly abundant material, Broom's description
is preliminary and incomplete, and the relationship to hving
species is unknown. In 1946, Broom {in Broom and Schepers,
p. 78) stated that: "The elephant shrew Elephantulus langi, or one
very closely allied, occurs in the Plesianthropus cave. The type is
from Schurveberg, Pretoria. It is common at Bolt's workings."
Presumably the species there referred to is E. antiquus.
PATTERSON: FOSSIL ELEPHANT SHREWS 309
Elephantulus rozeti (Duvernoy)
From an archaeological site at Redeyef, Tunisia, Gobert (1912)
recorded the presence of various genera of mammals, most of
which he believed to be referable to living species. Macrocelides
(sic) is among those listed. Three cultural levels occur at the site,
the two lower Paleolithic and the upper ranging from transitional
to Neolithic. Gobert gave no description of the mammalian
remains, which it would appear from the text were found in the
upper level. Romer (1928, pp. 100, 153, 161) lists this find as
Macroscelides rozeti. A little uncertainty attaches to the determi-
nation. Thomas (1901, 1913) split North African Elephantulus
into two species, E. rozeti and E. deserti, the former with three
and the latter with two subspecies. E. deserti is the more eastern
of the two and, if valid, the Redeyef material might therefore be
referable to it. More likely than not, however, subspecific distinc-
tion, at most, is involved. Although Thomas had stated, in 1901,
that deserti did not differ in size from rozeti, he claimed, in 1913,
that it was smaller; the very few published measurements do not
support the assertion. The differences appear to be confined to
pelage color.
Elephantulus rozeti, which dates from 1838, was long known as
Macroscelides rozeti, and numerous specimens so labeled found their
way into collections. Many of the labels were not changed when
Thomas and Schwann transferred rozeti to their new genus Ele-
phantulus. These labels have trapped trusting anatomists and
paleontologists. A number of accounts and illustrations in the
literature that purport to be of Macroscelides are actually of
Elephantulus, based on E. rozeti (e.g. Evans 1942, Fiedler 1953,
Grasse 1955, Saban 1956-1957, Van der Klaauw 1929, in part).
RHYNCHOCYONINAE
RhyncHOCYON Peters
Rhynchocyon clarki Butler and Hopwood
R. clarki Butler and Hopwood, 1957, pp. 4-11, figs. 2-3.
Horizon: Early Miocene.
Localities: Type from Songhor local fauna, Kenya; referred
material from Rusinga Island, Kenya, found in the upper and
lower Hiwegi beds and either in the Kiahera or in the lower part
of the Kathwanga beds.
This species, so well described by its authors, reveals, as they
point out, that the two surviving subfamilies had diverged prior to
310 bulletin: museum of comparative zoology
the Miocene. R. clarH ". . . as an early member of the Rhynchocyon
Hneage . . . is . . . nearer to the common ancestor of the two
groups, and possesses a number of primitive characters which have
been lost in Recent representatives of both subfamiUes." The
species is notably smaller than the living members of the genus,
which suggests a relatively recent increase in size within the
Rhynchocyon lineage.
MYLOMYGALINAE subfam. nov.
Diagnosis: I3, C, P4, M2. I1-3 small, subequal; C — P2 small,
single rooted; P4 — M2 large, hypsodont, crowns complex, flat, of
grinding type. Molars compressed anteroposteriorly, as wide as
long; protoconid and hypoconid angulate, directed antero-
externally; hypoconid and hypoconulid forming posterior lophid,
entoconid set off from posterior lophid by deep reentrant; re-
entrant between paraconid and metaconid situated on anterior
face of tooth; M2 large relative to Mi. Talonid of P4 fully molari-
form, trigonid larger, more elongate than in molars. Ventral
border of horizontal ramus strongly convex, alveolar border con-
cave beneath posterior cheek teeth.
Mylomygale Broom
Mylomxjgale Broom, 1948, p. 6.
Type species: M. spiersi Broom.
Distribution: Earlier Pleistocene, south Africa.
Diagnosis: Sole known genus of the subfamily diagnosed above.
Mylomygale spiersi Broom
(Fig. 3)
Mylomygale spiersi Broom, 1948, pp. 6-8, fig. 4, (1946, in Broom and Schepers,
p. 28, fig. 1 N-Qi).
Locality: "... a small cave about half a mile to the north of the
cave which yielded the Taungs man-ape skull." Approximately 80
miles N. of Kimberly, Bechuanaland, Union of South Africa.
Horizon: EarUer Pleistocene. ("This bone breccia is probably
of approximately the same age as [those in] the other caves.")
Diagnosis: Sole known species of the genus.
Discussion: This remarkable little macrosceUdid enjo3'^s the
distinction of being the only extinct genus correctly placed in the
' In this paper M. spiersi was figured and listed with the statement that: "It represents a
new family of the Menotyphla." No diagnosis or description was given.
PATTERSOX: FOSSIL ELEPHANT SHREWS
311
family by its describer. Discovery of Mylomygale revealed the
existence of an otherwise unknown division of the family, one that
evolved posterior cheek teeth that are as strikingly rodent-like as
those of the myohyracines are ungulate-like. Broom's remark to
the effect that had the molars been found isolated they would have
been regarded as belonging to some peculiar hystricomorph rodent
is no exaggeration.
The type specimen preserves five small alveoli followed by four
grinding teeth. Although he decided that the dental formula was
probably I3, C, P4, M2, Broom was in some doubt as to whether
the last two alveoli housed the roots of two teeth or of one. "As in
all the living jMacroscelids the anterior premolars are double-
rooted it might seem more probable that the two sockets held a
single premolar, but on the other hand if the anterior premolar
were double-rooted then there can only be three premolars, while
all living Alacroscelids have four. ... I think it more likely that
there were two small single-rooted premolars." This tentative
conclusion was, I believe, the correct one. Contrary to Broom's
statement, and as pointed out above, the roots of Pi are fused in
Macroscelides, in some species of Elephantulus, and in the extinct
Metoldobotes, while the roots of P2 are closely appressed in Macro-
scelides. The anterior portion of the horizontal ramus is short in
Mylomygale, and it is hence not surprising that the roots of P2 had
fused.
Figure 3. Mylomygale spiersi, lateral view of mandible and crown view of
dentition, X4; redrawn from Broom.
312 bulletin: museum of comparative zoology
P3, the most anterior tooth preserved, bears the same size rela-
tion to P4, and P4 to the molars, as in other members of the family.
P4 and the two molars are remarkable for their deep, narrow, and
persistent reentrant folds. Comparison with an unworn molar of
Macroscelides (Fig. 5h) permits an understanding of the cusp
pattern. The anteroexternal angle is composed of the protoconid
and the centrally situated paraconid. Hypoconid and hypoconulid
are joined to form the posteroexternal and posterointernal angles
and the posterior lophid. The metaconid makes up the antero-
internal angle and the entoconid the central internal. This degree
of independence of the entoconid is a departure from the usual
macroscehdid condition, in which entoconid and hypoconulid tend
to be connected (cf. Fig. la, b). The large size of M2 relative to Mi
is another character peculiar to Mylomygale within the family; in
all other known genera, even in the earhest, Metoldobotes, it is
decidedly smaller than its predecessor in the series. The size of this
tooth provides an example of the reversal of an evolutionary trend,
the reversal in this case being associated \Yith. the later trend
toward the acquisition of rodent-hke posterior cheek teeth.
MYOHYRACINAE
(= Myohyracidae Andrews 1914, Myohyracoidea Stromer 1926)
(Figs. 4b; 5a, b, c, f, i, j ; 6a, c, e, g, i)
Emended diagnosis: Macroscehdidae with complete dental
formula; Ij"^ large, wdthout enamel on Ungual faces; posterior
cheek teeth hypsodont; M3 greatly reduced; F^f submolariform,
Pt essentially molariform; P^ - M^ with moderately undulant
ectoloph, paracones and metacones with comparatively shallow
labial grooves between them, parastyles and metastyles prominent,
parastyles anteroexternal in P"' - :\I2; P^ - M^ with persisting
fossettes, those of molars arranged in anterior and posterior pairs;
P3 — M2 with two fossettids, one each in trigonid and talonid;
horizontal ramus deep beneath posterior cheek teeth, mental
foramen beneath P3.
Distribution: Early Miocene, east and southwest Africa.
Genera included: Myohyrax Andrews 1914, Protijpotheroides
Stromer 1922.
Discussion: Andrews (1914, pp. 169-171) described Myohyrax
oswaldi on a fragment of a ramus with P3 — ^U and some isolated
teeth, including an upper molar; this material gave no hint of the
vestigial nature of M3. With such evidence in hand it would
hardly occur to anyone to make a comparison with the macro-
scehdids, and Andrews did not do so. He referred the genus to a
PATTERSON: FOSSIL ELEPHANT SHREWS
313
new family of the Hyracoidea. All subsequent students have
looked at myohyracines in this light and some of them have com-
mented on how aberrant they are within that order. From the
work of Stromer (1926) and of Whit worth (1954), it is possible to
note that many of the characters in which they differ widely from
hyracoids are actually points of resemblance to macroscelidids.
a
Figure 4. Lateral views of skull and mandible of, a, Nasilio brachyrhynchus,
MCZ 43755, and, b, Myohyrax oswaldi, slightly modified from Whitworth;
b X2, a not to scale.
314 bulletin: museum of comparative zoology
The skull of Myohijrax, figured in outline by Whitworth, is not
dissimilar to those of other members of the family (Fig. 4). The
facial region in all is long, low and rather narrow; the cranium is
short and high ; the zygoma arises over the rear of M^ ; and the
glenoid cavity is situated high on the side of the skull. Whitworth
shows a slight notch between nasal and premaxilla ; I was unable to
detect this in the specimen. The palate is unfortunately not
visible. The myohyracine mandible, if allowance be made for the
increased depth beneath the hypsodont cheek teeth, is decidedly
macroscelidid in appearance. The symphysis is short and shallow;
the ascending ramus high and steeply rising, and the coronoid
process small; the condyle is high and not expanded transversely;
and the angle is hook-like and extended posterodorsally.
p2 _ ]Vp of Macroscelides and Nasilio resemble the correspond-
ing teeth of Myohyrax in a number of respects. The posterior cheek
teeth of myohyracines are somewhat bowed outwardly (Whit-
worth, 1954, pi. 6, fig. 2), P- — M^ are incHned backward and P3 —
M2 are inclined forward; the bowing is incipient and the pitching
definitely present in Macroscelides. M- of myohyracines is unre-
duced, in correlation with the retention of M3 ; in Nasilio, in which
M3 is retained, the posterior portion of A'P is less reduced than in
the other living forms. The great reduction of M3 is, of course, a
decided resemblance to the macroscelidids and a striking contrast
to the hyracoids. The crown pattern of the upper molars of the
myohyracines (Fig. 5) is basically macroscelidid and not hyracoid
in such characters as the large, external paracone and metacone,
the absence of a mesostyle (in this I agree with Andrews and with
Hopwood, believing Whitworth 's mesostyle to be the paracone),
and the position and relations of the lophs. In macroscelidines,
especially Macroscelides and Nasilio, the lophs are relatively high,
the protoloph going to the parastyle, the robust metaloph prima-
rily to the paracone; the protocone is connected posteroexternally
to the enlarged anteroexternal portion of the metaloph and the
posteroloph is transverse, connecting metastyle and hypocone, all
very much as in myohyracines. Between paracone, protoloph,
metaloph and protocone anteriorly, and between metacone,
metaloph and posteroloph posteriorly, two fossettes are isolated.
These are aligned anteroexternally to posterointernally, and are
the homologues of the anterior and posterior pairs of fossettes in
the upper molars of myohyracines. In unerupted or little worn
molars of Macroscelides tendencies toward division of each of these
fossettes into two may be seen.
Patterson: fossil elephant shrews
315
Figure 5. A comparison of myohyracine and macroscelidine cheek teeth.
Myohyrax oswaldi: a, P^ - M'; c, M^;/, dm^;;, dm4. Protypotheroides beeizi:
b, P2 - Ms; i, Ml. Macroscelides proboscideus: d, M' (unworn); e, M'; gr.dm^;
h, Ml (unworn); k, diUi. a, b, c, f, i redrawn from Stromer, j from Whitworth;
d, g, h, k MCZ 37022, e MCZ 37023. b X2; a, c, f, i X4; j X6.6; the rest not
to scale.
The crown structure of P4 and of the lower molars is close to that
of Palaeothentoides and Macroscelides (Fig. 5). The lingual re-
entrant between trigonid and talonid is less open than in these
forms, and this narrowing was probably brought about by a
swelling on the entoconid crest similar to but larger than that
present in Palaeothentoides. Macroscelides has fossettids in the
trigonids and talonids of unworn molars and these are closely com-
parable, although much shallower than those occurring in Protypo-
theroides.
316 bulletin: museum of comparative zoology
The fourth upper milk molar of Myohyrax (Fig. 5f), like the
permanent molars, is very similar to the corresponding tooth in
macroscelidines. The differences that exist between it and that of,
e.g., Macroscelides — parastyle less set off by grooves, more
prominent posteroloph, nearly straight lingual wall, protocone not
set off by an anterior groove — do not disguise the basic re-
semblance. Dm4 of Myohyrax (Fig. 5j), although considerably
worn, is again unmistakably macroscelidid in structure and quite
unlike that of hyracoids. As in all members of the family, it is very
long and low-crowned, with trigonid and talonid approximately
equal in length. The paraconid area is set off by external and
internal grooves from the large protoconid and the even larger
metaconid, which was almost certainly twinned as it is in Macro-
scelides. The paraconid was clearly anterocentral in position with
a short labial crest and a longer lingual one terminating in a para-
stylid. Lingual and labial reentrants between trigonid and talonid
are essentially as in Macroscelides. The hypoconid is very large;
the entoconid and hypoconulid have become united by wear, while
the groove between hypoconid and hypoconulid still persists. With
wear this would occur in Macroscelides. An entostylid is present
anterior to the entoconid, set off by grooves from it and from the
metaconid; a precisely similar structure occurs in dm4 of Petro-
dromus. I differ from Whitworth as regards cusp homologies in
this tooth. My parastylid is his paraconid, my entoconid and
hypoconuhd are regarded by him as a stylar development and my
entostylid is his entoconid. In hyracoids, dm'* is somewhat
narrower relative to length than is M^, which it otherwise resembles
very closely, and dm4 and Mi are nearly identical.
Stromer (1926) described and figured various postcranial frag-
ments, which he referred to Myohyrax. Except for an atlas, which
may not be correctly identified, all of these are decidedly macro-
scelidid in appearance. Figure 6 shows a selection of these frag-
ments, redrawn from Stromer, compared with corresponding parts
of a living member of the family. The resemblances are obvious
and do not need to be elaborated. One point may be stressed.
The astragalus is about as different as possible from that of
hyracoids. This element in the latter is as distinctive in its way as
are the corresponding bones of artiodactyls and perissodactyls. In
the hjTacoid astragalus the articular area of the trochlea con-
tinues distally over the medial side of the short neck into a curious,
step-like surface for the reception of the long, stout internal
malleolus of the tibia. This specialization had already been
attained by early Oligocene, Fayum forms, the earliest known
PATTERSON: FOSSIL ELEPHANT SHREWS
317
Figure 6. A comparison of myohyracine and rhynchocyonine postcranial
elements. Myohyrax oswaldi: a, c, e, g, i; Rhynchocyon cirnei: b, d, f, h, j.
a, b, proximal ends of humeri; c, d, proximal ends of ulnae; e, f, proximal ends
of femora; g, h, astragali; i, j, calcanea. a, c, e X2; g, i, X4; R. cirnei not to
scale. M. oswaldi redrawn from Stromer, R. cirnei MCZ 43735.
members of the order (Schlosser, 1911, p. 126, pi. 13, fig. 2). The
comparatively long-necked myohyracine astragalus shows no
trace of such a structure, and agrees in all essentials with those of
other macroscelidids (Fig. 6 g, h). It is regrettable that none of
these pieces is complete enough to give any idea of the degree of
fusion of the lower leg bones, not to mention the relative lengths
of the limb segments or of the fore and hind Hmbs. Whether or not
318 bulletin: museum of comparative zoology
the myohyracines were saltatorial, as are living forms^ remains
unknown.
Four species of early Miocene myohyracines have been de-
scribed: Myohyrax oswaldi Andrews 1914, Pr-otypotheroides beetzi
Stromer 1922, Myohyrax doederleini Stromer 1926, and Myohyrax
osborni Hopwood 1929. Of these, M. oswaldi and M. doederleini
are small and very similar, P. beetzi and M. osborni are much larger
and very similar. Whitworth recognizes but two species, oswaldi
and beetzi, and in this he is undoubtedly correct. He goes on to
synonymize Protypotheroides with Myohyrax, but here I am unable
to follow him. It appears to me that the differences between the
two valid species, summarized in the diagnoses below, are of
generic significance, as genera are defined in this family.
INIyohyrax Andrews
Myohyrax Andrews, 1914, p. 171.
Type species: M. oswaldi Andrews, 1914.
Distribution: Early Miocene, east and southwest Africa.
Emended diagnosis: Myohyracines with cement in fossettes of
cheek teeth; M^ single rooted; fossettids on P3 — M2 ephemeral.
Myohyrax oswaldi Andrews
Myohyrax oswaldi Andrews, 1914, pp. 169-171, pi. 28, figs. 4-6; Stromer, 1926,
pp. 123-124, pi. 41, figs. 26-28; Whitworth, 1954, pp. 26-40, text-figs.
9-15, pi. 5, figs. 3-4, pi. 6.
Myohyrax doederleini Stromer, 1926, pp. 120-123, text-fig. 19, pi. 41, figs.
1-23; Hopwood, 1929, p. 6, text-fig. 4.
Horizon: Early Miocene.
Localities: Kenya: Koru (type locaUty), Karungu, and^Rusinga
Island (definitely recorded from the lower Hiwegi beds) ; South-
West Africa: Elisabethf elder, a borehole some 37 km SSE of
Liideritzbucht (Stromer), and "south of Llideritz Bay" (Hop-
wood) .
Diagnosis: As for the genus. Intermediate in size between the
species of Petrodromus and those of the smaller macroscelidine
genera.
The combination of relatively persistent fossettes in the upper
molars and ephemeral fossettids in the lower molars is almost
precisely matched in Macroscelides. The species is common at
1 It is sometimes stated (e.g. Evans. 1942, p. 85) on the basis of observations by field workers
that, despite their evident saltatorial adaptations, macroscelidids do not hop but run on all four
feet. Run they obviously do but hopping has also been observed.
PATTERSON: FOSSIL ELEPHANT SHREWS 319
certain localities. In South-West Africa, Stromer records over
100 from Elisabethfelder, and Whitworth (1958, p. 47) lists 120
from Karungu in Kenya.
Protypotheroides Stromer
Protypotheroides Stromer 1922, p. 333.
Type species: P. beetzi Stromer, 1922.
Distribution: Early Miocene, southwest Africa.
Emended diagnosis: Myohyracines without cement in fossettes
of cheek teeth; M^ two rooted; fossettids on P3 — M2 deep.
Protypotheroides beetzi Stromer
Protypotheroides beetzi Stromer, 1922, p. 333; 1926, pp. 124-125, pi. 41, figs.
29-31.
Myohyrax osborni Hopwood, 1929, pp. 6-8, text-figs. 5-6.
Myohyrax beetzi Whitworth, 1954, p. 26.
Horizon: Early Miocene.
Localities: Langental, some 10 km NNE of Bogenfels (Stromer),
and "south of Llideritz Bay" (Hopwood), South-West Africa.
Diagnosis: As for the genus. A large species, for this family;
comparable in size to Rhynchocyon petersi the largest living form.
Neither Stromer nor Hopwood mention cement in P. beetzi, and I
was unable to detect any. The anterior wall of the alveolus of M^
is preserved in the type of "ilf . osborni'^ and shows the presence of
two roots. The fossettids extend nearly to the bases of the posterior
cheek teeth. P. beetzi is rare in comparison with AI. oswaldi, only
eight specimens having been recorded.
DISCUSSION
Intrafamilial Relationships
Among the living forms, Rhynchocyon stands apart in a number
of characters — e.g., largely or wholly edentulous premaxillae,
upper canines large and Pi caniniform, long and broad facial
region, large cranial table, no palatal fenestrae, backwardly sloping
ascending ramus, presence of chevron bones, radius and ulna not
fused, digit I lacking and digit V reduced in the manus. The
majority of authors agree in placing Rhynchocyon in a distinct
subfamily and this seems justifiable. The extinct R. clarki shows
that the subfamily was in existence by Miocene time, and in so
doing provides us with the nearest approach to a phyletic hneage
that we have. This species, as is not surprising, possesses some
320 bulletin: museum of comparative zoology
macroscelidine characters; divergence of the two subfamiUes may
date from earUer Ohgocene time.
The Macroscelidinae, with four living and two extinct genera,
form the core of the family, so far as present knowledge goes. The
various forms differ unevenly within rather narrow limits. The
heights of the cheek teeth range from brachyodont in Metoldohotes
and Petrodromus to subhypsodont in Palaeothentoides, Nasilio and
Macroscelides, yet two of the higher crowned forms, Palaeothen-
toides and Nasilio, are primitive in their retention of a vestigial M3
{Metoldohotes, the earUest known macroscelidid, had already lost
this tooth). Macroscelides stands alone in its possession of highly
inflated bullae and epitympanic sinuses. Nasilio, advanced as
regards molar height, has the tibia shorter relative to the femur
than in either Elephantidus or Macroscelides and resembles in this
respect the rather generahzed Petrodromus. The latter, in turn,
is specialized as regards the loss of the hallux, and so on. Palaeo-
thentoides and the living macroscelidines give the impression of
being terminal twigs of a once more numerous and varied group.
It is unsafe at present to assert that two or more members of the
subfamily are closer to each other than to the rest phylogenetically,
since characters in common could well have been achieved inde-
pendently.
The two extinct groups, Myohyracinae and Mylomygahnae,
best regarded for the present as subfamilies, are highly specialized
as regards their cheek teeth, the former in an unguIate-Uke, the
latter in a rodent-Uke direction. Unfortunately, we know them
only at moments in time — the two myohyracines in the earlier
Miocene, Mylomygale in the Pleistocene — and hence have no
direct evidence bearing on their phylogenies. The myohyracines,
which alone in the family retain M^ as well as M3, may have
branched off at an early date, possibly Eocene, the mylomygalines
perhaps somewhat later. The ancestry of both groups may have
lain in the IMacroscehdinae, but this is uncertain.
The only extra- African form that has been referred to the family
is Pseudorhynchocyon cayluxi Filhol (1892) from the Quercy
Phosphorites. This very unsatisfactorily known animal was based
on the posterior part of a left mandible, in which the alveoli of the
last molar provide the only trace of the dentition. The ascending
ramus is much inclined posteriorly, and Filhol saw in this a
resemblance to Rhynchocyon. In fact, however, Pseudorhynchocyon
in this respect goes far beyond conditions in the living form (Fig.
7). As Butler and Hop wood have pointed out, the ascending ramus
of the Miocene R. clarki is considerably less inclined than in the
PATTERSON: FOSSIL ELEPHANT SHREWS
321
-'--?
.-^
a
Figure 7. Posterior portions of mandibles of, a, Rhynchocyon clarki,
b, Rhynchocyo7i petersi, c, Pseudorhynchocyon cayluxi. X2. a modified from
Butler and Hopwood, b MCZ 22573, c redrawn from Filhol.
living species, which suggests that such inchnation is of relatively-
recent acquisition within the subfamily. Besides this, there is
nothing at all macroscelidid-like about the structure of the ascend-
ing ramus of Pseudorhynchocyon. The coronoid process is much
stouter than in any member of the family and extends well above
the condyle. The latter faces posteriorly and is level with the
cheek teeth, not far above them. The angle is low, and a prominent
masseteric crest leads downward and then upward from it, pro-
jecting down below the level of the ventral border of the horizontal
ramus. Butler and Hopwood conclude that "... the reference of
Pseudorhynchocyon to the Macroscelididae is most improbable."
They are quite right. Whatever the creature may be — and I can
offer no constructive suggestion on this score — it is not a member
of this wholly African family.
322 bulletin: museum of comparative zoology
The Food of Macroscelidids
The posterior cheek teeth of myohyracines are comparable as
regards degree of hypsodont}^ and crown complexity to those of
certain hypsodont notoungulates, or, in miniature, to those of late
Miocene or early Pliocene Equinae. As both Hopwood and Whit-
worth emphasize, such specialization can only be regarded as an
adaptation to a diet consisting in large part of harsh vegetation.
Mylomygale has posterior cheek teeth that are similar in height and
complexity to those of various hj^psodont rodents. Within the
Macroscelididae there have arisen two groups primarily adapted
to an abrasive vegetable diet. It thus becomes important to ascer-
tain if living members of the familj^ are to some extent herbi-
vorous.
Structurally, as has long been recognized, the macroscelidid jaw
is basically that of a herbi\'ore. The jaw muscles, especially AI.
temporalis, show resemblances to those of artiodactyls, although,
as Fiedler (1953, p. 161) has pointed out, the disposition of the
tendons (Sehnenskelet) is "insectivoran." The structure of the
cheek teeth, particularly of the higher-crowned living forms, is con-
sistent with a diet at least partially herbi^'orous. Is there e^•idence
that these animals do in fact eat plants? Regrettably, no thorough
study of the diet of any macroscelidid is available, and the anecdo-
tal literature is unsatisfactory. Unsubstantiated assertions to the
effect that macroscelidids are exclusively insectivorous are common.
Reports by collectors that insects were found in stomachs are
sometimes quoted, but insect remains are relatively easy to detect
and would likely be looked for (macroscelidids being "insecti-
vores"), whereas vegetable remains, if scanty, are more difficult to
recognize and might even be passed over ^^'ithout comment on the
supposition that they had been accidental!}^ ingested together with
the "prey. Nevertheless, a few statements do suggest that these
animals are omnivorous. Thus, Sclater (1901, p. 155) states, on the
authority of Francis, a collector, that Petrodromus sultan is ". . .
very partial to the droppings of the Livingstone buck (Nesotragus
livingstonianus) .'" Shortridge writes, of Elephantuhis (1934, p. 21),
". . . although mainly insectivorous [they] are to some degree
omnivorous, and may be caught in traps baited with meahes,
quaker oats, etc."; and, of E. intufi, "examined stomach contents:
insects and a small amount of vegetable matter" (p. 23). "The
smaller species . . . feed mainly on ants . . . supplemented by
tender shoots, roots and berries" (Walker et al. 1964, p. 134). The
PATTERSON: FOSSIL ELEPHANT SHREWS 323
most convincing statement is by Broom (1898, p. 68): "Macro-
scelides^, the Elephant-shrew, has a jaw which judging by analogy
would certainly be related to a herbivorous form and the molar
teeth would seem to be quite in harmony with this determination,
and yet though Macroscelides is largely a vegetable feeder [italics
mine] I have found in the stomach abundant remains of ants and
even of fairly large beetles." Although quite inadequate to reveal
how large a role vegetable food may play in the macroscelidid diet,
this small budget of information does reveal that plants in one
form or another are eaten. If early members of the family were
similarly omnivorous, the evolution of predominantly herbi-
vorous phyla is readily understandable.
The Systematic Position of the Macroscelididae
Real knowledge of the macroscelidids dates from 1829-, when
Macroscelides was described by Smith, and of the tupaiids from
1821 -, when Raffles proposed Twpaia. Neither group formed part
of Bowdich's Insectivora of 1821, based on Cuvier's "les insecti-
vores" of 1817, which included representatives of a majority of the
living families. Given the knowledge available in the earlier part
of the 19th century, however, it was inevitable that both families
would be placed in this order. As knowledge improved it was
equally inevitable that differences between them and the rest of the
Insectivora would become increasingly apparent. Peters (1864),
recognizing this, divided the order Insectivora into two major,
unnamed groups, one with, one without a caecum, and included
the colugos with the tupaiids and macroscelidids in the first.
Haeckel (1866, p. cix), excluding the colugos, gave to these groups
the formal names Menotyphla and Lipotyphla, the former based
equally on the Cladobatida ( = Tupaiidae) and the MacrosceUdea
( = Macroscehdidae) ^, the latter including the original Insectivora. ^
With this, a stage was set. Elephant shrews and tree shrews
became firmly associated in the minds of many investigators, and
discussions of affinities, particularly of the former group, fell into a
1 The old inclusive genus Macroscelides had not been subdivided at this date, and the
question therefore arises as to whether Broom was dealins; with the genus as now restricted.
It is known, however, that in 1897-98 he resided in Little Naniaqualand and hence was within
the range of M. proboscideus.
2 Macroscelides had been known since 1800, but disguised as Sorex proboscideus, Tupaia
since 1820, but disguised as Sorex glis.
3 Simpson (1931, p. 16n; 1945, pp. 176, 183) has stated that Menotyphla was based on the
macroscelidids; this does not appear to be the case.
4 There is some tendency nowadays to employ Lipotyphla as an ordinal name for the In-
sectivora minus the "Menotyphla." It needs to be emphasized, as McKenna (1963b, p. 4n) has
done, that "Lipotyphla" is to all intents and purposes strictly synonymous with Bowdich's
Insectivora.
324 bulletin: museum of comparative zoology
rut. The characters the two families had in common were hailed as
proof of close relationship, almost as though the possession of such
things as caeca and normal mammahan zygomatic arches and
pubic symphyses were pecuHar to them. Even after attention had
become focused on the evident resemblances to primates shown by
the tree shrews and strong doubts had been cast on the reahty of
Haeckel's Menotyphla, some students continued to associate the
two families closely, and even to waft the macroscelidids to the
primate heights as a kind of ill-fitting tail to the tupaiid kite.
Inclusion of the Tupaiidae in the Primates is a view becoming
more and more wadely accepted. There is no occasion here to trace
the development of this concept (Carlsson, 1909, 1922; Gregory,
1910; Le Gros Clark, 1934; Simpson, 1935, 1945; and others) or to
review the imposing body of data that favors it. Some items of
evidence that have not yet passed into the general literature may
be mentioned, however. Henckel (1928) and Roux (1947) con-
cluded that the chondrocranium of Tupaia excluded the family
from the order Primates. This opinion was largely based on the
absence of a septum interorbitale, a supposed hall mark of pri-
mates. Grasse (1955, p. 1649), who recognized Menotyphla in the
Haeckelian sense, utilized this to offset Saban's (1956-7) conclu-
sion, based on a thoroughgoing study of the adult skull, that
tupaiids were members of the order. Recently, Starck (1960, 1962),
working on a wide variety of primates, has found the septum
interorbitale to be a highly plastic structure without taxonomic
significance, its presence or absence largely depending on the
developmental stage under investigation. As he puts it, "Damit
verlieren die Hypothesen (Henckel), die Tupaia aus der Prima-
tenreihe ausschhessen woUen, ihre Hauptstiitze."^ The placenta-
tion of tupaiids was very poorly known until quite recently.
Meister and Davis (1956) have helped to fill this gap with their
description of three stages in Tupaia minor. They conclude that
"morphologically the placenta and other fetal membranes of
Tupaia are almost an ideal starting point from which to derive the
corresponding structures of the primates." Of particular interest
for the present study is their further conclusion, based on compari-
son with Van der Horst's drawings (1950) of Elephantulus, that
". . . the placenta and fetal membranes in these two forms differ
in almost every respect except placental type"; the data ". . . sup-
port the view that the tree-shrews and elephant shrews are not
1 Since this was sent to press, W. Spatz' study of the ontogeny of the cranium of Tupaia glis
(Morphol. Jahrb., 106: 321-416, 1964) has come to hand. He concludes that on this evidence
tupaiids are primates.
PATTERSON: FOSSIL ELEPHANT SHREWS 325
closely related." The little evidence available on ectoparasites
supports primate affinities for the former but not for the latter
(Patterson, 1957, pp. 23, 26). A few uncertainties remain concern-
ing the propriety of including tupaiids in the Primates. Some see
the structure of the hand as a bar to inclusion while others do not.
Buettner-Janusch and Buettner-Janusch (1964, p. 87) find that
the electrophoretic behavior of the hemoglobin differs from that of
other prosimians.^ Jane, Campbell and Yashon (1965) have shown
that in Tupaia the pyramidal tract occurs in the dorsal funiculus of
the spinal cord, whereas in insectivores it occurs in the ventral
funiculus and in primates in the lateral. ^ However, only one other
prosimian, Nycticehus, has thus far been studied in this connection.
A wider investigation may reveal that the distinction is not a clear
cut one. It should be recalled, to introduce a note of caution, that
until 1952 Tupaia could be said to differ from all primates in its
possession of an outer bar of Jacobson's cartilage. In that year
Eloff demonstrated the presence of the bar in Galago senegalensis.
Certain facts do, of course, await further assessment, but the
weight of the evidence now decidedly favors the ordinal reference ;
the burden of proof has shifted.
The Macroscehdidae, to anticipate a little, are without much
doubt a very ancient family that probably arose early in the
Cenozoic, possibly even toward the end of the Mesozoic. In some
respects specialized, they are nevertheless basically rather primi-
tive. It is not surprising therefore that they should to varying
degrees resemble other groups of mammals of more or less com-
parable antiquity. These resemblances, particularly those to the
Insectivora and to the tupaiid primates, have been interpreted as
indicative of close affinity to one or the other of these groups,
wrongly interpreted I now believe.
As regards the soft anatomy, Le Gros Clark (1933, p. 1004) has
stated of the brain that "... it would be difficult to conceive two
small mammalian brains which are more fundamentally different
and divergent in their structure than those of Macroscelides and
Tupaia.'' Stephan and Spatz (1962) and Stephan and Andy (1964)
also emphasize that the macroscelidid brain differs from those of
Insectivora in various ways, notably in the much larger mesen-
cephalon and hippocampus. The organ of Jacobson (Broom, 1902,
1 On the basis of serum protein comparisons Goodman (1963, p. 137) concludes: "Although
evidence for a definitive taxonomic assignment of the tree shrews has not been gathered, the
serological data demonstrate that the tree shrews have affinities with the Primates. (Elephant
shrews, placed by some taxonomists with tree shrews, do not show any primate or tupaiid
affinities.)"
2 In this, as in certain other respects, Tupaia may be simply primitive. The tract is dorsal
in monotremes, marsupials, edentates and rodents.
326 bulletin: museum of comparative zoology
1915) is of marsupial type, resembling that of Twpaia (presumably-
primitive in both groups) and very different from that of insecti-
vores. Carlsson, although concluding that macroscehdids were
close to erinaceids, did record certain characters in the musculature
in which they resembled the tupaiids (1909, p. 396). The testes
remain abdominal in elephant shrews, whereas the tree shrews
have a well developed scrotum. Both, of course, have a caecum,
but this is much larger in the elephant shrews. In sum, the evi-
dence of the soft parts would appear to oppose close relationship
to either the Tupaiidae or to the Insectivora.
The distinction between macroscehdids and tupaiids in placenta-
tion has been mentioned above (p. 324), as has the remarkable
number of eggs liberated from the ovaries of certain of the Macro-
scelidinae (p. 307). The two famihes also differ as regards the
young. In the elephant shrews these are decidedly precocial, being
born fully haired, with the eyes open, and capable of active locomo-
tion within a very short time (Hoesch and von Lehmann, 1956,
p. 17; ^Yalker et al. 1964, p. 135)^ In tree shrews — and also in
the Insectivora generally (Herter, 1957, p. 31) — this is not the
case. Uterine bleeding has been described by Van der Horst (1954,
and references there cited) in Elephantulus "myurus,^' with the
suggestion that this foreshadows the menstrual cycle of the higher
primates. The bleeding is of an unusual type, however. During
diestrum a polyp-like growth forms in one part of the uterus and
disintegrates at the end of the stage; coiled arteries, which are
''enormously developed," are confined to this part. In all likeli-
hood this is simply another macroscelidid peculiarity.
The dentition of macroscehdids, particularly the posterior cheek
teeth, is unlike that of any other group of mammals. Carlsson saw
resemblances to the teeth of Erinaceidae, but these are not close.
The Macroscelididae have at times been placed in the Insectivora
"Dilambdodonta" although there is nothing whatsoever dilambdo-
dont about their molars. Frechkop (1931) has stated that the
cheek teeth resemble those of ungulates more than those of any
other major group. This is correct — the myohyracines evolved
molars so ungulate-like as to mislead some very competent students
— but there is no detailed similarity to any particular group of
1 A very curious observation concerning the young of Elephantulus rupeslris has been
reported. Fitzsimons (1920, pp. 12-13) quotes an observer, van Musschenbroek, who noticed
that the two young ". . . were hanging on to something on top of the shoulder blades. On
examination I found they were two teats, one on either side. [The mother] carried them by
these teats [and] took good care to see that they were hanging on before she hopped away like
a miniature kangaroo." As regards the position of the teats confirmation is supplied by Burton
(1955), who states that Hayman found them to be situated high on the flanks just behind and
above the scapulae.
PATTERSON : FOSSIL ELEPHANT SHREWS 327
hoofed mammals. The resemblance is an interesting example of
convergence, as Friant (1935) concluded. On the basis of his
observations, Frechkop asserted that the macroscelidids were on
the ungulate road, just as the tupaiids were on the primate one. In
this, of course, he went too far. All, or practically all, ungulate
orders have surely emerged from the Condylarthra (to which the
Arctocyonidae properly belong), and there is nothing suggestively
condylarthran in the elephant shrews. Nevertheless, the myohyra-
cines do raise the possibility that placental mammals could achieve
"ungulate" grade independently of that order.^ The dentition of
the extinct macroscelidids gives no hint as to relations with any
other group. The earliest known form, Metoldobotes, had the typical
pattern and had lost the last molar.
The most recent study of the skeleton of the Alacroscelididae is
that of Evans (1942), who compared representatives of all genera
except Macroscelides (see p. 309) with those of Tupaia and Echino-
sorex. He concluded that the macroscehdids resembled Tupaia in
30 osteological features and Echinosorex in 13. Further, he
claimed that out of 40 lemuroid features mentioned by Gregory
and by Carlsson as occurring in Tupaia, the macroscelidids shared
in 32. This "simple morphological balance," as Simpson (1945,
p. 176) called it, has been seized on as evidence for the reality of
Menotyphla sensu Haeckel (e.g. Grasse, 1955, p. 1649; Heim
de Balzac and Bourhere, 1955, p. 1691). It does not provide such
evidence. Evans' study is in fact a classic example of the "rut
discussions" mentioned above. He has shown beyond question
that tupaiids and macroscelidids differ in a number of features,
some of them more or less similar, from Echinosorex, which, as a
true insectivore, has various characters not found in other major
groups of mammals. This is a far cry from proving a close rela-
tionship between elephant shrews and tree shrews, however.
Looking beyond the restricted prosimian-tupaiid-macroscelidid-
insectivore circle, it becomes apparent at once that few of Evans'
characters in common between tupaiids and macroscelidids are
confined to these families. Furthermore, the resemblance between
the two in many of the features cited is far from close. Some of the
common features, e.g. the relatively large braincase, were in all
probability independently acquired (the braincases at least house
very dissimilar brains); others, e.g. the structure of the orbito-
temporal region and of the zygomatic arch, are simply primitive
eutherian, or therian, characters inherited by both; others again.
1 The only other possible candidates for such a distinction that I can think of are the
notoungulates. I am not suggesting a macroscelidid-notoungulate relationship!
328 bulletin: museum of comparative zoology
e.g. the presence of a free centrale and of a third trochanter on the
femur, are characters so widely possessed as to be meaningless in
this context; yet others, e.g. large auditory bullae and slender
coronoid process of the mandible, are not sufficiently similar
structurally to quahfy as significant resemblances; and so on. The
same apphes to the features cited as occurring in common between
macroscehdids and lemuroids. Butler (1956, p. 476) has listed
certain cranial characters in which the Macroscelididae, Dermop-
tera, Tupaiidae and Lemuroidea resemble each other. Some of
these features are of the same sort as those cited by Evans. As
regards the JNIacroscehdidae, at least, they are equally open to
question, as Butler recognized. The evidence from the hard parts
seems to me to point in the same direction as that from the soft :
namely, that the macroscehdids are sharply distinct from both
Tupaiidae and Insectivora. Resemblances to ungulates do exist —
Rhynchocijon and Orycteropus are remarkably similar in the disposi-
tion of the bones in the orbitotemporal region (cf. figs. 124 and 177
in Gregory, 1920), the rostral and caudal entotympanics of elephant
shrews compare rather closely with those of notoungulates, fusion
of distal elements in the hmbs occurs in macroscehdids and in
hoofed mammals, the astragalus has a fairly long neck in some
small and primitive ungulates, etc. — but these appear to be
either convergent or simply primitive in both.
Evans concluded, on the basis of superficial resemblances, that
Anagale from the early Ohgocene of Mongolia "... is, in many
osteological features, intermediate between the Alacroscelidae and
the Tupaiidae and is either the common ancestor of the two
families or quite close to it." New evidence, derived from a hitherto
undescribed specimen of Anagale, and from Anagalopsis, reveals
that the cheek teeth are quite different from those of either family,
and that the tympanic forms the lateral portion of the bulla instead
of being a ring enclosed by the entotympanic, as Simpson (1931)
had supposed. Basing his conclusion on this evidence and on the
very peculiar structure of the unguals, AIcKenna (1963a) has re-
moved the Anagahdae from the Tupaioidea, where Simpson had
placed them, and hsted them as Eutheria incertae sedis. Whatever
the anagalids may prove to be, they are not related to the macro-
scehdids. Metoldobotes, it may be recalled, was contemporary with
Anagale.
As will by now be evident, I believe the elephant shrews to be
a group of mammals distinct from both the Insectivora and the
tupaiid primates. Their known distribution is exclusively African,
and they make their first appearance in the record in the earliest
PATTERSON: FOSSIL ELEPHANT SHREWS 329
adequately known mammalian fauna of that continent. They are
accompanied there by an array of mammalian groups unknown
elsewhere in deposits of earlier or similar date — hyracoids,
arsinoitheres, moeritheres, barytheres, proboscideans and catar-
rhine primates. So notable a degree of endemism argues a long
isolation of Africa^ (Darlington, 1957, pp. 365, 590; Patterson,
1957, p. 45), one lasting throughout much of the Eogene at least.
Macroscelidids may well have been members of this "old African"
fauna, survivors from the later Cretaceous beginnings of the
Eutheria. As a group they are more diversified and contain more
genera than almost one-third of the currently recognized orders of
eutherian mammals. They are, I believe, worthy of ordinal rank.
Butler (1956) has proposed for them the ordinal name Macro-
scelidea^ without definition. Such action seems preferable to
restriction of Menotyphla to the elephant shrews. ]\ienotyphla
has long had a proto-primate flavor, and since the tupaiid half of
the artificial assemblage almost surely is primate and the macro-
scelidid half assuredly is not, it hardly seems desirable to attempt
perpetuation of so ambiguous a name for the latter alone.
The order may be definied as follows:
MACROSCELIDEA
Dentition I°3^, C\ , Ft, M2I3 ; Ft large, molarif orm ; upper cheek
teeth without mesostyles; M3, when present, greatly reduced;
posterior cheek teeth brachyodont to hypsodont. Skull with
complete zygomatic arch; orbits large, open posteriorly; maxilla
not extending into orbital wall, palatine with orbital wing; auditory
bulla compound, ectotympanic, rostral and caudal entotympanics,
ahsphenoid, squamosal, periotic participating ; mandible with high
ascending ramus, condyle well above level of cheek teeth, coronoid
process small. Pelvis with pubic symphysis; humerus with
entepicondylar foramen; distal segments of legs longer than
proximal; radius and ulna fused or closely appressed, tibia and
1 When this isolation began and ended is of course uncertain. Darlington, on the basis of
the rather few northern forms that occur in the Fayum deposits, believes that a connection had
by then become established. This does not seem certain ; that only two or three of a great many
northern groups would have made their way over a land bridge had this been fully in existence
is rather unlikely. The Fayum rodents belong to a family not known in Eurasia; they may well
have descended from waif ancestors transported during the period of isolation.
2 Somewhat vaguely, however, since in the body of his paper (p. 479) he expressed doubt as
to the propriety of including the elephant shrews in the Insectivora ("Lipotyphla" in his
terminology), suggesting that they should either ". . . be included in the Primates as an out-
lying suborder, or a new order, Macroscelidea, should be created for them." In the summary
(p. 480) he simply remarked that they are ". . . placed in a new order, Macroscelidea." Haeckel's
prior — and invalid (by modern standards) — use of the same name for the family does not
constitute preoccupation. I was previously dubious about recognition of the order (1957, p. 23),
but with the increase in knowledge of the fossil record my doubts have disappeared.
330 bulletin: museum of comparative zoology
fibula fused; pollex and hallux reduced or absent; astragalar neck
moderately long. Proboscis long, flexible; organ of Jacobson of
marsupial type; brain with relatively large mesencephalon and
speciahzed hippocampus ; caecum relatively large ; testes abdominal.
One family, Macroscelididae, with four subfamilies: Macro-
scelidinae, Rhynchocyoninae, Mylomygahnae, jNIyohyracinae.
Known range: Early Oligocene to Recent, Africa.
SUMMARY
The Macroscelididae, a wholly African group so far as known,
includes four subfamihes, two of which are extinct. The Macro-
scelidinae date from the early Oligocene of the Fayum, where they
are represented by Metoldobofes, a form originally referred to the
insectivore family JNIixodectidae. Palaeothentoides of the early
Pleistocene (?), first described as a marsupial, is a valid member of
the subfamily. Extinct species of Elephantulus are known from the
Pleistocene. The early Miocene Rhynchocyon clarki provides the
only fossil record of the Rhynchocyoninae. The subfamily
Mylomygalinae is proposed for the Pleistocene Mylomygale, a
remarkable form with hypsodont posterior cheek teeth convergent
toward those of various rodents. The Myohyracinae, hitherto
placed in the Hyracoidea as Myohyracidae or ]\Iyohyracoidea, are
represented by the early Miocene Alyohyrax and Protypotheroides.
Their posterior cheek teeth are decidedly ungulate-like and com-
parable in complexity and degree of hypsodonty to those of
Equinae and hypsodont Notoungulata. Pseudorhynchocyon cayluxi
from the Quercy Phosphorites is not a member of the family.
The extinct subfamilies were beyond doubt predominantly
herbivorous. Some evidence indicates that the surviving forms are
to a degree omnivorous.
The affinities of the family are reviewed and the conclusion
reached that macroscehdids are not closely related either to the
tupaiid primates or to the insectivores. Resemblances to ungulates
are either convergent or primitive. Butler's order Macroscelidea is
recognized for the reception of the group and a definition is offered.
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PATTERSON: FOSSIL ELEPHANT SHREWS 331
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1955. Ordre des insectivores. Systematique. In Traite de Zoologie,
P.-P. Grasse, ed. Paris: Masson ct Cie. Vol. 17, Mammiferes,
fasc. 2: 1653-1697.
Henckel, K. O.
1928. Das Primordialcranium von Tupaiia und der Ursprung der
Primaten. Zeitschr. Anat. Entwickl., 86: 204-227.
Herter, K.
1957. Das Verhalten der Insectivorcn. In Handbuch der Zoologie,
J.-G. Helmcke, H. von Lengerken and D. Starck, eds. Berlin:
Walter de Gruyter u. Co. Bd. 8, lief. 9, 10(10): 1-50.
HoEscH, W. and E. von Lehmann
1956. Zur Saugetier-Fauna Sudwestafrikas. Bonn. Zool. Beitr., 7: 8-57.
PATTERSON: FOSSIL ELEPHANT SHREWS 333
HopwooD, A. T.
1929. New and little-kuowu maninials from the Miocene of Africa.
Amer. Mus. Novit., No. 344: 1-9.
Jane, J. A., C. B. G. Campbell and D. Yashon
1965. Pyramidal tract: a comparison of two prosimian primates.
Science, 147: 153-155.
Le Gros Clark, W. E.
1933. The brain of the Insectivora. Proc. Zool. Soc. London, 1932:
975-1013.
1934. Early forerunners of man. A morphological study of the evo-
lutionary origin of the primates. London and Baltimore: Balliere,
Tindall and Co.x, William Wood and Co. Pp. i-xvi, 1-296.
1959. The antecedents of man. An introduction to the evolution of the
Primates. Edinburgh: The University Press. Pp. i-ix, 1-374.
McKenna, M. C.
1963a. New evidence against tupaioid affinities of the mammalian family
Anagalidae. Amer. Mus. Novit., No. 215S: 1-16.
1963b. Primitive Paleocene and Eocene Apatemyidae (Mammalia,
Insectivora) and the primate-insectivore boundary. Ibid.,
No. 2160: 1-39.
Matthew, W. D.
1910. Schlosser on Fayum mammals. A preliminary notice of Dr.
Schlosser's studies upon the collection made in the Oligocene of
Egypt for the Stuttgart Museum by Herr Markgraf. Amer. Nat.,
49: 700-703.
1915. A revision of the Wasatch and Wind River faunas. Part IV — •
Entelonychia, Primates, Insectivora (part). Bull. Amer. Mus.
Nat. Hist., 34: 429-483.
Meister, W. and D. D. Davis
1956. Placentation of the pygmy treeshrew. Fieldiana: Zool., 35: 71-84.
Patterson, B.
1957. Mammalian phylogeny. In Premiere symposium sur la specificite
parasitaire des parasites des vertebres. Neuchatel: Paul Aldinger.
Pp. 15-48.
Peters, W.
1864. Ueber die Saugethier-Gattung Sulenodon. Abh. K. Akail Wiss.
Berlin, 1863: 1-22.
ROMER, A. S.
1928. Pleistocene mammals of Algeria. Fauna of the Paleolithic station
of Mechta-el-Arbi. Bull. Logan Mus., 1: 80-163.
Roux, G. H.
1947. The cranial development of certain Ethiopian insectivores and its
bearing on the mutual affinities of the group. Acta Zool., 28:
165-307.
Saban, R.
1956- Les affinites du genre Tiipaiia Raffles 1821, d'apres les caracteres
1957. morphologiques de la tete osseuse. Ann. Paleont., 42: 169-224
(1956); 43: 1-44 (1957).
334 bulletin: museum of comparative zoology
SCHLOSSER, M.
1910. tjber einige fossile Siiugetiere aus dem Oligociin von Agj^pten.
Zool. Anz., 35: 500-508.
1911. Beitrage zur Kenntnis der oligozanen Landsiiugetiere aus dem
Faj'um, Agypten. Beitr. Pal. Geol. Osterr.-Ungar. Orients, 24:
51-167.
SCLATER, \V. L.
1901. The mammals of South Africa. Vol. II. Rotlentia, Chiroptera,
Insectivora, Cetacea and Edentata. London: R. H. Porter.
Pp. i-xii, 1-241.
Shortridge, G. C.
1934. The mammals of South West Africa. A biological account of the
forms occurring in that region. Vol.1. Pp. i-xxv, 1-4157. London:
William Heinemann Ltd.
Simpson, G. G.
1931. A new insectivore from the Oligocene, Ulan Gochu horizon, of
Mongolia. Amer. Mus. Novit., No. 505: 1-22.
1935. The Tiffany fauna, upper Paleocene. II. Structuie and relation-
ships of Plesiadapis. Ibid., No. 816: 1-30.
1945. The principles of classification and a classification of mammals.
Bull. Amer. Mus. Nat. Hist., 85: i-xvi, 1-350.
Satrck, D.
' 1960. Das Cranium eines Schimpansenfetus {Pan troglodytes [Blumen-
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Korperform von Schimpansenfeten. (Beitrag zur Kenntnis
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Stephan, H. and O. J. Andy
1964. Quantitative comparisons of brain structures from insectivores to
primates. Amer. Zool., 4: 59-74.
Stephan, H. and H. Spatz
1962. Vergleichend-anatomische Untersuchungen an Insectivorengehii--
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PATTERSON: FOSSIL ELEPHANT SHREWS 335
1932. Palacothcntoides africanus nov. gen., nov. sp(>c., oiti erste.s
Iknitcltier aus Afrika. Ibid., 1931: 177-190.
Thomas, O.
1901. List of small mammals obtained by Mr. A. E. Pease, M.P., during
his recent expedition to Abyssinia, with descriptions of three new
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190B. The Rudd exploration of South Africa. V. List of mammals
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1944. Remarks on the systematics of Elephantulus. Jour. Mammal., 25:
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1950. The placentation of Elephantulus. Trans. Roy. Soc. S. Afr., 32:
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Van der Klaauw, C. J.
1929. On the development of the tympanic region of the skull in the
Macroscelididae. Proc. Zool. Soc. London, 1929: 491-560.
Wagner, P. A. and H. Merensky
1929. The diamond deposits on the coast of Little Namaqualand. With
an appendix on the palaeontology of the Namaqualand coastal
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Walker, E. P., F. Warnick, K. I. Lange, H. E. Uible, S. E. Hamlet,
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Whitwurth, T.
1954. The Miocene hyracoids of Ea.st Africa. Brit. Mus. (Nat. Hist.).
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1958. Miocene ruminants of East Africa. Ibid., No. 15: 1-50.
(Received 19 February 1965.)
Bulletin of the Museum of Comparative Zoology
HAEVARD UNIVERSITY
Vol. 133, No. 7
PANAMANIAN SPIDERS OF THE GENUS TMARUS
(ARANEAE THOAIISIDAE)
By Arthur M. Chickering
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
November 26, 1965
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Bulletin of the Museum of Comparative Zoology
HAEVARD UNIVERSITY
Vol. 133, No. 7
PANAMANIAN SPIDERS OF THE GENUS TMARUS
(ARANEAE THOMISIDAE)
By Arthur M. Chickering
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
November, 1965
Bull. Mus. Comp. Zool., Harvard Univ., 133 (7): 337-368, November, 1965
No. 7. — Panamanian Spiders of the Genus Tmarus
{Araneae, Thomisidae)
By Arthur M. Chickering
The Pickard-Cambridges (1889-1905) recognized seven species
of Tmarus from Panama. Petrunkevitch (1925) reported only a
single immature specimen of the genus from Panama. Banks
(1929) reported two species now regarded as T. ineptns O.P.-
Cambridge and T. studiosus O.P.-Cambridge. As a result of my
earlier study of the genus (1950) I was able to recognize eighteen
species. Four of these were known only from females, seven known
only from males, with the remaining seven probably known from
both sexes. Roewer (1954) listed twenty species known from
Panama. This list includes T. separatus Banks and T. interritus
Keyserhng, but this inclusion seems to have been an error. T.
separatus Banks is apparently known only from Mexico and
T. interritus Keyserling only from Brazil. Since the publication of
my previous paper on this genus (1950), I have collected spiders in
Panama during four additional periods as a result of which I have
accumulated a considerable number of specimens in this generic
group. These have been completely worked over with the result
that I am now compelled to recognize twelve additional species,
thus making a total of thirty species known from this small area.
It continues to be very difficult to match up males and females
correctly and I am obliged to state that I have some doubts con-
cerning the accuracy with which this has been done in the past.
As I have frequently stated in my published papers, I am deeply
grateful for the many privileges extended to me by the staff of the
Museum of Comparative Zoology, Harvard University, over a
period of many years. My studies could never have been con-
tinued without this aid and encouragement. Special acknowledge-
ments should be extended to Dr. Ernst Mayr, Director; Dr. P. J.
Darlington, Jr., Alexander Agassiz Professor of Zoology; and
Dr. Herbert W. Levi, Associate Curator of Arachnology.
Grant No. GB-1801 from the National Science Foundation
made it possible for me to spend seven months making collections
of spiders in the West Indies and Panama during the latter part of
1963 and the first five months of 1964. This grant is also making
it possible for me to continue my studies at the Museum of Com-
parative Zoology for a considerable period. A Guggenheim
Fellowship made it possible for me to collect in Jamaica, W. I., and
in Panama, in 1957 and 1958, and also to spend four months in the
340 bulletin: museum of comparative zoology
British ^Museum (Natural History) studying important collections.
Dr. G. Owen Evans and Mr. D. J. Clark, Department of Zoology
(Arachnida), British Museum (Natural History), have recently
loaned me important specimens of the genus Tmarus for study in
the preparation of this paper.
All types named in this publication together with all other
specimens of the genus Tmarus in my personal collection will be
deposited in the JNIuseum of Comparative Zoology, Harvard
University.
Genus Tmarus Simon, 1875
The type species of the genus is Tmarus piger (Walckenaer),
1802; widely distributed in Europe and Asia. The genus is cosmo-
pohtan in distribution and is most abundant in the neotropical
region.
In addition to the general family characteristics, generic features
common to the species from Panama may be stated as follows:
Chelicerae without teeth; only slightly porrect; essentially parallel.
Lip much longer than wide at base. Eyes: eight in two rows;
viewed from above, posterior row definitely recurved, anterior row
slightly so; lateral eyes larger than medians, on well defined
tubercles; posterior tubercles the larger; central quadrangle
usually wider behind than long and wider behind than in front.
Clypeus porrect, often conspicuously so; height equal to several
diameters of AME. Carapace usually abruptly declined to pos-
terior border; without a median thoracic groove or fovea. Legs:
spinose; usually 1243 in order of length but occasionally 2143 or
other minor deviation; one and two nearly equal in length and
longer than three and four which are also nearly equal in length.
Tarsal claws toothed; palpal claw in females also toothed. Ab-
domen often with a posterior tubercle or other modification; with
many short, stiff spines and spinules. Male palpi usually well
supplied with apophyses, and female epigyna usually well defined
and often complicated. To conserve space certain features pos-
sessed in common by all species under consideration will not be
specifically mentioned in the technical descriptions of new species
in this paper. For the same reason a more concentrated description
than usual in my published papers will be adopted in this and
succeeding publications. When eyes are referred to by the abbre-
viations Ar^IE, ALE, PME and PLE, the anterior medians, an-
terior laterals, posterior medians and posterior laterals are respec-
tively designated.
chickering: tmarus from Panama 341
A complete list of the species of the genus Tmarus from Panama
as they are now recognized may be given as follows: Tmarus
aculeatus Chickering; T. hucculentus Chickering; T. cognatus
Chickering; T. conforfus Chickering; T. corruptus 0. P.-Cambridge;
T. crefatus sp. nov. ; T. curvus Chickering; T. decens 0. P.-Cam-
bridge; T. decorus sp. nov.; T. hinnphreyi sp. nov.; T. impedus sp.
nov.; T. ineptus O. P.-Cambridge; T. innotus sp. nov.; T. innumus
sp. nov.; T. intentus 0. P.-Cambridge; T. levii sp. nov.; T. longus
sp. nov.; T. morosus Chickering; T. mundulus O. P.-Cambridge;
T. obsecus sp. nov.; T. parki Chickering; T. pauper 0. P.-Cam-
bridge; T. peregrinus Chickering; T. probus Chickering; T. pro-
dudus Chickering; T. protobius sp. nov.; T. rubinus sp. nov.; T.
sigillatus Chickering; T. stiidiosus 0. P.-Cambridge; T. vitusus
sp. nov. Ten of the species listed above are known only from
males ; ten species are known only from females ; the remaining ten
are probably known from both sexes.
Key to the males of known species of Tmarus from Panama
1. Species with embolus definitely and conspicuously curled either at
anterior end of bulb on ventral side or on retrolateral side {contortus,
curvus, irwrosus, produdus) 2
la. Species with embolus either extended more or less around the margin of
bulb and not definitely curled or, apparently, restricted to anterior end of
bulb {aculeatus, cretatus, decorus, humphreyi, ineptus, innotus, intentus,
mundulus, obsecus, parki, pauper, peregrinus, probus, sigillatus, studiosus,
vitusus) 5
2. Palpal tibial apophyses short, only ventral one strongly chitinized;
cymbium deeply excavate at retrolateral basal corner; embolus deeply
grooved, arises near middle of anterior border of bulb, makes a complete
retrolateral circle and then loops across middle of bulb again (fig. 4,
1950) T. contortus
2a. Palpal features not as given above {curvus, morosus, productus) 3
3. Palp: tibia deeply excavate retrolaterally and distally; with a strongly
cliitinized apophysis of moderate length ventral to excavation; embolus
makes a complete circle distal to anterior margin of bulb, then passes
retrolaterally to terminate in a finely dentate tip (fig. 7, 1950) . . T. curvus
3a. \\ ithout palpal features as given above {tnorosus, productus) 4
4. Palp: ventral, retrolateral, tibial apophysis strongly chitinized and dis-
tall}" knobbed; dorsal apophysis a sharply pointed spine; near anterior
margin of Ijulb the embolus turns toward base, then passes to retro-
lateral side and extends nearly to tip of cymbium as a long, slender
filament (figs. 13-14, 1950) T. morosus
4a. Palp: with a pair of long, slender, retrolateral, tibial apophyses almost
meeting distally, with a hook attached near base of the shorter apo-
physis; deeply grooved embolus makes a loop near distal margin of bulb,
342 bulletin: museum of comparative zoology
then a second loop near base of bulb, continues as a fine filament to
distal end of cymbium (fig. 25, 1950) T. produdus
5. Palp : species with at least the dorsal retrolateral, tibial apophysis elongated
and well developed (aculeatus, cretatus, decorus, hurnphreyi, parki, pere-
grinus, probus, studiosus, vitusus) 6
5a. Palp: species with retrolateral tibial apophyses less well developed; either
with but one apophysis or with two shorter and poorly developed
apophyses {ineptus, intentus, innotus, mundulus, obsecus, pauper, sigil-
latus) 14
6. Palp : with a retrolateral, dorsal, tibial apophysis extending nearly to tip
of bulb {aculeatus, decorus) 7
6a. Palp: with a retrolateral, dorsal, tibial apophysis extending at most little
beyond middle of bulb {cretatus, hurnphreyi, parki, peregrinus, probus,
studiosus, vitusus) 8
7. Palp: with the dorsal, retrolateral, tibial apophysis distally bifurcated
(fig. 1, 1950) T. aculeatus
7a. Palp: with the dorsal, retrolateral, tibial apophysis simple at tip, not
bifurcated (Figs. 7-8) T. decorus
8. Palp: with the dorsal, retrolateral, tibial apophysis subdivided near its
base into a short, pointed process and a long, pointed extension (figs.
29-30, 1950) T. studiosus
8a. Palp: without any basal division of the dorsal, retrolateral, tibial apophy-
sis {cretatus, hurnphreyi, parki, peregrinus, probus, vitusus) 9
9. Palp: with the dorsal, retrolateral, tibial apophysis definitely serrated
along its external edge near middle (fig. 23, 1950) T. probus
9a. Palp: without any serration along external edge of dorsal, retrolateral,
tibial apophysis {cretatus, hutnphrei/i, parki, peregrinus, vitusus) 10
10. Palp: with the ventral, retrolateral, tibial apophysis a relatively massive
structure, deeply notched at distal end (Fig. 10) T. hurnphreyi
10a. Palp: with the ventral, retrolateral, tibial apophysis relatively smaller
and unnotohed at its distal end {cretatus, parki, peregrinus, vitusus). .11
11. Palp: dorsal, retrolateral, tibial apophj-sis with two nearly right angles
in its total length (fig. 20, 1950) T. parki
11a. Palp: with donsal, retrolateral, tibial apophysis not provided with right-
angled bends {cretatus, peregrinus, vitusus) 12
12. Palp: dorsal, retrolateral, tibial apophysis flask -like in shape (fig. 22,
1950) T. peregrinus
12a. Palp: dorsal, retrolateral, tibial apophysis not flask-like in shape {cretatus,
vitusus) 13
13. Palp: dorsal, retrolateral, tibial apophysis elongated; ventral, tibial
apophysis definitely hammer-headed (Figs. 1-2) T. cretatus
13a. Palp: dorsal, retrolateral, tibial apophysi.3 robust, with a very slender
terminal spine; ventral, retrolateral apophysis short, broad, slightly in-
dented terminally T. vitusus
14. Palp: with a definitely recurved hook at base of tarsal bulb (fig. 28,
1950) T. sigillatus
14a. Palp: tarsal bulb without any definite recurved hook at its base {ineptus,
intentus, innotus, mundulus, obsecus, pauper) 15
chickering: tmarus from Panama 343
15. Palp: with a pair of short, pointed processes emerging from a rounded
depression at anterior end of bulb (fig. 10, 1950) T. ineptus
15a. Palp: bulb without such structures as given above {intentus, innotus,
mundulus, obsecus, pauper) 16
16. Palp: tarsal bulb with a single, bifurcated process emerging from a
rounded depression at anterior end (F. P. -Cambridge's fig. 26, table
10) T. intentus
16a. Palp: tarsal bulb without such a structure as given above (innotus,
mundulus, obsecus, pauper) 17
17. Palp: ventral, retrolateral, tibial apophysis a short, blunt process; con-
siderably longer than very short, dorsal, retrolateral apophysis (figs.
16-17, 1950) T. mundulus
17a. Palp: tibial apophyses not as given above (innotus, obsecus, pauper) . . 18
18. Palp: both tibial apophyses short, bluntly pointed processes (O. P.-
Cambridge's fig. 8, table 12) T. pauper
18a. Palp: retrolateral, tibial apophyses unlike those given above (innotus,
obsecus) 19
19. Posterior fourth of abdomen sharply constricted but with no discernible
tubercle (Fig. 27) T. obsecus
1 9a. Abdomen with no marked constriction; with a small but distinct dorsal
tubercle in last quarter T. innotus
No satisfactory key has yet been devised to aid in the identi-
fication of females of known species from Panama.
Tmarus aculeatus Chickering
Tmarus aculeatus Chickering, 1950. Bull. Mus. Comp. Zool., 103(4): 217,
fig. 1. The male holotype from Barro Colorado Island, C. Z., together with
several male paratypes from localities in the Panama Canal Zone and
nearby parts of Panama are all in the Museum of Comparative Zoology.
Roewer, 1954; Bonnet, 1959.
Since 1950 the species has been taken only on Barro Colorado
Island, C. Z., Summit, C. Z., and Arraijan, R. P. The female is
still unknown.
Tmarus bucculentus Chickering
Tmarus bucculentus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 220,
fig. 2. The female holotype from Ft. Randolph, Panama Canal Zone,
August, 1936, together with a female paratype, are in the Museum of Com-
parative Zoology. Roewer, 1954: Bonnet, 1959.
This species, known only from females, has been collected in
three different localities in the Canal Zone since my publication
in 1950.
344 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Tmarus cognatus Chickering
Tmarus cognatus Chickering, 1950, Bull. Mus. Comp. Zool. 103(4): 222,
fig. 3. The female holotype and two female paratypes from the Panama
Canal Zone Forest Preserve, August, 1939, are in the Museum of Compara-
tive Zoology. Roewer, 1954; Bonnet, 1959.
This species, also known only from females, has not been taken
since the original specimens were collected.
Tmarus contortus Chickering
Tmarus contortus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 224,
figs. 4-5. The male holotype, female paratype, one male paratype and three
immature specimens, all taken in the Madden Dam region, Panama Canal
Zone, August, 1939, are in the Museum of Comparative Zoology. Roewer,
1954; Bonnet, 1959.
A very small number of both sexes have been taken since the
establishment of the species and all have come from the Canal
Zone Forest Preserve.
Tmarus corruptus O. P.-Cambridge
Tmarus corruptus O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 95, pi. 12
fig. 10. The holotype female from BugaV)a, Panama, is in the British
Museum (Natural History). F. P.-Cambridge, 1900; Chickering, 1950;
Roewer, 1954; Bonnet, 1959.
This species remains poorly known and only from females. The
elder P.-Cambridge had the holotype from Bugaba, Panama;
F. P.-Cambridge beUeved that he had the species from Mexico.
It has not yet appeared in my collections. A female on loan from
the British jMuseum (Natural History) does not show the two
pairs of minute canals drawn so clearly by F. P.-Cambridge.
O. P.-Cambridge's figure 10c, plate 12, is nearer to what I see in
the epigynum than what is shown in F. P.-Cambridge's figure 32,
plate 10. In view of the uncertainties here and the scarcity of
material I am reserving further treatment of the species pending
the acquisition of more specimens for a comparative study.
Tmarus cretatus sp. nov.
Figures 1-6
The specific name is a Latin adjective suggested by the chalky
appearance of the body.
Male holotype. Total length, including slightly porrect cheli-
cerae, 3.84 mm. Carapace 1.49 mm long, 1.56 mm wide opposite
chickering: tmarus from Panama
345
interval between second and third coxae where it is widest;
0.47 mm tall and, therefore, less than }/^ as tall as wide. Eyes:
viewed from above, posterior row strongly recurved, anterior row
gently so; central ocular quadrangle wider behind than in front in
ratio of 13 : 10, longer than wide behind in ratio of 15 : 13;
posterior row occupies about 9/13 of width of carapace at that
level; ratio of eyes AME : ALE : PME : PLE = 5.5 : 12 : 4.5 : 13;
AjNIE separated from one another by nearly twice their diameter,
from ALE by a little more than twice their diameter; PME sepa-
rated from one another by a Uttle less than four times their
Tmarus cretatus sp. nov.
Fig. 1. Left palp of male, ventral view.
Fig. 2. Left palpal tibia, retrolateral view.
Fig. 3. Dorsal view of abdomen of male.
Fig. 4. Dorsal view of abdomen of female.
Fig. 5. Epigynum, from below.
Fig. 6. Spermathecae of female paratype, cleared and turned outward to
show internal arrangement.
346 bulletin: museum of comparative zoology
diameter, from PLE by six times their diameter ; laterals separated
by 23^ times the diameter of ALE. Height of clypeus equal to 43^
times the diameter of AAIE. Legs 1234. Palp: essential features
shown in Figures 1-2; tibial apophyses quite distinctive. Ab-
domen: posterodorsal tubercle, prominent in the female, appears
here as only a slight rise. Color in alcohol: general chalky appear-
ance important; first two pairs of femora with numerous white
flecks on prolateral surfaces; venter of abdomen with a median,
longitudinal, light brownish stripe and a narrow, white stripe on
each side (in some paratypes the median stripe is nearly black).
Female paratype. Total length, including bases of chelicerae
and slightly extended posterior spinnerets, 5.84 mm. Carapace
2.08 mm long, 2.21 mm wide opposite interval between second and
third coxae where it is widest. Clypeus somewhat more porrect
than in male; height equal to five times the diameter of AAIE.
Legs 1243. Abdomen: with a prominent posterodorsal tubercle as
indicated in Figure 4. Epigynum : essentially as shown in Figures
5-6. Color: essentially as in male; median, ventral, abdominal
stripe is dark brown.
Type locality. The male holotype is from Summit, Panama
Canal Zone, July, 1950 ; the female paratype was taken in the same
locality in August, 1950. One male paratype was taken at Summit,
C. Z., July, 1950; several female paratypes are in the collection
from Summit, C. Z., July and August, 1950, and Summit Gardens,
C. Z., August, 1954.
Tmarus curvus Chickering
Tmarus curvus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 228, figs.
7-8. The male holotype and female paratype were taken on Barro Colorado
Island, Panama Canal Zone, August, 1936. Paratypes of both sexes were
reported from several localities in the Canal Zone and in Panama proper; all
specimens are in the Museum of Comparative Zoology. Roewer, 1954;
Bonnet, 1959.
The species has been collected during two recent collecting
periods in the Canal Zone.
Tmarus decens O. P.-Cambridge
Tmarus decens O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 98, pi. 12,
fig. 9. The holotype female is in the British Museum (Natural History).
F. P.-Cambridge, 1900; Chickering, 1950; Roewer, 1954; Bonnet, 1959.
This species, known only from the female, has not yet appeared
in my collections and is known only from Bugaba, Panama.
chickering: tmarus from Panama 347
Tmarus decorus sp. nov.
Figures 7-8
The specific name is a Latin adjective suggested by the decora-
tive pattern on the cephalothorax and dorsal areas of the abdomen.
Male holotype. Total length, including somewhat porrect cheli-
cerae, to tip of anal tubercle 4.39 mm. Carapace 1.69 mm long;
1.5 mm wide opposite second coxae where it is widest; about
0.68 mm tall and, therefore, less than half as tall as wide. Eyes:
viewed from above, posterior row strongly recurved, anterior row
sUghtly so; viewed from in front, anterior row almost straight,
measured by centers ; central ocular quadrangle wider behind than
in front in ratio of about 16 : 11; about as wide behind as long.
Ratio of eyes AME : ALE : PME : PLE = 5:11 : 6.5 : 10. AME
separated from one another by nearly 2.5 times their diameter,
from ALE by three times their diameter. PAIE separated from
one another by slightly less than three times their diameter, from
PLE by nearly 4.5 times their diameter. Laterals separated by
nearly three times the diameter of PLE. Clypeus quite porrect;
height equal to about seven times the diameter of AME (only
chitinized area measured); apparently bearing only a single,
slender spine at each ventrolateral angle. Sternum: sternal suture
gently recurved; not extended between fourth coxae which are
separated by about three-fourths of their width. Legs: 1243; with
numerous long, slender spines. Palp: most essential features
shown in Figures 7-8; patella and tibia short; the latter with two
conspicuous apophyses. Abdomen: a slight swelhng may indicate
a more or less prominent posterodorsal tubercle in the female
when the latter is known ; with numerous long, slender spines ; only
a little more than twice as long as wide. Color in alcohol: carapace
yellowish with red and pink decorations; a narrow red marginal
band nearly encircles this part of the body from opposite the
interval between LE to posterior border where there is a central
gap in the colored margin; within the dorsal area there is an
irregular, bright red figure consisting of a series of short radii sur-
rounding a yellowish area; the clypeus bears a pair of relatively
large, irregular, reddish spots; the chelicerae are yellowish with a
central pink spot on each; legs basically yellowish but with many
small red or pink dots and larger reddish blotches; fourth leg and,
to a lesser extent, the third leg with the reddish dots united into
narrow stripes especially on the patellae and tibiae; abdomen
with a series of reddish and white transverse bands extending
laterally and posteriorly; the venter is yellowish with a dusty
brown center.
348 bulletin: museum of comparative zoology
Tyye locality. The holotype male is from Barro Colorado
Island, Panama Canal Zone, February, 1958. There are no para-
types and the female is unknown.
Tmarus humphreyi sp. nov.
Figures 9-13
For some time the males and females here regarded as repre-
senting one species were considered as two. Recently, however,
after studying the paratypes of both sexes it was decided that it
would be safe to combine them. The species is named in honor of
Richard L. Humphrey, M. D., a former student of mine and a
delightful companion on an extended collecting trip in Panama.
Male holotype. Total length, including somewhat porrect
chelicerae, 3.74 mm. Carapace 1.39 mm long, 1.19 mm wide
opposite anterior border of first pair of legs; 0.55 mm tall and,
therefore, a little less than half as tall as wide. Eyes : viewed from
above, posterior row rather strongly recurved, anterior row gently
so ; viewed from in front, anterior row nearly straight, measured by
centers; central ocular quadrangle wider behind than in front in
ratio of 25 : 18, longer than wide behind in ratio of 27 : 25;
posterior row occupies nearly four-fifths of width of carapace at
that level. Ratio of eyes AME : ALE : PIME : PLE = 5 : 10 :
6.5 : 9. AME separated from one another by twice their diam-
eter, from ALE by a slightly greater distance. PAIE separated
from one another by slightly more than twice their diameter,
from PLE by about 3.5 times their diameter. Laterals separated
by about twice the diameter of PLE. Clypeus: quite porrect; with
seven long, slender spines along ventral border; height, including
membranous border, equal to nearly six times the diameter of
AME. Chelicerae and maxillae essentially typical of the genus.
Sternum: bluntly terminated posteriorly and only slightly ex-
tended between fourth coxae which are separated by about two-
thirds of their width; otherwise essentially typical of the genus.
Legs: 1243. Palp: most essential features shown in Figures 9-11.
Abdomen: almost squarely truncate at anterior dorsal border; a
small posterodorsal tubercle is curved downward; dorsum with
numerous slender spines each arising from a small, basal tubercle.
Color in alcohol: carapace reddish brown with many irregular,
dark, oblique stripes along lateral sides; legs yellowish with fine,
black dotting; sternum brownish with darker spots extending from
coxae toward the center; abdomen a dull black dorsally with a
narrow, central, yellowish stripe extending through the anterior
half; lateral sides of abdomen with alternating, irregularly narrow,
chickering: tmarus from Panama
349
dark and light stripes; venter with a broad, nearly black, central
stripe and a narrow, irregular light stripe on each side.
Female paratype. Total length 5 mm, including bases of some-
what porrect chelicerae; carapace 1.76 mm long, 1.45 mm wide
opposite second coxae, 0.84 mm tall and, therefore, a Uttle more
than half as tall as wdde, with the usual supply of long, slender
spines. Eyes: central ocular quadrangle wider behind than in front
in ratio of 15 : 11; about as long as wide behind; ratio of eyes
Tmarus decorus sp. nov.
Fig. 7. Tibia and tarsus of left male palp, ventral view.
Fig. 8. Tibia and tarsus of left male palp, retrolateral view.
Tmarus humphreyi sp. nov.
Fig. 9. Tibia and tarsus of left male palp, ventral view.
Fig. 10. Tibia and tarsus of left male palp, retrolateral view.
Fig. 11. Tibia and tarsus of left male palp, dorsal view.
Fig. 12. Posterior end of abdomen of female.
Fig. 13. Epigynum from below.
350 bulletin: museum of comparative zoology
AAIE : ALE : PME : PLE = 5 : 10 : 6 : 8.5; AME separated from
one another by 2.6 times their diameter, from ALE by nearly three
times their diameter; PAIE separated from one another by sUghtly
less than three times their diameter, from PLE by nearly 14/3 of
their diameter; laterals separated by 2.2 times the diameter of
ALE. Clypeus: strongly porrect; height, including membranous
ventral border, equal to slightly more than six times the diameter
of AAIE. Legs: 1243. Abdomen: with a short posterodorsal
tubercle (Fig. 12); with the usual supply of slender spines each
arising from a minute tubercle. Epigynum: quite distinctive; with
a short, median scape; essentials shown in Figure 13; dissection
reveals a pair of conspicuously coiled spermathecae not seen in
external appearance. Color in alcohol : generally much lighter than
in male ; first and second legs yellowish with many reddish brown,
irregular spots; first and second femora with prolateral surfaces
conspicuously reddish brown with many fine, black dots each
associated with a stiff bristle or spine; third and fourth legs much
lighter; abdomen yellowish brown dorsally with lighter spots and
narrow stripes, with alternating light and darker lateral stripes,
ventrally with a broad, dark brown, median stripe.
Type locality. Alale holotype and female paratype from Gam-
boa, Panama Canal Zone, July, 1954. Paratypes of both sexes
collected at the following localities in the Canal Zone: Summit,
August, 1950; Summit Gardens, August and July, 1954; Gamboa,
July and August, 1954 and January, 1958; Corozal, July, 1954 and
December, 1957; Balboa, i\lay, 1964.
Tmarus impedus sp. nov.
Figures 14-15
The name of the species is an arbitrary combination of letters.
Female holotype. Total length, including the somewhat porrect
chelicerae, 6.11 mm. Carapace 2.41 mm long; 2.34 mm wide
opposite posterior border of second coxae where it is widest;
0.72 mm tall and, therefore, only one-third as tall as wide; ventral
margin regularly rounded from opposite PLE to posterior border;
other features as usual in females of the genus from Panama.
Eyes: viewed from above, posterior row moderately recurved,
anterior row slightly so; viewed from in front, anterior row nearly
straight, measured by centers; central ocular quadrangle wider
behind than in front in ratio of 4 : 3, wider behind than long in ratio
of about 10 : 9. Ratio of eyes AAIE : ALE : PAIE : PLE =
5 : 12 : 7 : 12. AME separated from one another by 3.6 times
chickering: tmarus from Panama
351
their diameter, from ALE by nearly four times their diameter;
PME separated from one another by nearly four times their
diameter, from PLE by nearly five times their diameter; laterals
separated by slightly more than 2.5 times their diameter. Clypeus
quite porrect; height, including membranous ventral border,
nearly equal to 8.5 times the diameter of AME; with the usual
slender spines. Chehcerae, maxillae, and lip, all essentially typical
of females of the genus from Panama. Sternum: scutiform;
moderately convex; longer than wide in ratio of 5 : 4; widest be-
tween second coxae; with many bristles; not extended between
Tmarus impedus sp. nov.
Fig. 14. Abdomen of female, lateral view, right side.
Fig. 15. Epigynum, from below.
Tmarus innotus sp. nov.
Fig. 16. Tibia and tarsus of left male palp, ventral view.
Fig. 17. Tibia and tarsus of left male palp, retrolateral view.
352 bulletin: museum of comparative zoology
fourth coxae which are separated by slightly more than half their
width. Legs: 1243; with numerous spines. Abdomen: very spiny;
anterior, dorsal border slightly raised; slightly anterior to middle
of dorsum a pair of small, conical tubercles arise, each topped by a
fairly robust spine; a little behind the middle of the dorsum 's a
low tubercle bearing numerous spines (Fig. 14). Epigynum: some-
what distinctive (Fig. 15). Color in alcohol: the carapace has a
triangular area which is yellow or whitish with small, irregular,
Hght, reddish brown dots and spots extending from PLE to the
top of the posterior decUvity; from the top of the dechvity a series
of radiating, irregular, whitish lines extend downward over a
yellowish brown background; legs yellowish with many reddish
spots; prolateral surfaces of first two pairs of femora mottled with
white spots; mouth parts yellowish; sternum hght yellowish;
abdomen with dorsum and dorsolateral sides a conglomerate of
reddish spots together with irregular, whitish spots, streaks, and
numerous black dots ; the pair of small dorsal tubercles are largely
black; venter with a median, light brownish stripe flanked on each
side by an irregularly dotted white area. It seems probable that
there would be much variation in the color pattern of a large
population of the species.
Type locality. The holotype female is from Summit, Panama
Canal Zone, August, 1950. No paratypes have appeared in my
collection and the male is unknown.
Tmarus ineptus 0. P.-Cambridge
Tmarus ineptus O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 94, pi. 13,
fig. 3. The holotype female is in the British Museum (Natural History).
F. P.-Cambridge, 1900; Chickering, 1950; Roewer, 1954; Bonnet, 1959.
The specimens which I have assigned to this species appear to
represent the most abundant of all of the species now known from
Panama. I have it from many localities in the Panama Canal Zone
and in Panama proper. The P.-Cambridges had the species only
from Bugaba, Panama. A comparison of the figures published by
the two P.-Cambridges, together with an examination of the single
female on loan from the British INIuseum (Natural History) and
the specimens in my own collection, poses serious questions regard-
ing my previous treatment of the species. The two P.-Cambridges
represented the epigynum quite differently in their respective
illustrations. It now seems quite possible that it will be necessary
to recognize the species in my collection as a new species. Until I
have an opportunity to study all of the specimens now in the
chickering: tmarus from Panama 353
P.-Cambridge collection and, if possible, to collect in the vicinity
of Bugaba I do not think it advisable to take a more positive stand
in regard to the matter.
Tmarus innotus sp. nov.
Figures 16-17
The name of the species is an arbitrary combination of letters.
Male holotype. Total length, including somewhat porrect cheli-
cerae, 2.8 mm. Carapace 1.1 mm long; 1.08 mm wide; 0.54 mm
tall. Eyes: viewed from above, posterior row quite strongly re-
curved, anterior row moderately so; viewed from in front, anterior
row slightly procurved, measured by centers; central ocular
quadrangle wider behind than in front in ratio of 26 : 17, wider
behind than long in ratio of 13 : 12; posterior row occupies about
seven -tenths of width of carapace. Ratio of eyes AME : ALE :
PME : PLE = 4:8:5:8. AME separated from one another by
2.5 times their diameter, from ALE by 2.75 times their diameter;
PAIE separated from one another by 3 times their diameter, from
PLE by slightly more than this; laterals separated by 2.25 times
their diameter. Clypeus: wdth a long, slender spine on each side
beneath interval separating AAIE from ALE; height equal to 4
times the diameter of AAIE (membranous ventral border not in-
cluded). CheHcerae, maxillae, lip and sternum all essentially
typical of males of the genus from Panama. Legs: 1243. Palp:
exposed parts much simpler than usual in the genus from Panama ;
tibial apophyses unlike others seen by me ; distinguishing features
shown in Figures 16-17. Abdomen: a little more than tw^ce as long
as wide; posterior end somewhat extended; with a small but
distinct dorsal tubercle probably indicating that the female has
a well developed posterodorsal tubercle extending above the
spinnerets. Color in alcohol: lateral sides of carapace a mottled,
medium reddish brown; clypeus and ocular region yellowish with
numerous light brown spots; dorsal area of carapace with a some-
what triangular, yellowish area with apex extending just over the
top of the posterior declivity and including brownish spots of
different sizes and form ; mouth parts and sternum yellowish with
variations. First and second femora with irregular, whitish spots
on prolateral surfaces; first and second metatarsi reddish. Abdomen
with an irregular whitish stripe in middorsal region; remainder of
dorsum and lateral regions yellowish with minute reddish dots at
bases of numerous spines; venter yellowish white.
354
bulletin: museum of comparative zoology
Type locality. Male holotype from Barro Colorado Island,
Panama Canal Zone, July 30, 1954. There are no paratypes and
the female is unknown.
Tmarus innumus sp. nov.
Figures 18-20
The name of the species is an arbitrary combination of letters.
Female holotype. Total length, including sUghtly porrect cheli-
cerae, 9.95 mm. Carapace 3.25 mm long, 2.89 mm wide, 1.37 mm
tall. Eyes : viewed from above, posterior row moderately recurved,
anterior row slightly so; viewed from in front, anterior row nearly
straight, measured by centers; central ocular quadrangle wider
behind than in front in ratio of about 4:3; wider behind than long
in the same ratio. Ratio of eyes AME : ALE : PME : PLE =
8 : 16 : 9.5 : 14. AME separated from one another by four times
Tmarus ituiumus sp. nov.
Fig. 18. Abdomen of female, dorsal view.
Fig. 19. Posterior end of abdomen of female, lateral view, right side.
Fig. 20. Epigynum, seen from below.
Tmarus levii sp. nov.
Fig. 21. Epigynum, seen from below.
chickering: tmarus from Panama 355
their diameter; from ALE by slightly less than this. PME sepa-
rated from one another by about five times their diameter; from
PLE by nearly six times their diameter. Laterals separated by
twice the diameter of ALE. Clypeus: moderately porrect; height,
including membranous ventral border, equal to nearly nine times
the diameter of AME. Chelicerae, maxillae, and lip all essentially
typical of females of the genus from Panama. Sternum : typical of
females of the genus from Panama; not extended between fourth
coxae which are separated by two-fifths of their width. Legs : 2143 ;
spines on first and second metatarsi shorter, more robust, and more
numerous than usual in females from Panama. Epigynum some-
what distinctive as shown in Figure 20. Carapace brownish with
three pairs of narrow stripes radiating over lateral sides ; posterior
decli\'ity yellowish at top but dark brown elsewhere ; a small brown
circular area surrounds each PAIE. Sternum light brown. Legs
yellowish with many black or gray dots and spots ; first and second
femora conspicuously dotted with black on prolateral and ventral
surfaces; first patellae nearly black ventrally. Abdomen con-
spicuously chalky white dorsally with irregular, blackish bars and
spots in posterior half; lateral sides irregularly white with brown
spots and irregular oblique rows of black dots and short bars;
venter with a light brown, median stripe flanked on each side by a
row of brownish dots.
Type locality. Female holotype from Summit Gardens,
Panama Canal Zone, July 19, 1954; one female paratype from
Summit, C. Z., July 21, 1950. Epigynum of paratype slightly
different from that of the holotype but not significantly so in my
judgment. The male is unknown.
Tmarus intentus 0. P.-Cambridge
Tmarus intentus O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 96, pi. 13,
fig. 1. The holotype male from Guatemala and the female from Bugaba,
Panama, are in the British Museum (Natural History). F. P.-Cambridge,
1900; Chickering, 1950; Roewer, 1954; Bonnet, 1959.
This appears to be a rare species in regions where I have col-
lected. It has not appeared in my collections since 1936 when I
took two females on Barro Colorado Island in July. Some uncer-
tainties appear to exist in respect to this species, however. Two
females on loan from the British Museum (Natural History) differ
quite markedly in general appearance although the epigyna are
closely similar. I hope to be able to study all specimens regarded
by the P.-Cambridges as belonging to this species and, if possible,
to collect in regions from which the species has been reported.
356 bulletin: museum of comparative zoology
Tmarus levii sp. nov.
Figure 21
This species is named after Dr. Herbert W. Levi, Associate
Curator of Arachnology, Aluseum of Comparative Zoology,
Harvard University.
Female holotype. Total length, including bases of chelicerae,
6.37 mm. Carapace 2.4 mm long; 2.05 mm wide; about 0.98 mm
tall; without a distinct thoracic groove; with numerous minute
tubercles each with a long, slender spine. Eyes : viewed from above,
posterior row moderately recurved, anterior row only slightly so;
viewed from in front, anterior row slightly procurved, measured by
centers. Central ocular quadrangle wider behind than in front in
ratio of 25 : 18; wider behind than long in nearly the same ratio.
Ratio of eyes AAIE : ALE : PAIE : PLE = 7 : 10.5 : 6.5 : 10.
AME separated from one another by about 3.5 times their di-
ameter, from ALE by about three times their diameter. PAIE
separated from one another by nearly six times their diameter,
from PLE by slightly less. Laterals separated by about three
times the diameter of PLE. Clypeus : with a row of six spines along
ventral border and with numerous bristles turned toward the
middle line; height equal to nearly eight times the diameter of
AME, including ventral membranous border. Chelicerae, maxil-
lae, and lip all essentially typical of females of the genus from
Panama. Sternum not extended between fourth coxae which are
separated by slightly less than one-third of their width. Legs:
1243; spines essentially as usual in females of the genus from
Panama; trichobothria observed on tibiae, metatarsi, tarsi and
also on palpal tibiae. Abdomen: anterior border with a row of
slender spines; dorsal surface with many slender spines, all ap-
parently erectile. Distinctive features of epigynum shown in
Figure 21. Carapace yellowish with light brown stripes radiating
irregularly from region just above beginning of posterior declivity
together with numerous brownish streaks and spots; posterior
declivity light brownish with nearly white, irregular border.
Clypeus yellowish with many reddish brown dots. Sternum, lip
and maxillae all yellowish. Legs yellowish with many reddish dots
at bases of spines; first and second femora conspicuously dotted
with red on a white background along prolateral surfaces. Ab-
domen: dorsum w^hitish with many fine, dark dots and many
larger, i-eddish dots at bases of spines; venter whitish along a
broad, central stripe and darker laterally with conspicuous,
narrow grooves and semitransparent dots.
chickering: tmarus from Panama
357
Type locality. Female holotype from Barro Colorado Island,
Panama Canal Zone, June, 1950. There are no paratypes and the
male is unknown.
Tmarus longus sp. nov.
Figures 22-23
The name of the species is a Latin adjective meaning extended,
suggested by the conspicuous posterior extension of the abdomen.
Female holotype. Total length 5.33, including somewhat porrect
chelicerae. Carapace 1.67 mm long; 1.36 mm wide; 0.64 mm tall.
Eyes: viewed from above, posterior row moderately recurved,
anterior row slightly so; viewed from in front, anterior row slightly
recurved, measured by centers; central ocular quadrangle wider
behind than in front in ratio of 5 : 3, wider behind than long in
ratio of 15 : 14; posterior row occupies about two-thirds of width
of carapace at that level. Ratio of eyes AME : ALE : PME : PLE
= 4 : 11 : 5.5 : 10. AME separated from one another by slightly
less than three times their diameter, from ALE by slightly less than
four times their diameter; PME separated from one another by
nearly 3.5 times their diameter, from PLE by nearly five times
their diameter; laterals separated by nearly twice the diameter of
ALE. Clypeus: moderately porrect; with a row of seven spines
near ventral margin with the second from each end being very long ;
height equal to about 7.5 times the diameter of AME. Chelicerae,
maxillae, and lip essentially typical of females of the genus from
Panama. Sternum quite convex; with many long, stiff bristles; not
Tmarus longus sp. nov.
Fig. 22. Posterior end of abdomen, lateral view, right side.
Fig. 23. Epigynum, seen from below.
358 bulletin: museum of comparative zoology
extended between fourth coxae which are separated by five-
thirteenths of their width. Legs: 1243; first two pairs of legs
unusually long ; spines are few in number and less conspicuous than
usual in the genus from Panama. Abdomen long and slender;
posterior end conspicuously extended (Fig. 22) ; with many spines.
Epigynum complicated ; with some parts quite obscure ; unlike any
other seen by the author (Fig. 23). Carapace with a somewhat
triangular, yellowish area extending from PLE to top of posterior
declivity which has a narrow white boundary and an incomplete,
white, longitudinal line through the middle; dorsolateral sides
light brownish with white lines and streaks extending ventrally in
somewhat radiate fashion; a fairly broad, white, longitudinal band
extends along lateral sides ; ocular area whitish with a pair of short,
yellowish, longitudinal stripes between PAIE. Palps and chelicerae
a mixture of yellowish and white; other mouth parts yellowish.
First two pairs of legs whitish with brown dots and larger spots,
the latter concentrated along prolateral surfaces of first two pairs of
femora; third and fourth legs yellowish with few reddish brown
dots and larger spots. Abdomen: with a fairly well defined folium
bounded laterally by an irregular, narrow, brown stripe extending
through about three-fourths of length of abdomen; lateral and
ventrolateral sides yellowish white with several narrow, longi-
tudinal folds; venter yellowish with a narrow, irregular, whitish,
ventrolateral border on each side.
Type locality. Holotype female from Panama Canal Zone Forest
Preserve, Panama Canal Zone, July, 1954; one immature paratype
taken in the same locality, January, 1958; the male is unknown.
Tmarus morosus Chickering
Tmarus jnorosus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 235, figs.
13-15. The male holotype and the female paratype are in the Museum of
Comparative Zoology. Roewer, 1954: Bonnet, 1959.
The holotype is from Barro Colorado Island, Panama Canal
Zone; the female paratype is from the Aladden Dam region, C. Z.;
paratypes are in the collection from Barro Colorado Island, C. Z.
Forest Preserve, Summit Gardens, Fort Sherman, and Chilibre, all
in the Canal Zone.
Tmarus mundulus 0. P.-Cambridge
Tmarus mundulus O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 95, pi. 12,
fig. 11. The holotype female from Bugaba, Panama, is in the British
Museum (Natural History). F. P.-Cambridge, 1900; Chickering, 1950;
Roewer, 1954; Bonnet, 1959.
chickering: tmarus from Panama 359
The P.-Cambridges had only females. What are believed to be
both sexes have appeared in my collection in large numbers from
many localities in the Canal Zone.
Tmarus obsecus sp. nov.
Figures 24-29
In spite of some lingering uncertainties it is regarded as fairly
safe to match the sexes as they are presented below. The name of
the species is an arbitrary combination of letters.
Male holohjpe. Total length, including bases of chelicerae,
4.49 mm. Carapace: 1.54 mm long; 1.32 mm wide; 0.57 mm tall.
Eyes : viewed from above, posterior row rather strongly recurved,
anterior row moderately so, not including strongly convex lenses;
viewed from in front, anterior row very slightly procurved, meas-
ured by centers. Central ocular quadrangle wider behind than in
front in ratio of 30 : 21 ; wider behind than long in ratio of 15 : 13.
PLE extend somewhat beyond lateral margins of carapace at their
level. Ratio of eyes AAIE : ALE : FME : PLE = 6:11 : 5 : 9.
AA'IE separated from one another by 11/6 of their diameter, from
ALE by 7/3 of their diameter. PAIE separated from one another
by four times their diameter, from PLE by slightly more than five
times their diameter. Height of clypeus, including membranous
ventral border, equal to 3.5 times the diameter of AME. Cheli-
cerae, maxillae and lip typical of males of the genus in Panama.
Sternum only slightly convex; posterior end bluntly pointed but
not extended between fourth coxae which are separated by nearly
one half their width. Legs: 1243. Palp: distinctive features shown
in Figures 24-26; both tibial apophyses short, blunt. Abdomen:
elongated; lateral sides nearly parallel throughout two-thirds of
length and then sharply narrowed; a sUght widening just before
constriction is taken to indicate a tendency to inflate as in the
female. Carapace with steep posterior declivity a fairly uniform
brown; radiating from top of posterior declivity are several
irregular, yellowish and brown stripes covering most of dorsal
surface and extending laterally to make a complicated color pat-
tern ; clypeus a dark brown ; anterior surface of chelicerae brownish
with yellowish stripes. Sternum brown with elongated darker
spots extending toward center from margin. Legs generally light
brown with many irregular, reddish and yellowish spots; first
femora with a dark brown stripe along prolateral surface; second
femora with a similar stripe but much broken into separate spots;
third and fourth legs much lighter. Abdomen: dorsum brownish
360
bulletin: museum of comparative zoology
in general but this results from a conglomeration of brown streaks
and spots on a light yellowish background ; a median lighter streak
extends through the entire length of the dorsum ; lateral sides with
narrow rows of elongated, brown spots and streaks alternating with
yellowish streaks; venter with a broad, brownish, median, longi-
tudinal stripe flanked on each side by a narrower yello^^dsh stripe.
Tmarus ohseciis sp. nov.
Fig. 24. Tibia and tarsus of left male palp, ventral view.
Fig. 25. Tibia and tarsus of left male palp, retrolateral view.
Fig. 26. Basal, palpal, tarsal apophysis, prolateral view.
Fig. 27. Abdomen of male, dorsal view.
Fig. 28. Abdomen of female, dorsal view.
Fig. 29. Epigynum, seen from below.
chickering: tmarus from Panama 361
Female paratype. Total length, including somewhat extended
chelicerae, 8.13 mm. Carapace 2.47 mni long; 2.2 mm wide; about
0.98 mm tall. Eyes: viewed from above, posterior row moderately-
recurved, anterior row shghtly so; viewed from in front, anterior
row straight, measured by centers. Central ocular quadrangle
wider behind than in front in ratio of about 3:2; wider behind
than long in about the same ratio. Ratio of eyes AME : ALE :
PAIE : PLE = 6 : 13 : 8 : 11. AME separated from one another by
shghtly more than three times their diameter, from ALE by about
3.5 times their diameter. PAIE separated from one another by
slightly more than four times their diameter, from PLE by about
4.75 times their diameter. Laterals separated by nearly twice the
diameter of PLE. Clypeus quite porrect; ventral margin with
numerous spines (some indicated only by scars) ; height, including
membranous ventral border, equal to slightly more than six times
the diameter of AME. Chehcerae, maxillae and lip typical of
females of the genus from Panama. Sternum moderately convex;
not extended between fourth coxae which are separated by a little
more than one-sixth of their width. Legs: 1243. Abdomen:
inflated laterally in posterior half as shown in Figure 28. Carapace
reddish brown with several narrow, yellowish lines and reddish
brown bands radiating from top of posterior declivity; lateral
sides irregularly spotted with reddish brown on a yellowish back-
ground ; steep posterior declivity nearly all reddish brown. Cheli-
cerae brownish with a yellowish stripe along median boundary.
Lip brown, lighter at distal end. Maxillae yellowish in general but
brownish in median halves. Sternum brown. First and second legs
brownish in general with irregular yellowish spots; femora of these
legs with a broad, brown, prolateral stripe throughout; third and
fourth legs much lighter with the femora also with a broad, pro-
lateral stripe much hghter in color. Abdomen: dorsum dark
colored with many small, red dots and larger reddish spots ; extend-
ing through the middle of the anterior half of the dorsum is a
narrow, irregular, dark stripe fringed with yellow ; this is continued
to the anal tubercle by a broken, median yellow stripe; the inflated
region has a broken, yellow bar extending across and down the
lateral sides ; lateral sides with a series of narrow, alternating, black
and reddish, longitudinal stripes; venter with a fairly broad, me-
dian, longitudinal, light brown stripe flanked on each side by a
broader, yellow stripe with dark, irregular dots.
Type locality. Male holotype from Summit, Panama Canal
Zone, August, 1950; female paratype from Barro Colorado
Island, C. Z., January, 1958; one immature female paratype from
Summit, C. Z., November, 1946 (N. L. H. Kraus).
362 bulletin: museum of comparative zollogy
Tmarus parki Chickering
Tmarus parki Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 242, figs.
19-20. The male holotype from Barro Colorado Island, Panama Canal
Zone, is in the Museum of Comparative Zoology. Roewer, 1954; Bonnet,
1959.
One additional male of this species was taken on Barro Colo-
rado Island, C. Z., January, 1958. This specimen has a much
brighter color pattern than was shown by the holotype; the cara-
pace is brightly colored somewhat like that of T. decorus sp. nov. ;
the dorsum of the abdomen has each spine arising from a red dot
surrounded by a yellowish ring. The female is unknown.
Tmarus pauper O. P.-Cambridge
Tmarus pauper O. P.-Cambridge, 1892, Arachnida-Araneida, 1: 96, pi. 12,
fig. 8. The male holotype is in the British Museum (Natural History).
F. P.-Cambridge, 1900; Chickering, 1950; Roewer, 1954; Bonnet, 1959.
The P.-Cambridges had only the male from Bugaba, Panama.
The species has not yet appeared in my collections and, so far as I
know, has not been reported since the original collections were
made.
Tmarus peregrinus Chickering
Tmarus peregrinus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 245,
figs. 21-22. The male holotype is in the Museum of Comparative Zoology.
Roewer, 1954; Bonnet, 1959.
The male holotype was taken on Barro Colorado Island, Panama
Canal Zone, July, 1934 and has not appeared in my collections since
that time. The female is unknown.
Tmarus probus Chickering
Tmarus probus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 247, figs.
23-24. The male holotype together with several paratype males from
several different localities in the Panama Canal Zone and Panama proper
are in the Museum of Comparative Zoology. Roewer, 1954; Bonnet, 1959.
The species has been collected on four different dates since the
publication of my paper in 1950, and all within the Canal Zone.
Dr. James Zetek contributed a specimen taken in El Cermeno,
Panama, January, 1940.
CHICKERING : TMARUS FROM PANAMA 363
Tmarus productus Chickering
Tmarus productus Chickering, 1950, Bull. Mus. Comp. ZooL, 103(4): 249,
figs. 25-26. The male holotype from Porto Bello, Panama, is in the Museum
of Comparative Zoology. Roewer, 1954; Bonnet, 1959.
The species has appeared in my collections only twice in recent
years: Summit, July, 1950, and Summit Gardens, July, 1954,
Panama Canal Zone. It seems possible that the female T. impedus
sp. nov. is the missing female for this species.
Tmarus protobius sp. nov.
Figure 30
The name for this species is an arbitrary combination of letters.
Female holotype. Total length 5.46 mm, including somewhat
porrect chehcerae. Carapace 1 .65 mm long ; 1 .36 mm wide ; 0.77 mm
tall. Eyes: viewed from above, posterior row rather strongly re-
curved, anterior row moderately so, exclusive of the very convex
lenses of ALE; viewed from in front, anterior row gently pro-
curved, measured by centers. Central ocular quadrangle wider
behind than in front in ratio of 29 : 19; slightly wider than long.
Ratio of eyes AME : ALE : PME : PLE = 4.5 : 11.5 : 7 : 10.5.
AME separated from one another by slightly more than twice
their diameter, from ALE by a little more than three times their
diameter. PAIE separated from one another by about 2.5 times
their diameter, from PLE by a little more than 3.5 times their
diameter. Laterals separated by twice the diameter of PLE.
Clypeus: moderately porrect; height, including membranous
ventral border, nearly six times the diameter of AME. Chehcerae,
maxillae, and lip essentially typical of females of the genus from
Panama. Sternum: moderately convex; posterior end sharply
pointed, not extended between fourth coxae which are separated by
five-thirteenths of their width. Legs: 1243. Abdomen : with a well
defined posterodorsal tubercle. Epigynum: somewhat distinctive
(Fig. 30) ; quite unlike that seen in any other species known to me.
Carapace in general hght yellowish but with irregular, brownish
lines radiating from a central light colored region out over dorso-
lateral sides ; posterior decUvity light brownish in center and light
yellowish on lateral sides. Chehcerae: each with a narrow, light
yellowish, lateral stripe and with anterior surface a mixture of
irregular light lines and light brownish spots. Other mouth parts
yellowish. Sternum: yellowish with an irregular row of red dots
along each lateral region. Legs: in general yellowish with many
364 bulletin: museum of comparative zoology
reddish dots, white spots and streaks ; first femora with an irregu-
lar, prolateral, brown stripe with red dots and streaks along
margins; second femora with a whitish, prolateral stripe and many
red dots; ventral surfaces of segments from patellae to tarsi are
similarly but less conspicuously colored. Abdomen : with a dorsal,
light brownish folium bordered by narrow, white lines ; lateral sides
whitish with fine dots arranged in irregular, longitudinal lines;
ventrolaterally there is a series of light brownish, irregular, narrow
stripes; venter hght yellowish with a pair of dotted Unes close to
midventral region and a single dotted line on each side near the
lateral border.
Type locality. Holotype female from Barro Colorado Island,
Panama Canal Zone, August, 1950. One paratype female from
Summit Gardens, C. Z., July, 1954. The male is unknown.
Tmarus rubinus sp. nov.
Figures 31-32
The name of the species is an arbitrary combination of letters.
Female holotype. Total length, including somewhat porrect
chelicerae, 6.11 mm. Carapace 1.96 mm long; 1.69 mm wide;
about 0.98 mm tall. Eyes: viewed from above, posterior row
moderately recurved, anterior row gently so; viewed from in front,
anterior row probably straight (raised cuticle makes it difficult to
measure accurately). Central ocular quadrangle wider behind than
in front in ratio of about 3:2; wider behind than long in ratio of
about 4 : 3. Posterior row of eyes occupies about five-sixths of
width of carapace. Ratio of eyes AAIE : ALE : PAIE : PLE =
5.5 : 12.5 : 7.5 : 10. AAIE separated from one another by slightly
more than three times their diameter, from ALE by about the same
distance. PAIE separated from one another by slightly less than
four times their diameter, from PLE by about the same distance.
Laterals separated by 1.8 times the diameter of PLE. Clypeus
moderately porrect; height equal to about six times the diameter
of AME. Chelicerae, maxillae, and lip essentially typical of
females of the genus from Panama. Sternum: only slightly convex;
not extended between fourth coxae which are separated only by
about one-fifth of their width. Legs: 1 = 234. Abdomen: with
posterodorsal tubercle quite conspicuously developed (Fig. 31);
otherwise essentially typical of females of the genus from Panama.
Epigynum: obscurely distinctive; essentials shown in Figure 32.
Carapace light brownish in general with whitish spots and streaks ;
dorsal radiations, so conspicuous in several species, are here
obscure and poorly defined; posterior decUvity with a whitish spot
chickering: tmarus from Panama
365
32
Tmarus protobius sp. nov.
Fig. 30. Epigynum, seen from below.
Tmarus rubinus sp. nov.
Fig. 31. Posterior end of abdomen of female, lateral view, right side.
Fig. 32. Epigynum, seen from below.
at top with remainder a variable brownish. Sternum hght yellow-
ish. Legs: yellowish in general with many small, reddish brown
spots ; first two pairs of femora conspicuously mottled with reddish
brown along the whole prolateral surface; dorsal and lateral sur-
faces of abdomen light yellowish with many small, reddish and
whitish dots, irregular spots and streaks; venter with a broad,
unicolorous, yellowish, median stripe; lateral sides of the median
stripe irregularly whitish.
Type locality. Female holotype from Summit Gardens, Panama
Canal Zone, July, 1954. There are no paratypes and the male is
unknown.
Tmarus sigillatus Chickering
Tmarus sigillatus Chickering, 1950, Bull. Mus. Comp. Zool., 103(4): 252,
figs. 27-28. The male holotype from the Panama Canal Zone Forest Pre-
serve, C. Z., is in the Museum of Comparative Zoology. Roewer, 1954;
Bonnet, 1959.
Several males have been added to the collection from Summit,
C. Z. Forest Preserve, and Summit Gardens, all in the Canal Zone
and all taken during the past fourteen years. The female remains
unknown.
Tmarus studiosus 0. P.-Cambridge
Tmarus sludiosus O. P.-Cambridge, 1892, Arachnida-Araneida, 1:97, pi. 12,
fig. 6. Types of both sexes are in the British Museum (Natural History),
from Bugaba, Panama. F. P.-Cambridge, 1900; Chickering, 1950; Roewer,
1954; Bonnet, 1959.
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bulletin: museum of comparative zoology
The collection now includes both sexes of this species from many
locaUties in the Canal Zone where it appears to be abundant.
Tmarus vitusus sp. nov.
Figures 33-35
The name of the species is an arbitrary combination of letters.
Male holotype. Total length 4.68 mm. Carapace 1.76 mm long;
1.69 mm wide; about 0.68 mm tall. Eyes: viewed from above,
posterior row quite strongly recurved, anterior row gently so
(exclusive of very convex lenses of ALE) ; viewed from in front,
anterior row almost straight, measured by centers. Central ocular
quadrangle wider behind than in front in ratio of 10 : 7; wider
behind than long in ratio of 5 : 4. Posterior row of eyes occupies
nearly full width of carapace. Ratio of eyes A]\IE : ALE : PME :
PLE = 5 : 11 : 7 : 10. AME separated from one another by 3.6
times their diameter, from ALE by three times their diameter.
PME separated from one another by 3.7 times their diameter,
from PLE by shghtly more than 4.4 times their diameter. Laterals
34
Tmarus vitusus sp. nov.
Fig. 33. Left palp of male, ventral view.
Fig. 34. Left palpal tibia, retrolateral view.
Fig. 35. Posterior end of abdomen, lateral view.
chickering: tmarus from Panama 367
separated by 2.5 times the diameter of PLE. Clypeus: with a row
of six long, slender spines near ventral border and a single similar,
medial spine below the interval separating AME. Chelicerae,
maxillae, and lip essentially typical of males of the genus from
Panama. Sternum moderately convex; squarely terminated half-
way between fourth coxae which are separated by slightly more
than half their width. Legs: 1243. Palp: essential features shown
in Figures 33-34. Abdomen: with a small but definite postero-
dorsal tubercle (Fig. 35), probably indicative of a more or less
prominent tubercle in the female of the species. Carapace with
a nearly triangular area outlined in yellowish enclosing a brownish
region just behind PME; just posterior to the apex of this region
is a somewhat rounded, yellowish area reaching a short distance
down the posterior decUvity; remainder of dorsal region, posterior
declivity and lateral sides are a mottled yellowish and brownish
color; interocular and clypeal region a dotted, yellowish coloration.
Legs yellowish with many brown dots, especially marked on pro-
lateral surfaces of femora, patellae, and tibiae of first and second
legs. Sternum yellowish with fine, brown, irregular dots. Abdomen :
dorsal and lateral sides in general grayish with a darker median
streak extending through anterior half and four narrow, darker,
cross bars poorly outlined; venter with a rounded, brownish spot
in front of genital groove and a median, longitudinal, light brown
stripe bordered by very light areas and extending nearly to the
spinnerets.
Type locality. Male holotype from Gamboa, Panama Canal
Zone, July 24, 1954. There are no paratypes and the female is
unknown.
BIBLIOGRAPHY
Banks, Nathan
1898. Arachnida from Baja California and other parts of Mexico.
Proc. California Acad. Sci., Ser. 3, Zool., 1(7): 205-309, 5 pis.
1929. Spiders from Panama. Bull. Mus. Comp. Zool., 69: 53-96, 4 pis.
Bonnet, Pierre
1959. Bibliographia Araneorum. Toulouse. Vol. 2 (5).
Cambridge, O. P.-, and F. P. -Cambridge
1889-1905. Arachnida-Araneida. Vols. I-II. In: Biologia Centrali-
Americana. Dulau & Co., London.
Chickering, A. M.
1950. The spider genus Tmarus (Thomisidae) in Panama. Bull. Mus.
Comp. Zool., 103 (4): 213-255, 4 pis.
Keyserling, Graf E. von
1880. Die Spinnen Amerikas. 1. Laterigradae. Baur & Raspe, Nurnberg.
368
bulletin: museum of comparative zoology
Petrunkevitch, Alexander
1911. A synonymic index-catalogue of spiders of North, Central, South
America, etc. Bull. Amer. Mus. Nat. Hist., 29: 1-809.
1925. Arachnida from Panama. Trans. Conn. Acad. Arts Sci., 27:
51-248.
ROEWER, C. Fr.
1954. Katalog der Araneae. Brussels, Vol. 2a: 1-923.
(Received 29 June 1965.)
INDEX TO SPECIES
aculeatus 343 levii 356
bucculentus 343 longus 357
cognatus 344 morosus 358
contortus 344 mundulus 358
corruptus 344 obsecus 359
cretatus 344 parki 362
curvus 346 -pauper 362
decens 346 peregrinus 362
decorus 347 probus 362
humphreyi 348 produdus 363
impedus 350 protobius 363
ineptus 352 rubinus 364
innotus 353 sigillatus 365
innumus 354 studiosus 365
intentus 355 vitusus 366
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 8
THE RELATIONSHIPS OF FOUR SMALL HISPANIOLAN
ELEUTHERODACTYLUS (LEPTODACTYLIDAE)
By Albert Schwartz
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
January 20, 1966
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Phylogeny and Evolution of Crustacea. Proceedings of a confer-
ence held at Cambridge, Mass., March 6-8, 1962. H. B. Whittington
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Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 8
THE RELATIONSHIPS OF FOUR SMALL HISPANIOLAN
ELEUTHERODACTYLUS (LEPTODACTYLIDAE)
By Albert Schwartz
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
January, 1966
Bull. Mus. Comp. Zool., Harvard Univ., 133(8): 369-399, January, 1966.
No. 8 — The Relationships Of Four Small Hispaniolan
Eleutherodactylus (Leptodactylidae)
By Albert Schwartz
10,000 SW S4th Street
Miami, Florida 33143
Perhaps the most confusing group of the genus Eleutherodactylus
in the West Indies is that composed of the small to minute frogs
of Hispaniola, none of which has a snout-vent length in excess of
25 mm. To this assemblage belong E. minutus Noble 1923,
E. abbotti Cochran 1923, E. audanti Cochran 1934, and E. haitianus
Barbour 1942 (= E. intermedins Cochran 1941, preoccupied).
E. abbotti was described from Laguna, Samana Province, RepubUca
Dominicana, E. audanti from Peak la Selle (= Mont la Selle),
Dept. de I'Ouest, Haiti. The remaining two forms were described
from the interior uplands of the Cordillera Central in La Vega
Province (Repiiblica Dominicana), minutus from near Paso Bajito,
Jarabacoa-Constanza Trail, and haitianus from Loma Rucilla,
8C00 to 10,0C0 feet. Although minutus and haitianus are still
known only from various localities in the Cordillera, abbotti has
been reported from many localities in the Repubhca Dominicana
and Haiti. E. audanti is here reported for the first time from
outside of the Massif de la Selle.
Shreve and Wilhams (1963: 320-323) discussed at some length
the situation of the species audanti and abbotti in the Port-au-Prince
region. Conclusions drawn from my own experience in the field
in both Haiti and the Republica Dominicana in 1962 and 1963
differ from theirs, and are drawn in part from a large body of fresh
material from critical localities both outside and within the range
of their particular study.
Through the courtesy and cooperation of the following curators,
I have been able to study specimens of this group of frogs : Ernest
E. Williams, Museum of Comparative Zoology (MCZ) ; Doris M.
Cochran, United States National Museum (USNM) ; and Charles
M. Bogert and Margaret Bullitt, American Museum of Natural
History (AMNH). Specimens in my own collection are designated
Albert Schwartz Field Series (ASFS). In the field in Hispaniola
I had the capable assistance of Patricia A. Heinlein and Ronald
F. Klinikowski, Dennis R. Paulson, David C. Leber, and Richard
Thomas. To all of them I express my sincere thanks for their
interest in these small frogs. The illustrations for the present
paper are the work of Klinikowski and Leber; they again merit my
gratitude for their endeavors.
372 bulletin: museum of comparative zoology
ELEUTHERODACTYLUS ABBOTTI AND ELEUTHERO-
DACTYLUS AUDANTI ON THE SOUTH ISLAND ^
Although E. abbotti was known from the south island by only
two doubtfully identified specimens (from Petionville and Fond-
des-Negres, Haiti) at the time of Cochran's monograph on the
herpetology of Hispaniola (1941), it is now known from many
localities there.
Shreve and Williams (1963) in a study of a large number of
small frogs from the La Selle region presented the following con-
clusions: 1) audanti is a subspecies of abbotti because "of the
presence in the lowlands of the Port-au-Prince region and on the
southwest peninsula and in Barahona of equivocal specimens
which appear to be in various grades and degrees intermediate
between audanti and abbotti"; 2) there are no absolute differences
between these two species; 3) at intermediate levels north of the
Massif de La Selle, such as Furcy, there exist populations of frogs,
some of which may be identified as abbotti, others as audanti, and
still others intermediate between these two species; and 4) mate-
rial from various lowland and highland localities (Peninsula de
Barahona, IMorne de Ca3'ette, Petionville, Fond-des-Negres,
Thiotte) cannot be referred with certainty to either abbotti or
audanti and are considered to be intergrades. An interesting
sidelight on Shreve and Williams' study was the discovery of
"the presence in the foothills of the La Hotte region ... of frogs
much more like abbotti than any others in the area south of the
Cul de Sac Plain." This analysis would suggest that abbotti and
audanti are conspecific, and that audanti is restricted to the
highest peaks of the La Selle, intergrades with abbotti at lower
levels, and is replaced by abbotti in the lowlands.
However, while collecting at Furcy in the summer of 1962, it
quickly became obvious that we were dealing with two types of
small frogs. Both are vocally very similar; the call of each is a
series of highpitched "tuck" 's, followed by a sharply ascending
"wheep" (although a series of "tuck-wheep, tuck-wheep" 's may
be interspersed in the normal call series, usually at its end or
beginning). The call of the smaller of these two frogs (audanti)
was distinctly higher than that of the larger (abbotti), but other-
wise the calls were identical in structure. However, these two
forms differ markedly from one another in that abbotti has a
1 The terms "south island" and "north island" are used in this paper as, for
example, by Wilhams, 1961.
SCHWARTZ : SMALL HISPANIOLAN ELEUTHERODACTYLUS 373
grayish green to tan ground color, is without definite leg bars and
postanal triangle (Fig. 1, left), and has an all yellow venter,
whereas audanti has a much more brightly colored dorsum, most
often some shade of reddish brown or buff, with a distinct and
clear-cut postanal triangle and leg bars (Fig. 1, right), and a gray
belly which is often spotted.
Fig. 1. Left: Eleuiherodactylus abbotti, adult female, ASFS X1649, Furcy,
5600', Dept. de I'Ouest, Haiti; snout-vent length 23.0 mm.
Right: Eleutherodactylus audanti audanti, adult female, ASFS X2362, 2.4 mi.
S Kenscoff, Dept. de I'Ouest, Haiti; snout-vent length 23.3 mm.
All the preserved specimens of these frogs collected by others
in the Furcy area can be separated into these two categories
without difficulty. Naturally, the yellow ventral coloration is no
longer present; the distinct leg and postanal markings of audanti
remain, however, and these are a ready means of differentiation
between the two species. Occasional "intermediates" (i.e., audanti
with slightly less clear-cut postanal triangle or abbotti with more
definite leg bands) I consider to be within the natural range of
variation of each of the two species involved; the Furcy popula-
tion is in no way composed of a large number of intergrades with
abbotti and audanti at the two extremes. It is, rather, composed
of frogs which are readily assignable to either audanti or abbotti
on the basis of pattern, with a very occasional specimen of each
whose markings slightly resemble those of the other species in
degree of clarity. Thus, at least at Furcy, audanti and abbotti
appear not as two races of one form, but as two sympatric species.
The following table (Table I) is based on a selection of ten
374 bulletin: museum of comparative zoology
specimens of each sex of each species (using as the criterion for
species the pattern described above) from the south island Haitian
uplands. These series include the largest member of each sex in
each case, and the type (a female) of audanti, as well as three other
adult female paratypes. jMeasurements and ratios indicate the
following: in males, abhotti reaches a larger size than audanti (in
females, the reverse appears to be true, but this is apparently an
artifact of the abhotti sample, since there are female abhotti from
medium elevations in the Cordillera Central which have a snout-
vent length equal to that of the largest audanti); measurements
of femur, tibia, and fourth toe are diagnostic, as is the tibia/
snout-vent ratio.
Table P
(measurements in milHmeters)
Tibia/snout-
Snout-vent Tibia Femur vent ratio
length aver- aver- aver-
(maximum) range age range age range age
abhotti d" 19.3 8.8-10.1 9.3 7.7- 8.5 8.0 48.6-55.4 52.2
audanti (d 18.4 7.2- 8.0 7.6 6.1- 7.0 6.6 40.8-46.8 43.8
abhotti^ 23.6 11.1-13.0 11.9 9.7-11.4 10.2 50.4-56.6 53.5
audanti 9 25.3 9.0-lO.S 10.1 8.0- 9.8 9.1 38.6-47.1 43.6
The following additional descriptive notes on the patterns of
the two species should be helpful. E. audanti is well illustrated
by Cochran (1941: 66). The heavily and distinctly crossbarred
limbs and the dark postanal triangle are clearly showTi. The leg
bars — one on the crus, one on the pes, and two incomplete bars
on the thigh — are usually outlined in preserved material by pale
bands, which set the bars off very distinctly from the ground
color. There is a single bar on the antebrachium, and another on
the wrist, again outUned by pale color. The dorsal pattern may
have a middorsal stripe. The major feature of the dorsal pattern
is a scapular X, the anterior limbs of which are usually fused to
the dark interocular bar; often the area between the interocular
bar and the anterior arms of the X is hkewise dark, thus giving a
1 Head measurements, not included in the table, are less clear-cut. In males
all these measurements overlap greatly, although abhotti averages consistently
higher. In females, abhotti averages larger in head length (8.2 vs. 8.1) and
naris to eye (2.4 vs. 2.2), but slightly less in head width (8.4 vs. 8.5) and
diameter of tj'mpanum (1.4 vs. 1.5); diameter of eye averages the same in
both species (3.0), although the eye of abhotti reaches a larger maximum size.
In no head measurements are the females of the two species separable.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 375
rather complex occipital and scapular figure. The sides may be
dark like the back, or may have a series of about five horizontal
bars before the groin. Two fresh specimens have a tan dorsal
band from snout to vent, with darker brown sides, the scapular X
faint but still present. The ventral dark pigmentation is variable,
but always present; there may be stippling, mottling, or even
blotching on the throat and belly; the underside of the hindUmbs
is always heavily stippled with brown. The pale snout of audanti
is a characteristic of the species.
E. abbotti, on the other hand, is generally paler than audanti
when preserved, and although it has the scapular X which is
joined to the interocular bar, the same number of Umb bands as
audanti, and even, at times, has the vertical side stripes, none of
the features is so bold and diagrammatic as in audanti. The
entire dorsal surface is irregularly mottled with darker color, and
the individual pattern elements are lost in the general obliterative
effect of the dorsal pigmentation. There is no clear-cut postanal
triangle, although the postanal area is somewhat darker than the
rest of the concealed surfaces. The limb bars are not set off from
the ground color by a distinct pale area; in fact, they have been
so much invaded by pale color that they are no longer conspicuous.
Usually the ventral surfaces are almost immaculate, although
there may be some diffuse stippling on the throat, and an occa-
sional individual has some belly stippling.
Cochran (1941 : 67-68) has described in great detail the palhd
dorsal coloration and asymmetrical spotting which occur in some
specimens of audanti. Apparently the entire dorsum loses its basic
pattern, and a blotchy, irregular, asymmetrical pattern is super-
imposed upon the now unicolor back, so that the preserved
specimen is pale, variously and irregularly mottled with dark
brown. Of the large series from Furcy available, only two indi-
viduals show this mottling, and even in them it is apparently in
its earliest stages. The back of one still retains some semblance
of pattern, but there are large licheniform patches on the hind-
limbs. The other is uniform pale pinkish dorsally, but the hind-
legs show expansion of the dark pigment from the crural bands
and elsewhere on the legs. This irregularly mottled state is more
common in specimens from the high La Selle, where large numbers
in any given series may be mottled. The faded back and increased
mottling is not a condition of age, since some tiny juveniles, as
well as adult and subadult males and females all show the mottled
state. Invariably, from any single locality, there are both "normal"
and mottled individuals in the sample.
376 bulletin: museum of comparative zoology
Shreve and Williams (1963: 322) stated that "nothing . . .
comparable to the orange or asymmetrically pigmented specimens
of audanti occurs in the abbotti populations north of the Cul de
Sac." However, there is a single specimen of abbot li from the
lowlands near Pimentel in the Repubhca Dominicana (and thus
well outside the range of audanti) which clearly shows this con-
dition. In addition, there are frogs (for example, one from Furcy,
MCZ 33549, one from Savane Zombi, :\1CZ 31953, and two from
10.5 miles south of Cabral, ASFS V71 and V83) which are typical
abbotti in size, proportion, and residual pattern, and show no
audanti influence in these respects, but which, nevertheless, have
the mottled condition of audanti. These could, perhaps, be con-
sidered intergrades. However, there are typical abbotti available
from the same localities, and, of the three localities, only at Furcy
do the two species occur together. Since abbotti, when far outside
audanti genetic influence, may manifest this sort of peculiar
spotting, I regard these specimens as within the chromatic varia-
tion of abbotti.
Since audanti and abbotti are members of the auriculatus group,
both have external vocal sacs, granular bellies, small patch-Uke
vomerine teeth, and enlarged digital discs. I am unable to dis-
tinguish the two species on any structural basis ; one possibiUty is
that the vomerine teeth of abbotti are slightly more oblique than
those of audanti, but this is at best a very subjective character.
Like most members of the auriculatus group, both audanti and
abbotti lack inguinal glands.
E. audanti is known from near the Dominico-Haitian border
(Foret des Pins) west to the vicinity of Pic Alacaya (foothills.
Massif de la Hotte). It is restricted to the uplands — the lowest
locality whose altitude is known and whence audanti has been
collected is Peneau, 5000 feet. A possibly lower locality is 2.4 miles
south of Kenscoff, but no altitude is available. The highest
locality is Mont Cabaio (7000 feet), although specimens are
recorded from Alont la Selle, without elevation given (the summit
of jNIont la Selle is slightly over 8000 feet). The specimens from
Pic Alacaya (MCZ 21551-53) represent the only Haitian audanti
outside of the Massif de la Selle ; they do not differ in pattern from
La Selle specimens. The adult female of the series has a tibia/
snout-vent ratio of 48.9, slightly greater than that reported above
for La Selle frogs.
In the La Selle region, abbotti occurs with audanti at elevations
up to 5600 feet (Peneau, Furcy, 2.4 miles south of Kenscoff).
There are specimens of abbotti from Savane Zombi (4200 feet) but
SCHWARTZ : SMALL HISPANIOLAN ELEUTHERODACTYLUS 377
none from Foret des Pins at 5800 feet where audanti has been
collected. E. abhotti has also been collected from the lower southern
slopes of the La Selle (one specimen, Thiotte, about 3000 feet);
from the northern lowlands of the eastern Tiburon Peninsula (one
specimen, IMorne de Cayette) ; from the western extremity of the
Tiburon, both north (one specimen, Marfranc, 120 feet) and south
(eight specimens, Camp Perrin, 1000 feet) of the ^Massif de la
Hotte; and from the intermediate southern slopes of this range
(five specimens, Carrefour Canon, 500 feet). (See Fig. 5 for
distribution of these two species.)
A large amount of fresh material from the Peninsula de Bara-
hona and the Sierra de Baoruco indicates that abhotti occurs there
as well. In this region it has been taken from near sea level
(La Cienaga) up to 3700 feet in the Sierra de Baoruco. In fact,
abhotti is the dominant small frog in this entire region. The
absence of audanti from the Sierra de Baoruco may be more
apparent than real. Since audanti in the La Selle has not been
taken lower than about 5000 feet, it may well not occur at lower
elevations in the Sierra de Baoruco. Most collecting in these
mountains has been in the Valle de Polo region, whose elevation is
less than that for the lowest record of audanti to the west.
In summary, E. abhotti is widespread throughout the south
island, occurring from about sea level to elevations of 5600 feet.
The species occurs not only in the lowlands, but in the La Hotte-
La Selle-Baoruco massif up to moderate elevations. E. audanti,
on the other hand, is known only from elevations above 5000 feet
in the massifs de la Hotte and la Selle, and in the latter range is
extremely abundant at these higher elevations. Its occurrence on
the Sierra de Baoruco requires confirmation. ^ Both species occur
together at elevations between 5000 and 5600 feet, at least in the
INIassif de la Selle.
1 Since the above was written, David C. Leber and Richard Thomas, in the
summer of 1964, succeeded in securing E. audanti in the Sierra de Baoruco.
Near the Dominico-Haitian border, eleven specimens of audanti were secured
between 4 and 11 kilometers northeast of Los Arroyos, Pedernales Province,
at elevations between 5600 feet and 7200 feet. In this same general region,
nine specimens of E. abbotti were also collected; these are from six localities
ranging in elevation from 2200 feet to 5800 feet. The latter high elevation
gives an increase of altitudinal overlap between audanti and abhotti of 200 feet
in the southern massifs. At one locality (5 km NE Los Arroyos) both species
were collected together. These new localities are not included on the map,
nor are these specimens included in the computations.
378 bulletin: museum of comparative zoology
ELEUTHERODACTYLUS ABBOTTI AND ELEUTHERO-
DACTYLUS AUDANTI ON THE NORTH ISLAND
E. abhotti has long been known to occur throughout much of
the north island. Described from Laguna on the Peninsula de
Samana, this frog was subsequently reported from many localities
in the Republica Dominicana (Cochran, 1941; Mertens, 1939),
from the Dominico-Haitian border east to the Peninsula de
Samana and the south shore of the Bahia de Samana. Its dis-
tribution in Haiti is poorly known ; it has been reported only from
near Limbe (Lynn, 1958), the Citadelle (Cochran, 1941: 61), and
Grande Riviere (Shreve and Williams, 1963: 322). The relatively
small number of specimens of abhotti in collections prompted
Shreve and Williams to suggest that abhotti w^as "nowhere very
abundant." On the contrary, abhotti is an extremely abundant
frog, somewhat more so in the uplands than in the lowlands. Not
only can it be collected at night, when huge choruses make the
forest resound, but can often be secured with ease during the day
in piles of coconut trash and old, rotting, and very wet piles of
cacao husks.
In the Cordillera Central abhotti occurs up to elevations of 6000
feet (Loma Vieja; 9.1 miles north of Constanza; 9.3 miles north
of Constanza). In the Sierra de Neiba it occurs at elevations as
high as 5000 feet (14.5 miles south of Elias Pina). The Cordillera
elevation is slightly in excess of the highest known records in the
Massif de la Selle, but not strikingly so. The altitude of major
abundance in the Cordillera is apparently about 3600-4000 feet,
where abhotti forms the largest portion of nocturnal frog choruses
in broadleaf gallery forest along rivers in pinewoods.
The measurements of three series of abhotti (ten males and ten
females each) from southern Haiti (La Selle), the interior Domini-
can uplands (Cordillera), and northern Republica Dominicana
are tabulated below (Table II) and reveal certain differences
among them. (Each series included the largest members of both
sexes; in each series most females were gravid.)
In coloration and pattern there appear to be no differences
among the various populations studied. The typical dorsal
ground color varies from gray to some shade of tan or light brown ;
there is a dark interocular bar, crossbars on the limbs, a yellowish
to whitish-gray belly, and a yellow \'ocal sac. Despite its wide
geographic and altitudinal distribution, abhotti has apparently not
differentiated into races.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 379
Table II. Means and extremes of nine measurements and one
ratio in three populations of Ekuthcrodactylus abbotti consisting of
the ten largest specimens of each sex from each region. (No races
are recognized.)
Republica
northern
Dominicana
Republica
Haitian uplands
uplands
Dominicana
lOd"
lOd"
lOcf
snout-vent length
17.9 (
16.4-19.3)
18.5 (17.9-18.9)
17.8 (17.0-18.8)
head length
6.5 (
6.1- 7.0)
6.4 ( 6.0- 6.9)
6.4 ( 6.0- 6.8)
head width
6.6 (
6.0- 7.1)
6.5 ( 6.3- 7.3)
6.6 ( 6.0- 7.4)
tympanum
1.3 (
1.1- 1.4)
1.2 ( 1.0- 1.3)
1.2 ( 1.0- 1.5)
eye
2.6 (
2.5- 2.7)
2.4 ( 2.2- 2.6)
2.7 ( 2.3- 3.0)
naris to eye
1.8 (
1.6- 2.0)
1.9 ( 1.7- 2.0)
1.8 ( 1.7- 2.0)
femur
8.0 (
7.7- 8.5)
7.7 ( 7.1- 8.3)
7.8 ( 7.5- 8.2)
tibia
9.3 (
8.8-10.1)
8.8 ( 8.5- 9.3)
8.8 ( 8.5- 9.3)
fourth toe
8.3 (
7.5- 9.3)
7.6 ( 6.6- 8.1)
7.5 ( 7.0- 7.8)
tibia /snout- vent ratio
43.8 (
40.8-46.8)
47.9 (45.5-51.1)
50.0 (45.7-52.5)
109
109
109
snout-vent length
22.2 (
20.9-23.6)
22.7 (20.4-25.4)
21.4 (20.9-22.8)
head length
8.2 (
7.4- 9.3)
8.1 ( 7.1- 8.8)
7.8 ( 7.5- 8.4)
head width
8.4 (
, 7.9- 9.2)
8.1 ( 6.9- 9.6)
7.9 ( 7.5- 8.7)
tympanum
1.4
; 1.3- 1.6)
1.5 ( 1.2- 1.7)
1.4 ( 1.2- 1.6)
eye
3.0
; 2.8- 3.4)
2.9 ( 2.5- 3.4)
3.0 ( 2.8- 3.2)
naris to eye
2.4 <
: 2.0- 2.8)
2.3 ( 1.8- 2.6)
2.3 ( 2.1- 2.7)
femur
10.2
; 9.7-11.4)
9.9 ( 8.6-10.8)
9.1 ( 8.3- 9.8)
tibia
11.9
;il. 1-13.0)
11.1 ( 9.7-12.5)
10.5 ( 9.9-11.2)
fourth toe
10.3
; 9.0-11.3)
9.4 ( 7.9-10.6)
8.8 ( 8.2- 9.7)
tibia /snout-vent ratio
53.5
(50.4-56.5)
48.9 (45.6-51.3)
49.2 (47.2-51.9)
Two samples of small frogs, from the Sierra de Neiba and the
Cordillera Central, merit special attention. The series from the
Sierra de Neiba consists of nine frogs from three localities ranging
in elevation from 4750 feet to 5950 feet; there are eight adult males
and one juvenile. The Cordillera series is made up of nine adult
male frogs taken while calling ; the elevation for part of this lot is
5000 feet (Valle de Culata). The specimens from 4 miles (7 km)
north of Constanza have no recorded elevation, but are from above
5000 feet. These two small series are distinctly different from
abbotti and are quite like audanti from the Massif de la Selle. The
bold postanal triangle and the conspicuously banded limbs ally
these small frogs with audanti; all are from high elevations. The
voice of the Cordillera specimens resembled the high-pitched calls
380 bulletin: museum of comparative zoology
of audanti more than the lower calls of abhotti. At Valle de Culata
these audanti and abhotti were heard calling in the same abandoned
field, which was grown up in Pteris and blackberries. The call of
audayiti at this locality reminded me very distinctly of that of
E. auricvlatvs in Cuba — a long series of telegraphic chcks
(= "tucks") with an occasional, almost inaudible "wheep" at the
end of the series. The abhotti call at this locality was that typical
for the species throughout its range, and was quite obviously
different from the call of audanti.
It is remarkable that, despite the large number of frogs which
have been collected from the Cordillera Central, audanti is pres-
ently known only from the Valle de Culata region. This valley is
not especially distinctive, being rather small and mostly cut over
or burned for pasture.
E. audanti is an upland species, with apparently isolated popu-
lations; the Sierra de Neiba and Cordillera populations differ not
only from one another but also from the La Selle form. For the
Sierra de Neiba population I propose the name :
Eleutherodactylus audanti notidodes ^ new subspecies
Holotijpe. MCZ 43204, an adult male, from 20 km (11.7 miles)
southwest Hondo Valle, elevation 5950 ft., Independencia Prov-
ince, Repiiblica Dominicana, one of a series taken 11 August 1963
by David C. Leber and Richard Thomas. Original number ASFS
V371.
Parafypes. ASFS V372-74, AAINH 71990-92, same data as
type; ASFS V385, 14.5 km (8.4 miles) SW Hondo Valle, 4750 ft.,
San Rafael Province, Repiiblica Dominicana, 11 August 1983,
R. Thomas; MCZ 43205, 25 km (14.5 miles) S Elias Pina, 5000 ft.,
San Rafael Province, RepiibHca Dominicana, 17 August 1963,
A. Schwartz.
Diagnosis. A subspecies of E. audanti characterized (in males;
females unknown) by larger size than a. audanti (male a. audanti
to 18.4 mm, male a. notidodes to 21.9), longer hind legs, ratio of
tibia/snout-vent length higher (46.5 in notidodes, 43.8 in audanti),
little or no dark ventral pigmentation, and hind leg crossbars
distinct but not prominently set off from ground color by pale
outlining.
Description of type. An adult male with the following measure-
ments (in mm) and ratio : snout-vent length, 21 .9 ; head length, 7.5 ;
' From the Greek noiis, notidis, moisture, wet, and -odes, dweller.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 381
head width, 7.4; diameter of tympanum, 1.2; diameter of eye, 2.7;
naris to eye, 2.2; femur, 7.9; tibia, 10.1; fourth toe, 8.6; tibia/
snout-vent length, 46.1. Head width equal to head length; snout
truncate with nares conspicuous at anterior end of canthus ros-
tralis ; diameter of eye greater than distance from naris to anterior
corner of eye; interorbital space 2.6, about equal to diameter of
eye; diameter of tympanum much less than diameter of eye,
distance from tympanum to eye equal to about three-quarters
diameter of tympanum. Digital discs present, that of digit three
the largest and equal to about two-thirds area of tympanum.
Fingers moderate in length, unwebbed, 3-4-2-1 in order of de-
creasing length; subarticular tubercles well developed, pale gray.
Toes moderate in length, unwebbed, 4-3-5-2-1 in order of decreas-
ing length ; subarticular tubercles dark gray and prominent. Heels
touch when femora are held at right angles to body axis. Dorsum
finely warty or shagreened with a raised median line from snout
to above vent. Throat and belly granular; vocal sac present,
large, extending posteriorly to between foreUmbs, heavily glandular
anteriorly. Inguinal glands absent. Posterior surface of thighs
with large juxtaposed rounded granules. Tongue small, oval,
entire, free behind, its greatest width about one-half that of floor
of mouth. Vomerine teeth in two sharply oblique patches, begin-
ning within the median border of the choanae, and separated from
the choanae by a distance equal to slightly less than the diameter
of a choana, and from each other by a distance equal to the length
of one tooth row.
Coloration of type in life. Dorsal ground color tan with a
darker brown interocular bar and a scapular X; snout, anterior to
interocular bar, slightly paler than back ; area between interocular
bar and two anterior arms of X suffused with darker brown ; back
in general rather uniformly tan, but on the sides this breaks down
into a series of about four lateral bars, separated from one another
by creamy gray from the belly. Hindlimbs tan with two crossbars
on the thigh, neither especially prominent and neither outlined
with pale color; a more conspicuous crural cross-bar, vaguely
outlined with pale color; and a single cross-bar on the pes. Fore-
limbs with one antebrachial bar and a wrist bar. A prominent
dark bar from the snout through the eye to the forelimb insertion
and covering the upper half of the tympanum. A prominent and
very dark brown postanal triangle, extending onto the underside
of the thighs. Ventral ground color creamy gray with scattered
dark brown stippling, most concentrated on the vocal sac and
undersides of the limbs. Iris silvery above.
382 bulletin: museum of comparative zoology
Variation. Variation in measurements and ratios is shown in
Table III. Structurally all the paratypes resemble the type. The
rather widely separated vomerine teeth and the angulated arrange-
ment of the two series in relationship to one another are common
features.
Table III. Means and extremes of three populations of Eleuthero-
daciylus audanti. (Females unknown from Sierra de Neiba and
Cordillera Central. Sample from Haiti consists of the ten largest
specimens of each sex.)
E. a. audanti
E. a. notidodes
E. a. melatrigonum
[Haitian uplands]
[Sierra de Neiba]
[Cordillera Central]
lOd"
8cf
9d^
snout-vent length
17.4 (16.2-18.4)
19.9 (17.9-21.9)
18.2 (17.6-20.3)
head length
6.1 ( 5.7- 6.4)
7.0 ( 6.5- 7.5)
6.5 ( 6.0- 6.9)
head width
6.3 ( 5.8- 6.6)
7.3 ( 6.3- 7.9)
6.6 ( 6.1- 7.3)
tympanum
1.1 ( 0.9- 1.3)
1.2 ( 1.0- 1.4)
1.2 ( 0.9- 1.4)
eye
2.4 ( 2.2- 2.5)
2.8 ( 2.4- 3.1)
2.6 ( 2.0- 3.0)
naris to eye
1.7 ( 1.5- 1.8)
2.2 ( 2.0- 2.4)
1.9 ( 1.6- 2.2)
femur
6.6 ( 6.1- 7.0)
7.7 ( 6.9- 8.5)
7.2 ( 6.4- 8.0)
tibia
7.6 ( 7.2- 8.0)
9.9 ( 8.4-10.1)
8.2 ( 7.8- 8.5)
fourth toe
6.6 ( 5.9- 7.1)
8.0 ( 7.2- 8.6)
7.0 ( 6.6- 7.5)
tibia/snout-vent
ratio
43.8 (40.8-46.8)
109
46.5 (44.7-48.3)
45.0 (40.9-47.4)
snout-vent length
23.1 (22.1-25.3)
head length
8.1 ( 7.4- 8.6)
head width
8.5 ( 7.3- 9.2)
tympanum
1.5 ( 1.2- 1.6)
eye
3.0 ( 2.7- 3.2)
naris to eye
2.2 ( 2.0- 2.4)
femur
9.1 ( 8.0- 9.8)
tibia
10.1 ( 9.0-10.8)
fourth toe
9.2 ( 8.4- 9.8)
tibia/snout-vent
ratio
43.6 (38.6-47.1)
In coloration, the paratypes are quite variable. All but one
were tan in life, but only a single specimen has the rather uniform
dorsal pigmentation of the type. In all, the leg bars are more
prominent than in the type, although never outlined boldly with
paler color. Five specimens have a dark tan dorsum, somewhat
irregularly pigmented, with a vague pair of pale reversed paren-
theses, dark interocular bar, scapular X, and pale snouts. Only
one of these normally colored specimens has any indication of
ventral mottling, and this is sparse and confined to the sides of
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 383
the abdomen. All the other individuals have only faint dark
stippling vcntrally. Two males have a pinkish orange dorsum
with irregular dark brown mottling on the limbs, sides, sacrum,
and head — the typical asymmetrical audanti pattern described in
detail by Cochran. Their dorsal blotching carries over onto the
venter, but is not heavy there. The smallest paratype (snout-
vent 11.1 mm) is pale pink, with prominent limb bars, but with
no irregular dark mottling.
Comparisons. E. a. notidodes requires comparison both with
the nominate race and with E. abhotti, which occurs with it. Com-
pared with a. audanti, notidodes reaches a larger size, and averages
greater in all measurements. There is virtually no overlap of
extremes in measurements of head length, eye, naris to eye,
femur, tibia, and fourth toe. E. a. notidodes has a longer tibia than
does a. audanti, although the overlap of the ratios is great. Both
subspecies are much alike in coloration and pattern; both show
some individuals with asymmetrically blotched dorsal pigmenta-
tion. Both have a dark postanal triangle and both show dark limb
bars, although those of notidodes are not outlined in paler color.
The ventral pigmentation is much heavier in a. audanti; the
notidodes with the heaviest ventral pigmentation is lighter than
the most lightly pigmented audanti.
Compared with male abhotti from the interior highlands, male
notidodes reach a larger size (21.9 vs. 18.9) and are longer snouted
(naris to eye 2.2 [2.0-2.4] in notidodes, 1.9 [1.7-2.0] in abbotti). All
other measurements show a great deal of overlap, although
notidodes averages higher in every measurement except tympanum
diameter (1.2 in both species) and femur (7.7 in both species).
The tibia/snout-vent length ratio averages greater (47.9) in
abbotti than in notidodes (46.5). None of these measurements is so
helpful as pattern for differentiating the two species ; by means of
the brown postanal triangle and the heavily banded legs, they can
be distinguished without difficulty.
The vomerine teeth may prove to be useful in differentiating
the two subspecies of E. audanti. The teeth in a. audanti, although
patch-like, seem to be arranged more horizontally than the series
in notidodes; in the latter subspecies the patches appear to be
more diagonal, the two series directed toward one another poste-
riorly in a broad V.
The small series of E. audanti from the Constanza region in the
Cordillera Central may be known as:
384 bulletin: museum of comparative zoology
Eleutherodactylus audanti melatrigonum ^ new subspecies
Holotype. UCZ 43206, from 7 km (4 miles) north of Constanza,
La Vega Province, Repiiblica Dominicana, one of a series taken
7 July 1963, by David C. Leber and Richard Thomas. Original
number ASFS X8774.
Paratypes. ASFS X8773, X8775-77, same data as type;
AININH 71993-96, 5.1 miles north of Constanza, Valle de Culata,
5000 ft., La Vega Prov., Republica Dominicana, 8 July 1963,
D. C. Leber, R. Thomas.
Diagnosis. A subspecies of E. audanti characterized (in males;
females unknown) by moderate size (male a. audanti to 18.4,
notidodes to 21.9, melatrigonum to 20.3 mm), intermediate length
of tibia, hindleg crossbars distinct and variously outlined in paler
color, although never so boldly as in a. audanti, and pinkish belly
with scattered dark chromatophores.
Description of type. An adult male with the following measure-
ments and ratio: snout-vent length, 20.3; head length, 6.9; head
width, 7.3; diameter of tympanum, 1.4; diameter of eye, 3.0;
naris to eye, 2.0; femur, 7.6; tibia, 8.3; fourth toe, 7.3; ratio of
tibia/snout-vent length, 40.9. Head width greater than head
length; snout truncate with nares conspicuous at anterior end of
canthus rostralis; diameter of eye greater than distance from naris
to anterior corner of eye ; interorbital space 2.8, slightly less than
diameter of eye ; diameter of tympanum much less than diameter
of eye, distance from tympanum to eye equal to about three-
quarters diameter of tympanum. Digital discs present, that of
digit three the largest and equal to about three-quarters area of
tympanum. Fingers moderate in length, unwebbed, 3-4-2-1 in
order of decreasing length; subarticular tubercles well developed,
dark gray. Toes moderate in length, unwebbed, 4-3-5-2-1 in
order of decreasing length, subarticular tubercles dark gray and
prominent. Heels touch when femora are held at right angles to
body axis. Dorsum very finely warty, warts most prominent on
upper eyehds and upper surface of hindlimbs ; a faint, fine, raised
line from snout to above vent. Throat and belly granular; vocal
sac present, large, extending posteriorly to between forelimbs,
heavily glandular, higuinal glands absent. Posterior surfaces of
thighs with large, rounded, juxtaposed granules. Tongue small,
oval, entire, free behind, its greatest width about one-half that of
floor of mouth. Vomerine teeth in two oblique patches, beginning
' From the Greek ryielas, black, and irigonon, triangle.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 385
well within the median border of the choanae, and separated from
the choanae by a distance equal to twice the diameter of a choana,
and from each other by a distance equal to slightly more than the
length of one tooth row.
Coloration of type in life. Dorsal ground color medium tan with
a black interocular bar, a black scapular X, its two anterior arms
forming with the interocular bar a slightly darker occipital tri-
angle ; a dark gray pair of dorsolateral lines and two rather diffuse
sacral spots; the two dorsolateral lines broken up posteriorly to
form a series of three or four dark dorsolateral spots which approach
the vent, forming a dark gray V above it; sides tan, heavily dotted
with dark gray. Hindlimbs tan with two faint crossbars on the
thigh, neither especially prominent and neither outlined with pale
color; a more conspicuous crural crossbar, vaguely outlined with
pale color; and a single dark crossbar indicated on the pes. Fore-
limbs with one antebrachial and one wrist bar. A prominent dark
brown bar from the snout through the eye to the forelimb insertion
and covering the upper half of the tympanum. Ventral ground
color pinkish with some dark gray stippling, especially on the
yellow vocal sac and undersides of limbs. Iris silvery above.
Variation. Variations in measurements and ratios are shown
in Table III (p. 382). The vomerine series are widely separated
and vary in angulation from almost straight to oblique, with the
latter the more common condition.
Three of the paratypes have median dorsal pale hairlines which
were creamy in life; all have some sort of dorsolateral dark mark-
ings, even if the dorsal ground color is dark brown, which separate
the dorsal color from the lateral dotting or spotting. The inter-
ocular bar, scapular X and sacral spots are common features,
although the latter is often almost completely obscured by the
dark dorsal pigmentation. The black to dark brown postanal
triangle is always conspicuous, and the leg and arm bars are like-
wise bold, and at times even outlined with pale color. The ventral
ground color was pinkish in life, with rather uniform dark brown
stipphng which never formed ventral blotches or mottling. No
specimen shows any indication of the pale dorsum and asymmet-
rical dark patches of the other two subspecies, although presumably
this condition occurs.
Comparisons. E. a. 7nelatrigonum is intermediate between a.
audanti and a. notidodes in all measurements except that of tym-
panum diameter. The ratio of tibia/snout-vent length is likewise
intermediate. The localities whence a. 7nelatrigonum is known are,
of course, not intermediate between those of a. audanti and
386 bulletin: museum of comparative zoology
a. notidodes. In both dorsal and ventral pigmentation and pattern,
a. melatrigonum resembles a. notidodes more closely than a. audanti.
The hindlimb bars of a. melatrigonum are slightly more prominent
than those of a. notidodes, and slightly less prominent than those of
a. audanti. No a. melatrigonum has the venter blotched, as is
usually the case in a. audanti. As noted above, there are no speci-
mens of a. melatrigonum available at present showing the pale
dorsum with asymmetrical blotching which occurs in both a.
audanti and a. notidodes.
From E. abbotti, with which E. a. melatrigonum occurs, the latter
can be best distinguished by its prominent postanal triangle and
more conspicuously banded limbs. Male melatrigonum reach a
larger size than do male abbotti. All measurements overlap, at
times rather widely; the measurement with least overlap is that of
the tibia (8.5-9.3 in upland abbotti, 7.8-8.5 in a. melatrigonum).
The ratio of tibia/snout- vent length averages less in a. melatrigonum
(45.0) than in abbotti (47.9), although the extremes overlap. Lower
ratios are consistently those of a. melatrigonum (40.9-47.4), while
higher ratios are those of upland abbotti (45.5-51.1).
Remarks. The discovery of two subspecies of E. audanti out-
side the La Selle-La Hotte massif in southern Haiti indicates
that this species probably occurs throughout the higher mountains
of much of the Repiiblica Dominicana. The Sierra de Neiba
appears to be a favored haven for high-mountain south island
species which have been able to cross the Cul-de-Sac-Valle de Neiba
plain, or to cross the intervening strait when the plain was flooded.
In addition to E. audanti, E. parabates (which is a north island
representative of the south island E. ventrilineatus-E . jugans group)
is another species of frog which has distinctly south island affini-
ties, and occurs only in the Sierra de Neiba. E. audanti, on the
other hand, has been able to extend its range farther north into
the Cordillera Central, and in so doing has been able to cross the
rather xeric but high Valle de San Juan. It is likely that E. audanti
occurs in the central Haitian mountains as well, since these are
continuous with the Dominican Sierra de Neiba and Cordillera
Central; the Haitian mountains are very poorly known herpe-
tologically. It is, of course, possible that audanti is a north island
species, having evolved in the uplands of the Cordillera, and
thence expanded southward into the Sierra de Neiba, across the
Cul-de-Sac into the jMassif de la Selle and thence into the Massif
de la Hotte. Neither explanation is better substantiated than the
other.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 387
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388 bulletin: museum of comparative zoology
ELEUTHERODACTYLUS MINUTUS AND ELEUTHERO-
DACTYLUS HAITIANUS
These two species present an even knottier problem than do
E. ahhotti and E. audanti. Regarded by Cochran as two distinct
species, minutus and haitianus are both known only from the
Cordillera Central in the Republica Dominicana. Both seem to be
small forms, with haitianus the smaller (but see further comments
below). Shreve and WiUiams (1963: 322) regarded haitianus as a
synonym of the prior minutus, and the latter as possibly an upland
subspecies of ahhotti. Cochran (1941 : 26) considered that minutus
had a smooth belly (and was in fact following Noble, 1923: 4, in
his original diagnosis of minutus) and haitianus (= intermedius,
sensu Cochran) had a granular belly. Her figures (pp. 47, 70) show
that rnimdus is a rather long-legged species, reaching a known
snout- vent length of 18 mm, and that haitianus is distinctly short-
legged, reaching a known snout-vent length (in the type) of 21 mm.
Cochran also commented in the text (1941: 48) that the "para-
type" of minutus which she examined had a slightly granular
belly. Shreve and Williams (1963: 323) later noted that the
venter of minutus is coarsely granular, in contrast to the more
finely granular venters of ahhotti and audanti.
Perhaps the best way to unravel this complicated situation is to
discuss the field situation, based on my recent collections only,
and then turn to the nomenclatorial problems. In the Cordillera
Central, primarily to the north of Constanza and between that
town and El Rio, there occurs a small frog with a maximum snout-
vent length of 17.0 mm in males and 19.4 mm in females. This
Eleutherodadylus is thus smaller in both sexes than ahhotti, and
males are smaller than male audanti melatrigonum from the same
general area. The species occurs from 3600 to 6100 feet, in upland
broadleaf forests; males call from herbaceous plants, often terres-
trial bromeliads, not more than one foot above the ground surface.
The call is a single, rising, high-pitched "wheep," almost a pulse,
very faint and insect-like. The dorsal ground color varies from
tan to brown, with darker sides, and occasionally there are reddish
dorsolateral lines separating the dorsal and lateral colors. The
dark postanal triangle is fairly distinct, the legs are banded, but
rather inconspicuously so at times, and there are remnants of a
scapular X. All dorsal pattern elements have a distinctly "muddy"
appearance, with no feature being especially clear-cut or prominent
(Fig. 3, left). Ventrally, the ground color varies from very pale
yellow or cream to gray, and the belly may be either immaculate
or have some dark gray dots scattered over it. In addition to the
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 389
scapular X, there may be a sacral chevron or smudge; the fore-
limbs are usually somewhat reddish orange.
To the southeast of Constanza, centering in the high Valle
Nuevo region, there occurs a small frog with a maximum snout-
vent length of 14.8 mm in males and 16.6 mm in females. This
species occurs from 5600 to 8000 feet, mainly in pine woods, where
it has been found under rocks and logs, and under sheets of moss
along a road cut. One calling male was taken about ten inches
above the ground on a herbaceous leaf near a small pool in the
woods. The call is a descending scale of staccato "beeps." The
dorsal ground color varies from tan to very dark brown, almost
black in many cases; the brachium is pale reddish, and the ankle
often has a reddish brown hue. The dorsal pattern consists of a
Fig. 3. Left: Eleutherodactylus minutus, adult female, ASFS X8938, 16 km
N Constanza, 6000', La Vega Province, Repiiblica Dominicana; snout-vent
length 19.4 mm.
Right: Eleutherodactylus haitianus, adult male, ASFS X8392, 9 km NNE
Valle Nuevo, La Vega Province, Republica Dominicana; snout- vent length
13.9 mm.
band usually outlined in dark brown to black, occasionally with
a median pale hairline. A common variant is the "dead leaf"
pattern — a series of obliterative pale and dark dorsal areas which
render the frog inconspicuous against a varicolored brown back-
ground. The snout is usually pale and sharply set off from the
balance of the back. A series of four dark lateral bars, radiating
from the sacrum, is commonplace (Fig. 3, right). The ventral
ground color varies from pale yellow (especially on the vocal sac
390 bulletin: museum of comparative zoology
in males) to clear white (not gray), and the belly and throat are
very heavily spotted with rather large dark brown to black spots
in almost all specimens. All dorsal pattern elements are sharp and
distinct, including the hindUmb bands which are usually outhned
with tan and stand out boldly from the ground color. There is a
dark postanal triangle which may be distinct or inconspicuous,
depending upon the intensity of the dorsal ground color.
The two forms are almost completely separable on the tibia/
snout-vent length ratio. The larger form from north of Constanza
averages 46.0 (44.5-49.7) in males and 47.3 (43.9-53.1) in females,
whereas the smaller, more southern form averages 41.3 (37.2-45.0)
in males and 40.4 (36.1-43.3) in females.
When all aspects of these frogs are considered — the coloration
and pattern, morphology, habitat, altitudinal distribution, and
vocalization — there seems little doubt that we are dealing with
two distinct species.
The nomenclatorial problems involved in allocating names to
them are somewhat more complex. I have examined the type of
E. minutus, but not that of haitianus; the latter is, however, well
illustrated by Cochran and some pertinent measurements are
given. I have had access to the large series of paratypes of haitianus
and these, although helpful, are so confusing in many ways that
they require special discussion.
The type locality of E. minutus is "near Paso Bajito, Jarabacoa-
Constanza Trail"; Paso Bajito is to the north of Constanza. The
elevation is not excessively high; we estimate it at about 4000 feet.
The tj^pe of minutus (AMNH 11404) is a gravid female with a
snout-vent length of 17.3 mm and a tibia length of 8.0 mm. The
frog is presently much faded, but dorsally there is a pale zone with
a faint scapular X and a faint sacral chevron, distinctly darker
sides, only the vaguest indications of leg bars and an interocular
bar, and a pale belly with very slightly darker spotting. Of the
two species discussed above, the type of minutus agrees best with
the larger form, which occurs to the north of Constanza, in pattern
and size. Additionally, the tibia/snout-vent length ratio in the
type (46.2) falls within the extremes of this ratio in the larger of
the two species (females, 43.9-53.1), and not within this ratio in
the smaller (females, 36.1-43.3). One other factor needs consider-
ation: Noble (1923: 4) described the belly of minutus as smooth.
Examination of the iypQ of minutus shows that the belly is, in fact,
rather faintly granular; in this, again, it agrees with the larger of
the two species discussed above. (Both the species under discus-
sion have granular belhes; that of the smaller species is more
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 391
coarsely and heavily granular than that of the larger.) In con-
sideration of the above data, I have no hesitancy in assigning the
larger species of frog, described in detail above, to minutus.
There remains the possibility, suggested by Shreve and Williams,
that minutus is an upland race of abhotti. Against this suggestion
is the fact that the two, although similar in pattern, are quite
different in intensity of dorsal pigmentation and in size (largest
upland abhotti male 18.9, female 25.4; largest minutus male 17.0,
female 19.4). Vocally they are distinct. Finally, they occur
together at two of our recent localities (9.1 miles north of Con-
stanza; 3.3 miles east of El Rio), and very close together at another
(9.3 miles north of Constanza), where abhotti was taken at 6000
feet, minutus at 5600 feet. There is no evidence of intermsdiate
specimens, although admittedly intergrades might be extremely
difficult to differentiate from the parent populations. Certainly,
however, when minutus and abhotti are collected at the same
locality, there is no difficulty in distinguishing one from the other.
The tibia/snout-vent length ratio of the two overlaps in both
sexes. The means are, however, higher in abhotti, averaging 47.9
(45.0-51.1) in males, 48.9 (45.6-51.3) in females; male minutus
average 46.0 (44.5-49.7), females 47.2 (43.9-53.1).
Turning now to the southern small frogs from the Valle Nuevo
region, it would seem quite logical to assume that these specimens
are haitianus. The type locality, Loma Rucilla, lies about 23.3
miles to the northwest of Valle Nuevo, and the elevation of the
type and Loma Rucilla paratypes is expressly stated as 8000 to
10,000 feet. There are three "lots of paratypes: USNAI 107567,
107569-74, .MCZ 23469-74 (17 specimens), Loma Rucilla; USNM
107575-76, Loma Vieja; USNAI 107578-85, AICZ 23495-500
(27 specimens), Valle Nuevo. The two paratypes from Loma
Vieja are clearly minutus in size, pattern, and tibia/snout- vent
length ratio, and need not be further considered. Ten of the 35
''haitianus" from Valle Nuevo are also minutus; if they are actu-
ally from Valle Nuevo itself, they represent a new altitude record
for minutus of 7600 feet. However, the paratypic series is labeled
as coming from 6000 to 8000 feet, so that the minutus may have
come from within the known altitudinal limits of that species.
Twenty-four of the Valle Nuevo paratypes are like the recent
Valle Nuevo material discussed above, and are remarkable only
in that the series contains females (up to 18.0 snout-vent) and
males (up to 15.8) which are larger than the more recently taken
specimens. The remaining paratype, a male, has a broad, pale
middorsal stripe, a feature not observed in the fresh specimens.
392 bulletin: museum of comparative zoology
The twenty-four paratopotypes from Loma Rucilla are extremely
puzzling. The type specimen was recorded by Cochran (1941 : 71)
to have a snout-vent length of 21, and thus exceeds all Valle Nuevo
specimens in size. Among the paratypes are large females with
snout-vent lengths from 18.0 to 19.7, bridging the gap in size
between the largest female from Valle Nuevo and the holotj^pe.
One of the six males (USNM 107572) is clearly E. minutus (thus
apparently raising the upper altitudinal limit of that species to at
least 8000 feet). Of the five remaining males, four are small and
the fifth very large (snout-vent 17.5), in fact larger than any other
male.
Loma Rucilla frogs lack the heavily spotted venters of the Valle
Nuevo frogs, and have throats which are dark with some paler
flecking. The larger frogs have unmarked venters and dark
throats, the smaller ones spotted venters; the latter group approxi-
mates the recently collected material from Valle Nuevo, although
the frogs are larger and have less ventral spotting. Cochran's
description of the type, "ventral surfaces . . . clouded with
minute darker dots," her drawing of the specimen, and its size
agree with the larger Loma Rucilla frogs in detail.
The nomenclatorial problem resolves itself into how many forms
(species or subspecies, if anj^) are involved. It seems rather unu-
sual, for example, that there should be such a large difference in
size in adult females between Valle Nuevo and Loma Rucilla.
None of the female Valle Nuevo specimens is equal in size to the
type or to the larger of the female paratypes from Loma Rucilla.
The difference in size, correlated with a difference in ventral pig-
mentation, noted among the Loma Rucilla specimens, strongly
suggests that we maj' be dealing with two species of frogs (the
larger species, E. haitianus). Aside from the ventral pigmentation
and size, I am unable to differentiate these two "forms" from one
another. The tibia/snout-vent length ratio of the larger specimens
falls neatly within the extremes of this ratio in specimens from
Valle Nuevo. The teeth appear identical, and there are no external
characteristics which differentiate them when preserved. Literest-
ingly, a single gravid female (MCZ 40813) from Paraje La Cienaga,
Manabao, in La Vega Province, has a snout-vent length of only
14.7, and is clearly much more like the Valle Nuevo frogs than
the larger Loma Rucilla frogs. La Cienaga hes about 7 miles east
of the peak of Loma Rucilla. This single small female suggests
very strongly that the larger Loma Rucilla specimens actually are
specifically distinct from the more southern frogs.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 393
Another possibility is that there are two subspecies of E. haiti-
anus involved, one on Loma Rucilla and the other to the south
and east, the Loma Rucilla frogs characterized by larger size and
much less heavily pigmented venters. Living specimens may be
very distinct, since many colors are evanescent in Eleutherodadylus.
The least hkely solution is that the presumed differences in ventral
pigmentation and size are happenstance artifacts of collecting;
the large number of specimens from both Valle Nuevo and Loma
Rucilla makes this very unlikely.
Without further data in life on the Loma Rucilla populations,
there seems only one course open, that of regarding all these frogs
as E. haitianus, while acknowledging rather important differences
between the northern and southern populations.
70'
-20*
18'-
9 9 tfi io 40
Km.
70'
L_
Fig. 4. Central Republica Dominicana, showing localities for E. minutus
(open circles) and E. haitianus (solid circles). Semi-solid circles represent
localities whence both species have been taken.
394
bulletin: museum of comparative zoology
There is no question that E. haitianus is distinct from E. minutus.
Aside from the habitat and vocal differences discussed above, the
two species are easily differentiated on the basis of the tibia/snout-
vent ratio; this and other meristic characters are shown in Table IV.
That the two species apparently occur together is based entirely
on old material with possibly less accurate and carefully taken
data than we have for more recently collected specimens; in our
experience they do not occur at precisely the same localities. This
is also true for the upper altitudinal limits of minutus; all the
higher elevations are based on old material. Until the older data
can be confirmed, it seems appropriate to regard minutus as having
a lower altitudinal distribution than haitianus, although there is
an overlap of 500 feet, according to freshly taken and carefully
documented material (see Fig. 4 for distribution). There is a need
for rather precise notation of elevation in the Cordillera Central.
A hike covering several thousand feet elevation may well encompass
the altitudinal limits of several species of frogs; the issue may
become quickly clouded if all specimens are labeled as coming
from between the two extremes in elevation without due regard for
more precise altitude.
Table IV. INIeans and extremes of E. minutus and E. haitianus.
For haitianus the ten largest of each sex from the Valle Nuevo
region were used; for minutus all adults of each sex. (Tibia/snout-
vent ratio computed for all specimens of each sex from Valle
Nuevo, regardless of maturity.)
E.
minutus
E. haitianus
S&
lOd"
snout-vent length
16.4 (
15.5-17.0)
13.5 (12.5-14.8)
head length
5.8 (
5.5- 6.1)
4.7 ( 4.4- 5.0)
head width
5.6 (
5.3- 6.0)
4.6 ( 4.2- 5.0)
tympanum
1.0 (
. 0.9- 1.3)
0.9 ( 0.8- 1.3)
eye
2.2 (
, 2.1- 2.4)
1.9 ( 1.7- 2.0)
naris to eye
1.4 (
, 1.2- 1.5)
1.1 ( 1.0- 1.3)
femur
6.6 (
6.3- 7.1)
5.1 ( 4.8- 5.6)
tibia
7.5 (
7.3- 7.7)
5.5 ( 5.0- 5.7)
fourth toe
6.5 (
, 5.8- 6.8)
4.9 ( 4.5- 5.6)
tibia/snout-vent ratio
46.0 (
44.5-49.7)
41.3 (37.2-45.0)
79
109
snout-vent length
17.9 (
16.0-19.4)
14.8 (12.0-16.6)
head length
6.4 (
5.9- 6.9)
5.1 ( 4.7- 5.8)
head width
6.1 (
5.5- 6.6)
5.0 ( 4.2- 5.6)
tympanum
1.1
: 1.0- 1.4)
1.1 ( 0.7- 1.2)
eye
2.4 (
2.1- 2.6)
1.9 ( 1.7- 2.2)
naris to eye
1.5 (
1.3- 1.8)
1.3 ( 1.1- 1.5)
femur
7.6 (
6.9- 8.0)
5.6 ( 4.7- 6.0)
tibia
8.5 (
7.9- 8.8)
5.9 ( 5.5- 6.5)
fourth toe
7.2 (
, 5.7- 7.7)
5.2 ( 4.8- 5.7)
tibia/snout-vent ratio
47.3
;43.9-53.1)
40.4 (36.1-43.3)
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 395
E. haitianus has not been collected with E. audanti. Our col-
lections also indicate that haitianus and abbotti do not occur
together, although the altitudinal ranges of the two overlap by
400 feet. There are specimens of haitianus and abbotti from Loma
Rucilla, the latter from an elevation of 6000 feet. The two species
can be separated by the much shorter tibia of haitianus and the
larger size of abbotti. The tibia/snout-vent length ratios of these
two species in the Cordillera uplands are: male haitianus 41.3
(37.2-45.0), male abbotti 47.9 (45.5-51.1), female haitianus 40.4
(36.1-43.3), female abbotti 48.9 (45.6-51.3).
There are four other small Eleutherodactylus specimens from the
Cordillera Central which are of interest. These are two males and
one female from 11 km (6.4 miles) east of Paso Bajito, 4500 feet
(ASFS X8839-41), and a female from Valle Nuevo (AICZ 23498).
The two males are in some ways very Uke E. abbotti, except that
they are distinctly longer-legged than any male montane abbotti
(tibia/snout- vent length 55.9 and 54.1). They also appear to be
more broad headed. Both have leg bars of the abbotti style, and
an inconspicuous postanal triangle Uke abbotti. They differ from
abbotti in ventral pigmentation, since both have a series of dark
spots along the lower jaw, and additionally one has some dark
ventral dotting. Of the females, one is gravid and has a stiout-
vent length of 18.1, which is rather small for gravid abbotti. These
two frogs, from two widely separated localities (AICZ 23498 is
a paratype of haitianus), are very similar dorsally in that the
pattern consists of a pale snout, a very dark chestnut triangle from
the interocular bar onto the sacrum where it meets, apex to apex,
with another triangle which has its base across the groin. The
ground color is a dull orange-tan. There is a prominent postanal
triangle, but no other hindlimb markings except dark ankles.
Ventrally, these frogs have some faint stippling on the throat and
a series of dark spots along the lower jaw margin. Whether these
two females are correctly associated with the two males above is
unknown. At least the Paso Bajito female has a tibia/snout- vent
length ratio of 48.1, a figure which is included within the known
range of Cordillera abbotti. The Valle Nuevo female, on the other
hand, has a ratio of 40.0, which is far below all Cordillera abbotti
females, and within the extremes of female haitianus, a species
with which it is definitely not associated.
I have not assigned any of the above four specimens to any
species. They may represent one or two new species of small
Cordilleran frogs, or they may be aberrantly long-legged or
pigmented individuals of well-known upland species.
396 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
RELATIONSHIPS OF SMALL HISPANIOLAN FROGS
Of the four species discussed in the present paper, three are
clearly members of the auriculatus group — i.e., E. abbotti, E. aud-
anti, and E. minutus. As in other members of the group, the
presence of a granular venter, enlarged digital discs, short patch-
Uke vomerine series, and an external vocal sac indicate their
affinities. E. haitianus probably should hkewise be considered a
member of this assemblage. It differs from the others in having
much smaller digital discs and a somewhat more squatty habitus.
All four species lack inguinal glands; all four have the peritoneal
covering of the testes completely pigmented with jet black chroma-
tophores, while the peritoneal covering of the ovaries has scattered
black to gray chromatophores. This dark gonadal pigmentation
is a phenomenon which occurs sporadically in Antillean Eleuthero-
dactylus, without apparent regard for the affinities of the species.
For instance, it occurs in the dimidiatus group {jugans, parabates,
ventrilineatus) , in the ricordi group {zugi), as well as in the auricu-
latus group.
There is a possibility that E. haitianus should be assigned to
the varleyi group, in which are presently included (Shreve and
Williams, 1963: 339): varleyi, glandulijeroides, and cubanus. E.
glanduliferoides and varleyi, in addition to having the varleyi group
characters of feebly developed discs, short vomerine series, and
small size, also have prominent inguinal and popliteal glands;
Shreve and Williams commented that varleyi did not possess these
structures, but they are prominent and orange colored in freshly
collected material. I do not know if cubanus possesses glands.
E. varleyi has a pectoral vocal sac; the condition of the vocal sac
is unknown in glanduliferoides, and cubanus apparently lacks a
vocal sac. E. cubanus and glanduliferoides are smooth ventrally;
varleyi was diagnosed as having a granular belly, although the
specimens before me appear to have smooth venters.
E. haitianus resembles the varleyi group members in small size,
pattern, and short vomerine series. However, the venter is
coarsely granular, there are no inguinal or femoral glands, and
there is a vocal sac. Black testes are not found in the species
varleyi, at least. On the basis of all characters, I prefer to regard
haitianus as a member of the auriculatus group, somewhat more
divergent than its near relatives in the Cordillera.
Of the four species involved in the present discussion, abbotti
and audanti are more closely related to one another than to
minutus and haitianus. Likewise, minutus and haitianus are more
closely related, with minutus more hke the abbotti-audanti pair.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 397
As noted before, audanti is presumably an upland derivative of
ahbotti (which is widespread throughout the island) ; audanti has
either evolved in the Massif de la Selle and migrated thence to
the La Hotte, on the one hand, and to the Sierra de Neiba-
Cordillera Central, on the other, or has evolved in the Cordillera
and has moved thence south into the La Selle. More mesic con-
ditions in the lowlands would have faciUtated such movement.
Both minutus and haitianus, occurring, as far as is known, only in
the Cordillera, may well represent a sequential series abbotti-
minutus-haitianus (if audanti was not developed from abbotti in
the Cordillera but is an immigrant there), or abbotti-audanti-
minutus-haitianus (if audanti developed in the Cordillera and is an
immigrant to the La Selle).
Specimens examined
Eleutherodadylus abbotti: Haiti, Dept. du Sud, ASFS X2797,
X2917-22, Camp Perrin; ASFS X3347-51, Carrefour Canon, 500';
MCZ 37729, Marfranc; Dept. de VOuest, ASFS X1363-80, X2006-
08, X2011, Peneau, 5000'; ASFS X1649, X1795-812, X1818,
X1899, i\ICZ 33546-50, 33552-60, 33562-63, 33565, 33568-73,
33576-79, 33581, 33586-87, 34212, 34221, 34223, 34226, 34229,
34231-32, 34234, 34242-46, 34249, 34250-53, 34256, 34258,
34261-62, 34264, 34272, 34275-76, 34278-81, 34283-84, 34287-89,
34291-92, 34295, 34301-02, 34304-05, + 16 unnumbered speci-
mens, 31729, 31730-32, 31797, Furcy, 5600'; ASFS X3868-69,
2.4 mi. S Kenscoff; MCZ 33280, Morne de Cayette; MCZ 36742,
Thiotte; I\ICZ 31952-53, Savane Zombi, Foret des Pins; Dept. du
Nord, MCZ 3100, 3526, Grande Riviere du Nord. Republica
Dominicana, Barahona Prov., ASFS X9642-43, 3.3 mi. NE La
Cienaga; ASFS X9791-94, 0.6 mi. N Las Auyamas, 3000'; ASFS
X9809-15, 1.8 mi. N Las Auyamas, 3400'; ASFS X9914-15, 8 km
NELas Auyamas, 2600'; ASFSX9919, V71-83, 10.5 mi. S Cabral,
3500'; ASFS V152, 24 km SW Barahona, 3700'; MCZ 35779-90,
del Monte's finca, nr. Barahona; MCZ 35791-96, Herrmann's
finca, nr. Paraiso; Dajabon Prov., ASFS V1623-24, 12 km S Loma
de Cabrera, 2400'; Valverde Prov., ASFS V1241, 9 km N La Cruz
de Guayacanes, 1600'; Puerto Plata Prov., ASFS V1689, 8 km E
Imbert, 1100'; I\ICZ 23545-46, 25 km S Puerto Plata; Santiago
Prov., MCZ 23451-55 (8 specimens), Pico Diego de Ocampo;
Espaillat Prov., ASFS V1698, 6 km SE Sabaneta de Yasica; ASFS
V1955, 2 km N Puesto Grande, 2200'; Maria Trinidad Sanchez
Prov., ASFS V1S60, 2 km S El Factor; Samand Prov., ASFS
398 bulletin: museum of comparative zoology
V1914, 11 km E Sanchez; MCZ 23530-31, Sanchez; ASFS V1976-
82, 5 km W Samana; Duarte Prov., ASFS V1823-24, 9 km NW
Pimentel; El Seiho Prov., ASFS X7835-36, X7975-81, 3.5 mi.
S Sabana de la Mar; ASFS X7902-07, 3.3 mi. SW Miches; ASFS
X9267, 2.3 mi. SE Miches; ASFS X9337, 1.4 mi. SE Miches;
La Vega Prov., ASFS X8564-601, X8880-84, 4 km SW EI Rio,
4000'; ASFS X8116, 11.5 mi. E El Rio, 3800'; ASFS X9162-63,
23 km E El Rio, 3050'; ASFS X9197-98, X9225-32, X9240, 6 km
E El Rio, 3600'; ASFS V1735, 14 km SW La Vega, 1600'; ASFS
V1792, 4 km NW La Vega; ASFS V2021, 12 km NE Jarabacoa,
2000'; MCZ 40812, Paraje La Cienaga, IManabao, Municipio
Jarabacoa; MCZ 31129, 40815-18, Constanza; ASFS X8249-54,
1 mi. S Constanza, 4000'; ASFS X8244-48, 7.2 mi. S Constanza,
5000'; ASFS X9085, 11.5 mi. SE Constanza, 5800'; MCZ 23520-21,
Loma Vieja, 6000'; ASFS X8754-64, 5.1 mi. N Constanza, Valle
de Culata, 5000'; MCZ 40811, La Cienaga, Culata; MCZ 30588,
Aserradero Bermiidez, Constanza; ASFS X9796, 9.1 mi. N Con-
stanza, 6000'; ASFS X8949, 16 km N Constanza, 6000'; ASFS
X8826, X8829-30, 6 km W. Constanza, 4250'; ASFS X8892,
X8897, Tireo Abajo; MCZ 40806, 43458-65, La Raima, Con-
stanza; MCZ 23481-82, Loma Rucilla; ASFS X81 26-27, 1.2 mi.
SE Monsenor Nouel, 700'; San Rafael Prov., MCZ 31170-71,
40814, Rancho de la Guardia; ASFS V380-84, 14.5 km SW Hondo
Valle, 4750'; ASFS V532, V536, 25 km S Elias Pifia, 5000'; ASFS
V537, 19 km S Elias Pina, 4000'; ASFS V543-46, 15 km S Elias
Pina, 3400'; San Juan Prov., ASFS V393, 7 km W Vallejuelo,
2600'.
Eleutherodactylus audanti audanti: Haiti, Dipt, du Sud, MCZ
21551-53, foothills, IMassif de la Hotte; Dept. de VOuest, MCZ
34208-11, 34213-20, 34222, 34224-25, 34227-28, 34230, 34233,
34235-41, 34247-48, 34254-55, 34257, 34259-60, 34263, 34265-71,
34273-74, 34277, 34282, 34285-86, 34290, 34296-300, 34303,
34306-07, + 33 mmumbered specimens, 33551, 33561, 33564,
33566-67, 33574-75, 33580, 33582-85, 37728, ASFS X1813-17,
X1819-98,X1900-02, Furcy, 5600'; ASFS X1362,X2009-10, X2012-
13, Peneau, 5000'; ASFS X1313-21, 2.5 mi. S Kenscoff, 5600';
ASFS X2362-64, X3870-72, 2.4 mi. S Kenscoff; MCZ 24280
(5 specimens), USNM 72595-97, Morne Cabaio, 7000'; MCZ
21576-89 + 39 unnumbered specimens, La Visite, La Selle range;
MCZ 19704-08, USNM 95111-13, Mont la Selle; USNM 85009,
"Morne la Selle"; ASFS X1922, X3920-27, Foret des Pins, 5800';
MCZ 31954-63, Marie Claire, Foret des Pins; MCZ 24586-88,
Bois Pin, nr. Marigot.
Eleutherodactylus audanti melatrigonum: Republica Dominicana,
La Vega Prov., MCZ 43206, ASFS X8773, X8775-77, 7 km N
Constanza; AMNH 71993-96, 5.1 mi. N Constanza, 5000'.
SCHWARTZ: SMALL HISPANIOLAN ELEUTHERODACTYLUS 399
Elenfherodactylus audanti notidodes: Repuhlica Dominicana, San
Rafael Prov., MCZ 43204, ASFS V372-74, AMNH 71990-92,
20 km SW Hondo Valle, 5950'; ASFS V385, 14.5 km SW Hondo
Valle, 4750'; MCZ 43205, 25 km S Elias Pina, 5000'.
Ele^dherodaciylus haitianus: Repuhlica Dominicana, La Vega
Prov., USNM 107567, 107569-71, 107573-74, Loma Rucilla,
4000-10000'; MCZ 23469-74 + 11 untagged specimens, Loma
Rucilla; USNM 107578-85, Valle Nuevo, 6000-8000'; MCZ
23499-500 + 14 untagged specimens, 31588-89, Valle Nuevo;
MCZ 40813, Paraje La Cienaga, Manabao, Municipio Jarabacoa;
ASFS X8294-99, X8465, 9 km NE Valle Nuevo, 7400'; ASFS
X8392-94, 9 km NNE Valle Nuevo; ASFS X8339-40, 3 km
NNE Valle Nuevo; ASFS X8461-62, 11 km SE Valle Nuevo,
8000'; ASFS X8994, 5.3 mi. SE Valle Nuevo, 8000'; ASFS X9070-
76, 8.4 mi. SE Valle Nuevo, 7900'; ASFS X8676, 8.9 mi. SE Valle
Nuevo, 8000'; ASFS X9083, 15 km SE Constanza; ASFS X8929,
16 km SE Constanza, 5600'; ASFS X9153, 11.8 mi. SE Constanza,
5800'.
Eleutherodactylus rninutus: Repiiblica Dominicana, La Vega
Prov., USN]\I 107572, Loma Rucilla, 4000-7000'; AMNH 11404,
MCZ 9338, Paso Bajito; ASFS X9241, 12 km E El Rio, 3600';
ASFS X8713, X8795, 9.1 mi. N Constanza, 6000'; ASFS X8790,
X8938-47, 16 km N Constanza, 6000'; ASFS X9145, 12.6 mi. SE
Constanza, 6100'; USNM 107575-76, Loma Vieja, 6000'; MCZ
23495-97 + 6 untagged specimens, Valle Nuevo.
LITERATURE CITED
Cochran, Doris M.
1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177:i-vii,
1-398, 12 pis., 120 figs.
Lynn, W. Gardner
1958. Some amphibians from Haiti and a new subspecies of Eleuthero-
dactylus schmidti. Herpetologica, 14(3):153-57.
Mertens, Robert
1939. Herpetologische Ergebnisse einer Reise nach der Insel Hispaniola,
Westindien. Abh. senckenberg. naturf. Ges., 449:1-84, 10 pis.
Noble, G. K.
1923. Six new batrachians from the Dominican RepubUc. Amer. Mus.
Novit., No. 61:1-6.
Shreve, Benjamin, and Ernest E. Williams
1963. The herpetology of the Port-au-Prince region and Gonave Islands,
Haiti. Part II. The frogs. Bull. Mus. Comp. Zool., 129(5) :302-42,
5 pis.
Williams, Ernest E.
1961. The evolution and relationships of the Anolis semilineatus group.
Breviora, Mus. Comp. Zool., No. 136: 1-7, map.
(Received 7 April 1965.)
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 9
TWO NEW FISHES OF THE MYCTOPHID GENUS
DIAPHUS FROI\I THE ATLANTIC OCEAN
By Basil Nafpaktitis
Museum of Comparative Zoology, Harvard University
CAMBRIDGE, MASS., U.S.A.
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Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 9
TWO NEW FISHES OF THE MYCTOPHID GENUS
DIAPHUS FROM THE ATLANTIC OCEAN
By Basil Nafpaktitis
Museum of Comparative Zoology, Harvard University
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
January, 1966
Bull. Mus. Comp. ZooL, Harvard Univ., 133(9): 401-424, January, 1966.
No. 9 — Two New Fishes of the Mydophid Genus Diaphus from
the Atlantic Ocean
By Basil Nafpaktitis
INTRODUCTION
During the course of a revisionary study of the myctophid genus
Diaphus, representatives of two undescribed Atlantic species were
found. These two, Diaphus bertelseni and Diaphus lewisi, are
described below.
Of the few authors who have seriously studied Diaphus, a
specialized myctophid offshoot, some thought it necessary to split
the complex into various combinations of genera and subgenera
(Goode and Bean, 1895; Fraser-Brunner, 1949; Bolin, 1939,
1959a). Others have maintained the vast array of species within
the single genus Diaphus (Brauer, 1906; Taning, 1918, 1928, 1932;
Parr, 1928, 1929; Kulikova, 1961). I believe that the former ap-
proach has been adopted because of inadequate material and the
taxonomic complexity of the group. For reasons which will be
presented later in this paper, the inclusion of all the sp3cies (except
two or, possibly, three) within the single genus Diaphus is followed
here.
The taxonomic study of the "Diaphus complex" is not simple,
but a reasonable approach can be made. Some of the difficulties
involved are: 1) the close similarity among several species of the
group; 2) the limited number of reliable diagnostic characters;
3) the relatively high degree of intraspecific variation; 4) the frail
nature and generally poor state of preservation of these meso-
pelagic fishes; 5) the frequent differences between the luminous
organs on the anterior part of the head (important diagnostic
characters) of mature specimens and juvenile individuals of the
same species; and 6) the sexual dimorphism frequently found in
the size and occasionally in the number of these head luminous
organs. As a result, the number of nominal species is now close to
one hundred. We do not now know which of these are valid, but
the number of synonyms may prove to be large.
I am indebted to the Smithsonian Institution and to Dr. Robert
Gibbs of the U. S. National Museum (USNM) for providing funds
and facilities for the study of material deposited in that institution.
Dr. E. Bertelsen of the Danish Marine Biological Laboratory
(DMBL) and Dr. C. Richard Robins of the University of Miami
Marine Laboratory (UMML) generously made available funds and
facilities for study at their respective institutions. Dr. Richard
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bulletin: museum of comparative zoology
Backus of the Woods Hole Oceanographie Institution (WHOI),
Dr. William J. Richards of the Washington Bureau of Commercial
Fisheries (WBCF), and Dr. Gerhard Krefft of the Institut fiir
Seefischerei, Hamburg, have kindly provided material in their care.
I am grateful to Dr. Rolf L. Bolin of Stanford University and
Dr. Giles W. Mead of the Museum of Comparative Zoology
(MCZ), Harvard University, for their kindness in reviewing the
manuscript and for offering valuable advice and criticism. Partial
financial support from National Science Foundation Grant
GF 147 is gratefully acknowledged.
Abbreviations of names of luminous organs on the head, and of
body photophores (Fig. 1), are as follows: Dn: dorsonasal; Vn:
So.
fiOpostenor
Fig. 1. Diagram of a generalized Diaphus showing distribution of the
luminous organs of the head and body photophores. Abbreviations after
Tuning, 1928, slightly modified.
ventronasal; So: suborbital; Ant: antorbital, a small triangular
luminous organ present in several species and lying between the
Dn and the anterodorsal aspect of the orbit, not shown in Figure
1; Br: branchiostegal; Op: opercular; PO: thoracic (or pectoral);
PVO: subpectoral; PLO: suprapectoral ; VO: ventral; VLO:
supraventral; SAO : supra-anal ; AOa: anterior anal; AOp: posterior
anal; Pol: posterolateral; Pre: precaudal.
Measurements were made as follows: standard length (s. 1.):
shortest distance between tip of snout and end of hypural; head
length: from tip of snout to extreme posterior margin of opercular
flap; length of upper jaw: from anterior tip of premaxillary to its
posterior end; eye diameter: horizontal distance between opposite
margins of bony orbit; depth of head: vertical through posterior
margin of orbit; depth of body: vertical through base of upper ray
of pectoral fin; depth of caudal peduncle: least vertical depth;
predorsal: shortest distance between tip of snout and origin of
NAFPAKTITIS: NEW MYCTOPHID FISHES 405
dorsal fin; preventral: shortest distance between tip of snout and
base of outermost ray of ventral fin; preanal: shortest distance
between tip of snout and origin of anal fin.
All measurements were made with a pair of dividers and re-
corded in tenths of millimeters. Most measurements were made
under a dissecting microscope. Initial values given in the lists of
measurements are arithmetic means of all specimens measured;
values within parentheses represent extremes. Methods of taking
and presenting measurements conform to those used by Bolin
(1939).
DiAPHUS BERTELSENi new species
Figures 2-5
Holotype. A 49.0 mm standard length specimen, MCZ 43121,
R/V CHAIN, WHOI, cruise 17, station RHB 801, 26 April, 1961,
00°15'S, 18°35'W to 00°15'S, 18°45'W, 10-ft. Isaacs-Kidd Mid-
water Trawl (IKMT), 0-85-0 m depth.
Paratypes. One, 61.0 mm s.l, MCZ 43122, R/V CHAIN,
WHOI, cruise 35, station RHB 971, 22 February, 1963, 02°00'S,
24°57'W to 01°48'S, 24°54'W, 10-ft. IKMT, 0-295-0 m depth.
Two, 21.0-23.0 mm s.l., Dana Collections, R/V DANA station
1223 V, 1 February, 1922, 22°06'N, 84°58'W, ring-trawl, open,
conical net, 300 cm in diam. at opening (E300), 600 m wire out.
Additional material examined. One, 9.8 mm s.l., R/V DANA
station 1191 I, 14 December, 1921, 17°49'N, 64°54'W, 600 m wire
out; one, 11.7 mm s.l, R/V DANA station 1231 II, 6 February,
1922, 24°30'N, 80°00'W, 600 m wire out; one, 8.5 mm s.l., R/V
DANA station 1243 III, 16 February, 1922, 21°04'N, 73°48'W,
300 m wire out; four, 9.0-10.5 mm s.l, R/V DANA station
1256 IV, 4 March, 1922, 17°43'N, 64°56'W, 300 m wire out; one,
9.0 mm s.l., R/V DANA station 1257 IV, 6 March, 1922, 17°43'N,
64°56'W, 300 m wire out; one, 9.0 mm s.l., R/V DANA station
1274 III, 27 March, 1922, 17°43'N, 64°56'W, 600 m wire out. All
these specimens were captured with stramin-nets, open, conical,
200 cm in diam. at opening (S200).
Description. Body deep and short, its depth 3.5-3.8 in standard
length; head large, its length 2.9-3.0 in standard length; diameter
of eye 3.4-3.6 in length of head, 2.3-2.5 in length of upper jaw;
length of snout equal to or slightly shorter than half the diameter
of the eye; upper jaw 1.4-1.5 in length of head and extending less
than one diameter of the eye behind posterior margin of orbit;
posterior margin of operculum moderately pointed, the point
406
bulletin: museum of comparative zoology
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E
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o
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c
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OS
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o
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00
NAFPAKTITIS: NEW MYCTOPHID FISHES
407
above PLO. Origin of dorsal fin somewhat anterior to base of
ventral fin ; origin of anal fin behind end of base of dorsal fin ; pec-
toral fin short, its length 1 .3 in distance between its base and that of
ventral fin; ventral fin just reaching anus; adipose fin somewhat
anterior to end of base of anal fin.
Luminous organs. Dn round, in cup-shaped recess, entirely-
above nostril and well separated from that of opposite side (Fig. 3) ;
Fig. 3. Diaphus bertelseni, front view; semidiagrammatic representation
of the luminou.s organs of the head.
Vn considerably larger than Dn, widest at anteroventral aspect of
orbit, extending slightly around and under the nostril anteriorly,
hardly reaching vertical through anterior margin of lens posteriorly.
A conspicuous strip of darkly pigmented tissue present between
anterior margin of orbit and luminous organs of head. Body
photophores large, close to each other (particularly so in juveniles)
and all well below lateral line; distance between PLO and lateral
line 3-33^ times the distance between PLO and base of upper ray
of pectoral fin; distance between VLO and lateral line 21/^-23^
times the distance between VLO and base of outermost ventral fin
ray; SAO equidistant from each other and on a straight, sub-
vertical line ; distance between lateral line and SAO3 three times the
diameter of that organ; AOai elevated, its lowest margin on or
slightly above line through upper margins of next two organs of
same series; AOao and AOae on a gentle curve with Pol; distance
between lateral line and Pol three times the diameter of that organ;
AOpi above end of base of anal fin ; Prc4 slightly detached from rest
408
bulletin: museum of comparative zoology
of organs in same series and well below lateral line. Large tri-
angular luminous "scale" associated with PLO (especially well
preserved on paratype, MCZ 43122).
Dentition. Both jaws with inner, irregular series of sharp teeth
which are conspicuously larger than those in outer series; those on
posterior part of premaxillary definitely curved forward (Fig. 4).
\
i>.o.
Fig. 4. Lateral view (right side) of posterior parts of jaws, showing den-
tition in Diaphus bertelseni (from holot\pe).
Counts and measurements. The data which follow are based
on the holotype and the three paratypes. Data from non-type
material are identified as such. Dorsal fin 14-15; anal fin 15;
pectoral fin 11; ventral fin 8; gill rakers on first arch 5 + 1 + 12;
scales in lateral line 34-35 ; AO 6 in anterior series and 4 in posterior
series. Eight additional specimens, 8.5-11.7 mm s.l., had 6 + 4 AO,
and a ninth had 7+3 on the left and 6+4 on the right side.
Measurements, in per cent of standard length, are as follows:
length of head 34.4 (33.3-34.8); diameter of eye 9.9 (9.6-10.2);
length of upper jaw 23.8 (23.0-24.4); depth of body 27.0 (26.0-
28.5); depth of caudal peduncle 12.7 (12.2-13.0); predorsal 46.6
(44.9-47.8); preanal 69.9 (67.2-71.4); preventral 50.5 (48.4-52.4).
Affinities. In general shape of body and in size and arrangement
of photophores, Diaphus bertelseni (especially juveniles, Fig. 5) is
quite similar to D. brachycephalus Taning. This similarity is
superficial, however, and there are many striking differences, such
as the lack in D. bertelseni of a So, which is so prominent in D.
brachycephalus; the number and arrangement of the AOa (6, with
the first raised in D. bertelseni, as opposed to 5 in a straight line) ;
the presence in D. bertelseni of a large luminous "scale" at PLO,
the same structure being completely absent in Taning 's species,
NAFPAKTITIS: NEW MYCTOPHID FISHES 409
which also possesses strongly curved and broad-based posterior
premaxillary teeth.
Differences between Diaphus berielseni and D. coeruleus (Klun-
zinger) are found primarily in the shape, size and position of the
Dn; in the head and body dimensions (length of head 2.9-3.0 in
standard length, as opposed to 3.5-3.7 in D. coeruleus; body depth
3.5-3.8 in standard length, as compared to 4.6-5.0 in D. coeruleus) ;
in the length of the upper jaw which, in D. bertelseni, extends less
than one eye diameter behind the posterior margin of the orbit,
while in D. coeruleus "Upper jaw . . . hinder end . . . surpasses the
eye by more than one eye diameter" (Weber and Beaufort, 1913,
p. 169) ; in the number of AOp (4 in D. bertelseni, 5 in D. coeruleus).
/:;^«'feSv?^:'::'>:,..„^., ^ff0>'-^:^ _
Fig. 5. Diaphus bertelseni, juvenile, 9.8 mm standard length; R/V DANA
station 1191 I, 17°49'N, 64°54'\V, 600 m wire out.
From Diaphus taaningi Norman, D. bertelseni differs mainly in
the size and arrangement of photophores (larger and considerably
further below lateral line in D. bertelseni than in D. taaningi), and
in the number of organs in the AO series (6+4, as opposed to 5 + 5
in D. taaningi).
In all cases discussed above, differences in fin ray and lateral line
counts are omitted as they are considered by the writer to be of
minor diagnostic importance.
Distribution. Pattern and limits of distribution of this species
can not now be inferred. The positions of the few stations in which
Diaphus bertelseni was taken (Fig. 6) suggest that the species is a
South Atlantic one. UtiUzing the South Equatorial Current,
members of this group may enter the North Atlantic while remain-
ing confined to the South Atlantic Central Water Mass. It is also
possible that very young individuals are swept away from the
northwestern boundaries of the South Atlantic Central Water
Mass by the North Equatorial Current and are carried further
west and north. The fact that only juveniles were taken in the
Antillean and Florida waters, whereas the two south equatorial
stations yielded adults, may be purely accidental.
410
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
This species is the same as that Hsted as "Diaphus (sensu Fraser-
Brunner) sp. A" by Backus et al., 1965.
The species is named in honor of Dr. E. Bertelsen, Director of
the Danish Marine Biological Institute, Charlottenlund.
Fig. 6. Chart showing the stations at which Diaphus berielseni (solid dots)
and Diaphus lewisi (rectangles) were taken. The solid dot at about 17°N,
64°\V, represents four DANA stations (1191 I, 14 December, 1921; 1256 IV, 4
March, 1922; 1257 IV, 6 March, 1922; 1274 III, 27 March, 1922). Areas of
high productivity are stippled (reproduced to scale from Sverdrup et al., 1942).
Diaphus lewisi new species
Figures 7-11
Holotype. Male, 27.5 mm standard length, Dana Collections,
R/V DANA station 4005 XI, 12 Alarch, 1930, 13°31'N, 18°03'W,
stramin-net, open, conical, 200 cm in diam. at opening (S200),
50 m wire out.
Paratypes. Two, 20.0-28.5 mm s.L, Dana Collections, data as
above for holotype.
Four, 15.6-32.0 mm s.L, MCZ 44000, R/V CHAIN, WHOI,
cruise 35, station RHB 972, 23 February, 1963, 00°03'N, 25°00'W
to 00°15'N, 25°00'W, 10-ft. IKMT, 0-87-0 m depth.
NAFPAKTITIS: NEW MYCTOPHID FISHES 411
One hundred, 18.5-31.8 mm s.L, R/V GERONIMO, WBCF,
cruise 2, station 138, BCF Cat. No. 370, 19 August, 1963, 09°15'S,
07°06'E, mid- water trawl, depth (?). Eighty specimens in MCZ,
MCZ 44001, 20 specimens in USNM, USNM 259155-Fl.
Additional material exainined. One, 22.0 mm s.l., R/V DANA
station 1159 I, 29 October, 1921, 17°55'N, 24°35'W, ring-trawl,
open, conical, 300 cm in diam. at opening (E300), 5000 m wire out;
133, 11.0-20.0 mm s.L, R/V DANA station 4003 VI, 9 March,
1930, 08°26'N, 15°11'W, stramin-net, open, conical, 150 cm in
diam. at opening (S150), 1000 m wire out; one, 14.5 mm s.l.,
R/V DANA station 4004 IV, 11 March, 1930, 10°21'N, 17°59'W,
stramin-net, open, conical, 200 cm in diam. at opening (S200),
100 m wire out; two, 25.0-26.8 mm s.L, R/V CHAIN, WHOI,
cruise 17, station IKT4, 1 April, 1961, 02°10'S, 17°25'W, 10-ft.
IKMT, 1800 m wire out; two, 23.0-25.2 mm s.L, R/V CHAIN,
WHOI, cruise 17, station IKT5, 3 April, 1961, 00°35'S, 11°30'W,
10-ft. IKMT, depth (?); three, 15.2-15.7 mm s.L, R/V CHAIN,
WHOI, cruise 17, station IKT9, 20 April, 1961, 07°15'N, 14°00'W,
10-ft. IKMT, 400 m wire out; one, 20.0 mm s.L, R/V CHAIN,
WHOI, cruise 17, station RHB 801, 26 April, 1961, 00°15'S,
18°40'W, 10-ft. IKMT, 0-85-0 m depth; three, 17.5-26.0 mm
s.L, R/V CHAIN, WHOI, cruise 17, station RHB 803, 1 May,
1961, 09°27'N, 27°45'W, 10-ft. IKMT, 0-275-0 m depth; 94,
10.2-26.0 mm s.L, R/V CHAIN, WHOI, cruise 17, station RHB
804, 1-2 May, 1961, 10°55'N, 29°30'W, 10-ft. IKMT, 0-42-0 m
depth; three, 25.5-28.7 mm s.L, R/V GERONIMO, WBCF,
cruise 2, station 166, BCF Cat. No. 181, 25 August, 1963, 05°49'S,
10°00'E, mid-water trawl, depth (?); six, 21.5-29.0 mm s.L, R/V
GERONIMO, WBCF, cruise 3, station 130, BCF Cat. No. 971,
26 February, 1964, 04°58'N, 00°30'W, mid-water trawl, depth (?);
one, 23.5 mm s.L, R/V GERONIMO, WBCF, BCF Cat. No. 324,
03°28'S, 00°14'W; one, 30.0 mm s.L, R/V PILLSBURY, UMML,
station 10, 25 May, 1964, 05°55'N, 02°52'E to 05°58'N, 02°50'E,
IKMT, gear depth 0-655-1065-0 m; two, 15.0-18.0 mm s.L,
R/V PILLSBURY, UMML, station 36, 29 May, 1964, 03°50'N,
02°37'W, IKMT, gear depth 0-750-0 m; one, 15.2 mm s.L,
R/V PILLSBURY, UMML, station 37, 29 May, 1964, 04°00'N,
02°46'W to 04°05'N, 02°50'W, IKMT, gear depth 0-480-490-0 m;
five, 23.5-26.8 mm s.L, R/V WALTHER HERWIG, Institut
fiir Seefischerei, station 103, 24 March, 1964, 14°30'N, 22°45'W,
IKMT, depth (?); one, 31.0 mm s.L, R/V WALTHER HERWIG,
Institut fiir Seefischerei, station 109, 25 March, 1964, 14°30'N,
19°42'W, IKMT, depth (?); one, 29.2 mm s.L, R/V WALTHER
412 bulletin: museum of comparative zoology
HERWIG, Institut fur Seefischerei, station 129, 4 April, 1964,
09°14'N, 16°00'W, IKMT, depth (?).
Of the additional material examined, those specimens taken by
R/V DANA are deposited in the Dana Collections, Charlotten-
lund, Denmark; the specimens collected by R/V CHAIN and
R/V GERONBIO are deposited in the AICZ; those caught by
R/V PILLSBURY and R/V WALTHER HERWIG are to be
found in the UMjNIL and the Institut fiir Saefischerei, Hamburg,
respectively.
None of the collections examined appears to have been made
with closing nets.
Description. Small form; depth of body 3.8-4.3 in standard
length; length of head 2.9-3.1 in standard length; diameter of eye
3.4-4.2 in length of head and 2.3-2.9 in length of upper jaw; snout
longer than half the diameter of the eye; anterior spine-hke end of
supraorbital ridge extending forward and downward nearly
reaching the posterodorsal aspect of the nostril; upper jaw 1.4-1.5
in length of head and extending one diameter of eye behind poste-
rior margin of orbit ; pterotic spine conspicuous ; posterior margin
of operculum rounded dorsally, moderately pointed posteriorly,
the point at or slightly below PLO. Origin of dorsal fin directly
above or slightly anterior to base of ventral fin; orighi of anal fin
behind end of base of dorsal fin; upper rays of pectoral fin reach
base of ventral fin; ventral fin nearly reaching origin of anal fin;
adipose fin above last 4-5 rays of anal fin.
Luminous organs. Dn apparently fused with Vn, the compound
structure beginning at or somewhat higher than the dorsal margin
of nostril, extending between latter and anterior margin of orbit
and expanding ventrally, its posteroventral tip not reaching
vertical through anterior margin of lens; a strip of black tissue
separates the preorbital luminous complex from the anterior margin
of orbit and extends along the ventral aspect of eye; its posterior
end expands into a small pocket-like structure which protrudes
into the iris behind the vertical through the middle of lens ; inside
the pocket-like structure there is a small, round So. Tha suborbital
organ first appears in individuals 16.0-17.0 mm s.l., and in well
preserved adult specimens it appears connected with tha Dn-Vn
complex by means of a very narrow isthmus of luminous tissue.
Body photophores of medium size and separated from one another
by an interspace at least as wide as the diameter of these organs;
PLO and VLO distinctly closer to pectoral and ventral fin bases,
respectively, than to lateral line; SAO series of three unequally
NAFPAKTITIS : NEW MYCTOPHID FISHES
413
Ik M
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a
o
o
o
C3
+-*
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e
03
lO
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u
l>^
c3
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414
bulletin: museum of comparative zoology
spaced photophores forming a slightly curved and steeply ascend-
ing line ; distance between SAOi and SAO2 1 3^-2 times that between
SAOi and VO5, distance between SAO2 and SAO3 at least 13^ times
that between SAOi and SAO2, SAO3 about its own diameter below
lateral hne; AOai 1-2 times its diameter anterodorsad to A0a2,
last AOa in line with or slightly raised above preceding three or
four organs of same series; Pol directly above last AOa and 3^^-!
times its own diameter below lateral line; AOpi above posterior
end of anal fin base ; Prc4 distinctly detached from rest of organs of
same series and well below lateral hne. A triangular luminous
"scale" present at PLO.
Dentition. Upper jaw with inner series of strongly recurved,
broad-based teeth (Fig. 8) ; lower jaw with inner irregular series of
teeth feebler than those of upper jaw and only slightly curved
forward.
Sexual dimorphism. Alales are easily distinguished from females
by the presence in the former of a conspicuous, roughly triangular
antorbital luminous organ (Ant) immediately above and in contact
with the Dn-Vn complex (Figs. 7 and 9). This sexual character
first appears as a small, diffuse patch of luminous tissue in individ-
uals 18.0-19.0 mm standard length, and attains its maximum size
...■■■■■ ■■ ...-;;^s#gi^P
•■■■ ....--i^s^iiiilt'
i!s^
V^;^^.
^1^
■aS:^
%^^^
^
Fig. 8. Lateral view (right side) of posterior parts of jaws, sliowing den-
tition in Diaphus lewisi.
NAFPAKTITIS: NEW MYCTOPHID FISHES
415
and its definitive outlines at sexual maturity which, in this small
fish, is attained at a size of 27.0-28.0 mm standard length.
Counts and measurements. Meristic data based on 127 speci-
mens are as follows: dorsal fin 13-14 (usually 14); anal fin 14-15
(usually 15); pectoral fin 10-11; ventral fin 8; gill rakers on first
arch 7-8 + 1 + 14-15, total 22-24; scales in lateral line 35-36;
.^^lifet^--......
w
■:0%"---^.
■■ -■ ; -■•.•.*
.i \
b.n.
:iviv;.itv-<.-.v
^^"^^iW^i^m^m^^^^^s,...,
■"^?**f%
■■■■' ■" ■■■"■" ■■■ ■ '-'w^t-'
:y^ ^c X
■o-n.
y6
Fig. 9. Sexual dimorphism in the luminous organs of the head in Diaphus
lewisi. A: male, 32.0 mm standard length; B: female, 27.0 mm standard
length.
416 bulletin: museum of comparative zoology
AOa 5-6 (very rarely 7), AOp 4-5 (very rarely 6). The AO series
show considerable variation in number and arrangement not only
among individuals but also between two sides of the same indi-
vidual, the most common patterns being those shown in Figures 7
and 10.
Fis;. 10. Caudal region of Diaphus leuisi, showing variation in the num-
ber and arrangement of the photophores in the AOa series.
Measurements, in per cent of standard length, based on 40
specimens 15.6-32.0 mm s.l., as folloAvs: length of head 33.6
(32.1-35.0); diameter of eye 8.9 (8.1-10.3); length of upper jaw
23.3 (22.0-24.2) ; depth of body 24.4 (23.0-26.2); depth of caudal
peduncle 10.9 (10.0-12.6); predorsal 46.5 (44.8-48.9); preanal
65.4 (62.0-67.0); pre ventral 47.1 (44.8-48.5).
J>.n.
Fig. 11. Diaphus lewisi, juvenile, 11.5 mm standard length; R/V CHAIN,
WHOI, cruise 17, station RHB 804, 10°55'N, 29°;^0'W, depth 42 m.
Affinities. Diaphus lewisi is a small fish, probably not growing
larger than 32-33 mm in standard length. It belongs to the
Diaphus rafmesquei group which includes D. rafincsquei (Cocco)
and D. holti Taning in the northeastern North Atlantic, D. mollis
Ta,ning, D. brachycephalus Taning, and probably D. theta Eigen-
mann and Eigenmann in the southwestern North Atlantic and, in
NAFPAKTITIS: NEW MYCTOPHID FISHES 417
addition, several other species elsewhere. The main characters
common to the members of this group are the three distinct
luminous organs on the head (Dn, Vn and So) and the strongly
recurved, broad-based premaxillary teeth. D. lewisi, although
deviating from the typical rafinesqiiei pattern of luminous organs
on the head in the fusion of the Dn and Vn, does possess the
peculiar premaxillary dentition characteristic of the group.
Diaphus suborbitalis Weber from the Indo-Pacific area has
luminous organs on the head which seem to be intermediate
between those of the rafinesquei group and those found in D. lewisi.
In Weber's species the Dn and Vn tend to, but have not fused
completely. In many other respects D. suborbitalis is, according to
Weber (1913, p. 91), closely related to D. fulgens Brauer, a typical
species of the rafinesquei group from the Pacific Ocean and one
possibly identical with D. mollis Taning. Further discussion of
possible relationships among the species mentioned in this para-
graph must await a direct comparison of pertinent material.
Distribution. Available data suggest that the area occupied by
Diaphus lewisi is rather well defined (Fig. 6). Within its range
this species is quite common, and the pattern of its distribution is
obviously influenced by the hydrography along the west coast of
Africa. The cold Benguela Current and the continuous upwelling
from depths of 200-300 m account for the remarkably high pro-
ductivity, the relatively low temperature, and the low salinity of
the coastal surface water. These conditions stand in contrast to
those of surface waters of the general South Atlantic tropical and
sub-tropical areas which are warmer, more sahne and low in
nutrient content (Raymont, 19G3). The belt of fertile water
extends approximately 200 km offshore, forming tongues of
gradually diminishing plankton density extending outward from
the coast (Fig. 6). These tongues correspond to the main water
movements that flow away from the coast, as indicated by the out-
ward extension of the isotherms (Sverdrup et al, 1942). The high
productivity of the coastal Benguela Current is reflected in the
rich zooplankton with which, in turn, may be associated the
abundance of marine vertebrate and invertebrate forms (Raymont,
1933).
With a knowledge of the hydrography of the area concerned, the
distribution of Diaphus lewisi and, no doubt, of other mesopelagic
forms becomes meaningful. Relatively abundant along the con-
tinental slope of the African west coast and particularly so in areas
of high productivity, D. lewisi becomes less and less common
towards the west. The northernmost limits of its distribution
418 bulletin: museum of comparative zoology
(about 18°N) more or less coincide with the southern limits of the
North Atlantic Central Water Mass. A rather marked drop in
water temperature at about 18-20°S and in depths of 200-400 m
possibly constitutes a barrier to the spreading of the species further
south along the west coast of Africa.
The species is named in honor of Dr. Robert E. Lewis, Curator
of the i\Iuseum of Natural History, American University of Beirut,
Lebanon.
DISCUSSION
Taxonomy. The taxonomy of the "Diaphus complex," hke that
of other difficult groups, has been arranged and repeatedly re-
arranged, especially on the generic level, with little success in
achieving a stable system. Inadequate material may have been
one of the main reasons for the existence in the literature of so many
genera and subgenera. With the increase in the number and size
of collections, some students of this group began realizing that they
were dealing with the potentially unlimited inventiveness of nature
as regards the size, shape and complexity of the luminous organs
of the head. They also saw that such morphological features as
the "theta" (Greek letter 6) configuration of the body photophores
and the number and arrangement of some of the series of these
organs (PVO, PO, VO and Pre) seemed, with very few exceptions,
highly constant throughout the group. As a result one single genus,
Diaphus, was accepted and used by them.
]\Iyctophid taxonomy has been heavily dependent on the number
and distribution of photophores. It is undoubtedly true that these
characters are of great importance in distinguishing the genera
within the family. However, at the species level the value of these
characters is often questionable. Their inadequacy, in the Diaphus
group at least, becomes obvious as one grows familiar with: a) the
close morphological similarity shown by several species of the group,
and b) the relatively high degree of intraspecific variation which
often masks interspecific differences.
In contrast to the generally conservative diaphid morphology,
the luminous organs of the head show an astonishing variety of
sizes, shapes, patterns and numbers. In most cases the role of these
organs in interspecific differentiation is decisive.
Of the few diagnostically important characters available to the
student of the Diaphus group, some have been consistently over-
looked. To cite one example: In most studies of Diaphus very
little, if any, attention has been paid to dentition, a character the
NAFPAKTITIS: NEW MYCTOPHID FISHES 419
importance of which has been repeatedly emphasized by Bolin
(1939, p. 124; 1959, p. 20). In his detailed description of the genus
Diaphus, as exemphfied by Diaphus rafinesquei, Bolin (1939, p. 124)
calls attention to the pecuUar dentition of this species and suggests
that: "Probably most of the other species now placed in the genus
will require re-allocation since examination of several forms has
failed to reveal a dentition similar to that found in Diaphus
rafinesquei."
Bohn's suggestion subsequently found support in the observation
that those forms with sharply recurved premaxillary teeth also had
three well-defined luminous organs on the head (Dn, Vn and So).
These two features, namely dentition and pattern of luminous
organs, seemed important enough to justify generic distinction of
the species which possessed them. However, further examination
of large numbers of specimens belonging to different diaphid species
has revealed that the premaxillary teeth display a series of gradu-
ally changing shapes. Between the two extremes, straight and
sharply recurved, there exists a series of more or less curved pre-
maxillary teeth. Furthermore, the young of some species (e. g.
Diaphus elucens, D. prohlematicus, etc.) have curved teeth, especi-
ally on the posterior end of the premaxillary. These teeth are
apparently lost and replaced by straight or less-curved ones in
older individuals of the same species. Finally, a generic hne
becomes even more difficult to draw if one considers the existence
of forms like Diaphus lewisi and, probably, D. suborhitalis Weber.
The former species (I have not personally examined D. suborhitalis,
hence I cannot speak with certainty about it), while deviating
considerably from the rafinesquei pattern of luminous organs on
the head, does show the rafinesquei kind of dentition.
The gradual change in shape displayed by the premaxillary teeth
does not at all diminish the diagnostic usefulness of dentition,
especially in the case of juveniles of morphologically very similar
diaphid species. Young individuals of Diaphus fragilis Taning and
D. elucens Brauer, for instance, are very difficult to differentiate.
With the numbers and arrangement of their body photophores
strikingly similar, luminous organs of the head easily confused
(before attainment of their definitive pattern and size), and with
horizontal distributions considerably overlapping, the juveniles of
these two species can be identified on the basis of premaxillary and
particularly dentary teeth shape and size (e. g. lower jaw with
inner series of very large, Avidely but regularly spaced teeth in
Diaphus fragilis, lower jaw with inner irregular series of denser and
much smaller teeth in D. elucens).
420 bulletin: museum of comparative zoology
Turning once again to the character of the luminous organs of
the head, we are faced with a situation somewhat analogous to that
of the teeth. These organs, in their great variety of designs, can,
with some imagination, be arranged in one or more series of pro-
gressively increasing complexity. This has certainly been at-
tempted in the past (Parr, 1928, p. 140). However, sound infer-
ences about evolutionary trends of these organs cannot be made on
the basis of external morphology alone. A careful histological
study of their structure and innervation could probably help us
gain some insight into the evolution of the numerous patterns and
thus, perhaps, lead us to a better understanding of the phylogenetic
relationships among the various species in the group.
Professor Rolf Bolin is of the opinion (personal communication)
that there are several divergent lines within the Diaphus group.
Although I fully agree with his opinion, I feel inclined to disagree
with the idea that these divergent lines are, at the present state
of our knowledge at least, clear enough to justify a division of the
group into more than two genera. An attempt to formalize these
lines would lead either to a large number of genera or, at best, to a
reasonable number of genera but with several species "suspended"
in between. On the basis of the arguments presented above, it is
suggested here that all but the species assigned to the genus
Lobianchia Gatti (e.g. Aethoprora Goode and Bean, Panthophos
Jordan and Hubbs, Lamprossa Jordan and Hubbs, Cavelampus
Whitley) be included within the single genus Diaphus, without
formal division into subgenera (e.g. Hypcrphotops, Panthophos and
Lamprossa, all three erected by Fraser-Brunner in 1949).
Ecology, speciaiion and phenotypic similarity. Some speculation
on the phenotypic similarity among several myctophid forms is in
order here, and should start with a brief consideration of some
aspects of oceanic ecology and of possible ways of speciation
among high-seas forms. Ecological conditions in the sub-tropical
and tropical oceans seem to have been quite stable in geological
time (Marshall, 1963, p. 182). Relative differences in physico-
chemical factors, i. e. temperature, salinity and density, between
adjacent water masses and, below the thermocline, between bathy-
metrically contiguous layers are usually very small. Repeated
invasions of these adjacent environments by populations of a given
species probably result in some of the more adaptable individuals
becoming physiologically adjusted to the small differences. Fol-
lowing this adjustment, which may not require more than slight
adaptive changes in the reproductive physiology of the invaders.
NAFPAKTITIS: NEW MYCTOPHID FISHES 421
colonization of the new habitat, or habitats, is under way. Subse-
quently, one might expect the gene flow between the colonizers and
the parent species to decrease. Finally, perhaps through the
development of preferential mating and competition "... between
[the] contiguous populations in the zone of contact and the subse-
quent elimination of the less well adapted intermediates in this
zone. . . ." (Ebeling, 1962, p. 149, from Fryer, 1959), the initial,
partial isolation of the populations may be substantially reinforced.
Now, if this is the approximate sequence and extent of the major
evolutionary events leading to the formation of new species, then
it would be reasonable to infer that speciation in many deep-sea
fishes does not necessarily involve conspicuous or even readily
noticeable structural modifications.
In considering the great morphological similarity among several
myctophid forms one should also take into account such evolu-
tionary processes as convergence and parallel adaptations. Hubbs
(1941, p. 190) says: "The general tendency of fishes to speciate
along parallel courses in correlation with the temperature and
sahnity of the water is being repeatedly indicated. ..." Finally,
in his discussion of the phenomenon of sibling species, Mayr (1963,
p. 57) calls attention to the evidence, found in recent work in
developmental genetics, indicating that there is a selective pre-
mium on the maintenance of the phenotype. "Any disturbance of
the developmental process by a gene mutation will result in a
selection pressure in favor of other genes that restore development
along the normal, time-tested channels" (Mayr, 1963, from
Lerner, 1954, and Waddington, 1956).
The rather strong emphasis placed above on adaptation to
apparently minor differences in physico-chemical factors seems to
be in conflict with the known ability of adult myctophids to
tolerate wide extremes in temperature, salinity and, perhaps,
several other ecological factors. In the course of their extensive
(several hundred meters) diurnal vertical migrations, these fishes
"may weU be subjected to much greater differences in temperature
than those marking the limits of their distribution" (Fraser-
Brunner, 1949, p. 1020). This conflict may be resolved if we
assume that during a certain period, or periods, of the year these
fishes become, in terms of reproductive physiology, specifically
stenothermal and stenohaline. It may also be that survival of the
gametes, successful fertihzation, or early development fail in the
absence of optimal, species-specific physico-chemical conditions.
If these assumptions are correct we should expect these optimal
422 bulletin: museum of comparative zoology
ecological conditions to delimit the breeding area and hence the
distribution of the species. But, "certainly in the sea we have
many suggestions that the distribution of the species is much
wider in many instances than the distribution of the breeding
population" (Ebeling, 1962, p. 139, from Bullock, 1958). This
brings us to another potent environmental factor, namely the
oceanic current patterns, and their role in the phenomenon of
"expatriation."
Expatriation. Within a breeding area the larvae are restricted
to surface or near-surface waters. Just before, or at the very
beginning of metamorphosis, these larval forms move to deeper
layers. Based on his studies of larval fishes, Taning (1918, p. 20)
writes: "There can . . . hardly be any doubt that the Scopelids
[myctophids], when metamorphosis sets in, undergo a thorough
change, acquiring a different specific gravity to that which they
had as postlarvae, and consequently move down, either actively or
passively, to water layers of a specific gravity suitable to their
requirements during metamorphosis, and later, after the meta-
morphosis is completed, ascend once more to the upper layers.
In other words, we have here a decided instance of (passive or
active) ontogenetic migration." Both as larvae prior to their
"ontogenetic migration" and as young right after their ascent to
the upper layers, myctophid and other forms with similar life
histories are at the mercy of the prevailing current systems in the
area concerned. The larvae and young may either be carried away
from the breeding area or they may be kept restricted within it
(current gyres, eddies, etc.). In the first case the individuals may
spread over a wide range, far away from the breeding area of the
species to which they belong. Sexual maturity will probably be
attained while in environments which, although favorable in
terms of satisfaction of purely indi\'idual needs, are far from meet-
ing the species-specific reproductive requirements. These indi-
viduals are thought to be "expatriated" and reproductively lost to
their populations (Ebeling, 1962, p. 139).
Concerning the fate of the "expatriates," Eknian (1953, p. 317)
remarks that ". . . it remains questionable whether the species is
able to exist independently in the unfavorable region or whether
it wou!d not die out there if it were not continuously reinforced
from the more favorable regions," and ". . . the unfavorable
region is . . . outside the real home of the species, and it [is] . . .
possible to contrast the autochthonous main mass of the species
which lives in the reproductive area with an allochthonous sterile
expatriated contingent in an expatriation area." Bolin (1959b,
NAFPAKTITIS : NEW MYCTOPHID FISHES 423
p. 142), also, points out that "while straggling adults may exist for
long periods in waters far beyond the normal range of the species,
permanent populations are restricted to the proximity of the areas
where spawning can be successful." During my study of the distri-
bution of the diaphid species in the Atlantic Ocean I have found
considerable evidence indicating that many of the diaphid speci-
mens captured off the northeastern coast of the United States are
"expatriates" belonging to species the breeding areas of which are
well within the tropical and sub-tropical waters.
I am well aware of the weaknesses in my assumptions and argu-
ments. I would hke to conclude with a well-worn but nevertheless
appropriate statement — a great deal of work remains to be done,
especially with regard to the distribution and bionomics of this
scientifically and, perhaps, economically important group of
mesopelagic fishes.
LITERATURE CITED
Backus, R. H., G. W. Mead, R. L. Haedrich and A. W. Ebeling
1965. The mesopelagic fishes collected during cruise 17 of the R/V
Chain, with a method for analyzing faunal transects. Bull.
Mus. Comp. ZooL, 134(5): 139-157.
BoLiN, R. L.
1939. A review of the mj'ctophid fishes of the Pacific coast of the United
States and of lower California. Stanford Ichthyol. Bull., 1(4):
89-160.
1959a. Iniomi. Myctophidae from the "Michael Sars" North Atlantic
deep-.sea expedition, 1910. Rep. "Michael Sars" N. Atl. Deep-
sea Exped., 4(7): 1-45.
1959b. Differential bipolarity in the Atlantic and Pacific as expressed by
the myctophid fishes. In International Oceanographic Congress,
Preprints, 31 August - 12 September 1959, Mary Sears, ed.,
American Association for the Advancement of Science, Washing-
ton, D. C, pp. 142-143.
Brauer, a.
1903. Die Tiefsee-Fische. I. Systematischer Teil. Wiss. Ergeb. Deutsch.
Tiefsee-Exped., Valdivia, 1898-99, 15, 432 pp.
Ebeling, A. W.
1962. Systematica and zoogeography of the species in the bathypelagic
fish genus Melamphaes Glinther. Dana-Report No. 58, 1962,
Carlsberg Foundation, Copenhagen, 164 pp.
Ekman, S.
1953. Zoogeography of the sea. Sidgwick and Jackson, London, 417 pp.
Fraser-Brunner, a.
1949. A classification of the fishes of the family Myctophidae. Proc.
Zool. Soc. London, 118: 1019-1106.
424 bulletin: museum of comparative zoology
GooDE, G. B. and T. H. Bean
1895. Oceanic ichthyology. Special Bull. U. S. Nat. Mus., Washington,
XXXV + 553 pp.
HuBBs, Carl L.
1941. The relation of hydrological conditions to speciation in fishes.
Sj^iip. Hydrobiol., Univ. Wisconsin Press, pp. 182-196.
Kltjnzinger, C. B.
1870. Fische des Rothen Meeres. I. Theil. Percoiden-Mugiloiden.
Wien, 248 pp.
KULIKOVA, E. B.
1961. On the lantern fishes of the genus Diaphus (Scopelidae) from the
western Pacific. Trudy Inst. Okeanol. Moskva, 43: 5-39. (In
Russian.)
Marshall, N. B.
1963. Diversit}', distribution and speciation of deep-sea fishes. In
Speciation in the sea, J. P. Harding and Norman Tebble, eds.,
Symp. Syst. Ass., Pub. 5, London, pp. 181-195.
Mayr, E.
1963. Animal species and evolution. Harvard Univ. Press, Cambridge,
Massachusetts, 797 pp.
Norman, J. R.
1930. Oceanic fishes and flatfishes collected in 1925-1927. Discovery
Repts., Cambridge, 2: 261-370.
Parr, A. E.
1928. Deep-sea fishes of the order Iniomi from the waters around the
Bahama and Bermuda Islands. Bull. Bingham Oceanogr. Coll.,
New Haven, 3 (Myctophidae): 47-156.
1929. Notes on the species of myctophine fishes represented by type
specimens in the United States National Museum. Proc. U. S.
Nat. Mus., 76: 1-47.
Raymont, J. E. G.
1963. Plankton and productivity in the oceans. Pergamon Press,
Macmillan Co., New York, 660 pp.
SvERDRUP, H. U., M. W. Johnson and R. H. Fleming
1942. The oceans, their physics, chemistry and general biology. Prentice-
Hall, Inc., New Jersey, 1087 pp.
Taning, a. V.
1918. Mediterranean Scopelidae. Repts. Danish Oceanogr. Exped.
1908-10, Kj0benhavn, 2: 1-154.
1928. Synopsis of the scopelids in the North Atlantic. Vidensk. Medd.
Dansk. Naturh. Foren. Kbh., Kj0benhavn, 86: 49-69.
1932. Notes on scopelids from the Dana expeditions. Vidensk. Medd.
Dansk. Naturh. Foren. Kbh., Kj0benhavn, 94: 125-146.
Weber, M.
1913. Die Fische der Siboga-Expedition. Siboga Repts., Leyden,
57 (32): XII + 710.
Weber, M. and L. F. Beaufort
1913. The fishes of the Indo-Australian archipelago. Leyden, 2:
XX + 404.
(Received 5 April 1965.)
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, Xo. 10
THE AMEIVA (LACERTILIA, TEIIDAE) OP
HISPANIOLA.
IT. GEOGRAPHIC VARIATION IN AMEIVA
CHRYS0LAE3IA COPE
By Albert Schwartz and
Ronald F. Klinikowski
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
March 16, 1900
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Phylogeny and Evolution of Crustacea. Proceedings of a confer-
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Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 10
THE AMEIVA (LACERTILIA, TEIIDAE) OF
HISPANIOLA.
II. GEOGRAPHIC VARIATION IN AMEIVA
CHRYSOLAEMA COPE
By Albert Schwartz and
Ronald F. Klinikowski
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
March, 1966
Bull. Mus. Comp. Zool., Harvard Univ., 133(10): 425-487, March, 1966.
No. 10 — The Ameiva {Lacertilia, Teiidae) of Hispaniola
II. Geographic Variation in Ameiva Chrysolaema Cope
By Albert Schwartz
10,000 SW 84th Street, Miami, Florida 33143
AND
Ronald F. Klinikowski
127 Spring St., Reading, Pennsylvania
INTRODUCTION
The largest of the three species of Hispaniolan ground lizards,
Ameiva chrysolaema Cope, was described in 1868 with the type
locality ''Gonave Island." Three other names were then ap-
plied to this species in rapid succession: vittipunctata Cope
1871, affinis Fischer 1883, and regularis Fischer 1888. Of these,
vittipunctata, as Cochran (1941: 275-276) has made clear, was
apparently described by Cope from a young specimen (snout-
vent length 88 mm) which was part of the same series from
which he himself had taken the type of chrysolaema. Cope, how-
ever, gave a different type locality — ' ' city of Santo Domingo ' '
— for vittipunctataA
In her revision of the species in ' ' Herpetology of Hispaniola, ' '
Cochran (1941: 275-292) considered vittipunctata Cope a strict
synonym of A. chrysolaema Cope. A. regularis Fischer was re-
garded as a strict synonym of affinis Fischer, which was accepted
as a valid mainland subspecies (a new status since affinis had
previously been regarded either as a full species or a synonym).
Prior to 1941 two subspecies of A. chrysolaema had been de-
scribed from satellite islands of Hispaniola : A. c. woodi Coch-
ran from He de la Tortue and A. c. ahhotti Noble from Isla Be-
ata. These Cochran considered recognizable. A third subspecies,
A. c. hoekeri Mertens (1938: 338), however, from the mainland
at Fondo Negro, Republica Dominicana, was rejected as a syno-
nym of A. taeniura.
1 Cochran (1941 : 245) noted that the type locality of Cele^tus f= Diploglos-
SU8) weinlandi, also described by Cope and said by him to have been collected
on He de la (ionave, was incorrect and the s|ipcimen actnallv came from the
mainland within 25 miles of Port-au-Prince, Haiti. Since A. C. Younglove who
collected the type of C. tveinlanrti in 1868 also collecred the type of A. rhrysn-
laema, it is appropriate, as Dr. Cochran has done, to restrict the type locality
of A. chryftolaema to "within 2.t miles of Pnrt-aii-I'rince." If the type of A.
vittipunctata is actually part of the same series as the type of A. chrysolaema,
the type locality of the former should likewise be considered the same as that of
the latter, despite Cope's statement that it came from the city of Santo Domingo.
428 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
The discussion below departs radically from Cochran's revi-
sion in rejectinpr affinis while recoonizing" many other mainland
subspecies. In large part this has been due to the much greater
amount of material available to us.
Six hundred and fifty-five carefully documented and noted
specimens of A. chrysolaema have been collected by ourselves
and associates betvreen June 1962 and September 1964. Large
fresh series of this lizard are now availal)le to us from southern
and central Haiti, from He de la Tortue and Gonave, and from
the whole of its range in the Republica Dominicana. These speci-
mens are in the Albert Schwartz Field Series (ASFS) and the
collection of Richard Thomas (RT). They have been amassed
through the efforts of ]\Iiss Patricia A. Heinlein, and Messrs.
David C. Leber and Richard Thomas. All deserve our commen-
dations and thanks, most especially Messrs. Leber and Thomas
who made special efforts on our liehalf to secure topotypes of
A. c. ahboffi on Lsla Beata. Li addition, we have examined 417
specimens in the following collections : American ^Museum of
Natural History (AMXH), Carnegie ]Museum (C^I), ]\Iuseum
of Comparative Zoology (MCZ), Museum of Zoology, University
of Michigan (U]\OIZ), and United States National Museum
(USNM). To the curators and their assistants — Charles M.
Bogert and Miss Grace M. Tilger, Neil D. Richmond, Ernest E.
Williams, Charles F. Walker and George R. Zug, Doris M. Coch-
ran and James A. Peters — we wish to express our appreciation
for allowing us to examine pertinent specimens under their care.
Paratypes of new forms have been deposited in the Museum of
Natural History, University of Kansas (KU), and the University
of Illinois Museum of Natural History (UIMNH), as well as in
the above collections. The Harvard collections once again have
been indispensable ; through the efforts of Dr. "Williams, large
and well preserved series from northern Haiti have been made
available to us; without these our interpretation of the north-
western Haitian situation would have been not merely difficult,
but rather impossible. In the matter of literature, Edmond V.
Malnate has been most helpful and we are grateful for his co-
operation. The illustrations are the work of the junior author.
SYSTEMATIC ACCOUNT
The Species as a Whole
Amciva clirysolacma may be defined as follows: 1) a large
species of the genus Amcii'a with snout-vent length to 160 mm in
males and 134 in females; 2) dorsal caudal scales keeled and
SCHWARTZ AND KLINIKOWSKI : AMEIVA 429
straig-lit; 3) ventrals in 10, 11, or 12 transverse rows and in 33
to 41 long-itndinal rows; 4) fourth toe subdipital seales from 66
to 101; 5) femoral pores 24 to 52; 6) fifteenth eandal verticil
with 30 to 52 scales; 7) dorsal pattern consistino- of one of the
following: a) a series of dorsal yellow to huffy longitudinal
lines on a brown, tan, grayish tan, greenish, or lilackish ground
color, the lines (straight or wavy) at times modified into dashes,
dots, or fused with one another to give ultimately a median dor-
sal longitudinal band, b) a rather uniform covering of brightly
colored (blue, orange, yellow) spots on a dark background,
c) a pale ground with dark vermiculations and tigroid vertical
lateral bars, or d) completely or almost unicolor dorsally without
any striking jiattern elements; and 8) hemipenis extending to
about the seventh to ninth caudal verticil, sulcate surface naked,
sulcus bifurcating apically, the branches ending in two poorly
defined scalloped apical discs, non-sulcate surface entirely
flounced, the flounces extending to the margins of the sulcate sur-
face, a small smooth triangular area on the non-sulcat(> side which
divides the flounces for about one-third the length of the organ
into two fields of flounces which correspond to the apical discs.
The center of the distriliution of A. chrysolacma is in the (Jul
de Sac-Valle de Xeiba region of llispaniola; the species is rep-
resented by large series and from numerous localities in this gen-
eral region. From here, A. cJirysoIacDia extends westward to the
vicinity of Leogane on the Tiburon Peninsula, and eastward as
far as San Pedro de Macoris in the Pepublica Dominicana. From
Leogane to San Pedro de ]\lacoris, there is a set of localities
which implies a more or less continuous distribution. From this
basic center, populations extend northAvestwarcl along the shore
of the Golfe de la Gonave into the valley of the Riviere de I'Arti-
lionite, and in the central portion of Haiti at least as far north
as the vicinity of Mirebalais. In the Kepublica Dominicana, there
is practical continuity of the main southern mass of the species
north of the Sierra de Xeiba in the Valle de San Juan and
thence to the Dominico-Haitian 1, order near Pedro Santana and
immediately across the border at Cerca-la-Source in Haiti. A
second major center lies in the northwestern portion of Haiti and
extends thence eastward into the Valle de Cibao in the Republica
Dominicana. Whether these two major populations are com-
pletely isolated from one another is unknoAvn ; there is at least
no contact in the Republica Dominicana, since the Cordillera
Central stands between the two main regions, and the central
valley along the eastern slopes of this range is unoccupied by
chrysolaema.
430 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
There are several apparently disjunct populations in southern
Haiti and the Republiea Dominicaua ; in the former country
there is a single specimen from Aquin to the west along the
Tiburon Peninsula, far removed from the nearest records from
Leogane. In the Republiea Dominicana there are specimens from
Juanillo near the eastern extremity of the island. Although there
are no records from between San Pedro de Macoris and Juanillo,
the species has presumably occurred in that region fairly re-
cently (and may still occur as isolated populations), since it oc-
curs on Isla Catalina and Isla Saona. It is at present unknown
from the adjacent mainland in each case. There were two speci-
mens reported (Cochran, 1941:282) from Ile-a-Vache but these
cannot now be located ; although in two extended visits to this
island we secured only the very abundant A. tacniura Cope, it is
not altogether impossible that A. chrysolaema occurs there as
well. Finally, there are populations south of the Sierra de Bao-
ruco-Massif de la Selle which are completely cut off by these
ranges from their more northern relatives ; this same phenomenon
has now been noted in several other species of reptiles from this
area. These mountains, as well as the virtually non-existent
coastal plain along the eastern shore of the Peninsula de Bara-
hona, form an inescapable trap for several reptiles in the south-
ern portion of the Peninsula.
For the remaining satellite islands, A. chrysolaema is repre-
sented by abundant material from He de la Gonave and occurs
also on He de la Tortue, Isla Beata, and the Siete Hermanos
islands off the northwestern portion of the Republiea Domini-
cana. Of the three species of Hispaniolan Anieiva, none has so
broad a range as does A. chrysolaema. Nonetheless, there is geo-
graphic evidence that even this species is retracting its range ;
the apparent absence of chrysolaema from much of the south-
eastern portion of the Republiea Dominicana, but its occurrence
on off-shore islands in this region, suggests a formerly more
widespread distribution in this area.
A. chrysolaema is absent from the distal third of the Tiburon
Peninsula in Haiti, and apparently from much of the central i^or-
tion of that country as well. In the Republiea Dominicana it does
not occur in the Cordillera Central nor the Sierra de Neiba, and
is absent from the central and eastern portions of the country-
except along the southeastern coast. Interestingly, despite its
occurrence in the Valle de Cibao, it does not occur along the
northcentral coast of the Republiea Dominicana. The Cordillera
SCHWARTZ AND KLINIKOWSKI : AMEIVA 431
Septentrional acts as an effective barrier in this region. Much
of the eastern Republica Dominicana is mesic (in fact, the area
of highest rainfall in the country occurs in this region), and
since A. chrysolaema is distinctly a lizard of xeric habitats, this
one factor may well have prevented its expansion into this region
and onto the Peninsula de Samana. However, its absence along
the coast from Cabo Engaiio westward is strange, since this coast
is arid and appears suitable for these lizards. Considering the
disjunct nature of the populations of the species in extreme east-
ern Hispaniola, it is possible that it never occurred in this region
or that it has already retreated from this suitable coastal area.
A. chrysolaema lives in xeric regions. It is abundant in the
Cul-de-Sac plain below sea level and on the Peninsula de Bara-
hona. It occurs also in the dry Valle de San Juan, at elevations
of about 1000 feet. Although more tolerant of less xeric situa-
tions than A. lineolaia, the two often occur together, with A.
Uncolata inhibiting more open, cactus-studded, sandy regions,
and A. chrysolaema preferring slightly more shady areas, such
as adjacent copses or thickets of Acacia. If lowland woods are
present, A. chrysolaema may invade them; the woods may not be
dense nor with abundant ground cover. Maritime deciduous for-
ests along the mangrove border (but usually not the mangroves
themselves) offer a suitable habitat. Scrub-lands and open
beaches with some cover are often adetiuate. In its relationships
with the other two species, A. chrysolaema most often occurs with
A. lineolata as noted above. On occasion, however, A. chrysola-
ema occurs with the shade-loving A. taeniura. In such instances,
chrysolaema appears to be tlie secondary invader of a habitat
which is the preferred habitat of taeniura; in one such case near
Oviedo on the Peninsula de Barahona, taeniura kept strictly to
the open dry forest, whereas chrysolaema occurred almost exclu-
sively along the edges of the woods where they abutted on a dry
mangrove flat. The latter is likely the more preferred habitat
of chrysolaema, but during the heat of the day this species was
not averse to foraging in marginal forested situations.
Characters studied
We have examined a total of 1072 specimens of A. chryso-
laema (in contrast to 42 examined by Barbour and Noble, 1915,
and 198 by Cochran, 1941) ; of these, 655 are specimens collected
by ourselves and parties at various times, and upon which we
have extensive data on coloration and pattern. Of the races
432 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
recognized and described in the present paper, we have seen liv-
ing or freshly killed specimens of all but two (the race from
northwestern Haiti, and the race from extreme eastern Repvib-
lica Dominicana). We feel that data on coloration and pattern
are absolutely indispensable for any modern worker on the genus
Ameiva. Old, faded, or (even worse) badly discolored specimens
are completely useless except for scale counts and measure-
ments, and analyses of populations must rest heavily and se-
curely upon data from living or freshly killed animals.
We have taken counts of rows of longitudinal and transverse
ventrals, fourth toe subdigital scales, femoral pores, and scales
in the fifteenth tail verticil (see Tables 1-4). Of these, only the
numlier of transverse rows of ventrals can be used (partially) to
characterize subspecies — i.e., having either 10 or 12 transverse
rows of ventrals. Xo ]K)pulation has all specimens with 10 or all
with 12 rows. However, there is most often a preponderance of
one or the othei- in any particular sample, and we have used this
modal number as typical of the race in question (Table 2), un-
less the sample is rather small or the two categories differ by only
a very few individuals.
Of least value systematically is the number of longitudinal
rows of ventrals. In the entire lot of A. chrysolacma examined,
this figure varies from 33 to 41. The means for the 15 popula-
tions described herein vary from 38.3 to 36.5. Data for longitu-
dinal rows are presented in each case, but these data are not em-
phasized.
The counts of fourth toe scales, femoral pores, and fifteenth
verticil scales show some trends, although in almost all cases the
amount of overlap is rather large between most populations. Dif-
ferences of means, however, between those races which rank first
and last in each category may be rather striking (see Tables 2.
3 and 4). The largest difference between the highest and lowest
populations is in fourth toe scales, where the high population has
a mean of 91.0, and the low 77.8 — a difference of 13.2 scales.
For facility Ave have combined in all cases the fourth toe scales
from both feet and the femoral pore counts for both legs into
one figure for each specimen ; we do not feel that this weakens
the use of the data and it may well intensify slight differences,
which would otherwise be almost unnoticeable. We have given
the means and extremes for these three scale counts for each
subspecies ; the differences, if any, obviously are mean differ-
ences, since overlap of ranges is great in most cases.
schwartz and klinikowski : ameiva 433
The Recognizable Subspecies
Ameiva chrysolaema chrysolaema Cope, 1868
Ameiva chrysolaema Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 20:127
(type locality — "He de la Gonave" = within 25 miles of Port-au-
Prince, fide Cochran, 1941:275).
Ameiva vittipunctata Cope, 1871, Proc. Acad. Nat. Sci. Philadelphia,
22:220 (type locality — "city of Santo Domingo" = within 25 miles
of Port-au-Prince; see Cochran, 1941:275-76 for discussion of rationale
for this restriction).
Ameiva affinis Fischer, 1883, Beschreibungen neuer Eeptilien, [Separat-
Abdruk aus dem] Osterprogramm des akademischen Gymnasiums, Ham-
burg, p. 1 (type locality — "Haiti").
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of very large size (males to 160 mm, females to 130
mm snout-vent length), usually 12 transverse rows of ventrals,
moderate number of fourth toe subdigital scales, high number of
femoral pores and of scales in the fifteenth tail verticil ; dorsal
pattern consisting of about six longitudinal yellow lines and/or
yellow dots arranged in series (Fig. 1, left), and a black gular
band which may involve the chest and undersides of the arms.
Distribution: From St. Marc (and including the "Artibonite
Valley") on the north, southeast along the shore of the Golfe de
la Gonave, east throughout the Cul-de-Sac plain to the environs
of Etang Saumatre (Manneville, Ganthier, Fond Parisien), and
west on the Tiburon Peninsula as far as the vicinity of Leogane
(Pere) ; an isolated specimen from Aquin, Dept. du Sud, Haiti
(Fig. 11).
Discussion : A. c. chrysolaema is distinctly the largest and most
bulky of the races of the species. Males reach a snout-vent length
of 160 mm and females 130 mm. Color notes on a series from
Eaux Gaillees in the Haitian Cul de Sac show the situation as far
as coloration and pattern are concerned. Males from Eaux Gail-
lees were noted as dark brown to reddish brown dorsally, espe-
cially reddish on the shoulders and head (which may also be
grayish). Lores and cheeks with gray blotches. The back has
either a series of six yellow lines and yellow dots in the inter-
spaces between the lines, or has six rows of lemon yellow spots.
The lateral fields are black with or without a longitudinal series
of yellow spots. The lower sides have large yellow spots as
well, and the lateralmost belly plates are blue-spotted. The
ventral ground color is dull blue-gray, the throat pale orange
(Maerz and Paul, 1950, PL 9D7). There is a black gular band,
434
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 1. Left, Ameiva chrysolaema chrysolaema, ASFS X2162, 3.9 mi.
NW Ganthier, Dept. de 1 'Quest, Haiti. Right, Ameiva c. umiratilis, holo-
type, MCZ 77231, Baraliona, Barahona Prov., Eepublica Dominicana.
which may expand posteriorly to cover the chest and under-
sides of the arms. Some adult males have the shoulders and
neck blackened, so that in these regions the yellow lines are
much dulled and obscured. The females are like the males except
that the lines are yellow and prominent anteriorly. The venter
is dull blue-gray. The black gular collar is present but less pro-
nounced than that of the males, and the throat is orange but
paler than that of males.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 435
A series from Diquiiii to the ^vest of Port-au-Prince was col-
ored much as the Eaiix Gaillees material. The dorsal ground
color in males was brown with yellow dots arranged in lines or
with yellow lines additionally present. There is a black lateral
field with yellow spots. The sides of the head and axillae had
vivid and prominent blue to blue-green blotches. The heads were
dull reddish brown to dull orange, with orange-pink throats. The
ventral ground color was grayish to orange with blue spots on the
sides of the abdomen. The hindlimbs were dotted with yellow,
the forelimbs with blue-green. Females resemble the males, but
the dorsal lines or dots are less bright and prominent.
From the above descriptions it is obvious that, despite some
diiferences in details, these two populations (as well as many
others throughout the range of chrysolaema) share a communitj'
of dorsal markings — the longitudinal series of yellow^ lines
and/or dots. Cochran (1941: pi. 8E) showed a dorsal view of
the type of A. chrysolaema. The six dorsal lines, in this case
partially fragmented into series of longitudinal dashes, are
quite distinct. There is no doubt that the type of A. chrysolaema
did indeed originate in the vicinity of Port-au-Prince, since only
this subspecies occurs anywhere near that city. Specimens from
He de la Gonave are much duller, less prominently marked, lack
fragmentation of the dorsal lines, and are not referable to the
nominate form.
The longitudinal ventrals vary between 35 and 40 (mean
37.7) and these scales are most often arranged in 12 transverse
rows (67.9 per cent), with 29.2 per cent having 10 transverse
rows and 2.9 per cent having 11 rows. The variation in number
of transverse rows depends primarily on whether the lateralmost
of the enlarged ventral scales is sufficiently large to be con-
sidered a ventral; we have so considered it if its length (longi-
tudinally) is equal to that of the next inner adjacent row, and
if its width (transversely) is equal to at least half that of the
next inner adjacent row. Occasional specimens may also have one
or two rows of the belly plates divided, thus attaining counts
of 11 or 12 in another fashion.
The fourth toe subdigital scales range from 76 to 101 (mean
86.7) and the femoral pores range from 33 to 50 (mean 43.7).
The scales in the fifteenth caudal verticil vary between 37 and
51 (mean 44.4).
Within the range ascribed to A. c. chrysolaema, there are vari-
ous relatively minor pattern variants which we consider as
436 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
part of the normal variation of the subspecies. Specimens
from St. Marc at the northwesternmost extreme of the range are
very boldly lined longitudinally; the same is true of two indi-
viduals from Manneville. In both cases there are specimens from
adjacent localities or from the same locality which have more
typical chrysolaema markings.
The single individual from Aquin (USNM 72614) is unique
in several (possibly significant) features. The longitudinal lines
are composed of dots which also have a more or less transverse
arrangement, so that the back has a rather conspicuous trans-
versely banded appearance, a condition seen in no other A. c.
chrysolaema. The black lateral field is obscured, and there is no
black gular band, the entire throat, chest and undersides of the
arms being unmarked. The specimen is an adult male with a
snout-vent length of 146 mm, ventrals in 38 longitudinal and 12
transverse rows, 95 subdigital fourth toe scales, 41 femoral pores,
and 43 scales in the fifteenth caudal verticil. None of these counts
will distinguish the specimen from A. c. chrysolaema, although
we attach no particular significance to this fact. Considering
the wide gap between the known localities of A. chrysolaema
between Pere near Leogane and Aquin (a distance of some 90
kilometers) and the fact that the Aquin specimen comes from
the southern, in contrast to the northern, side of the Tiburon
Peninsula, it is likely that this single individual comes from a
population which is distinct from A. c. chrysolaema. AVithout
additional material, and especially lacking careful data on col-
oration and pattern in life, we are unwilling to name this
single Aquin specimen as distinct from A. c. chrysolaema.
The character ascribed by Cochran (1941:277) to differenti-
ate A. c. affinis — i.e., the interparietal being larger than the
adjacent scales — we find to be completely untenable. Dr. Coch-
ran has also shown (1941:291) that the scale counts of affinis
fall within the known range of A. c. chrysolaema. Specimens
which she assigned to affinis were reported (1941:292) from
Momance, Manneville, and Pere in Haiti. Other specimens (p.
282) from Manneville were assigned to c. chrysolaema. In a
series of twenty-seven specimens from Fond Parisien and the
eastern Cul de Sac, for example, six have the interparietal larger
than adjacent scales and the balance have the interparietal
smaller. The same situation applies to series from other locali-
ties within the range of A. c. chrysolaema, and if we accept
affinis as differentiated by this character alone, then the races
SCHWARTZ AND KLINIKOWSKI : AMEIVA 437
chrysolaema and affinis are broadly and randomly sympatric.
No features of pattern or coloration will distinguish specimens
with smaller interparietals from those with larger interparietals,
and we cannot detect any other constant scale feature which will
distinguish two forms in this region. For this reason we con-
sider affinis Fischer as a synonym of chrysolaema Cope. The
reasons for considering vittipunctata Cope as a synonym of
chrysolaema have been outlined in the introduction.
Specimens examined: Haiti, Dept. de I'Artihonite, "Artibonite
Valley" (not mapped), 1 (USNM 75921) ; St. Marc, 5 (USNM
59079, MCZ 58012-13, 65351, AMNH 49766) ; Bept. de VOuest,
2.2 mi. (3.5 km) SW Trou Forban, 1 (ASFS X1927) ; between
Arcahaie and Trou Forban, 1 (MCZ 51433) ; 6.3 mi. (10.1 km)
NE Arcahaie, 3 (ASFS X1928, X1930, X1946) ; 13 mi. (20.8
km) SW Arcahaie, 1 (ASFS X1938) ; Port-au-Prince, 12
(AMNH 49637-38, MCZ 13839, 59495-502, 69420) ; Carrefour
Feuille, Port-au-Prince (not mapped), 1 (MCZ 65810) ; Delmas,
2 (MCZ 65808-09) ; Petionville, 1 (USNM 59078) ; 10 mi. (16 km)
SW Port-au-Prince, 1 (UMMZ 92197) ; 3.5 mi. (5.6 km) E Croix
des Bouquets, 15 (ASFS X2197-211) ; Eaux Gaillees, 33 (ASFS
X1651-83) ; Manneville, 11 (MCZ 8621-23, 8625, 8629-33, 8614,
8618) ; 3.9 mi. NW Ganthier, 18 (ASFS X2153-70) ; 1.3 mi. (2.1
km) NW Fond Parisien, 3 (ASFS X2174-76) ; 0.4 mi. (0.6 km)
SE Fond Parisien, 7 (ASFS X2189-95) ; Hatte Latham (not
mapped), 1 (MCZ 51424); Diquini, 36 (ASFS X2381-407,
MCZ 6292, 8706, 8649-51, 8653-54, 8658-59) ; Momance, 8 (MCZ
8634-35, 8638-41, 8649, 20875) ; Qa Ira, 2 (MCZ 64919-20) ;
Pere, 3 (MCZ 13271-73) ; Dept. du Sud, Aquin, 1 (USNM
72614).
Ameiva chrysolaema umbratilis,^ new subspecies
Ilolotype: MCZ 77231, a subadult female, from Barahona,
Barahona Province, Republica Dominicana, taken 25 July 1963,
one of a series taken by native collectors. Original number
X9721.
Paratypes: All from the Republica Dominicana, as follows:
MCZ 81000-04, USNM 152558-60, KU 79861-64, UIMNH 56886-
89, RT 738-39, same data as holotype; ASFS X9568-69, Bara-
hona, Barahona Prov., 24 July 1963, native collector; AMNH
37943-49, 38133-39, Barahona, iBarahona Prov., 12 October 1922,
1 From the Latin for "remaining in the shade."
438 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
G. K. Noble; AMNH 63191, 63193, Barahona, Barahona Prov-
ince, 10-19 July 1932, W. G. Hassler; MCZ 58019, Barahona,
Barahona Prov., 13 July 1932, W. G. Hassler; MCZ 43813-14,
Barahona, Barahona Prov., 18 July 1932, W. G. Hassler ; ASFS
V199-200, 4 km NW, 1 km SW Barahona, Barahona Prov., 5
August 1963, A. Schwartz, R. Thomas; AMNH 49837-38, "Palo-
mino Spring's, nr. Barahona" (not mapped), Barahona Prov.,
15 August 1935, W. G. Hassler.
Associated specimens : Rcpuhlica Dominicana: Independencia
Prov., 6.5 mi. (10.5 km) NE Jimani, 1 (ASPS X9507) ;
4.4 mi. (7.0 km) SE Jimani, 3 (ASFS X9515-17) ; 13 km
SW La Descubierta, 3 (ASPS X9364-66) ; 5 km E La Descu-
bierta, 5 (ASFS X9354-58) ; Las Baitoas, 1 (MCZ 58776) ; 22
km SE Duverge, 7 (ASFS X9928-34) ; 1 km W El Naranjo, 1000
feet, 2 (ASFS X9943-44) ; northv^^est side, Laguna del Rincon, 1
(MCZ 58779) ; Guayabal, 9 km N Postrer Rio, 2 (MCZ 57732-
33) ; Baoruco Prov., Jaragua, 5 (ASFS X9469-72, RT 713) ;
0.7 mi. (1.2 km) E El Estero, 2 (ASFS X9467-68) ; 0.8 mi.
(1.3 km) SW Neiba, 15 (ASFS V252-64, RT 775-76).
Diagnosis: A subspecies of A. chrysolacma characterized by a
combination of moderate size (males to 130 mm, females to 112
mm snout-vent length), usually 10 (but often 12) transverse
rows of ventrals, moderate number of fourth toe subdigital
scales, low number of femoral pores, and high number of scales
in the fifteenth caudal verticil ; dorsal pattern consisting of dull
grayish brown to greenish black dorsal ground color with a series
of eight to ten dorsal longitudinal lines composed of small and
numerous dull yellowish to tan dots (Fig. 1, right), and a black
gular band which in adult males may involve the chest and
undersides of the arms.
Distribution : The Valle de Neiba from just east of Jimani to
the vicinity of tlie city of Barahona, Republica Dominicana
(Fig. 11).'
Description of type: A subadult female with the following
measurements and counts : snout-vent length 93 mm, tail 184
mm ; ventrals in 37 longitudinal and 10 transverse rows ; fourth
toe subdigital scales 39 and 41 (total 80) ; femoral pores 18
and 17 (total 35) ; 43 scales in the fifteenth caudal verticil.
Dorsal ground color grayish brown in life, head gray, shoulders
greenish ; ten rows of dull yellowish dorsal dots, the dots in each
series virtually confluent with one another, giving a vague wavy
line ; lateral fields obscure darker gray with scattered buffy dots.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 439
Throat pale purplish orange, followed by a black gular band
which extends slightl}' onto the chest and onto the underside of
the f orelimbs ; ventral ground color grayish blue ; tail dull grayish
brown above and grayish blue below, with some scattered darker
scales dorsally.
Variation: See tables. The characters of umhratilis are best
expressed in populations from the eastern section of the Valle de
Neiba ; however, even the most western specimens from the vicin-
ity of Jimani are in no way comparable to Haitian A. c. chryso-
laenia. The dorsal ground color was noted in life as being green-
ish black (Jimani, La Descubierta, Jaragua), greenish brown
(Duverge, El Naranjo), brown (El Estero), and grayish brown
(Barahona). The dots in the dorsal longitudinal lines are tiny
and very often confluent, giving almost a vermiculate appearance
to the dorsal band; the dots vary in color from greenish
(Jimani), creamy (La Descubierta), pale yellow to pale green
(Duverge), yellow (El Estero), grayish yellow (Jaragua), or
dull yellowish to tan (Barahona).
The lateral fields are usually dull and inconspicuous, hardly
darker than the lateral coloration ; they often include a row
of yellow to creamy spots and are not outlined either above or
below by pale and prominent longitudinal lines. The ventral
coloration varies from pinkish gray and grayish orange to
grayish blue, with specimens having the brighter colors known
from the western extremity of the range. The throats are dull
pinkish gray and grayish orange to dull purplish or dull orange.
The l)lack gular band is invariably present and may, in adult
males, expand to cover much of the chest and anterior abdomen
and underside of the forelimbs. The upper surfaces of the
limbs are usually unspotted, but if there are a few scattered dots
these are blue on the forelimbs and yellow on the hindlimbs.
Comparisons: The coloration and pattern of chrysolaema and
xmhratilis are strikingly different; even in the western portion
of the range of umhratilis, no specimen approaches closely the
vivid dorsal coloration and pattern of the nominate race. The
extreme condition in the eastern Valle de Xeiba contrasts
strongly with the condition at Fond Parisien, for instance, and
specimens from Jimani and La Descubierta are much more like
individuals from Barahona in having fine dorsal dotting and
generally more drab colors than they are to specimens from
Fond Parisien. Umhratilis is a smaller lizard; no specimen of
cither sex of this race achieves the much bulkier and larger
440 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
size of chrysolaema. This is certainly not a sample artifact since
large series of both forms are at hand. The tendency for vmhra-
tilis to have 10 versus 12 transverse rows of ventrals, as in
chrysolaema, is of interest, although in umhratilis the specimens
are almost equally divided between the 10- and 12-row condi-
tions. In number of longitudinal ventral rows, the two races
are comparable {chrysolaema 37.7, umhratilis 37.1). In number
of femoral pores these two races differ strongly, with a mean of
43.7 in chrysolaema and 35.6 in umhratilis. Chrysolaema aver-
ages slightly higher in counts of femoral pores and fifteenth
verticil scales.
Remarks: The occurrence of two very distinct races of A.
chrysolaema in the Cul de Sac-Valle de Neiba complex is sur-
prising. Aside from the more mesic eastern and western ends of
this long xeric valley, the conditions throughout are quite com-
parably severe. Interestingly, the Valle de Neiba is greatly con-
stricted just to the east of Jimani ; it is possible that this narrow
neck (7-10 km) has been effective in separating these two races.
Specimens from the Republica Dominicana to the northwest of
Jimani may well be assignable to A. c. chrysolaema.
Ameiva chrysolaema boekeri Mertens, 1938
Amciva chrysolaema hoeTceri Mertens, 1938, Senckenbergiana, 20:338
(type locality — south of Foiido Negro, lower Rio Yaque del Sur, Bara-
hona Province, Republica Dominicana).
Diagnosis: A subspecies of A. chrysolaema characterized by
a combination of moderate size (males to 126 mm, females to
111 mm snout-vent length), usually 10 transverse rows of ven-
trals, moderate number of fourth toe subdigital scales, low num-
ber of femoral pores, and high number of scales in the fifteenth
verticil ; dorsal pattern of two phases : ( 1 ) back yellowish brown,
grayish tan, to olive, and without pattern and often without any
indication of lateral fields, or (2) dorsum colored as above
but with faint paler marblings or longitudinal lines and a fairly
prominent black to dark gray lateral field (Fig. 2), and a
black gular band which may involve the chest and underside
of the arms.
Distribution: North of the Rio Yaque del Sur in extreme
eastern Valle de Neiba, north and east to north of Azua and
east to the vicinity of Bani, in the Llanos de Azua, Republica
Dominicana ; intergrades with the next subspecies to the north-
west in the vicinity of Hato Nuevo, Azua Province (Fig. 11).
SCHWARTZ AND KLINIKOWSKI : AMEIVA
441
Fig. 2. Left, Ameiva c. hoekeri, ASFS X7811, 10 mi. NW Bani, Peravia
Prov., Eepiibliea Dominicana. Eight, Ameiva c. boeTceri, ASFS V689, 15.2
mi. S San Jose de Ocoa, Peravia Prov., Eepubliea Dominicana.
Discussion: A. c. hoekeri was described on the basis of four-
teen lizards from Fondo Negro. Of these, seven males were
dorsall}^ patternless (including the type), four males showed
a " chriisolaema"-\ike pattern, and the final male was considered
by Mertens (1939:72) to resemble the Beata race ahhotti — i.e.,
it was dorsally dotted. The two paratypic females were " chryso-
toema "-like as well. We have examined a single paratype of
hoekeri (MCZ 44757) and eighty-six other specimens from the
range ascribed by us to hoekeri above. Of ten localities, only
442 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
two have "pure" or almost "pure" hoekeri populations (i.e.,
patternless), viz., a series of five specimens from the west side
of Punta Martin Garcia, Barahoua Province, and a series of
twenty-one from 10 mi. XW Bani, Peravia Province. Additional
specimens (four) from the eastern side of Punta Martin Garcia
show the typical duality of dorsal pattern, however, and thus
the uniformity of this small series of five is not significant. The
large series from Bani, on the other hand, has only a single
adult male which shows any pattern; this population is thus
almost completely patternless. A fresh series of five topotypes
from Fondo Negro has four individuals with patterns, and one
without pattern. Thus, although the patternless condition pre-
dominates at some localities (Bani), elsewhere (and including
the type locality) both types of dorsal pattern occur.
The two types of dorsal patterns, as delimited by jMertens,
grade into one another. The back may be longitudinally lined with
from six to ten tan to yellowish lines of fine dots, or these may
be very obscure centrally and more prominent at the sides of
the dorsal field. Some individuals have the back finely marbled.
The lateral fields are well developed and enclose a series of buffy
to cream dots; the lateral fields are often outlined below by a
longitudinal yellowish line, and a similar line may border these
fields above. The lower sides may be dotted with blue or bluish
green. The dotted condition of the back, considered by Mertens
as being ahhotti-like, is not at all comparable to the large and
brilliant sky-blue spotting on a black ground of that race. There
is some similarity between patterned hoekeri and nmhratUis. No
nmhratilis however is unpatterned.
In patternless lizards, the dorsal ground color was recorded
as yellowish brown (Fondo Negro), brown (San Jose de Ocoa),
grayish tan (Punta Martin Garcia), black (Barreras), and
reddish brown (Bani). The lateral fields may be completely
absent or may be indicated by a somewhat grayer longitudinal
lateral stripe, without any sort of included or adjacent pale
dots. The ventral ground color is blue-gray, purplish blue, blue,
light olive, gray, or orange-gray. The throat is likewise variable,
but is some shade of dull orange; females have throats which
are typically more grayish orange than males. The tails are
brown to grayish tan above, and gray below.
Scale counts for the series (including intergrades from Hato
Nuevo) are: longitudinal ventrals 34-40 (mean 37.2), trans-
verse ventrals in 10 (82.4 per cent) or 12 (17.6 per cent) rows,
SCHWARTZ AND KLINIKOWSKI : AMEIVA 443
fourth toe scales 73 to 98 (mean 84.6), femoral pores 31-41
(36.5), fifteenth verticil 37-48 (mean 42.7).
The much smaller size and faded pattern of those hoekeri
which have patterns, as well as the patternless individuals, can
easily be distinguished from A. c. chrysolaema. Chrysolaema is
likewise characterized by 12 rather than 10 rows of ventrals.
In counts of fourth toe scales, femoral pores, and fifteenth verti-
cil scales, hoekeri averages less than chrysolaema, the most
striking difference being in femoral pores {chrysolaema 43.7,
hoekeri 36.5). Patternless hoekeri can be easily differentiated
from all umhratilis, since this race is never patternless. Pat-
terned hoekeri are much like umhratilis. In both, the dorsal pat-
tern is faded and not well demonstrated. One feature is sugges-
tive ; patterned hoekeri have the lateral fields prominent and
often outlined both above and below, whereas the typical umhra-
tilis condition is an obscure lateral field, not set off by longi-
tudinal pale lines. Both umhratilis and hoekeri usually have 10
rows of ventrals, although umhratilis has a much higher fre-
quency of 12-row individuals. In counts of fourth toe scales
and femoral pores, hoekeri averages slightly higher than um-
hratilis; the means for fifteenth verticil scales are identical.
Because of the similarities of umhratilis and patterned hoekeri,
we have considered the possibility that the name hoekeri should
be applied to Ameiva from the Yalle de Neiba. To be counted
against this conception is the fact that of 61 umhratilis, none
is unpatterned, whereas 56.3 per cent of the specimens (hoekeri)
from north of the Kio Yaque del Sur are patternless. There is
no indication of this patternless condition in specimens from
Barahona, nor from elsewhere in the range of umhratilis. We
prefer to regard hoekeri as a separate entity, distinct from um-
hratilis to the south.
A. c. hoekeri is approached geographically by three adjacent
races; of these it is known to intergrade only with the race to
the northwest in the Valle de San Juan (these intergrades will
be discussed later). From umhratilis the range of hoekeri is sep-
arated by the lower reaches of the Rio Yaque del Sur and by
the extremely mesic conditions of much of the eastern portion
of the Valle de Neiba. Although hoekeri is not presently known
to intergrade with the race next to the east along the southern
coast of the Republiea Dominicana, it may well do so. It is cer-
tainly significant that hoekeri occupies the western Llanos de
444 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Azua in the vicinity of Ban! ; just to the east of that city, con-
ditions become more mesic, and the region is occupied by another
race. Intergradation likely takes place where these two regions
come into contact.
Specimens examined: Repuhlica Dominicana, Barahona Prov.,
Fondo Negro, 6 (ASFS X9703-07, MCZ 44757); west side,
Punta Martin Garcia, 5 (ASFS V84-88) ; Azua Prov., 3 km E
Barreras, 2 (ASFS V3164-65) ; 2 km W Puerto Viejo, 2 (ASFS
V3183-84) ; 22 km NW Azua, 3 (ASFS V465-67) ; 1.8 mi. (2.9
km) W, 1.1 mi. (1.8 km) N Azua, 18 (ASFS X8002-18, X8103) ;
1.8 mi. (2.9 km) W, 2.7 mi. (4.3 km) N Azua, 10 (ASFS
X8019-28); Peravia Prov., 8.9 mi. (13.9 km) S San Jose de
Ocoa, 1300 feet (430 m), 1 (ASFS V714) ; 15.2 mi. (24.3 km)
S San Jose de Ocoa, 9 (ASFS V687-95) ; 10 mi. (16 km) NW
Bani, 23 (ASFS X7801-21, RT 613-14). Intergrades between
A. c. hoekeri and the race to the northwest were examined from :
RepuMica Dominicana, Azua Prov., Hato Nuevo, 10 (ASFS
X437-46).
Ameiva chrysolaema alacris,^ new subspecies
Holotype: MCZ 77232, an adult male, from 10 km SE San
Juan, San Juan Province, Republica Dominicana, one of a series
taken 9 August 1963 by Albert Schwartz and Richard Thomas.
Original number V283.
Paraiypes: All from the Republica Dominicana, as follows:
ASFS V284-97, same data as holotype ; RT 778-79, 10 km S San
Juan, San Juan Prov., 9 August 1963, R. Thomas ; MCZ 81005-06,
USNM 152561-64, AMNH 92842-43, KU 79865-66, UIMNH
56890-93, 2.5 km W, 4.4 km S San Juan, San Juan Prov., 9
August 1963, D. C. Leber, R. F. Klinikowski ; KU 79867-68,
AMNH 92844, 2.5 km W, 5.4 km S San Juan, 9 August 1963,
D. C. Leber, R. F. Klinikowski ; ASFS V389-90, 10 km E Valle-
juelo, San Juan Prov., 12 August 1963, R. Thomas; USNM
152565, 3 km E Las Matas, San Juan Prov., 9 August 1963, R.
Thomas.
Associated specimens: Haiti, Dept. du Nord, Cerca-la-Source,
1 (USNM 76780) ; Repuhlica Dominicana, San Rafael Prov., 3.8
mi. (6.1 km) SE Sabana Cruz, 1 (ASFS V329) ; Guayabal, 1
(MCZ 58672); Azua Prov., Tubano (= Padre las Casas), 3
(USNM 66729-31) ; 0.7 mi. (1.1 km) NW Villarpando, 9 (ASFS
V419-27).
1 Prom the Latin for "lively."
SCHWARTZ AND KLINIKOWSKI : AMEIVA
445
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of moderate size (males to 126 mm, females to 109
mm snout-vent length), usually 10 transverse rows of ventrals,
moderate number of fourth toe subdigital scales, very low num-
ber of femoral pores, and high number of scales in fifteenth
caudal verticil ; dorsal pattern consisting of five to seven bold
longitudinal dorsal lines (the lateralmost forming a strong upper
border to the prominent black lateral fields with their enclosed
bright yellow dots), the longitudinal lines never broken into
dots and lines as in c. chrysolaema and always conspicuous and
discrete (Fig. 3, left), and a black gular band which rarely in-
volves also the chest and undersides of the arms.
Fig. 3. Left, Ameiva c. alaeris, holotype, MCZ 77232, 10 km SE San
Juau, San Juan Piov., Republica Dominicana. Eight, Ameiva c. proeax,
holotype, MCZ 77233, Santo Domingo, 2.2 km SW Rio Ozama, Dist. Nac,
Eepublica Dominicana.
446 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Distribution: From east central Haiti (Cerca-la-Source) south-
eastward through the Valle de San Juan (Fig. 11) ; intergrading
with A. c. hoekeri at Hato Nuevo, Azua Province, and with A. c.
chrysolaema in the vicinity of Mirebalais, Dept. de I'Ouest,
Haiti (see discussion below).
Description of type: An adult male with the following meas-
urements and counts: snout-vent length 116 mm, tail 200 mm;
ventrals in 37 longitudinal and 12 transverse rows; fourth toe
subdigital scales 42 and 44 (total 86) ; femoral pores 16 and 16
(total 32) ; 46 scales in the fifteenth caudal verticil. Dorsal
ground color brown with seven longitudinal pale yellow lines,
the lateralmost bordering above the black lateral fields with their
isolated yelloAv dots; lateral fields bordered below by a slightly
duller longitudinal line which contains a series of bright yellow
dots; lower sides dotted with yellow. Throat gray, venter dull,
dirtj^ orange. A black gular band, not extending onto the chest
or undersides of the arms. Tail brown above, gray below, with
an indistinct proximal and lateral area of yellowish dots; top of
tail with some darker brown scales. Fore- and hindlimbs with
pale scattered small dots, bluish on forelimbs and yellowish on
hindlimbs.
Variation: See tables. A. c. alacris presents a constant as-
semblage of coloration and pattern elements throughout its
range. The dorsal ground color is alwaj^s brown, with from
five to seven pale or dull yellow longitudinal lines, these
lines always forming a conspicuous pattern. The lines are
entire and not fragTiiented or modified into series of longitud-
inal dots, although in some specimens the more central lines,
especially posteriorly, may be broken into dashes. In general,
however, the integrity of the lines (even when fragmented) is
maintained. The black lateral fields are bold, set off by pale
longitudinal lines above and below, and enclose a single series
of scattered yellow dots ; the lower line bordering the lateral
field may have superimposed upon it a series of bright yel-
low dots, thus rendering the black lateral field even more con-
spicuous. The lower sides are dotted with yellow. The throat
varies from gray to very pale orange, and the venter likewise
varies between these two extremes.
A. c. alacris intergrades to the southeast with A. c. hoekeri
and to the southwest with A. c. chrysolaema. A series of ten
specimens from Hato Nuevo, Azua Province (ASFS V437-46),
SCHWARTZ AND KLINIKOWSKI : AMEIVA 447
shows the intergradatiou with hoekeri. Of this series, five are un-
patterned hoekeri, and five represent the patterned phase of that
race. These five patterned lizards have the lateral fields darker
than most patterned hoekeri, and there is a distinct tendency to
have the dorsal lines more boldly (brighter yellow) displayed
anteriorly, although the posterior dorsal pattern is fainter and
very like "typical" patterned hoekeri. The distance from Hato
Nuevo {alacris X hoekeri intergrades) to Villarpando {alacris)
is only ten kilometers, yet the series from the latter locality is
typical of alacris in all waj^s and has no patternless individuals.
From the vicinity of Mirebalais, Dept. de I'Ouest, Haiti, we
have examined specimens from the following localities: 3.4 mi.
(5.4 km) NE Barrage de Peligre, 2 (ASFS X2217-18) ; 1.1 mi.
(1.8 km) S Mirebalais, 3 (ASFS X2237-39) ; Mirebalais, 1 (MCZ
68510) ; La Tombe, nr. Mirebalais, 8 (MCZ 68517-24) ; Boudou,
nr. Mirebalais, 2 (MCZ 69387-88) ; Duvie, nr. Mirebalais, 1 (MCZ
68478). Of these, the last three places named. La Tombe, Boudou,
and Duvie, cannot be located ; they have not been mapped. Taken
as a whole, this lot of lizards is intermediate between chrysolaema
and alacris, although they are closer to alacris than to chrys-
olaema. Three lizards (ASFS X2237, ASFS X2217, MCZ 68510)
show the disintegration of the dorsal lines into series of yellow
spots, a typical chrysolaema feature. Several male specimens
are larger than alacris, with snout -vent lengths of 133 to 145 mm
(five lizards) ; two females have snout-vent lengths of 109 mm
(the upper extreme of alacris females), and another has a snout-
vent length of 110. In life, our specimens from Barrage de
Peligre and Mirebalais had yellow lines and a greenish wash on
the neck — the latter a chrysolaema character. Finally, some
individuals have a discrete black gular band as in alacris, whereas
others have the band expanded onto the chest and arms as in
chrysolaema. We consider this entire lot of specimens inter-
gradient between alacris and chrysolaema.
Comparisons: A. c. alacris is easily distinguished from the
three previously described races on the basis of dorsal pattern ;
the discrete, bold, and undotted longitudinal lines of alacris
contrast with the patternless or weakly patterned races hoekeri
and umhratilis, and with the larger and dorsally dotted and lined
chrysolaema. Alacris is a race with ten transverse rows of
ventrals as are umhratilis and hoekeri, in contrast to the twelve-
rowed chrysolaema. In fourth toe scales, alacris (84.8) averages
close to hoekeri (84.6), slightly higher than umhratilis (83.0)
448 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
and lower than chrysolaema (86.7). In femoral pores, alacris
has the lowest average (33.8) of any race of A. chrysolaema;
of the described forms, it is approached by umhratiUs (35.6)
and hoekeri (36.5) and is far below chrysolaema (43.7). In
scales in the fifteenth caudal verticil, on the other hand, both
alacris and chrysolaema are high (44.0 and 44.4), with hoekeri
and unih rat ills (both 42.7) lower.
Remarks: A. c. alacris occupies the high and xeric Valle de
San Juan and associated upland foothills (i.e., Tubano). It is
likely that it is more widespread in east-central Haiti than the
present evidence of one specimen from Cerca-la-Source indicates.
Presumabl}^ the race follows down the valley of the Riviere de
I'Artibonite, and in the vicinity of Mirebalais has genetic con-
tact with the more southern chrysolaema. Such genetic con-
tinuity may come across the Montagues de Trou d'Eau from the
Cul de Sac (although there is no obvious means of penetration
of this mountain mass), or chrysolaema may reach Mirebalais
via the valley of the Artibonite from the St. Marc area. The
only evidence for the latter is the single specimen noted under
A. c. chrysolaema, from the "Artibonite Valley"; this individual
may have come from some undetermined locality which is inter-
mediate between St. Marc and Mirebalais. Further collecting in
these areas should easily reveal the precise place of contact
between these two races.
AmEIVA CHRYSOLAEMA PROCAX^ lieW SUbspCcics
Holotype: MCZ 77233, an adult male, from Santo Domingo,
2.2 km SW of the Rio Ozama, Distrito Nacional, Repiibliea
Dominicana, one of a series taken 14 June 1963 by Ronald F.
Klinikowski, David C. Leber, and Richard Thomas. Original
number X7714.
Paratypes: All from the Republiea Dominicana, as folloAVS:
ASFS X7711-13. X7724-26, MCZ 81007-10, USNM 152566-70,
KU 79869-71, UIMNH 56894-97, RT 605-08, same data as holo-
type ; ASFS X9254-56, Santo Domingo, old airport, Distrito
Nacional, 17 July 1963, D. C. Leber, R. Thomas; AMNH 92845-
49, 5.9 km W Santo Domingo, Distrito Nacional, 20 June 1964.
D. C. Leber, R. Thomas.
Associated specimens: Rcpuhlica Dominicana, San Cristobal
Prov., 8.4 mi. (13.6 km) NE Sabana Grande de Palenque, 2
1 From the Latin for "bold."
SCHWAETZ AND KLINIKOWSKI : AMEIVA 449
(ASFS X8167-68) ; 4.2 mi. (6.7 km) NE Sabana Grande de
Palemiue, 22 (ASFS X8149-66, RT 643-46).
Diagnosis: A subspecies of A. chrysolaenia characterized by a
combination of large size (males to 141 mm, females to 116 mm
snout-vent length), usually 10 (but often 12) transverse rows
of ventrals, moderate number of fourth toe subdigital scales, low
number of femoral pores, and high number of scales in the
fifteenth caudal verticil ; dorsal pattern a series of six or seven
longitudinal yellow lines in a reddish brown field, the lines usu-
ally wavy or broken into a series of longitudinal dashes (Fig. 3,
right), and a black gular band which may be so expanded as to
involve the entire ventral surface including the undersides of
the arms.
Distribution: The coastal regions of San Cristobal Province
and the Distrito Nacional, from the Rio Ozama on the east to the
vicinity of Sabana Grande de Palenque on the west (Fig. 11) ;
presumed to occur inland, since individuals were seen crossing
the road near the city of San Cristobal.
Description of type: An adult female with the following
measurements and counts : snout-vent length 114 mm, tail 275
mm ; ventrals in 38 longitudinal and 12 transverse rows ; fourth
toe subdigital scales 46 and 46 (total 92) ; femoral pores 18 and
20 (total 38) ; 44 scales in the fifteenth caudal verticil. Dorsal
ground color rich reddish brown in life, with a series of seven
dull longitudinal lines, the median line rather obscure, the lateral
lines broken into a series of wavy dashes, the lateralmost lines
bordering above the black lateral fields, which have a series of
tiny yellow dots inclosed within them. Sides of head gray with a
creamy subocular patch and some pale irregular blue blotches.
Throat fleshy gray. Ventral color dark blue-gray. Hindlimbs
heavily spotted with yellow, forelimbs faintly spotted with blue.
A dark gray gular band not involving the chest and underside of
the arms. Tail reddish brown above, dark gray below.
Variation: See tables. The series of A. c. procax is remark-
ably uniform in both coloration and pattern. The dorsal ground
color is always some shade of reddish brown, and there may be
a yellowish green wash over the shoulders. The longitudinal
lines are conspicuous, although the median line may be reduced
or faint. Only in young and subadult individuals are the lines
entire, and even in these specimens there is a strong tendency
toward wavy fragmentation. The lateral fields are black and
the yellow dots included therein are regularly very tiny; in
450 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
females these included dots are less well differentiated than in
the males. The gular band is present, and in large males may be
very extensive, including most of the belly. The venter is blue
in juveniles and females, and gray to solid black in males.
Comparisons: The reddish brown dorsal coloration and the
wavy dorsal lines separate procax from all other races. Some
specimens of chrysolacma were noted in life as reddish brown, but
this is not the usual dorsal coloration of the nominate race. In-
dividuals of chrysolaema with this coloration are readily differen-
tiated from procax on the basis of the very different dorsal pat-
terns of the two subspecies.
In size, procax is closest to chrysolaema, although procax is dis-
tinctly the smaller of the two. Chrysolacma is typically a subspe-
cies with 12 rows of ventrals, whereas procax, although it has in-
dividuals with 12 rows, has a modal condition of 10 rows. Procax
averages fewer (84.8) fourth-toe scales than chrysolaema (86.7),
the same as alacris, and more than hoekeri (84.6) and umhratilis
(83.0). In number of femoral pores, procax (36.1) averages far
lower than chrysolacma (43.7), and slightly lower than hoekeri
(36.5), but slightly higher than umhratilis (35.6) and alacris
(33.8) ; procax is one of the races with a low^ number of femoral
pores. In fifteenth verticil scales, procax averages less (42.8)
than chrysolaema (44.4) and alacris (44.0), and is about equal to
hoekeri and umhratilis (42.7).
Remarks: A. c. procax is presently not known to intergrade
either with hoekeri to the west or the race next to the east along
the southern Hispaniolan coast. The easternmost hoekeri locality
(where the population incidentally is almost completely made up
of non-patterned individuals) and the westernmost locality of
procax are separated by onh' 35 kilometers. As noted in the dis-
cussion of hoekeri, Bani lies about on the dividing line between
the xeric Llanos de Azua to the west and more mesic conditions
on the east. Procax inhabits these eastern more mesic regions,
whereas hoekeri is restricted to the xeric Llanos de Azua.
Ameiva CHRYSOLAEMA PARVORis,^ new subspecics
Holotype: MCZ 77234, an adult male, from 0.9 mi. (1.4 km)
E Boca Chica, Distrito Naeional, Republica Dominicana, one of a
series taken 23 x\ugust 1963 by Ronald F. Klinikowski, David C.
Leber, and Richard Thomas. Original number V649.
1 From the Latin parvum (small) and os, oris (mouth), a translation of Boca
Chica. the type locality.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 451
Paratypes: All from the Republica Dominicana, as follows:
MCZ 81011-14, USNM 152571-74, A:MNH 92850-55, KU 79872-74,
UIMXH 55638-39, RT 789-90, same data as liolotype; ASFS
V669-79, Boca Cliica, eastern edge, Distrito Nacional, 23 August
1963, R. F. Klinikowski, D. C. Leber, R. Thomas.
Associated specimens: Repuhlica Dominicana, San Pedro de
Macoris Pro v., 0.5 mi. S San Pedro de Macoris, at lighthouse, 11
(ASFS X8181-91) ; La RomanaProv., Isla Catalina, western end,
4 (ASFS V554-57).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of large size (males to 137 mm, females to 113 mm
snout-vent length), usually 12 transverse rows of ventrals, mod-
erate number of fourth-toe subdigital scales and femoral pores,
and high number of scales in the fifteenth tail verticil; dorsal
pattern of tan to blackish brown ground color with dull yellow
spots which may be either discrete or confluent, giving a reticu-
late appearance (Fig. 4, left), lateral fields present and black, or
broken to give a tigroid effect, and a black gular band which may
be expanded to involve the chest and the undersides of the arms.
Distribution: Coastal southeastern Republica Dominicana,
from Boca Chica on the east to San Pedro de Macoris on the west,
and including Isla Catalina ; range as here described apparently
discontinuous, and A. chrysolaema unknown from the mainland
opposite Isla Catalina (Fig. 11).
Description of type: x\n adult male with the following meas-
urements and counts : snout-vent length 129 mm, tail 282 mm ;
ventrals in 38 longitudinal and 10 transverse rows; fourth-toe
subdigital scales 41 and 42 (total 83) ; femoral pores 19 and 19
(total 38) ; 44 scales in the fifteenth tail verticil. Dorsal ground
color dull blackish brown, with the entire back from the neck to
the sacrum covered with dull yellow spots, not aligned into linear
series ; lateral fields black, not bordered above or below, and in-
vaded by the brown dorsolateral coloration, the entire sides spot-
ted with yellow dorsally and pale blue ventrally. Throat gray,
ventral ground color gray. A black gular band which extends
slightly onto the chest and also the underside of the arms. Fore-
limbs faintly spotted with bluish, hindlimbs spotted with dull
yellow. Tail grayish brown dorsally, gray ventrally.
Variation: See tables. The dorsal ground color of A. c. par-
voris varies from blackish brown (type locality) to tan and
brown (San Pedro de Macoris). The spotted condition of
the black is typical of most specimens, including small juveniles.
452
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 4. Left, Ameiva c. parvoris, holotype, MCZ 77234, 0.9 mi. E Boca
Chica, Dist. Xac, Eepublica Domiuicana. Eight, Ameiva c. jacta, holotype,
MCZ 75267, Juanillo, La Eouiana Prov., Eepublica Dominicana.
but others show a more lineate pattern somewhat like that
described for procax — the dorsal lines broken into dashes
giving a wavy appearance. The lateral fields maj^ be obscured
as in the type, or may be slightly more prominent, with much
encroachment of brown to give an irregular and indefinite edge ;
at the same time the black lateral field pigment may extend
up onto the sides of the back, thereby giving a distinctly tigroid
appearance to the sides and dorsolateral regions. In some indi-
viduals the dorsal spots are confluent, thereby increasing the
tigroid effect by transverse dorsal pale markings. The throat is
SCHWARTZ AND KLINIKOWSKI : AMEIVA 453
grayish to dull orange and the ventral ground color varies from
grayish blue to dull, deep orange.
The small series from Isla Catalina resembles the mainland
specimens in dorsal pattern and in extent of the gular band,
which in parvoris may involve the chest and undersides of the
arms. The dorsal dots are conspicuously confluent, the lateral
fields are obsolete, and the ventral coloration is pale bluish with
an orange tint. The most obvious difference in coloration is that
the dorsal surfaces of the hindlimbs are rusty — a feature found
in no mainland parvoris. The Catalina series is composed of one
adult male with a snout-vent length of 126 mm and thus a large
lizard, and three females (one of which is a small juvenile), the
largest of which has a snout-vent length of 112 mm, again a large
lizard. The transverse ventrals are 10 in three specimens and 12
in one. In all other features of scalation the Catalina lot falls
within the known range of mainland parvoris. Additional speci-
mens from Isla Catalina ma}- well reveal that it is inhabited by
still another distinctive race ; the strikingly rusty hindlimbs are
indicative of at least one major color difference between Isla
Catalina specimens and mainland parvoris.
Comparisons: No other race of A. chrysolaema has the back
with irregularly arranged and at times confluent spots, and
tigroid sides. This feature alone will distinguish parvoris from
the previous races. In size, parvoris is much smaller than chryso-
laema and slightly smaller than procax, but larger than the re-
maining subspecies. In fourth toe scales, parvoris averages less
(83.2) than other races except umhratilis, which is comparable
(83.0). Pari^om averages less (38.2) than chrysolaema (43.8) in
femoral pores, but exceeds the remaining races, all of which have
low femoral pore counts. In fifteenth verticil scales, parvoris has
the same mean as hoekeri and umhratilis (42.7), less than chry-
solaema and alacris (44.4 and 44.0), and almost the same as
procax (42.8).
Remarks: A. c. parvoris is not known to intergrade with
procax on the west nor with the following subspecies to the
east. The easternmost locality for procax is separated by 35
kilometers from the westernmost parvoris record. It is possible
that the Rio Ozama may divide these two races of A. chrysolaema.
The area occupied by parvoris does not differ in any obvious way
from that inhabited by procax.
454 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Ameiva chrysolaema jacta^ new subspecies
Holotype: MCZ 75267, an adult male, from Juanillo, La Ro-
mana Province, Republica Dominicana, taken 29 March 1963, by
Clayton E. Ray and Robert R. Allen.
Paratypes: MCZ 75268-69, same data as holotype.
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of large size (males to 134 mm snout-vent length;
females unknown), usually 12 transverse rows of ventrals, low
number of fourth toe subdigital scales, moderate number of femo-
ral pores, and very low number of scales in the fifteenth caudal
verticil; a dorsal pattern of dark brown to black tigroid mark-
ings on a grayish tan background (in preservation), the pattern
extending in a diluted fashion onto the neck, a bold, checker-
board-patterned tail (Fig. 4, right), and a black gular band
which extends onto the chest and underside of the arms.
Dist)ihidion: Known only from the type locality in extreme
eastern Republica Dominicana (Fig. 11).
Description of type: An adult male with the following meas-
urements and counts : snout-vent length 134 mm, tail 304 mm ;
ventrals in 38 longitudinal and 12 transverse rows; fourth-toe
subdigital scales 41 and 41 (total 82) ; femoral pores 21 and 22
(total 43) ; 35 scales in the fifteenth caudal verticil. Dorsum
(preserved) tannish gray with a dark brown, almost black pat-
tern consisting of bold vertical tigroid markings on the sides and
about five wide, dark, longitudinal lines on the back, the latter
much confused and joined by the lateral vertical markings, giv-
ing a rather complete and complex dark brown reticulum, which,
although present on the neck, is much paler gray. Lateral fields
completely absent, no dotting on sides or back whatsoever. Upper
surface of forelimbs with obscure grayish lines and blotches,
hindlimbs with a dark brown reticulum enclosing large spots
which are pale centrally and darker peripherally. Tail with a
bold checkerboard pattern of grayish tan, white, and dark brown,
this pattern becoming obsolescent and absent on the distal half of
the tail. Belly and throat presently dull grayish orange, with
some lateral ventral scales with very dark gray blotches, thereby
giving the belly somewhat of a faint checkerboard appearance
laterally. Black gular band obsolete but indicated, and anterior
ten rows of ventrals clouded with dark gray.
1 From the L.itin for "thrown," an nllusion to the far flung distribution of this
subspecies.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 455
Variation: See tables. In coloration and pattern the two
paratypes are much like the type and require little comment.
The major difference is that the paratypes have a bold, black,
gular band which involves the chest and the underside of the
arms. Neither lizard has any indication of lateral fields, and
the sides and back have the tigroid vertical bars and brown to
black reticulum, as well as the checkerboard tail, just as de-
scribed for the type.
Comparisons: A. c. jacta does not need detailed comparison of
pattern with any other described subspecies ; the boldly and viv-
idly marked back with its light ground color will distinguish
jacta from the remaining races. A. c. parvoris is closest to jacta
in pattern, but the differences are so striking that the similarity
between these two subspecies is not very great.
In size, jacta is smaller than chrysolaema, procax and parvoris,
and larger than the remaining forms. All other races have a
higher numl^er of fourth-toe scales and fifteenth verticil scales.
Jacta has a high mean of femoral pores, having less than chryso-
laema, and more than the balance of the subspecies.
Remarks: No intergradation is known between jacta and par-
voris to the southwest; the easternmost mainland locality for
parvoris is separated by 145 kilometers from that of jacta. We
have attempted to secure specimens of A. chrysolaema between
San Pedro de Macoris and Juanillo at several localities along
the coast (La Romana, Boca de Chavon, Boca de Yuma) as well as
inland in this eastern region, without success. Typical xeric chry-
solaema habitats here are occupied by A. taeniura. Considering
the likeness of jacta to the race from Isla Saona, described below,
it is probable that this boldly marked type of lizard was at one
time (and still is?) abundant locally in extreme eastern Hispan-
iola. Presently, the hiatus between jacta and parvoris and the
apparent absence of the species in this eastern region suggests
strongly that the populations are relict wdth a disjunct distribu-
tion.
AmEIVA CHRYSOLAEMA RICHARDTHOMASI ^ UCW SUbspecicS
Holotypc: :\ICZ 77235, an adult male, from the environs of
Mano Juan, Isla Saona, Repiiblica Dominicana, taken 19 July
1964 by Richard Thomas. Original number V3018.
1 Named for the collector.
456
BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
Paratypes: ASFS V3019-30, USNM 152575-76, AMNH 92856-
58, KU 79875-77, UIMNH 56898-99, RT 935, same data as holo-
type.
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of large size (males to 137 mm, females to 124 mm
snout-vent length), usually 10 (but often 12) transverse rows of
ventrals, high number of fourth-toe subdigital scales and femoral
pores, and moderate number of scales in the fifteenth caudal ver-
ticil; dorsal pattern of two phases: (1) back gray-green with
Fig. 5. Left, Amelia c. richardthomasi, holotype, MCZ 77235, environs
of Mano Juan, Isla Saona, Eepublica Dominicana. Eight, Ameiva c. ridtard-
thomasi, ASFS Y3019, environs of Mano Juan, Isla Saona, Eepublica
Dominicana.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 457
only an indistinct mottling of gray-brown in the area of the lat-
eral fields or (2) back gray-green with heavy black mottling, no
lateral fields, and tigroid vertical bars on the sides, the dorsal
mottling not extending onto the shoulders and neck (Pig. 5),
and without a black gular band.
Di.strihufion: Known only from the type locality, but pre-
sumed to occur throughout Isla Saona (Fig. 11).
Description of type: An adult male with the following meas-
urements and counts : snout-vent length 137 mm, tail 146 mm,
twice broken and regenerated ; ventrals in 37 longitudinal and 12
transverse rows ; fourth-toe subdigital scales 45 on left side ;
femoral pores 22 and 20 (total 42) ; 38 scales in the fifteenth tail
verticil. Dorsal ground color gray-green (Maerz and Paul: pi.
22F1), becoming finely mottled laterally with a series of \Qvy
faint grayish brown vertical bars in the region of the lateral
fields ; lower sides putty colored. Throat and venter orange with
no black gular band. Fore- and hindlimbs unicolor with back,
and patternless. Tail grayish tan without obvious checkerboard
pattern above, putty colored below.
Variation: See tables. Sixteen specimens of richardthomasi
(including the type) are patternless dorsally and have a finely
filigreed, grayish brown area in the region of the lateral fields.
The tails are gray to tan without a prominent checkerboard pat-
tern as in the patterned lizards. The venters are orange to drab
gray, generally slightly more orange in the pectoral region, and
grayer posteriorly. Throat and underside of forelimbs are
mottled to nearly unicolor orange, sometimes in discrete flecks.
The younger specimens have light gray throats. The underside
of the hindlimbs and tail is gray to putty colored with some
orange on the anterior surface of the femur, and in some speci-
mens on the entire underside of the hindlimb. Two small juve-
niles (snout-vent length 47 and 51 mm) are in this patternless
phase.
Eight specimens (adults and subadults of both sexes) have
patterned backs. In this phase, the dorsal ground color is gray-
green with a heavy black mottling and strikingly' tigroid barred
sides. The black dorsal x^attern quickly fades at the shoulders
and is absent or very suppressed on the neck. The dorsal blotch-
ing is much as that described for jacta, i.e., a vermiculate or at
times longitudinally arranged configuration of black on a lighter
ground. In no specimens are the lateral fields apparent and
there is no spotting or dotting on the sides. The bellies of these
458 BULLETIN: MUSEUM OF COMPARATR^ ZOOLOGY
patterned lizards were the same as those of patteruless speci-
mens; females of the patterned lizards have the extent and in-
tensity of the orange not so great as do the males. The checker-
board tail is a common feature.
As in A. c. hoekeri, we have no doubt that the two phases in
richardthomasi represent two basic patterns, and in no way
should be interpreted as an adult phase versus a juvenile and
subadult phase. Although the only two juveniles at hand are
patternless, there is an intermingling of sizes of both sexes insofar
as the two phases are concerned. Just as in hoekeri, which is rep-
resented by a much longer series, there are no intermediates be-
tween the two conditions; each lizard is distinctly in one phase
or the other .
Comparisons: The patterned phase of A. c. richardthomasi re-
quires comparison only with A. c. jacta to which race the former
is obviously closely allied. The two can easily be differentiated in
that jacta has a black gular band and the dorsal pattern con-
tinues anteriorly onto the neck, whereas richardthomasi lacks a
gular band and has the pattern faded anteriorly. Jacta also is
not known to have a patternless phase
The patternless phase of richardthomasi requires comparison
with patternless hoekeri. The two are much alike, but richard-
thomasi differs in having the filigreed or mottled lateral field
area whereas hoekeri has an obsolete lateral field and no mottling
in this region. Also the dorsal hues of hoekeri populations are
usually not greenish.
In size, richardthomasi is smaller than chrysolaema and procax,
equal to parvoris, and larger than the remaining races. In having
a mean of 87.6 fourth-toe scales, richardthomasi averages higher
than all previously named races. In femoral pore counts, it is
higher than all races except chrysolaema. Considering fifteenth
caudal verticil scales, richardthomasi is exceeded by all forms ex-
cept jacta.
Remarks: The close alliance of richardthomasi with jacta is
obvious. If we assume that extreme eastern Hispaniola is (was)
inhabited by a population with heavy dorsal mottling and
marbling, this population must also have given rise to the Saonan
subspecies. We have no evidence that jacta or a related form
still occurs on the adjacent Hispaniolan mainland. Certainh'
richardthomasi is an insular derivative of an extreme eastern
heavilv marked form which was likelv similar to jacta.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 459
Ameiva chrysolaema leberi^ new subspecies
Holohjpe: MCZ 77236, an adult male, from 5 km E Peder-
nales, Pedernales Province, Republica Dominicana, one of a series
taken 25 June 1964 by David C. Leber and Richard Thomas.
Original number V2509.
Paratypes: All from the Republica Dominicana, Pedernales
Prov., as follows: ASFS V2510-19, USNM 152577-82, RT 932,
KU 79878-81, same data as holotype; MCZ 81015-18, UIMNH
56900-05, Pedernales, 3 July 1964, D. C. Leber, R. Thomas;
AMNH 92859-61, 1 km E Pedernales, 25 July 1963, R. F. Klini-
kowski ; AMNH 92862-63, 12 km E Pedernales, 25 June 1964, R.
Thomas.
Associated specimens: Haiti, Dept. d l' Quest, Tean, nr. Sal-
trou (not mapped), 4 (MCZ 69389-92) ; Saltrou, 7 (AMNH 50000-
04, 50007-08). BepuMica Dominicana, Pedernales Prov., Oviedo,
3 (MCZ 58674-76).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of small size (males to 111 mm, females to 104 mm
snout-vent length), 10 transverse rows of ventrals, moderate num-
ber of fourth-toe subdigital scales and femoral pores, and low
number of scales in the fifteenth caudal verticil ; a completely pat-
ternless rusty brown dorsum, no lateral fields (Pig. 6, left), a
deep orange-red belly, and a black gular band which may involve
the chest and underside of the arms.
Distribution : To the south of the Massif de la Selle and Sierra
de Baoruco, from the vicinity of Saltrou in Haiti, east onto the
Peninsula de Barahona, to 12 kilometers southeast of Pedernales
(Fig. 11). The record from Oviedo is discussed below.
Description of type: An adult male with the following meas-
urements and counts : snout-vent length 103 mm, tail 233 mm ;
ventrals in 34 longitudinal and 10 transverse rows; fourth-toe
subdigital scales 40 and 42 (total 82) ; femoral pores 21 and 20
(total 41) ; 40 scales in fifteenth caudal verticil. Dorsum uniform
rusty brown anteriorly, becoming gray-brown posteriorly; sides
of head reddish brown. Lateral fields absent. Throat orange, ven-
tral ground color brick red, lower sides (and lateralmost two
rows of ventral scales) blue. A black gular Imnd, extending onto
the first four or five rows of ventrals and onto the undersides of
the arms. Tail gray above, off-white below. Dorsum and top of
tail completely unpatterned.
1 Xamed for one of the collectors.
460
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 6. Left, Ameiva c. leheri, holotype, MCZ 7720(5, '> km E Pederualcs,
Pedeniales Prov., Kepubliea Dominicana. Bight, Ameiva c. ficta, holotype,
MCZ 77237, 13.1 mi. SW Euriquillo, Pedernales Prov., Eepubliea Dominicana.
Variation: See tables. The dorsal ground color varies from
rusty brown to reddish brown, and the ventral ground color from
gray with small amounts of red to orange-red, fading posteriorly,
to brick red. The gular band is present in all specimens, and in
only one male does it not extend onto the chest. The lower sides
and lateral two rows of ventrals on each side are blue, or at least
have blue flecking, but one small male has the lateral ventrals
orange-red like the balance of the venter. The lores and cheeks
are unspotted pinkish gray. The tails are gray, unmarked above,
SCHWARTZ AND KLINIKOWSKI : AMEIVA 461
and the undersides of the tails are a grayish off-white. There is
no obvious sexual dichroniatism.
Comparisons: By virtue of its patternless dorsum, leheri can
easily be differentiated from all races except patternless hoekeri
and patternless richardthomasi. The different dorsal hues of
leheri and ricJiardfhomasi (rusty brown versus greenish gray)
and the presence of a gular band in the former and its absence in
the latter, as well as the larger adult size of richardthomasi, all
make this distinction easy. From patternless hoekeri, leheri dif-
fers in smaller adult size, and in lacking any expression of the
lateral fields, whereas boekeri retains the fields as obsolete grayish
longitudinal smudges. The vivid venters of leheri are not found
in hoekeri, and the brighter dorsal ground color of leheri con-
trasts strongly with the more drab tones of hoekeri.
Leheri is the smallest race of A. chrysolaema. In number of
fourth-toe scales, leheri averages lower (85.5) than richard-
thomasi (87.8) and chrysolaema (86.7), and is higher than the
remaining subspecies. In femoral pores, leheri again averages
less (41.3) than chrysolaema (43.7) and richardthomasi (42.6),
the same as jacta, and more than in the other races. In fifteenth
verticil scales, leheri is exceeded by all subspecies except jacta.
Remarks: A. c. leheri is not known to intergrade with either
chrysolaema to the north (from whose range it is completely
separated by the Massif de la Selle) or with the form to the east
on the Peninsula de Barahona. The three specimens from
Oviedo noted in "Associated Specimens" above, will be discussed
in detail in the treatment of the following subspecies.
AmeIVA CHRYSOLAEMA FICTA^ UCW SUbspCcicS
Holotype: MCZ 77237, an adult male, from 13.1 mi. (20.8 km)
SW Enriquillo, Pedernales Province, Republica Dominicana, one
of a series taken 22 July 1963 by Albert Schwartz and Richard
Thomas. Original number X9401.
Paratypes: All from the Repiiblica Dominicana, Pedernales
Province, as follows: ASFS X9402-09, same data as holotype;
ASFS X9950, same locality as holotype, 30 July 1963, R.
Thomas; ASFS V197-98, same locality as holotype, 4 August
1963, D. C. Leber, R. Thomas.
Associated specimens: Repuhlica Dominicana, Pedernales
Prov., 30 km from Oviedo, road to Pedernales, 1 (MCZ 58673) ;
1 Prom the I-atin for "invented, devispfl," in allusion to resemblances to abbntti.
462 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Oviedo, 15 (MCZ 58677-80, 58682-90, 58692-93) ; 5 mi. (8 km)
NE Oviedo, 11 (ASFS X9957-58, V273-80, RT 752) ; Barahona
Prov., 3 km SW Enriquillo, 1 (ASFS V290) ; Enriquillo, 2
(MCZ 58777-78).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of moderate size (males to 121 mm, females to
113 mm snout-vent length), 10 transverse rows of ventrals, low
number of fourth-toe subdigital scales, moderate number of
femoral pores and scales in the fifteenth tail verticil; dorsum
tan to brown, spotted with pale blue, lateral fields obsolescent
and often replaced by vertical blackish bars or vermiculations
(Fig. 6, right), belly rust colored, a prominently checkerboarded
tail, and a black gular band often extending onto the chest and
underside of the arms.
Distribution: The Peninsula de Barahona from (apparently),
30 km NW Oviedo in the west, east to the east coast in the vicin-
ity of Oviedo, and thence north to Enriquillo (Fig. 11) ; see
however Remarks below.
Description of type: An adult male with the following meas-
urements and counts : snout-vent length 112 mm, tail 287 mm ;
ventrals in 36 longitudinal and 10 transverse rows; fourth toe
subdigital scales 38 and 39 (total 77) ; femoral pores 21 and
20 (total 41) ; 38 scales in the fifteenth caudal verticil. Dorsal
ground color brown, with six longitudinal series of more or less
discrete pale blue spots, the lateralmost series bordering above
the remnants of the lateral fields, below which is another longi-
tudinal series of pale blue spots; lower sides with alternating
vertical black and pale blue bars ; f orelimbs vaguely spotted with
small dots, hindlimbs boldly marked Avith large rusty spots
dorsally. Throat grayish orange, belly rust colored; an exten-
sive black gular band which includes the first eight rows of
ventrals and extends onto the undersides of the arms. Tail
brown above, with blue spots on the first nine verticils dorsally,
and additionally somewhat checkerboarded; tail ivory below.
Variation: See tables. The tan to brown dorsum with pale
blue discrete spots characterizes A. c. ficta. The spots, on
occasion, may be greenish anteriorly or tan posteriorly, but
in general they are pale blue. The lateral fields are obsolete
or almost completely obliterated by vertical black bars alter-
nating wdth blue bars on the sides. The throat and ventral
ground color are grayish orange and rust, with bellies of females
slightly paler than those of males. In some individuals the
dorsal spots are distinctly lineate in appearance, and in a few
SCHWARTZ AND KLINIKOWSKI : AMEIVA 463
the back has a more or less complete finely filigreed appear-
ance, although this is not the norm. The spots themselves vary
in size, distribution, and density ; they may be much smaller
than in the type and much more closely appressed to one an-
other, or the paramedian rows (if rows are discernible) may
be fused to form a pair of paramedian pale blue lines. The
checkerboard tail with blue spotting on its basal portion is a
common feature.
Comparisons: Only one other race thus far discussed, A. c.
parvoris, has a spotted dorsum, although A. c. chrysolaema has
a pattern of spots and lines. In neither of these two races are
the dorsal spots pale blue, but are rather some shade of yellow.
In actuality, the pattern of chrysolaema, although dotted, bears
little resemblance to that of ficta; the pattern of parvoris is
similar but the coloration and general aspect of the lizards of
these two races are quite distinctive. Parvoris lacks a conspicu-
ously checkerboarded tail.
Compared to the described races, jicta is exceeded in size
by all other forms except leheri, which is still smaller. In
number of fourth-toe scales, ficta is exceeded by all races except
jacta. Chrysolaema, richardthomasi, jacta, and leheri exceed
ficta in mean number of femoral pores, and in this character ficta
exceeds the balance of the races. Ficta averages higher in
fifteenth verticil scales than richarclthomasi, leheri, and jacta,
and lower than the other races.
Remarks: The distribution of A. c. ficta encompasses the east-
ern shore of the Peninsula de Barahona from Enriquillo south
to Oviedo, and thence inland toward Pedernales for a distance of
30 kilometers. Ficta is not known to intergrade with either
imibratilis to the north or leheri to the west (but see below). The
northernmost station for ficta is 40 kilometers from the closest
record of umhratilis ; we presume that these two races do not
come in contact because of the, at best, narrow and intermittent
nature of suitable habitats for chrysolaema along the east coast
of the Peninsula de Barahona.
There are three specimens (MCZ 58674-76) from Oviedo which
are clearly leberi and in no way resemble ficta. Assuming
that these specimens did indeed come from Oviedo, they present
a problem. They are the only specimens from the entire eastern
coast of the Peninsula de Barahona which are patternless ; in our
considerable collecting experience in the Oviedo region, we never
encountered nor collected any leheri-\ike individuals. There are
464 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
several possibilities ; all of which depend upon the assumed cor-
rectness of the locality data for these three specimens: 1) they
represent a Ze&en-phase of ficta; 2) leheri and ficta are not both
subspecies of chrysolaema; 3) leheri and ficta are both chrysolaema
derivatives but one has been so long separated from the parent
stock that the two forms act as species, with a rather broad (30
km) region of sympatry.
In defense of our arrangement of these two forms as subspecies
of A. chrysolaema, the following comments are pertinent: 1)
There is no incontrovertible evidence that these three lizards are a
patternless phase of ficta. All of our own Oviedo material is pat-
terned, and there is no indication that there exists a patternless
phase oi ficta (although of course there is this possibility). 2) Since
neither Ichcri nor ficta intergrades with any other subspecies for
reasons of geography, and since both have apparently been long
isolated from chrysolaema and umhraiilis to the north, one or both
might be logically regarded as a distinct species (if so, then
ahbotti and ficta would compose one species, or leheri could be
so regarded). We feel that leheri, despite its complete isolation
from chrysolaema, is so like patternless boekeri and richard-
thomasi that to regard it as a distinct species would be mislead-
ing and obscure its clear relationships to the balance of A.
chrysolaema. A somewhat stronger case may be made for sep-
arating ahhotti and ficta at the species level; here again, how-
ever, the resemblance of both these spotted forms to parvoris for
example (as well as the overall similarities of ahhotti- ficta to
the more northern subspecies) tends in our opinion to negate
removing these two forms from the species chrysolaema. 3) The
most appealing interpretation is that one (leheri) of the two in-
volved forms has been long separated from its parent stock (A. c.
chrysolaema), and that once contact between it and another
subspecies (ficta) has been re-established, the two forms do not
intergrade but act as separate species. The present lack of
contact between leheri and chrysolaema and between ficta and
umhratilis suggests that both forms may well have had long inde-
pendent histories. It is even not improbable that ficta has been
derived from aMotti, rather than the reverse, and thus ahhotti
may have been insularly isolated from leheri. Such a combination
of situations might argue for species status for both leheri and
ahhotti-ficta and we have considered this possibility. On the other
hand, such a decision obscures the obvious relationships of these
two forms to A. chrysolaema (in contrast, for instance, to A.
taeniura or A. lineolata) .
SCHWARTZ AND KLINIKOWSKI : AMEIVA 465
Finally, and probably the most important point is that the
region between Oviedo and Pedernales still remains little known
herpetologically ; there is alwaj^s the possibility that the pre-
sumed leheri from Oviedo are in actuality from farther west and
thus from within the known range 0/ leheri. We have tried to
adhere in this ease to a via media, and rather than make assump-
tions from inadequate data, we consider both leheri and ficta sub-
species of A. chrysolaema, although admitting that the situa-
tion is not completely clear. Exclusive of these three question-
able lizards, the ranges of jicta and leheri approach, very
closely; the distance between the nearest localities for the two
races is only 15 kilometers.
AmEIVA CHRYSOLAEMA ABBOTTI Noble, 1923
Ameiva ahhotti Noble, 1923, Amer. Mus. Novitates, 64:1 (type locality
— Isla Beata, Eepublica Dominicana).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of small size (males to 117 mm, females to 108 mm
snout-vent length), usually 12 transverse rows of ventrals, high
number of fourth-toe subdigital scales and femoral pores, and
moderate number of scales in the fifteenth verticil; dorsum
black with a pattern of isolated spots which are orange or
yellowish, becoming blue anteriorly, lateral field absent, the sides
spotted with sky-blue spots (Fig. 7, left), venter deep brick red
to orange posteriorly, and a black gular band which expands
to cover the chest and underside of the arms.
Distrihution: Known only from Isla Beata, off the tip of Cabo
Beata (Fig. 11).
Disciissio7i: A. c. ahhotti is the most brilliantly colored and
striking of the races of A. chrysolaema. The combination of black
dorsal coloration, orange to yellowish spots middorsally, becom-
ing blue anteriorly, and the vivid blue lateral spots provides a
particularly colorful lizard. The forelimbs are black to brown
(distally) with blue spots, and the hindlimbs black with prox-
imally blue and distally orange spots. The venter is brick red,
grading to orange or pinkish posteriorly, and the lateralmost
ventrals are invaded by blue and white spotting. The heads
are tan to orange with white or bluish spots on the sides. There
is a bold black pectoral band which extends onto the chest and
even onto the venter and the undersides of the arms. The
underside of the hindlimbs is orange on the thighs and orange
466
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 7. Left, Ameiva c. abbotti, ASFS V2743, Isla Beata, Eepublica
Dominicana. Eight, Ameiva c. secessa, holotype, MCZ 77238, Etroits, He
de la Gonave, Haiti.
to light tan on the crura. The upperside of the tail is checker-
boarded black and white or cream; the underside of the tail is
gray to greenish on its proximal half to two-thirds, and uniform
light tan to pinkish distally. The dorsal spots are invariably
discrete and there is neither a tendency for them to become
lineate or to be fused into longitudinal lines. The females are
colored and patterned like the males; a juvenile lacks the bright
orange ventral color and the black pectoral area.
A. c. ahhoiti is so very distinctive in its dorsal coloration and
pattern that it is hardly necessary to compare it with any other
SCHWARTZ AND KLINIKOWSKI : AMEIVA 467
subspecies. Closest, at least in pattern, are parvoris and ficta;
the former, although a dorsally spotted race, is not so gaudy and
lacks the discrete spotting so characteristic of abhotti. The ad-
jacent race ficta, on the mainland, resembles ahhotti in basic
pattern, but differs in having the back brown rather than
black, in having the dorsal spots pale blue rather than orange
to yellowish, in having the spots at times arranged into lines,
and in having the spotting on the back quite variable in density.
In contrast, ahhotti is surprisingly constant in density of dorsal
spotting. Ficta is primarily a race with 10 rows of ventrals.
whereas ahhotti usually has 12.
Variation: See tables. A. c. ahhotti has the highest mean (89.6)
of fourth-toe scales of any race described to this point, and is ap-
proached only by richardthomasi (87.8) ; of all races of A. chry-
solaema, ahhotti has the highest average of femoral pores (43.8),
although it is closely approached by chrysolaema (43.7). AVith a
mean of 40.4 fifteenth verticil scales, ahhotti exceeds ficta, richard-
thomasi, leheri and jacta, and has less verticil scales than the
other races.
The relationships of abhotti are obviously with ficta on the
mainland. Whether the latter occurs on the southern tip of
the Peninsula de Barahona is unknown, but it certainly is a
more likely candidate there than the drab and patternless leheri,
if we assume that abhotti was derived directly from the adjacent
mainland.
Specimens examined: Repuhlica Dominicana, Isla Beata, 39
(ASFS V2743-69, MCZ 28571-73, 37578-79, 37581-82, 17676-77,
57049, RT 934, UMMZ 83098).
Ameiva chrysolaema secessa^ new subspecies
Holotype: MCZ 77238, an adult male, from Etroits, He de
la Gonave, Haiti, taken 17 July 1962 by Elie Cyphale. Original
number X2447.
Paratypes: All from He de la Gonave, as follows: ASFS
X2440-46, X2448-59, UIMNH 56906-09, USNM 152583-87, AMNH
92864-69, KU 79882-86, same data as holotype ; USNM 80377-78,
Pointe Quest, 21 March 1930, L. H. Parish and W. Perrygo;
USNM 77062-69, MCZ 25539-48, Pointe a Raquette, August
1927, W. J. Eyerdam; MCZ 80251-78, Pointe a Raquette, sum-
mer 1964, G. Whiteman; MCZ 80231-36, Nan Palmiste, 4 km
1 From the Latin for "distant, removed."
468 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
from Pointe a Kaquette, summer 1964, G. Whiteman; MCZ
80237-50, Ti Roche, 0.5 km from Pointe a Raquette, summer 1964,
G. Whiteman; USNM 80359-68, 80370-76, UMMZ, 92196, Anse a
Galets, 23 March 1930, L. H. Parish and W. Perrygo; MCZ
37568-77, Anse a Galets, 9 April 1934, T. Barbour; USNM
76803, Nan Cafe, March 1929, A. J. Poole and W. Perrygo.
Associated specimens: He de la Gondve (no other locality),
3 (CM 8133, MCZ 12870-71).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of large size (males to 134 mm, females to 111 mm
snout-vent length), usually 12 transverse rovs^s of ventrals, high
number of fourth-toe subdigital scales, and moderate number
of femoral pores and scales in the fifteenth caudal verticil ; dor-
sum reddish brown with six or seven dull buffy longitudinal
lines, grayish brown (rather than black) lateral fields with
isolated buffy dots enclosed therein (Fig. 7, right), and with the
black gular band usually absent, or at least very restricted,
seldom involving the anterior ventrals but at times extending
onto the underside of the arms.
Distribution: He de la Gonave, Haiti (Fig. 11).
Description of type: An adult male with the following meas-
urements and counts: snout-vent length, 117 mm, tail 134 mm,
broken; ventrals in 39 longitudinal and 10 transverse rows;
fourth-toe subdigital scales 45 and 45 (total 90) ; femoral pores
21 and 21 (total 42) ; 42 scales in the fifteenth caudal verticil.
Dorsal ground color reddish brown with seven dull buffy longi-
tudinal lines, the median line somewhat broken and indistinct;
head dull tan, neck greenish ; sides of head gray with whitish
blotches. Lateral fields grayish brown with an enclosed series
of buffy dots, more distinct posteriorly than anteriorly, the
lateral fields set off above by the lateralmost dorsal lines, and
below by a series of bluish spots; lower sides spotted with blue.
Throat dirtj^ pinkish gray, gular band absent, belly gray. Fore-
limbs with scattered pale greenish blue spots, hindlimbs pro-
fusely dotted with pale yellow. Tail reddish brown above, grayish
blue below, with some blue scales on the sides.
Variation: See tables. The large series of A. c. secessa at hand
shows little variation in pattern ; the entire animal invariably is
quite dull, and the lateral fields are never conspicuous. The longi-
tudinal lines do not contrast especially strongly with the dorsal
ground color and in some topotypes are very obscure and are
seen with some difficulty. In many specimens there are six
SCHWARTZ AND KLINIKOWSKI : AMEIVA 469
(rather than seven) dorsal lines, the median line being absent. In-
variably the gular band is poorly developed or completely absent ;
if the band is present, it does not involve the anterior ventrals
but may send some pigment onto the underside of the arms. The
coloration of the venter varies from gray and bluish gray to dull
orange-gray. There is no sexual dichromatism.
Comparisons: A. c. secessa is so dull and drab compared to
all other races that no comparison is really necessary. It differs
from the spotted races parvoris, ficta, and ahhotti in being lon-
gitudinally lined, and from the patternless races hoekeri, richard-
thomasi and leheri in having a pattern. It is much duller pat-
terned, and likewise differently patterned, from the other lined
races — clirysolaema, mnhratilis, alacris, and patterned hoekeri.
From jacta and richardthomasi (in the patterned phase), secessa
differs in lacking the lateral tigroid markings and in having a
quite different dorsal pattern. Perhaps the most cogent compari-
son is with clirysolaema which occupies all the adjacent mainland
about the Golfe de la Gonave. From clirysolaema, secessa can
at once be differentiated by its much more drab coloration and
pattern, and by the lack of dotting in combination with lines on
the dorsum. Both chrysolacma and secessa are typically 12-row
lizards.
In fourth toe scales, secessa has a higher mean than any other
race, being approached most closely by ahhotti (89.6). In num-
ber of femoral pores, secessa is exceeded only by ahhotti, chryso-
laema and richardthomasi; in fifteenth verticil scales, secessa,
exceeds ficta, richardthomasi, leheri and jacta, and is exceeded by
the means of the balance of the subspecies.
Remarks: Although A. c. secessa, has presumably evolved from
the adjacent A. c. clirysolaema, in dorsal pattern it grossly
resembles alacris and procax, but is quite distinct in several
features, notably the obscure lateral fields and the lack of
a gular band. It seems likely that the nominate race has
carried the pattern evolution — i.e., disintegration of the longi-
tudinal lines into a series of dots — farther than has the isolated
secessa which has become faded and pale in contrast to its more
brightly colored neighbor. Another possible origin of secessa is
discussed below.
470 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Ameiva chrysolaema DEFENSOR^ iiew subspecies
Holotype: MCZ 63379, an adult male, from Mole St. Nicholas,
Dept. du Nord Quest, Haiti, one of a series taken 24-29 July
1960 by A. S. Kand and J. D. Lazell, Jr.
Paratypes: All from Haiti, Dept. du Nord Quest, as follows:
MCZ 63368-72, 63374-78, same data as holotype; MCZ 63364-
67, Jean Rabel, 26 July 1960, A. S. Rand and J. D. Lazell, Jr. ;
AMNH 49856-57, Port a I'Ecu, 1 April 1935, W. G. Hassler;
USNM 59925, Bale des Moustiques, 3 May 1917, W. L. Abbott ;
AMNH 49851-55, river just W Port-de-Paix, 2 April 1935, W. G.
Hassler ; MCZ 58014, river just W Port-de-Paix, 2 August 1935,
W. G. Hassler.
Associated specimens: Haiti, Dept. du Nord Quest: Bombar-
dopolis, 1 (MCZ 63381) ; Dept. de VArtibonite, Gros-Morne, 1
(MCZ 63380).
Diagnosis: A subspecies of A. c. chrysolaema characterized by
a combination of moderate size (males to 126 mm, females to
106 mm snout-vent length), 10 transverse rows of ventrals, low
number of fourth toe subdigital scales and scales in the fifteenth
caudal verticil, and moderate number of femoral pores ; dorsal
pattern a series of six or seven dull longitudinal lines on a tan to
brown background, lateral fields dull brown, not especially con-
trasting with the dorsal ground color and often with the included
light spots in the lateral field much reduced or completely absent,
a checkerboard tail pattern (Fig. 8, left), and no indication of a
black gular band, black on the anterior ventrals, or extension of
black pigment onto the underside of the arms.
Distribution: The northwest peninsula of Haiti, from Bom-
bardopolis in the south to the vicinity of Port-de-Paix in the
northeast, and thence south to Gros-Morne (Fig. 11).
Description of type: An adult male with the following meas-
urements and counts: snout-vent length 118 mm, tail 242 mm,
partially regenerated; ventrals in 38 longitudinal and 12 trans-
verse rows; fourth-toe subdigital scales 44 and 44 (total 88), fem-
oral pores 19 and 18 (total 37) ; 38 scales in the fifteenth verticil.
Dorsal ground color (in preservative) dull brown with a series
of seven tan longitudinal lines, the median line the least conspic-
uous, all lines disappearing on the neck. Lateral fields brown,
with included tan dots only in their posterior thirds, the anterior
1 For the Latin for "defender" in allusion to the English fort at Mole St.
Nicholas which guarded the Windward Passage.
SCHWARTZ AND KLINIKOWSKI : AMEIVA
471
Fig. 8. Left, Ameiva c. defensor, holotype, MCZ 63379, Mole St. Nicholas,
Dept. du Nord Quest, Haiti. Bight, Ameiva c. woodi, unnumbered specimen
from MCZ 37583-92, He de la Tortue, Haiti.
two-thirds being without dots. Forelimbs grayish tan, vaguely
dotted, hindlimbs brown with large pale spots, leaving almost a
reticulum of dark brown surrounding the large pale areas. Ven-
tral ground color (including throat) bluish gray, no black pig-
ment on throat, chest, or undersides of arms. Lower sides with
gray and blue markings which are almost tigroid. Tail tan, heav-
ily checkerboarded with dark brown above, blue-gray marked
with cream below, and with black and some cream on sides.
472 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Variation: See tables. We are somewhat handicapped in
discussing A. c. defensor since we have not seen this subspecies
in life. Judging from the material at hand, especially the
fresh specimens from the Museum of Comparative Zoology
(old material in the American Museum from Port-de-Paix
and Port a I'Ecu is so discolored that it is completely worth-
less insofar as coloration is concerned but does still retain
some evidences of pattern), defensor is typically a dull and
drab lizard with dorsal coloration of tan to brown with six
or seven longitudinal buffy lines. The lateral fields are brown
and have the included dots much reduced (often absent an-
teriorly) or completely absent. In the latter case the lateral
field presents an unbroken brow^n lateral band. In some speci-
mens, the longitudinal lines have become more obscure than in
the type, due to light pigmented areas in the interline regions,
and in one extreme case (IMCZ 63378 — snout-vent 119, and
thus not the largest male) the entire back is marbled with dark
and light and the sides are tigroid, the latter a condition noted
to a slighter degree in some other individuals. No specimen has
any indication of black on the chest or undersides of the arms,
and the gular black band is lacking completely. The prom-
inently checkerboarded tail is a constant feature and is plainly
discernible even in old and discolored individuals.
Comparisons: A. c. defensor most closely resembles A. c. seccssa
from He de la Gonave. However, the reduced or absent dotting
in the lateral fields, smaller size, and the generally somewhat
brighter dorsal pattern (although defensor is nonetheless a
rather drab lizard) will distinguish the two races. Secessa usu-
ally has 12 rows of ventrals, defensor usually has 10. From
the patternless races, def elisor can be distinguished in having a
pattern, and from the spotted subspecies by having a dorsal pat-
tern of longitudinal lines. From the other lined races, def elisor
differs in the lack of a black lateral field with included yellow
dots, and lack of a black gular band.
In fourth-toe scales, defensor (82.4) exceeds only ficta (81.9)
and jacia (79.7) ; in number of femoral pores, defensor (37.2)
exceeds only loekeri (36.5), procax (36.1), umlratiUs (35.6), and
alacris (33.8). In fifteenth verticil scales, defensor (38.0) ex-
ceeds only leheri (37.9) and jacta (35.7). In counts of fourth-
toe scales and fifteenth verticil scales, defensor is quite low in
the series of subspecies.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 473
Remarks: A. c. defensor is not known to intergrade with any
other subspecies ; there are two wide hiatuses, however. The
closest approximation of records for defensor (Gros-Morne) and
chrysolacma (St. Marc) is 83 kilometers. To the east, there are
no specimens available between Port-de-Paix and Cap-Ha'itien,
a distance of 70 kilometers.
Although A. c. secessa is closer geographically to A. c. chryso-
laema, the former is much more similar to defensor than to the
nominate race. Such a similarity may be merely convergence
and may not reflect direct relationships. It is possible, on the
other hand, that defensor has reached the northwest peninsula
from GonPive ; it seems hardly likely that the reverse is true —
i.e., that Gonave has lieen colonized from the north — consider-
ing the proximity of Gonave to the adjacent mainland (21 kilom-
eters at its closest point) and its distance from the northwest
peninsula (72 kilometers at its closest point).
Ameiva chrysolaema woodi Cochran, 1923
Ameiva chrysolaemea woodi Cochran, 1923, Occ. Papers Boston Soc. Nat.
Hist., 8:181 (type locality — He Tortue, Haiti).
Diagnosis: A subspecies of A. chrysolaema characterized by
a combination of large size (males to 141 mm, females to 126 mm
snout-vent length), 10 transverse rows of ventrals, low number
of fourth toe subdigital scales, moderate number of femoral pores
and scales in the fifteenth verticil ; dorsum very dark brown with
three to five dull yellowish to buffy stripes or a median dorsal
buffy longitudinal band ; the stripes may be variously joined and
modified to give rather complex dorsal figures which are derived
from the simple five lines (Figs. 8, right; 9) ; sides with vertical
tigroid markings, and no black gular band or extensions thereof
onto the chest and underside of the arms.
Distribution: He de la Tortue, Haiti (Fig. 11).
Discmsion: The most strikingly patterned and at the same
time most variable of the races is A. c. ivoodi. Basically the
dorsal pattern is a series of three to five broad, dull yellowish
to buffy longitudinal lines on a very dark brown ground. This
basic pattern may be modified in that the area between the two
paramedian lines may be filled in with a buffy color so that the
back has a lateral pair of pale lines and a middorsal pale zone.
From this condition, the balance of the back may be filled in
with paler, so that the entire back is marked with a single broad
pale middorsal zone. In two individuals, the pale lines have
474
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Fig. 9. Left, Ameiva c. woodi, ASFS X2270, Palmiste, He de la Tortue,
Haiti. Eight, Ameiva e. woodi, ASFS X2267, Palmiste, He de la Tortue,
Haiti.
grossly fragmented and joined randomly, to form a bizarre,
longitudinally reticulate pattern -which is quite distinctive. The
heads are dull gray-green, with the lores gray. The sides are
tan to grayish brown ; this lateral color invades the dark brown
back to give a series of alternating dark brown and tan vertical
tigroid bars, the tan bars often faintly brick colored at their
dorsal points. The throats are dirty pale orange, those of the
females usually lighter than those of the males (although one
female has a brighter orange throat than any other specimen
SCHWARTZ AND KLINIKOWSKI : AMEIVA 475
examined in life). The venter is gray, occasionally with a pale
orange wash. No specimen has any indication of a black gular
band or anj- black on the chest and undersides of the arms, al-
though there may be some isolated black flecking on the chest.
The tails are tan dorsall3^ wdth prominent black markings, giv-
ing a strong checkerboard effect. The undersides of the tails are
gray and often have almost as prominent checkerboarding as
the upper surfaces.
Variation: See tables. In pattern, no other race of A. chryso-
laema is comparable to tvoodi; the three to five longitudinal lines
are fewer than in the pattern regularly noted in other subspecies,
and the peculiar middorsal broad band, either with or without
two dorsolateral lines, and the irregular fragmentation and join-
ing of the lines on a dark brown ground are all features which
woodi shares with no other race. In lacking a gular band, woodi
is comparable only to the adjacent defensor, and seccssa and
richardtJiomasi. All other forms have the band present. A. c.
ivoodi is a remarkably distinct form; it resembles none of the
mainland races and is particularly unlike the adjacent mainland
defensor.
A. c. ivoodi is a large subspecies, being exceeded only by
chrysolaema in size, although procax is equal in snout-vent
length. Woodi, in having a low mean (80.5) of fourth-toe scales,
exceeds only jaeta (79.7) in this count. In femoral pore count,
woodi is exceeded by abhotti, chrysolaema, ricliardthomasi , and
secessa, and has a mean femoral pore count equal to those of jacta
and leheri. The moderate fifteenth verticil coimt (38.8) of
woodi exceeds only richardthomasi, defensor, leheri, and jacta,
and is equal to that of ficta.
The derivation of woodi must certainly be from the adjacent
defensor of the mainland. The lack of a gular band, and black
on the chest and undersides of the arms, and the ten rows of
ventrals are features in common between the two races. There
the resemblances cease, however, since woodi is a boldly and color-
fully patterned lizard, whereas defensor is dull and drab. De-
fensor likewise does not exhibit any patterns which are reminis-
cent of those of vwodi, although the lack of dots in the lateral
fields may foreshadow the absence of these fields entirely in
woodi.
Specimens examined: Haiti, He de la Tortiie, Palmiste. 10
(ASFS X2267-76) ; no specific locality on the island, 29 (MCZ
37583-92 -f 19 unnumbered specimens).
476
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Ameiva chrysolaema regularis Fischer, 1888
Ameiva regularis Fischer, 1888, Jahrb. Wiss. Anst. Hamburg, 5:26 (type
locality, Sans Souci, Haiti; this locality is the palace of the same
name near Milot, Dept. du Nord, Haiti).
Diagnosis: A subspecies of A. chrysolaema characterized by a
combination of large size (males to 132 mm, females to 128 mm
snout-vent length), usually 12 transverse rows of ventrals, very
low number of fourth-toe subcligital scales, moderate number of
femoral pores, and high number of scales in the fifteenth verticil ;
dorsal pattern a series of five to seven pale yellow lines on a tan
to browm ground color, occasionally with a clear tan middorsal
zone, neck greenish and dorsal ground color often suffused with
Fig. 10. Ameiva c. regularis. ASFS Y-1215, 9 km NW Villa Vasquez,
Monte Cristi Prov., Republiea Dominicana.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 477
blackish, lateral fields black with an included row of yellow
dots (Fig. 10), and black gular band present or absent, when
present seldom invading the chest or extending onto the under-
side of the arms.
Distribution: North central Hispaniola, from Cap-Haitien
and Grande Riviere du Nord in the west, east to Fort Liberte,
Haiti, and thence to Monte Cristi and throughout the Valle de
Cibao as far east as the vicinity of Santiago, Republica Domini-
cana; also the Siete Hermanos islands (known from Isla Muer-
tos, Toruru, Monte Chico, and Tercero) and Isla Cabras to the
north of Monte Cristi (Fig. 11).
Discussio7i: Cochran (1941 : pi. 8, figs. B and D) has illustrated
two phases in the dorsal pattern of A. c. regularis from Cap-
Ha'itien, which is near the type locality of the subspecies. Gen-
erally, throughout its wide range on the mainland, regularis is
fairly consistent in dorsal pattern. Specimens from the Valle
de Cibao, Monte Cristi, and Pepillo Salcedo, which we have
seen in life, were brown dorsally, often suffused with blackish,
with a greenish wash on the neck, and had five to seven longi-
tudinal pale yellow lines. At times there is a clear tan mid-
dorsal zone resulting from fusion of stripes and filling in of
the interspaces with tan. The lateral field is black and prom-
inent with a longitudinal series of yellow dots. The lower sides
are dotted with blue-green, and the sides of the belly are bright
blue. The ventral ground color varies from grayish to deep dull
orange, and the throat from yellowish to gray-orange. The fore-
limbs are spotted with blue-green, the hindlimbs with yellow.
The gular band may be either present or absent ; if present, it is
not extensive and seldom encroaches upon the chest or extends
onto the undersides of the arms.
There are variants of the above basic pattern, such as that
shown by Cochran (1941: pi. 8, fig. D), in which there are faint
filigreed lines in a middorsal zone. Some specimens show acces-
sory dots between the longitudinal lines, especially posteriorly;
in the series from near Villa Vasquez, Monte Cristi Province,
two in life clearly showed a secondary dorsal condition, similar
to that typical of adult A. c. cJirysolaema, in which the dorsal
lines are supplanted by a series of bright yellow dots overlying
the fainter longitudinal lines. In general, specimens from near
Cap-Haitien seem somewhat darker than those from the xeric
Valle de Cibao, but in features of pattern they are not remark-
ably different from those from the Valle de Cibao. Occasional
478 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
specimens from the western portion of the range of regularis
have the yellow dots in the lateral fields very tiny, and the field
thus appears, grossly, to be immaculate, as is characteristic of
some defensor.
A series of three adult male lizards from Isla Cabras, off
the coast just north of Monte Cristi differs from mainland ma-
terial in having dark brown lateral fields, and yellow dots on
the lower sides. These three specimens also have extensive black
gular bands involving the chest and the undersides of the arms.
Neither in size nor scalation do there seem to be any differences
betw^een these lizards and those from the adjacent mainland. We
consider them as regularis since in most features of pattern and
coloration they are very close to that race.
There are 17 specimens from the Siete Hermanos, a group of
seven islets off the mouth of the Kio Yaque del Norte. Of these
lizards, ten are from Isla Muertos, two from Isla Tercero, two
from Isla Toruru, and three from Isla Monte Chico. In colora-
tion, pattern, and scalation they do not differ from mainland
specimens, and we regard them as regularis.
Variation : See tables. A. c. regularis may be differentiated by
its lined pattern from the patternless races — hoekeri, leheri,
richardtJiomasi — and those which have distinct patterns (dots,
vermiculations, etc.) — parvoris, jacta, ricJiarcWwmasi, fief a, ab-
hotti. From the lined races, regularis differs in lacking a dotted
and lined dorsum in combination (chrysolaema), in having a
dark ground color dorsally and complete and black lateral fields
(defensor, secessa), in almost always lacking a pattern of a mid-
dorsal tan zone and at times having a black gular band (woodi),
and in having twelve rather than ten rows of ventrals (hoekeri,
alacris, procax, umhratilis). Begularis most closely resembles
alacris and procax; compared with alacris, regularis reaches a
larger size and averages fewer fourth-toe subdigital scales (77.8
versus 84.8). From procax, regularis differs in smaller adult size
and in often lacking a black gular band, which procax regularly
possesses. The two subspecies differ in fourth-toe scales, with
regidaris having a lower mean (77.8) than procax (84.8).
The north central subspecies is not known to intergrade with
either of its neighbors ; there are no specimens to the west between
Cap Ha'itien (regularis) and Gros-Morne (defensor) , a distance of
70 kilometers. The closest approximation of regularis (Fort
Liberte) and alacris (Cerca-la-Source) is 58 kilometers. The
intervening mountains almost certainly completely separate these
two subspecies.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 479
The distribution of A. c. rcgularis along the northeastern coast
of Haiti and thence into the Valle de Cibao in the Republica
Dominicana suggests that this form has evolved along the north-
ern coast and thence has penetrated into the xeric cul-de-sac of
the Valle de Cibao. The affinities of rcgularis with alacris sug-
gest strongly that the parent stock has been the latter, yet we
cannot visualize any means of dispersal of alacris to the north
Haitian coast ; presently at least the Massif du Nord and the
Cordillera Central form insurmountable barriers. One route of
dispersal suggests itself : the valleys of the Grande Riviere du
Nord and the Riviere Bouyaha (the latter a member of the
Artibonite system whose upper valleys are occupied by alacris)
approach each other in the Departement du Nord. These valleys
and their approximation may have offered a means of ingress
for Ameiva from the south into the northern Haitian littoral.
Specimens examined: Haiti, Dipt, du Nord, Cap-Haitien, 14
(USNM 74075-86, MCZ 37593-94) ; Ti Guinin nr. Cap-Haitien,
(not mapped), 24 (UMMZ 122819 [=12 specimens], MCZ 66527-
38) ; Grande Riviere du Nord, 46 (UMMZ 122820 [=12 speci-
mens], MCZ 63353-63, 66514-26); Fort Liberte, 6 (USNM
76770-75) ; RepuMica Dominicana, Monte Cristi Prov., Laguna
de Salodillo, 7 km S Pepillo Salcedo, 1 (ASFS V1430) ; 4 km
E Pepillo Salcedo, 8 (ASFS V1149-55, V1166) ; Isla Cabras,
3 (ASFS V1372-74) ; Monte Cristi, 1 (MCZ 58018) ; 2 km SE
Monte Cristi, 5 (ASFS V1210-12, V1284-86) ; 9 km NW Villa
Vasquez, 14 (ASFS V1214-25, RT 811-12) ; 5 km W Guayubm, 15
(ASFS V1494-508) ; 7 km N Guayubin, 13 (ASFS V1471-83) ;
Valverde Prov., 9 km N Los Quemados, 1 (ASFS V1766) ; 7 km
E Valverde, 10 (ASFS V2931-40) ; 2 km E Esperanza, 5 (ASFS
V1755-59) ; Santiago Prov., 7 km W Santiago, 2 (ASFS V2925-
26) ; Santiago and vicinity, 6 (MCZ 58665-66, 58668-71) ; Siete
Hermanos, Isla Muertos, 10 (ASFS V1590-95, RT 826, USNM
76733-35) ; Isla Monte Chico, 3 (USNM 76715-17) ; Isla Ter-
cero, 2 (USNM 76736-37) ; Isla Toruru, 2 (ASFS V1573-74).
DISCUSSION
Before proceeding to a discussion of the variation and possible
history of Ameiva chrysolaema in Hispaniola, we would like to
bring out several facts which seem especially worthy of mention.
The distribution of patternless races, or at least races which
have some patternless members (leheri, hoekeri, richardthomasi),
480
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
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SCHWARTZ AND KLINIKOWSKI : AMEIVA 481
is especially interesting. Of these subspecies, one (leheri) is iso-
lated on the south shore of His^Daniola below the La Selle-Baoruco
massifs; leheri approaches, insofar as known without intergrada-
tion, the very different patterned race ficta on the Peninsula de
Barahona. Another (hoekeri) has patterned individuals, and is
surrounded by three patterned races, of which it intergrades
with one {alacris), is separated from another (umhratilis) by
the Rio Yaque del Sur, and from the third {procax) by a
distinct and dramatic change of environment. Finally, richard-
fhomasi on Saona has patterned and patternless individuals;
this subspecies is related most closely to jacta.
Three races are dorsally spotted : ahhotti and ficta on Beata
and the Peninsula de Barahona, respectively, and parvoris on
the southeastern coast of Hispaniola. Parvoris is separated from
its neighbor to the west, procax, by the Rio Ozama, and from
its eastern neighbor, jacta, by a wide gap which is apparently
})resently uninhabited by A. chrjjsolaema. The occurrence of
parvoris on Isla Catalina, to the east of the known mainland
distribution of that race, is noteworthy.
Of the striped races, chrysolaema stands alone in its large size
and its style of dorsal patterning, a combination of lines and
dots. Chrysolaema is known to intergrade with alacris, another
striped race, in the vicinity of Mirebalais, but no intergrades are
known between chrysolaema and unihratilis in the Cul de Sac-
Valle de Neiba. rmhraiilis resembles the striped phase of hoekeri
to some extent ; the two are not known to intergrade. The races
procax, ala<;ris, and regularis are all comparably striped ; of
them, alacris and procax have 10 rows of ventrals, whereas
regularis has 12 rows of ventrals. The range 0/ alacris is sep-
arated from that of procax by the intervening and quite different
hoekeri. Defensor, by virtue of its pallid coloration and drab pat-
tern, stands alone among the mainland races, but it is approached
somewhat by the drab secessa from Gonave ; secessa has 12 rows
of ventrals. rkfensor 10. The Tortue subspecies woodi is very
distinctive, I)ut logically must have been derived from either
defensor or regularis, the only two races on the north coast.
The Sierra de Xeiba and the Montagues du Trou d'Eau form
the northern boundary of the Cul de Sac-Valle de Neiba plain,
which is in actuality a fossil strait that once separated His-
paniola into two distinct islands, the north and south islands
The south side of the same plain is bounded by the Massif de la
Selle and its associated northern ranges (Morne I'Hopital, ]\Iont
482 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
des Enfants Perdus) and the Sierra de Baoruco. Considering
only this region, one is struck by the amazing diversity of the
four races of A. chrysolaema associated with it : chrysolaema in
the northwest, umhratilis in the northeast, ficta in the southeast
and leberi in the southwest. Here we are involved with four
races whose patterns are radically different — chrysolaema dark
with longitudinal lines and dots, umbratilis pale with lines, ficta
with large dorsal spots, and leheri without pattern.
It has become customary to catalogue, if possible, Hispaniolan
reptiles and amphibians into either north or south island species
— i.e., depending upon their present and presumed past distribu-
tion. Although this is somewhat difficult in the present case,
we feel that A. chrysolaema is a north island species. Except for
the isolated occurrence of A. c. chrysolaema at Aquin on the
Tiburon Peninsula (based on a single specimen), this south-
western extremity of Hispaniola lacks the species; the otherwise
westernmost record is from Pere, near Leogane. The occurrence
of the very different race ficta on the Peninsula de Barahona
(and its relative ahhotti on Beata), as well as leheri to the west
along the south coast, indicates that the A. chrysolaema stock was
long isolated on the southeastern portion of the south island,
where ficta evolved in isolation from the northern mass of the
species. The presence of leheri along this south shore is most
puzzling, especially since it is geographically closest to ficta (see
Remarks under A. c. ficta for additional comments), and since
it also resembles hoekeri far to the northeast. Perhaps leheri is
the more ancient of the two mainland south island races, and its
present rather restricted distribution a mere remnant of a range
which was once more extensive, especially to the west toward
Jacmel. If such is the case, leheri might be regarded as a sub-
species derived from (pre) chrysolaema; a possible source of
colonization from the northern shore of the Tiburon Peninsula
might be the Vallee de Trouin, the low pass between the north
and south shores of the peninsula. It is of course quite possible
that additional collecting in the vicinity of Jacmel or between
that city and Saltrou will reveal the presence of leheri; it is
also possible that chrysolaema or chrysolaema X leheri inter-
grades may l)e found in the Vallee de Trouin as well.
On the north island, and including the Cul de Sac-Valle de
Neiba, we visualize the old coast inhabited by two races, chryso-
laema and umhrotilis, much as today procax and parvoris occur
along the south shore of the eastern Republica Dominicana. With
SCHWARTZ AND KLINIKOWSKI : AMEIVA -483
the closure of the strait, each of these races has expanded into
the resulting xeric plain, although to the northwest along the
Golfe de la Gonave, chrysolaema still occupies the narrow coastal
plain and adjacent xeric foothills much as it may formerly have
done farther south. Once across the plain, chrysolaema has ex-
tended its range inland to some extent (Petionville), and to the
west (Pere, Momance). Vmhratilis, on the other hand, has not
been able to penetrate far into the adjacent mountains, although
it does reach an elevation of 1000 feet near El Naranjo. The
range of nnihratilis is bounded on the north by the valley of the
Rio Yaque del Sur.
To the east of the Rio Yaque del Sur are a series of four, more
©r less coastal, races. Of these, hoekeri, immediately to the north
and east of the Yaque, is rather like umbratilis in its patterned
phase. We consider hockcri as a direct derivative of umhratilis
and restricted to the Llanos de Azua. The next three races —
procax, parvoris, jacta — show increasingly scattered patterns
of distribution to the east, with jacta apparently the most iso-
lated. As noted previously, procax and hoekeri approach one
another in the vicinity of Bani, precisely in the area of rapid
transition from the xeric Llanos de Azua to the more mesic
coastal areas to the east. The Rio Ozama separates procax from
parvoris, which is known from only two localities on the main-
land and from a slightly differentiated population on Isla Cata-
lina. We consider hoekeri, procax, and parvoris as a more or less
sequential coastal series still maintaining its integrity in response
to environmental and geographical influences.
Jacta, on the other hand, along with richardthomasi, represents
a very different sort of lizard. We feel that the jacta-richard-
thomasi populations at one time (and perhaps still) occupied
most of the extreme eastern end of the island. The presently
restricted and scattered distribution and records for not only
jacta but also parvoris, as well as the isolated occurrence of
parvoris on Catalina, suggest strongly that the range of A.
chrysolaema in this region is retracting, leaving isolated outliers
which may be indicative of former populations. The absence of
records of the species between San Pedro de Macoris and Juan-
illo, as well as only two general localities of parvoris on the
mainland, add substance to this supposition. The eastern dis-
tribution of Ameiva lineolata tends also to bear out this conten-
tion.
484 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Of the remaining- races from the southern part of the north
island, only the strii)ed alacris is left. Alacris is so like procax
in many features that it is difficult not to associate the two. On
the other hand, they are separated presently by hoekeri, with
which race alacris intergrades, as it does with chrijsolaema on
the west. Tt is possible that alacris and procax were at one time
confluent, but that bockcri has pushed between them, thereby
severing- any direct genetic continuity between the two. An-
other possibility is that alacris has been derived from chryso-
laema, either across the Montagues de Trou d'Eau or along the
valley of the Riviere de I'Artibonite. Continued evidence of
intergradation between these two races near ]Mirebalais lends
support to this possibility.
A. c. defensor on the northwestern peninsula has obviously
been long isolated from its more southern relatives. Presumably
it has been derived from chrysolaema. The Gonave race secessa
resembles defensor in pattern and cohn-ation, and it is possible
that Gonave was colonized from tlie north (defensor) rather than
from the adjacent mainland [chrysolaema) . The latter, however,
seems more likely both on the basis of proximity and what is
presently known of the origin of the Gonave herpetofauna.
The relationships of regularis seem closest to the procax-alacris
pair; possibility of origin of regularis from alacris via the Arti-
bonite system and thence to the Grande Riviere du Xord has
already been discussed. Other possibilities are an old origin
from defensor (although this is not particularly appealing) or
an origin from procax via the central valley to the east of the
Cordillera Central in the Republica Dominicana. The only evi-
dence against this is that A. chrysolaema does not occupy this val-
ley today, although procax occurs at its southern end and
regularis at its northern end uear Santiago.
Finally, A. c. woodi on Tortue, although closer geographically
to defensor than to regularis, seems closer in some characteristics
to regularis than to defensor. There are, however, tendencies of
pattern in defensor which herald the extreme peculiarities of
woodi patterns. On the other hand, occasional regularis have
dorsal patterns like some woodi. It seems more likely that ivoodi
is a direct derivative of defensor.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 485
LITERATURE CITED
Barbour, Thomas, and G. Kingley Noble
1915. A revision of the lizards of the genus Ameiva. Bull. Mus. Comp.
Zool., 59(6):417-479.
Cochran, Doris M.
1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177:
i-vii + 1-398, 12 pis., 120 figs.
Maerz, a., and M. Rea Paul
1950. A dictionary of color. New York, McGraw-Hill Book Co.,
pp. i-vii, 1-23, 137-208, 56 pis.
Mertens, Robert
1938. Aniphiliien und Reptilien aus Santo Domingo, gesanimelt von
Dr. H. Boker. Senckenbergiana, 20: 332-42, 6 pis.
1939. Herpetologische Ergebnisse einer Reise naeh der Insel Hispan-
iola, Westindien. Abh. senckenberg. naturf. Ges., 449: 1-84,
10 pis.
Schwartz, Albert
(in press) The Ameiva (Reptilia, Teiidae) of Hispaniola. I. Ameiva
lineolata Dumeril and Bibron. Caribbean Jour. Sci.
(Received February 25, 1965.)
486
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Subspecies
chrysolaema
woodi
procax
richardthomasi
parvoris
secessa
jacta
regularis
umbratilis
boeheri
alacris
defensor
ficta
abbotti
leberi
160
141
141
137
137
135
134
132
130
126
126
126
121
117
111
130
126
116
124
113
111
128
112
111
109
106
113
108
104
Table 1. Subspecies of Ameiva chrysolaema ranked according to snout-
vent length (in mm) of largest male for each race.
Transverse ventrals
Subspecies
N
Mean and extremes
(mo(
secessa
50
91.0
(80-99)
12
abbotti
32
89.6
(84-97)
12
richardthomasi
22
87.8
(80-92)
10
chrysolaema
106
86.7
(76-101)
12
leberi
48
85.5
(78-96)
10
alacris
48
84.8
(77-94)
10
procax
51
84.8
(76-98)
10
boelceri
86
84.6
(73-98)
10
parvoris
42
83.2
(76-92)
12
umbratilis
61
83.0
(73-100)
10
defensor
24
82.4
(77-90)
10
ficta
41
81.9
(75-92)
10
woodi
39
80.5
(66-93)
10
jacta
3
79.7
(77-82)
12
regularis
140
77.8
(67-88)
12
Table 2. Subspecies of Ameiva chrysolaema ranked according to means of
number of fourth-toe subdigital scales; each race is also characterized by
the modal number of transverse rows of ventral scales (but see discussions
of umbratilis, procax, and richardthomasi). N ^ number of specimens ex-
amined.
SCHWARTZ AND KLINIKOWSKI : AMEIVA 487
Subspecies
abbotti
chrysolaema
richardtJiomasi
seoessa
jacta
leberi
woodi
ficta
parvoris
regularis
defensor
boeTceri
procax
umbratilis
alacris
Table 3. Subspecies of Ameiva chrysolaema ranked according to mean
number of femoral pores; N = same as in Table 2.
Femoral pores
Mean and extremes
43.8
(35-52)
43.7
(33-50)
42.6
(39-47)
41.5
(36-46)
41.3
(39-43)
41.3
(35-45)
41.3
(36-46)
40.9
(33-47)
38.2
(24-45)
37.6
(32-44)
37.2
(30-41)
36.5
(31-41)
36.1
(30-43)
35.6
(28-42)
33.8
(30-39)
Scales in 15th caudal verticil
Subspecies Mean and extremes
chrysolaema
alacris
regularis
procax
boeTceri
umbratilis
parvoris
abbotti
secessa
ficta
woodi
richardthomasi
defensor
leberi
jacta
Table 4. Subspecies of Ameiva chrysolaema ranked according to mean num-
ber of scales in fifteenth caudal verticil ; N = same as in Table 2.
44.4
(37-51)
44.0
(39-49)
43.1
(30-52)
42.8
(39-48)
42.7
(37-48)
42.7
(38-48)
42.7
(38-46)
40.4
(36-47)
40.1
(36-44)
38.8
(36-46)
38.8
(34-43)
38.6
(36-41)
38.0
(34-41)
37.9
(34-43)
35.7
(35-36)
Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 11
*N
A NEW ATTEMPT TO CONSTRUCT LIFE TABLES FOR
KENT ISLAND HERRING GULLS
By
Raymond A. Paynter, Jr.
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
May 18, 1966
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Bulletin of the Museum of Comparative Zoology
HARVARD UNIVERSITY
Vol. 133, No. 11
A NEW ATTEMPT TO CONSTRUCT LIFE TABLES FOR
KENT ISLAND HERRING GULLS
By
Raymond A. Paynter, Jr.
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
May, 1966
Bull. Mus. Comp. ZooL, Harvard Univ., 133 (11): 489-528, May, 1966.
No. 11 — A New Attempt to Construct Life Tables for Kent Island
Herring Gulls'^
By
Raymond A. Paynteb, Jr.
INTRODUCTION
An earlier attempt (Paynter, 1949) to construct a life table
for the population of Herring Gulls {Larus argentatus sniith-
sonianus) on Kent Island, New Brunswick, yielded results which
seemed in conflict with the observed status of the colony. The
life table indicated that the population was in a steep decline,
although the size of the colony was thought to be constant or
possibly even producing a population surplus which was con-
tributing to the general increase in Herring Gulls that had been
noted in northeastern North America for nearly half a century.
To account for the discrepancy between the life table and what
was believed to be the actual status of the population, it was
suggested that, (1) the 1935 class (erroneously cited as "1936"),
which was utilized to construct the postfledging portion of the
table, had been banded too short a time (11 years) to yield all
potential recoveries, thereby producing a truncated table; (2)
that the 1935 year class may have suffered unusually severe post-
fledging mortality or that the 1947 year class, which provided
data for the egg and nestling portion of the table, may also have
been unrepresentative, or possibly both situations had prevailed ;
(3) that there may have been a loss of bands, particularly among
older birds, that caused the calculated survival rate and life ex-
pectancy to be lowered.
Fifteen years have elapsed since that study. Additional birds
from the 1935 class have been recovered and it is now possible
to test the hypothesis that older recoveries might be sufficient to
raise the calculated life expectancy and survival rates to levels
commensurate with the presumed status of the population. Re-
coveries from bandings in other years at Kent Island have also
accumulated, allowing comparisons between several year classes.
Finally, there have been published three additional life tables
for the species (Paludan, 1951; Hickey, 1952; Olsson, 1958) and
these provide valuable comparative data. It is the purpose of this
1 Contribution No. 32 from the Bowdoin Scientific Station, Kent Island, Grand
Munau, Xew Brunswick, Canada.
492 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
paper to re-examine the dynamics of the Kent Island Herring
Gull population in the light of these developments.
MATERIAL AND METHODS
In calculating ages of banded gulls it has been the custom at
Kent Island (e.g., Paynter, 1947; 1949) to begin the year on
July first, which is about the earliest chicks are large enough
to band. To avoid the inclusion of prefledging mortality in the
banding recoveries, bands were removed from any chicks dying
before fledging and were placed on other young.
In some banding studies it is necessary to make adjustments
for bias resulting from the disproportionate recovery of newly
fledged birds in the vicinity of their place of birth (see Earner,
1955). This correction is not required of the Kent Island data
because the gulls leave the island shortly after they are able to
fly and disperse over a wide area (Gross, 1940) where, presum-
ably, they are no more likely to be recovered than older birds.
Banding began at Kent Island in 1934. The terminal date for
data used in this study is 30 June 1963. Thus, the oldest poten-
tial recovery would be a bird in its twenty-eighth year. To date
the oldest record of a gull banded as a fledgling is an individual
caught at Kent Island in its twenty-sixth year and released bear-
ing a new band. The oldest record of a bird dying while still
wearing its original band is a gull in its twenty-fourth year ; the
next oldest is a twenty-second year bird. For purposes of this
study only young banded in the six year classes from 1934
through 1939 will be considered. The maximum potential age for
birds in the most recently banded year class is twenty -three years,
which seems close to the maximum age of recovery that may be
expected in the Kent Island population (see p. 514). A few birds
older than twenty-six almost certainly will be found in future
years, but thej^ probably play an insignificant part in the dy-
namics of this population.
Heeding Hickey's warnings (1952) of clerical and other errors
in the banding records kept by the Fish and Wildlife Service,
microfilms of all Kent Island banding schedules and recovery
records were obtained through the courtesy of Allen J. Duvall.
These were carefully compared with the files at Kent Island and
all erroneous and questionable data were eliminated. I have no
illusions about the complete accuracy of the resulting material,
but considering the fact that hundreds of people have been in-
volved in reporting and handling these data during three decades,
3,646
125
3.43
10,748
352
3.27
6,665
254
3.81
4,652
146
3.14
2,983
77
2.58
3,000
145
4.83
HERRING GULL LIFE TABLES 493
no further refinements seem possible. It is believed that the quan-
tit}^ of data is sufficient to offset whatever deficiencies in quality
remain.
TABLE I
Kent Island Fledglings Banded and Admissible i Eecoveries
Year Banded Eecoveries Per cent
1934
1935
1936
1937
1938
1939
Total 31,694 1,099 3.47
1 See below for deflnition of admissible recoveries.
From 1934 through 1939 nearly 32,000 fledgling gulls were
banded on Kent Island (Table I). Discrepancies between the
yearly totals in Table I and those published previously (Paynter,
1947) result from the re-analysis of the Fish and Wildlife Serv-
ice records.
These 32.000 bandings have yielded 1,206 recoveries and re-
turns. It is necessary, however, to remove from consideration
records in certain categories. Because the Kent Island life table
is to be constructed from a mortality series, no records of living
birds ("returns") are admissible. As a consequence, those in-
dividuals which were trapped in later years at Kent Island and
found bearing bands are eliminated, as are those birds which
were reported to the Fish and Wildlife Service as having been
"captured and released." On the grounds of uncertainty,
whether the banded bird was living or dead, Hickey (1952, p. 94)
probably would exclude those records reporting "no informa-
tion," "observed," "found ill," "caught by fisherman," etc.
Some of these reports doubtless pertain to gulls not yet dead
when their bands were read, but it seems unlikely that many of
these records could have been obtained from birds that were not
already weakened from illness or injury and soon to die. For
this reason I include these records in the mortality series.
Also eliminated from the study are gulls collected in connec-
tion with research at Kent Island and those shot elsewhere under
494 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
gull control permits. Neither of these hazards operated con-
sistently from year to year.
Lack (1954, p. 91) notes that the inclusion of birds which have
been recovered by means of shooting (i.e. "normal" shooting —
not systematic control) may introduce bias if this category repre-
sents a substantial portion of the recovery sample and if juve-
niles are more easily shot than adults. In certain instances this
may be correct, but if a given cohort is hunted with equal inten-
sity throughout its life span, shooting must be considered to be
merely another one of the hazards acting upon a population, such
as the stress of weather and the toll of predators. The fact that
young birds may be more easily shot than older ones does not
bias the sample any more than does the fact that adverse weather
or predators may claim more young than adults. On the other
hand, banded birds which are shot, or for that matter killed
tlirough any human agency, probably are more likely to be re-
ported than birds dying of disease or other natural causes. Hu-
man activity, therefore, may appear as a disproportionately high
cause of death in the sample of recoveries and may also increase
the total number of recoveries. But, as long as the human activity
causing death operates consistently throughout the life of a co-
hort, the recoveries need not be excluded from the mortality
series used in preparing a life table.
About five per cent of all Kent Island recoveries are of indi-
viduals reported as shot ; the true rate is probably higher but,
because the species is protected by law, is concealed within the
reports as "found," "no information," etc. The actual percent-
age is certainlj^ not nearly so high as in Europe, where the
species is unprotected and where about 60 per cent of all reports
are of gulls which have been shot (Paludan, 1951 ; Olsson, 1958).
There is no reason to suspect that the distribution by age or year
class of recoveries of Kent Island gulls shot but reported in other
categories differs from that of birds accurately recorded as shot.
Taking the shooting reports at face value, we find considerable
variation in the percentage of these recoveries within various age
classes, as well as within year classes. For example, in the 1935
cohort, about 6.5 per cent of the gulls were reported as shot ;
there were no records of birds shot beyond their seventh year and
the percentages of reports from the first year through the seventh
year are 8.4, 2.9, 6.5, 4.1, 16.0, 0, and 5.2. In contrast, in the
1936 year class, only 3.6 per cent of the records are of birds
shot ; the oldest was in its twelfth year and the percentages by
age classes run 4.8, 3.0, 0, 0, 5.2, 0, 0, 0, 0, 50.0, 0, and 50.0. The
HERRING GULL LIFE TABLES 495
distribution of shooting reports illustrated here is similar in all
six cohorts. It is of interest that while shooting occurs only in
the first half of the maximum potential life span (there are no
records beyond the thirteenth year), there is no clear-cut pattern
within this period. One might have expected to find shooting
relatively more common among immature (and inexperienced)
birds.
Birds recovered after having been rebanded are excluded. If
Kent Island gulls lose bands, which they doubtless do, and if the
loss is a function of the age of the bands rather than a random
occurrence throughout the life span of the birds, this may be evi-
dent in the life table, provided it is not obscured by other phe-
nomena, as a declining survival rate. The inclusion of rebanded
birds in the study, while possibly giving a truer picture of age-
specific survival rates, would introduce another source of bias.
Unfortunately, there has been an insufficient number of reband-
ings to allow an analysis of birds in this category.
The Kent Island life tables are begun with the laying of the
first egg, which occurs in late May, roughly one-tenth of a year
earlier than the July 1 date taken as the start of the year when
calculating the ages of banded birds. Because there were no
studies of the survival of the eggs and chicks at Kent Island
during the period 1934-1939, the records of the 1947 year class
(Paynter, 1949) are used, creating composite tables. The mean
incubation period is taken at 28 days, or 0.07 years. In 1947 it
was found that 28.6 per cent of the eggs failed to hatch ; the
mean age of the young at fledging was calculated to be 43 days
(0.12 years), and 48.5 per cent of the chicks are believed to have
failed to fledge, giving a production of 0.92 fledglings per breed-
ing pair.
KENT ISLAND LIFE TABLES
Composite life tables, using the 1947 prefledging data, for the
six-year classes from 1934 through 1939, as well as a table for
the combined year classes, are presented in Table II. Semi-loga-
rithmic survivorship curves (Ix) are shown in Figures 1 and 2.
The table and graph for the combined six-year classes (Table
II; Fig. 2) indicate that about 63 per cent of the eggs laid fail
to produce fledged young (see Paynter, 1949, for discussion).
Then there is heavy mortality (45 per cent) from the time of
fledging until the following July 1, a somewhat lessened rate for
the second year of life (35 per cent), and finally a lower average
496 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
rate for the bulk of the remaining life span (26.8 per cent for
the years 3 through 15). This is the expected pattern for most
long-lived avian populations. It suggests that young (post-
fledging) gulls are less able to cope with the hazards of their
environment and that birds surviving the first two years of life
have either learned to avoid more of the hazards, or the less adept
ones have been eliminated through natural selection. If data for
the oldest age groups were more abundant one might find the
mortality rate increasing because of the senility. On the other
hand, it is unlikely many animals survive this long in nature and
that we shall ever have sufficient data from senile individuals to
document this phenomenon, if indeed it does occur.
Examined individually, the life tables and survivorship curves
(Table II; Fig. 1) show interesting similarities as well as dif-
ferences, which one expects of natural populations in a variable
environment. For example, in four of the six cohorts the mor-
tality rate declined the year following fledging, but in the 1936
and 1938 year classes it rose. The first year postfledging mor-
tality rate for the 1936 cohort was 36.2 per cent, which is un-
usually low and as a result was slightly below the rate of the
second year (39.5 per cent), which was only a little in excess of
the average (35 per cent). The same pattern was displayed, in a
more exaggerated manner, in the 1938 cohort. The mortality rate
in the first year was a comparatively low 37.7 per cent, but in
the second year it jumped to 52.1 per cent, the highest observed.
Similar yearly fluctuations occur throughout the life tables. An-
nual variations in the severity of the weather are presumably
responsible for many of these swings, but the problem has not
been studied.
Next to be considered is the reproductive potentiality of the
colony as indicated by the life tables. North American gulls are
assumed to breed in their fourth year (year 3-4) or, in other
words, three summers after hatching. This may not be an exact
assumption for Drost, Focke, and Freytag (1961) found in a
German colony of European Herring Gulls {L. a. argentatus)
that 20 per cent bred in their third year, 25 per cent in their
fourth year, and 55 per cent in their fifth year, with the mean
breeding age slightly in excess of four years. Paludan (1951),
however, concluded that Danish gulls regularly breed in their
third 3^ear, basing this on a single observation of a bird of this
age in a breeding colony. The gull had some dark areas on the
tail and on the lesser wing coverts. From this Paludan assumed
that fully adult plumage is attained in the third year and that
HERRING GULL LIFE TABLES 497
TABLE II
Composite Life Tables for Kent Island Cohorts 1934-1939, Utilizing 1947
Prefiedging Mortality Data of Paynter (1949)
1934 Cohort
X
1
d
l,000q^
e„
X
X
X
X
Age 1
^0. surviving to
No. dying in
Mortality rate
Expectation of life
in
start of age
age interval
per 1,000 alive
remaining to those
years
interval out of
out of 1,000
at start of age
attaining the age
1,000 eggs laid
eggs laid
interval
interval (in years)
0.00-0.07
1,000.0
286.0
286.0
.97
0.07-0.19
714.0
346.3
485.0
1,28
0.19-1.10
367.7
182.4
496.1
2.31
1.10-2.10
185.3
44.1
238.0
3,23
2.10-3.10
141.2
26.5
187.7
3.09
3.10-4.10
114.7
20.1
175.2
2.68
4.10-5.10
94.6
38.4
405.9
2.15
5,10-6.10
56.2
23.7
421,7
2.28
6.10-7.10
32.5
11.8
363.1
2.57
7.10-8.10
20.7
8.9
429.9
2,76
8.10-9.10
11.8
0
0
3,47
9.10-10.10
11.8
3.0
254.2
2,47
10.10-11.10
8.8
3.0
340.9
2,14
11.10-12.10
5.8
0
0
1,98
12.10-13.10
5.8
2,9
500.0
,98
13.10-14.10
2.9
2.9
1,000.0
.50
1935 Cohort
0.00-0.07
1,000.0
286.0
286.0
.94
0.07-0.19
714.0
346.3
485.0
1,23
0.19-1.10
367.7
176.5
480.0
2.21
1.10-2.10
191.2
70.0
366.1
2.92
2.10-3.10
121.2
30.3
250,0
3.33
3.10-4.10
90.9
23.0
253.0
3.27
4.10-5.10
67.9
20.9
307.8
3.21
5.10-6.10
47.0
13.6
289.4
3.41
6.10-7.10
33.4
10.5
314.4
3.60
7.10-8.10
22.9
4.2
183.4
4.02
8.10-9,10
18.7
4.2
224.6
3.81
9.10-10.10
14.5
2.1
144.8
3.76
10.10-11.10
12.4
2.1
169.3
3.32
11.10-12.10
10.3
4.1
398.1
2.90
12.10-13.10
6.2
1.0
161.3
3.50
13.10-14.10
5.2
2.1
403.8
3.08
14.10-15.10
3.1
1.0
322.6
3.84
15.10-16.10
2.1
0
0
4.43
16.10-17.10
2.1
0
0
3.38
17.10-18.10
2.1
1.1
523.8
2.38
18.10-19.10
1.0
0
0
3.50
19.10-20.10
1.0
0
0
2.50
20.10-21.10
1.0
0
0
1,50
21.10-22.10
1.0
1.0
1,000,0
.50
498 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
1936 Cohort
X
'x
d
X
1,000 q^
\
0.00-0.07
1,000.0
286.0
286.0
1.04
0.07-0.19
714.0
346.3
485.0
1.38
0.19-1.10
367.7
133.1
362.0
2.50
1.1O-2.10
234.6
92.7
395.1
2.75
2.10-3.10
141.9
39.2
276.2
3.23
3.10-4.10
102.7
23.1
224.9
3.27
4.10-5.10
79.6
27.5
345.5
3.07
5.10-6.10
52.1
18.8
360.8
3.42
6.10-7.10
33.3
8.7
261.3
4.08
7.10-8.10
24.6
4.4
178.9
4.35
8.1O-9.10
20.2
5.8
287.1
4.19
9.10-10.10
14.4
2.9
201.4
4.68
10.10-11.10
11.5
1.4
121.7
4.74
11.10-12.10
10.1
2.9
287.1
4.33
12.10-13.10
7.2
1.4
199.4
4.87
13.10-14.10
5.8
1.5
258.6
4.93
14.10-15.10
4.3
0
0
5.49
15.10-16.10
4.3
1.4
325.6
4.56
16.10-17.10
2.9
0
0
5.41
17.10-18.10
2.9
0
0
4.41
18.10-19.10
2.9
0
0
3.41
19.10-20.10
2.9
1.5
517.2
2.41
20.10-21.10
1.4
0
0
3.50
21.10-22.10
1.4
0
0
2.50
22.10-23.10
1.4
0
0
1.50
23.10-24.10
1.4
1.4
1,000.0
.50
1937 Cohort
X
'x
''x
1,000 q^
%
0.00-0.07
1,000.0
286.0
286.0
.85
0.07-0.19
714.0
346.3
485.0
1.10
0.19-1.10
367.7
176.5
480.0
1.96
1.10-2.10
191.2
52.8
276.1
2.45
2.10-3. 10
138.4
37.8
273.1
2.19
3. 10-4. 10
100.6
22.6
224.6
1.83
4.10-5.10
78.0
30.2
387.2
2.24
5.10-6.10
47.8
20.1
420.5
2.34
6.10-7.10
27.7
5.0
180.5
2.68
7.10-8.10
22.7
10.1
445.0
2.16
8.10-9.10
12.6
0
0
2.50
9.10-10.10
12.6
5.0
396.8
1.50
10.10-11.10
7.6
2.5
328.9
1.16
11.10-12.10
5.1
5.1
1,000.0
.49
HERRING GULL LIFE TABLES 499
1938 Cohort
X
1
d
1,000 q
e
X
X
' ^x
x
0.00-0.07
1,000.0
286.0
286.0
,89
0.07-0.19
714.0
346.3
485.0
1.16
0.19-1.10
367.7
138.6
376.9
2.09
1.10-2.10
229.1
119.4
521.2
2.16
2.10-3.10
109.7
33.4
304.5
2.97
3.10-4.10
76.3
14.3
190.0
3.06
4.10-5.10
62.0
9.6
154.8
2.65
5.10-6.10
52.4
19.1
364.5
2.04
6.10-7.10
33.3
9.5
285.3
1.92
7.10-8. 10
23.8
9.5
399.2
1.50
8.10-9.10
14.3
4.8
335.7
rj6
9.10-10.10
9.5
9.5
1,000.0
.49
1939 Cohort
x
'x
"'x
1,000 q^
\
0.00-0.07
1,000.0
286.0
286.0
1.04
0.07-0.19
714.0
346.3
485.0
1.38
0.19-1.10
367.7
182.8
497.1
2.49
1.10-2.10
184.9
71.0
384.0
3.60
2.10-3.10
113.9
32.0
280.9
4.54
3.10-4.10
81.0
10.1
124.7
5.18
4.10-5.10
70.9
12.7
179.1
4.85
5.10-6.10
58.2
7.6
130.6
4.79
6.10-7.10
50.6
7.6
150.2
4.44
7.10-8. 10
43.0
7.6
176.7
4.13
8.10-9.10
35.4
2.5
70.6
3.91
9.10-10.10
32.9
7.6
231.0
3.18
10.10-11.10
25.3
5.1
201.6
2.98
11.10-12.10
20.2
7.6
376.2
2.61
12.10-13.10
12.6
0
0
2.88
13.10-14.10
12.6
7.6
603.1
1.88
14.10-15.10
5.0
2.5
500.0
2.98
15.10-16.10
2.5
0
0
4.48
16.10-17.10
2.5
0
0
3.48
17.10-18.10
2.5
0
0
2.48
18.10-19.10
2.5
0
0
1.48
19.10-20.10
2.5
2.5
1,000.0
.48
500 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Combined 1934-1939 Cohorts
X
'x
"x
1/000 q^
^
0.0O-0.07
1,000.0
286.0
286.0
.95
0.07-0.19
714.0
346.3
485.0
1.25
0.19-1.10
367.7
165.5
450.1
2.26
1.10-2.10
202.2
71.7
354.6
2.82
2.10-3. 10
128.5
33.4
260.0
3.27
3.10-4.10
95.1
20.4
214.5
3.30
4.1O-5.10
74.7
23.7
317.3
3.07
5.10-6.10
51.0
15.3
300.0
3.26
6.10-7.10
34.7
9.0
259.4
3.56
7.10-8.10
25.7
6.4
249.0
3.63
8.10-9.10
19.3
3.3
171.0
3.67
9.10-10.10
16.0
4.0
250.0
3.33
10.10-11.10
12.0
2.4
200.0
3.27
11.10-12.10
9.6
3.6
375.0
2.95
12.10-13.10
6.0
1.0
166.7
3.43
13.10-14.10
5.0
2.3
460.0
3.02
14.10-15.10
2.7
.7
259.3
4.18
15.10-16.10
2.0
.3
150.0
4.50
16.10-17.10
1.7
0
0
4.23
17.10-18.10
1.7
.3
176.5
3.23
18.10-19.10
1.4
0
0
2.86
19.10-20.10
1.4
.7
121.4
1.86
20.10-21.10
.7
0
0
2.29
21.10-22.10
.7
.4
571.4
1.28
22.10-23.10
.3
0
0
1.33
23.10-24.10
.3
.3
1,000.0
.33
the bird was merely retarded. This appears to be an unneces-
sarily complicated interpretation. It would seem more logical to
conclude that this was a case of an early breeder. Lacking data
for the Kent Island gulls, we shall accept the fourth year as the
mean breeding age.
In 1947 the average clutch size at Kent Island was nearly 2.5
eggs (Paynter, 1949). Observations (unpublished) in 1948 indi-
cated a somewhat higher mean for first clutches (ea. 2.75) and a
lower mean (ca. 2.00) for clutches replacing those lost through
predation, but the average for all final clutches was again close
to 2.5 eggs.
Taking the life table for the combined years 1934-1939, we find
that from an initial cohort of 1,000 eggs 95.1 birds survive to
July 1 of the third summer after hatching. It will be recalled
that July 1 is the time when banding of the chicks is first practi-
cal and that egg-lajdng begins about one-tenth of a year earlier,
HERRING GULL LIFE TABLES 501
in late May. We must, therefore, take into account birds which
are alive at the time of laying but which do not survive until
July 1. From the life table it is seen that 33.4 birds die during
the third year. If the mortality rate is constant throughout the
year, one-tenth, or 3.3 birds, die between the time of laying and
the first of July. These individuals may then be added to those
alive on July 1, giving a total of 98.4 birds which survive to
breed for the first time from 1,000 eggs laid four summers earlier.
If the sex ratio is equal, and there is no evidence that it is not,
there is an average of 49.2 pairs alive at the start of the first
breeding season. At this point these birds have a mean life ex-
pectancy of about 3.3 years. If they breed annually, which they
proba])h'- do for the greater part of their life spans, and lay a
yearly average of 2.5 eggs per clutch, the 49.2 pairs are capable
of producing 405.9 eggs. This is only 41 per cent of the number
required to maintain a stable population.
The only year class with a life table differing materially from
the combined 1934-1939 table is the 1939 cohort, with its gen-
erally better survival rate among older birds. The calculated
number of birds at the start of the first breeding season is 84.2.
The mean life expectancy is about 5.2 years. This would allow
for a lifetime production of approximately 547 eggs, which again
is still far short of the 1,000 eggs needed for a stable population.
If these life tables accurately represent the dynamics of the
Kent Island population, it is obvious that the size of the colony
should have been rapidly declining in the 1930 's and early
1940 's, and that the magnitude of the decline would be so great
that it would be apparent to field observers. Only constant, large-
scale immigration could obscure this phenomenon. TTnfortu-
nately, the colony was not accurately censused until 1940 (Crys-
tal, 1941), when approximately 12,000 nests were counted, and
the presence of 3,000 more was estimated. Allowing for the
inclusion of "play nests" (Goethe, 1937), there must have been
a breeding population well in excess of 20,000 birds. During my
field Avork from 1946 through 1948 the colony was densely popu-
lated and I estimated it to contain 25,000 to 30,000 individuals
(Paynter, 1949). The late Ernest Joy informed me that he had
noticed no change in the size of the colony during his tenure as
warden from 1935 to 1948. Thus, we must conclude that the
Kent Island population was stable, or at least not noticeably un-
stable, from at least 1935 through 1948. The possibility remains
that the colony was failing to reproduce itself and was dependent
502
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
1,000
<
to
o
LLl
Q.
o
>
>
=>
to
I I I I I I ' I I I ' I ' I ■ I I
I.I 3.1 5.1 7.1 9.1 11,1 13.1 15,1 17.1 19.1 21.1 23.
AGE IN YEARS
Fig. 1. Survivorship curves for Kent Island cohorts 1934-39.
HERRING GULL LIFE TABLES
503
1,000 .
Kent Id., 1934-'39
Fenno-Scandia, l925-'54
N. Amer., l925-'30
Denmark, 19l7-'42
100 _
Z3
o
UJ
a.
CO
o
>
1 _l
AGE IN YEARS
Fig. 2. Survivorship curves for four populations.
504 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
on immigrants to maintain a constant size, but this seems im-
probable. The Herring Gull has been increasing along the east-
ern seaboard, as well as in Europe, since the turn of the century.
It would appear unlikely that the Kent Island colony was not
contributing to this general increase. A more reasonable con-
clusion is that the six life tables are deficient. Before speculating
on the possible reasons for this, life tables for other populations
of the species will be examined.
OTHER LIFE TABLES
Hickey (1952) has considered the first three life tables pre-
pared for North American Herring Gulls (viz., Marshall, 1947 ;
Paynter, 1947, 1949), commenting on their errors and discrepan-
cies; they need not be reconsidered here. Then using recoveries
of gulls banded at various colonies in North America from 1925
through 1930, Hiekej^ constructed an abridged life table for a
TABLE III
Life Table for Theoretical Population of North American Gulls Banded
1925-1930 (504 Eecoveries) (after Hickey, 1952)
X
1
X
d
X
1,000 q
X
e
X
0-1
1,000.0
599.2
599.2
2.00
1-2
400.8
117.1
292.1
3.25
2-3
283.7
59.5
209.7
3.39
3-4
224.2
61.5
274.3
3.15
4-5
162.7
41.7
256.2
3.16
5-6
121.0
45.6
376.8
3.08
6-7
75.4
16.1
213.5
3.63
7-8
59.3
7.9
133.2
3.48
8-9
51.4
11.9
231.5
2.95
9-10
39.5
9.9
250.6
2.69
10-11
29.6
5.9
199.3
2.42
11-12
23.7
9.9
417.7
1.90
12-13
13.8
4.0
289.8
1.91
13-14
9.8
2.0
204.1
1.48
14-15
7.8
5.9
756.4
.73
15-16
1.9
1.9
1,000.0
.47
HERRING GULL LIFE TABLES 505
theoretical population of 504 birds, beginning the table with re-
ports received subsequent to 31 August of the year in which the
gulls were hatched. No reports were available for gulls older
than their sixteenth year. Although the table is probably fore-
shortened, recoveries beyond the sixteenth year are so infrequent
that the resulting table is doubtless close to what would have
been found if it had not been terminated until all potential re-
coveries had been obtained. Adjustments were made for yearly
variations in the number of birds banded and for the fact that
some birds had not been banded sufficiently long to yield recov-
eries in the year 15-16 (see Hickey, 1952, p. 11). This life table,
recalculated to form a cohort of 1,000, is presented in Table III,
and the survivorship curve is plotted in Figure 2.
The first year mortality rate is about 60 per cent, in contrast
to an average of 45 per cent at Kent Island from the time of
fledging in August to the following June 30, and the second year
mortality is 29 per cent, compared to the Kent Island figure of
35.5 per cent. The mean annual mortality rate for the next ten
years is 25.6 per cent, which is almost exactly that at Kent
Island (Table IV).
Because this table is begun in September, neglecting the egg,
nestling, and earliest postfledging mortality, a different tech-
nique from that used with the Kent Island data must be em-
ployed to assess the reproductive potential of the population.
TABLE IV
Mean Mortality Kates
Age
Population
0-1
1-2
2-12
Kent Id., 1934- '39
45.01
35.5
26.0
North Anier. (Hickey, 1952)
59.9^
29.2
25.6
Denmark (Paludan, 1951)
62.32
21.7
15.6
Fenno-Scandia (Olsson, 1958)
56.8^
31.9
33.8
iFirst interval 0.9 year, i.e. fledging to 30 June following year; 1 July to 30
June thereafter.
^Year begins 1 September.
3 Year begins 1 August.
506 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
This is done by totaling the l^ column (survivors) from the
year 3-4 (first breeding year) to the end of the table, dividing
the total by two to give the number of pairs of breeding birds,
and, finally, dividing the initial cohort (1,000) by the number
of pairs of breeding birds (see Hickey, 1952, pp. 94-95). The
result is the average number of young per pair of adults which
need be raised to the first of September of the year of hatching in
order to maintain a stable population. Applying this to Hickey 's
table, we find that an average of 2.44 young must be fledged and
survive to September 1. This is only slightly below the mean
clutch size found at Kent Island (2.5) and, therefore, allows
almost no egg or chick mortality. Even if the mean clutch is
assumed to be three, which is the maximum number of eggs laid
by Herring Gulls, except for an exceedingly rare clutch of four
(about 0.6 per cent of all clutches [Paludan, 1951, p. 49]), this
would also allow for but 19 per cent mortality between the laying
of the first egg and September 1. Such a mortality rate seems
much too low in the light of observations at Kent Island, where
more than 60 per cent of the eggs failed to yield fledged young.
This life table cannot, therefore, be accepted.
The first life table for a European population of Herring
Gulls is that constructed by Paludan (1951) for 966 recoveries,
including those shot (ca. 60 per cent), from 11,689 birds banded
in Denmark from 1917 through 1942 (Table V). The oldest recov-
ery is that of an individual in its twenty-sixth year ; the oldest po-
tential recovery at the time the table was compiled would have
been a thirty-fourth year bird. Adjustments, similar to those of
Hickey (1952), w^ere made to compensate for the fact that only 8
of the 26 year classes had been banded sufficiently long to yield
recoveries in the twenty-sixth year. The mean recovery rate for
these eight cohorts was 8.54 per cent, versus only 3.47 per cent
for the six cohorts at Kent Island. The marked difference in re-
coveries is doubtless due to the frequency with which Danish
gulls are shot, which enhances the chances that a banded bird will
be reported, and also probably due to a better retention of
bands, which will be considered subsequently. From this table
(Table V), which starts on September 1 following hatching, it
is seen that the first year mortality is 62.3 per cent, closely ap-
proximating Hickey 's finding in North America. The second
year mortality drops to 21.7 per cent, which is considerably lower
than that in Hickey 's table (29.2 per cent) or the Kent Island
table (35.5 per cent). The mean annual mortality for the next
ten years is only 15.6 per cent, in contrast to the North American
HERRING GULL LIFE TABLES 507
TABLE V
Life Table for Danish Gulls Banded 1917-1942 (966 Eecoveries) (Paludan,
1951)
X
1
d
1,000 q
e
X
X
X
X
0-1
1,000
623
623
2.72
1-2
377
82
217
5.39
2-3
295
56
190
5.75
3-4
239
37
155
5.98
4-5
202
28
129
5.98
5-6
174
30
172
5.83
6-7
144
18
125
5.95
7-8
126
21
167
5.73
8-9
105
17
162
5.79
9-10
88
14
159
5.83
10-11
74
6
81
5.84
11-12
68
15
221
5.35
12-13
53
4
75
5.68
13-14
49
6
122
5.12
14-15
43
7
163
4.72
15-16
36
6
167
4.56
16-17
30
8
267
4.42
17-18
22
2
91
4.96
18-19
20
2
100
4.37
19-20
18
3
167
3.81
20-21
15
0
0
3.37
21-22
15
3
200
2.37
22-23
12
7
583
1.83
23-24
5
0
0
2.50
24-25
5
0
0
1.50
25-26
5
5
1,000
0.50
and Kent Island rates of about 26 per cent (Table IV). This low
rate gives a survivorship curve (Fig. 2) which is strikingly dif-
ferent, after the second year, from those of the Kent Island and
North American populations.
Applying the same methods for determining the required
productivity of this population as were used with Hickey's data,
508 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
it is found tliat to maintain a constant population it is necessary
for a pair of gulls to raise 1.3 young- to September first follow-
ing hatching. While this would necessitate about 40 per cent
greater productivity than recorded (Paynter, 1949) at Kent
Island (0.92 young per pair raised to fledging in early August),
it may be a reasonable expectation. On the other hand, the pre-
fledo'ing mortalitv in Paludan's colonv was considerably higher
than that at Kent Island. In the Danish colony in 1943 there was
about a 10 per cent loss of eggs (vs. 28.6 per cent at Kent Island)
and an 80 per cent loss of chicks (vs. 48.5 per cent at Kent
Island). With an average clutch of three eggs this heavy mor-
tality would result in a net production of about 0.5 fledgling for
each pair of adults or, in other words, only a maximum of 39 per
cent of the required productivity as calculated from the life
table.
Paludan believed that fledgling production nearly as low as
that observed in the Danish population would be sufficient to
maintain the population if certain deficiencies in the life table
were corrected. He reasoned that the number of birds which
breed for the first time should equal the number that die during
that year. From the life table he found the mean annual mor-
tality from the second year onward to be about 15 per cent. This
would mean that in a population of 1,000 birds, 150 would die
during the year and 150 should begin to breed for the first time,
if the population is to remain stable. Using the life table figure
of 62.3 per cent mortality for the first year and 15 per cent mean
annual mortality thereafter, he calculated a production of ap-
proximately 600 fledglings per 1,000 adults (1.2 per pair) would
be necessary to yield about 150 birds two and a half years later,
when he thought breeding began. This would require more than
twice as many survivors as he had observed (0.54 per pair).
However, Paludan reasoned that the observed first year mor-
tality rate was higher than it should be owing to bias in favor of
recoveries near the breeding colonies ; a reduction in the first
year mortality would increase the number of birds surviving to
breeding age and, hence, reduce the required production of fledg-
lings per pair. Also, he believed the prefledging mortality that he
had recorded was in excess of that which is normal in Denmark.
He concluded that the yearly production of between 0.5 and 1
fledgling per pair of adults would be sufficient to maintain the
population, and even allow for the general increase that had been
noted throughout Europe.
HERRING GULL LIFE TABLES 509
One significant probable error is the assumption that the en-
tire population begins to breed in the third year. As we have
seen above, a few gulls (20 per cent) do breed this early, but
the average age is somewhat in excess of the fourth year. Wlien
Paludan's statistics are adjusted for this later breeding age, it is
found that a pair of gulls would have to produce roughly 1.4
fledglings, rather than 1.2 fledglings.
TABLE VI
Life Table for Daaiish Gulls, Utilizing Kent Island Prefiedging Data and
Assuming First Year Recoveries to be Half of Those Actually Recorded
(see i>. 508 for explauation)
X I d 1,000 q e
XX XX
O-0.07 1,000.0 286.0 286.0 1.43
0.07-0.19 714.0 346.3 485.0 1.92
0.19-1.10 367.7 166.6 453.0 3.56
1.10-2.10 201.1 43.8 217.8 5.37
2.10-3.10 157.3 30.0 190.7 5.73
3.10-4.10 127.3 19.9 156.3 5.96
4.10-5.10 107.4 14.9 138.7 5.97
5.10-6.10 92.5 16.0 173.0 5.85
6.10-7.10 76.5 9.6 125.3 5.97
7.10-8.10 66.9 11.2 167.4 5.75
8.10-9.10 55.7 9.1 163.4 5.81
9.10-10.10 46.6 7.5 160.9 5.85
10. 10-n.lO 39.1 3.2 81.8 5.88
11.10-12.10 35.9 7.5 208.9 5.36
12.10-13.10 28.4 2.2 77.5 5.64
13-10-14.10 26.2 3.2 122.1 5.07
14.10-15.10 23.0 3.8 165.2 4.71
15.10-16.10 19.2 3.2 166.7 4.54
16.10-17.10 16.0 4.3 268.7 4.35
17.10-18.10 11.7 1.1 94.0 4.77
18.10-19.10 10.6 1.1 103.8 4.22
19.10-20.10 9.5 1.6 168.4 3.65
20.10-21.10 7.9 0 0 3.29
21.10-22.10 7.9 1.6 202.5 2.29
22.10-23.10 6.3 3.7 587.3 1.73
23.10-24.10 2.6 0 0 2.50
24.10-25.10 2.6 0 0 1.50
25.10-26.10 2.6 2.6 1,000.0 .50
510 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
If normal fledgling production in the Danish population is
assumed to be one per pair of adults, which is the maximum sug-
gested by Paludan, it is necessary for the first year mortality
to drop from the observed rate of 62.3 per cent to approximately
52 per cent, in order to replace the 15 per cent loss of breeding
birds. This requirement seems reasonable if, as believed by Palu-
dan, there was a disproportionately high recovery of first-year
birds in the sample.
It is instructive to note (Table VI) that if the production of
fledglings is assumed to be the same as at Kent Island (0.92
young per pair of adults) and if the recovery of first year gulls
is halved, the first year mortality rate would be 45.3 per cent,
which is almost exactly that found at Kent Island.^ x\t sexual
maturity there then would be nearly 64 pairs of birds, with a life
expectancy of almost six years. If these birds laid an average of
2.5 eggs per clutch, there would be produced about 950 eggs, or
nearly sufficient production to maintain the population. If there
were three eggs per clutch there would be a net production of
about 1,140 eggs, adequate productivity to permit about a five
per cent annual increase in fledged birds.
From these calculations one is tempted to conclude that Palu-
dan's first-year recoveries were twice as abundant as they would
have been had the birds dispersed as widely as they do in suc-
ceeding years and that Paludan was correct in suggesting that
breeding pairs need produce only about one fledged chick per
year. However, there are certain peculiarities in the recovery
data that require examination.
Paludan (1951, Table 29, pp. 108-109) presents a convenient
tally of the annual recoveries for the year classes from 1917
through 1942. These records, in contrast to the Kent Island
data, show extremely wide annual variations. For example, in
the first year after banding the percentage of the total banded
birds recovered ranged from 1.0 per cent in 1939 to 26.7 per cent
in 1929, and in the second year the range was from 0 in five
year classes to 4.0 per cent in 1937. Doubtless some of these great
fluctuations are caused by sampling vagaries owing to the small
number of birds banded each year and also to wide variations
even within these small numbers (15 to 1,291 bandings per year).
Nevertheless, there would seem to be some fluctuations which can-
not be dismissed as sampling errors. These are well illustrated
1 For these calculations it is necessary to assume that the Danish records begin
on 1 .Tuly ami that the lirst year recoveries are made in nine-tenths of a year,
and thus are comparable with the Kent Island data.
HERRING GULL LIFE TABLES 511
by the second year recoveries for the year classes of 1935, 1940,
1941, and 1942, years in Avhich the number of birds banded did
not differ greatly. The number of gulls banded in these years
was 765, 811, 861, and 710, respectively, and the number of sec-
ond j^ear recoveries, in the same order, were 0, 11, 1, and 1. Ex-
pressed differently, the recovery rates were 0, 1.36, 0.12, and
0.14 per cent.
We need not know the reasons for these fluctuations to ap-
preciate what profound effects they have on a demographic study.
The life table constructed by Paludan shows a mortality rate in
the second year of nearly 22 per cent, a rate, as has been pointed
out, considerably lower than that recorded for any other popu-
lation. This rate was obtained from data which indicate that
about 0.77 per cent of all Danish recoveries were made in the
second year. This is the average for all 26 year classes. If, how-
ever, we eliminate the year classes for which there were no re-
coveries during the second year the average rises to 0.96 per
cent. Using this figure in the life table would increase the second
year mortality rate to 27.6 per cent, and if w^e employ the figure
of 1.36 per cent, which is the percentage of second year re-
coveries in 1940, the mortality rate rises to nearly 35.8 per cent,
which is almost exactly that of the Kent Island population. Con-
comitant with these increases in the second year mortality rate
are decreases in that of the first year, so that in the early inter-
vals the Danish life table becomes quite similar to that of Kent
Island.
These calculations should suffice to illustrate that the Danish
life table is based on small and highly variable samples, and that
even seemingly minor changes in the data may have large-scale
effects. This should also warn against assuming that Paludan 's
life table is more nearly correct than others merely because it
"balances" when certain data are borrowed from other popu-
lations.
The second life table for European gulls is that prepared by
Olsson (1958), for 1,222 recoveries, including those shot (60 per
cent), from about 12,700 birds banded in Sweden, Norway, and
Finland (=Fenno-Scandia) from the mid-1920 's to the mid-
1950 's (Table VII). The average rate of recovery was about 9.5
per cent, which is nearly three times that at Kent Island and
about ten per cent higher than that in Denmark. The table
starts on August 1 following hatching. The oldest recovery is a
gull in its 17th year; the oldest potential recovery at the time
the study ended would be an individual in its 29th year. The raw
512 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
data have been adjusted to compensate for the fact that some
year classes had not been banded long enough to yield all their
potential recoveries. It may be seen (Table IV) that the first
year mortality of 56.8 per cent is somewhat lower than that found
in Hickey's North American and Paludan's Danish populations,
and that the second year rate is slightly higher than that of the
North American gulls, and considerably higher than that of the
Danish birds, but still not so great as that of the Kent Island
population. For the next ten years the mean annual mortality is
33.8 per cent, which is more than twice the rate recorded by
Paludan and moderately higher than that of the American and
Kent Island populations.
Olsson had no records of the fledgling production in the Fenno-
Scandian colonies and did not, therefore, attempt to test the accu-
racy of his life table against such data. Using the method devised
by Hickey, it is found that 2.86 fledglings must be raised to
TABLE VII
Life Table for Fenno-Scandian Gulls Bauded 1925-1935 (1,122 Kecoveries)
(Olsson, 1958)
X
I
d
1,000 q
e
X
X
X
X
0-1
1,000
568
568
1.93
1-2
432
138
319
2.80
2-3
294
81
276
2.88
3-4
213
66
310
2.79
4-5
147
45
306
2.81
5-6
102
24
235
2.83
6-7
78
26
333
2.55
7-8
52
15
289
2.58
8-9
37
11
297
2.42
9-10
26
6
231
2.23
10-11
20
n
550
1.75
11-12
9
5
556
2.28
12-13
4
0
0
3.50
13-14
4
0
0
2.50
14-15
4
2
500
1.50
15-16
2
0
0
1.50
16-17
2
2
1,000
0.50
HERRING GULL LIFE TABLES 513
August 1 by each pair of breeding gulls if the population is to
remain static. Even with a mean clutch of three eggs, such a high
rate of nesting success appears improbable and it must be con-
cluded that the Fenno-Scandian life table is also inaccurate.
POSSIBLE ERRORS IN THE KENT ISLAND TABLES
From the fact that the Kent Island colony was observed to
have maintained its size from at least 1935 to 1948 (see p. 501) it
seems reasonable to conclude that the population was either rela-
tively stable during the 1930 's and 1940 's or, even more likely,
that it was increasing and its surplus overflowed the colony and
added to the expanding North American population. The life
tables, however, indicate that the population should have been
decreasing at a catastrophic rate. If we assume that the life
tables are deficient, we must look for a source, or sources, of
error within the data used in constructing the tables.
There are three kinds of error which might distort the life
tables, causing them to indicate that the population was rapidly
declining. First, banding recoveries may have been accumulated
for too short a period (23 through 28 years), thereby failing to
cover the full life spans of the six cohorts studied. Second, the
1947 egg and prefledging mortality rates, which were used in con-
structing the initial intervals of all six tables, may have been
uncharacteristically high. Third, bands may have been recovered
more readily in the early postfledging years than later in the life
spans, causing an apparent increase in the mortality rates for the
early intervals of the tables.
The first hypothesis, i.e. that a study allowing for recoveries
for a maximum period of between 23 and 28 years does not cover
the full life span of the Herring Gull, is correct, but probably
only to a small and insignificant degree. For example, a Kent
Island gull has been found alive in its twenty-sixth year, and a
bird banded as an adult was recovered 24 years later meaning it
was a minimum of twenty-seven years old when it died. There
are also records of captive Herring Gulls which have lived
nearly fifty years (Gross, 1940) ; it is conceivable that some wild
individuals may attain an equally advanced age. Nevertheless,
it seems that very few gulls live beyond their twenty-third year,
which is the oldest potential recovery within the youngest cohort
studied at Kent Island, and those birds that do live longer doubt-
less have little influence on the reproductive rate of the popula-
tion because of their relatively insignificant numbers and possibly
(unproven) because of sterility brought about by senility.
514 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Support for the argument that the life tables adequately cover
the life spans of the gulls may be found in the recovery data.
Of the 954 recoveries from the five oldest cohorts, with a maxi-
mum potential age ranging from 24 to 28 years, only one indi-
vidual (0.16 per cent) reached the age of 24, none exceeded that,
and but three lived more than 18 years. Expressed differently,
less than one-fifth of one per cent of the recoveries were of birds
older than 18 years. In Paludan's Danish study (1951) there
were eight cohorts with a maximum potential age between 26 and
34 years. Of the 236 recoveries from these cohorts there was just
one bird in its twenty-sixth year and the recovery rate between
the eighteenth and twenty-sixth year was only 0.8 per cent of
the total sample. The maximum potential age of birds in Olsson's
(1958) Fenno-Scandian population was twenty-nine years, but
the oldest recovery was a single seventeenth-year individual.
There is, of course, the possibility that a loss of bands could
account for the failure to recover gulls even older than those
now known. But it is believed that while the frequency of re-
coveries may be reduced because of band loss, there is little likeli-
hood that the maximum span of life is significantly greater than
that which has been recorded. We must, therefore, seek another
explanation for the failure of the life tables to document the
population dynamics of the Kent Island colony.
The second hypothetical source of error in the life tables, i.e.
that the i^refledging mortality in 1947 was unusually high, is al-
most certainly void. It was found that nearly 29 per cent of the
eggs failed to hatch and about 48.5 per cent of the young died
before fledging, resulting in 63.2 per cent mortality, or a net
production of 0.92 fledglings per breeding pair (Paynter, 1949).
This falls well within the range recorded at other Herring Gull
colonies. For example, Paludan (1951) who admitted that his
calculations are imprecise, estimated a production of 0.5 fledged
young per pair, or 83 per cent prefledging mortality. In a study
on the Summer Isles in the Irish Sea, Darling (1938) found five
colonies (ranging from 6 to 150 individuals) which had pre-
fledging mortality rates between 58.3 and 88.9 per cent, and
which yielded from 0.78 to 0.96 fledged young per nest. Drost,
Focke, and Freytag's (1961) German colony, which started with
two pairs of birds and in 12 years built up to 139 pairs, had an
average prefledging mortality rate of about 75 per cent, resulting
in the fledging of an average of 0.7 gulls from each nest. On
Skokholm, an island off the coast of Wales with a colony of 300
pairs of gulls, Lockley (1947) estimated that less than one
HERRING GULL LIFE TABLES 515
fledgling per adult pair was produced. In another Welsh colony
of 440 pairs, Harris (1964) reported a production of about one
fledgling for each two nests.
Other species of gulls laying three-egg clutches show a similar
range of nesting failures. A colony of Ring-billed Gulls (L.
delaivarensis) in Michigan suffered 88 per cent prefledging losses,
producing 0.67 young per pair of adults (Emlen, 1956). The
California Gull {L. calif ornicus) ^ in a colony in Utah, was found
to have unusually low prefledging mortality, suffering a loss of
only about 40 per cent of its eggs and young, and fledging 1.77
chicks per nest (Behle and Goates, 1957). In British Columbia a
colony of Glaucous-winged Gulls {L. glaucescens) , studied for
two years, produced 1.0 and 1.7 young per nest, and had a mor-
tality rate of 64 and 52 per cent, respectively (Vermeer, 1963).
Four small colonies of Lesser Black-backed Gulls (L. fuscus) in
the Summer Isles produced about 1.5 young from each nest, with
a prefledging mortality rate of approximately 48 per cent (Dar-
ling, 1938).
In summary, the prefledging mortality rate at Kent Island in
1947 was below that which has been found in most other Herring
Gull colonies and in a colony of L. delaivarensis, and somewhat
higher than that recorded for populations of L. californicus, L.
glaucescens, and L. fuscus. From this it is concluded that the
1947 prefledging survival data are in the right order of magni-
tude and probably are not responsible for the failure of the
composite life tables to indicate that the population was stable or
increasing.
The third type of error that could distort the Kent Island life
tables, causing the survival rates for breeding birds to appear too
low to maintain the population, is a disproportionately high re-
covery of bands in early age classes. This could be brought about
through some circumstance which allows young birds to be more
readily recovered than older individuals, through a loss of bands
among older birds which would reduce their rate of recovery, or
through a combination of these factors. Both phenomena are
difficult to detect but almost certainly at least one is the reason
that the life tables are not reconcilable with the observed status
of the population.
Paludan (1951, p. Ill) believed that the mortality rate for the
first year class of the Danish gulls was unrepresentatively high
because newly fledged birds died near the natal colony, where
they are more likely to be recovered and reported than older
birds which range more widely. At Kent Island, however, the
516 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
fledglings quickly leave the colony and winter far south of New
Brunswick. Each successive year the length of the migration is
lessened until as adults the birds winter only a few hundred
miles from the colony (Gross, 1940). This migration pattern re-
duces the opportunity for any recoveries at Kent Island during
the winter months. Moreover, the records indicate that very few
immature gulls are recovered at the colony during any season.
There seems no possibility that the recovery sample is biased in
favor of immature birds because they die in the vicinity of the
natal colony.
There remains to be considered the possibility that the wide
dispersal of young birds during the winter, and their failure to
return to the colony during the breeding season, might in some
way enhance their chances of recovery over those of the less wide-
ranging adults. If the immature gulls were in closer proximity
than the adults to urban areas, w^here dead birds are more likely
to be found, this possibility would exist. However, during the
winter the younger birds are scattered from the heavily popu-
lated northeastern United States south to the more sparsely in-
habited (tulf Coast and Central America, and in the breeding
season return to the Northeast. On the other hand, the adults
winter in the heavil^y populated Northeast and breed where ob-
servers are constantly alert for dead birds wearing bands. Bias
in favor of the recovery of immature gulls seems improbable
under these circumstances.
Having considered and rejected as improbable all other poten-
tial sources of error, we are left with only band loss to account
for the discrepancies between the life tables and the observed
status of the Kent Island colony.
All Kent Island gulls were marked with aluminum butt-end
l)ands, rather than with the locking, clip-type bands now used in
Europe. In a British study, using butt-end and clip bands on
dilferent samples plus supplementary durable plastic bands,
Poulding (1954) found that during the first year after banding
as fledglings, Herring Gulls with locking bands were recovered
with twice the frequency of birds wearing butt-end bands.
Breaking down the analysis further, it was noted that when the
gulls wore butt-end bands only about four per cent of the year's
total recoveries were made in the second half of the year, whereas
nearly 41 per cent of the recoveries occurred in the second six
months when the birds wore clip-type bands. Thus the data indi-
cate that 50 per cent of the butt-end bauds are lost in the first
year and nearly all of this loss takes place before the bands are
HERRING GULL LIFE TABLES 517
six months old. Observations of living birds confirmed the 50 per
cent loss of butt-end bands and showed a complete retention of
clip bands.
This evidence strongly suggests that the heavy initial loss of
butt-end bands may be caused by the removal of the bands by
the gulls, rather than by a weakening of the bands through wear.
Differences in the strength of individual bands, and possibly
variations in the manner of closing them, could well account for
the rapid loss of half of the bands while some of those in the
remaining half are retained in good condition for many years.
If this type of band loss occurs among Kent Island birds, the
life tables would be significantly affected. It would mean, dis-
regarding for the moment the normal reduction in the size of
cohort caused by death, that at the time of fledging the banded
sample would be twice the size it is when entering the second
year of life. It follows that the number of recoveries in the
second year, and all subsequent years, would be half of the total
had there been no loss of bands and that the recoveries in the
first year would be somewhat lower, but not a full 50 per cent
lower because the bands are retained for part of the first half
year. Viewed from a different aspect, recoveries in the first year
would be disproportionately more numerous than those of the
remaining year classes. The effect on the life table Avould be a
reduction in the calculated number of birds surviving beyond the
first year and, of course, a marked decrease in the apparent num-
ber of individuals which survive to the breeding age.
We have no evidence that Kent Island gulls lose half their
bands during the first six months, but if we adjust the life table
to compensate for such a loss and the table is then reconcilable
with the apparent true status of the population, we shall have
good circumstantial evidence that this is the source of error. If,
for the sake of simplicity, we assume that the mortality rate is
constant during the entire first year and that 50 per cent of the
bands are lost at the end of the first six months and none in the
second half year, one-third of the recoveries would occur in the
second six-month period. Doubling this figure will give the num-
ber of recoveries w^hich would have been made if the initial
banded sample had been half its original size and there was no
subsequent loss of bands. For the six combined cohorts there
were 1,09!) recoveries, 494 of which occurred during the first
year. Making the proposed adjustments for the first year, the
first year recoveries would drop to 329 and the total for the en-
tire life span to 934.
518 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Using these data to construct a new life table, it is found that
the number of survivors reaching the breeding age of year 3-4
rises to about 116, versus 98.4 in the uncorrected table. The life
expectancy at this age remains 3.3 years. AVith a breeding cohort
of this size and with this life expectancj^ a total of 477 eggs
could be produced, which is an increase of 71 eggs over the total
(406) calculated for the uncorrected life table. Nevertheless, this
is still less than half the number required to maintain a level
population.
If we were to assume that early band loss is the sole source of
error in the life table, in order to achieve sufficient production at
the breeding stage it would be necessary to adjust the recoveries
in the first year so that their total would fall well below that of
the second year. Such a low rate of mortality is obviously spu-
rious. We must conclude that while early band loss is a distinct
possibility in the Kent Island population, there must be addi-
tional losses later in the life span.
Band loss maj^ fall into three broad patterns. The first, and the
most expected, is loss which is correlated with the age of the
bands. One would expect bands to become progressively weaker
through wear and as a result be lost with increasing frequency.
The second type is a proportional, or constant, loss. This is most
likeh" to take place if the bands are continually removed by the
birds or if they merely drop off at random. The third pattern,
which may be uncommon, is an inconsistent, or fluctuating, loss.
This may occur because of variations in behavior, such as shifts
in feeding ranges from less saline to more saline water, which
might have a variable effect on the durability of the bands, or
because the new bands are not of uniform strength or are not all
fastened securely. The large initial loss of bands noted by
Poulding (1954) is an example of an inconsistent pattern which
seems attributable to the latter cause.
If we are able to determine the pattern of band loss among
Kent Island gulls it may then be possible, in certain instances, to
adjust the raw data to compensate for the losses and to construct
a life table which will document the colony's demography. Be-
cause the Kent Island birds were all marked with similar bands
there is no control group against which the various cohorts may
be tested, as in Poulding 's study. We must, therefore, see if any
evidence of band loss may be detected within the data available.
Semi-logarithmic graphs of survivors (Ix) offer a means of ap-
proaching the problem.
HERRING GULL LIFE TABLES 519
If survivors (Is) are plotted against age on a semi-logarithmic
graph, the points will form a straight line if the rate of survival
is constant. If the survival rate should decrease, or if there is an
accelerating loss of bands which would create an apparent de-
crease in the rate of survival, the line will assume a sigmoid
shape, being deflected downward. As may be seen in the plot for
the six combined cohorts at Kent Island (Fig. 2), starting with
the fourth year (year 3-4) of life, the survival rate is relatively
constant for at least ten years (to year 12-13). There is then a
downward trend, but there are so few recoveries one cannot be
certain that the pattern is not an artifact. From this it is evident
that if there is band loss that is correlated with age, it does not
begin until sometime after age thirteen, by which time the cohort
has dwindled to a small fraction of its initial size.
A further test for a correlation between a loss of bands and the
time which they have been worn may be made by utilizing the
recoveries of birds banded as adults. Between 1936 and 1947 a
total of 1,856 gulls in adult plumage were banded; 97, or 5.2
per cent, were recovered up to 30 June 1963 (Table VIII).
TABLE VIII
Kent Islaud Adults Banded and Admissible ^ Eecoveries
Year
1936
1937
1938
1939
1941
1946
1947
Total 1,856 97 5.23
1 Set' 1). 4!);j for (li>liiiit:(iii of ailniissible recoveries.
Nothing is known of the ages of these birds beyond the fact that
they were in adult plumage and, therefore, were at least in their
fourth year of life when banded. The maximum time available
for recoveries from the youngest cohort is 16 years and from the
oldest cohort it is 28 years. The oldest recovery was a bird
banded 24 years earlier ; the next oldest were two gulls which had
mded
Recoveries
Per ce
200
2
1.00
196
13
6.63
611
46
7.53
100
5
5.00
155
5
3.23
497
21
4.22
97
5
5.15
520 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
borne their bands for 14 and 17 years, respectively. A semi-
logarithmic survivorship curve for the 97 recoveries from the
combined seven cohorts is shown in Figure 3. Owing to the
paucity of older recoveries, no adjustment has been made for the
fact that after age 16 the number of potential recoveries de-
creases because not all cohorts have been banded sufficiently long
to yield recoveries between ages 17 and 24.
In Figure 3 a comparison is made between the semi-logarithmic
survivorship curve for the birds banded as adults and the curve,
starting at the fourth (adult) year, for those gulls banded as
fledglings. If there is a positive correlation between the age of
the bands and their loss, one would expect the survival rate for
the latter group to decrease, and the curve to become deflected
three years sooner than for the former group, because the birds
banded as fledglings had borne their bands three years longer. If
band loss begins during the first year following banding, the
curves w^ould diverge immediately. If it starts some years later
the two lines would remain parallel until band loss begins, when
the curve for the group with older bands would descend more
rapidly ; the plot for the group with the newer bands would con-
tinue in a straight line (a constant annual survival rate) for
three additional years and then it too would begin to fall away.
As may be seen in Figure 3, the two curves remain nearly
parallel for ten years, or in other words, until the group banded
as fledglings has reached its thirteenth year. Then, as has been
discussed, the curve for the group with the older bands begins
to decline at a more rapid rate. This is not follow^ed three years
later by a decline in the other curve, which one would expect if
bands began to be lost at an increasing rate after they had been
worn thirteen years. In both groups, and particularly in the
group banded as adults, there are so few recoveries in the older
age categories it would be imprudent to attempt at this time to
read any significance into these differences. All we may safely
conclude is that if band loss is a function of age it almost cer-
tainly does not begin until the bands liaA'e been worn at least
thirteen years. Because this means that the birds could have bred
for ten seasons prior to the beginning of accelerated band loss,
there seems little likelihood that this type of band loss would
have a significant effect on the life tables.
This test also reveals an interesting fact regarding early band
loss. It indicates that gulls banded as adults do not suffer a
greater loss of bands in the first year than in later years, which
is contrary to what one would have expected from Poulding's
HERRING GULL LIFE TABLES
521
study (1954) of British fledglings. This is probably an indica-
tion that the North American butt-end bands are more uniformly
durable than the British butt-end bands. This also seems to ex-
plain why the attempt to compensate for disproportionately
1,000 —
100 _
<
Z3
O
l/J
o
>
>
3
I/)
Banded as adults
10 _
Banded as fledglings
l_
[ I I I I ' I ■ I ■ I ■ 1 ■ I M
6 8 10 12 14 16 18 20 22
' I ■ I
Adults 0 2 4
Fledglings 3.1 5.1 7.1 9.1 II. I 13.1 15.1 17.1 19.1 21.1 23.1 25.1
AGE IN YEARS
Fig. 3. Survivorship curves for Kent Island gulls banded as fledglings
and as adults.
522 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
heavy band loss within the first year class of Kent Island gulls,
banded as fledglings, failed to produce a satisfactory life table.
Nevertheless, one could argue that adults do not remove their
bands with as great a frequency as fledglings, explaining this
difference by assuming that the legs of fledglings are more sensi-
tive than those of adults and that the irritating bands are re-
moved. Or, possibly, one could reason that in the process of
learning to detect what is food, and what is not, the fledglings are
attracted by the shiny bands and peck at them until they are
loosened and lost. However, the fact that the Kent Island life
table is not appreciably improved by adjustments made to com-
pensate for heavier band loss in the first year of life seems to
negate such arguments. The simple explanation that British
butt-end bands are of more variable durability than those used at
Kent Island seems the most satisfactory explanation.
The only pattern of band loss yet to be considered is that which
occurs at a constant, i.e. proportional, rate. Having concluded
that band loss is not positively correlated with the age of the
band, and that bands are not lost with a greater frequency at any
particular time during the bird's life, it is almost certain that
the Kent Island population suffers a steady loss of bands during
its entire life span. This would progressively reduce the number
of recoveries in each successive age interval, thereby depressing
the survivorship curve and reducing the calculated expectation
of life throughout the life table.
We do not know the rate at which bands are lost and whether
this rate is sufficiently low to allow at least a portion of the
population to retain its bands until it has lived its full life span.
However, because a living gull was found at Kent Island which
had borne a band for 26 years, and because there are records
from there of four birds Avhich died between the ages of 20 and
24, it is evident that the rate of band loss must be comparatively
low. For example, if the annual loss ran as high as 20 per cent,
the roughly 32,000 individuals in the Kent Island sample would
have dwindled to about 455 banded birds by the beginning of
the twentieth year, even without considering attrition owing
to mortality. If the average annual mortality were 10 per cent,
making a cumulative annual reduction in the marked population
of 30 per cent, there would have been just 37 banded survivors
at the start of the twentieth year, certainly too few to have
yielded five records of birds 20 years and older. Assuming that
losses and mortality are each 10 per cent annually, there would
be approximately 445 banded survivors after nineteen years, or
HERRING GULL LIFE TABLES 523
about 1.4 per cent of the initial sample, which seems ample to
allow for the recovery of five individuals in the next seven years.
Althoug-h the Herring Gull is a long-lived species, 10 per cent
average annual mortality would appear rather low; Drost et al.
(1961) calculated a rate of 10 per cent for adult birds, but our
calculations must include immatures within the average. If we
continue to allow for 1.4 per cent survival at age 20, any increase
in the mortality rate would have to be balanced by a decrease in
the rate of band loss. Thus, 15 per cent annual mortality, which
is about the figure found by Paludan for birds between the ages
of 2 and 12, would mean a band loss of five per cent at the most.
Crude as these calculations may be, it seems reasonable to con-
clude that Kent Island gulls lose bands at an average rate some-
where in the vicinity of 5 or 10 per cent per year and that the
average annual mortality rate must range betw^een 10 and 15
per cent.
These speculations are based on the premise that bands are
recovered throughout the life span of the birds and are not
totally lost before the oldest gulls die. There can be no proof
that this is correct, but empirically it would seem that the oldest
recoveries probably very nearly represent the potential natural
longevity of the Kent Island Herring Gull. A larger cohort, more
durable bands, and additional decades of observation surely
would produce recoveries older than those now known, but it is
difficult to imagine that these could be more than an insignificant
fraction of the total sample.
CONCLUSIONS
It is now evident that a continued loss of bands accounts for
the failure of the Kent Island life tables to reconcile with the
observed status of the population (i.e. either a stable or increas-
ing population). Unfortunately, in spite of our estimate that
the loss amounts to about 5 or 10 per cent annually, there is no
way by which the raw data may be adjusted to compensate for
these losses. We know, for example, that there were 494 re-
coveries for the combined cohorts in the first nine-tenths of a
year, and that without a ten per cent loss of the banded sample
there would have been about 549 recoveries. In the second year
there were 220 recoveries, but this number, without band losses
during the second year, would have been approximately 243, plus
an unknown number of individuals which lost bands in the first
vear but survived to die during the second year. The difference
524 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
between the recorded recoveries and the number of recoveries
there would have been had there been no band loss, increases, of
course, in each successive age interval. This also has the effect of
accelerating the descent of the survivorship curve and probably
accounts for the difference between the Kent Island curve and
that of the Danish population (Fig. 2).
From this analysis we must conclude that because of band
losses the Kent Island banding records are nearly valueless as a
means of investigating the dynamics of this population of
Herring Gull. Their only use in studies of this sort, if one is
willing to accept the premise that some bands are sufficiently
durable to be retained through the life span of at least a few of
the longest living gulls, is to demarcate the maximum potential
longevity of the species. All North American Herring Gulls have
been marked with similar butt-end bands, and band loss is with-
out doubt the reason Hickey (1952) also failed in his attempt to
construct an accurate life table. Because aluminum butt-end
bands are used on almost all birds banded in North America, any
data obtained from this source is suspect. Future workers should
be particularly alert to the possibility of band loss before in-
vesting time and effort in this type of research.
Locking bands were used on nearly all gulls banded in Fenno-
Scandia (Olsson, 1958) . However, the material used in the manu-
facture of the bands evidently was not very durable, for Olsson
estimated that about 5 per cent of their original weight was lost
annually. Such rapid wear, and the resulting loss of bands, is
presumably the reason for the apparent reduced maximum lon-
gevity of the Fenno-Scandian gulls when compared with North
American and Danish birds. There can be little doubt that the
similarity between the survivorship curves for Fenno-Scandian
and North American birds is caused by band losses.
Paludan (1951) does not mention the type of band used on
Danish gulls, but, according to Poulding (1954), butt-end bands
were first used and later abandoned in favor of locking liands.
Possibly some of the yearly variations in recovery rates that have
been noted (p. 510) are attributable to changes in the type of
bands employed. Nevertheless, this cannot be one of the main
causes for these variations because one would expect the recovery
rates for the more recent year classes to be consistently higher
than those for the older year classes, but no such pattern is
evident.
Band loss almost certainly occurred in the Danish population.
Even if locking bands eliminate losses earh* in the life span,
HERRING GULL LIFE TABLES 525
wear surely accounts for band failures in the later years of this
long-lived gull. Bias in favor of recoveries during the first year
of life, and band losses during the latter part of the life span,
must be the primary reasons for the deficient Danish life table
and its dissimilarity to other life tables. Fundamental differences
between the mortality rates of the North American and Fenno-
Scandian populations, on one hand, and the Danish population,
on the other hand, as proposed by Paludan (1951) and Olsson
(1958), seem most unlikely.
ACKNOWLEDGMENTS
I am grateful to Charles E. Huntington for making available
the banding records of the Bowdoin Scientific Station, and to
Allen J. Duvall for his assistance in obtaining microfilms of the
Fish and Wildlife Service files. Da\4d W. Norton was of great
assistance in collating these records. I thank Barry Margolin for
spending much time pondering the statistical aspects of the
study, my wife, Elizabeth S. Paynter, for preparing the figures,
and Charles H. Blake for his helpful reading of the manuscript.
SUMMARY
1. At Kent Island, New Brunswick, 31,694 Herring Gull
{Lams argentatus smithsonianiis) fledglings were banded with
aluminum butt-end bands between 1934 and 1939; by 30 June
1963 there were 1,099 (3.47 per cent) recoveries suitable for use
in a mortality series (Table I) .
2. The maximum potential age for a recovery in the oldest co-
hort is 28 years, and in the youngest cohort it is 23 years; the
two oldest birds at death were in their twenty-fourth and twenty-
second years; one gull w^as captured alive in its twenty-sixth
year.
3. About five per cent of all recoveries are of birds which
have been shot ; rates vary within the six year classes from 3.6 to
6.5 per cent; probably more birds are shot than are reported
because the species is protected by law ; no gull older than thir-
teen has been reported shot ; there is no indication that immature
birds are more readily shot than adults.
4. Using prefledging mortality data for 1947 (Paynter, 1949),
composite life tables are constructed for each of the six cohorts
and for the combined cohorts (Table II) ; survivorship curves
526 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
are also drawn (Figs. 1 and 2) ; there are minor yearly varia-
tions, but the patterns are generally similar, showing heavy mor-
tality the first year, lessened mortality the second year, and a
lower, relatively constant, rate thereafter.
5. Assuming breeding begins in the fourth year (year 3-4),
that the average clutch is 2.5 eggs, and that each nesting results
in the production of 0.92 fledglings (Paynter, 1949), it is found
that an average of about 98.4 gulls attain breeding age from
each 1,000 eggs laid, and these have a life expectancy of 3.3
years, enabling them to produce 405.9 eggs, or about 41 per
cent of the number required to maintain a stable population.
6. The population is believed to liave been stable, or possibly
expanding, from at least 1935 to 1948. The life tables must,
therefore, be faulty.
7. Three types of error leading to distorted life tables are
possible: 1. recoveries were accumulated for too short a period
(23 to 28 years) to document the potential life span; this is
probably a minor and insignificant source of error. 2. The 1947
egg and prefledging mortality rates were excessively high ; this
is rejected because comparable rates have been found in this
and related species. 3. There is a disproportionately high re-
covery of bands in the early age classes because (a) young birds
are more readily recovered, (b) older birds lose bands and are
lost from the sample, or (c) both factors are operative.
8. It is certain that band loss, resulting in a disproportionately
high recovery of young birds, must account for the failure of
the life tables to document the demography of the Kent Island
population.
9. Band loss could be (a) positively correlated with the age
of the band, (b) proportional (i.e. constant) with respect to the
size of the sample, or (c) fluctuating; it is concluded that bands
are lost at a relatively constant rate throughout the life span.
10. Assuming that some bands are retained long enough to
document the maximum potential life span, mortality and band
loss combined seem not to exceed 20 per cent annually ; it is sug-
gested that band loss averages around 5 or 10 per cent per year
and that the average annual mortality ranges between 10 and 15
per cent.
11. Hickey's (1952) life table (Table III; Fig. 2) for North
American gulls proved faulty presumably because of band losses
similar to those at Kent Island.
12. Olsson's (1958) life table (Table VI; Fig. 2) for Fenno-
Scandian gulls is similar in pattern to those for North America
HERRING GULL LIFE TABLES 527
and Kent Island; locking bands probably eliminated losses
throng'h mechanical failnre, but rapid wear (five per cent an-
nually) doubtless resulted in band loss correlated with age and
eventually to a total loss of marked birds by the eighteenth year,
foreshortening the apparent maximum life span.
13. Paludan's (1951) life table (Table V; Fig. 2) for Danish
gulls, with its higher survival rate, most nearly fits what is
known of the demography of the species but bias in favor of re-
coveries during the first year, wide variations (of unknown
cause) in yearly recovery rates, and probably band losses to-
ward the end of the life span distort the data.
14. Suggestions of essential differences between the mortality
rates of the various populations are considered spurious.
LITEEATUEE CITED
Behlb, W. H., and W. a. Goates
1957. Breeding biology of the California Gull. Condor 59:235-246.
Crystal, F. H.
1941. Herring Gull census. Ann. Eep. Bowdoin Sei. Sta. 5:37-38
[mimeo.].
Darling, F. F.
1938. Bird flocks and the breeding cycle. Cambridge U. Press, Lon-
don. X + 124 p.
Drost, E., E. Focke and G. Freytag.
1961. Entwicklung und Aufbau einer Population der Silbermowe,
Larus argentatus argentatus. J. Ornithologie 102:404-429.
Emlen, J. T., Jr.
1956. Juvenile mortality in a Eing-billed Gull colony. "Wilson Bull.
68:232-238.
Earner, D. S.
1955. Birdbanding in the study of population dynamics. In A. Wolf-
son [ed.], Eecent studies in avian biology. U. Illinois Press,
Urbana. Pp. 397-449.
Goethe, F.
1937. Beobachtungen und Untersuchungen zur Biologie der Silber-
mowe {Larus a. argentatus Pontopp.), auf der Vogelinsel Mem-
merstand. J. Ornithologie 86:1-119.
Gross, A. O.
1940. The migration of Kent Island Herring Gulls. Bird-Banding
11:129-155.
Harris, M. P.
1964. Aspects of the breeding biology of the gulls Larus argentatus,
L. fuscus and L. marinus. Ibis 106:432-456.
528 BULLETIN : MUSEUM OF COMPARATI\^ ZOOLOGY
HiCKEY, J. J.
1952. Survival studies of banded birds. U.S. Dept. Interior, Fish and
Wildlife Ser., Spec. Sci. Eep., Wildlife No. 15. 177 p.
Lack, D.
1954. The natural regulation of animal numbers. Oxford XJ. Press,
London, viii + 343 p.
LOCKLEY, E. M.
1947. Letters from Skokholm. J. M. Dent and Sons Ltd., London.
X + 243 p.
Marshall, H.
1947. Longevity of the American Herring Gull. Auk 64:188-198.
Olsson, V.
1958. Dispersal, migration, longevity and death causes of Strix aluco,
Buteo buteo, Ardea cinerea and Larus argentatus. A study
based on recoveries of birds ringed in Fenno-Scandia. Acta
Vertebratica 1:91-189.
Paludan, K.
1951. Contributions to the breeding biology of Larus argentatus and
Larus fuscus. Vidensk. Medd. fra Dansk naturh. Foren. 114:1-
128.
Paynter, E. a., Jr.
1947. The fate of banded Kent Island Herring Gulls. Bird-Banding
18:156-170.
1949. Clutch-size and the egg and chick mortality of Kent Island
Herring Gulls. Ecologj' 30:146-166.
POULDING, E. H.
1954. Loss of rings by marked Herring Gulls. Bird Study 1 : 37-40.
Vermeer, K.
1963. The breeding ecology of the Glaucous-winged Gull {Larus glau-
cescen^) on Mandarte Island, B. C. Brit. Columbia Prov. Mus.
Occas. Papers 13, 104 pp.
(Eeceived September 2, 1965.)
Harvard MCZ LIbrai
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