HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. LXXXII, No. 1 THE BRAINCASE OF THE CARBONIFEROUS CROSSOPTERYGIAN MEGALICHTHYS NITIDUS By Alfred S. Romer CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE MUSEUM April, 1937 No. 1. — The Braincdsr of the Carboniferous Crossopterygian Megalichthys nitidus By Alfred S. Romer INTRODUCTION Below is given an account of the braincase and closely associated structures of the Permo-Carboniferous rhipidistian crossopterygian Megalichthys nitidus (Cope). At a later time I propose to describe the remainder of the cranial anatomy of this fish. The rhipidistian crossopterygians are of great phjlogenetic impor- tance, since they are unquestionably the closest known relatives of the ancestors of land vertebrates. But despite the great advances made during recent years by Watson, Stensio and others in our knowledge of the anatomy of paleozoic fossil fishes their braincase is still very inadequately known. A complete summary of the existing literature is given by Holmgren and Stensio (1936). In 1919 Bryant showed that the Eusthenopteron braincase was composed of two major seg- ments, pictured the main outlines of these elements, but found little interpretable detail. Watson and Stensio have since published revised restorations of the palate in which some features of the braincase are visible and the latter has published a side view of the braincase with a number of foramina identified. Watson has published a lateral view of the braincase of Ostcolcpis, a photograph of a natural cast of the ear cavities and brief description of other salient features of this genus. This comprises our entire knowledge of the rhipidistian braincase; the internal structure is as yet almost unknown, although Stensio has promised a future study based on sections of Eusthenopteron material. The coelacanths, the only other group of crossopterygians (since Polypterus is now generally acknowledged not to belong to this stock) are obviously aberrant and degenerate, but ne^•ertheless of interest. A number of late paleozoic and mesozoic genera have been described by Stensio, Watson and Aldinger. These show a great reduction in ossification and their interpretation depends upon comparison with more primitive members of the group. For knowledge of such primi- tive types we are dependent upon two Devonian specimens described by Stensio. One is a nearly complete braincase of Diplocercides, which has been described by Stensio upon several occasions. He has figured the lateral surface and several cross-sections and promises a fuller 4 bulletin: museum of comparative zoology future account of the knowledge derived from the sections. A second specimen is the anterior part of a skull described as Dictyonostcus ardicus, which agrees in general with Diplocercides; this form is of somewhat uncertain phylogentic position but, as Stensio has shown, it shows coelacanth affinities. Megalichthys nitidus was described many years ago by Cope, but until recently material has been exceedingly rare and consequently little morphological information was available. During the past few years however a considerable quantity of material has been collected by field parties under the writer's direction, and the description given is based upon this material. All the specimens are from the various formations of the Wichita group of the Texas "redbeds," a group which lies close to the boundary between the Carboniferous and Per- mian and which the writer (1936) has recently argued is to be regarded as late Carboniferous, essentially Stephanian, in age. The description is based mainly upon the following specimens, all except one being in the collections of the Museum of Comparative Zoology, Harvard University : No. 6494. A skull nearly complete and little damaged except for weathering in the rostral and nasal capsule region. Belle Plains for- mation, valley of Little Wichita River, Baylor Co. This has been sec- tioned vertically at half-millimeter intervals by the "peel" method which, as I have recently noted (1936a), I have used for vertebrate material with success. No. 6495. A skull, crushed flat and eroded dorsally, sectioned at much closer intervals in a horizontal plain. Moran Formation, head of Cottonwood Creek, Archer Co. No. 6496. Posterior half of braincase. Admiral Formation, Rattle- snake Canyon, Archer Co. No. 6497. Anterior half of braincase. Probably Moran Formation. West of Anarene, Archer Co. No. 6. Walker Museum. Posterior half of braincase. Probably Admiral Formation; 12 miles northeast of Wichita Falls, Wichita Co. No. 6498. Anterior half of braincase. Probably Moran Formation. West of Anarene, Archer Co. No. 6499. A number of specimens, mainly somewhat immature, Belle Plains Formation, Tit Mountain, Archer Co. The figures are a composite, based for internal structures on No. 6494 and for the external surfaces on Nos. 6496 and 6497, except for details in which these specimens were imperfect. The size is that of the sectioned specimen, which appears to be that of the average romer: braincase of megalichthys o adult. Most specimens appear to have been within 10% of the size represented; a few, apparently immature, as mueii as 20% smaller. Otico-occipital, dorsal aspect From above (fig. 1) the posterior moiety of the braincase shows three main areas: (1) a central region lying under the "parietals," covering the braincase and the most medial portion of the ear capsule; (2) a backwardly directed occipital region; (3) on either side an ex- panded otic region. In the central region the braincase extends upward to reach the under side of the "parietal" bones, to which it is tightly fused. Because of this fusion, the surface has been seen only in section, and small details cannot be made out. I have found no foramina piercing the dorsal surface. There is a slight hollowing medially somewhat back of the middle of the "parietal" area. Posteriorly small vessels (not indi- cated in the figure) appear to extend forward beneath the "parietals" to the ossification centers of these elements. Posteriorly the dorsal surface dips to terminate in a triangular shelf overhung slightly by the posterior ends of the "parietals." The sur- face here is irregular, the details varying from specimen to specimen; there are indications of the point of entrance of small blood vessels on to the shelf from the postero-lateral borders. On the surface of the shelf rested the anterior margins of the extrascapulars. It must be emphasized in connection with the often assumed homo- logization of these bones with the tetrapod tabulars and dermal- supra-occipitals, that the connection is a loose one; they are not in any sense integral parts of the skull proper. Laterally a thin layer of bone continuous with the perichondral layer of the braincase runs out imder the "supratemporal" and is fused to that element. In the figure this has been cut away on the right side to show the extent of the fossa lying beneath it and above the otic capsule. Half way forward, close to the division between "supratemporal" and "intertemporal" a solid ridge extends outward from the braincase proper; this contains the anterior vertical canal of the ear. There is little evidence of any extension from the brain-case roofing the fossa which lies beneath the "intertemporal" element anterior to this ridge. Overlying the foramen for the trigeminus nerve there is a short lateral dorsal projection from the margin of the brain- case. Anteriorly the dorsal surface terminates in a thin irregular edge beneath the anterior edge of the "parietals." 6 bulletin: museum of comparative zoology Occipital Region (Figs. 1, 2, 4, 5). This may be defined as the constricted portion of the braincase lying posterior to the foramina for nerve X, containing the posterior portion of the brain cavity and notochordal canal, offer- ing points of articulation with the vertebral column and, laterally, affording areas of attachment for the most anterior myomeres. Centrally situated on the posterior aspect is the comparatively small foramen magnum. Above, a median ridge, separating right and left portions of the axial musculature, leads upward to the dorsal shelf described earlier. Below is seen the posterior opening of a large canal which runs forward beneath the cranial cavity. This cavity has been generally believed to house an unconstricted anterior portion of the notochord. More recently Stensio (1932) has suggested that it housed a set of muscles which pulled the anterior segment of the braincase downward on the otico-occipital segment. The present material shows, I think conclusivelv, that the earlier idea is the correct one. The notochord in the vertebral region of Megalichthys was surrounded laterally by bony central elements whose diameter was similar to that of the canal now discussed. Although in situ articulation of vertebral column and skull is not seen in my material, it is certain that the most anterior central elements fitted upon the posterior borders of the canal. There is no trace of, or space for a notochordal pit above the canal and below the endocranial cavity and an abrupt anterior ter- minus for a large notochord is quite unknown in any vertebrate. It seems obvious that the notochord continued forward through this subcranial canal ; further evidence for this view will be given later. A conspicuous feature of the sides of the occipital region is the pres- ence of a series of dorsoventral ridges which divide the surface into three antero-posterior segments. It seems certain that these represent the imprint of three successive myomeres, and suggest the incor- poration into the skull of the corresponding skeletal materials. The forward and upward slant of the posterior end of the occiput, however, indicates that there has not been a complete incorporation of the skeletal materials corresponding to the third myomere; presumably some sort of pro-atlantal structure may have been present dorsally. The presence of three occipital myomeres agrees with the condition in Ceratodus (Greil, 1913) while the other living lungfish have three somites in the embryo but only two myomeres persist. Two to three myomeres appear in general in the development of amphibians. The presence of three occipital myomeres may well ha\'e been a characteris- romer: braincask of iMEGALICHTHYS Fig. 1. Dorsal view of the Megalichthys braincase, x 3/2. On the right side a thin sheet of bone overlying the supraotic fossa has been removed. 8 bulletin: museum of comparative zoology tic of the common ancestors of lungfish, crossopterygians and tetra- pods. Within the area of attachment of each of the first two myomeres is seen the opening of a small canal. These canals, as noted elsewhere, were for nerves which seem highly comparable to the two hypoglossal roots found in many amniotes, and the term hypoglossals may, I think, be applied to them, rather than the terminology applied to the variable occipital nerve elements of other fish groups. No such nerve is found associated with the third muscle segment; its nerve presumably emerged posterior to the foramen magnum. Between the areas of the second and third muscle segments is found the external opening of a canal which passes up within the substance of the bone from the ventral aortic groove. This obviously carried an intersegmental artery. From its upper opening varying radiating grooves indicate its branches. No such canal is found between the first and second segments; presumably this single artery carried the entire blood supply to the occipital area. The great depth of the brain case in the occipital region is, of course, due to the presence of the huge notochordal canal; this extends for- ward a distance beneath the otic region; in side view a portion of the lateral wall of its anterior end is visible. The ventral surface of the occipital region is nearly flat. Just inside the lateral margins is seen a pair of deep grooves, diverging anteriorly and terminating at about the junction of occipital and otic regions. These grooves obviously carried the lateral aortae. The abrupt an- terior termination of the canals indicates that at this point the aortae, followed forward, curved sharply downwardly and somewhat laterally. Otic region, dorsal aspect Seen dorsally (fig. 1) the otic region extends outward from the main stem of the braincase as a triangular structure, with a median base and the apex at a prominent lateral projection. This has been termed the parotic process, and the name may be retained for des- criptive purposes, although it is by no means certain that it is homo- logous with all of the processes so named in other vertebrate groups. The otic region is divided above into two areas by a series of struc- tures running inward from the parotic process. The most medial of these structures is the ridge, noted earlier, which surrounds the most antero-lateral portion of the anterior vertical semicircular canal; this ridge is fused to the overlying dermal bones. Halfway out (the canal romer: braincase of megaliciithys 9 Fig. 2. (right) Lateral view of the braincase. Fig. 3. (left). Longi- tudinal section. Posteriorly the section is sagittal; anterior to the lines A-A it follows the left olfactory tract to the nasal capsule. Both x 3/2. 10 bulletin: museum of comparative zoology here having curved far ventrally) the ridge ceases, and the fossae anterior and posterior to the ridge are in communication through a large fenestra. I know of no important structure which might have passed through this fenestra, although, for example, there may have been some small artery which may have been the predecessor of the important (pseudo) temporal artery of the anurans. The fenestra was bridged above not only by the dermal roof but also, apparently, by a thin film of bone pertaining to the braincase itself. Distal to the fenestra the braincase again rises to form a dorsal buttress to the parotic process. This buttress is again fused to the dermal roof, in the "supra-temporal" region. Although I have not completed a study of the dermal elements, it would appear that the dermal shoulder girdle attached near this point. If this be true, it may be concluded that the overlying area of the dermal roof cor- responds in general to the tabular region of the tetrapod skull and that this region of the crossopterygian braincase is homologous with part, at least, of that portion of the otic region which in primitive amphibians lies beneath the tabular area. This process is pierced by a small foramen, lying beneath the course of the lateral line canal. It was probably traversed by the hypotic ramus of the facial nerve, which supplied this canal. Posterior to the series of structures just described lies a large fossa, roofed by a thin sheet of bone. Postero-medially it is bounded by a ridge running outward to cover the distal part of the posterior vertical semicircular canal. Since the fossa overlies most of the internal ear it may be termed the supraotic fossa. It opens out postero-laterally, the ventral margin of the opening being formed by a curved ridge which covers the arc of the horizontal semicircular canal and terminates antero-laterally at the dorsal parotic buttress. Half way along the course of this ridge is a small but well marked tubercle which may have served as a point of muscular attachment. Within the lateral portion of the floor of the fossa at least one spec- imen shows a faint groove curving backward and inward from the small foramen which presumably carried the hypotic ramus of nerve VII. It presumably follows the course of the ramus communicans VII — X found in many fishes. I know of no important structures which might have occupied this fossa. Opisthotic region (Figs. 2, 4) The lateral apex of the parotic process is formed by a dorso-ventral ridge which separates the lateral aspect of the otic romer: braincase of megalichthys 11 Fig. 4. Ventral view of the braincase. x 3/2. 12 bulletin: museum of comparative zoology region into an anterior temporal or prootic region facing rather an- teriorly, and a posterior region, facing as much posteriorly as laterally. This latter area, here described, corresponds roughly to the opisthotic region of tetrapods. It is bounded anteriorly by the ridge just men- tioned, dorsally by the sharp ridge below the supraotic fossa, ventrally by a sharp line of demarcation setting off the under surface of the braincase, and posteriorly by a vertical line beyond which the brain- case turns abruptly back into the occipital region. Part way down the posterior side of the vertical ridge of the parotic process" is the pos- terior opening of a large canal, obviously the jugular canal. In addi- tion to carrying the vena capitis lateralis, the sections show that the canal carrying the hyomandibular trunk of nerve VII opens into the bottom of the jugular canal and that this nerve also must have emerged to the surface here. While there is no positive proof, compar- isons make it certain that the orbital or "external carotoid" artery, homologous with the temporal or stapedial of higher forms, passed forward through the same opening. Dorsal to this opening is a depressed area, lacking a perichondral bony surface, and obviously articular in nature. Ventral to the canal is another large area of like nature. As noted below, these are the points of attachment of the hyomandibular. Farther back along the ventral margin is a third articular area, probably for the first branchial arch. Posterior to the jugular canal the course of the vena capitis lateralis is well defined by a depression bounded dorsally and ventrally by ridges. This groove terminates well back along the opisthotic surface ; here the contours indicate that the vein turned somewhat laterally and ventrally on its way to the ductus cuvierii. Overhanging the posterior end of the venous groove is a massive process, the tip of which was frequently unossified. It was obviously an important point of attachment. The sectioned specimen indicates an articulation with a posterior member of the branchial arch series. Although I do not feel competent at the present time to discuss this matter, this process suggests analogy to and possibly homology with the "paroccipital process" of primitive reptiles, and this term may be provisionally used for the process under discussion. The ridge bordering the dorsal margin of the channel for the vena capitis lateralis commences posteriorly at the "paroccipital process" and runs somewhat ventrally as well as anteriorly. Possibly it may have formed a point of attachment for the opercular muscles (cf. Griel 1913). Above it a rounded groove runs backward and upward between the "paroccipital process" and the posterior rim of the supra- romer: braincase of megalichthys 13 otic fossa. This groove may have carried a blood supply to the muscles of the area lateral to the occiput. The corresponding ventral ridge, on the other hand, fades out posteriorly; anteriorly it curves downward sharply behind the ventral hyomandibular articulation. Anterior to the ridge there is a small area between it and the ventral hyomandibular articular area through which the orbital artery may have emerged from the ventral surface of the braincase. This is an appropriate place for the passage of this artery. The ventral ridge pos- terior to this position prevents the consideration of a more posterior niatp ^^^ cot2n ms Fig. x3/2. 5. Posterior view of the otico-occipital portion of the braincase. course. It is highly reasonable to expect this artery to originate in the region of the hyoid arch ; in some fishes, for example, it actually arises from the upper end of the second arch rather than from the dorsal aorta. Between the ridge and the posterior ventral articular area there is a deep but smoothly rounded groove, the function of which I do not know. Possibly it marks the course of a hyoid vein. At the ventral margin of the posterior end of the venous groove is found a foramen which is seen in the sectioned specimen to lead out of the cavity of the internal ear, well posteriorly and at about the point of junction of saccular and utricular regions. As noted later, I believe this to be the external opening of nerve IX, with which may have been associated an incipient fenestra rotunda. Along the ventral margin of the lateral opishtotic surface are two small tubers; coinparison with Ceratodus suggests that they afforded origin for branchial arch levator muscles. 14 bulletin: museum of comparative zoology At the posterior margin of the opisthotie region are found two foramina which mark the external openings of canals leading from the endocranial cavity. One is situated behind the base of the ridge en- closing the posterior vertical canal. The second is situated more ventrally and posteriorly behind the paroccipital process; it is the larger of the two. It appears certain that they represent the points of exit of the nerves of the vagus group. I think it probable, as noted in the discussion of the brain, that the upper carried the posterior lat- eralis nerve, the lower the remaining components of the vagus. The lower in addition may have included a vena cerebralis posterior, although the latter cannot have been of any great size. Both foramina may vary in showing a small accessory opening, somewhat dorsal to the main one or an intermediate condition in which this is represented by a notch in the margin of the main opening. These smaller openings presumably were for dorsal rami. The anterior opening of the notochordal canal is found centrally beneath the opisthotie region. Laterally there is a broad expanse of bone underlying the saccular region; sections show part of this bony layer to be quite thin. About opposite the parotic process there is a low transverse ridge facing rather anteriorly and probably forming a base for ligaments which would have attached anteriorly to the noto- chordal sheath or posterior margins of the parasphenoid, thus strength- ening the union of the two halves of the braincase. Presumably these ligaments were derived from the same superficial membranous sheet in which, in other fishes and in tetrapods, the posterior portion of the parasphenoid ossifies. Structural features of the hyoviandibular region Although I have not completed my study of the visceral skeleton, the tentative restoration of the hyomandibular is of interest. This large element was found close beside the parotic region in the sec- tioned specimen, and actually articulated on one side. The head of the bone is divided into two subcircular articular areas which fitted the two corresponding depressions on the parotic crest. The two are connected anteriorly by a thin crest of bone which was in line with the ridge terminating the parotic process ; the anterior surface of the hyo- mandibular appears to have continued the plane of the anterior surface of the process, running parallel to the adjacent palatoquadrate. Between the two heads is the proximal end of a canal which ran outward along the shaft and emerged distally on the dorso-external romer: braincase of isiegalichthys 15 surface of the bone. One is tempted to suggest that this is the opening for a "stapedial" (here orbital) artery, as in the tetrapod stapes. But the external opening of the canal is placed too far distally for this to have been the case. Obviously it was a canal for the hyomandibular nerve, as in actinopterygians. The relations of the hyomandibular to the otic region and the jugular canal tend to shed light on several interesting morphological problems. The contrasting attachment of the hyomandibular in existing fish groups — ventral to the vena capitis lateralis in elasmobranchs, dorsal to the vein and commonly to the nerve in actinopterygians— has been a source of puzzlement to many morphologists ; compare, for example, the discussion in Goodrich (1930 p. 416 ff.). Of the various suggestions made in this regard we may note three of interest. (1) Stensio has pointed out that a change in position of a single insertion might take place by the head travelling along the outside of the jugular canal. (2) De Beer has suggested on theoretical ground that both dorsal and ventral attachments were primitively present. (3) Goodrich suggests that the presence of a canal in the hyomandibular of holosteans and at least some palaeoniscids may aid in accounting for the varied position of the nerve in forms which lack this opening. The situation seen in Megalichthys indicates that all three of these suggestions may be essentially correct, if we may (not unreasonably, I think) assume that the conditions seen here are really primitive. The hyomandibular attachment is along the edge of the jugular canal and migration of the attachment would be readily possible. In the attach- ments of the Megalichthys element we have a concrete example of DeBeer's theoretical double dorsal and ventral attachment. Loss of the dorsal head could give the elasmobranch relation to the vein, loss of the ventral head the actinopterygian condition. The presence here, as in primitive actinopterygians, of a perforation of the bone for the nerve suggests that this canal may have been general in early fishes. The canal enters the bone proximally between the two heads; loss of either head would make the disappearance of the canal a relatively simple matter. Of interest is the light shed by the Megalichthys hyomandibular on the development of the stapes (or columnella, s.l.) of tetrapods. The morphology of the tetrapod middle ear has been thoroughly and clearly' summarized by Goodrich (1930, Ch. VIII). The most important features are: (1) the course of the hyoman- dibular nerve or its equivalent backward over and down l)ehind the 16 bulletin: museum of comparative zoology stapes; (2) a dorsal process abutting against the otic region and above the nerve and lateral head vein (the process well developed in reptiles, degenerate in mammals); (3) the stapedial or temporal ( = orbital) artery running upward through an opening in the stapes (in mammals; in most reptiles the columnella is single headed, and the artery loops up and over it from below and behind). In the stapes of the early reptile Captorhimis recently described by Price (1935) these features are clearly seen. The basal portion is per- forated for the stapedial artery, and there is a well developed dorsal process. In contrast with Megalichthys there is no nerve foramen. This substitution of openings is readily correlated with the develop- ment of the foot plate. The primitive orbital artery curved up close beside the ventral hyomandibular articulation. With the expansion of the foot plate, the ventral head of the bone would tend to become stouter and include the artery within its substance. Wider separation of the two heads, for functional "reasons," would tend to eliminate the bony tissue between them and release the nerve from its canal. Temporal Region (Figs. 2, 4). The lateral portion of the braincase anterior to the "parotic process" corresponds closely to the area of the prootic ossifi- cation of primitive tetrapods. The lateral surface of this "temporal" region faces nearly as much anteriorly as laterally. The median boundary is formed by the wall of the braincase proper, the posterior boundary by a line running outward to the parotic process. The sur- face is essentially concave both in dorso-ventral view and in vertical section. The anterior opening of the jugular canal lies at the ventro-posterior corner of the temporal area. Its opening is in great measure concealed from view by the presence of a high thin shelf which runs from this point forward almost to the anterior end of the bone. This is essentially the posterior part of the subocular shelf of De Beer (1926), although the term is not appropriate here. Within this shelf is a long, well- marked trough which obviously was occupied by the vena capitis lateralis running backward from the orbital region to the jugular canal. Beneath the trigeminal region a large canal opens into this trough from behind. This issues from a space, noted later, beneath the brain cavity and indicates the point of emergence of the middle cerebral vein, here following the original course of the vena capitis medialis, although with an opposite direction of flow. The size of its opening romer: braincase of megalichthys 17 suggests that this vein drained the major part of the endocranial cavity. Above the anterior opening of the juguhir canal there is a concave area on the anterior face of the parotic process which is partially roofed by the expanded dorsal end of this process and in some cases partially separated ventrally from the jugular vein by a thin bridge of bone extending in from the lateral shelf of the jugular trough (this process is absent in the sectioned specimen). The spir- acular opening appears to have been situated just lateral and anterior to the summit of the parotic process and it is reasonable to assume tiiat the pocket formed here was occupied by a spiracular sense organ after the fashion of living dipnoans. About a third of the way forward along the course of the jugular trough a small tubercle projects outward over the medial side. From this point a ridge leads up and back to a second tubercle a short dis- tance in front of the fenestra leading to the supraotic fossa. I do not know with what attachments these structures were concerned. Behind the ridge there opens postero-laterally the small canal which ob- viously carried the hypotic ramus of the facialis. From its opening a smooth triangular area indicates the course of branches of this nerve outward to the spiracular organ and backward and upward to the small foramen in the parotic crest leading back beneath the lateral line to the supraotic fossa. Two small openings, one a short distance anterior to the hypotic VII foramen, a second farther forward and higher, are the entrances for small canals which, as discussed below, appear to have carried veins which drained inward from the temporal fossa to the cerebellar region. Well anteriorly two larger foramina are found opening out above the jugular trough. The posterior one is the smaller, and is directed rather anteriorly and slightly dorsally from beneath an overhanging ridge. This may be interpreted as an opening for the lateralis elements of the facial nerve. The more anterior opening is larger and faces more laterally. This presumably carried V2 and V3. Anterior to this foramen a ridge descends from a lateral projection of the dorsal surface which has been seen in the dorsal view. A thinner bony area in front of this ridge terminates the lateral surface of the otico-occipital. In front is a gap between the two halves of the braincase. This afforded a point of exit for the profundus and very likely nerve VI as well. A series of ridges along the outer face of the lateral boundary of the jugular trough may have given attachment to fascia or musculature. They mark the lateral boundary of the somewhat concave ventral 18 bulletin: museum of comparative zoology surface of the prootic region. On this surface well laterally is found the forwardly directed opening for the palatine branch of the facial. In the temporal region are found the elements which in actinop- terygians make up the typical trigemino-facialis chamber of Allis' descriptions (1919 etc.). They are, however, here widely dispersed, and cannot be said to form a chamber in any sense. The lateral wall of the actinopterygian chamber is confined here to the outer margin of the jugular trough and the far-laterally placed outer wall of the jugular canal. The pars ganglionaris (recess) is well separated from the diffuse area representing the pars jugularis. As noted earlier the notochordal canal opens out ventrally far back in the otic region, and continues forward as an open channel; there is no ventral connection between the two prootic regions, although the median walls swing down ventrally so as to cover a part of the sides of the canal. These walls are somewhat imperfect, even in the sec- tioned specimen, and I cannot be sure that the outline as figured is correct. As Stensio has noted, this ventral opening in itself is not a true (posterior) basicranial fenestra; it is merely an opening into the notochordal canal. Between the canal and brain cavity a partition persists forward to a point somewhat anterior to the level of the parotic processes. In front of this is a large true basicranial fenestra closed in front only by the posterior margin of the anterior segment of the braincase. Eihmo-sphenoid, dorsal aspect (Fig, 1) The anterior moiety of the braincase is narrow behind, where it lies beneath the median portion of the "frontals": anteriorly it expands rapidly, underlies the elements of the rostral shield, sends out marked antorbital processes, and terminates anteriorly in widely separated nasal capsules and a slightly projecting rostral region. It is solidly fused to the overlying elements (it is, however, only loosely attached to the premaxillae). In consequence this dorsal surface is drawn from reconstructions of the sectioned specimen and the finer details are imcertain. In general the dorsal surface exhibits a gentle slope downward and forward, with a slight lateral downward curvature in the antorbital and nasal region and a slight concavity in the middle of the ethmoid region. The pineal, although not penetrating the dermal roof, reaches the roof of the braincase beneath the "frontals." Grooves underlying the romer: braincase of megalichthys 19 thickened area of the elements bearing the supraorbital lateral line canals run forward and laterally over the antorbital region, gradually fading out distally. Several openings lead upward into this groove from the vmderlying lateralis nerve, and there are a number of open- ings presumably for vessels leading to the centers of ossification of the overlying dermal elements; as noted below, their restoration is none too certain and I have probably missed several of these small canals. Parasphenoid, basilar articulation and related structures (Figs. 2, 4) Although a dermal bone, the parasphenoid is, as usual so integral an element of the braincase complex and so difficult to separate from it, even arbitrarily, that it must be considered here. Along much of the ventral mid line of the ethmosphenoid the paras- phenoid bears an elongate oval plate thickly studded with small but sharp conical teeth. The anterior termination of this tooth-bearing plate projects well below the level of the braincase, with which it is connected by a longitudinal ridge composed of dermal bone. An- teriorly the parasphenoid is superficially in contact with the "vomers"; but a thin radiating sheet of bone which appears to be continuous with the parasphenoid spreads far forward over the lower surface of the inter-nasal area of the braincase. Posteriorly a flange develops along the sides of the parasphenoidal rostrum, formed ventrally of dermal bone and dorsally of cartilage replacement bone. The median edge of the palatoquadrate complex fits onto the upper surface of this flange so that (as in primitive amphibians and even Scymouria) there is practically no interpterygoid vacuity. Posteriorly the flanges flare out widely, and the postero- ventral termination of this half of the braincase has the section of a hemi-cylinder, the core composed of endochondral bone, the surface of parasphenoidal tissue which gradually fades out far dorsally along the posterior end of the sphenoid region. On either side two slit-like openings penetrate the parasphenoid, curving dow^n and in toward the midline. Here within the substance of the parasphenoid the two channels communicate with each other and with the floor of the pituitary fossa through a network of sinuses difficult to interpret in section. It seems certain that the posterior opening is that of the internal carotid. The anterior one obviously is the homologue of the tetrapod palatine artery. It cannot be considered as the ophthalmic (in the sense of Goodrich and De Beer) since it is ventral to the trabecular region. Radiating lines indicate the branch- 20 bulletin: museum of comparative zoology ing of the palatine external to its opening. Surface markings running down to the carotid opening from far up around the sides of the pos- terior end of the sphenoid region indicate that more posteriorly the carotid was situated at a comparatively high level and lateral to the expanded anterior end of the notochord. The broad posterior portion of the lateral flanges of the paras- phenoid curve widely upward and outward to encircle on either side a stout process formed of endochondral bone; the circular outer faces of these processes are turned somewhat anteriorly as well as laterally. These articulated movably with the palatoquadrate complex and are highly comparable with the basipterygoid processes of primitive rep- tiles and amphibians. It is frequently assumed that the basilar connection between brain- case and primary upper jaw was primitively a connection between the cartilages or replacing bony elements of these structures, and that the presence of dermal bones in this articulation represents a secondary condition. But it will be noted that the present process contains not only an endochondral core, but a dermal outer layer. The connection is already a compoimd one, and this double participation in the basal articulation is ob\iously an ancient character. Jnicrorhital Region The ethmo-sphenoid terminates posteriorly in a series of structures discussed in a later section: ventrally the concavity for the anterior end of the notochord; half way up a pair of posteriorly directed articular processes; dorsally a similar pair. Between these two sets of articu- lations the lateral margin is formed by a rounded pillar of bone. This has been termed the "alisphenoid wulst" by Stensio in comparison with a seemingly similar region of the actinopterygian skull. The term, however, is misleading, since in neither case are we dealing with a homologue of the true alisphenoid region of mammals. The term laterosphenoid pillar seems more appropriate. There is little lateral development of this region and no apparent evidence dorsally of the palato-quadrate articulation found here in coelacanths. In relation to this fact, the pillar is not traversed by a canal for the superficial ophthalmic, nor is there any opening for the profundus, which surely emerged behind the pillar and passed forward lateral to it. Anterior to the laterosphenoid pillar, the dorsal half of the lateral wall of the braincase is a fairly smooth sheet of bone, somewhat con- romer: braincase of megalichthys 21 cave above, somewhat expanded below over the side of the mid- and forebrain regions. A foramen for nerve IV should be present here, but I am unable to find it. A tiny foramen of this sort is easy to miss in section, and my material does not show the surface at all perfectly in this region. A shallow groove extends backward along the lateral margin of the sphenoid region from the top of the basipterygoid process. This obviously marks the position of the anterior end of the vena capitis lateralis on its way to enter the trough in the temporal region. This groove descends anteriorly into a deep pocket partially concealed by the basipterygoid process. This presumably was occupied, in part at least, by the posterior end of an infra-orbital sinus. A large foramen leads from this pocket into the pituitary fossa and was obviously traversed by the pituitary vein. I can find no opening for the ophthal- mic vein which in most fishes passes outward dorsal to the trabeculae and well posterior to the orbit. It it existed it must have used this same opening. The large optic foramen emerges on the anterior side of a prominence which may have served as the point of origin for the rectus muscles. To my embarrassment I have been ixnable to discover, either in sec- tions or whole specimens, an oculomotor foramen. It is obvious that there is not the slightest development of the myodome characteristic of the actinopterygians. The muscles con- cerned are well forward of their assumed point of entrance into the skull via the pituitary vein channel. Ethmoid Region The more anterior regions of the braincase are best seen in ventral view (fig. 4). In the mid-ventral area the lateral margins of the flanges of the parasphenoid region curve out anteriorly, leaving a broad and rather flat subrostral area between them: as noted pre- viously much of this area appears to have a film of parasphenoidal bone on its surface. In the midline is a groove which may have re- ceived processes from the dorsal surface of the medial edges of the "vomers." At the ventral anterior margin is a depression which lodged the palatal processes of the premaxillae with their large teeth. Laterally are a pair of pockets which appear to have been open to the roof of the mouth between premaxillae and "vomers" and received the tips of a large pair of lower jaw teeth. On the dorsal surface of the braincase a pair of canals emerge 22 bulletin: museum of comparative zoology medial to the nasal capsules ; they presumably carried bloodvessels and nerves to the rostral region; the details of their distribution are variable and none too clear. The ventral part of the lateral ethmoidal surface, below the ex- pansion caused by the underlying olfactory tract, is occupied by a series of pockets, variable in their arrangement. Possibly they were for the most part occupied by glandular structures. Anteriorly, beneath the median side of the nasal capsule, the series terminates in a deeper cavity which, unlike the others, lacks an ossified surface layer. In the sectioned specimen this pocket is occupied by the anterior end of the palato-quadrate; this is obviously the ethmoidal articulation. It will be noted that from here back to the basal articulation the palato- quadrate complex is in practically continuous contact with the base of the braincase. If this condition is at all primitive, it is easy to see how intermediate types of articulation may have evolved. Dorsally the braincase sends out a thin antorbital process; the lateral ethmoidal region gradually thickens anteriorly to the nasal capsule. On the under side two canals enter this process. The more posterior and dorsal one surely carried the lateralis nerve; the more anterior one, entering at a lower level, presumably carried the pro- fundus and accompanying blood vessels forward to the nasal and snout region. The nasal capsules are widely separated, with a thick mass of bone between them. The olfactory canal enters the posterior portion of the median wall of the capsule by a large circular opening, not well seen in the figures. Anterior to it, and quite distinct from its opening, is an equally large circular pocket in the median wall, subdivided by ir- regular ridges. This presumably lodged a median diverticulum of the nasal cavity, possibly homologous with the tetrapod Jacobson's organ. In none of my specimens are the thin dorsal and lateral walls of the nasal capsule completely preserved, and the outlines of the opening of the capsule given in the figures are a composite of which I am none too confident. It is certain, however, that the capsule opened widely antero-laterally toward the external nostril, and ventrally to the choana; there was no bony solum nasi, although, of course, such a structure might have been developed in cartilage. Notochord; intra-cranial kinetics Some evidence has been previously cited to show that the great ventral canal beneath otico-occipital was occupied by the undiminished romer: brainxase of megalichthys 23 anterior end of the notochord. In median section (fig. 3) this canal and related structures may be clearly seen. The canal continues forward with perfectly smooth walls beneath the posterior portion of the otic region. Farther forward the ventral wall disappears; still farther forward the dorsal wall. But an open cavity of equal diameter con- tinues forward beneath the anterior end of the otico-occipital. An- teriorly this cavity terminates in a hemispherical pocket in the posterior end of the ethmo-sphenoid; the diameter of this pocket is almost exactly that of the canal, with which it is almost exactly in line. It is difficult to believe that these areas were occupied by any structure other than an undiminished anterior continuation of the notochord. Still further proof is afforded by the fact that (as seen in section) the anterior termination of the notochordal space is, in the fashion of primitive vertebrates, just posterior to the pituitary. A final proof is offered by the nature of the bony materials sur- rounding the notochordal space. The bone of the braincase consists in general of a lightly ossified endochondral reticulum covered in almost every case by a thin perichondral layer. In the lining of the noto- chordal canal, however, we find a third type of tissue, a dense bone of heavily fibrous nature. This layer terminates abruptly at the anterior end of the notochordal canal, dorsally as well as ventrally. This tissue reappears in the cavity which obviously received the anterior end of the notochord. It is found nowhere else in the skull. But it is found, again, as would be expected, in one other locality — the internal surface of the vertebral centra. In this tissue we are definitely dealing with an ossification in a fibrous sheath surrounding the notochord. We may then confidently restore the notochord in Megalichthys. In the posterior part of the otico-occipital it was tightly attached to the braincase, and it was again tightly bound anteriorly to the ethmo- sphenoid. Between, in the "prootic" region, it was unattached; its flexibility would tend to allow a certain degree of motion between the two portions of the braincase. Presumably this movement was re- stricted by a dorsal ligamentous union of "parietals" and "frontals" as well as more indirect lateral dermal connections. Further restric- tions were placed in this movement by two sets of articulations be- tween otico-occipital and ethmo-sphenoid moieties, best seen in the section, but also visible in part in other figures. One pair of connections lies lateral to the floor of the endocranial cavity. On the "prootic" the medial wall of the jugular trough is continued forward beneath the lateral gap between the two halves of the braincase, so as to overlap the adjacent area of the "sphenoid;" 24 bulletin: museum of comparative zoology this projection is concave internally. Into the concavity fits a rounded, horizontal ridge on the "sphenoid," which projects back somewhat 3zn I at sofen coijzn vap svom jca 2npal Fig. 6. Anterior view of the otico-occipital portion of the braincase. X 3/2. from the general posterior margin of the bone at the level of the floor of the braincase. The two opposed surfaces lack perichondral bony coverings; presumably they formed a closely knit joint. .■vnos2 bptp Vp2 pas etha Fig. 7. Posterior view of the ethmo-sphenoid portion of the braincase. x3/2. At the dorsal end of the pillar terminating the "prootic" anteriorly there is developed on either side a rounded depression which faces downward and outward. The upper end of each laterosphenoidal romer: braincase of megalichthys 25 pillar is expended into a rounded knob which faces upward and back- ward, fitting into the sockets on the posterior bone. The opposed sur- faces are smoothly finished by perichondral bone; there is little sug- gestion of any close connection, and conditions suggest that a bursa intervened. This whole set of characters, large notochord and two sets of articular surfaces, suggests, as other writers have noted, a kinetic mechanism adapted to lessening the jars which would otherwise be transmitted to the posterior half of the braincase in the seizing and biting of food by these highly predaceous fishes. Presumably the "normal" situation was one in which the anterior segment of the braincase was depressed and a gap present between the upper pair of articular surfaces. When the jaws were snapped on to the prey, the anterior end of the braincase, (as well as the primary and secondary jaws) would "give" iipward in the flexure of the notochord beneath the "prootic" region. The ethmo-sphenoid would swing upward with the lower pair of articulations as the pivot; the upper surfaces would swing together, but the shock would be buffered by interposed bursae. With release of pressure, the ventral ligaments would presumably pull the snout downward again without need of any strong musculature for the purpose. Endocranial Cavity The brain cavity is displayed in the longitudinal section and recon- structed "moulds" of the internal cavities are shown in figs. 8-10. The portion of the brain cavity enclosed in the otico-occipital ap- pears essentially to be that of the medulla and cerebellum. Pos- teriorly, in the region of the spinal medulla, it is a small tubular structure, Avhich gradually expands in breadth and increases in height toward the anterior end of the otico-occipital. The broad medullary floor is widely open below through the large basicranial fenestra. In the central part of this opening the brain was presumably separated from the underlying notochord only by its meninges. Laterally in this area, however, appear to have been the principal points of exit of venous blood from the brain. Beaneth the lateral walls of the fenestra there is found on either side a large pocket which presumably contained a venous sinus. Outward from this pocket runs a large canal which turns forward and then upward, carrying the middle cerebral vein into the vena capitis lateralis. From the lateral margin of the floor beneath the vagus region a 26 bulletin: museum of comparative zoology canal descends into the substance of the bone lateral to the noto- chordal canal. Here it branches and ends blindly. It is surely a nutrient vessel. Presumably other such vessels are present, but they are not well enough seen in section to be restored with confidence. The dorsal surface of the posterior portion of the cavity appears to show a step-wise anterior increase in the height of the brain. A first upward step in the region of nerve X presumably gave the medullary walls their full height ; a second step presumably allowed the develop- ment of a posterior chorioid plexus, giving way anteriorly to the cerebellum. Farther forward on the dorsal surface is found a complex system of grooves and small canals. On both sides, above the roots of the facial nerve, is a curved longitudinal groove, from either end of which a small canal leads outward to the temporal region, where the external foramina have been noted. These are too far dorsal to have carried nervous structures. I interpret them as carrying veins inward from the surface to the intra-cranial system. O'Donaghue (1920) has noted the presence of similar vessels in Sphenodon and as he notes, Watson finds comparable openings in a number of fossil tetrapod types. Possibly similar structures would be found rather generally if search were made for them. Still farther forward, behind a transverse ridge in the roof which may mark the anterior end of the cerebellum, small canals are seen to enter the roof, running dorsally in an anterior or antero-lateral direction. I have found them difficult to trace. It is probable that they reach the region of the dorsal articulation between the two halves of the braincase and supply this area with blood vessels. Lateral openings from the walls of the medullary cavity are numer- ous. Posteriorly two small canals run out laterally from the lower margin of the medulla. These appear be "occipital" nerves. Their ventral orgin indicates that they are surely somatic motor in nature, and despite their distinct external openings I feel no hesitation in terming them hypoglossals. More anterior and more dorsal in position, are a pair of large canals which constitute the X-XI complex; their external openings have been noted earlier. The more dorsal (and externally more anterior) of the two arises from a groove high up on the lateral face of the brain- case; the more ventral from a larger pocket beneath this. While I feel none too sure, its position suggests that the more dorsal one was occupied by the posterior lateralis nerve, the groove leading to it indicating its roots from the lateralis tract of the medulla. Presumably romer: braincase of megalichthys 27 niatp X Fig. 8. Endocranial cavities in dorsal view, x 3/2. Zb bulletin: museum of comparative zoology the ventral opening carried the other vagal elements, and in all prob- ability a small posterior cerebral vein. Leaving aside for the time openings into the otic capsule (including nerve IX) there remain for consideration a series of openings pertain- ing to nerves V-VII-VIII, lying along the inner face of the prootic wall and constituting essentially the acustico-trigemino-facialis recess of Allis (1919). The most posterior of these is a large canal which runs out laterally into the bone of the prootic region. It soon divides. The larger posterior branch, evidently carrying the entire acoustic nerve, enters the otic capsule. The smaller division, obviously for the main stem of the facialis, runs on laterally just anterior to the auditory capsule, from which it is not completely separated. The main, hyomandibular, ramus runs outward to emerge in the floor of the jugular canal which it undoubtedly traversed to its posterior opening. Proximal to this point a small canal, for the palatine ramus, turns sharply forward to open on the ventral surface beneath the jugular trough. Next anterior, much smaller and more dorsally situated is a canal for the hypotic ramus of the facial. This runs outward just beneath the surface of the temporal fossa to open above the jugular trough half way to the parotic process. Its probable further course has been noted previously. Third in the series, also rather dorsally situated, is a fairly large opening which pierces the lateral wall, here thin, in an anterior as well as lateral direction. From its position and direction it presumably carried the anterior lateralis elements (ophthalmicus superficialis and buccalis VII). Next anterior, somewhat more ventral, and larger is an outwardly directed opening in the wall which I interpret as carrying the tri- gemis proper (maxillary and mandibular). Within the foramen is a small branch which runs up dorso-anteriorly in the lateral wall of the vertical pillar found here. This may have transmitted a small sensory branch to the joint region of the braincase. There is a well-marked oval lateral gap between the two halves of the braincase. Since there is no opening anterior to this for the pro- fundus, it must have left the endocranial cavity at this appropriate point; possibly nerve VI as well. In contrast with the breadth of the medullary region, the more an- terior portions of the brain cavity are elongate but narrow. The mid- brain region is but little expanded. The diencephalic region is indi- cated by the large pineal diverticulum. romer: braincase of megalichthys 29 Fig. 9. Endocranial cavities in ventral view. x. 3/2. 30 bulletin: museum of comparative zoology The position of the pituitary is of interest. The "dorsum sellae" is not at all dorsal ; in contrast to the orthodox vertebrate situation, the floor of the brainease slopes down and forward rather gently into the infundibular region; on the other hand, there is a marked "dorsum" over the anterior part of the fossa. These conditions may be inter- preted in the light of the peculiarly large size of the anterior end of the notochord; the pituitary has been pushed forward into an unusual position. The ventral arterial openings and the lateral opening for the pituitary vein and possibly the ophthalmic artery have been noted earlier. On one side of the sectioned specimen a small channel runs upward from the front of the pituitary sac into the base of the brain cavity below the assumed position of the optic nerve. Presumably this carried the optic artery, the independent channel developing because of the unusual position of the pituitary. However I could not trace this channel through on the other side of the same specimen and it appears to be absent in a second. The optic foramina are far forward, terminating channels which gradually separate from the under side of the cavity of the forebrain. At a point above the middle of the pituitary fossa and just back of the tip of the pineal diverticulum the brain cavity divides into two, indicating the position of the lamina terminalis. Obviously there can have been but little development of an unpaired telencephalon. For- ward there extend from this point two markedly divergent tubes lead- ing to the nasal capsules. The tubes are of nearly uniform size through- out and there are few definite indications of subdivision into regions containing hemispheres, olfactory lobes and more distal olfactory "nerves," although I infer that a marked constriction about two- thirds of the way along each tube marks the end of the olfactory lobe. There are marked differences in the details of the walls of these tubes from specimen to specimen and even between the two sides of the same specimen. In one case, at least, there is a groove running along the median wall for a considerable distance beyond the point of bifurcation; presumably this was for a blood vessel. There is a number of small canals in the ethmoidal region, some of which are shown in the figures. I have found them hard to interpret because it is difficult in section to trace these tiny structures through the spongy bone, and a further difficulty lies in the fact that the sectioned specimen is somewhat imperfect on the left side; in the figures I have restored the missing areas on the basis of the right side. In consequence there are probably errors both of omission and com- mission. romer: braincase of megalichthys 31 Fig. 10. (right) Endocranical cavities in lateral view; the otic capsule removed. Fig. 11 (left). Attempted restoration of brain in side view. Both X 3/2. 32 bulletin: museum of comparative zoology From the "olfactory tubes" a number of small canals branch out to emerge on the surface of the braincase beneath the centers of over- lying dermal elements on the ethmoidal shield; presumably they carried nutrient vessels. Several diverge from a common pocket in the region which I assume to have contained the olfactory lobe. The large median "postrostral" is supplied from both sides. On the right side of the sectioned specimen a small canal splits off from the "olfactory tube" dorsally and runs forward to emerge into the capsule at its dorsal margin over the olfactory nerve. Presumably this carried blood vessels. The left side is imperfect and in the figures I have restored this region as a mirror image of the right side. How- ever in another specimen this separate canal was absent and condi- tions here appear to have been variable. In the lateral ethmoidal region a canal which runs forward and in- ward beneath the position of the lateral line obviously carried ophthal- micus superficialis VII. There are several branches upward from this canal to the cranial roof. It appears to anastomose with the vascular canal mentioned in the last paragraph. Probably it continued antei;o- medially from this point, but I have been unable to trace it satis- factorily. A second canal, farther forward on the lateral ethmoid region opens almost immediately into the dorso-posterior part of the nasal capsule. Presumably this carried the profundus nerve and associated blood vessels. A groove leading anteriorly and medially over the medial wall of the nasal capsule appears to mark the course of the median branch of the profundus to a canal which goes to a "foramen apicale" on the snout. Nervous system In figs. 11 and 12 I have attempted to restore the brain and portions of the cranial nerves. The data upon which the restoration is based have been given above, and I need not comment upon it in detail. Such features as, for example, the grooving of the inner surface of the braincase bv the vessels above the medulla, indicate that the brain rather fully filled the endocranial cavity. But the comparative thick- ness of my sections, together with minor imperfections in the endo- cranial walls, have rendered it difficult to find many details in contours which might have been significant in setting off the regions of the brain. The division between mid- and hind brain areas seems clearly marked, and the position of the various nerves and of epiphysis and romer: braincase of megalichthys 33 Fig. 12. Attempted restoration of the brain and proximal portions of the nerves in dorsal view. On the right side the ramus communicans VII-X is indicated, the obvious general external course of the ophthalmic nerves is shown in dotted line, and the deduced paths of the rami of nerve VIII are figured. X 3/2; 34 bulletin: museum of comparative zoology pituitary renders it certain that in most respects the divisions indicated cannot be far from correct. As noted above, the forebrain region presents the greatest difficulties. In general I have restricted the restoration of nerves to those por- tions found enclosed in bone; however, I have indicated the obvious general external course of the ophthalmics and hypotic ramus of VII. On the right side I have indicated, from data given below, the probable course of the subdivisions of the auditory nerve. Because of lack of satisfactory evidence I have omitted the eye muscle nerves. It is obvious that the brain as restored is essentially similar to that seen in dipnoans on the one hand and amphibians on the other; thus the neurological evidence, as far as it goes, agrees with all other lines of work tending to indicate the close relationship of crossopterygians with these two groups. Of particular interest is the fact that the fore- brain here is highly evaginated as in amphibians and lungfish, not merely Protoptcrus and Lepidosuren but Ceratodus as well (Holmgren and van der Horst 1925) and in contrast to the other fish groups (cf. Herrick 1921). The two hemispheres diverge markedly, instead of lying close beside each other as in both amphibians and lungfish. This may reasonably be considered a primitive condition on the assumption that the developing hemispheres invaded the canals diverging immediately from the old forebrain termination toward the olfactory capsules. The hemispheres lack the peculiar ventral expan- sions of the living dipnoans; the large lingula of the Ceratodus para- physis cannot have been developed. The large diameter of the pineal diverticulum suggests that "bolsters" containing the habenular ganglia may have been present as in Ceratodus. The comparatively small optic lobe region is in correlation with the small eyes of Megalichthys. Some of the details of the nerve openings, as the apparently distinct development of anterior and posterior lateralis nerves, suggests a rather primitive condition. Circidatory Systein In the description of the braincase a number of facts have been noted which shed light upon part of the cranial circulation. These may be summarized here. We have noted a groove on the under side of the occipital region which was undoubtedly occupied by the lateral aorta, and an occipital artery which passed up from this point to supply the area lateral to the posterior part of the skull. The aorta obviously was curving upward romer: braincase of megalichthys 35 and backward to enter the groove, suggesting that the arches at least as far back as the tetrapod systemic were placed anterior to the groove. The orbital artery has left no definite imprint, but its probable course up beside the ventral attachment of the hyomandibular and thence forward through the jugular canal is readily inferred. The forward course of the internal carotid must have been along the under side of the prootic wall, median to the palatine foramen, rising up alongside the lateral border of the anterior end of the notochord and then curving downward on the parasphenoid to enter its foramen through that bone to the floor of the pituitary fossa. We have noted the re-emergence from the parasphenoid of a palatine artery branch- ing from the carotid within that bone. It has been suggested that small foramina leading upward from the front end of the pituitary fossa indicate an occasional independent course for the intra-cranial part of the optic arteries. There is no positive evidence of the presence of an ophthalmic artery or the related pseudobranchial. In most respects the arterial circulation thus inferred agrees well with that of other fishes. In many cases the ventral vessels are more deeply embedded in the base of the braincase. The orbital artery here seems to be identical in its relations with that of most actinoptery- gians. In many selachians and Polypterus it is, however, entirely lateral to the less completely developed "temporal" region and, further, rises up antero-medial to the palatine nerve. In some rela- chians it is embedded in the braincase, but the relations to the pala- tine are the same; the foramen is a ventral one. On the other hand, there is nothing known which would prevent the development of the arterial system into that seen in the tetrapods. The development of the palatine artery, not normally present in fishes, is a distinct indication of dipnoan and tetrapod affinities. As I have noted earlier the course of the orbital is just that to be expected in a morphological predecessor of the temporal of lower tetrapods and the mammalian stapedial. As has been seen, the course of the vena capitis lateralis is clearly marked; from the pocket in the orbital region containing the pituitary vein backward over the basipterygoid process, along the trough in the lateral margin of the temporal region, through the jugular canal, and back along the outer side of the otic capsule to the level of the vagus. The vein is as usual ventral to the trigeminal, but dorsal to the pala- tine and hyomandibular divisions of the facial. I have found no anterior cerebral vein, and the posterior cannot have been large. The main drainage from the skull was surely through the middle cerebral. 36 bulletin: museum of comparative zoology as described; its course lies beneath the facial, in proper position for the embryonic vena capitis medialis. The conditions are in harmony with those found in fishes in general. The prominence of the middle cerebral agrees with conditions in amphibians. In addition to major vessels we have noted various small vascular canals presumably of nutrient nature and small veins draining the temporal and internal ear regions. Internal Ear The otic capsule is well ossified internally, and the outlines of the main structures are clearly indicated (Figs. 8, 9, 13). The otic region lies beneath the supraotic fossa and the areas immediately adjacent to it; as has been noted, portions of the canals are enclosed in ridges bounding this fossa. The ventral surface of the ear cavity is separated from the under surface of the otic region by a thin sheet of bone. The ear region is remarkably large by comparison with most other vertebrate types. It is able to gain in depth without projecting ventrally owing to the development of the large notochordal canal; the saccular area is lateral to the canal and extends far below the level of the brain cavity. The main sacculo-utricular cavity may be de- scribed as having the shape of a half-filled sack, with a broad base resting on the floor of the otic region, the upper part constricting as it rises inwardly and posteriorly to an apex in the region of the crus commune. Part way up the sack there is a constriction on the outer wall which appears to mark the division between saccular and utricular areas. The saccular area has a large subcircular and nearly flat floor, which tilts downward toward the outer and anterior margins. Postero- externally there is a depression in the floor which indicates the position of the lagena. In both sides of the sectioned specimen there is pre- served a large otolith which covers much of the floor of the saccular cavity. Over much of the surface it is a lens-shaped structure. In the lagenar region it sends down a ventral process which follows the con- tours of this depression. Along the back wall of the saccular cavity there is a slight out-pocketing suggesting the presence of a papilla basilaris. There is no positive evidence of a papilla neglecta or amphibiorum, although presumably some structure of this sort must have been present. There is no evidence from the external contours concerning romer: braincase of megalichthys 37 an utricular recess; it certainly could not have attained the large size of the recess of dipnoans. A small utricular otolith was found on the left side of the sectioned specimen in the region of the crus commune. Presumably this is a post-mortem displacement from an originally more anterior position. aca vcl ccom D send smrp Fig. 13. The ear region of Megalichthys. A, B. sections through the left otic capsule, the cut being a vertical one made diagonally from a point just in front of the jugular canal postero-medially into the braincase in the vagus region. In A the view is in an antero-medial direction; in B, the reverse. C, D, reconstructions of the cavities of the left otic capsule as positive casts. C, lateral view; D, medial view. All x 3/2. Three semicircular canals are clearly outlined, occupying their nor- mal positions and without obvious peculiarities. Conjoined areas for the ampullae of the anterior and horizontal canals are seen anteriorly. The horizontal and posterior vertical canals have a common area of posterolateral origin from the end of a ridge which bounds the back border of the utricular area, the end of the horizontal canal lying over the assumed area of the ampulla of the posterior vertical. 38 bulletin: museum of comparative zoology For the most part the inner ear is separated from the brain cavity by stout bony walls. There is, however, a remnant of the embryonic con- nection here in the shape of a long slit which starts rather low an- teriorly behind the foramen for nerves VII-VIII and slants backward and upward, widening above, to terminate high up medial to the area of the cms commune. The lower part of this opening appears to have been closed in life by a thin film of bone, remains of which are seen in the sections. By comparison with the usual amphibian condition, it would be assumed that the posterior ramus of nerve VIII entered the capsule through this slit. But the course of the nerve can be better accounted for on the assumption given below. The upper part of the opening from the internal ear lacks any trace of a wall. This region extends somewhat over the roof of the brain cavity. Here there is developed a pocket which extends a short distance back into the substance of the cranial roof, which is as much a part of the endocranial as of the otic cavity. It may reasonably be assumed that this pocket represents the position of an endolymphatic sac or at least the posterior part of one. There is no evidence of any canal upward to the surface for an external endolymphatic opening. From this general dorsal terminal area of the otic region a small canal runs back into the base of the canal for the posterior lateralis nerve. It is very clearly seen ori the right side; on the left the region is imperfect. I know of no comparable structure. Presumably it carried a small vessel, vein or lymphatic, out from the otic region. I cannot account for it under any theory of origin of perilymphatic structures. The acoustic nerve enters the otic capsule at its anterior median corner, shortly after it separates from the trunk of the facial. It ap- pears to have divided into two portions almost immediately. One slants steeply upward along the anterior wall to presumably reach the ampullae of the horizontal and anterior vertical canals and the utricu- lar macula. The larger subdivision appears to have remained at a low level and enters the floor of the saccular area. In the sections there are low ridges which appear to radiate over the saccular floor and indicate the distribution of the nerves to the saccular and lagenar maculae; the possession of a common otolith suggests that these two sensory areas were more or less fused. A groove along the median wall of the saccular floor suggests the position of a posterior ramus which rose posteriorly to the posterior canal and the assumed maculae neglecta and basilaris. A foramen, noted previously, pierces the outer wall of the capsule at about the junction of saccular and utricular areas close to the postero- romer: braincase of megalichthys 39 lateral corner of the otic cavity. At the ventral postero-internal corner of the cavity a canal passes into the floor of the braincase anterior to the opening which I assume to have carried the non-lateralis elements of the vagus. The internal opening seems rather certainly to mark the position of the tetrapod perilymphatic connection with the endocranial cavity. The external opening appears to be the same as that in the coela- canth Diplocercides w^hich Stensio suggested might be a rudimentary fenestra ovalis. This can hardly be the case; it is far removed from the future stapes and far posterior in position. Its external opening is close beneath the course of the vena capitis lateralis. This suggests that it might have carried a vessel outward from the ear region, a condition unusual but not exceptional, since an analogous opening is found in the venous hypotic duct of some chelonians (Nick 1912). A possibility involving both these openings which I adopt, although with some hesitation, is that they transmitted the glossopharyngeal nerve (less its usual lateralis component), its course lying along the hind wall of the otic capsule between inner and outer openings. The nerve traverses the capsule in many vertebrates and the external opening is in an appropriate position. The low position of the internal opening into the brain cavity causes some difficulty. Current theories regarding the development of the tetrapod perilym- phatic openings imply that the perilymphatic system first penetrated internally into the base of a jugular foramen of amphibian type (con- taining IX and X as well as the posterior cerebral vein) and then spread outward along this canal to the surface of the braincase. The fenestra rotunda is assumed to have originated by splitting off from the jugular foramen and rotating forward. If the conditions in Megalichthys are representative of those of the tetrapod ancestor, it may well be that this is nearly the reverse of the truth. The external opening of the glossopharyngeal may have been a primitive fenestra rotunda in essentially its definitive position, the internal opening of the glossopharyngeal the primitive perilymphatic foramen. The varied amphibian conditions may be ascribed to a secondary withdrawal of nerve IX from the otic capsule, with consequent, Init varied "at- tempts" of the perilymphatic system to follow the nerve. General and evihryological considerations Before attempting to compare the Megalichthys braincase with that of other forms it seems advisable to attempt to resolve the adult braincase into its probable embryological antecedents (fig. 14). In 40 bulletin: museum of comparative zoology doing so I have in general followed the terminology and schemata of Goodrich (1930) and DeBeer (1926, etc.). The general position of the trabeculae seems clear. They obviously terminated posteriorly alongside the pituitary region and ran forward below the f orebrain region, below the optic foramen. If the para- sphenoid were removed, the pituitary fossa would still open ventrally to the roof of the mouth as the remains of the early fenestra hypo- physeos. The internal carotids enter this opening in typical fashion, below the trabeculae, the pituitary vein emerges above them. The basilar articulation may, as generally, be assumed to represent the approximate position of the polar cartilage. The broad anterior end of the adult braincase would at first sight suggest a typically platybasic condition of the embryo. But it will be noted that the sphenoidal region is quite narrow below. The tra- beculae must have lain quite close together in early stages. The condi- tion would thus have been an intermediate one, from which either platybasic or tropibasic conditions might easily have been derived. The transverse bar of bone which posteriorly terminates the ethmo- spheniod in its ventral half, is interposed between the pituitary and the tip of the notochord. The morphological importance of this area has been recently emphasized by DeBeer (1926). The present bar, the "dorsum sellae," is obviously a development of the acrochordal cartilage. Here, as DeBeer believes to be the case generally, the acrochordal is obviously quite distinct in origin from the parachordals, from which it is separated centrally by the posterior basicranial fenestra and laterally by the lower intra-cranial joint. I shall attempt no analysis of the more anterior supra-trabecular structures of the nasal and ethmoid regions, which appear to vary greatly from group to group. More posteriorly the upper part of the lateral wall of the braincase is obviously developed from the orbital or sphenolateral plate of the embryo. The taenia marginalis, which generally connects orbital and otic regions along the dorsal edge of the braincase is interrupted by the upper intra-cranial articulation. DeBeer (1926) has pointed out the importance of the pila prootica (or antotica), found generally in vertebrates except most actinop- terygians and mammals, as a landmark. It runs down from the posterior end of the orbital cartilage, posterior to nerves II and III and the pituitary vein, anterior to the prootic incisure or foramen and hence anterior to nerves V and VII (except that the profundus may be embedded in the pila). It generally extends upward the line of the dorsum sellae. romer: braincase of megalichthys 41 This is an exact definition of the region which I have termed the laterosphenoid pillar, bounding the ethmo-sphenoid posteriorly above the notochord. A pila lateralis of actinopterygian type should be outside this area, lateral to the head vein, profundus and anterior lateral line nerve. Obviously this structure is non-existent in Megalichthys. My uncertainty as to the course of nerve III renders the position of the pila metoptica uncertain. pan-t ^marg snlat Fig. 14. Attempted analysis of the embryonic components of the Megali- chthys braincase. Just as the trabeculae appear to form the essential structure of the ethmo-sphenoid half of the adult Megalichthys skull, so the para- chordals are obviously the structural base of the otico-occipital moiety. Posteriorly they have obviously fused both below the notochord and above it to form a basal plate; anteriorly they are widely separated by the persistent basicranial fenestra, through which the middle cerebral vein enters the braincase. Posteriorly the presence of two hypoglossal elements suggests that three arches have fused to form the occipital region. The general region of the embryonic otic capsule is obvious. The metotic fissure has, of course, closed except for the foramina of nerves IX and X. Dorsally otic and occipital structures have fused to form a solid roof to the braincase. As has been seen, the otic capsule still retains part of its original internal opening. Anterior to the otic capsule proper the "prootic" region has a com- plicated structure. Much of the upper margin of this region may be assumed to be formed by the posterior part of the taenia marginalis extending forward from the capsule; the lower by the anterior ends of the parachordals. Here lie a large number of openings and canals, the osseous divisions between which may be analyzed in terms of commissures. The prefacial commissure is by definition that part of the cranial wall lying between the facialis (proper) and the trigeminal (including 42 BULLETIN": MLTSEUM OF COMPARATIVE ZOOLOGY profundus) and lateralis VII. In Megalichthys the facial (excluding lateralis) extends far out laterally in the temporal region before emerg- ing by various exits. Hence the prefacial commissure in a broad sense includes much of the surface of the "temporal" fossa; its basic portion may be regarded as the area of the side wall of the braincase just back of the opening for the lateralis nerve. With the development of the prefacial commissure we have the embryonic prootic foramen divided into two major areas. In Me- galichthys we find both of these subdivided to an extent not seen, as far as I am aware, in any other type of fish (except the Polypterini). The anterior region is subdivided by two further commissures, one between the lateralis opening and the trigemenus proper, the second, terminating the lateral wall of the temporal region anteriorly, be- tween V2 & V3 and the lateral gap behind the pila antotica through which VI and VI presumably emerged. The three posterior components of the facialis (otic, hyomandibular and palatine) are embedded in a solid mass of bone continuous with the anterior wall of the otic capsule. It is difficult to analyse this mass in terms of commissures. The lateral part of the temporal wall may be in part the postpalatine commissure. The area of bone separating the otic ramus from the main trunk of the facial may be considered in theory as till another, unnamed, commissure. The most lateral part of the otic region, the outer face of the parotic process is, by definition, the lateral commissure of DeBeer. As has been seen above, the two moieties of the adult Megalichthys braincase correspond essentially to the two major regions of the em- bryonic vertebrate chrondrocranium ; the trabeculae and polar car- tilages and overlying structures anteriorly, and the parachordals and related elements posteriorly. Further, this adult subdivision appears to represent a basic distinction between two modes of origin of skeletal material. For both the observations of Miss Piatt and the experi- mental work of Stone indicate that the trabecular region is derived from the neural crest and thus related in embryonic origin to the visceral arches, while the parachordals and more posterior structures are derived from typical axial mesenchyme. Dr. W. K. Gregory has suggested to the writer that the condition in rhipidistians might be considered a retention of a primitive vertebrate character. This sug- gestion is a stimulating one and worthy of serious consideration. At present, however, the weight of paleontological evidence is in the other direction. For the time at least we must, I think, adopt the opposite hypothesis, that the intracranial joint and associated structures are a romer: braincase of megalichthys 43 specialization which has arisen within the group as a retention in the adult of an essentially embryonic condition. In an analysis of skull structure such as that of the earlier part of this section, the general procedure is to erect a simple framework and then fill out the edifice by the addition of commissures, taeniae, etc. This is essentially a description of actual embryological history. Im- plicit however in much of this type of discussion is the assumption that these processes are also a recapitulation of phylogeny; that the primitive gnathostome had a rather simple cranial structure which was gradually expanded and complicated in various later forms. But our increasing knowledge of early vertebrate history shows that in many cases groups have not "advanced" structurally, but have degenerated, as Watson, for example, has repeatedly shown. Further, as especially illustrated by the work of Stensio, the oldest vertebrates show no tendency to exhibit the theoretically simple structure of an "archetype." Existing evidence points strongly toward the point of view that the primitive gnathostome had a braincase of expanded and complicated structure. I shall return to this point in a concluding section. • Comparison with other crossopterygians Such knowledge as we possess of the braincase of other crossoptery- gians may be reviewed in the light of the structure seen fairly com- pletely in Megalichthys. A complete summary' of our earlier knowl- edge, with references to all earlier literature is given in a recent work by Holmgren and Stensio (1936) and detailed references are omitted here. Watson in 1926 figured a lateral view of the braincase of Osteolepis macrolepidotus. The general topography appears to be similar to that in Megalichthys, although the specimen is incomplete and many details could not be made out. The openings for nerves I, II and super- ficial ophthalmic VII are properly identified, as is the jugular canal. The general area of the hyomandibular attachment is indicated, but the details appear not to have been visible and only a single head is indicated. The opening marked "VII" is perhaps that for the otic ramus of that nerve. A foramen for nerve III is shown in a position which suggests the opening for the pituitary vein. The groove bound- ing the vena capitis lateralis anterior to the jugular canal is shown, but not identified. Stensio has published an emended copy of this figure. In it the pituitary vein is, I believe, correctly identified and a small foramen unlabelled by Watson is presumably correctly assigned 44 bulletin: museum of comparative zoology to the superior ophthalmic. Other changes made in the identifications appear to be less happy. The anterior opening of the jugular canal is suggested as the opening for the otic ramus of VII. In addition to the figure mentioned, Watson described a number of other features of the brain case of Osteolepis. With few exceptions this description could be applied word for word to Megalichthys. Bryant's pioneer work of 1919 on Eusthenopteron was of funda- mental importance in establishing the double nature of the braincase, but he was unable at that time to interpret its structure in any detail. Subsequently Stensio has figured the braincase of Eusthenopteron foordi in side view and has promised a future description of serial section work in progress on this genus. The side view as figured is com- parable in many respects to Megalichthys. Openings for the oculo- motor nerve and the arteria ophthalmica magna, which I have failed to find, are indicated. All three areas of attachment in the hyomandi- bular region are figured, but only the dorsal area is labelled. The antero-lateral region of the otico-occipital above the jugular trough and the area between the two moieties of the braincase are assumed to have been filled with a solid sheet of cartilage. This is improbable; since Eusthenopteron appears, as far as our knowledge goes, to resemble Megalichthys markedly in other features of the brain- case, it is probable that better material would show that the construc- tion here was the same. Except for a few scattered observations, I have cited all work done on rhipidistian crossopterygian braincase structure. The facts are scanty, but such as they are they suggest that Megalichthys may be taken as a typical member of the group. Of the more aberrant crossopterygians, the coelacanths, many of the forms described are Mesozoic types which are obviously specialized and degenerate and need not be discussed here. Of great interest, however, is the Devonian Diplocercides. This has a well ossified braincase which has been described by Stensio on several occasions; serial sections have been made, but only preliminary notes of the re- sults are as yet published. The ethmosphenoid of this coelacanth is in many ways comparable to that of Megalichthys. In the orbit, in addition to the openings found in the latter form, Stensio shows foramina for nerve III and the ophthalmica magna. A pronounced point of difference is the develop- ment of a large "antotical process" at the upper posterior border for an articulation with the ascending process of the palatoquadrate, an articulation which he considers a secondary one. The presence of this romer: braincase of megalichthys 45 process results in the backward extension of the furrow lying beneath the supraorbital lateral line canal, and in the enclosure of much of the superficial ophthalmic in a canal through the substance of this process. A seemingly important difference between conditions here and in Megalichthys and, it would seem, other rhipidistians, is the perforation of this process by a canal which Stensio interprets as transmitting the profundus nerve. This fact, presumably, has led him to the conclu- sion (193G p. 347) that the split in the braincase comes at the back end of the trigeminus opening and hence that in such a form as Diplo- cercides part of the otic region is included in the ethmosphenoid half of the braincase. But presumably the inclusion of the profundus in the "laterosphenoid pillar" or pila antotica is also related to the develop- ment of the upper palato-quadrate articulation; as De Beer (1926) notes, the profundus, although emerging from the braincase posterior to the pila antotica (prootica) may become enclosed in that element. In Stensio's specimen the occipital region is apparently missing. A single opening is indicated in the vagal region, rather than the two seen in Megalichthys. An external opening from the otic capsule com- parable to that seen in Megalichthys is interpreted as possibly a fen- estra ovalis; nerve IX is stated to emerge close beside but separately from the opening. A typical jugular canal seems to be present. It is assumed that the facial nerve emerged to the surface at about the point where the otic ramus emerges in Megalichthys and that the hyomandibular ramus then turns backward into the jugular canal. Possibly, however, the hyomandibular ramus may have followed its internal course as in rhipidistians. There appears to be no outer rim to the jugular furrow. Antero-laterally the otico-occipital is poorly ossified, for the area whence normally issue all branches of the trig- eminus, the anterior lateralis and the middle cerebral vein is repre- resented by a single large gap, uncrossed by any osseous subdivisions. Dictyonosteus articus is a second Devonian form described by Stensio; it is represented only by an imperfect ethmo-sphenoid. Its taxonomic position is somewhat uncertain, but as its describer early pointed out, it seems to be of coelacanthid affinities. As far as its structure has been interpreted it agrees well with Diplocercides . The earliest coelacanths appear to show a braincase structure very similar to that of their rhipidistian cousins; the only differences of im- portance appear to be associated with the presence in coelacanths of a highly developed articular area in the "antotic" region. The two groups have obviously departed but little from a common ancestral type. 46 bulletin: museum of comparative zoology Comparison with tetrapods The structure of the crossopterygian braincase is of major morpho- logical importance in relation to the evolution of the early tetrapods. Watson in 1926 made a number of interesting comparisons despite the unsatisfactory knowledge of crossopterygians which existed at that time. The comparison may now be carried further. The kinetic peculiarities of the crossopterygian braincase — intra- cranial joint and over-enlarged intra-cranial notochord — are generally assumed to have been absent in the tetrapod ancestors or, if ever present, had been lost before the tetrapod stage had been attained, although it is of interest that in many Paleozoic tetrapods there is a marked zone of weakness and reduction of bony connection between the two regions. In order to facilitate comparisons I have therefore, in fig. 15a, illustrated the Megalichthys braincase so modified that these features have been eliminated. The parasphenoid has been extended back to cover the ventral gap and it is assumed that cartilage lay beneath this area about a smaller notochord; the articular areas are assumed to be fused. If kineticism is secondary such a braincase might be fairly representative of that expected in a tetrapod ancestor and perhaps representative of the general stock from which lungfish, crossopterygians and tetrapods have descended. In comparing with tetrapods, we encounter difficulties in the fact that there are no living tetrapods which can be considered at all primitive in braincase structure. All living amphibians are highly modified in skull structure ; indeed even a lizard or Sphenodori is closer to primitive conditions in a number of features. For primitive condi- tions we must turn to the extinct labj-rinthodonts. Here we encounter further difficulty in the fact that much of our knowledge of them is derived from rachitomous and stereospondylous forms which, as shown by Watson (1919, etc.), already exhibit specializations similar to those seen in existing amphibians. We are forced to rely upon our knowledge of the carboniferous Embolomeri, as described mainly in Watson's important contribution of 1926. Even here we are hampered by the fact that available material is none too good and in consequence many features of importance are obscure and the internal structure of the braincase is almost unknown. If the crossopterygian and embolomerous braincases are com- pared (fig. 15a, c) a striking difference in proportionate size of parts is noticeable. If the basipterygoid articulation, for example, be used as a point of reference, it will be noted that in the crossopterygian this romer: braincase of megalichthys 47 structure is in front of the mid point of the braincase; in the amphibian it is back nearly three-fourths of the distance from snout to occiput. The same shift backward holds true for the position of the outlets for nerves II and V, the position of the pineal opening dorsally and, in- ternally, the position of the pituitary. It is obvious that in the amphibian there has been a great elongation of the ethmoid region (rather than the possible converse assumption of posterior shortening). This elongation and consequent disparity of proportions may be interpreted in terms of relative development of brains and jaws. The elongation of the "face" of early amphibians as contrasted with their piscine relatives is probably related in part to changed food habits and elongation of the jaws, with a necessary elongation of the braincase. In great measure, however, this elongation is probably related to size differences in the forms considered. Crossopterygians investigated are fishes of modest size; the Embolomeri whose brain- cases are known are mostly far larger. The results of change of size upon proportions of animals are so obvious that they are usually overlooked (but cf. Watson 1930). If, for example, an animal doubles in length, its necessary food intake is (roughly) cubed, and a disproportionate growth of mouth parts tends to result. On the other hand, it is obvious on functional grounds that brain and sense organs need to increase but little; large animals are proportionately small brained. The brain could not be stretched out to any extent and hence would come to be concentrated in the posterior portion of an elongated skull. There would obviously be a tendency for retention of their original relations on the part of the surrounding skeletal structures. The only place that the braincase could adapt itself to facial elongation would be in front of the brain, in the ethmoid region. In accordance with these obvious facts I have in fig. 15b dia- grammed the Megalichthys braincase further modified to show the pro- portions which might be expected in a long-jawed form of large size. With this change, not only do the various regions fall into line but also the various structural features are clearly comparable. The nasal capsule region is not preserved in known Embolomeri and was presumably present only in cartilage. The nostrils were widely separated in embolomeres and hence we may assume that the anterior end of the braincase was broad, and narrowed rapidly backward much as in Megalichthys. Presumably the anterior end of the palatoquadrate retained an ethmoidal connection here as in Megalichthys, antecedent 48 bulletin: museum of comparative zoology to the connection of the "pterygoid process" with this region seen in anurans, coecilians and such a urodele as Menopoma. The braincase in the ethmoidal region between, roughly, the anterior end of the parasphenoidal rostrum and the optic foramen, is short in the one case, extremely long in the other, but structurally almost identical. In either form there is a narrow ventral edge capped by the parasphenoid. Above the parasphenoid the braincase is for some distance upward essentially an interorbital septum in both forms, although this feature is more pronounced in the more "tropibasic" embolomere than in the crossopterygian. The dorsal part is more expanded in both forms, to contain the anterior end of the endocranial passages, consisting in both of paired tubes anteriorly, converging posteriorly to the anterior end of the brain cavity proper. In the case of the crossopterygian I have assumed that much of the length of these tubes was occupied by the olfactory lobes and even the anterior ends of the hemispheres. In the readjustment of parts correlated with elongation of the skull in amphibians this was probably not the case. Presumably the paired canals were occupied by the olfactory "nerves." This reduction in necessary bulk of the dorsal part of the region has allowed a slimmer development of the septal region. The primary upper jaws in both types were closely applied, but not fused, to the braincase in the shelf above the parasphenoid at the base of the interorbital septum, a condition which might easily give rise either to a separation and the development of the typical interptery- goid vacuities of amphibians or to the fusion of the adjacent elements characteristic of many reptiles. In the sphenoid region, the basiptery- goid process of the amphibian is quite similar to that of the crossop- terygian. Both include in their composition not only endochondral bone but also a dermal component from the parasphenoid. The seemingly lower position of the basipterygoid process in amphibians is to be in part correlated with the expanded notochord back of this region in crossopterygians and consequent differences in the level of the posterior part of the floor of the brain case. The large opening which Watson interprets as mainly for the optic nerve in Embolomeri lies, as in rhipidistians, ventral to the anterior prolongation of the endocranial cavity ; here, however, the reduction of this cavity anteriorly leads to this opening becoming a perforation through the posterior end of the high, thin septum. Conditions in later types (cf. Watson 1919, fig. 12, Capitosaurus) render it certain that the vena capitis lateralis passed backward over the basipterygoid process in embolomeres as in rhipidistians. Watson romer: braincase of megalichtiiys 49 notes grooves indicating that the internal carotid curved down and forward around the process to its foramen, as in crossopterygians. The course of the ophthahnic artery and pituitary vein is doubtful; 3ffllat pocp? fenov" vp+vmm +211 lat ppar? J3JJ pocp? vp+^aoph Fig. 15. A, Side view of the braincase of Megalichthys so modified, by the fusion of the two moieties and the extension of the parasphenoid backward over the exposed notochordal area, as to eliminate the kinetic speciahzations. B, The same, further modified in an amphibian direction, by a change of pro- portions, to introduce an elongated "sphenethmoid" region. C, Diagram of the side view of a primitive embolomerous amphibian, from Watson's figures and descriptions. analogy with other lower tetrapods suggests a position in front of the basipterygoid process in a position similar to the foramen which I assume to have carried them in Mcgalichihys. A bar passing backward 50 bulletin: museum of comparative zoology and downward between the assumed openings for the optic and trig- eminal nerves seems certainly to represent the pila antotica (prootica) which I have termed the laterosphenoid pillar. This marks the pos- terior termination of the crossopterygian ethmo-sphenoid. Above and below the ends of this bar there is a solid connection of the two halves of the braincase in the position of the upper and lower articulations in the crossopterygian. The ventral gap is covered by the parasphenoid in the embolomeres. It is of interest, however, that in many other early tetrapods, there is an unossified gap in the floor of the braincase in an area comparable to the fenestra ventrally separating the two halves of the crossopterygian structure. Even though the cross- opterygian subdivision be assumed to have been absent in tetrapod ancestry, early land vertebrates show a strong development of fea- tures which might have led to the origin of such a condition. In the prootic region a large anterior opening undoubtedly trans- mitted the entire trigeminal nerve and the superficial ophthalmic; the commisures separating the various components have disappeared, leading to the conditions seen here in most tetrapods. The position of the facialis is uncertain. From the situation in other paleozoic tetra- pods, however, I feel justified in assuming that the hyomandibular and palatine rami left the braincase farther posteriorly and gained the surface (as tentatively indicated) in a fashion comparable to that in crossopterygians. The posterior margin of the trigeminal opening is probably the prefacial commisure, and the foramen had not the in- clusiveness of the embryonic (and anuran) prootic foramen. A marked change in the lateral aspect of the otic region is due to the disappearance of the embryonic lateral commissure, the jugular canal and the trough anterior to it, freeing the vena capitis lateralis and the orbital artery (temporal, stapedial) to take an anterior-posterior course lateral to the braincase in the "cranio-quadrate passage." I have elsewhere called attention to the obvious development of the crossopterygian type of hyomandibular into the primitive tetrapod stapes. In this connection it may be recalled that Watson points out that the fenestra ovale had not yet developed in the Embolomeri studied, the point of ventral attachment of the "stapes" still being merely a depression in the outer wall of the braincase, as in cross- opterygians. Watson finds a foramen for the vagus in an appropriate position. He does not describe any openings such as those which I have tenta- tively assigned to a posterior lateralis nerve and the glossopharyngeal. Possibly, as in modern amphibians, these nerves have gathered into a romer: braincase of megalichthys 51 common foramen. I have elsewhere noted the possible implications of such a change on the development of the perilymphatic system. The dorsal part of the otic region is poorly preserved in known Embolo- meri, and needs careful restudy in other early tetrapods. In conse- quence I am uncertain as to the fate of the supra-otic fossa and ad- joining structures. A seemingly logical assumption is that the parotic process has become the "paroccipital" process found at the back of the labyrinthodont skull, and that this in turn has become the "paroc- cipital" process of many permo-carboniferous reptiles. I feel, however, none too sure of any of these homologies, and am inclined to believe that the reptilian paroccipital, at least, is a new development from the process to which I have tentatively given this name in Megalichthys. Further data is needed. The occipital regions seem fairly comparable in the two groups. The hypoglossal opening has not been identified in embolomeres as yet, but its presence in other early amphibians makes it certain that was present here, as in Megalichthys, but presumably with a single external opening. There is little development of a true condyle in Megalichthys, the major connection with the column being afforded by the notochord itself. It is easy, however, to see how the single large circular condyle of the primitive amphibian developed with the reduction of the notochord. The roof of the braincase is in both cases unbroken by foramina except that for the pineal. In the crossopterygian this opening is far forward; in the embolomere far toward the back. This seeming contrast, however, is correlated with the changes in proportions noted above; it is not so much a backward movement of the pineal as an elongation of the region anterior to it. In this connection brief comment may be made concerning the homologies of the roofing bones in the two cases. I have recently (1936) tended to take the despairing attitude that it is perhaps im- possible to make positive comparisons between the two groups ; but the obvious similarities here noted in the case of the braincase gives renewed hope. A stumbling block in the identification of the important median longitudinal elements lies in the seeming contrast in the position of the pineal opening in the two cases. In tetrapods it lies far back between the definitive parietals; in crossopterygians far forward between the bones generally identified as frontals. Morphologists have been in general loath to believe that the foramen would change its position relative to the dermal elements. Save-Soderberg has recently (1935) 52 bulletin: museum of comparative zoology attempted to account for the seeming shift by a complicated cutting apart and pasting together of fragments of the elements of this region, for which there is little evidence. The writer (1936) pointed out that the "extrascapulars" of crossopterygians are not homologous with the dermal supraoccipitals of amphibians, with which they have been gen- erally correlated; but this only seemed to make the problem more difficult of solution. A ray of light on this seemingly difficult situation is the recent sug- gestion of WestoU (1936), which he promises to elaborate in a future paper, that the elements concerned with the pineal in the two cases are actually the same; the "frontals" of the fish are equivalent, in part, at least, to the true parietals. At first sight this seems highly improb- able, for it leaves the large "parietals" of the fish as homologues of the comparatively small dermal-supraoccipitals of the tetrapod. But in the light of the history of the braincase it seems that WestoU 's solution is the correct one. The differences in seeming position of the pineal and true parietals are not absolute but in the main relative and due to differences in length of the facial region. Anterior elongation would reduce the supposed fish "parietals" to reasonable comparative size, and we may assume further that these true dermal-supraoccipitals have begun the process of reduction which eventually resulted in their disappearance from later tetrapod skulls. Comparatively little is known of the internal structure of the em- bolomere braincase. A few points concerning the matter have been mentioned earlier. The pituitary fossa, in the absence of the large crossopterygian notochord, is of "normal" construction, with an overhanging sella. The general agreement of the internal soft parts of crossopterygians with those of modern amphibians suggests that in general the cavities containing these structures in embolomeres may have been comparable to the crossopterygian structure. I have pre- viously commented on the seeming contrast in the mode of exit from the brain cavity of the posterior branch of the auditory nerve and of the glossopharyngeal. Apart from the kinetic features — intracranial joint and large noto- chord— the crossopterygian braincase is not only a possible morpho- logical ancestor of that of the embolomere, but approaches the latter closely in many features. To convert the one into another we need only assume: (1) elongation of the ethmoid region, (2) loss of the jugular canal, (3) fusion of nerve exits in a few cases. romer: braincasp: of megalichthys 53 Comparison ivlth Dipnoaus In the following comparisons with various fish groups, as was the case with tetrapods, it will be assumed that the ehondrocranial sub- division and large notochord of the known crossopterygians are secondary characters which will be disregarded for comparative purposes. It is unquestionable that crossopterygians and lungfish are allied stocks. But comparison of their endocranial structures is difficult. Part of this is due to the marked contrast in jaw structure and articu- lations. But further difficulties are due to the fact that living lungfish have surely departed widely from the ancestral types in braincase structure, as they are known to have done in the case of their dermal skeleton. Of older dipnoans we know as yet only an incomplete brain- case of Diptcrus described by Watson and Day (1915); a promised description of a second paleozoic braincase by Stensio will be awaited with interest. The posterior region of the modern lungfish skull, with its incor- poration of numerous vertebral elements and the enormous backward expansion of the parasphenoid, is particularly difficult to compare with that of a crossopterygian. Here, however, Watson and Day's work indicates that early conditions were similar. Their figure of the ventral surface of the occipital region of Diptcrus shows a condition in the aortic grooves, etc., highly comparable with a crossopterygian. Their description of the remainder of the post-otic region reads almost exactly like a description of the occipital region of Megalichthys. The condition in the orbitotemporal and otic regions, as interpreted on embryological grounds, is basically different in structure. There is no outer wall (lateral commissure) in the parotic region. Instead the region is enclosed laterally by an otic process of the palatoquadrate which, however, in the embryo serves the same functional purpose of enclosing a ''cranio-quadrate passsage" the contents of which (vena capitis lateralis, orbital arter\-, hyomandibular ^TI) are exactly those of the jugular canal of Megalichthys (or Acipenser). There is, of course, no floor to the passage; but here again there is analogy to the structure of the jugular canal in the embryonic Amia before the for- mation of the post-palatine commissure. In later embryonic and larval life there occurs a great further development of cartilage in the area from the otic ramus of the palatoquadrate back along the outer surface of the otic region. Not 54 bulletin: museum of comparative zoology only does the "cranio-quadrate passage" become floored, but its ele- ments are separated; hyomandibular VII is far removed at its outlet from the artery and vein; the orbital artery and internal carotid are enclosed by cartilage back beyond their point of division, along the dorsal aortic channel ; the vein not only has a separate channel through the region comparable in position to the jugular canal, but is almost completely enclosed by cartilage to a point behind the vagus ; and the vagus (except for lateralis components) is completely enclosed to a point below its crossing the head vein. In the orbital region there are further contrasts associated, again, with the fusion of palatoquadrate with braincase. A solid attachment is found between the two elements not only at the site of the basiptery- goid process but also by a bar running dorso-ventrally external to the profundus and head vein. Although jaw and braincase arise as a unit in Lepidosirien and Protopterus (Agar 1906) and fuse at a very early stage in Ceratodus, this bar is interpreted as pertaining to the palato- quadrate (as an ascending process) rather than to the chondrocranium (as a pila lateralis). Anterior to the basipterygoid region, palatoquadrate and trabeculae are indistinguishably fused. Internally the endocranial cavity, nerve exits and otic region are quite similar in the two groups ; the writer's restoration of the nervous system as inferred from its cavities, for example, are closely comparable with Pincus' description of Protopterus (1895). If we attempt to reconstruct the skull of a common ancestor of a dipnoan and a crossopterygian, a major concern is with the relations of palato-quadrate and braincase. First, is the "autostyly" of the dipnoan primary or secondary? To this question the orthodox answer is that it is secondary and we will, for the moment, assume that this answer is correct and that movable articulations were present in the common ancestor. A basal articula- tion would certainly have been present and the close apposition of jaw and braincase in the ethmoid region of the crossopterygians could readily change to the fusion seen here in the lungfish. The dipnoan palatoquadrate is further attached to the skull by otic and ascending processes. No such articulations are present in rhipidistians. They are present in modern amphibians. There, how- ever, these attachments are surely secondary, as is shown particularly by the work of Watson (1919) and Sushkin (1927); the palatoquadrate lies close to the braincase at these points in embolomeres and fusion follows. Since the crossopterygian palatoquadrate has a similar rela- romer: braincase of megalichthys 55 tion to the braincase, it is possible to imagine that the dipnoan connections are likewise secondary. But an "otic" connection of palatoquadrate with braincase is found in early shark types and the chimaeras. Hence this may well be a primitive vertebrate character, present in the common ancestors of lungfish and crossopterygians and lost in the latter. If so, it would be expected that in this common ancestor the parotic process would have had an expansion of the lateral commissure forward anterior to the spiracle for this attachment; essentially a postorbital process. This development would necessarily imply that the jugular canal reached farther forward than in known crossopterygians and that instead of an open "temporal" region there would have been a structural ar- rangement highly comparable to the jugular part of an actinopterygian trigeminofacialis chamber; here more properly a trigeminal chamber, since the facial nerve would probably have been embedded in the posterior wall. The articulation of the ascending ramus, however, may be reason- ably interpreted as secondary. It is unknown in fishes apart from the forms here discussed and is surely secondary in tetrapods. In cros- sopterygians it is present in the coelacanths. But since it appears to be a progressive development, and since the basipterygoid attachment disappears in the course of this development, it is reasonable to assume that we have the replacement of an old ventral attachment by a new dorsal one. I have noted that the Ceratodus braincase is more expanded laterally in the otic region than is the case in crossopterygians, so that it more fully encloses the various nerves and vessels. This may be secondary; but (as noted below) the great degree of development seen here in a number of other fish types suggests that the condition in Ceratodus is essentially primitive. From the above discussion of dipnoans we may postulate that the chondrocranium in a form antecedent to known crossopterygians would, if found, differ from them in the absence of the kinetic mechan- isms, the presence of an "otic" articulation of the palato-quadrate, the presence of a trigeminal chamber, and probably a greater development of the region lateral and ventral to the otic capsule. It has become increasingly apparent during the past decade that a great phylogenetic cleft divides the "higher" fishes into two groups, the actinopterygians on the one hand, lungfishes and crossopterygians on the other; with the latter would of course be included such hypo- thetical and hoped for creatures as that just described and the actual 56 bulletin: museum of comparative zoology ancestor of the tetrapods, structurally approached but not attained by the rhipidistians. For this group I know of no appropriate term. Tate Regan (1928) has used Crossopterygii for these forms; but this is too wide an extension of a familiar term. Save-Soderbergh (1934, 1935) has proposed the term Choanata as that of a super-class (?) to include these forms and their tetrapod descendents as part of a phylogenetic scheme of rather unusual character. The term is hardly necessary in any phylogenetic and taxonomic scheme for which there is adequate support, although it is highly useful in a more "popular" sense, com- parable to "tetrapod," "amniote" etc. To cover the piscine members of this category, I propose the related term Choanichthyes. The classification of the gnathostome fishes would thus be as follows : Class Placodermi Class Chondrichthyes (Elasmobranchii s. 1.) Class Actinopterygii Class Choanichthyes The relations of the first two groups may be a matter of debate for many years to come. Comparison ivith Actinopterygians Since the actinopterygians are a group widely divergent from the choanate fishes, it would be expected that they would possess many features unconnected with the earlier history of the choanate stock. Under this category would come the myodome and the associated structural modifications. Apart from these, however, they exliibit a number of characters similar to those seen in the crossopterygians and give many suggestions as to the earlier nature of the ancestors of the Choanichthyes. Far more primitive than any of the existing higher actinopterygians are the carboniferous palaeoniscids in which the braincase has been described by Watson (1925). The myodome was already developed, and the characteristic high thin interorbital septum of the actinoptery- gians present; as Watson points out these features are obviously divergencies from primitive conditions related to the sensory dom- inance of the eyes. The pila prootica, lost in more advanced members of the class, was still present. A basal articulation with the palato- quadrate is found in Watson's material ; this is lost in almost all later types, except Lepidosteus. Neither in these nor in any other known actinopterygians is there any "otic" articulation; the loss of this con- nection, surely a primitive one, is to be associated with the develop- romer: braincase of megalichthys 57 ment of hyostyly. There is no ventral articulation of the hyomandihii- lar; presumably a reduction associated with function. The original area of "otic" palatocjuadrate attachment seems clearly marked out by the large postorbital process, so greatly expanded that it surrounds the spiracular area to form a canal. There is a highly developed trig- eminofacialis chamber with an associated jugular canal, forming an arrangement such as I have pointed out would be expected in an ancestor of the choanate fishes. The posterior part of the braincase is highly developed. The head vein is not enclosed to the extent seen in Ceratodns, although it is over- himg by a ridge rather as in McgaUchthys. On the other hand, the ventral surface is more developed, enclosing within its substance much of the dorsal aorta. It is of interest that in Watson's carboniferous material there is no subdivision of the completely ossified braincase; presumably a primitive condition. Despite the undoubtedly wide separation of the two higher fish stocks, it will be seen that thev exhibit fundamental similarities. Indeed, the structure deduced earlier for the braincase of a choanate ancestor would seem essentially to have been that from which the actinopterygian braincase might have been derived as well. If we assume the development of large eyes and jaw suspension by the hyomandibulars the remaining actinopterygian specializations follow as a logical conseciuence. Comparison with shark-like fishes If we attempt to trace the history of the choanate braincase down- ward into the lower gnathostomes, we gain little satisfaction from mod- ern sharks and rays. The braincases in these forms is markedly different from that expected in an ancestor of the Choanichthyes, or of an actinopterygian for that matter. The differences lie not merely in the widely different proportions but in many structural features, such as the absence of a functional basal articulation of the palato- quadrate and the almost universal absence of an otic articulation, the absence of a trigeminofacialis chamber and of a jugular canal (except for a slight development of this sort in Squalus) and, in general, the comparatively "undeveloped" state of the braincase so that many of the blood vessels and nerves which in bony fishes are frequently en- closed in bony channels are here free from the braincase or enclosed only to a slight degree. Indeed, the chimaeras, despite their speciali- 58 bulletin: museum of comparative zoology zations, afford a more satisfactory basis for comparison, since both basal and otic palatoquadrate connections are present. Among paleozoic sharks, however, we meet with better success. I have been engaged for a number of years in the study of the braincase of permo-carboniferous pleuracanths. Unfortunately the work is as yet unfinished and I have published but a few incomplete and dia- grammatic figures (1933). In these sharks there is a well developed "postorbital" process, enclosing an area which is essentially a trigem- ino-facialis chamber, and pierced by the structures which con- stitute the contents of the jugular canal or "cranioquadrate passage;" the palato-quadrate (as has long been known) articulates with this process ; the ventral surface of the braincase encloses a long reach of the dorsal aorta, etc. So different were these forms from modern sharks that I have been forced to compare them with bony fishes instead for an interpretation of their structure, although it was impossible that there should be any close connection between pleuracanths and "higher" fish types. It was therefore pleasing to me to see the recent preliminary account by Stensio (Holmgren and Stensio 1936) of his findings in a Cladodus specimen from Wildungen. The material is fragmentary (for example, the skull was probably more elongate than his restoration would indicate) but shows a condition almost exactly like that in pleura- canths. It is evident that we are dealing here with a braincase form which was widespread among early sharks. We have here, then, a type of shark skull which comes fairly close to the type inferred as ancestral to later bony fishes. But we are not yet at the meeting point of these varied lines. There is in these forms no basipterygoid articulation, for example, although the chimaeras show that such an articulation can exist in the chondrichthyes. Further, all trends in modern work on fishes, particularly that of Sten- sio, indicates that the common ancestors would have had an ossified braincase and a roof of dermal bones, whereas these types have cartilaginous (although well calcified) braincases and there is no positive evidence of any roofing bones in cladoselachians or pleura- canths. These forms are primitive sharks, but they are definitely on the shark side of the bifurcation; degeneration has been initiated. A further step downward is seen in Macropetalichthys, in which the braincase has been described by Stensio (1925); as shown by the post- cranial skeleton this form is a shark. Its skull structure, although at first glance seemingly grotesque, can readily be interpreted in terms of pleuracanths and cladodonts on the one hand and the hypothetical romer: braincase of megalichthys 59 ancestor of the higher fish groups on the other. The orbit is confluent with a trigeminal chamber. A huge lateral process may well have served as the point of attachment for an otic ramus of the palato- quadrate as well as the hyomandibular. The elements of the jugular canal group traverse this long process as in pleuracanths and dado- donts and the rest of the head vein is partially concealed. The aortic region is buried beneath the bone or, posteriorly, lodged in a deep groove. There is no definite evidence of a basal articulation; but it will be noted that there is a reasonable point of articulation at a point beneath a "preorbital process" which is at one and the same time ethmoidal and basal in position. The basal articulation may have evolved as an "emphasized" posterior termination of a once single articular area between future ethmoid and basipterygoid regions of attachment, the separation taking place in relation to elongation of the "sphenethmoid" region. As indicating the probability of this, it may be noted that the palatoquadrate is continuously in close contact with or fused to the braincase between these two regions of attachment in forms as far apart as chimaeras, lungfish, crossopterygians, embolo- meres and even Scymouria. Finally and most important is the fact that in Macropetalichthys the braincase is ossified and is roofed with dermal bones. We have thus travelled back to the stage in the development of skeletal materials to be expected in the common ancestor of all living jawed vertebrates. I do not claim that Macropetalichthys and his relatives are the actual ancestors of later vertebrates; the acanthodians, for example, when better known, may prove to be even more generalized and primitive. But it is, I think, clear that in braincase structure Macropetalichthys is very close to conditions expected in such a common ancestor. In this connection I wish to protest against the frequent assumption (as in Holmgren and Stensio 1936, pp. 324-335, DeBeer and Moy- Thomas 1936, p. 309, etc.) that the presence of certain structural features in Macropetalichthys argues for their being structures charac- teristic of arthrodires, and particularly the assumption that similarities between Macropetalichthys and sharks indicate a close relationship between arthrodires and sharks. These assumptions rest upon the premise that Macropetalichthys is an arthrodire; this premise is con- trary to the known facts (Heintz 1932). Until recent years this devonian fossil was regarded as an arthrodire for the reason that it possessed a bony skull roof. The presence of such a structure in a shark was contrary to theories of vertebrate evolution then prevalent and Macropetalichthys was disposed of by including in 60 bulletin: museum of comparative zoology the arthrodires, although there was no positive ground for such in- clusion. This belief was still prevalent when, in 1925, Stensio wrote his excellent paper on the braincase of this fish and he was hence justified at the time in using Macropetalichthys as a basis for arguing that arthrodires were closely related to sharks. In 1933 the writer in his "Vertebrate Paleontology" took the (then) bold step of removing this form from the arthrodires and placing it among the sharks (in a broad sense), using Smith Woodward's term Stegoselachii to cover provisionally this and other primitive sharks in which dermal elements were still present. This step was almost imme- diately justified by Broili's discovery (1933) of Macropetalychthyids in which the body was preserved; these showed that the entire skeleton was almost typically shark-like, and not arthrodian in nature. Macro- petalichthys cannot be included in the Arthrodira or the broader group, Placodermi, without stretching these terms to an unwarranted degree. Unquestionably placoderms and sharks (Elasmobranchii, s. 1.= Chondrichthyes) have a common ancestor, and presumably the arthrodires and other placoderms, although aberrant, have retained many primitive features to be expected in a shark ancestor. But Macropetalichthys cannot be used as an argument on the placoderm side of the question; it is definitely a shark, while the typical arthro- dires go back to a primitive stock, described especially by Heintz (1929), much closer to the antiarchs and indicating a pedigree widely divergent from the Macropetalichthys shark group. Notes on Early Gnathostome Cranial History As noted earlier, consideration of the phylogenetic history of the fish cranial region on embryological grounds leads to the assumption that a cartilagenous condition of the skeleton was a primitive one; further that the original chondrocranium was a simple structure and separate from the jointed jaws and other visceral arches; complexity of structure and fusion of parts are assumed to be secondary. Any inquiry, such as the present one, into the pedigree of a fish skull type, leads to almost diametrically opposite conclusions if modern paleontological research be kept in mind. In the first place, the work of both Watson and Stensio has made it clear that in most instances a cartilagenous condition of skeletal parts is secondary. The latter, particularly, has shown the widespread occur- rence of endochondral as well as dermal bone among the oldest verte- romer: braincase of megalichthys ()1 brates. That a bony structure is to be expected in an ancestral gnathostome is a thesis which I feel to be so firmly established that I have accepted it in the preceding discussion without question. Secondly, primitive vertebrates were not simple in cranial structure, but as complex as, or more complex than many of their descendants. In the days when paleontology was in its infancy, morphologists fondly evolved hypothetical ancestral forms to fill the gaps between living groups, "archetypes" of simple and diagrammatic structure. Today many of these gaps are filled by actual fossils. But these are never the simple types imagined; often they are quite the reverse, showing many unexpected quirks in structural evolution. So in the present case. In the search for a cranial type ancestral to that of crossopterygians, we have found no simplicity. The compara- tively simple and "undeveloped" modern shark braincase shows little resemblance to that of bony fishes; the more complex, more "ex- panded" type of older sharks is much more similar. It is obvious, I think, that the primitive gnathostome braincase was developed to a degree almost unknown in modern forms, to include within its substance many structures usually external to it. Neglecting many details, these inclusions were of two sorts. 1. The dorsal aorta and related structures were probably enclosed in the base of the braincase from the internal carotoid region back into or beyond the aortic arch region. 2. The jugular vein was probably enclosed in a canal all along the side of the skull from the orbit to a point opposite the occiput. The anterior opening of this canal would be the primitive trigeminus cham- ber; back of this the jugular canal is the remnant of an originally longer structure. The orbital artery would have traversed this canal in its anterior part, and not only V and VII, but nerves IX and X may have opened into this cavity; it would have been essentially an elon- gated "cranio-quadrate passage." While for purposes of simplification I have spoken of a common canal for these structures, it is more probable, from the "shark" evidence, that vein, nerve and artery were actually separately en- closed, although adjacent; the nerves may have emerged close to but not in continuity with the vein canal, and the orbital artery (as is often the case) likewise may have occupied a separate but adjacent canal. With regard to visceral arch articulations, the Megalichthys double dorsal and ventral attachment of the hyomandibular leads to interest- ing possibilities. I assume that this is primitive. The argument of the 62 bulletin: museum of comparative zoology Fig. 16. A series of possible stages in the early history of the vertebrate skull, in descending order. A, diagram of hypothetical primitive shark in which branchial arches and jaws articulate with the braincase dorsally and ventrally, an expanded "jugular canal" region extended backward along either margin of the braincase from a primitive "trigemino-facialis chamber", and the anterior end of the aorta and related vessels were enclosed on the under side of the braincase. A\ cross section of the same, through the otic region. B, the same figure so modified as to suggest an earlier "autostylic" condition with all visceral arches fused with skull. The broken line indicates the division between skeletal materials of "visceral" (neural crest) origin and normal mesoderm. B', cross section. Axial mesodermal area shaded. C, still further modification of the same with shortening of the trabecular region and further fusion of the branchial arches, in a condition comparable to that of ostra- coderms. romer: braincase of megalichthys 63 previous section suggests that in a primitive gnathostome there were two palatoquadrate articulations; an anterior ventral one with the trabecular region, from which later ethmoid and basipterygoid articu- lations arose, and one postero-dorsally with the combined postorbital and otic process. These two attachments are comparable to those of the hyomandibular. Can a similar situation have held farther back? There is evidence suggesting, although not at all proving, that primi- tively the branchial arches were attached to the side of the braincase by double dorsal and ventral articulations (cf. Schmalhausen 1923 and Sushkin 1927, the latter citing Woskoboinkoff). I have, in fig. 16a, shown in diagrammatic and vastly over-simplified fashion, the conditions which may have prevailed in an early gnatho- stome. As noted previously, Macropetalichthys shows essentially the braincase characters suggested; there is little paleontological evidence regarding the visceral arches. A further theoretical step downward in skull history concerns the attachment of the visceral arches to the skull. It is generally assumed, particularly by those approaching the subject from the embryological side, that the jaws, and presumably the other arches, were without question separate elements in primitive vertebrates (cf. for example, de Beer and Moy-Thomas 1935, p. 307, footnote), and that a fused autostylic condition of the jaws is secondary. There is however a great amount of evidence which points in the other direction. The fusion of jaws and braincase in amphibians is certainly secondary. On the other hand, as we have noted, the dipnoan situation is less clear cut, and in the chimaeras jaws and brain- case are a unit at this first appearance; there is no evidence of a secondary condition. Further Moy-Thomas (1936, etc.) notes that sharklike forms with fused jaws were common in the Paleozoic, in contrast with their subsequent rarity. There are certainly strong sug- gestions that in primitive gnathostomes a fused condition of the upper jaws was a general, and not improbably a really primitive condition. Some, if not all, of the modern forms with this type of jaw support may be secondary; but this "reversion" may be readily explained by a return to paths of embryonic development not long abandoned (even the amphibians, it will be noted, were "reverting" by the beginning of the Permian). If Agnathous forms be considered the evidence for a primitively fused cranial structure is overwhelming. In living cyclostomes the visceral apparatus, except for the specialized "tongue" is solidly united. And, finally and most conclusively, Stensio's splendid work of 1926 on 64 bulletin: museum of comparative zoology the cephalaspids has logically forced him to adopt without question the thesis that the primitive condition in vertebrates was one in which visceral arches, braincase and dermal covering formed a single solid structure. One may be justified, therefore, in assuming that in a primitive gnathostome the upper jaws and upper ends of the other arches were fused to the skull, as shown in diagram in fig. 16b. In cross section (fig. 16b') I have shaded the braincase component of the fused mass, in contrast to the visceral arch portion. In such a solid structure the line of demarcation, being of no functional im- portance, might vary somewhat. This leads to interesting theoretical possibilities. For example, if the separation in the region of the otic process of the palato-quadrate took place at the point indicated in the diagram, there would remain on the braincase a lateral commissure enclosing a jugular canal and forming the lateral wall of a trigeminal chamber. If the line of division between embryonic visceral and axial components were slightly more medial, these structures would be absent from the braincase and the outer wall incorporated in the palatoquadrate; the enclosure would be an antrum petrosum lateralis, rather than trigeminal chamber and jugular canal. DeBeer (1926, p. 359 f. f.) is correct, on embryological grounds, in criticising AUis' com- parison (1914) of the "antrum" of Ceratodus with the "chamber" of Actinopterygians. But under the hypothesis here entertained the two cavities may be phylogenetically identical, and Allis' comparison essentially sound. In the dorsal view of the hypothetical fused stage discussed, the broken line indicates, laterally, the division between arch and braincase components; anteriorly this line crosses the braincase at the level of the division between the trabecular and parachordal regions of the skull. This is in accord with the findings of Piatt and Stone, noted earlier, dividing the cranial mass into its two embryological compo- nents. The trabecular region arises like the visceral arches from neural crest material, while the posterior part of the braincase arises in "normal" fashion. This embryological division of the early braincase leads to a final hypothetical suggestion with regard to early vertebrate cranial history. All lines of evidence suggest that the anterior region of the head — this visceral, trabecular region — has been elongated during the phylogenetic history of vertebrates. Much of the evidence has to do with skeletal structures; but there is, as is well known, a great array of material concerning the pituitary complex which can only be explained by such romer: braincase of megalichthys 65 a process; while the history of the brain is best explained on the assumption of a gradual forward expansion of the forebrain. With these facts in mind, I have ventured, in fig. 16c to further modify my diagrammatic skull by shortening the anterior part of the \nsceral region. With this shortened state might reasonably be asso- ciated a coalesced condition of the nasal sacs and various modifications of associated internal structures. In such an assumed stage we see a cranial pattern which in its pro- portions might well have been that of the less specialized lower Devonian arthrodires (cf. Heintz 1929, etc.), whose internal structure is as yet unknown. Finally, we have here a theoretically basic gnathostome stage which is not separated by any great or unbridgeable structural gap from the known ostracoderms. Even such a seemingly specialized type as Cephalaspis shows many resemblances to my hypothetical prognathostome. The anterior visceral region — the "snout" — is differently developed in the two cases, nostrils and hypophysis being situated dorsally in Cephalaspids; at present the primitive conditions here cannot be deduced with certainty. I have not attempted to interpret the ventral aspect of the visceral region. But in general contours there is a marked similarity, and reference to Stensio's figures will show that although the internal structure of cephalaspids is, of course, highly complex, the situation of the main arteries, the main head veins and the proximal portions of the nerves is fundamentally similar to that which I have deduced for a gnathostome ancestor. Possibly the anaspids, if their internal structure were known, would be even closer, for they obviously lacked such specializations as the presumed electric fields of the Osteostraci. To sum up in ascending rather than descending order, I assume the major stages in cranial evolution to have been as follows : 1. An agnathous stage with arch and braincase components fused to one another and to the overlying dermal roof, as postulated by Stensio and as represented in great measure by anaspids and cephalaspids. 2. A primitive gnathostome stage, in which the lower halves of the jaws and other arches were movable, the upper halves still fused, the anterior visceral region of the skull and jaws still short. This stage presumably represented in a general way by the lower Devonian arthrodires. 3. A primitive "shark" in which the anterior portion of the skull (and jaws) have elongated; the visceral arches perhaps freed from they braincase but the upper jaws still fused and a dermal skull roof rev 66 bulletin: museum of comparative zoology tained. Represented, except that the palatoquadrate has been freed, by such an shark as Macropetalichthys and possibly the acanthodians, were these forms better known. 5. Subsequent to this stage began the differentiation of the later fish groups. Almost universally the palatoquadrate becomes freed from the braincase; the originally broad and deep braincase tends to con- tract, releasing to the surface many of the structures originally embedded in it; the originally solidly ossified braincase tends to break up into separate elements or, as in many "higher" types as well as sharks, to degenerate to a cartilagenous structure; in sharks the dermal roof is lost. In the theory of vertebrate skull evolution advanced above there is little actually new; many of the suggestions made here have already been advanced by other workers and all the elements of this assumed sequence are indicated by modern work on paleozoic fishes. I have not attempted to work out the full story of possible structural develop- ments, and have confined myself to a few major features. In attempt- ing to illustrate the evolutionary story by simple diagrams, I have laid myself open to the criticism which maybe made of all "archetypes," for surely even the most primitive vertebrates (as illustrated in cephalas- pids) were highly complex. The theory advanced will undoubtedly prove to be incorrect in many details, not improbably in many major features. But I have, at least, attempted to erect a working hypothesis which is far more consistent with the known paleontological facts than those based primarily on embryology which are now current. SUMMARY The braincase of Megalichthys is described in some detail, and a restoration of the cranial nervous system, internal ear and certain parts of the vascular system attempted. These soft parts appear to be com- parable to those of dipnoans on the one hand and amphibians on the other. Megalichthys appears to be representative of the rhipidistian crossopterygians in braincase structure. Except for "kinetic" features the crossopterygian braincase is a reasonable morphological antecedent of that of primitive tetrapods and shows a close approach to tetrapod conditions in such features as (for example) the hyomandibular ( = stapes). In Dipnoi and Crossopterygii the braincase is reducible to a common pattern. The class term Choanichthyes is proposed to cover these two romer: braincase of megalichthys 67 groups and hypothetical rohitcd types. A search for the antecedents of the Crossopterygian braincase leads to the conclusion that in the primitive devonian shark Macropctalichthys we have a form morpholog- ically ancestral in braincase structure to both sharks and all "higher fishes." Consideration of fish history suggests that in the primitive gnathostome the braincase was a highly expanded structure to which the upper ends of visceral arches were attached. Such a condition in great measure bridges the structural gap separating gnathostomes from the older jawless ostracoderms. 68 bulletin: museum of comparative zoology BIBLIOGRAPHY Agar, W. E. 1906. The Development of the Skull and Visceral Arches in Lepidosiren and Protopterus. Trans. Roy. Soc. Edinburgh 45, pp. 49-64. Allis, E. p. 1914. The pituitary fossa and trigemino-facialis chamber in Ceratodus fosteri. Anat. Anz. 46, pp. 625-637. 1919. The myodome and trigemino-facialis chamber of fishes, and the corresponding cavities in higher vertebrates. Jour. Morph. 32, pp. 207-326. 1928. Concerning the pituitary fossa, the myodome and the trigemino- facialis chamber in recent gnathostome fishes. Jour. Anat. 63, pp. 95-141. Broili, F. 1933. Ein Macropetalichthyide aus den Hunsriickschiefern. Sitz. Bayer. Akad. Wiss. Miinchen, 1933, pp. 417-437. Bryant, W. C. 1919. On the Structure of Eusthenopteron. Bull. Buffalo Soc. Nat. Sci. 13, pp. 1-23. DeBeer, G. R. 1926. Studies on the Vertebrate Head. II. The orbito-temporal region of the skull. Quart. Jour. Micr. Sci. 70, pp. 263-360. DeBeer, G. R., and J. A. Moy-Thomas. 1935. On the Skull of Holocephali. Phil. Trans. Roy. Soc. London 224B, pp. 287-312. Goodrich, E. S. 1930. Studies on the Structure and Development of Vertebrates. London, XXX +837 pp. Greil, a. 1913. Entwicklungsgeschichte des Kopfes und des Blutgefassystems von Ceratodus forsteri. Semon's Zoologische Forschungreisen in Australien I, pt. 2, pp. 935-1492. Heintz, a. 1929. Acanthaspida. Skrift. Svalbard Ishavet, Oslo, Nos. 23 and 23a, 81 and 20 pp. 1932. The Structure of Dinichthys. In: Bashford Dean Memorial Volume, Archaic Fishes, ed. by E. W. Gudger, Amer. Mus. Nat. Hist., New York, pp. 115-224. romer: braincase of megalichthys 69 Herrick, C. J. 1921. A Sketch of the Origin of the Cerebral Hemispheres. Jour. Comp. Neurol. 32, pp. 429-454. Holmgren, N. and E. A. Stensio 1936. Kranium und Visceralskelett der Acranier, Cyclostomen und Fische. In: E. Bolk and others, Handbuch der Vergleichenden Anatomic der Wirbeltiere, vol. 4, pp. 345-353. Holmgren, N. and C. J. van der Horst. 1925. Contribution of the Morphology of the Brain of Ceratodus. Acta Zool., 6, pp. 59-166. Moy-Thomas, J. A. 1936. The Structure and Affinities of the Fossil Elasmobranch Fishes from the Lower Carboniferous Rocks of Glencartholm, Eskdale. Proc. Zool. Soc, London 1930, pp. 761-788. Nick, L. 1912. Das Kopfskelet von Dermochelys coriacea L. Zool. Jahrb. Abt. Anat., 33, pp. 1-238, pis. 1-12. O'Donoghue, C. H. 1920. The blood vascular system of the Tuatara, Sphenodon punctatus. Philos. Trans. Roy. Soc. London 210B, pp. 175-252. Price, L. I. 1935. Notes on the Brain Case of Captorhinus. Proc. Boston Soc. Nat. Hist., 40, pp. 377-386. Regan, C. T. 1929. Article: Fishes, in Encyclopedia Britannica, 14th ed., vol. 9, pp. 305-328. Romer, A. S. 1933. Vertebrate Palenotology. Chicago, 491 pp. 1936. Early History of Texas Redbeds Vertebrates. Bull. Geol. Soc. Amer., 46, pp. 1597-1658. 1936a. Studies on American Permo-Carboniferous Tetrapods. Problems of Paleontology, 1, pp. 85-93. 1936b. The Dipnoan Cranial Roof., Amer. Jour. Sci., 32, pp. 241-256. Save-Soderbergh, G. 1934. Some points of view concerning the evolution of the vertebrates and the classification of this group. Arkiv for Zoologi, 26A. No. 17, 20 pp. 1935. On the dermal bones of the head in labyrinthodont Stegocephalia and primitive Reptilia. Medd. om. Greenland 98, no. 3, 195 pp. 70 bulletin: museum of comparative zoology SCHMALHAUSEN, J. J. 1923. Der suspensorial apparat der Fische und das Problem der Ge- horknochelchen. Anat. Anz. 56, pp. 534-543. Stensio, E. a. 1922. Uber zwei Coelacanthiden aus dem Oberdevon von Wildungen. Pal. Zeitschr. 4, pp. 167-210. 1922a. Notes on Certain Crossopterygians. Proc. Zool. Soc, London 1922, pp. 1241-1271. 1925. On the Head of the Macropetalichthyids. Publ. Field Mus. Nat. Hist., Chicago no. 232, Geol. Ser. 4 no, 4, pp. 89-197. 1926. The Downtonian and Devonian Vertebrates of Spitsbergen. Part I Family Cephalaspidae. Skrifter om Svalbard og Nordishavet 12, 391 pp. 1932. Triassic Fishes from East Greenland. Medd. om Greenland 83, no. 3, 305 pp. Stjshkin, p. p. 1927. On the modifications of the mandibular and hyoid arches and their relations to the brain case in the early tetrapoda. Palaeont. Zeitschr. 8, pp. 263-321. Watson, D. M. S. 1919. The Structure, Evolution and Origin of the Amphibia. — The "Orders" Rachitomi and Stereospondyli. Philos. Trans. Roy. Soc. London. 209 B, pp. 1-73. 1925. The structure of certain palaeoniscids and the relationships of that group with other bony fish. Proc. Zool. Soc. London 1925, pp. 815-868. 1925a. The internal ear of Osteolepis. Jour. Anat. 59, pp. 385-386. 1926. Croonian Lecture — The Evolution and Origin of the Amphibia. Philos. Trans. Roy. Soc. London, 204 B, pp. 189-257. 1930. Adaptation. Report 97th meeting British Assoc. Adv. Sci., pp. 89-99. Watson, D. M. S. and Henry Day. 1916. Notes on Some Palaeozoic Fishes. Manchester Memoirs 60, no. 2, 48 pp. Westoll, T. S. 1936. On the Structures of the Dermal Ethmoid Shield of Osteolepis. Geol. Mag. 73, pp. 157-171. romer: braincase of megalichthys 71 EXPLANATION OF ABBREVIATIONS USED IN FIGURES For many portions of the nervous and circulatory system the same abbre- viation may represent both the structure and foramina or canals containing it. aa Dorsal end of aortic arches. aca Ampulla of anterior vertical semicircular canal. ace Ampulla of horizontal semicircular canal. acp Ampulla of posterior vertical semicircular canal. aop Optic artery. aoph Ophthalmic artery. aor Orbital artery. ap Palatine artery. bal? Probable point of attachment of first branchial arch. bart Basal attachment of palatoquadrate to skull. bptp Basipterygoid process. bral First branchial arch. ca Anterior vertical semicircular canal. ccom Crus commune of utriculus. ce Horizontal semicircular canal. cer Cerebellum. cfa Canal reprsenting foramen apicale. ch Cerebral hemispheres. ci Internal carotid artery. clat Lateral commissure. cot VII Canal carrying the hypotic ramus of facial nerve. cp Posterior vertical semicircular canal. cpref Prefacial commissure. da Anterior end of dorsal aorta. daa Dorsal process articulating with otico-occipital. dap Cup to receive articular surface of ethmo-sphenoid. dper Perilymphatic duct. ds "dorsum sellae" between pituitary and anterior notochord attachment. en Position of external naris. endc Endocranial cavity. etha Pocket forming ethmoidal articulation with palatoquadrate. fa Foramen apicale. fbc Basicranial fenestra. "fenov" Position of future fenestra ovale, fm Foramen magnum, fr? Possible homologue of fenestra rotunda, ga Groove for lateral aorta, gh Habenular ganglion, gj Groove marking course of "jugular" vein. 72 bulletin: museum of comparative zoology gu Groove underlying lateral line. hy Hyoid arch. typ Hypophysis. hmd Dorsal articulation with hyomandibular. hmv Ventral articulation with hyomandibulae. inf Infundibulum. jc Jugular canal. "jc" Hypothetically expanded jugular canal area. jca Anterior opening of jugular canal. jcp Posterior opening of jugular canal. jt Trough in temporal region occupied by head vein. 1 Lagena. lo Olfactory lobe. It Position of lamina terminalis. mb Macula of pars basilaris. mn Macula neglecta. ms Ridges indicating speta between muscle segments on occiput, msc Mesencephalon. msl Maculae of sacculus and lagena. my Utricular macula. na Nutrient artery. nc Position of notochord. neap Nasal capsule. ncm Medial "pocket" in nasal capsule. nlatp Posterior lateralis nerve. nve Small foramina furnishing blood supply to ethmoid shield. oa Occipital artery. oart Otic (dorsal) attachment of palatoquadrate to braincase. oca Occipital arches. ocap Otic capsule. ol Optic lobes. one Orbito-nasal canal. orbc Orbital (sphenolateral) cartilage. pant Pila antotica (prootica). parch Parachordal. pas Parasphenoid. peart Polar cartilage. pep Posterior chorioid plexus. pin Pineal region. pit Pituitary fossa. pl Laterosphenoid pillar. peep? Possible homologue of paraccipital process. ppar Parotic process. pq Palatoquadrate. rcom Ramus communicans n. VII-X. romer: braincase of megalichthys 73 s Sacculus. send Endolymphatic sac. sof Supraotic fossa. sofen Fenestra between temporal and supraotic regions. sot Saccular otolith. spso Location of spiracular sense organ. svom Venous sinus beneath brain draining into middle cerebral. t Tubercles presumable for branchial arch muscles. tfc Primitive trigemino-facial chamber. tmarg Tanenia marginalis. trab Trabecula. u Utriculus. uot Utricular otolith. vaa Ventral process articulating with otico-occipital. vap Groove for articulation with ethmo-sphenoid. vcm Middle cerebral vein. vdac Dorsal tube presumable carrying vein or lymphatic from ear to cavity of posterior lateralis nerve, vj Anterior end of trough carrying vena capitis lateralis, vmd Small dorsal vessels in cerebellar region, vp Pituitary vein. vpr Small veins penetrating braincase from temporal region. I Olfactory nerve. II Optic Nerve. III Oculomotor nerve. IV Abducens nerve. Vmm Maxillary and mandibular rami of trigeminus. Vp Profundus nerve. Vp2 Canal in nasal region for profundus nerve. Vllhm Hyomandibular ramus of facial nerve. Vlllat Anterior lateralis nerve. VIIos Superficial opthalmic nerve. VIIos2 Canal in lateral ethmoid region for superficial ophthalmic nerve. Vllpal Palatine ramus of facial nerve. Vllot Hypotic ramus of facial nerve. VIII Auditory nerve. Vllira Anterior ramus of auditory nerve. Vllirp Posterior ramus of auditory nerve. Vlllrsl Ramus of auditory nerve to sacculus and lagena. IX Glossopharyngeal nerve. X Vagus nerve (less lateralis component). XII Hypoglossal nerve. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. LXXXII, No. 2 THIRD LIST OF ANTILLEAN REPTILES AND AMPHIBIANS By Thomas Barbour CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM November, 1937 No. 2. — Third List of Antillean Reptiles and Amphibians By Thomas Barbour INTRODUCTION I published a Second List of Antillean Reptiles and Amphibians in 1935 (Zoologica, 19, no. 3). Since that time much new information has accumulated. I have, therefore, prepared a third list, departing from the general custom not always consistently followed, of designating all or most island forms binominally. The practice of using trinomials for races that are obviously closely related has become so general, that I present herewith some attempts to show relationship in this way. There have been many groups in which I have not yet felt that our knowledge is complete enough to do this, and these I have allowed to stand as'in the previous lists, simply bringing the information concern- ing distribution up to date. Doctor Charles Schuchert in his noteworthy Historical Geography of the Antillean-Caribbean Region (New York, John Wiley & Sons, 1935, pp.I-XXVI and 1-811) writes on page 107 as follows: The writer agrees with Barbour and others that the Antillean faunas are too homogeneous throughout, and have too many phyla with deKcate organisms, to have reached the islands by flotsam and jetsam dispersal. On the other hand, very few species are now common to Antillia and Central America, and this means long isolation. "The time of the older migration is best seen in the snails, which have had a long and prolific evolutionary history, with a vast specific and generic differen- tiation. They can not be drowned out during submergences so long as there are islands left, and probably more islands existed than the paleogeographic maps show. Pilsbry thinks that the snail faunas of the Greater Antilles are cer- tainly as old as the early Eocene and probably go back as far as the Cretaceous, and Simpson holds that the migration was not only toward the east but back again as well to the land of origin. The older migration is also indicated by the few primitive mammals (insectivores only), but mammals do not proliferate into species as do the snails. Thirty of the 50 genera, according to Anthony, and 83 of the 97 species of Antillean mammals are restricted to the islands; the outstanding feature of the fauna is its endemic nature, with relationships to South America via Central America. "Most zoogeographers see continual land contacts in two places: an older, greater, and longer-enduring bridge from Honduras-Nicaragua across to Jamaica and Hispaniola, and a much younger, evanescent one from Yucatan to Cuba and Hispaniola. The writer does not see the evidence for the latter 78 bulletin: museum of comparative zoology bridge, and thinks that the mountain islands of Cuba may have been the asylums which retained the older fauna and received waifs from the younger one. Barbour holds that the younger bridge lasted the longer, but this does not seem to be a necessary postulate, if the mountain islands were of long endur- ance. The island of Hispaniola was clearly the meeting ground for both sets of migrants, and the center from which they radiated west into Cuba and east into Puerto Rico, etc. To the writer these two sets of migrations are best ex- plained as follows: It appears to him that no bridge existed from Cuba to Yucatan after Triassic time, and more especially none during the later Ceno- zoic, since the latter land was then widely beneath the sea, and as for Mesozoic connection, it also appears improbable for the same reason. The only bridge that seems probable, from the geological evidence, is that from Honduras- Nicaragua to Jamaica and Hispaniola. The latter island, however, has two faunas that are more or less separated by a mountain barrier, a northern assemblage with Cuban affinities and a southern one whose relations are dis- tinctly with Jamaica. As the writer sees the physical evidence, the Antillean basin broke down from the Gulf of Mexico southwest across the Central Ameri- can geanticline, first cutting off Cuba from Central America, then sending its waters east to the south of Cuba, next separating Jamaica from Cuba but not from Haiti, and eventually cutting Cuba off from Hispaniola; thus, the Hon- duras-Jamaica-Hispaniola bridge was the last part of the Antillean geanticline to break down." It seems to me that this statement covers the whole situation very satisfactorily. The details concerning the time and place of "land bridges" will always be subject to perfectly reasonable differences of opinion, inasmuch as proof often is impossible and the evidence may be variously interpreted. The writer feels, however, that Doctor Schuchert has made out a very strong case for his own view. There has been so much of what may be called "rippling" along the whole length of the broken land mass which includes Cuba on one end and the Virgin Islands on the other, that there can be little doubt but that most or all of these lands have been under water at least once or perhaps more often. The important point being that they have never all been completely under at the same time. This statement is probably true, in part at least, for the Lesser Antillean Islands as well. It is a general thesis that large islands support a fauna of many species and small islands do not. The abundance of different types present seems in many cases to be definitely a function of the size of the island. This plays a part in explaining conditions as we see them now. The fauna of Cuba, however, today may represent the combined population of descendants of the fauna of a considerable archipelago. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 79 I have discussed elsewhere (Proc. Boston Soc. Nat. Hist. 40, Feb. 1935, p. 351 et seq.) the fauna of the Bahama Islands. Since this was written a correspondent on the Island of Exuina has sent to the Mu- seum of Comparative Zoology a box of bones found in a small, undis- turbed corner of a cave from which cave earth was being taken for fertilizer. This find consisted of the remains of several hundred indi- viduals of a Gcocapronu/s, besides the remains of some extraordinarily large hawks and owls, representing new extinct genera and species. These have been studied by Doctor Alexander Wetmore.^ Each re- curring find of this sort emphasizes the extraordinary changes which have taken place in the Bahamas during the last couple of centuries, or less. These giant hawks lived, beyond doubt, in a high forest and, in- deed, Columbus speaks of the big trees which he found upon landing at San Salvador. Of course Columbus may have been accustomed to a landscape in Spain, arid and with little forest, even 400 years ago; hence we may perhaps take his speaking of the forests on San Salvador with a grain of salt. On the other hand I suspect that he spoke accu- rately. The only remnant of forest and, indeed, it can hardly be called that today, is a stand of really large, old Gumbo limbo {Bursera sima- ruba) trees which still stand, sheltered by a low ridge called the Vic- toria Hills. I do not believe this covers over 50 acres, perhaps not two- thirds as much, and here, and here only, are to be found the small population of Cmfurus nycanus. I very much doubt whether there are over 40 or 50 of these birds and should this bit of woodland go the Woodpeckers would go too. This provides a vivid demonstration of what has happened in the past to the forest fauna of the Bahamas. While these islands mav have been directlv connected with the Greater Antilles, the present poverty of their fauna cannot be used as an argument to support this view for it is increasingly clear that the fauna has not always been as poor in species as it is now. It seems to me, in the final analysis, in speculating concerning the origin of the fauna of the various island groups that two major premises must be kept in mind; first, to consider all of the animals of each island and not simply to consider the evidence based on the conditions in one group alone. Then second, the solution which most easily explains any given situation is inherently the most probable one. There has been undoubtedly some dispersal by flotsam and jetsam and some dispersal by winds and some transport by migrating birds and a good many types have been carried by man, both primitive and civilized. One of 'Bull. M. C. Z., 52, 12, Oct. 1937, p. 427-441, pi. 1. 80 bulletin: museum of comparative zoology my colleagues argues for hurricane dispersal, all sorts of creatures often being carried in the rolled up "boots" of royal palm leaves. These, however, would mostly, by probability, if ever really blown away, have landed in the sea and when they did — if ever — crash to land on some distant island — or one near at hand for that matter, the passengers would have to be sturdy indeed to withstand this method of landing. Nevertheless the theory is ingenious and intriguing and may occasion- ally have functioned — but it is not fair to discuss his ideas before he has even had a chance to publish them. They may well turn out to be more generally popular than my own. However, to conclude that all of the animals of an island, such as any one of the Greater Antilles, have been derived by any or all of these causes is to support an ex- planation which is to my way of thinking infinitely less probable than to postulate extensive changes in land form in a region where so much tectionic movement is evident on every hand. That the separation of the Greater Antilles took place a long time ago is certain for Cadea was not derived from Amphisbaena nor Cricocaura differentiated from its Xantusiid forebears except in a very long time. Professor Daly's ("The Changing World of the Ice Age," Yale Univ. Press, New Haven, 1934, pp. I-IXX and 1-271) ingenious and con- vincing theorem that vast bodies of water have been removed from oceanic circulation and tied up in the form of poiat ice during the vari- ous periods of glaciation, thus reducing the general level of the surfaces of the oceans, would throw most of the Bahaman archipelago into a few, vast islands and even if the amount of water so tied up was only as much as Daly postulates and far less than that presumed by Shepard (Zeit. fiir Geomorph., 9, 1935, pp. 99-105), great changes in topography would be brought about, and there is no reason to suppose that in a region where upthrust and downthrust block faulting seems to be prevalent that many channels between the islands may have been much shallower than they are now or even non-existent but a short time ago. Four hundred and seventy-nine named forms are listed in this paper. In my West Indian Zoogeography of 1914 I listed two hundred and eighty-one forms from the area covered by this paper, in which the Swan Island forms are not listed, as they were in 1914. For much pertinent comment and useful information I have first and foremost heartily to thank Major Chapman Grant. My colleagues Arthur Loveridge and Benjamin Shreve have also frequently and generously discussed problems and given me much useful advice. I have also had generous help from Dr. Stejneger and Dr. Dunn. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 81 SYSTEMATIC TABLE OF CONTENTS A. Class AMPHIBIA Order SALIENTIA Family HYLIDAE Hyla septentrionalis septentrionalis (Boulenger) 94 Hyla septentrionalis dominicensis (Tsehudi) 94 Hyla septentrionalis brunnea (Gosse) 94 Hyla vasta Cope 94 Hyla lichenata (Gosse) 95 Hyla pulchrilineata Cope 95 Hyla wilderi Dunn 95 Hyla marianae Dunn 95 Hyla heilprini Noble 95 Hyla rubra Daudin 95 Family BUFONIDAE Bufo longinasus longinasus (Stejneger) 96 Bufo longinasus dunni (Barbour) 96 Bufo longinasus ramsdeni (Barbour) 96 Bufo peltaeephalus Tsehudi 96 Bufo empusus (Cope) 96 Bufo gutturosus Latreille 97 Bufo lemur lemur (Cope) 97 Bufo lemur turpis (Barbour) 97 Bufo marinis (Linne) 97 Family LEPTODACTYLIDAE Eleutherodactylus auriculatus auriculatus (Cope) 97 Eleutherodaetylus auriculatus sonans (Dunn) 97 Eleutherodactylus auriculatus auriculatoides (Noble) ... 98 Eleutherodactylus auriculatus portoricensis Schmidt .... 98 Eleutherodactylus cooki Grant 98 Eleutherodactylus audanti Cochran 98 Eleutherodactylus wetmorei Cochran 98 Elevitherodactylus jugans Cochran 98 Eleutherodactylus armstrongi Noble & Hassler 98 Eleutherodactylus jamaicensis Barbour 99 Eleutherodactylus lentus lentus (Cope) 99 82 bulletin: museum of comparative zoology Eleiitherodactylus lentus weinlandi (Barbour) 99 Eleutherodactylus lentus richmondi (Stejneger) 99 Eleutherodactylus lentus schmidti (Noble) 99 Eleutherodactylus glandulif er Cochran 99 Eleutherodactylus glanduliferoides Shreve 99 Eleutherodactylus brevirostris Shreve 100 Eleutherodactylus inoptatus (Barbour) 100 Eleutherodactylus darlingtoni Cochran 100 Eleutherodactylus ruthae Noble 100 Eleutherodactylus urichii (Boettger) 100 Eleutherodactylus martinicensis (Tschudi) 100 Eleutherodactylus brittoni Schmidt 100 Eleutherodactylus abbotti Cochran 101 Eleutherodactylus bakeri Cochran 101 Eleutherodactylus montanus Schmidt 101 Eleutherodactylus semipalmatus Shreve 101 Eleutherodactylus pictissimus Cochran 101 Eleutherodactylus femur-laevis Cochran 101 Eleutherodactylus minutus Noble 101 Eleutherodactylus rufifemoralis Noble & Hassler 101 Eleutherodactylus orcutti Dunn 102 Eleutherodactylus cunctator Dunn 102 Eleutherodactylus nubicola Dunn 102 Eleutherodactylus luteolus (Gosse) 102 Eleutherodactylus gossei Dunn 102 Eleutherodactylus pantoni Dunn 102 Eleutherodactylus junori Dunn 102 Eleutherodactylus cundalli Dunn 102 Eleutherodactylus grabhami Dunn 102 Eleutherodactylus andrewsi Lynn 103 Eleutherodactylus alticola Lynn 103 Eleutherodactulus varleyi Dunn 103 Eleutherodactylus parvus Barbour & Shreve 103 Eleutherodactylus atkinsi atkinsi Dunn 103 Eleutherodactylus atkinsi orientalis Barbour & Shreve . . 103 Eleutherodactylus varians (Gundlach & Peters) 104 Eleutherodactylus eileenae Dunn 104 Eleutherodactylus dimidiatus (Cope) 104 Eleutherodactylus emiliae Dunn 104 Eleutherodactylus albipes Barbour & Shreve 104 Eleutherodactvlus intermedins Barbour & Shreve 104 BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 83 Eleutlierodactylus pinarensis Dunn 104 Eleutherodactylus greyi Dunn 104 Eleutlierodactylus brevipalmatus Schmidt 105 Eleutherodactylus sierrae-maestrae Schmidt 105 Eleutherodactylus turquinensis Barbour & Shreve 105 Eleutherodactylus ricordii (Dumeril & Bibron) 105 Eleutherodactylus cuneatus (Cope) 105 Eleutherodactylus gundlachii Schmidt 105 Eleutherodactylus casparii Dunn 105 Eleutherodactylus gryllus Schmidt 105 Eleutherodactylus cochranae Grant 106 Eleutherodactylus locustus Schmidt 106 Eleutherodactylus cramptoni Schmidt 106 Eleutherodactylus antillensis (Reinhardt & Liitken) .... 106 Eleutherodactylus wrightmanae Schmidt 106 Eleutherodactylus unicolor Stejneger 106 Eleutherodactylus monensis (Meerwarth) 106 Eleutherodactylus flavescens Noble 106 Eleutherodactylus karlschmidti Grant 106 Leptodactylus fallax Muller 107 Leptodactylus dominicensis Cochran 107 Leptodactylus albilabris (Giinther) 107 Leptodactylus validus Garman 107 Family BRACHYCEPHALIDAE Sminthillus limbatus limbatus (Cope) 107 Sminthillus Hmbatus orientalis Barbour & Shreve 107 Class REPTILIA Order SQUAMATA Suborder SAURL\ Family GEKKONIDAE Gymnodactylus fasciatus Dumeril & Bibron 108 Gonatodes albogularis (Dumeril & Bibron) 108 Gonatodes notatus (Reinhardt & Liitken) 108 Gonatodes fuscus (Hallowell) 108 Phyllodactylus spatulatus Cope 108 Phyllodactylus martini Van Lidth de Jeude 109 Hemidactylus maljouia (Moreau de Jonnes) 109 84 bulletin: museum of comparative zoology Hemidactylus brookii Gray 109 Hemidactylus turcicus (Linne) 109 Thecadactylus rapicaudus (Houttuyn) 109 Aristelliger praesignis (Hallowell) 109 Aristelliger lar Cope 110 Aristelliger expectatus Cochran 110 Aristelliger cochranae Grant 110 Aristelliger barbouri (Noble & Klingel) 110 Tarentola ciibana Gundlach & Peters 110 Sphaerodactylus roosevelti Grant 110 Sphaerodactylus decoratus Garman 110 Sphaerodactylus stejnegeri Cochran 11 Sphaerodactylus gibbus Barbour 11 Sphaerodactylus torrei Barbour 11 Sphaerodactylus cinereus Wagler 11 Sphaerodactylus mariguanae Cochran 11 Sphaerodactylus oxyrrhinus Gosse 11 Sphaerodactylus armstrongi Noble & Hassler 11 Sphaerodactylus difficilis Barbour 11 Sphaerodactylus altavelensis Noble & Hassler 112 Sphaerodactylus notatus Baird 112 Sphaerodactylus macrolepis Giinther 112 Sphaerodactylus danforthi Grant 112 Sphaerodactylus grandisquamis Stejneger 112 Sphaerodactylus monensis (Meerwarth) 112 Sphaerodactylus townsendi Grant 112 Sphaerodactylus richardsoni Gray 112 Sphaerodactylus becki Schmidt 113 Sphaerodactylus inaguae Noble & Klingel 113 Sphaerodactylus gilvitorques Cope 113 Sphaerodactylus nigropunctatus Gray 113 Sphaerodactylus caicosensis Cochran 113 Sphaerodactylus corticolus Garman 113 Sphaerodactylus festus Barbour 113 Sphaerodactylus goniorhynchus Cope 113 Spaerodactylus argus argus (Gosse) 114 Sphaerodactylus argus bartschi (Cochran) 114 Sphaerodactylus argus argivus (Garman) 114 Sphaerodactylus anthracinus Cope 114 Sphaerodactylus copei Steindachner 1 14 Sphaerodactylus scaber Barbour & Ramdsen 114 BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 85 Sphaerodactylus sanianaensis ("ochruii 114 Sphaerodactylus funtasticus Duineril & Bibron 114 Sphaerodactylus pictus Garman 115 Sphaerodactylus sputator (Sparrmaii) 115 Sphaerodactylus elegantulus Barbour 115 Sphaerodactylus microlepis Reinhardt & Liitken 115 Sphaerodactylus klauberi Grant 115 Sphaerodactylus gaigeae Grant 115 Sphaerodactylus vincenti Boulenger 115 Sphaerodactylus nicholsi Grant 116 Sphaerodactylus monilifer Barbour 116 Family IGUANIDAE Iguana iguana iguana (Linne) 116 Iguana iguana rhinolopha (Wiegmann) 116 Iguana delicatissima Laurenti 116 Chamaeleolis chamaeleonides (Dumeril & Bibron) 117 Xiphocercus valenciennesii (Dumeril & Bibron) 117 Xiphocercus darlingtoni Cochran 117 Chamaelinorops barbouri Schmidt 117 Chamaelinorops wetmorei Cochran 117 Audantia armouri Cochran 117 Deiroptyx vermiculata (Dumeril & Bibron) 117 Deiroptyx bartschi Cochran 118 Anolis equestris equestris Merrem 118 Anolis equestris luteosignifer (Xoble & Hassler) 118 Anolis equestris noblei Barbour & Shreve 1 18 Anolis equestris hassleri Barbour & Shreve 118 Anolis cuvieri Merrem 118 Anolis roosevelti Grant 118 Anolis ricordii Dumeril & Bibron 119 Anolis garmani Stejneger 119 Anolis porcatus porcatus (Gray) 119 Anolis porcatus maynardi (Garman) 119 Anolis porcatus brunneus (Cope) 119 Anolis porcatus smaragdinus (Barbour and Shreve) 119 Anolis porcatus fairchildi (Barbour and Shreve) 120 Anolis porcatus longiceps Schmidt 120 Anolis bohorucoensis Noble & Hassler 120 Anolis chloro-cvanus Dumeril & Bibron 120 86 bulletin: museum of comparative zoology Anolis allogus allogus (Barbour & Ramsden) 120 Anolis allogus mestrei (Barbour & Ramsden) 120 Anolis allogus ahli (Barbour) 120 Anolis bimaculatus Sparrman 120 Anolis evermanni Stejneger 121 Anolis krugi acutus (Hallowell) 121 Anolis krugi krugi (Peters) 121 Anolis krugi wattsi (Boulenger) 121 Anolis krugi fori-esti (Barbour) 121 Anolis krugi gingivinus Cope 121 Anolis gundlachi Peters 121 Anolis sabanus Garman 122 Anolis leachii leachii Dumeril & Bibron 122 Anolis leachii antiquae (Barbour) 122 Anolis leachii lividus (Garman) 122 Anolis leachii barbudensis (Barbour) 122 x\nolis leachii terrae-altae (Barbour) 122 Anolis leachii alliaceus (Cope) 122 Anolis leachii nubilus (Garman) 123 Anolis asper Garman 123 Anolis richardii Dumeril & Bibron 123 Anolis cristatellus cristatellus (Dumeril & Bibron) 123 Anolis cristatellus wileyi Grant 123 Anolis cristatellus cooki Grant 123 Anolis cristatellus monensis (Stejneger) 123 Anolis alutaceus alutaceus (Cope) 124 Anolis alutaceus clivicolus Barbour and Shreve 124 Anolis spectrum Peters 124 Anolis cyanopleurus Cope 124 Anolis semilineatus Cope 124 Anolis olssoni Schmidt 124 Anolis hendersoni Cochran 124 Anolis poncensis Stejneger 125 Anolis pulchellus Dumeril & Bibron 125 Anolis latirostris Schmidt 125 Anolis stratulus Cope 125 Anolis coelestinus Cope 125 Anolis distichus distichus (Cope) 125 Anolis distichus distichoides (Rosen) 125 Anolis distichus dominicensis (Reinhardt & Liitken) .... 126 Anolis distichus caudalis (Cochran) 126 BARBOUR. ANTILLEAN REPTILES AND AMPHIBIANS 87 Anolis distichus wetmorei (Cochran) 126 Anolis distichus altavelensis (Noble & Hassler) 126 Anolis distichus juHae Cochran 126 Anolis sagrei sagrei (Dumcril & Bibron) 126 Anolis sagrei ordinatus (Cope) 126 AnoHs monticola Shreve 127 Anolis luteosignifer Garman 127 Anolis lineatopus Gray 127 Anolis homolechis homolechis (Boulenger) 127 Anolis homolechis rubribarbus (Barbour & Ramsden) . . . 127 Anolis homolechis quadriocellifer (Barbour & Ramsden) . 127 Anolis homolechis patricius (Barbour) 127 Anolis greyi Barbour 128 Anolis cybotes cybotes (Cope) 128 Anolis cybotes doris (Barbour) 128 Anolis cybotes longitibialis (Noble) 128 Anolis angusticeps angusticeps Hallowell 128 Anolis angusticeps oligaspis Cope 128 Anolis isolepis Cope 128 Anolis lucius Dumeril & Bibron 129 Anolis argenteolus Cope 129 Anolis argillaceus Cope 129 Anolis bremeri Barbour 129 Anolis loysiana Cocteau 129 Anolis leucophaeus leucophaeus (Garman) 129 Anolis leucophaeus albipalpebralis (Barbour) 130 Anolis leucophaeus mariguanae Cochran 130 Anolis leucophaeus sularum Barbour and Shreve 130 Anolis roquet roquet (Lacepede) 130 Anolis roquet marmoratus (Dumeril & Bibron) 130 Anolis roquet luciae Garman 130 Anolis roquet vincentii Garman 130 Anolis roquet gentilis Garman 131 Anolis roquet extremus (Garman) 131 Anolis opalinus Gosse 131 Anolis iodurus Gosse 131 Anolis grahami grahami Gray 131 Anolis grahami conspersus Garman 131 Norops ophiolepis (Cope) 131 Cyclura figginsi Barbour 131 Cyclura portoricensis Barbour 132 88 bulletin: museum of comparative zoology Cyclura mattea Miller Cyclura pinguis Barbour Cyclura cornuta cornuta (Bonnaterre) Cyclura cornuta stejnegeri (Barbour & Noble) Cyclura cornuta nigerrima (Cope) Cyclura coUei Gray Cyclura carinata carinata (Harlan) Cyclura carinata bartschi Cochran Cyclura nuchalis Barbour & Noble Cyclura rileyi Stejneger ; . . Cyclura inornata Barbour & Noble Cyclura baeolopha Cope Cyclura caymanensis Barbour & Noble Cyclura macleayi Gray Cyclura ricordii (Dumeril & Bibron) Leiocephalus carinatus carinatus (Gray) Leiocephalus carinatus armouri Barbour & Shreve . . . . Leiocephalus carinatus coryi Schmidt Leiocephalus carinatus hodsdoni Schmidt Leiocephalus carinatus punctatus Cochran Leiocephalus carinatus picinus Barbour & Shreve . . . . Leiocephalus carinatus helenae Barbour & Shreve . . . . Leiocephalus carinatus virescens (Stejneger) Leiocephalus carinatus varius Garman Leiocephalus melanochlorus Cope Leiocephalus schreibersii (Gravenhorst) Leiocephalus personatus personatus (Cope) Leiocephalus personatus aureus Cochran Leiocephalus personatus mentalis Cochran Leiocephalus personatus scalaris Cochran Leiocephalus personatus louisae Cochran Leiocephalus eremitus Cope Leiocephalus cubensis Gray Leiocephalus greenwayi Barbour & Shreve Leiocephalus psammodromus Barbour I^eiocephalus raviceps Cope Leiocephalus loxogrammus loxogrammus (Cope) Leiocephalus loxogrammus parnelli Barbour & Shreve. Leiocephalus macropus Cope Leiocephalus inaguae Cochran Leiocephalus semilineatus Dunn 132 132 132 132 132 132 132 133 133 133 133 133 133 133 134 134 134 134 134 134 134 135 135 135 135 135 135 136 136 136 136 136 136 136 137 137 137 137 137 137 137 BARBOUR ANTILLEAN REPTILES AND AMPHIBIANS 89 Leiocephalus barahonensis Schmidt 138 Leiocephalus beatanus Noble 138 Leiocephalus vinculum Cochran 138 Hispaniolus pratensis Cochran 138 Family ANGUIDAE Celestus de la sagra de la sagra (Cocteau) 138 Celestus de la sagra nigropunctata Barbour & Shreve. . . . 138 Celestus rugosus Cope 138 Celestus costatus (Cope) 139 Celestus badius Cope 139 Celestus maculatus (Garman) 139 Celestus occiduus (Shaw) 139 Celestus impressus Cope 139 Celestus pleii (Dumeril & Bibron) 139 Sauresia sepoides Gray 139 Wetmorena haetiana Cochran 140 Family XANTUSTIDAE Cricosaura typica (Gundlach & Peters) 140 Family TEIIDAE Kentropyx intermedins Gray 140 Ameiva aquilina Garman 140 Ameiva fuscata Garman 140 Ameiva cineracea Barbour & Noble 140 Amieva atrata Garman , 141 Ameiva pluvianotata Garman 141 Ameiva erythrops Cope 141 Ameiva griswoldi Barbour 141 Ameiva erythrocephala (Daudin) 141 Ameiva garmani Barbour 141 Ameiva pleii Dumeril & Bibron 141 Ameiva corvina Cope 142 Ameiva polops Cope 142 Ameiva wetmorei Stejneger 142 Ameiva eleanorae Grant and Roosevelt 142 Ameiva maynardi maynardi Garman 142 Ameiva maynardi uniformis Noble & Klingel 142 90 bulletin: museum of comparative zoology Ameiva maynardi parvinaguae Barbour & Shreve 142 Ameiva alboguttata Boulenger 143 Ameiva birdorum Grant 143 Ameiva exsul Cope 143 Ameiva vittipimctata Cope 143 Ameiva taeniura Cope 143 Ameiva chrysolaema chrysolaema Cope 143 Ameiva chrysolaema abbotti Noble 144 Ameiva chrysolaema juliae Cochran 144 Ameiva barbouri Cochran 144 Ameiva thoracica Cope 144 Ameiva dorsalis Gray 144 Ameiva auberi Cocteau 144 Ameiva rosamondae Cochran 144 Ameiva beatensis Noble 145 Ameiva navassae Schmidt 145 Scolecosaurus alleni alleni (Barbour) 145 Scolecosaurus alleni parviceps Barbour 145 Gymnophthalmus pleei Bocourt 145 Family AMPHISBAENIDAE Cadea palirostrata Dickerson 145 Cadea blanoides Stejneger 146 Amphisbaena fenestrata Cope 146 Amphisbaena bakeri Stejneger 146 Amphisbaena caeca Cuvier 146 Amphisbaena manni Barbour 146 Amphisbaena innocens Weinland 146 Amphisbaena cubana Peters 146 Amphisbaena caudalis Cochran 146 Family SCINCIDAE Mabuya mabouia (Dumeril & Bibron) 147 Mabuya lineolata Noble & Hassler 147 Suborder OPHIDIA Family TYPHLOPIDAE Typhlops tenuis Salvin 147 Typhlops rostellatus Stejneger 148 BARBOUR: AXTILLEAN REPTILES AND AMPHIBIANS 91 Typhlops richardii Dumeril & Bibron 148 Typhlops pusillus Barbour 148 Typhlops domiiiicana Stejneelly scales. Sphaerodactylus gaigeae Grant Mountains between Maunabo and Yabacoa, Puerto Rico. A small dark colored species known only from the collection of Major Chapman Grant whose unbounded industry has made him the peerless authority on the herpetology of the Puerto Rican area. Sphaerodactylus vincenti Boulenger St. Vincent. No information available as to present status. 116 bulletin: museum of comparative zoology Sphaerodactylus nicholsi Grant Puerto Rico. Said to be somewhat similar to the species from St. Vincent. A chance resemblance no doubt. Sphaerodactylus monilifer Barbour Dominica. Probably abundant but I have no real information about this species. Family IGUANIDAE Iguana iguana iguana (Linne) St. Thomas, Water Island, Hassel Island, Tortola, Peter Island, Guana Island, St. John, Saba, Grenada, Tobago, Trinidad, tropical lowlands of South Amer- ica from western Panama to Brazil. Dr. Dunn has recently examined all available material of the genus Iguana and this arrangement is based on his conclusions. (Copeia, 1934, p. 1.) Iguana iguana rhinolopha (Wiegmann) ?St. Kitts, ?St. Lucia, Swan Island, lowlands of tropical Central America from Costa Rica northward in rain forest areas to the states of Guerrero and Vera Cruz, Mexico. The Swan Island specimens are unstable and many possess and many lack the nasal spines. The Antillean specimens are probably based on specimens incorrectly labelled as to locality. If there really ever were iguanas on these islands, the mongoose has exterminated them. There is what may be an iguana egg from St. Lucia in the Mus. Comp. Zool. It is so labelled, and it was taken many years ago. Iguana delicatissima Laurenti Anguilla, St. Martins, St. Bartholomew, St. Eustatius, Nevis, Guadeloupe, Les Saintes. This species has been recorded from Swan Island, where it is not now found and from the Caymans where it is either very rare or occa- sionally brought in by the very widely seafaring people. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 117 Chamaeleolis chamaeleonides (Dumeril & Bihron) Cuba. The most peculiar of all the offshoots from the Anoline stock. A rare species and beyond doubt a monotypic genus, in spite of several names applied with the idea of multiplying the forms. XiPHOCERCUS VALENCIENNESII (Dunieril & Bihron) Jamaica. Not uncommon in woods and fruit plantations. It may be related to Phenacosaurus of Colombia or be simply a chance offshoot from Anolis in Jamaica and Haiti and only fortuitously similar to the South American genus. XiPHOCERCUS DARLINGTONI Cochran Haiti. A surprising discovery, made in 1935 by Dr. Darlington of Harvard at Roche Croix, Massif de la Hotte, 5,000 ft. Another Jamaican genus in Hispaniola. Chamaelinorops barbouri Schmidt Navassa. Not found during the careful exploration of Clench, Schevill and Rehder during January, 1930. Possibly exterminated by introduced animals. Chamaelinorops wetmorei Cochran Hispaniola. The unique type is from near Miragoane, Haiti. Audantia armouri Cochran Haiti. Recently discovered on the Morne La Selle. It resembles Plica or Leiocephalus superficially but more probably it represents the stock of the following genus. Found by Dr. Darlington also on Morne La Hotte. Deiroptyx vermiculata (Dumeril & Bibron) Cuba. Bank of streams of Pinar del Rio Province, taking refuge in the water and hiding among submerged rocks and stones when pursued. lis bulletin: museum of comparative zoology Deiroptyx bartschi Cochran Cuba. Long unrecognized but not rare in western Cuba. Anolis equestris equestris Merrem Havana Province to Western Oriente, Cuba. The finest and largest form in the genus. Rather uncommon every- where but wide ranging. Less common than its allies, A. garmani of Jamaica and A. ricordii of Hispaniola, and about equally abundant with A. cuvieri of Puerto Rico. These are the "Giant Anoles" of the Antilles and they may be related to the A. insigiiis group of Central America. Anolis equestris luteosignifer (Noble & Hassler) Western Cuba. Replaces the preceding form in the Pinar del Rio area east to about San Antonio de los Baiios in Havana province. Anolis equestris noblei Barbour & Shreve Eastern Cuba. Replaces the typical forms from Nipe Bay to the Mantanamo basin. Anolis equestris hassleri Barbour & Shreve Island of Pines. The representative form on this island. Anolis cuvieri Merrem Puerto Rico and Vieques. A rather uncommon member of the series of "Giant Anoles." Recorded from Tortola but Major Grant doubts its occurrence there. Anolis roosevelti Grant Culebra. Apparently a very fine and distinct form. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 119 Anolis ricordii Dumeril & Bibron Hispaniola. One of the "Giant" series. Found throughout the whole Island and next to A. garmani of Jamaica the most abundant of the tribe. Anolis garmani Stejneger Jamaica. The beautiful great green or barred "Venus Lizard" of Jamaica. A common woodland form, by far the most abundant of the group of the "Giant iVnoles." Anolis porcatus porcatus (Gray) Cuba and Isle of Pines. A very abundant species. The "Chamaeleon" now sold iniquitously by thousands at "the circus." It has replaced its ally, our southern "Chamaeleon," A. carolinensis (Voight) in this hateful traffic. Anolis porcatus maynardi (Garman) Grand Cayman. This extraordinary lizard, the most extreme member of the long- headed series, is by no means common. Anolis porcatus brunneus (Cope) Crooked Island, and the neighboring islands, and probably also Watlingg Island. A fine series of topotypes defines this beautiful species, long confused for lack of topotypes. Anolis porcatus smaragdinus (Barbour and Shreve) Bahamas. The species which has been called A. porcahis and A. hnmnens by recent authors but which is a perfectly distinct species inhabiting the islands of the Great Central Bahama Bank, Andros, New Providence, Eleuthera, Long, etc. The common green anole of the Central Ba- hamas. 120 bulletin: museum of comparative zoology Anolis porcatus fairchildi (Barbour and Shreve) Cay Sal Group, Bahamas. A green ancle of the poreatus-principalis-smaragdinus-brunneus series, perfectly distinct and confined to this isolated group of islets. Anolis porcatus longiceps Schmidt Navassa. Apparently the only species at present to be found in any number on this Island. Anolis bohorucoensis Noble & Hassler San Domingo. A fine species apparently confined to the Sierra de Bohoruco, south- ern San Domingo. Anolis chloro-cyanus Dumeril & Bibron Hispaniola. A widespread and not uncommon form. Anolis allogus allogus (Barbour & Ramsden) Cuba. This fine form has a wide distribution in the mountains of eastern Cuba. Anolis allogus mestrei Barbour & Ramsden Cuba. A rather rare species of the higher woods in the limestone hills of western Cuba. Anolis allogus ahli Barbour Cuba. Confined to the mountains between Trinidad and Cienfuegos. Not uncommon in high damp woods. Anolis abatus Ahl probably belongs here. Anolis bimaculatus Sparrman St. Eustatius, St. Kitts and Nevis. Abundant. A strictly arboreal species. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 121 Anolis evermanni Stejneger Puerto Rico. A highland species which may be related to A. Icucophaeus of Inagua. An abundant form. Anolis krugi acutus (Hallowell) St. Croix. This is still an abundant form. I have just received a fine series. Major Grant agrees that A. newtoni belongs here as a synonym. Anolis krugi krugi (Peters) Puerto Rico. A small species belonging to what I call the rupicolous as against the arboreal Lesser Antillean series. Anolis krugi wattsi (Boulenger) St. Kitts, Nevis, St. Eustatius and Antigua. A pretty little species found on the outcrops of igneous rock and, insofar as my experience goes, not in trees. It is one of the A. acutus allies. Anolis krugi forresti (Barbour) Barbuda. Mr. Parker has recently let me see more material from this island. The form is close to the preceding but, I think, quite valid. Anolis krugi gingivinus Cope St. Martins, St. Barts, Anguilla and St. Eustatius. Common. A member of the series of small sized Lesser Antillean species. Anolis gundlachi Peters Puerto Rico. Apparently an abundant species. 122 bulletin: museum of comparative zoology Anolis sabanus Garman Saba. A most remarkably differentiated form, a rock lizard, pure and simple. The males with. really leopard-like spotting. It is so well defined that I think it had best stand alone. . Anolis leachii leachii Dumeril & Bibron Guadeloupe. This form having the oldest name heads the series comprising most of the large arboreal Lesser Antillean Auoles. Anolis leachii antiquae (Barbour) Antigua. A beautiful and common arboreal species. Anolis leachii lividus (Garman) Montserrat. All the lizards are said still to be common on this Island. Anolis leachii barbudensis (Barbour) Barbuda. Mr. Parker of the British Museum has just allowed me to examine some specimens of this form hitherto known from the type only. It now appears that this race is very close if not really indistinguishable from the form on Antigua. More material from both islands is needed to settle the question. Anolis leachii terrae-altae (Barbour) Les Saintes; near Guadeloupe. A fine big species which Noble found abundant in 1914. Anolis leachii alliaceus (Cope) Dominica. I was surprised in 1929 to find that this species seemed much less conspicuous and common than its allies on other islands nearby. So much for what may have been a most erroneous conclusion drawn from the visit of a few days only. It is, however, by no means rare. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 123 Anolis leachii nubilus (Garman) Redonda. A beautiful great lizard; one of the finest in the genus. It is known only from the original series. Anolis asper Garman Marie Galante. A bizarre and gorgeous species common on the old mango trees — about the only trees still standing over a large part of this hurricane- stricken isle. This form is so distinct and so highly specialized that it must surely stand as a full species although no doubt it belongs in this category as far as ancestry is concerned. Anolis richardii Dumeril & Bibron Grenada and Tobago. A splendid great lizard; a strict tree-dweller. Anolis cristatellus cristatellus (Dumeril & Bibron) Puerto Rico, Vieques, St. Thomas, St. John, St. James, Anegada, Fallen Jeru- salem, Tortola, Virgin Gorda, Guana Island, Peter Island, Water Island and Mosquito Island. A common and handsome species. It has been suggested that a separate genus be established for the fin-tailed species, but as a matter of fact this character appears in \'arious phyla and it may not always be a token of relationship. Anolis cristatellus wileyi Grant Culebra. A form differing in color, and apparently constantly, from the typical race and found on Culebra and the surrounding Cays. Anolis cristatellus cooki Grant Southwestern Puerto Rico. A well defined race confined to the desert area about La Brea Point. Anolis cristatellus monensis (Stejneger) Mona. Apparently a common species. 124 bulletin: museum of comparative zoology Anolis alutaceus alutaceus (Cope) Cuba and Isle of Pines. Known from all parts of the Island but nowhere abundant. A species of the low scrublands. Anolis alutaceus clivicolus Barbour and Shreve Eastern Cuba. A mountain form which in several areas seems* to intergrade with the preceding race. Anolis spectrum Peters Cuba. A not uncommon lizard in woodlands during the rainy season. It disappears completely during the dry portion of the year. It may tie in with one of the A. semilincatus, A. olssotii, A. hendersoni series of Haiti, Anolis cyanopleurus Cope Cuba. A marvelously beautiful species which Dr. Ramsden has rediscovered in the old type locality, the mountains about Guantanamo. I suspect from the habit that it must be terrestrial. It is said to be local and uncommon. Anolis semilineatus Cope Hispaniola. An abundant cursorial grass-living form. It is not improbable that trinominal designation may be indicated if the ranges of this and the two following forms can be shown not to overlap. Anolis olssoni Schmidt Hispaniola. Apparently a not uncommon member of the group of slender terres- trial species long confused with A. semilincatus and allied to A. spec- trum of Cuba. Anolis hendersoni Cochran Hispaniola. A small terrestrial species mostly, if not wholly, from the western portion of the Island. BARBOUR: AXTILLEAN REPTILES AND AMPHIBIANS 125 Anolis poncensis Stejneger Puerto Rico. A rare local species. One which is terrestrial and almost Norops-like in habit. Anolis pulchellus Dumeril & Bibron Puerto Rico, Vieques, Culebra, St. John, St. James, Virgin Gorda, Tortola, Peter Island, Guana Island, Anegada, St. Thomas, St. Croix, Just van Dyke. A common ground-living species. Doubtfully recorded from Haiti. Anolis latirostris Schmidt Navassa. Known from the unique type only. Now apparently extinct. Possi- bly a terrestrial form, hence a prey to the cats left when the lighthouse was made automatic and the keepers were moved away. Most lizards and all snakes have probably gone from Navassa except Anolis longi- ceps which is strictly arboreal. Anolis stratulus Cope Puerto Rico, Vieques, Culebra, St. John, St. Thomas, Tortola, Peter Island, Guana Island, Fallen Jerusalem and Just van Dyke. A common lowland species. Anolis coelestinus Cope Hispaniola. I have seen this form from Haiti only. Anolis distichus distichus (Cope) Bahama Islands. Common on the ceiba trees on New Providence Island. It occurs on Eleuthera, Long Island, Rum Cay and Watlings Island as well. Mr. Shreve is of the opinion that the Rum Cay form may be distinct but I only got a single specimen there in 1934. Anolis distichus distichoides (Rosen) Andros Island. A poorly defined form replacing A. distichus. It is very abundant. 126 bulletin: museum of comparative zoology Anolus distichus dominicensis (Reinhardt & Liitken) Hispaniola. This species is not uncommon in Haiti but the stock seems to be rare on La Gonave. I secured a small series in 1929 — but in a very dry time. Anolis distichus caudalis (Cochran) . La Gonave Island. Representative of a plastic stock on La Gonave. Anolis distichus wetmorei (Cochran) Beata Island. Confined to this island where it seems to be very rare. Beata is now swarming with feral dogs, cats and goats — - fauna and flora are suffer- ing as one might expect. Ground lizards with whole tails are now rare — as soon the lizards themselves will be also. Anolis distichus altavelensis (Noble & Hassler) Alta Vela Island. A rather poorly defined form. Anolis distichus juliae Cochran Isle Vache. A recently discovered form. Anolis sagrei sagrei (Dumeril & Bibron) Cuba and Isle of Pines; probably introduced into Jamaica and Belize. The commonest Anolis and, as its range is wide in Cuba, perhaps this form has the largest species population in the genus. The common- est fence, house-wall and brush lizard in Cuba, by far. Anolis sagrei ordinatus (Cope) Bahamas. Known from Turks Island to New Providence. Common every- where. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 127 Anolis monticola Shreve Haiti. Found by Dr. Darlington in the eastern foothills of Morne La Hotte. Said to be related to A. sagrei and perhaps should be trinominally designated. Anolis luteosignifer Garman Cayman Brae. Probably as abundant as it ever was. Anolis lineatopus Gray Jamaica. The common fence lizard of the dry Liguanea Plain about Kingston. It swarms here but occurs nowhere else, so far as anyone knows at present. Anolis homolechis homolechis (Boulenger) Cuba and Isle of Pines. A widespread and not uncommon species found in wooded ravines or lowland woods and heavy scrub. Anolis homolechis rubribarbus (Barbour & Ramsden) Cuba. Known only from a very few specimens from Puerto Cananova on the north coast of the oriental province. Anolis homolechis quadriocellifer (Barbour & Ramsden) Cuba. Known only from the Cape San Antonio region of extreme western Cuba. Anolis homolechis patricius (Barbour) Cuba. Only known from a series taken by Dr. Ramsden at Mina Piloto, near Sagua de Tanamo, northern coast of Oriente Province. 128 bulletin: museum of comparative zoology Anolis greyi Barbour Cuba. Only known from a small number taken in the town of Camaguey and in the Cubitas range of hills not far away. Anolis cybotes cybotes (Cope) Hispaniola. Common as are the allies of A. sagrci wherever they occur. This is one of a series of dominant and successful races. Anolis cybotes doris (Barbour) La Gonave. I have now seen a good many specimens of this lizard. We may follow Miss Cochran in giving it subspecific rank. Anolis cybotes longitibialis (Noble) Beata Island. I have found this lizard rare on several visits to Beata. Anolis angusticeps angusticeps Hallowell Cuba and Isle of Pines. I consider this a really rare species in both western and eastern Cuba. It is more abundant in the Isle of Pines. Anolis angusticeps oligaspis Cope Bahamas. Found upon New Providence (Hog Id. type), Andros I., (U.S.N.M.) and Long Island (Barbour). It is the rare representative of A. angusti- ceps of Cuba. It may occur also upon other islands. Much intensive herpetological work remains to be done in the central and southern Bahama Islands. Anolis isolepis Cope Cuba. An excessively rare species. It occurs in the mountains of Oriente Province. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 129 Anolis LUCIUS Dumeril & Bihron Cuba. The abundant lizard of the Hmestone cliffs and open caves of central Cuba from Matanzas and Santa Clara Provinces, especially. Anolis argenteolus Cope Cuba. Found in the Province of Oriente. Far from rare, it occurs on rocks, cliffs, and often also on building walls and fences. I have taken it on the trunks of the great Ficus nitida (Sp. Laurel de la India) trees which used to stand in the Plaza at Santiago. Anolis argillaceus Cope Cuba. I have never seen this species in life. Dr. Ramsden says it is not uncommon in the old coffee plantations high in the mountains about Guantanamo. Anolis bremeri Barbour Cuba. A fine, striking species, known only from the type which I took years ago at Herradura in Pinar del Rio Province. One of the most distinct species in Cuba. Its great maroon-brown gular fan is wholly unlike that of any other Anole. Anolis loysiana Cocteau Cuba. A rare and bizarre little lizard. It is found sparingly all over Cuba on trees having a light colored bark. It is extraordinarily like rough bark in appearance. Some believe that the genus Acantholis proposed to contain this species is really valid. It becomes more common during the summer rains than it is in the dry season, our winter. Anolis leucophaeus leucophaeus (Carman) Inagua. A common species. 130 bulletin: museum of comparative zoology Anolis leucophaeus albipalpebralis (Barbour) Turks and Caicos Islands. This species seems plastic like A. dominicensis. Anolis leucophaeus mariguanae Cochran Mariguana Island. Another good representative race. Anolis leucophaeus sularum Barbour and Shreve Atwood's Cays, Bahamas. A race, about as good as the others, which Mr. Greenway recently found on West Booby Cay in the Atwood's Cay group. Anolis roquet roquet (Lacepede) Martinique. This heads the lot of the smaller Lesser Antillean races. They are less well defined in general than the races of A. leachii. In some cases they can only be told apart while living, their colors then being quite diagnostic. They usually frequent the beach grape and poison wood trees about the shores of the Island. The larger races inhabit the inland forests. To this stock belongs also A. acneus Gray of Trinidad and Anolis honairicnsis Ruthven. Anolis roquet marmoratus (Dumeril & Bibron) Desirade. I know nothing of this form. Garman found it abundant in 1882. Anolis roquet luciae Garman St. Lucia. Apparently, like so many Antillean species, whether from one reason or another much less common than formerly. Anolis roquet vincentii Garman St. Vincent. Like most of the reptiles of this Island, this species is now rare. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 131 Anolis roquet gentilis Garman Grenada and the Grenadines. A rather small, inconspicuous lizard which is still abundant. Anolis roquet extremus (Garman) Barbados. A color race only. Anolis opalinus Gosse Jamaica. A rather rare, woodland species, most often seen in western Jamaica. Anolis iodurus Gosse Jamaica. A beautiful and not uncommon little woodland species. It is found widely distributed on the Island. Anolis grahami grahami Gray Jamaica. Common in the woods of eastern Jamaica. Anolis grahami conspersus Garman Grand Cayman. It is not common. NoROPS oPHiOLEPis (Cope) Cuba and Isle of Pines. A common terrestrial species usually found hiding in the heavy tufts or bunches of pasture grasses. Cyclura figginsi Barbour Bitter Guana Cay, near Great Guana Cay, Exuma group. This little colony is now, I learn, almost certainly exterminated. 132 bulletin: museum of comparative zoology Cyclura portoricensis Barbour Puerto Rico. Extinct but relatively recent bones found in several caves. Cyclura mattea Miller St. Thomas. Recently extinct, known from recent osseous remains only. Cyclura pinguis Barbour Anegada. Rare. Cyclura cornuta cornuta (Bonnaterre) Hispaniola, La Gonave, Petit Gonave and Beata Island. Persisting only in isolated colonies on the larger island but common on Beata, although only old individuals are now to be seen. The eggs are dug up by feral dogs and if any young hatch they are devoured by the feral cats. Cyclura cornuta stejnegeri (Barbour & Noble) Mona. Another rare species. This may be the same as C. cornuta. Cyclura cornuta nigerrima (Cope) Navassa. Extinct. I am not sure that this was really distinct from C. cornuta', in fact, I rather doubt it, but material is lacking to settle the question. Cyclura collei Gray Jamaica. Almost extinct. There are a few on Goat Island, off the Bushy Park property, and a few on the Cays about Montego Bay. Cyclura carinata carinata (Harlan) Turks Island. Abundant still on some Cays near Turks Island and in the Caicos group. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 133 Cyclura carinata bartschi Cochran Booby Cay, east of Mariguana Island. Said to be more or less intermediate between the preceding and following species. Cyclura nuchalis Barbour & Noble Fortune Island; North Cay, Fish Cay in Acklin's Bight. Tracks also seen on Guana Cay of the same group. Abundant on Fish Cay but rare on the other islets of Acklin's Bight. Cyclura rileyi Stejneger Cays and west and south shores of the lagoon of Watlings Island ; (Green Cay and White CajO- Still common. Cyclura cristata Schmidt (type loc. White Cay) seems to be a synonym. Mr. Armour collected a series on Green Cay during the 1934 cruise of the Utowana. Cyclura inornata Barbour & Noble U Cay in AUen's Harbor near Highborn Cay, Bahamas. Once widespread, no doubt now extirpated through use by the negroes for food. This was the only specimen which Maynard could find — a relict on a tiny islet. Cyclura baeolopha Cope Andros Island. Reported to be considerably decreased in numbers. Cyclura caymanensis Barbour & Noble Cayman Brae and Little Cayman. Reported still to be not uncommon. Cyclura macleayi Gray Cuba and Isle of Pines. Persisting in wild and inaccessible districts. 134 bulletin: museum of comparative zoology Cyclura ricordii (Dumeril & Bibron) Hispaniola. Long known from the type only, until rediscovered by Dr. W. L. Abbott. Now known to be not uncommon in a few scattered localities in San Domingo. Leiocephalus carinatus carinatus (Gray) Cuba, Isle of Pines, and Cayman Brae. Widespread about rocky shores, headlands and sea cliffs. So far as I am aware, seldom or never seen inland, certainly never in Cuba. With its tail tightly curled over its back this lizard jumps and hops about its haunts in a most unreptilian manner. The Cayman Brae specimens may represent a separate form but material is too scant to be sure. Leiocephalus carinatus armouri Barbour & Shreve North Bahamas. A distinct race confined to Grand Bahama, the Abacos and nearby Cays. Leiocephalus carinatus coryi Schmidt Bemini Islands. A small race related to L. c. armouri. Leiocephalus carinatus hodsdoni Schmidt Long Island. Another Bahaman race quite distinct and related to the two forms mentioned above. Leiocephalus carinatus punctatus Cochran Acklin's Island, Crooked Island and the Cays in Acklin's Bight. A good, distinct form, probably a species rather than a subspecies. Leiocephalus carinatus picinus Barbour & Shreve Atwood's Cay, Bahamas. An apparently strictly localized form. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 135 Leiocephalus carinatus helenae Barbour & Shre\e Mira por vos Cays. Another very local race. Leiocephalus carinatus virescens (Stejneger) Green Cay, Bahamas. Known from the type series. Leiocephalus carinatus varius Garman Grand Cayman. I have been several times to Grand Cayman for short visits and never saw this species at all. Its allies are all companion denisons of the beach plant association. Leiocephalus melanochlorus Cope Hispaniola. Known from Jeremie in southwest Haiti to Puerto Plata in northern San Domingo. Leiocephalus schreibersii (Gravenhorst) Hispaniola. A common species on Haiti. We have not seen it from San Domingo. Leiocephalus personatus personatus (Cope) Hispaniola. Allied to L. cuhensis. Miss Cochran informs me that the typical race of this species is from southwestern Haiti. I suspect L. Ihrrminicri (Dumeril & Bibron) to be a synonym of this species. It was said to have come from Trinidad and Martinique, L'herminier, and Plee collectors, but both these gentlemen caused confusion on more than one occasion by either labelling their material incorrectly or else by shipping the results of a visit to several islands home to Paris in one lot shipment, after receipt of which the whole consignment was entered in the records of the Jardin des Plantes as having been collected at the point of shipment. This sort of thing has caused confusion for modern workers on a host of occasions. 136 bulletin: museum of comparative zoology Leiocephalus personatus aureus Cochran Haiti. Known only from the region about Jacmel. Leiocephalus personatus mentalis Cochran San Domingo. Apparently confined to the eastern portion of the Republic. Leiocephalus personatus scalaris Cochran Haiti. From the wet, heavily forested part of northern Haiti. Leiocephalus personatus louisae Cochran Saona Island. Confined to this small island. Leiocephalus eremitus Cope Navassa. Not found by Beck or the Clench party last year. Cats and dogs, now feral, may be to blame for the disappearance of this and other species. Leiocephalus cubensis Gray Cuba and Isle of Pines. The common lizard of the canefields. I believe all species with similar habits are highly beneficial in controlling insects which are injurious to the cane. Leiocephalus greenwayi Barbour & Shreve Plana Cays, Bahamas. A very distinct form abundant on East Plana Cay, and probably the same form occurs on the western island. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 137 Leiocephalus psammodromus Barbour Turks Island. A common species and one which I at first called L. arctian'us but found that that name had been obscurely given by Tschudi to a Peru- vian species that apparently had escaped all notice of subsequent authors. Leiocephalus raviceps Cope Cuba. I once doubted the validity of this species but it seems to be really well defined and confined to eastern Cuba. Leiocephalus loxogrammus loxogrammus (Cope) Rum Cay, Bahamas. This species w^ill probably prove to be much more widespread than we now know it to be. Leiocephalus loxogrammus parnelli Barbour & Shreve Watlings Island, Bahamas. A well defined local race. Leiocephalus macropus Cope Cuba. A species found abundantly throughout the Province of Oriente but, so far as we now know, not westward of, let us say, a vertical line drawn north and south and passing about through Holguin. Leiocephalus inaguae Cochran Great Inagua. Common around the coastal region of the island. Leiocephalus semilineatus Dunn Hispaniola. Known only from Thomazeau, Haiti. 138 bulletin: museum of comparative zoology Leiocephalus barahonensis Schmidt Hispaniola. Known only from the southeastern portion of San Domingo. Leiocephalus beatanus Noble Beata Island. Common and the only representative of the genus which either Noble or I. was able to find on the Island. Leiocephalus vinculum Cochran Gonave Island, Haiti. Apparently far from abundant — at least about Anse a Galets. HiSPANiOLUS pratensis Cochran Hispaniola. Taken by Milles at St. Michel, Haiti. Familv ANGUIDAE Celestus de la sagra de la sagra (Cocteau) Western and central Cuba. A widespread but excessively rare and perhaps disappearing species. Celestus de la sagra nigropunctata Barbour & Shreve Eastern Cuba. A well defined color variant. Celestus rugosus Cope Hispaniola. Whether or not this species is really valid remains to be determined when more material comes to hand. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 139 Celestus costatus (Cope) Hispaniola. This species may be the same as C occiduus of Jamaica. These species all change greatly during growth and are rather in confusion taxonomically. Celestus badius Cope Navassa. This species may still occur on Navassa. I have a specimen taken but a few years ago. It may be identical with C costatus. Celestus maculatus (Garman) Cayman Brae. A rather poorly defined but, I think, valid form apparently known from the type only. Celestus occiduus (Shaw) Jamaica. A form which was once common and of which old adults reached a great size — like Tiliqua of Australia or Corucia of the Solomon Islands. No such giants now occur and the species is rare. Celestus impressus Cope Jamaica. A smaller and commoner species than C. occiduus but still one of which we know very little. Celestus pleii (Dumeril & Bibron) Puerto Rico. A species which is much like its Cuban congener but abundant rather than rare. Sauresia sepoides Gray Hispaniola. I once sunk this genus into Celestus but the consensus of opinion is that I was wrong. It seems really to be not uncommon. 140 bulletin: museum of comparative zoology Wetmorena haetiana Cochran Hispaniola. Known from a few examples taken by Wetmore in the higher regions of the La Selle massif in Haiti. Family XANTUSIIDAE Cricosaura typica (Gundlach & Peters) Cuba. Confined to the area, of a few square miles at most, between Belig and Cabo Cruz, Oriente, Cuba. Family TEIIDAE Kentropyx intermedius Gray Northern South America, Barbados. This species apparently was formerly common on Barbados but it is now wholly extinct on that Island. Garman named (K. copei) but did not describe this species. I have recently seen material from Demarara and there is no doubt as to the identity of the Barbados lizards with those from British Guiana. It may have been artificially introduced into Barbados. Ameiva aquilina Garman . St. Vincent and Grenada. Extinct on St. Vincent but still persisting on Grenada. Ameiva fuscata Garman Dominica. Owing to the absence of the mongoose this, the finest of all the Antil- lean Ameivas, is still a common species. Ameiva cineracea Barbour & Noble Guadeloupe. Extirpated except for a few individuals which persist on the tiny islets off the coast. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 141 Amieva atrata Garman Redonda. A black species superficially like .1. corvina and living under similar conditions. It has not been collected recently, probably only because the Island is now almost never visited. Ameiva pluvianotata Garman Montserrat. I have just learned that this species is still very common all over the Island. • Ameiva erythrops Cope St. Eustatius. Peters found this form abundant in 1922. Ameiva griswoldi Barbour Antigua, Nevis and Barbuda. Extinct on Nevis, it is also almost gone on Antigua where it persists only right in the town of St. John in yards and gardens. Mr Parker has recently let me see ground lizards from Barbuda which he believes belong to this species. I think he is correct but the material is not exactly comparable. Ameiva erythrocephala (Daudin) St. Kitts. Extirpated from the wilder parts of the Island; it still occurs in the gardens and yards of Basseterre. Here it is safe from the mongoose. Ameiva garmani Barbour Anguilla. • This species is still abundant. It is closely allied to A. pleii. Ameiva pleii Dumeril & Bibron St. Barts and St. Martin. We have again no recent information to indicate that this is not still an abundant species. 142 bulletin: museum of comparative zoology Ameiva corvina Cope Sombrero. A black form which, like so many Lacertids and some species of Cnemidophorus and indeed another Ameiva, has this peculiar colora- tion associated with isolation on a very small, arid, sunbaked and rocky island. Ameiva polops Cope St. Croix. Extinct, but very few specimens have been preserved. Ameiva wetmorei Stejneger Puerto Rico. Rare and confined to the arid zone about Guanica. This species also belongs to the lineolata-maynardi-polops stock, which thrives only in arid areas. Ameiva eleanorae Grant and Roosevelt Caja de Muertos. A rather ill-defined form confined to this tiny islet off the coast of Puerto Rico. Ameiva maynardi maynardi Garman Great Inagua. A beautiful species of the A. lineolata series, north and west coasts of Inagua. A. leucomelas Cope 1894 is a synonym. Ameiva maynardi uniformis Noble & Klingel Great Inagua. Found commonly from Southwest Point to Couch Shell Point, re- placing the typical form. Ameiva maynardi parvinaguae Barbour & Shreve Little Inagua. A form of well marked and peculiar coloration. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 143 Ameiva alboguttata Boulenger Mona Island. According to recent accounts still abundant. Closely related to the Puerto Rican form next following. Ameiva birdorum Grant Diablo Key near Fajardo, Puerto Rico. A good, distinct form confined to a tiny island of but about ten acres, but what a horrid name it bears ! Ameiva exsul Cope St. Thomas, Water Island, St. John, St. James, Peter Island, Buck Island, Guana Island, Vieques, Anegada, Tortola, Anguilla, St. Croix and Puerto Rico. Now exterminated on St. Thomas. I have always doubted the St. Croix record. It is common where it still occurs at all. Ameiva vittipunctata Cope Hispaniola. A very beautiful and apparently not very common form. Ameiva taeniura Cope Hispaniola. When Dr. Noble and I prepared our Revision of Ameiva in 1915, I think I was principally to blame for concluding that this species was the young of A. lineolata. Miss Cochran has shown that this is untrue and that the species is perfectly valid. Ameiva chrysolaema chrysolaema Cope Hispaniola, La Gonave. A very common and widely spread species. A large series taken last year at Anse a Galets, La Gonave Island. 144 bulletin: museum of comparative zoology Ameiva chrysolaema abbotti Noble Beata Island. Common on this beautiful and usually uninhabited Island. Ameiva chrysolaema juliae Cochran Haiti, Isle Tortue. Ameiva barbouri Cochran La Gonave Island: La Source. Taken only by Eyerdam in 1927. I did not find it when on La Gonave in 1929 and November, 1934. Although I secured a great number of Ameivas, all were A. chrysolaema chrysolaema. Ameiva thoracica Cope Bahama Islands. Now known to be widespread in the northern and central portion of the Bahama archipelago. Ameiva dorsalis Gray Jamaica. Formerly abundant, then, after the mongoose came, pretty well reduced — almost exterminated. Now recovering slightly in numbers in the cities and settlements where the mongoose population is kept in hand. Ameiva auberi Cocteau Cuba and Isle of Pines. Nowhere abundant but very widely distributed. Perhaps most fre- quently seen along railway embankments. Ameiva rosamondae Cochran Saona Island. A most beautiful and very distinct species. The most brilliantly colored of the entire genus. It is distinctly a rare form. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 145 Ameiva beatensis Noble Beata Island. I found this species much less common than A. chrysolacma abbotti on recent visits to Beata. Ameiva navassae Schmidt Navassa. Known from the type only, taken by R. H. Beck in 1917. Not found by the last collectors in 1930. Scolecosaurus alleni alleni (Barbour) Grenada. A distinct and not uncommon species of the wet spice gardens. This little creature is most commonly found under heaps of half decayed cocoa pods. Scolecosaurus alleni parviceps Barbour Cannouan Island. Known from a single specimen taken by Dr. David Fairchild while on the Utowana. The genus probably occurs on all the Grenadines. Gymnophthalmus pleii Bocourt St. Lucia and Martinique. Extinct on Martinique. Excessively rare on St. Lucia. Whether G. ludkenii, also of Bocourt, from "St. Lucia" is really dis- tinct or whether it ever came from St. Lucia w^ill, in part, be solved finally only by examination of the type. Only pleei was found on these two islands by Garman, who took a good series before it was extermi- nated. Parker, who records the one specimen taken in 1932, remarks that its characters tend to confirm the supposition that there is only one West Indian species. Family AMPHISBAENIDAE Cadea palirostrata Dickerson Isle of Pines. A very distinct and abundant species. 146 bulletin: museum of comparative zoology Cadea blanoides Stejneger Cuba. Rare and confined to Matanzas, Havana and Pinardel Rio Provinces. Amphisbaena fenestrata Cope Tortola, St. Thomas, St. Croix and St. John. This form may be found to be still more Widely distributed. Amphisbaena bakeri Stejneger Puerto Rico. Rare and local. Amphisbaena caeca Cuvier Puerto Rico. Common. Amphisbaena manni Barbour Hispaniola. This form seems to be about equally abundant with innocens. Amphisbaena innocens Weinland Hispaniola. Not uncommon in Haiti. Amphisbaena cubana Peters Cuba. Common in Central Cuba. Best found by following plows. Amphisbaena caudalis Cochran Grande Cayemite Isl., Haiti. Known from but two examples taken by Eyerdam in 1927. It is allied to A. innocens. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 147 Family SCINCIDAE IVIabuya mabouia (Dumeril & Bibron) From Mexico and the Bahamas through the West Indies and on the mainland south to Trinidad and Patagonia. Absent from Cuba. Any number of races have been recognized and named from time to time, some confined to single islands and others to island groups, but with large series all of these forms break down. Incipient races there are beyond doubt but apparently the inherent fluidity or variability within the species has prevented these races from becoming fixed. My friend. Professor E. R. Dunn, has revised this situation in Proc. Acad. Nat. Sci. Phila., 1935, p. 533-557. He recognizes two races within the species but statistical studies based on vastly more material are needed before one can really settle the question of races within this plastic, wide-ranging and perhaps oft artificially introduced form. Skinks are zoological tramps. Skinks are apparently extinct on the following islands where once they were known to occur: St. John, St. Lucia, St. Vincent, Grenada, Barbados, Martinique. Mabuya lineolata Noble & Hassler San Domingo. A fine distinct species which has recently been found. It must be very rare to have eluded collectors for so long. The mongoose is abundant in San Domingo to be sure, but the early collectors all failed to find the skink. Suborder OPHIDIA Family TYPHLOPIDAE Typhlops tenuis Salvin Mexico, Guatemala and Andros Island. Rosen got what he called this species at Mastic Point in 1910. I have never felt very sure that it was not an undescribed form wrongly identified. 148 bulletin: museum of comparative zoology Typhlops rostellatus Stejneger Puerto Rico. Seems to be related to T. dominicana. Perhaps other species remain to be uncovered in the Lesser Antilles. Typhlops richardii Dumeril & Bibron St. Thomas, Tortola, St. John. Typhlops pusillus Barbour Hispaniola. Not uncommon in Haiti. Typhlops dominicana Stejneger Dominica and Guadeloupe. The specimens from Martinique should belong here, one would suppose, rather than to T. jamaicensis. More material is highly de- sirable from all of the islands. Typhlops platycephalus Dumeril & Bibron Puerto Rico, Vieques, Culebra, Caja de Muertos, Cayo Luis Peiia. Apparently fairly well differentiated though long confused with T. jamaicensis. Typhlops sulcatus Cope Navassa. May not really be a valid species. It has not been found by the recent collectors. Typhlops jamaicensis (Shaw) Jamaica. A common form. Typhlops monensis Schmidt Mona Island. Member of the T. lumbricalis series. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 149 Typhlops lumbricalis (Linne) Cuba, Hispaniola, Andros, New Providence and Abaco. Common everywhere and no doubt fortuitously introduced into the Bahamas. Typhlops granti Ruthven & Gaige Caja de Muertos, 18 miles off Ponce, Puerto Rico. Family LEPTOTYPHLOPIDAE Leptotyphlops albifrons (Wagler) Watlings Island, Antigua, Grenada and with a wide range in tropical America: This tiny burrowing snake has an erratic distribution and has prob- ably been carried about by primitive man, being occasionally intro- duced with material intended for garden planting. Leptotyphlops bilineata (Schlegel) Martinique, St. Lucia, Guadeloupe and Barbados. This, another tiny species, may have a considerably wider range among islands than we now know. Family BOIDAE Epicrates angulifer Bibron Cuba and Isle of Pines. Formerly common everywhere, now confined to the wilder regions, although individuals occasionally stray into the cultivated areas. The great extension of cane cultivation has decimated this species. Every cane cutter carries a machete all the time and uses it on every snake. Epicrates striatus striatus (Fischer) Hispaniola. This form seems to be really uncommon. 150 bulletin: museum of comparative zoology Epicrates striatus strigilatus (Cope) Andros and New Providence in the Bahamas. The fowl snake of the Bahamas was formerly abundant and may still be found but it is ruthlessly killed by the natives on account of its fondness for poultry. Stull believes these two forms to be separable. Epicrates striatus chrysogaster (Cope) Turks Island. Of this form I have no recent information, except that it is said to be rather common on some of the Turks Island Cays. Epicrates striatus relicquus (Barbour & Shreve) Sheep Cay off Gt. Inagua Island, Bahamas. This is no doubt the extirpated boa of Great Inagua, persisting on this islet to which no feral animals have been carried. Perhaps the trinominal best suggests the affinity of this distinct form. Epicrates inornatus inornatus (Reinhardt) Puerto Rico. Now a really rare species and one which is related to the large boas of Cuba, Jamaica, and Hispaniola. Epicrates inornatus granti Stull Tortola and Guana Island. Known from the single specimen taken by Major Chapman Grant on Tortola. He learned that it occurs in the rocky cliffs of Guana Island also. Epicrates fordii fordii (Giinther) Hispaniola. A very rare snake, which is noteworthy since generally speaking Antillean boids are abundant, except where artificially reduced in number. Possibly the mongoose is responsible for its rarity but it seems to have seldom been collected even before the introduction of the mongoose into Hispaniola. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 151 Epicrates fordii monensis Zenneck Mona. A very little-known species but one which I believe to be most closely allied to E. fordii. This combination of names is by Stull, the most recent reviser of the Boidae. Epicrates subflavus Stejneger Jamaica. I had supposed this species gone in Jamaica itself but Mr. Frank Cundall of the Institute of Jamaica at Kingston has one alive, from the southeast part of the Island. It persists on Goat Island off the south coast, in small numbers. Epicrates gracilis (Fischer) Hispaniola. I have never seen a specimen of this form in all the Haitian material which has passed through my hands. As described it has a very pe- culiar and unique color pattern but modern material would be very welcome. Boa cookii grenadensis (Barbour) Grenada. I may not have been justified in separating this form from B. cookii. I am, however, inclined to believe that it is fairly well differentiated and stabilized. Boa hortulana Linne St. Vincent, Grenada, The Grenadines and Trinidad, widespread on the main- land. The species still occurs on Grenada and may, being arboreal, persist on St. Vincent. This, however, I am inclined now to doubt. Constrictor constrictor orophias (Linne) St. Lucia, Dominica. The "tete chien" is rare on St. Lucia but still occurs — and even, occasionally at least, eats a mongoose. On Dominica it is less un- common. There is a Zoological Park (Phila.) record for St. Kitts 152 bulletin: museum of comparative zoology which I beheve to be incorrect; captive snakes get carried far and wide and dealers convey notoriously inaccurate locality records. There are also records from Trinidad but my friend, Mr. Urich, a most compe- tent resident authority, told me that the species does not occur in Trinidad. It is confined to two islands only. Tropidophis maculatus maculatus (Bibron) Western Cuba and Isle of Pines. Found sparingly in western Cuba and the Isle of Pines. I follow Miss StuU's conclusions in the taxonomy of this genus. Tropidophis maculatus pilsbryi Bailey Central and Eastern Cuba. Bailey has recently described this form on a number of specimens from the mountains of Santa Clara and Oriente provinces. Tropidophis maculatus jamaicensis Stull Jamaica. Excessively rare, almost extinct, since the introduction of the mon- goose. Tropidophis maculatus haetianus (Cope) Hispaniola, La Gonave and Isle Tortue. Not uncommon all over the Island. Tropidophis pardalis pardalis (Gundlach) Cuba and Isle of Pines. The Abaco records which have caused such worry were evidently due to the wrong copying of field data by me. (Cf. Bailey Proc. New Eng. Zool. Soc, Vol.' 16, 3 May, 1937, p. 46). Tropidophis pardalis canus (Cope) Great Inagua, Eleuthera Islands, Cat Island, and Long Island. Common on Eleuthera but now very rare on Inagua. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 153 Tropidophis pardalis curtus (Garman) New Providence, Bahamas. A common form. It occurs under stones of walls and in the rocks heaped about the orange trees. Since it at times sallies forth after heavy rains, it is locally called "thunder snake." Like all its congeners, it is nocturnal. Tropidophis pardalis barbouri Bailey Eleuthera, Cat and Long Island, Bahamas. My colleague Shreve and I considered describing this species but he did not consider it sufficiently distinct. It is, however, valid if not strikingly well defined. Tropidophis pardalis androsi StuU Andros Island. Apparently abundant but I have never happened to see a specimen. Tropidophis pardalis greenwayi Barbour & Shreve Ambergris Cay, Caicos Island. Probably widespread in this group of islands. Tropidophis bucculentus (Cope) Navassa. Known from but three specimens, it has not been found by recent expeditions. Bailey believes that this represents a distinct species and not a sub-species of pardalis as Stull concluded. Tropidophis wrighti Stull Cuba. Known, so far as I am aware, from the type only. This was taken by Charles Wright, the botanist, who collected for a long time in the Guantanamo Basin and, I think, nowhere else in Cuba. 154 bulletin: museum of comparative zoology Tropidophis nigriventris Bailey Camaguey Prov., Cuba. Just described and known as yet from but two specimens. Tropidophis melanurus (Schlegel) Cuba and Isle of Pines. The largest member of the genus, reaching a length of nearly a yard. It is abundant and widespread. It feeds on frogs, lizards and birds. Although more inclined to be arboreal than the other species of the genus, it is equally nocturnal and perhaps the most abundant of them all. Tropidophis semicinctus (Gundlach & Peters) Cuba. Widespread but distinctly uncommon. Family COLUBRIDAE Natrix compressicauda Kennicott Cuba, Florida Keys, extreme southwestern Florida. My finding this species in mangroves near Caibarien on the north coast of Cuba established the specific identity of the excessively rare Cuban Natrix and relegated several long questioned names to a definite synonymy. Tretanorhinus variabilis variabilis Dumeril & Bibron Cuba. Not uncommon in fresh-water ponds and rivers. A nocturnal species. Its mainland ally, T. nigroluteus, is rather partial to mangrove swamps. Tretanorhinus variabilis insulae-pinorum (Barbour) Isle of Pines. This species seems to have regularly 19 rows of scales while the Cuban snakes have 21. This is, at first sight, a trivial character but one which is apparently really diagnostic. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 155 Drymobius boddaerti bruesi (Barbour) St. Vincent and Grenada. Extinct on St. Vincent but still to he found on Young's Island off its coast and very rare in Grenada. Mr. Shreve believes that with more material from Young's Island another race might be named. My friend, Mrs. Gaige, advised me to resurrect my name bruesi for this race which I first applied with the idea that the Grenadian snake was an Alsophis. Uromacer oxyrhynchus Dumeril & Bibron Hispaniola and Isle Tortue. A form found all over the Island, i.e., both Haiti and San Domingo. I have seen it from Port au Prince and Samana. Uromacer frenatus (Giinther) Hispaniola and Isle Tortue. We now have a fine series of this species. Uromacer wetmorei Cochran Beata Island. A valid form related to the preceding. Uromacer catesbyi (Schlegel) Hispaniola and La Gonave. A widespread but rather rare species. Uromacer scandax Dunn Isle Tortue, near Haiti. An abundant ally of U. catesbyi. Uromacer dorsalis Dunn La Gonave Island. Apparently a derivative of the Haitian U. frenatus. 156 bulletin: museum of comparative zoology Alsophis anomalus (Peters) Hispaniola and Isle Tortue. I have but little information to give concerning this species. Dr. G. M. Allen took one at Port au Prince in 1919. I took one on Isle Tortue during the Utowana cruise of 1934, besides which I have received no other recent specimens. Alsophis leucomelas leucomelas (Dumeril & Bibron) Guadeloupe and Marie Galante. Extinct on both islands. Alsophis leucomelas sanctorum (Barbour) Les Saintes Is. near Guadeloupe. No doubt abundant still. Alsophis leucomelas sibonius (Cope) Dominica. With no mongoose on this island, the species should be abundant still. There are still great areas of wild land on Dominica. Alsophis leucomelas manselli Parker Montserrat. Still to be found. Alsophis leucomelas antiguae Parker Antigua. Extinct. St. Croix. Extinct. Alsophis melanichnus Cope Hispaniola. We await more information concerning this snake with great interest. Its non-appearance in any of the collections which have come before me is perhaps indicative that it is fast disappearing. Alsophis sanctae-crucis Cope BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 157 Alsophis ater (Gosse) Jamaica. Very rare indeed. A species which has suffered fearfully from the ravages of the mongoose. Dunn has shown that this is related to A. melatiicknus Cope of Haiti. Alsophis rijgersmaei Cope St. Martins, St. Barts and Anguilla. No herpetologist has visited St. Martins in recent years, but Dunn has re-examined the types and considers that Garman's name of Alsophis cinereiis cannot stand as valid. Alsophis variegatus (Schmidt) Mona Island. Probably still abundant. Alsophis portoricensis (Reinhardt & Liitken) Puerto Rico, Desecheo and Caja de Muertos Island. A distinctly rare form. Alsophis anegadae Barbour Anegada. I still feel that this form warrants recognition as valid. Its peculiar pattern is characteristic of every Anegada specimen which I have seen, even though it occurs very sporadically elsewhere, where other patterns are the place mode. Alsophis antillensis (Schlegel) Vieques, St. Thomas, St. James, Salt Island, Peter Island, St. John, Tortola, Virgin Gorda, Culebra, Pinero and Dog Island. Extinct on St. Thomas, rare on Puerto Rico, elsewhere abundant. Major Grant doubts the records for Puerto Rico. Alsophis rufiventris (Dumeril & Bibron) Saba, St. Kitts, St. Eustatius and Nevis. Still abundant on Saba and St. Eustatius but extinct on the other two islands. 158 bulletin: museum of comparative zoology Alsophis vudii vudii Cope Bahama Islands. This racer is common throughout most of the middle group of Bahama Islands : — New Providence, Eleuthera, Long Island, Green Cay, tlie Exuma Cays, Andros Ids. and no doubt upon many others. Alsophis vudii aterrimus Barbour & Shreve Grand Bahama. A black racer, not brown or grayish, perhaps confined to this little- known island. Alsophis vudii picticeps Conant Bimini Islands. Related to the two preceding races but well defined. Alsophis vudii raineyi Barbour & Shreve Crooked Isl., Bahamas. A well defined local form. Alsophis vudii utowanae Barbour & Shreve Sheep Cay off Great Inagua Isl., Bahamas. Another distinct relict on Sheep Cay which was no doubt common on Great Inagua before the introduction of so many domesticated animals which have become feral. Alsophis angulifer angulifer (Bibron) Cuba and Isle of Pmes. A very common species in all open plains, pastures and savannas. Alsophis angulifer fuscicauda (Garman) Cayman Brae, and Little Cayman. A well defined race. Mr. Roger Conant has recently shown me a specimen from Little Cayman, which he has recorded recently. (Proc. New Eng. Zool. Club, Vol. 16, Oct. 4, 1937, p. 81). BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 159 Alsophis angulifer caymanus (Garman) Grand Cayman. I have never seen sufficient material to decide whether this form is really different from that of Cuba. Dromicus andreae andreae (Reinhardt & Liitken) Cuba. A common snake at pastures and open fields. I follow Professor E. R. Dunn in suppressing the genus Leimadophis. Dromicus andreae nebulatus (Barbour) Isle of Pines. Another common form. It is closely related to the foregoing species, indeed closely similar specimens occur also in extreme eastern Cuba. We should probably recognize three races or abandon this name. Dromicus callilaemus Gosse Jamaica. Small and more retiring, this species is not so near extermination as L. ater. Nevertheless it is a distinctly rare snake. Dromicus funereus Cope . Jamaica. This form long buried in the synonymy has been shown by Major Grant to be valid. Dr. Stejneger agrees. Two out of the three original types has been found in Washington. Dromicus juliae juliae (Cope) Dominica. Probably still not uncommon. Dromicus juliae copeae Parker Guadeloupe. Extinct. 160 bulletin: museum of comparative zoology Dromicus melanotus (Shaw) Grenada, Trinidad and Venezuela. Extinct apparently on Grenada but common elsewhere. Dromicus perfuscus Cope Barbados. Extinct. Marie Galante. Extinct. St. Lucia. Extinct. Martinque. Extinct. Dromicus mariae (Barbour) Dromicus ornatus (Garman) Dromicus .cursor (Lacepede) Dromicus anegadae (Barbour) Anegada. We have no recent information concerning this form but no reason to suppose that it is not still abundant. Dromicus exiguus Cope St. Thomas, St. John, Tortola and Just van Dyke, and Hassel II. Extinct on St. John and St. Thomas, it is not uncommon on the other islands. Major Grant doubts the Culebra records. Dromicus stahli (Stejneger) Puerto Rico. Still not uncommon, widely distributed and confined to this Island. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 161 Dromicus alleni (Dunn) La Gonave Island. A distinct and striking island form. Dromicus parvifrons parvifrons (Cope) Hispaniola. One of several races which appear to be common, reasonably well localized in southwest Haiti and probably valid. Dromicus parvifrons niger (Dunn) Hispaniola. This form inhabits most of San Domingo. Dromicus parvifrons protenus (Jan) Hispaniola. A common widespread form. Known from many localities in north- ern and central Haiti and the higher plateau of San Domingo. Dromicus parvifrons lincolni (Cochran) Beata Island. A slightly differentiated form. Dromicus parvifrons tortuganus (Dunn) Isle Tortue. Another well marked form of which we took a good series during the visit of the Utoicana to this island in 1934. Dromicus parvifrons rosamondae Cochran Isle Vache. A fairly well defined form based on a good series of specimens. 162 bulletin: museum of comparative zoology Hypsirhynchus ferox Giinther Hispaniola. This species is strictly nocturnal and oviparous. In my experience, it is restricted apparently to the Cul de Sac area not far from Port au Prince. Dunn has discarded the genus Hypsirhynchus. I believe that this sluggish, nocturnal form is well worthy of generic distinction. Arrhyton taeniatum Giinther Cuba. An uncommon species, like its fellow, found by day under stones or while plowing. At night it is sometimes met with abroad. Arrhyton redimitum (Cope) Eastern Cuba. Material recently received has proved the validity of this form, so Mr. Benjamin Shreve assures me. Arrhyton vittatum (Gundlach & Peters) Cuba. These snakes are probably allies of Contia of the mainland. Darlingtonia haetiana Cochran Haiti. An extraordinary new genus recently found by Dr. Darlington of Harvard at Roche Croix, in the northeastern foothills of Morne La Hotte, at 5,000 ft. altitude. Its affinity may be with the preceding genus but it is very well defined. PsEUDOBOA CLOELIA (Daudin) Dominica, St. Lucia, Grenada, Trinidad and tropical America generally. This species is surely extinct in St. Lucia, probably excessively rare on Grenada and its status on Dominica is still, no doubt, unchanged. I have never, however, seen or heard of recent specimens from any of the islands. Nevertheless, I think the records are really based on valid wild-caught specimens. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 163 PsEUDOBOA NEUWEIDII (DuiTieril & Bibron) Grenada, Trinidad and with a wide range in tropical America. Garman took three examples on Grenada during the Blake Expedi- tion about 1883. So far as I can learn it has never been taken before or since. Ialtris dorsalis (Giinther) Hispaniola, Isle Vache. A large and uncommon species which has been found in both Haiti and San Domingo. It seems to have no close allies among Antillean reptiles and to be very rarely collected indeed. Ialtris parishi Cochran Haiti. Known only from southwestern Haiti. Family CROTALIDAE BoTHROPS ATROX (Linue) Martinique and St. Lucia. Whatever may be the origin of the Fer-de-lance's appearance on these islands, one thing Amaral has definitely proved — the snake is the common wide-ranging form of tropical America. Order CHELONIA Family TESTUDINIDAE Testudo tabulata Walbaum Tropical South America, feral on Lovango Cay and Water Island, near St. Thomas. Carried, from time to time, to most of the islands from South America. Not a native element of the Antillean fauna. 164 bulletin: museum of comparative zoology Family EMYDIDAE Pseudemys felis Barbour Cat Island. I wonder if the significance of the finding of a Pseudemys in tlie Bahamas has been fully appreciated. It seems to me that in this connection the following facts are worthy of note. There are innumerable mangrove swamps throughout the archi- pelago. These would be suitable homes for Malaclemys but none are to be found. Malaclemys, confined to salt water marshes and mangrove swamps, are, or were but a few years ago, abundant from Cape Cod to the Florida keys. Pseudemys, so far as I know, never goes into salt or brackish water and yet, of the innumerable islands of the Bahamas, there's only one which has a few poor little fresh water ponds, or per- haps better mud holes, in which it supports a very considerable popu- lation of Pseudemys whose habits have become highly specialized to meet the peculiar conditions under which they live. What the mathematical probability would be which would bring these ponds and Pseudemys together by, let us say, the carrying of young turtles by a hurricane is, of course, utterly incalculable. It seems to me that some more plausible reason must be found for their being there. Perhaps not long ago when the Bahamas were higher there were larger lakes, more of them and more Pseudemys. What we see now, unsuspected until a few years ago, no doubt represents an- other disappearing remnant of fauna and of these there are many in the Antilles. It is still a little difficult to figure how Pseudemys got into these ponds even if they were once much larger, for fresh water turtles seem to be particularly unfitted for chance dispersal. It is hard to believe that they swam over, if no Malaclemys ever has. That looks as if such turtles did little sea swimming and that they were picked up by winds of hurricane force and dumped in a place where they could survive seems to presuppose a toughness of fiber on the part of the turtle which is contrary to our knowledge of the beasts and again the mathe- matical chances against picking up, carrying and then not landing in an unfavorable spot would, I should suppose, be millions to one. The Cat Island turtle is, I believe, rather better differentiated than are the turtles of the several different Greater Antilles on which they occur. Cuba. BARBOUR: ANTILLEAN REPTILES AND AMPHIBIANS 165 PsEUDEMYS DECussATA Gray What, I believe, to be the undoubted type, in the British Museum, has been photographed by Mr. Parker and I feel reasonably sure that this represents a Cuban form. The type localities for this and the following form are unknown. PsEUDEMYS RUGOSA Shaw Cuba. Thanks to Dr. Stejneger and Mr. Parker I have photographs of the type in the Museum of the Royal College of Surgeons and suspect this also to be a Cuban species. PsEUDEMYS STEJNEGERi Schmidt Puerto Rico. A small, possibly distinct form. Major Grant doubts the validity of this species. I am inclined to agree with him but let it stand, pending a general revision of the turtles of all the islands which I hope to undertake, or help someone else do, when I have much more material. PsEUDEMYS SSp. Cuba, Haiti, Jamaica. There are other pond turtles on these islands but their systematic status is as yet in doubt. Order LORICATA Family CROCODYLIDAE Crocodylus rhombifer Cuvier Cuba and Isle of Pines. Found in the Zapata Swamp in Cuba and no doubt still also in the Cienaga of the Isle of Pines. Specimens more than six feet long are now much less often seen than a generation ago. 166 bulletin: museum of comparative zoology Crocodylus acutus Cuvier Cuba, Jamaica, and Hispaniola; as well as extreme southern Florida and the Keys and Central America. Crocodylus intermedius Graves Orinoco Basin. Accidental in Grenada, September 6, 1910. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. LXXXII, No. 3 LLST OF THE FISHES, TYPES OF POEY, IN THE MUSEUM OF COMPARATIVE ZOOLOGY By Luis Howell y Rivero Museo Poey University of Havana CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE MUSEUM July, 1938 oi Comp, ^h'^' Zoology ""'-^ JUL 14 1938 M C^ No. 3. — List of the Fishes, Types of Poey, in the Museum of Comparative Zoology By Luis Howell y Rivero INTRODUCTION The present paper presents a list of some of the types of fishes de- scribed by FeHpe Poey, viz. those which are deposited in the collection of the Museum of Comparative Zoology. Felipe Poey (1799-1891), who during his entire life studied Cuban fishes, sent to the Museum of Comparative Zoology, between the years 1861 and 1870, several collections of alcoholic fishes, as well as a great number of skeletons and mounted specimens. Among these were a large number of his t^^pes, and it is- the purpose of this paper to discuss such of these as can at present be located. A careful checking has been made of ever}^ specimen sent by Poey to determine which are t.vpes. I have been helped in every instance by his various publications, the original invoices which accompanied the lots, now deposited in the Library of the Museum of Comparative Zoology, besides many notes brought with me from Cuba. Attention is called to the fact that more than once the invoice number of the specimens sent by Poey has been taken as the original "species num- ber," and this has caused confusion. Every species described or identi- fied by him pertaining to the Cuban fauna as far as he knew it, bears a "species number" in all his papers, regardless of the number of speci- mens used or seen. The writer has a complete list of Poey's "species numbers" and these are given with each species herein as No. Poey after the M. C. Z. catalogue number. In this paper, the name by which the species was described is given first, followed by the name which gives the actual status of the species. In the synonymy, reference is given to all the publications of Poey in which the species is mentioned. After this, reference is given to the original description of the species in those cases in which Poey's species is considered a sjiionym. Last comes the reference to the work of Jordan & Evermann (Fishes of North and Middle America, 1896- 1898), as the most complete account of the fishes of the West Indies. In all possible cases reference will be given to the latest work concern- ing any major group or family. To save space by avoiding a certain amount of repetition, all refer- ences have been abbreviated, giving a complete and detailed bibli- 170 bulletin: museum of comparative zoology ography at the end of the paper, where attention is called to the publi- cation dates of the fascicules of the Memorias and Repertorio of Poey. Lack of accuracy in this matter has been the cause of more than one error. All of the holotypes and many of the paratypes and cotypes are figured in Poey's unpublished manuscript "Ichthyology of Cuba," but some of these figures have already appeared in his various publications. Since it is understood that the general locality is Cuba, the locality reference is omitted, except in cases where a specific locality is stated in the description, e.g. Jardin Botanico, Habana. (for Girardinus metal- licus). It is a satisfaction to find that of the 389 species described by Poey, the types of 188 species are here preserved. I regret to say that a few of his types are lost, although in some instances he states in his in- voices or original description that he has sent them to the Museum of Comparative Zoology; in other cases they are cited as being in this Museum in the work of Jordan & Evermann (loc. cit.), among these Chauliodus richardsoni Poey, Gramma loreto Poey and Physiculus kaupi Poey. Most of the specimens sent by Poey, especially the types, are in a very good state of preservation. In this paper, the general an-angement of the "Check List of Fishes and Fish-like Vertebrates of North and Middle America," 1930 (Bull. U. S. Fish Comm.) has been followed. All measurements given refer to the total length of the specimens, unless something else is specifically indicated. SQUALIDAE Etmopterus Etmoptcrus Rafinesque, 1810a:14 {Etmopterus aculeatus Rahnesque = Squalus spinax Linnaeus) 1, Spinax hillianus Poey = Etmopterus hillianus (Poey) S2mia.r MlUamis Poey, 1861 :340, 359, pi. 19, fig. 13-14. Squalus hiUiamis, Poey, 1868:454. Spinax spinax Poey, 1876-203. Etmopterus pusillus, Jordan & Evermann, 1896:55. Etmopterus hillianus, Garman, 1913:224, pi., 10, fig. 1-4. Holotype: M. C. Z. 1025, female. No. Poey 617. HOWELL Y RIVERO : TYPES OF POEY FISHES 171 CLUPEIDAE Sardinia Sardinia Poey, 1860:311 {Sardinia pseudo-hispanica Poey) 2. Sardinia pseudo-hispanica Poey = Sardinia anchovia (Cuvier & Valenciennes) Sardinia pseudo-hispanica Poey, 1860:311; 1861:384; 1868:419; 1876:148. SardincUa anchoina Cuvier & Valenciennes, 1847:269; Jordan & Evermann, 1896:429. Cotypes: M. C. Z. 17768 and 17771. No. Poey 34. In the first lot there are five specimens, 100 to 145 ram., the one 130 mm. long figured by Poey in his MS; the second number includes only one specimen. Harengula. Harengxda Cuvier & Valenciennes, 1847:280 {Harengula latutus Cuvier & Valenciennes) 3. Harengula sardina Poey Harengida sardina Poey, 1860:310; 1861:384; 1868: 418; 1876:147. SardineUa sardina, Jordan & Evermann, 1896:430. Holotype: M. C. Z. 17868. Paratypes: M. C. Z. 17736, two speci- mens 128 and 153 mm. No. Poey 40. Chirocentrodon Chirocentrodon Giinther, 1868:463 {Chirocentrodon taeniatus Gunther) 4. Pellona bleekeriana Poey = Chirocentrodon bleekeriana (Poey) Pellona bleekeriana Poey, 1867:242; 1868:419; 1876: 148. Ilisha bleekeriana, Jordan & Evermann, 1896:436. Cotypes: M. C. Z. 17845, seven specimens in such bad condition that no measurements could be made. No. Poey 537. ENGRAULIDAE Anchoviella Anchoviella Fowler, 1911:211 {Engraulis perfasciatus Poey) 172 bulletin: museum of comparative zoology 5. Engraulis perfasciatus Poey = Anchoviella perfasciata (Poey) Engraulis ijerfasciatus Poey, 1860:312; 1861:385; 1868:422, 460; 1876:149. Stolephorus perfasciatus, Jordan & Evermann, 1896: 441. Types: M. C. Z. 17955, nine specimens, 82 to 103 mm., among which the one 100 mm. long is the holot^-pe. No. Poey 422. 6. Engraulis cubanus Poey = Anchoviella cubana (Poey) Engraulis cubanus Poey, 1868:420, 460; 1876:149. Stolephorus cubanus, Jordan & Evermann, 1896:442. Types: M. C. Z. 17958, four specimens, the largest of which is the holotjy-pe. No. Poey 23. Anchovia Anchovia Jordan & Evermann, 1896:449 (Engraulis microlepidotus Kner & Steindachner) 7. Engraulis productus Poey = Anchovia clupeoides (Swainson) Engraulis productus Poey, 1866a :380; 1868:422; 1876:149. Engraulis clupeoides Swainson, 1839 :388. Stolephorus clupeoides, Jordan & Evermann, 1896:447. Stolephorus productus, Jordan, & Evermann, 1896:447. Types: M. C. Z. 17961, two specimens, the largest of which is the holotype. No. Poey 36. Cetengraulis Cetengraulis Giinther, 1868:383 {Engraulis edentulus Cuvier) 8. Engraulis brevis Poey = Cetengraulis edentulus (Cuvier) Engraulis brevis Poey, 1866a :379; 1868:422. Cetengraulis brevis, Poey, 1876:148. Engraulis edentulus Cuvier, 1829:323. Cetengraidis edentulus, Jordan & Evermann, 1896:450. Holotype: M. C. Z. 24296. No. Poey 716. HOWELL Y RIVERO: TYPES OF POEY FISHES 173 CONGRIDAE Conger Conger Cuvier, 1817:231 {Muraena conger Linnaeus) 9. Conger esculentus Poey = Conger conger Linnaeus. Conger esculentm Poey, 1861:346, 385; 1867:246; • - 1868:424; 1876:151. Muraena conger Linnaeus, 1758:245. Leptocephalus conger, Jordan & Evermann, 1896:354. Holotype: M. C. Z. 9328. No. Poey 581. 10. Echelus caudilimbatus Poey = Conger caudilimbatus (Poey) Echelus caudilimbatus Poey, 1867:249, pi. 2, fig. 8. Ophiosoma caudilimbatus, Poey, 1868:424. Conger caudilimbatus, Poey, 1876:152. Leptoconger caudilimbatus, Jordan & Evermann, 1896:355. Holotype: M. C. Z. 9324. No. Poey 176. ECHELIDAE Myrophis Myrophis Liitken, 1851:1 (Myrophis punctatus Liitken) 11. Myrophis microstigmius Poey Myrophis microstigmius Poey, 1867:250, pi. 3, fig. 4; 1868:425; 1876:153. Myrophis imnctatus, Jordan & Evermann, 1896:371. Holotype: M. C. Z. 33440. No. Poey 653. This species has long been considered a synonym of Myrophis punctatus Liitken, but it proves to be distinct (Rivero, 1934:341). OPHICHTHYIDAE Myrichthys Myrichihys Girard, 1859:58 {Myrichthys tigrinus Girard) 12. Ophisurus latemaculatus Poey = Myrichthys oculatus (Kaup) 174 bulletin: museum of comparative zoology Ophisurus latemaculatus Poey, 1867:252, pi. 3, fig. 1; 1868:425. Pisodonophis latimaculatus, Poey, 1876:153. Ophichthys latimaculatus, Poey, 1880:10. Pisoodonophis oculatus Kaup, 1856:22. Myrichthys oculatus, Jordan & Evermann, 1896:376. Holotype: M. C. Z. 27223. No. Poey 606. 13. Ophisurus longus Poey = Myrichthys acuminatus (Gronow) Ophisurus longus Poey, 1867:254; 1868:425. Pisodonophis longus, Poey, 1876:154. Ophichthys longus, Poey, 1880:11. Muracna acuminata Gronow, 1854:21. Myrichthys acuminatus, Jordan & Evermann, 1896: 377. Holotype: M. C. Z. 9155. No. Poey 180. Ophichthus Ophichthus Ahl, 1787:5 (Muraena ophis Linnaeus) 14. Ophisurus chrysops Poey = Ophichthus gomesii (Castelnau) Ophisurus chrysops Poey, 1861:321, 385. Ojihichthys chrysops, Poey, 1868:425; 1876:154. Ophisurus gomesii Castelnau, 1855:84, pi. 44, fig. 2. Ophichthus gomesii, Jordan & Evermann, 1896:384. Holotype: M. C. Z. 27212. No. Poey 604. Mystriophis Mystriophis Kaup, 1856:10 {Ophisurus rostellatus Richardson) 15. Conger MORDAX Poey = Mystriophis intertinctus (Richardson) Conger mordax Poey, 1860:319; 1861:385. Macrodonophis mordax, Poey, 1867:252; 1868:425. Crotalophis mordax, Poey, 1876:153. Ophisurus intertinctus Richardson, 1844a :102. Mystriophis intertinctus, Jordan & Evermann, 1896: 386. Holotype: M. C. Z. 9220. No. Poey 339. HOWELL Y RIVERO: TYPES OF POEY FISHES 175 MURAENIDAE Gymnothorax Gymnoihorax Bloch, 1795:83 {Gijvmothorax muraena Bloch) 16. Gymnothorax obscuratus Poey Gymnothorax obscuratusYoey , 1870b :320; 1876:159. Lycodontis obscuratus, Jordan & Evermann, 1896:398. Holotype: M. C. Z. 27217. No. Poey 736. Type locality: Havana. Channomuraena Channomuraena Richardson, 1844a:96 (Ichthyophis viitatus Richardson) 17. Channomuraena cubensis Poey = Channomuraena vittata (Richardson) Channomuraena cubensis Poey, 1867:266, pi. 3, fig. 6; 1868:428. Ichthyophis vittatus Richardson, 1844b :1 14, pi. 53, figs. 7-9. Channomuraena vittata, Poey, 1876:160; Jordan & Evermann, 1896:404. Holotype: M. C. Z. 9191. No. Poey 230. SYNODONTIDAE Trachinocephalus Trachinocephalus Gill, 1861:53 (Saurus myops Cuvier & Valenciennes) 18. Saurus brevirostris Poey = Trachinocephalus myops (Forster) Saurus brevirostris Poey, 1860:305; 1861:384. Trachinocephalus brevirostris, Poey, 1868:415; 1876: 144. Salmo myops Forster, in Bloch & Schneider, 1801 :421. Trachinocephalus myops, Poey, 1868:415; Jordan & Evermann, 1896:533. • Holot^-pe: M. C. Z. 6895. No. Poey 488. Synodus Synodus Gronow, 1777:449 {Synodus snyodus Linnaeus) 176 bulletin: museum of comparative zoology 19. Saurus spixianus Poey = Synodus foetens (Linnaeus) Saurus spixianus Poey, 1860:304; 1861:384. Synodus spixianus, Poey, 1868:413; 1876:144. Salmo foetens Linnaeus, 1766:513. Synodus foetens, Jordan & Evermann, 1896:538. Holotype: M. C. Z. 6884. No. Poey 588. SUDIDAE SUDIS Sudis Rafinesque, 1810a :60 (Sudis hyalina Rafinesque) 20. Paralepis intermedius Poey = Sudis intermedius (Poey) Parelepis intermedius Poey, 1868:416; 1876:142. Sudis intermedius, Jordan & Evermann, 1896:600. Holotype: M. C. Z. 32931. No. Poey 710. Type locality: Matanzas. MYCTOPHIDAE DiAPHUS Diaphus Eigenmann & Eigenmann, 1890:3 (Scopelus engraulis Giinther) 21. Myctophum nocturnum Poey = Diaphus dumerilli (Bleeker) Mydophum nocturnum Poey, 1861:426; 1868:416; 1876:145. Colletia nocturna, Jordan & Evermann, 1896:567. Scopelus dumerilli Bleeker, 1856:66. Diaphus dumerilli, Parr, 1928:126, fig. 23. Holotype: M. C. Z. 6871. No. Poey 297. CYPRINODONTIDAE RiVULUS Rivulus Poey, 1860:307 (Rivulus cylindraceus Poey) 22. Rivulus cylindraceus Poey Rivulus cylindraceus Poey, 1860:308; 1861:383; 1868: 412; 1876:140, pi. 8, fig. 4; 1880:5, pi. 5, fig. 4; Jordan & Evermann, 1896:662; Myers, 1927:121. HOWELL Y RIVERO : TYPES OF POEY FISHES 177 Types: M. C. Z. 6423, the female being the holotvpe, the male para- type. Paratj-pes: M. C. Z. 6395. No. Poey 366. T^-pe locality: Arroyo Mordazo, near Habana. Cyprinodon Cyprinodon Lacepede, 18031) :486 (Cyprinodon variegatus Lacepede) 23. Trifarcius felicl\nus Poey = Cyprinodon variegatus riverendi (Poey) Trifarcius felicianus Poey, 1868:412; 1876:140. Trifarcius Riverendi Poey, 1856:306. Cyprinodon felicianus, Jordan & Evermann, 1896:676. Cotypes: M. C. Z. 6402 and 6410; in the first lot one specimen 49.4 mm. long; in the second 3 specimens 37-39.6 mm. No. Poey 719. POECILIDAE Gambusia Gamhusia Poey, 1854:382 (Gambusia punctata Poey) 24. Gambusia punctata Poey Gambusia -punctata Poey, 1854:384, pi. 31, fig. 18, pi. 32, fig. 5-9; 1861:383; 1868:410; 1876:140. Jordan & Evermann, 1896:676; Hubbs, 1926:35. Cotypes: M. C. Z. 6393-6394. All the specimens in the same bottle, 22 males and 39 females. No. Poey 505. Type locality : Rio Alraendares, near Habana. 25. Gambusia puncticulata Poey Gamhusia puncticulata Poey, 1854:386, pi. 31, fig. 6-7; 1861:383; 1868:410; 1876:141; Jordan & Evermann, 1896:680; Hubbs, 1926:37. CotjTJes: M. C. Z. 6391, 6401, and 6397, 12 males and 24 females. No. Poey 510. T^-pe locality: Moats of the old city wall of Havana City. Glaridichthys Glaridichthys Garman, 1896:232 (Girardinus uninotatus Poey) 26. Girardinus uninotatus Poey = Glaridichthys uninotatus (Poey) 178 bulletin: museum of comparative zoology Girardinus uninotatus Poey, 1860:309; 1861:383; 186^11; 1876:142. Heterandria u7iinotata, Jordan & Evermann, 1896:687. Glaridichthys uninotatus, Hubbs, 1926:59. Cotypes: M. C. Z. 6406, four males and 21 females, M. C. Z. 6243, 2 males and 4 females, M. C. Z. 6405, 3 females. No. Poey 522. Type locality: Rio Taco Taco, Pinar del Rio. Girardinus Girardinus Poey, 1854:383 {Girardinus metallicus Poey) 27. Girardinus metallicus Poey Girardinus metallicus Poey, 1854:387, pi. 31, fig. 8-11; 1861:383; 1868:411; 1876:141; Hubbs, 1926:60. Heterandria metallica, Jordan & Evermann, 1896:687. Cotypes: M. C. Z. 6407, 1 male and 10 females, M. C. Z. 6414, 7 females, and 1412a, 1 male and 13 females. No. Poey 506. Type locality: Brook in the Botanical Garden, Habana. LiMIA Limia Poey, 1854:388 (Limia cubensis Poey = Poecilia vittata Guichenot) 28. Limia cubensis Poey = Limia vittata (Guichenot) Limia cubensis Poey, 1854:388, pi. 31, fig. 12-13, pi. 32, fig. 10-11; 1861:383; 1868:411; 1876:141; 1880:5. Poecilia vittata Guichenot, in Ramon de la Sagra, 1853:146, pi. 5, fig. 1; Jordan & Evermann, 1896:692. Limia vittata, Hubbs, 1926:75. Cotypes: M. C. Z. 6404, 1 male and 4 females, M. C. Z. 6403, 18 males and 40 females. No. Poey 347. Type locality: Moats of the old city wall of Havana. 29. Limia pavonina Poey = Limia vittata (Guichenot) Limia pavonina Poey, 1876:142. Poecilia pavonina, Jordan & Evermann, 1896:692. Poecilia vittata Guichenot, in Ramon de la Sagra, 1853:146, pi. 5, fig. 1; Jordan & Evermann, 1896:692. Cotypes: M. C. Z. 6400, 2 males and 1 female. No. Poey 549. HOWELL Y RIVERO: TYPES OF POEY FISHES 179 BELONIDAE Strongylura Strongylura Van Hasselt, 1824:374 {Strongylura caudimaculata Van Hasselt = Belone strongylura Van Hasselt) 30. Belone notata Poey = Strongylura notata (Poey) Belone notata Poey, 1860:293; 1861:376; 1867:166; 1868:382; 1876:120. Tylosurus notatus, Jordan & Evermann, 1896:710. Holotype: M. C. Z. 32933. No. Poey 413. Tylosurus Tylosurus Cocco, 1829:18 {Tylosurus contraini Cocco =Sphyraena acus Lacepede) 31. Belone melanochira Poey = Tylosurus raphidoma (Ranzani) Belone melanochira Poey, 1860:294; 1861:376; 1868: 382; 1876:120. Belone raphidoma Ranzani, 1842:359, pi. 37, fig. 1. Tylosurus raphidoma, Jordan & Evermann, 1896:715. Cotypes: M. C. Z. 624, 2 specimens 490 & 530 mm. No. Poey 541. 32. Belone crassa Poey = Tylosurus raphidoma (Ranzani) Belone crassa Poey, 1860:291; 1861:376; 1867:165; 1868:381; 1876:120. Belone raphidoma Ranzani, 1842:359, pi. 37, fig. 1. Tylosurus raphidoma, Jordan & Evermann, 1896:715. Paratype: M. C. Z. 623; the largest of the three specimens men- tioned in the original description, the median one, the holotype, missing. No. Poey 435. 33. Belone latimana Poey = Tylosurus acus (Lacepede) Belone latimana Poey, 1860:292; 1861:376; 1867:166; 1868:382; 1876:120. Sphyraena acus Lacepede, 1803b :6, pi. 1, fig. 3. Tylosurus acus, Jordan & Evermann, 1896:717. Holotype: M. C. Z. 622. No. Poey 353. 180 bulletin: museum of comparative zoology HEMIRAMPHIDAE Euleptorhamphus Euleptorhamphus Gill, 1859b: 156 (Euleptorhamphus brevoorti Gill) 34. Euleptorhamphus velox Poey Euleptorhamphus vclox Poey, 1868:383; 1876:121; Jordan & Evermann, 1896:724. Holotype: M. C. Z. 8779. No. Poey 722. Hyporhamphus Hyporhamphus Gill, 1859a :131 {Hyporhamphus tricuspidatus Gill = Hemir- hamphus unifasdatus Ranzani) 35. Hemirhamphus fasciatus Poey = Hyporhamphus unifasciatus (Ranzani) Hemirhamphus fasciatus Poey, 1860:299; 1861:377; 1867:167. Hemirhamphus pocyi, Poey, 1868:383; 1876:121. Hemirhamphus unifasciatus Ranzani, 1842:326. Hyporhamphus unifasciatus, Jordan & Evermann, 1896:720. Cotypes: M. C. Z. 32934, 2 specimens. No. Poey 194. Type locality : Habana Bay. Hemiramphus Hemiramphus Cuvier, 1817:186 {Esox brasiliensis Linnaeus) 36. Hemirhamphus filamentosus Poey = Hemiramphus brasiliensis (Linnaeus) HemirhamphMs filamentosus Poey, 1860:297; 1861 :377. 1868:382; 1876:121. Esox brasiliensis Linnaeus, 1758:314. Hemiramphus brasiliensis, Jordan & Evermann, 1896:722. Types: M. C. Z. 8775, the largest being the holotype, the other two paratypes, 290 & 320 mm. No. Poey 50. BOTHIDAE BOTHUS Botkus Rafinesque, 1810a:23 {Bothus rumulo Rafinesque) HOWELL Y RIVERO: TYPES OF POEY FISHES 181 37. Rhomboidichthys spinosus Poey = BoTHUS OCELLATUS (Agassiz) Rhomboidichthys spinosus Foey, 1868:409; 1870:139. Platophrys spinosus, Jordan & Evermann, 1898:2662. Rhombus occllatus Agassiz, 1831:85, pi. 46. Bothus ocellatus, Norman, 1934:222. Holotype: M. C. Z. 11345. No. Poey 669. Syacium Syacium Ranzani, 1840:20 (Syacium micrurum Ranzani) 38. HipPOGLOssus OCELLATUS Poey = Syacium micrurum Ranzani Hippoglossus occllatus Poey, 1861 :314, 385. Hemirhombus occllatus, Poey, 1868:407; 1876:138. Syacium micrurum Ranzani, 1840:20, pi. 5; Jordan & Evermann, 1898:2672; Norman, 1934:132. Holotype: M. C. Z. 11188. Paratypes: M. C. Z. 11144, 11192, 11194. No. Poey 515. CiTHARICHTHYS Cithanchthys Bleeker, 1862a :427 (Citharichthys cayennensis 'Bleeker = Citha- richthys spilopterus Giinther) 39. Hemirhombus fuscus Poey = CiTHARICHTHYS SPILOPTERUS Giinther Hemirhombus fuscus Toey, 1868:406; 1876:138. Citharichthys spilopterus Giinther, 1862:421; Jordan & Evermann, 1898:2685; Norman, 1934:149. Cotypes: M. C. Z. 11251. No. Poey 227. POLYMIXUDAE Polymixia Polymixia Lowe, 1838:198 (Polymixia nobilis Lowe) 40. Dinemus venustus Poey = Polymixia lowei Giinther. Dinemus venustus Poey, 1860:161, pi. 14, fig. 1; 1861 :366. Polymixia lowei Giinther, 1859:17; Poey, 1868:297; 1875:35; Jordan & Evermann, 1896:854. Cotypes: M. C. Z. 21812, 2 specimens 165 & 250 mm. No. Poey 160. 182 bulletin: museum of comparative zoology HOLOCENTRIDAE OSTICHTHYS Ostichthys Jordan & Evermann, 1896:846 (Myripristis japonicus Cuvier & Valenciennes) 41. Myriopristis fulgens Poey = Ostichthys trachypomus (Giinther) Myriopristis fulgens Voey, 1860:160; 1861:366. Myriopristis tr achy pom a Giinther, 1859:25; Poey, 1868:301; 1875:38; Jordan & Evermann, 1896:- 846. Cotypes: M. C. Z. 21910, 2 specimens, 180 & 190 mm. total length, No. Poey 296. Myripristis Myripristis Cuvier, 1829:150 {Myripristis jacobus Cuvier & Valenciennes) 42. Myriopristis lychnus Poey = Myripristis jacobus Cuvier & Valenciennes Myriopristis lychnus Poey, 1860:159; 1861:366; 1868:301; 1875b :38. Myripristis jacobus Cuvier & Valenciennes, 1829a :121 ; Poey, 1865:274; Jordan & Evermann, 1896:846. Types: M. C. Z. 10982, 8 specimens, the holotype 206 mm., the paratype 150 mm. long. No. Poey 204. Holocentrus Holocentrus Bloch, 1790:61 {Holocentrus sogo Bloch) 43. Holocentrus matejuelo Poey = Holocentrus ascensionis (Osbeck) Holocentrus matejuelo Poey (not of Bloch and Schnei- der), 1860:155, pi. 13, fig. 13-14; 1861:366; 1868:298; 1875b :35. Perca ascensionis Osbeck, 1765:388. Holocentrum longipinne, Poey, 1865:274. Holocentrus ascensionis, Jordan & Evermann, 1896: 848. Cotypes: M. C. Z. 21915-18; specimens 220 to 260 mm. No. Poey 26. 44. Holocentrum vexillarium Poey = Holocentrus vexillarius (Poey) HOWELL Y RIVERO: TYPES OF POEY FISHES 183 Holocentrum vexUlarium Poey, 1860:158; 1861:366; 1868:299; 18751) :37. Holocentrus vexillarius, Jordan & Evermann, 1896:852. Holotype: M. C. Z. 10935. No. Poey 303. 45. Holocentrum riparium Poey = Holocentrus VEXILLARIUS (Poey) Holocentrum riparium Voey, 1875b :37. Holocentrus vexillarius, Jordan & Evermann, 1896:852. Types: M. C. Z. 10934, the holotype being the largest of the four specimens. Paratypes: M. C. Z. 10936, 15 specimens, 43.5 to 60 mm. No. Poey 542. 46. Holocentrum perlatum Poey = Holocentrus osculus (Poey) Holocentrum perlatum Poey, 1860:157; 1861:366; 1868:298; 1875b :36. Holocentrum osculum, Poey, 1860:156. Holocentrus osculus, Jordan & Evermann, 1896:853. Holotype: M. C. Z. 10938. No. Poey 535. Flammeo Flammeo Jordan & Evermann, 1898:2871 {Holocentrum marianus Cuvier & Valenciennes) 47. Holocentrum rostratum Poey = Flammeo marianus (Cuvier & Valenciennes) Holocentrum rostratum Poey, 1860:157; 1861:366; 1868:298; 1875b :38. Holocentrum marianus Cuvier & Valenciennes, 1829a: 164. Holocentrus marianus, Jordan & Evermann, 1896:852. Flammeo marianus, Jordan & Evermann, 1898:2871. Cotypes: M. C. Z. 10969, 2 specimens 143 & 150 mm. total length. No. Poey 446. Plectrypops Plectrypops GiU, 1862b :237 (Holocentrum retrospinis Guichenot) 48. Holocentrum prospinosum Poey = Plectrypops retrospinis (Guichenot) Holocentrum prospinosum Poey, 1861 :343. My riopristis prospinosum , Poey, 1861:366. 184 bulletin: museum of comparative zoology Plecirypops prospinosus, Poey, 1868:302; 1875b :38. Holocentrum rctrospinis Guichenot, 1853:35; pi. 1, fig. 3. Plecirypops rctrospinis, Jordan & Evermann, 1896: 853. Lectotype: M. C. Z. 21884. No. Poey 534. Species first described based on Guichenot's figure. SYNGNATHIDAE Syngnathus Syngnathus Linnaeus, 1758:336 (Syngnathus acus Linnaeus) 49. Syngnathus brachycephalus Poey Syngnathus brachycephalus Poey, 1868:444; 1876:375. Siphostoma brachyccphalum, Jordan & Evermann, 1896:769. Types: M. C. Z. 11726; the male Holotype, the female Paratype. No. Poey 685. ATHERINIDAE Hepsetia Hepsetia Bonaparte, 1837 (no pages) (Atherina boyeri Risso) 50. Atherina laticeps Poey = Hepsetia stipes (Miiller & Troschel) Atherina laticeps Foey, 1860:265; 1861:380; 1868:390; 1876:100; Jordan & Evermann, 1896:790. Atherina stipes Miiller & Troschel, 1847:671; Jordan & Evermann, 1896:790. Hepsetia stipes, Jordan & Hubbs, 1919:34. Holotype: M. C. Z. 18227. Paratypes: M. C. Z. 18228, 17 specimens 50 to 68 mm.; M. C. Z. 18229, 4 specimens, 49 to 75 mm.; M. C. Z. 18230, 6 specimens, 65 to 80 mm. No. Poey 582. MUGILIDAE Mugil Mugil Artedi, in Linnaeus, 1758:316 (Mugil cephalus Linnaeus) 51. Mugil lebranchus Poey = Mugil brasiliensis Agassiz. HOWELL Y RIVERO: TYPES OF POEY FISHES 185 Miigil hhrauchis Poey, 1860:260, pi. 18, fig. 3; 1861:379; 1868:388; 1876:98. Mugil hnisiUensis Agassiz, 1829:134, pi. 72; Jordan & Evermann, 1897:810. Cotypes: M. C. Z. 34160-61. Complete unmounted skeletons of two specimens, about 540 and 640 mm. long. No. Poey 193. Type locality : Cienfuegos, Sta. Clara Prov. JOTURUS Joturus Poey, 1860:263 {Joturus pichardi Poey) 52. Joturus pichardi Poey Joturus pichardi Poey, 1860:263, pi. 18, fig. 4-5; 1861:380; 1868:390; 1876:99; Jordan & Ever- mann, 1896:821. Holotype: M. C. Z. 23886. No. Poey 518. Type locality : Rio Almendares, near Habana. SPHYRAENIDAE Sphyraena Sphyraena Rose, in Artedi, 1793:52 {Esox sphyraena Linnaeus) 53. Sphyraena picudilla Poey Sphyraena picudilla Poey, 1860:162; 1861:372; 1868: 359; 1876: 96; Jordan & Evermann, 1896-824. Cotype: M. C. Z. 25967, specimen 365 mm. long. No. Poey 361. GEMPYLIDAE Gempylus Gempylus Cuvier & Valenciennes, 1831b :152 {Gempylus serpens Cuvier & Valenciennes) 54. Gempylus ophidianus Poey = Gempylus serpens Cuvier & Valenciennes Gempylus ophidianus Poey, 1860:246, pi. 18, fig. 1; 1861 :373. Prometheus ofidianus, Poey, 1868:364. Nealotus ophidianus, Poey, 1876:94. Gempylus serpens Cuvier & Valenciennes, 1831b :152; Jordan & Evermann, 1896:884. Holotype: M. C. Z. 8586. No. Poey 408. 186 bulletin: museum of comparative zoology NOMEIDAE NOMEUS Nomeus Cuvier, 1817:315 {Gobius gronovii Gmelin) 55. Nemeus oxyurus Poey = NoMEUS GRONOVII (Gmelin) Nomeus oxyurus Poey, 1860;236; 1868:376; 1876:91. Gobius gronovii Gmelin, 1788:1205. Nomeus gronovii, Jordan & Evermann, 1896:949. Types: M. C. Z. 16963, two specimens, the largest of which is the Holotype. No. Poey 492. Type locality: Habana. CARANGIDAE Hemicaranx Hemicaranx Bleeker, 1862b :135 (Hemicaranx marginatus Bleeker) 56. Caranx heteropygus Poey = Hemicaranx amblyrhynchus (Cuvier & Valenciennes) Caranx heteropygus Poey, 1861:344, 373; 1867:164; 1875b :77. Caranx amblyrhynchus Cuvier & Valenciennes, 1833: 76, pi. 248; Poey, 1866a :328. Carangops amblyrhynchus, Poey, 1868:366. Hemicaranx amblyrhynchus, Jordan & Evermann, 1896:912. Holotype: M. C. Z. 17254. No. Poey 605. Type locality: Habana. Elaphotoxon Elaphotoxon Fowler, 1905:76 (Scomber ruber Bloch) 57. Caranx cibi Poey = Elaphotoxon babtholomaei (Cuvier & Valenciennes) Caranx cibi Poey, 1860:224. Carangoides cibi, Poey, 1866b :15; 1868:366; 1875b :76. Caranx bartholomaei Cuvier & Valenciennes, 1833:75; Jordan & Evermann, 1896:919. Holotype: M. C. Z. 17252. No. Poey 540. HOWELL Y RIVERO: TYPES OF POEY FISHES 187 XUREL Xurel Jordan & Evermann, 1927:505 (Caranx vinctus Jordan & Gilbert) 58. Caranx secundus Poey = Xurel fasciatus (Cuvier & Valenciennes) Caran.r secundus Foey, 1860:223; 1861:373. Carangops secundus Foey, 1866b:15; 1868:367; 1875b: 78. Caranx fasciatus Cuvier & Valenciennes, 1833:53. Hemicaranx secundus, Jordan & Evermann, 1896:914. Cotypes: M. C. Z. 16702-03, two specimens 335 and 385 mm. total length.. No. Poey 186. 59. Caranx frontalis Poey = Xurel lugubris (Poey) Caranx frontalis Foey, 1860:222; 1861:373. Caranx lugubris Poey, 1860:222; 1861:373; Jordan & Evermann, 1896:924. Carangus lugubris, Poey, 1868:365; 1875b :76. Cotype: M. C. Z. 17348, specimen 470 mm. long. No. Poey 552. Hynnis Hynnis Cuvier & Valenciennes, 1833:145 pi. 257 {Hynnis goreensis Cuvier & Valenciennes) 60. Hynnis cubensis Poey Hynnis cubensis Poey, 1860:235; 1861:374; 1868:368; 1875b :79; Jordan & Evermann, 1896:932. Type: M. C. Z. 34155, a complete unmounted skeleton, which, judg- ing by its size and that of the specimen described (770 mm.), might be the holotype. No. Poey 457. SERIOLIDAE Elagatis Elagatis Bennett, 1840:283 (Seriola bipinnulata Quoy & Gaimard) 61. Seriola pinnulata Poey = Elagatis bipinnulatus (Quoy & Gaimard) Seriola pinnulata Foey, 1860:233. Dccaptcrus pinnulatus, Poey, 1861:374. Elagatis pinmdatus, Poey, 1868:373. 188 bulletin: museum of comparative zoology Seriola bipimiulata Quoy & Gaimard, 1824:363, pi. 61, fig. 3. Elagatis hipimndatus, Poey, 1875b :83; Jordan & Evermann, 1896:906. Cotypes: M. C. Z. 17228, 1 specimen 315 mm. long. No. Poey 349. APOGONIDAE Apogon Apogon Lac^pede, IS02 All {Apogonruberhacepede = Mullusimberbisljmna,eus) 62. MoNOPRiON maculatus Poey = Apogon maculatus (Poey) Monoprion maculatus Poey, 1860:123; 1861:362; 1875b :34. Amia maculata, Poey, 1867:235; 1868:304. Apogon macuhitus, Jordan & Evermann, 1896:1109. Holotype: M. C. Z. 9745. Paratype: M. C. Z. 9755. No. Poey 436. 63. Monoprion pigmentarius Poey = Apogon pigmentarius (Poey) Monoprion pigmentarius Poey, 1860:123; 1861:362; 1875b :35. Amia pig7ne?itaria, Poey, 1867:235; 1868:305. Apogon pigmentarius, Jordan & Evermann, 1896:1109. Type: M. C. Z. 9753, probably the holotype, specimen very much dried up, with tip of tail broken. No. Poey 270. 64. Amia binotata Poey = Apogon binotatus (Poey) Amia binotata Poey, 1867:234; 1868:305. Monoprion binotatus, Poey, 1875b :35. Apogon binotatus, Jordan & Evermann, 1896:1109. Cotypes: M. C. Z. 9750, 3 specimens 48.5 to 52.5 mm. No. Poey 660. Apogonichthys Apogonichthys Bleeker, 1854:321 (Apogonichthys perdix Bleeker) 65. Apogonichthys puncticulatus Poey Apogonichthys puncticulatus Poey, 1867:233; 1868: 305; 18V5b:35; Jordan & Evermann, 1896:1111. Holotype:M. C. Z. 9695. No. Poey 643. HOWELL Y RIVERO: TYPES OF POEY FISHES 189 CENTROPOMIDAE Centropomus Centropomus Lac6pede, 1803a :248 {Sciaena undecimalis Bloch) 66. Centropomus appendiculatus Poey = Centropomus undecimalis (Bloch) Centropomus apprndlcidatus Poey, 1860:119, pi. 13, fig. 1; 1861 :362; 1868:280; 187.5b:32. Sciaena undecimalis Bloch, 1792:60; Centropomus undecimalis, Poey, 1865:194; Jordan & Evermann, 1896:1119. Cotypes: M. C. Z. 10274, specimen 2.30 mm. long; 10306, specimen 3.30 mm. long, and 34156, unmounted complete skeleton, about 540 mm. long. No. Poey 51. 67. Centropomus pedimacula Poey Centropomus pedimacula Poey, 1860:122, pi. 13, fig. 4-5; 1861:362; 1868:280; 1875b :33; Jordan & Evermann, 1896:1119. Holotype: M. C. Z. 10273. Paratype: M. C. Z. 10272. No. Poey 560. Type locality: Cienfuegos, Sta. Clara Prov. 68. Centropomus parallelus Poey Centropomus parallelus Poey, 1860:120, pi. 13, fig. 2-3; 1861:362; 1868:280; 1875b :33; Jordan & Evermann, 1896:1122. Holotype: M. C. Z. 10271. No. Poey 134. Type locality: Cienfuegos, Sta. Clara Prov. 69. Centropomus ensiferus Poey Centropomus ensiferus Poey, 1860:122, pi. 12, fig. 1; 1861:362; 1868:280; 1875b :33; Jordan & Ever- mann, 1896:1125. Cotypes: M. C. Z. 10299 and 10300, two specimens 260 and 310 mm. No. Poey 561. - EPINEPHELIDAE Petrometopon Petrometopon GiU, 1865:105 {Perca guttaius Toey = Sparus cruentatus Lac4p6de) 70. Serranus apiarius Poey = Petrometopon cruentatus (Lacepede) 190 bulletin: museum of comparative zoology Serranus apiarius Poey, 1860:143; 1861:364. Petrometopon apiarius, Poey, 1868:288; 1875b :20. Sparus cruentatus Lacepede, 1802:156. Petrometopon cruentatus, Jordan & Evermann, 1896: 1141. Holotype: M. C. Z. 10158. No. Poey 210. ' Menephorus Menephorus Poey, 1871:50 {Serranus dubius Poey) 71. Menephorus punctiferus Poey Menephorus punctiferus Poey, 1875b:21. Bodianu^ punctiferus, Jordan & Evermann, 1896:1147. Holotype: M. C. Z. 10019. No. Poey 309. Epinephelus Epinephelus Bloch, 1793:11 (Epinephelus marginalis) 72. Serranus mystacinus Poey = Epinephelus mystacinus (Poey) Serranus mystacinus Poey, 1852:52, pi. 10, fig. 1; 1861:364; 1867:154. Schistorus mystacinus Poey, 1868:287; 1875b :18. Epinephelus mystacinus, Jordan & Evermann, 1896: 1151. Holotype: M. C. Z. 9991. No. Poey 39. 73. Serranus conspersus Poey = Epinephelus niveatus (Cuvier & Valenciennes) Serranus conspersus Foey, 1860:139; 1861:364; 1867: 157. Serranus niveatus Cuvier & Valenciennes, 1828:285. Epinephelus niveatus, Poey, 1868:286; 1875b :15; Jordan & Evermann, 1896:1156. Cotypes: M. C. Z. 10161, three specimens, 112 to 180 mm. No. Poey 503. Promicrops Promicrops Gill, in Poey, 1868:287 (Serranus guasa Poey = itaiara Lichtenstein) 74. Serranus guasa Poey = Promicrops- itaiara (Lichtenstein) HOWELL Y RIVERO: TYPES OF POEY FISHES 191 Serranuf) guasa Poey, 1860:141, pi. 13, fig. 8; 1861 :354, 363; 1867:154. Promicrops guasa Poey, 1868:287; 1875b :13. Serranus itaiara Lichtenstein, 1821:278. Promicrops guiiatus, Jordan & Evermann, 1898:1162. Type: M. C. Z. 34157, complete unmounted skeleton of paratype, specimen mentioned in Memorias, 1861:354 (720 mm.); besides there is a skull measuring 335 mm. No. Poey 138. Trisotropis Trisotropis Gill, 1865:104 {Johnius gutlatus Bloch & Schneider = Perca venenosa Linnaeus) 75. Serranus bonaci Poey = Trisotropis bonaci (Poey) Serranus honaci Poey, 1860:129; 1861:363; 1867:155. Trisotropis bonaci Foey, 1868:283; 1870a :306; 1875b: 13. Mycteroperca bonaci, Jordan & Evermann, 1896:1174. Cotypes: M. C. Z. 10000 (2 spec), 10032-34, 10195; 6 specimens, 98 to 358 mm. No. Poey 388. 76. Serranus brunneus Poey = Trisotropis bonaci (Poey) Serranus brunneus Poey, 1860:131; 1861:363; 1867: 156. Trisotropis brunneus Poey, 1868:282; 1870a :305; 1875b :13. Serranus bonaci Poey, 1860:121. Mycteroperca bonaci, Jordan & Evermann, 1896:1174. Types: M. C. Z. 10184, 2 specimens, the largest of which is the holotype. No. Poey 208. 77. Serranus cyclopomatus Poey = Trisotropis bonaci (Poey) Serranus cyclopomatus Poey, 1861 :353, 363; 1867:156. Trisotropis cyclopomatus Poey, 1868:284. Trisotropis brunneus Poey, 1870a :305; 1875b :13. Serranus bonaci Poey, 1860:121. Mycteroperca bonaci, Jordan & Evermann, 1896:1174. T^'pes: M. C. Z. 10180, 2 specimens, the largest the holotype, the other a paratype. No. Poey 517. 192 bulletin: museum of comparative zoology 78. Serranus dimidiatus Poey = Trisotropis dimidiatus (Poey) Serranus dimidiatus Poey, 1860:129; 1861:363. Trisotropis diviidiatus, Poey, 1868:285; 1870a :308; 1875b :14. Myderoperca dimidiata, Jordan & Evermann, 1896: 1179. Holotype: M. C. Z. 26953. No. Poey, 350. Mycteroperca Myderoperca Gill, 1862b :236 {Serranus olfax Jenyns) 79. Serranus falcatus Poey = Mycteroperca falcata (Poey) Serranus falcatus Foey, 1860:138; 1861:363. Trisotropis falcatusToey,18dS:285; 1870a :309; 1875b: 15; Jordan & Evermann, 1896:1184. Cotypes: M. C. Z. 10188, 10014, 10183, three specimens 270 to 405 mm.; M. C. Z. 34158, complete unmounted skeleton, about 630 mm. long. No. Poey 43. 80. Mycteroperca calliura Poey Mycteroperca calliura Poey, 1865:181; 1866a :409. Trisotropis calliurus Poey, 1868:284; 1870a :309; 1875b :14. Mycteroperca calliura, Jordan & Evermann, 1896: 1186. Cotype: M. C. Z. 10011, specimen 320 mm. long. No. Poey 65. 81. Serranus felinus Poey = Mycteroperca tigris (Cuvier & Valenciennes) Serraiius felinus Foey, 1860:134; 1861:363; 1867:155. Trisotropis felinus Poey, 1868:283; 1870a :307. Serranus tigris Cuvier & Valenciennes, 1833 :325. Trisotropis tigris, Poey, 1875b :14. Mycteroperca tigris, Jordan & Evermann, 1896:1187. Holotype: M. C. Z. 10071. No. Poey 576. 82. Serranus repandus Poey = Mycteroperca tigris (Cuvier & Valenciennes) Serranus repandus Poey, 1860:135; 1861:363; 1867: 155 Trisotropis felinus, Poey, 1868:283. HOWELL Y RIVERO: TYPES OF POEY FISHES 193 Serranus tigris Cuvier & Valenciennes, 1833:325. Trisotroins tigris, Poey, 1875b :14. Myderopcrca tigris, Jordan & Evermann, 1890:1187. Holotype: M. C. Z. 10076. No. Poey 568. Chorististium Chorististium Gill, 1862a :15 (Liopropoma rubre Poey) 83. Liopropoma rubre Poey = Chorististium rubrum (Poey) Liopropoma rubre Poey, 1861 :418. Chorististium rubrum, Poey, 1868:291; 1875b :32; Jordan & Evermann, 1896:1136. Holotype: M. C. Z. 9691. No. Poey 417. SERRANIDAE Prionodes Prionodes Jenyns, 1842:46 {Prionodes fasciatus Jenyns) 84. Centropristes fusculus Poey = Prionodes fusculus (Poey) Centropristes fusculus Poey, 1861 :342, 365. Haliperca fusculus, Poey, 1868:281; 1875b :22. Prionodes fusculus, Jordan & Evermann, 1896:1211. Holotype: M. C. Z. 10015. No Poey 300 85. Serranus phoebe Poey = Prionodes phoebe (Poey) Serranus phoebe Poey, 1852:55, pi. 2, fig. 3. Centropristes phoebe, Poey, 1861 :365. Haliperca phoebe, Poey, 1868:281 ; 1875b :22. Prionodes phoebe, Jordan & Evermann, 1896:1211. Cotypes: M. C. Z. 36949, 200 mm. total length; 31437, 163 mm. total length. No. Poey 322. 86. Serranus jacome Poey = Prionodes tabacaria (Cuvier & Valenciennes) Serranus jacome Poey, 1852:57, pi. 2, fig. 1. Haliperca jacome, Poey, 1875b :22. Centropristes tabacarius Cuvier & Valenciennes, 1829a :33; Poey, 1861:365. Haliperca tabacaria, Poey, 1868:282. Prionodes tabacarius, Jordan & Evermann, 1896:1215. 194 bulletin: museum of comparative zoology Cotypes: M. C. Z. 10008, 10067, two specimens 145 mm. long No. Poey 216. 87. Serranus praestigiator Poey = Prionodes tigrinus (Bloch) Serranus praestigiator Poey, 1852:58, pi. 2, fig. 2. HaUpcrca pracstigiaior, Poey, 1868:282; 1875b :22. Holoccntrus tigrinus Bloch, 1790:77. Serranus tigrinus, Poey, 1861 :363. Prionodes tigrinus, Jordan & Evermann, 1896:1214. Holotype: M. C. Z. 10241. No. Poey 324. Hypoplectrus Hypopledrus Gill, 1862b:236 (Pledropoma puella Cuvier & Valenciennes = variety of Perca unicolor Walbaum) 88. Hypoplectrus maculiferus Poey = Hypoplectrus unicolor (Walbaum) Hypoplectrus maculiferus Poey, 1871:78, pi. 1, fig. 2; 1875b :24. Perca unicolor Walbaum, 1792:352. Hypoplectrus unicolor, Jordan & Evermann, 1896:1192 Holotype: M. C. Z. 9783, specimen 120 mm. long. No. Poey 390. 89. Hypoplectrus pinnivaria Poey = Hypoplectrus unicolor pinnivarius (Poey) Hypoplectrus pinnivaria Poey, 1868:291; 1875b :24. Hypoplectrus unicolor pinnivarius, Jordan & Ever- mann, 1896:1192. Holotype: M. C. Z. 34035. No. Poey 642. 90. Plectropoma guttavarium Poey = Hypoplectrus unicolor guttavarius (Poey) Plectropoma guttavarius Poey, 1852:70; 1854:441; 1861:364. Hypoplectrus guttavarius, Poey, 1868:291; 1875b :24, Hypoplectrus unicolor pinnivarius, Jordan & Ever- mann, 1896:1192. Cotypes: M. C. Z. 9781, two specimens 123 & 130 mm. long. No. Poey 323. 91. Plectropoma nigricans Poey = Hypoplectrus unicolor nigricans (Poey) HOWELL Y RIVERO: TYPES OF POEY FISHES 195 Pledropoma nigricans Toey, 1852:71; 1861:364. Hypoplcdrus nigricans, Poey, 1868:290; 1875b :24. Hypoplccirus iinicolor nigricans, Jordan & Evermann, 1896:1193. Holotype: M. C. Z. 34036. No. Poey 430. 92. Plectropoma bovinum Poey = Hypoplectrus unicolor bovinum (Poey) Plectropoma hovinum Poey, 1852:69; 1861:364. Hypoplectrus hovinus, Poey, 1868:290; 1875b :23. Hypoplectrus unicolor bovinum, Jordan & Evermann, 1896:1193. Cotypes: M. C. Z. 9795, two specimens 125 & 132 mm. long. No. Poey 129. PRIACANTHIDAE Priacanthus Priacanthus Cuvier, 1817:281 {Priacanthus macrophthalmus Cuvier) 93. Priacanthus catalufa Poey = Priacanthus arenatus Cuvier & Valenciennes Priacanthus catalufa Voey, 1863:182; 1865:274; 1868: 302; 1875b :38. Priacanthus arenatus Cuvier & Valenciennes, 1829:75; Poey, 1865:274; Jordan & Evermann, 1896:1237. Types: M. C. Z. 9766, three specimens, the one bearing Poey's original number label being the holotype. No. Poey 637. PEMPHERIDAE Pempheris Pempheris Cuvier, 1829:195 {Kurtus argenteus Bloch & Schneider = Pemp/iem touea Cuvier 94. Pempheris mulleri Poey Pempheris millleri Poey, 1860:203; 1861:371; 1868: 358; 1876:94; Jordan & Evermann, 1896:978. Types: M. C. Z. 17318, two specimens, the larger of which is the holotype. No. Poey 415. 196 bulletin: museum of comparative zoology LUTIANIDAE LUTIANUS Lutianus Bloch, 1790:105 (Lutianus lutianus Bloch) 95. Mesoprion caudanotatus Poey = Lutianus buccanella (Cuvier & Valenciennes) Mesoprion caudanotatus Poey, 1854:440, pi. 3, fig. 3; 1861:365; 1867 ;158. Mesoprion buccanella Cuvier & Valenciennes, 1828: 344; Poey, 1868:295. Lutjanus buccanella, Poey, 1875b :27. Neomenis buccanella, Jordan & Evermann, 1898:1261. Cotypes : M. C. Z. 9804; 9823, 9870, 9932, 9888, from 150 to 260 mm. No. Poey 217. 96. Mesoprion profundus Poey = Lutianus vivanus (Cuvier & Valenciennes) Mesoprion profundus Foey, 1860:150; 1861:365; 1865: 267; 1867:157; 1868:294. Lutjanus profundus, Poey, 1875b :28. Mesoprion vivanus Cuvier & Valenciennes, 1828:343. Neomenis vivanus, Jordan & Evermann, 1898:1262. Cotypes: 9966, 9990, two specimens 340 & 415 mm. long. No. Poey 28. 97. Mesoprion campechanus Poey = Lutianus campechanus (Poey) Mesoprion campechanus Poey, 1860:149; 1861:365; 1868:294; 1875b :29. Neomenis aya, Jordan & Evermann, 1898:1264. Types: M. C. Z. 9982, two specimens the largest of which is the holotype. No. Poey 71. 98. Mesoprion ambiguus Poey = Lutianus ambiguus (Poey) Mesoprion ambiguus Poey, 1860:152, pi. 12, fig. 4, pi. 13, fig. 18; 1861:365.^ Ocyurus ambiguus, Poey, 1868:295. Lutjanus ambiguus, Poey, 1875b :30. Neomenis ambiguus, Jordan & Evermann, 1898:1272. Holotype: M. C. Z. 9951. No. Poey 151. HOWELL Y RIVERO: TYPES OF POEY FISHES 197 99. Mesoprion ojanco Poey = LuTiANUS MAHOGONi (Cuvier & Valenciennes) Mesoprion ojanco Poey, 1860: 150, pi. 13, fig. 10; 1861: 365; 1868:295. Lutjanus ojanco, Poey, 1875b :28. Mesoprion mahogoni Cuvier & Valenciennes, 1828:338. Neomenis mahoqoni, Jordan & Evermann, 1898:1272. Cotypes: M. C. Z. 9917, 9939, 9950, 9952, 8 specimens 220 to 320 mm. No. Poey 86. Rhomboplites Rhomboplites Gill, 1862b :237 (Centropristis aurorubens Cuvier & Valenciennes) 100. Mesoprion elegans Poey = Rhomboplites aurorubens (Cuvier & Valenciennes) Mesoprion elegans Poey, 1860:153; 1861:365; 1867: 158. Rhomhopliies elegans, Poey, 1868:295; 1875b :31. Centropristis awrorw^en* Cuvier & Valenciennes, 1829a: 34. RJwinhopliies aurorubens, Jordan & Evermann, 1898: 1277. Cotypes: M. C. Z. 22138, 3 specimens 290 to 330 mm. No. Poey 273. Tropidinius Tropidinius Gill, 1868:296 (Mesoprion arnillo Poey =Apsilus dentatus Guichenot) 101. Mesoprion arnillo Poey = Tropidinius dentatus (Guichenot) Mesoprion arynllo Poey, 1860:154; 1861:365. Tropidinius arnillo, Poey, 1868:296; 1875b :30. Apsilus dentatus Guichenot, 1853:29, pi. 1 fig. 2; Jordan & Evermann, 1898:1278. Types: M. C. Z. 9954, 2 specimens, the largest of which is the holo- type. No. Poey, 142. Pristipomoides Pristipomoides Bleeker, 1852:574 (Pristipomoides typus Bleeker) 102. Mesoprion vorax Poey = Pristipomoides macrophthalmus (Miiller & Troschel) 198 bulletin: museum of comparative zoology Mesoprion vorax Poey, 1860:151; 1861:365. Platyinius vorax, Poey, 1868:292; 1875b :31. Centropristis macro phthalrnus Miiller & Troschel, 1848:666. Aprion macro phthalrnus, Jordan & Evermann, 1898: 1280. Holotype: M. C. Z. 26479. No. Poey 472. VERILIDAE Verilus Verilus Poey, 1860:124 {Verilus sordidus Poey) 103. Verilus sordidus Poey Verilus sordidus Poey, 1860:125, pi. 12, fig. 6; 1861: 362; 1868:291; 1875b :32; Jordan & Evermann, 1898:1284. Holotype: M. C. Z. 21764. No. Poey 141. HAEMULIDAE Haemulon Haemulon Cuvier, 1829:175 {Haemulon elegans Cuvier = /Sparus sciurus Shaw) 104. Haemulon notatum Poey = Haemulon bonariensis Cuvier & Valenciennes Haemulon notatum Poey, 1860:179; 1861:369; 1868: 317; 1875b:46. Haemulon bonariensis Cuvier & Valenciennes, 1830a: 174; Jordan & Evermann, 1898:1297. Cotype: M. C. Z. 10472, 288 mm. long. No. Poey 348. 105. Haemulon carbonarium Poey Haemulon carbonarium Poey, 1860;176; 1861:369; 1868:318; 1875b:44; Jordan & Evermann, 1898: 1300. Cotypes: M. C. Z. 10502, 3 specimens, 255 to 280 mm. No. Poey 367. 106. Haemulon dorsale Poey = Haemulon melanurum (Linnaeus) HOWELL Y RIVERO: TYPES OF POEY FISHES 199 Haemulon dorsale Poey, 1860:179; 1861:369; 1868: 318; 1875b :44. Perca mclanura Linnaeus, 1758:292. Haemulon melanurum, Jordan & Evermann, 1898: 1302. Cotype: M. C. Z. 10590, 205 mm. long. No. Poey 364. 107. Haemulon luteum Poey = Haemulon sciurus (Shaw) Haemulon luteum Poey, 1860:174; 1861:354; 369; 1868:317; 1875b :44. Spams sciurus Shaw, 1803:439, pi. 64. Haemulon sciurus, Jordan & Evermann, 1898:1303. Cotype: M. C. Z. 1078a, 192 mm. long. No. Poey 511. 108. Haemulon multilineatum Poey = Haemulon sciurus (Shaw) Haemulon multilineatum Poey, 1860:178; 1861:369; 1868:318; 1875b :44. Sparus sciurus Shaw, 1803:439, pi. 64. Haemulon sciurus, Jordan & Evermann, 1898:1303. Holotype: M. C. Z. 10478. No. Poey 376. 109. Haemulon arara Poey = Haemulon plumieri (Lacepede) Haemulon arara Poey, 1860:177; 1861:369; 1868:318; 1875b :45. Labrus plumieri Lacepede, 1802:480, pi. 2, fig. 2. Haemulon plumieri, Jordan & Evermann, 1898:1304. Holotype: M. C. Z. 10545, specimens very much dried up. No. Poey 291. Brachygenys Brachygenys Scudder, in Poey, 1875b :47 {Haemulon taeniatum Foey = Haemu- lon chrysargyreum Giinther) 110. Haemulon taeniatum Poey = Brachygenys chrysargyreus (Giinther) Haemulon taeniatum Poey, 1860: 182; 1861:369; 1868: 319. Brachygenys taeniata, Poey, 1875b :47. Haemulon chrysargyreus Giinther, 1859:314. Brachygenys chrysargyreus, Jordan & Evermann, 1898: 1307. 200 bulletin: museum of comparative zoology Types: M. C. Z. 10482, 3 specimens, the largest of which is the holo- type. No. Poey 369. Bathystoma Bathystoma Scudder in Putnam, 1863:12 (Haemulon jeniguano Poey =Haemu- lon aurolineatum Cuvier & Valenciennes) 111. Haemulon jeniguano Poey = Bathystoma aurolineatum (Cuvier & Valenciennes) Hacvmlon jeniguano Poey, 1860:183; 1861:369; 1868: 319; 1875b :47. Haemulon aurolineatum Cuvier & Valenciennes, 1830a: 176. Bathystoma aurolineatum, Jordan & Evermann, 1898:1310. Holotype: M. C. Z. 1080. No. Poey 420. 112. Haemulon quinquelineatum Poey = Bathystoma striatum (Linnaeus) Haemulon quinquelineatum Voey, 1861:419; 1867:161. Haemulon quadralineatum Poey, 1868:319; 1875b :47. Perea striata Linnaeus, 1758:293. Bathystoina striatum, Jordan & Evermann, 1898:1310. Holotype: M. C. Z. 10542. No. Poey 211. Anisotremus Anisotremus Gill, 1861:105 {Sparus virginicus Linnaeus) 113. Haemulon obtusum Poey = Anisotremus surinamensis (Bloch) Haemulon obtusum Poey, 1860:182; 1861:369. Anisotremus obtusus, Poey, 1868:312; 1875b :43. Lutjanus surinamensis Bloch, 1790:98. Anisotremus surinamensis, Jordan & Evermann, 1898:1318. Cotype: M. C. Z. 34159, complete unmounted skeleton, about 540 mm. long. No. Poey 170. 114. Pristipoma spleniatum Poey = Anisotremus spleniatus (Poey) Pristipoma spleniatum Poey, 1860:187. Anisotremus spleniatus, Poey, 1861:368; 1875b :43; Jordan & Evermann, 1898:1321. Cotype: M. C. Z. 21778, 65 mm. total length. No. Poey 49. HOWELL Y RIVERO: TYPES OF POEY FISHES 201 POMADASYS Po7nadasys Lacepede, 1803a :516 (Sciaena argentea Forsk&l) 115. Pristipoma productum Poey = P0MADASYS PRODUCTUS (Poev) Pristipoma producium Poey, 1860:186; 1861:368; 1868:311; 1875b :42. Pomadasys product us, Jordan & Evermann, 1898:1332. Cotype: M. C. Z. 21889, 245 mm. long. No. Poey 418. Rhonciscus Rhonciscus Jordan & Evermann, 1895:387 {Pristipoma crocro Cuvier & Valenciennes) 116. Pristipoma cultriferum Poey = Rhonciscus crocro (Cuvier & Valenciennes) Pristipoma cultriferum Poey, 1860:185; 1861:368; 1868:310; 1875b:41. Pristipoma crocro Cuvier & Valenciennes, 1830a :197. Pomadasys crocro, Jordan & Evermann, 1898:1333. Cotypes: M. C. Z. 10594, 10592; 145 and 140 mm. No. Poey 84. SPARIDAE calamus Calamus .Swainson, 1839:171, 221 {Pagellus calamus Cuvier & Valenciennes) 117. Pagellus orbitarius Poey = Calamus calamus (Cuvier & Valenciennes) Pagellus orbitarius Poey, 1860:201; 1861:367. Sparus orbitarius, Poey, 1868:308. Calamus orbitarius, Poey, 1872:179, pi. 6, fig. 2; 1875b :56. Pagellus calamus Cuvier & Valenciennes, 1830b :152, pi. 152. Calamus calamus, Jordan & Evermann, 1898:1349. Holotype: M. C. Z. 21849. No. Poey 149. 118. Pagellus caninus Poey = Calamus bajonado (Bloch & Schneider) 202 bulletin: museum of comparative zoology Pagellus caninus Poey, 1860:199; 1861:367; 1867:160. Sparus bajonado Bloch & Schneider, 1801:284; Poey, 1868:308. Calamus bajonado, Poey, 1872:176, pi. 6, fig. 1; - 1875b :55; Jordan & Evermann, 1898:1352. Cotypes: M. C. Z. 21835, 21834, 390 and 430 mm. No. Poey 468. 119. Calamus macrops Poey Calamus macrops Poey, 1872:181, pi. 7, fig. 3; 1875b: 56; Jordan & Evermann, 1898:1354. Holotype: M. C. Z. 21839. No. Poey 221. 120. Pagellus humilis Poey = Calamus penna (Cuvier & Valenciennes) Pagellus humilis Poey, 1868:308. Grammateus humilis, Poey, 1872:182; 1875b :56. Pagellus penna Cuvier & Valenciennes, 1830b :154. Calamus penna, Jordan & Evermann, 1898:1354. Cotypes: M. C. Z. 21843-44, 195 & 185 mm. total length. No. Poey 288. 121. Grammateus medius Poey = Calamus medius (Poey) Grammateus medius Poey, 1872:183, pi. 7, fig. 4; 1875b :56. Calamus medius, Jordan & Evermann, 1898:1356. Paratype: M. C. Z. 21842. No. Poey 192. Salema Salema Jordan & Evermann, 1895:390 {Perca unimaculala Bloch) 122. Sargus caribaeus Poey = Salema rhomboidalis (Linnaeus) Sargus caribaeus Poey, 1860:197; 1861:367; 1868:309; 1875b :56. Perca rhomboidalis Linnaeus, 1758. Archosargus unimaculatus, Jordan & Evermann, 1898:1359. Cotype: M. C. Z. 21713, 3 specimens 175 to 260 mm. No. Poey 571. Type locality : Batabano. HOWELL Y RIVERO: TYPES OF POEY FISHES 203 DiPLODUS Diplodus Rafinesque, 1810b:54 (Sparus annularis Linnaeus) 123. Sargus caudimacula Poey = Diplodus argenteus (Cuvier & Valenciennes) Sargus caudimacula Foey, 1860:198; 1861:367; 1868: 310; 18751) :57. Sargus argenteus Cuvier & Valenciennes, 1830b :44. Diplodus argenteus, Jordan & Evermann, 1898:1363. Cotype: M. C. Z. 21715, 275 mm. long. No. Poey 589. KYPHOSIDAE Kyphosus Kyphosus Lacepede, 1802:114 {Kyphosus bigibbus Lacepede =/uscus Lacepede) 124. Pimelepterus flavo-lineatus Poey = Kyphosus incisor (Cuvier & Valenciennes) Pimelepterus flavo-lineatus Foey, 1866a :319; 1868:324; 1875b :65. Pimelepterus incisor Cuvier & Valenciennes, 1831a: 198; Poey, 1866a :319. Kyphosus incisor, Jordan & Evermann, 1898:1386. Holotype: M. C. Z. 21712. No. Poey 371. GERRIDAE Eugerres Eugerres Jordan & Evermann, 1927:506 {Gerres piumim Cuvier & Valenciennes) 125. Gerres patao Poey = Eugerres brasilianus (Cuvier & Valenciennes) Gerres patao Poey, 1860:192; 1868:320; 1875b :50. Gerres hra^silianus Cuvier & Valenciennes, 1830b :344, 458; Jordan & Evermann, 1898: 1378. Cotype: M. C. Z. 23151, 290 mm. long. No. Poey 173. MULLIDAE Upeneus Upeneus Cuvier, 1829:157 (Upeneus bifasciatus Lacepede) 204 bulletin: museum of comparative zoology 126. Upeneus flavovittatus Poey = Upeneus martinicus Cuvier & Valenciennes Upeneus flavovittatus Poey, 1853:224, pi. 17, fig. 4; 1861:367. Mulloidcs flavovittatus, Poey, 1868:307; 1875b :34. Upeneus martinicus Cuvier & Valenciennes, 1829a: 356; Jordan & Evermann, 1896:859. Holotype: M. C. Z. 21819. No. Poey 281. SCIAENIDAE CORVULA Corvula Jordan & Evermann, 1886:377 {Johnius batabanus Poey) 127. Johnius batabanus Poey = Corvula batabana (Poey) Johnius batabanus Poey, 1860:184; 1861:370; 1868: 324; 1875b :49. Corvula batabana, Jordan & Evermann, 1898:1430. Cotype: M. C. Z. 10926-27, 21957, three specimens 184 to 228 mm. No. Poey 85. Type locality: Batabano, Havana Pro v. MALACANTHIDAE Caulolatilus Caulolatilus Gill, 1862b:240 (?Latilus chrysops Cuvier & Valenciennes) 128. Caulolatilus cyanops Poey Caulolatilus cyanops Poey, 1866a:311, 312; 1868:330; 1876:95; Jordan & Evermann, 1898:2278. Cotype: M. C. Z. 12826, 265 mm. long. No. Poey 412. CHAETODONTIDAE Prognathodes Prognathodes Gill, 1862b :238 (Chelmo pelta Guniher = Chelmo aculeatus Poey) 129. Chelmo aculeatus Poey = Prognathodes aculeatus (Poey) HOWELL Y RIVERO: TYPES OF POEY FISHES 205 Chelnw arideaim Poey, 1860:202; 1861 :371. Proqnathodcs acitlratus, Poey, 1868:354; 1875b :63; " Jordan & Evermann, 1898:1671. Holotype: M. C. Z. 16253. No. Poey 56. Type locality: Matanzas. Chaetodon Chaetodon Linnaeus, 1758:272 {Chaetodon capistratus Linnaeus) 130. Sarothrodus amplecticollis Poey = Chaetodon bimaculatus Bloch Sarothrodus amplecticollis Poey, 1868:353. Sarothrodus amplexicollis, Poey, 1875b :63, pi. 7, fig. 1-3 Chaetodon bimaculatus Bloch, 1790, pi. 219, fig. 1; Poev, 1861:371. Sarothrodus bimaculatus, Poey, 1868:353; 1875b :62. Chaetodon ocellatus, Jordan & Evermann, 1898:1674. Holotype: M. C. Z. 16250. No. Poey 665. 131. Chaetodon sedentarius Poey Chaetodon sedentarius Poey, 1860:203; 1861:371; Jordan & Evermann, 1898:1675. Sarothrodus sedentarius, Poey, 1868:354; 1875b :62. Types: M. C. Z. 16198, two specimens the largest of which is the holotype. No. Poey 247. Type locality : Cienfuegos, Sta Clara Prov. 132. Sarothrodus ataeniatus Poey = Chaetodon ataeniatus (Poey) Sarothrodus ataeniatus Poey, 1868:353; 1875b :63. Chaetodon ataeniatus, Jordan & Evermann, 1898:1676. Holotype: M. C. Z. 16251. No. Poey 250. POMACANTHUS Pomacanthus Lacepede, 1803a:517 {Chaetodon arcuatus Linnaeus) 133. Chaetodon littoricola Poey = Pomacanthus paru (Bloch) Chaetodon littoricola Poey, 1868:351; 1875b :60. Chaetodon paru Bloch, 1787:57. 206 bulletin: museum of comparative zoology Pomacanthns arcuatus, Jordan & Evermann, 1898: 1679. Poviacanthus parn, Jordan & Evermann, 1898:1680. Types: M. C. Z. 16165, two specimens, the largest of which is the holotype. No. Poey 577. ACANTHURIDAE ACANTHURUS Acanthurus Forskil, 1775:59 {Chaetodon unicornis), etc. 134. Acanthurus tractus Poey = Acanthurus bahianus Castelnaii Acanthurus tractus Voey, 1860:208; 1861:372; 1868: 356; 1875b :67. Acanthurus bahianus Castelnau, 1855:24, pi. 11, fig. 1. Teuthis hahianus, Jordan & Evermann, 1898:1691. Holotype: M. C. Z. 21856. No. Poey 447. SCORPAENIDAE scorpaena Scorpaena (Artedi) Linnaeus, 1758:266 (Scorpaena porcus Linnaeus) 135. Scorpaena rascacio Poey = Scorpaena plumieri Bloch Scorpaena rascacio Poey, 1860:169; 1861:366; 1868: 303; 1875b :40. Scorpaena plumieri Bloch, 1798:234; Jordan & Ever- mann, 1898:1848. Cotypes: M. C. Z. 13948-50, 13957, 4 specimens 165 to 305 mm. No. Poey 359. 136. Scorpaena occipitalis Poey = Scorpaena inermis Cuvier & Valenciennes Scorpaena occipitalis Foey, 1860:171; 1861:366; 1868: 303; 1875b :41. Scorpaena inermis Cuvier & Valenciennes, 1829b :228; Jordan & Evermann, 1898:1853. Types: M. C. Z. 13894, 5 specimens, the largest of which is the holo- type. No. Poey 474. HOWELL Y RIVERO: TYPES OF POEY FISHES 207 PERISTEDIIDAE VULSICULUS Vulsiculus Jordan & Evermann, 1895:489 (Peristedion imberbe Poey) 137. Pterystedion imberbe Poey = Vulsiculus imberbe (Poey) Pterystedion imberbe Poey, 1861 :367, 389 (Note No. 25) Peristedioji imberbe, Poey, 1867:158; 1868:304, 462. Peristedion micronemus Poey, 1870b :321. Vidsiculus imberbis, Jordan & Evermann, 1898:2181. Holotype: M. C. Z. 13566. No. Poey 523. POMACENTRIDAE FURCARIA Furcaria Poey, 1860:194 {Furcaria puncta Foey = HeUases multilineatus Guichenot) 138. Furcaria cyanea Poey Furcaria cyanea Poey, 1860:196, pi. 14, fig. 4; pi. 13, fig. 21-22; 1861:370; 1868:330; 1876:104. Chromis cyaneus, Jordan & Evermann, 1898:1547. Holotype: M. C. Z. 14670. No Poey 460. 139. Furcaria puncta Poey = Furcaria multilineata (Guichenot) Furcaria puncta Poey, 1860:195; 1861:370; 1867:161; 1868:329; 1876:104. Helia-ses multilineatus Guichenot, 1853:76, pi. 2, fig. 2. Chromis multilineatus, Jordan & Evermann, 1898: 1547. Holotype: M. C. Z. 14664. No. Poey 209. Eupomacentrus Eupomacentrus Bleeker, 1877:73 {Pomacentrus lividus Bleeker) 140. Pomacentrus xanthurus Poey = Eupomacentrus leucostictus (Miiller & Troschel) Pomacentrus xanthurus Poey, 1860:190; 1861:370; 1868:326; 1876:101. 208 bulletin: museum of comparative zoology Pomacentrus leucostictus Miiller & Troschel, 1848:674. Eupomacentrus leucostictus, Jordan & Evermann, 1898:1555. Cotypes: M. C. Z. 14677a, 3 specimens 90 to 108 mm. No .Poey 481. 141. Pomacentrus caudalis Poey = Eupomacentrus leucostictus (Miiller & Troschel) Pomacentrus caudalis Poey, 1868:328, 1876:102. Pomacentrus leucostictus Miiller & Troschel, 1848:674. Eupomacentrus leucostictus, Jordan & Evermann, 1898:1555. Types: M. C. Z. 14682, 7 specimens from 18.5 to 48 mm., the largest of which is the holotype. No. Poey 546. 142. Pomacentrus analis Poey = Eupomacentrus analis (Poey) Pomacentrus analis Foey, 1868:327; 1876:101. Eupomacentrus analis, Jordan & Evermann, 1898: 1554. Holotype: M. C. Z. 14678. No. Poey 587. 143. Pomacentrus partitus Poey = Eupomacentrus partitus (Poey) Pomacentrus partitus Poey, 1868:327; 1876:102. Eupomacentrus partitus, Jordan & Evermann, 1898: 1558. Holotype: M. C. Z. 14680. No. Poey 702. 144. Pomacentrus obscuratus Poey = Eupomacentrus adustus (Troschel) Pomacentrus obscuratus Poey, 1876:101. Pomacentrus adustus Troschel, 1865:633. Eupomacentrus adustus, Jordan & Evermann, 1898: 1551. Types: M. C. Z. 14681, 4 specimens, the largest of which is the holotype. No. Poey 586. Stegastes Stegastes Jenyns, 1842:63 (Stegastes imbricatus Jenyns) 145. Pomacentrus denegatus Poey = Stegastes ciirysurus (Cuvier & Valenciennes) Pomacentrus denegatus Foey, 1860:190; 1861:370. HOWELL y RIVERO: TYPES OF POEY FISHES 209 Ghjpkisodon chrysurus Cuvier & Valenciennes, 1830a:- 350. Micros pat hodon chrysurus, Poey, 1868:329; 1876:103; Jordan & Evermann, 1898:1567. Cotypes: IVI. C. Z. 14672, 14675, two specimens 135 & 142 mm. No. Poey 391. LABRIDAE BODIANUS Bodianus Bloch, 1790:31, 33 (Bodianus bodianusBloch = LabrusrufusLmnsie\is) 146. CossYPHUS PULCHELLUS Poey = Bodianus pulchellus (Poey) Cossi/phus pulchellus Poey, 1860:208; 1861:378. Bodianus pulchellus, Poey, 1868:332, 459; 1876:105. Harpe ptdchclla, Jordan & Evermann, 1898:1584. Holotype: M. C. Z. 14292. No. Poey 419. Decodon Decodon Giinther, 1862:101 {Cossyphus puellaris Poey) 147. Cossyphus puellaris Poey = Decodon puellaris (Poey) Cossyphus puellaris Foey, 1860:210; 1861:378. Decodon puellaris, Poey, 1868:332; 1876:107; Jordan & Evermann, 1898:1584. Types: M. C. Z. 14339, two specimens, the largest of which is the holotype. No. Poey 385. CORIDAE Iridio Iridio Jordan & Evermann, 1895:412 {Julis dimidiotus Agassiz = Lahrus cyano- cephalus Bloch) 148. Julis humeralis Poey = Iridio bivittata (Bloch) Julis humeralis Poey, 1860:212; 1861:378. Choerojulis humeralis, Poey, 1868:335; 1876:108. Lahrus bivittatus Bloch, 1791:133. Iridio birittattis, Jordan & Evermann, 1898:1595. Types: M. C. Z. 14200, 3 specimens, the largest of which is the holotype. No. Poey 397. 210 bulletin: museum of comparative zoology 149. Julis internasalis Poey = Iridio cyanocephalus (Bloch) Julis internasalis Poey, 1861:421. Chocrojulis iniernasalis, Poey, 1868:334; 1876:108. Labrus cyanocephalus Bloch, 1791:139. Iridio cyanocephalus, Jordan & Evermann, 1898:1594. Cotypes: M. C. Z. 14252, 3 specimens 210 to 245 mm. No. Poey 258. 150. Julis cinctus Poey = Iridic garnoti (Cuvier & Valenciennes) Julis cinctus Poey, 1860:211, pi. 13, fig. 19; 1861:378. Choerojulis cinctus, Poey, 1868:334; 1876:108. ' Julis garnoti, Cuvier & Valenciennes, 1839a :285. Iridio garnoti, Jordan & Evermann, 1898:1593. Holotype: M. C. Z. 14265. No. Poey 338. 151. Julis ruptus Poey = Iridic garnoti (Cuvier & Valenciennes) Jidis ruptus Poey, 1860:212, pi. 13, fig. 20; 1861:378. Choerojulis ruptus, Poey, 1868:334. Choerojulis cinctus, Poey, 1876:108. Julis garnoti, Cuvier & Valenciennes, 1839a :285. Iridio garnoti, Jordan & Evermann, 1898:1593. Holotype: M. C. Z. 14284. No. Poey 275. SPARISOMIDAE Cryptctomus Cryptotomus Cope, 1870:462 {Cryptotomus roseus Cope) 152. Callicdcn retractus Poey = Cryptotomus retractus (Poey) CaUiodon retractus Poey, 1868:345; 1876:116. Cryptotomus retractus, Jordan & Evermann, 1898: 1623. Holotype: M. C. Z. 14461. No. Poey 558. Spariscma Sparisoma Swainson, 1839:227 (Scarus abilgardii Bloch) 153. Scarus lacrimcsus Poey = Spariscma radians (Cuvier & Valenciennes) HOWELL Y RIVERO: TYPES OF POEY FISHES 211 Scarus lacrimosus Foey, 1861:422; 1868:343; 1876:113. Scams radians, Cuvier & Valenciennes, 1839b :153. Sparisoma radians, Jordan & Evermann, 1898:1631. Holotype: M. C. Z. 14538. No. Poey 632. 154. ScARUS MiNiOFRENATUS Poey = Sparisoma aurofrenatum (Cuvier & Valenciennes) Scanis miniofrcnatus Poey, 1861:379, 393 (Note No. 61); 1867:164; 1868:337; 1876:111. Scarus aurqfrenatus, Cuvier & Valenciennes, 1839b: 142; Poey, 1866a :374. Sparisoma aurofrenatum, Jordan & Evermann, 1898: 1634. Neotypus: M. C. Z. 14545, 14547, 14550, three specimens, 225 to 227 mm. Although the name given by Poey was a substitute for that of Cuvier, these specimens were sent labeled as miniofrenatus by Poey. No measurements have ever been given. No. Poey 365. 155. Scarus distinctus Poey = Sparisoma distinctum (Poey) Scarus distinctus Foey, 1861:423; 1867:163; 1868:341; 1876:114. Sparisoma distinctum, Jordan & Evermann, 1898: 1635. Cotypes: M. .C. Z. 14513, 14540, 14544, 200 to 250 mm. No. Poey 333. 156. Scarus lateralis Poey = Sparisoma chrysopterum (Bloch & Schneider) Scarus lateralis Poey, 1860:219; 1861:379; 1867:162; 1868:337; 1876:112. Scarus chrysopterus Bloch & Schneider, 1801:286; pi. 57; Poey, 1866:373,375. Sparisoma chrysopterum, Jordan & Evermann, 1898: 1636. Holotype: M. C. Z. 14520. No. Poey 462. 157. Scarus squalidus Poey = Sparisoma flavescens (Bloch & Schneider) Scarus squalidus Poey, 1860:218; 1861:379; 1868:338. Scarus flavescens Bloch & Schneider, 1801:290; Poey, 1876:113. 212 bulletin: museum of comparative zoology Sparisoma flavescens, Jordan & Evermann, 1898: 1639. Cotypes: M. C. Z. 14519, 14522. two specimens, 320 & 360 mm. long. No. Poey 463. 158. ScARUS circumnotatus Poey = Sparisoma rubripinne (Cuvier & Valenciennes) Scarus circumnotatus Poey, 1861:423; 1868:340; 1876:114. Scarus riibripinne Cuvier & Valenciennes, 1839b :147; Poey, 1866a :374. Sparisoma rubripinne, Jordan & Evermann, 1898: 1640. Holotype: M. C. Z. 14512. No. Poey 279. 159. Scarus brachialis Poey = Sparisoma brachiale (Poey) Scarus hrachialis Foey, 1861:345,379; 1868:337; 1876: 113. Sparisoma brachiale, Jordan & Evermann, 1898:1641. Holotype: M. C. Z. 14555. No. Poey 607. SCARIDAE Scarus Scarus ForskM, 1775:25 (Scarus croicensis Bloch) 160. PSEUDOSCARUS GNATHODUS Poey = Scarus gnathodus (Poey) Pscndoscarus gnathodus Poey, 1867:240; 1868:350; 1876:119. Scarus gnathodus, Jordan & Evermann, 1898:1650. Cotype: M. C. Z. 14578, 270 mm. long. No. Poey 608. 161. PsEUDOSCARUS LINEOLATUS Poey = SCARUS CROICENSIS Bloch Pseudoscarus lincolatus Poey, 1867:240; 1868:350; 1876:119. Scarus croicensis Bloch, 1790:27; Jordan & Evermann, 1898:1650. Cotypes: M. C. Z. 31448, 14565, 182 & 185 mm. total length. No. Poey 282. HOWELL Y RIVERO: TYPES OF POEY FISHES 213 ELEOTRIDAE DORMITATOR Dormilator Gill, 1862b :240 {Dormitaior gundlachi Poey) 162. Eleotris omocyaneus Poey = Dormitator maculatus (Bloch) Eleotris omocyaneus Poey, 1860:269; 1861:381; 1867: 167. Dormitator omocyaneus, Poey, 1868:396; 1876:128. Sciaena maculata Bloch, 1785, pi. 299, fig. 2. Dormitator maculatus, Jordan & Evermann, 1898: 2196. Holotype: M. C. Z. 13372, male; paratype: M. C. Z. 13371, female. No. Poey 298. 163. Eleotris gundlachi Poey = Dormitator MACULATUS (Bloch) Eleotris gundlachi Poey, 1860:272; 1861:381. Dormitator gundlachi, Poey, 1868:396; 1876:128. Sciaena maculata Bloch, 1785, pi. 299, fig. 2. Dormitator maculatus, Jordan & Evermann, 1898: 2196. Paratype: M. C. Z. 13374, 200 mm. long. No. Poey 553. Erotelis Erotelis Poey, 1860:273 (Erotelis valenciennesi Poey = Erotelis stnaragdus Cuvier & Valenciennes) 164. Erotelis valenciennesi Poey = Erotelis smaragdus (Cuvier & Valenciennes) Erotelis valenciennesi Poey, 1860:273; 1861:381; 1868:396; 1876:127. Eleotris smaragdus Cuvier & Valenciennes, 1837b :173. Erotelis smaragdus, Jordan & Evermann, 1898:2204. Holotype: M. C. Z. 12567. No. Poey 203. GOBIIDAE Bathygobius Bathygobius Bleeker, 1 878 :54 (Gobius nebulo-pundatus Riippell = Gobius fuscus Ruppell) 214 bulletin: museum of comparative zoology 165. GoBius mapo Poey = Bathygobius soporator (Cuvier & Valenciennes) Gobius mapo Poey, 1860:277; 1861:380; 1868:392. Gobius soporator Cuvier & Valenciennes, 1837b :42; Poey 1876:124; Jordan & Evermann, 1898:2216. Types: M. C. Z. 13116, 13117; both numbers in the same bottle. five specimens from 46 to 110 mm., the largest of which is the holotype, No. Poey 498. 166. Gobius lacertus Poey = Bathygobius soporator (Cuvier & Valenciennes) Gobius lacertus Poey, 1860:278; 1861:380; 1867:167; 1868:392; 1876:125. Gobius soporator Cuvier & Valenciennes, 1837b :42; Poey, 1876:124; Jordan & Evermann, 1898:2216. Types: M. C. Z. 13114, 9 specimens from 40 to 92 mm., the largest of which is the holotype. No. Poey 583. 167. Gobius brunneus Poey = Bathygobius soporator (Cuvier & Valenciennes) Gobius brunneus Poey, 1868:393; 1876:125. Gobius soporator Cuvier & Valenciennes, 1837b :42; Poey 1876:124; Jordan & Evermann, 1898:2216. Types: M. C. Z. 13110, two specimens, of which the largest is the holotype. No. Poey 650. GOBIONELLUS Gobionellus Girard, 1858:168 (Gobionellus hastatus Girard) 168. Smaragdus costalesi Poey = Gobionellus lyricus (Girard) Smaragdus costalesi Poey, 1860:280; 1861:380. Gobionellus costalesi, Poey, 1868:394; 1876:126. Gobius lyricus Girard, 1858:169; Jordan & Evermann, 1898:2224. Holotype: M. C. Z. 13109. No. Poey 613. Type locality : Rio Almendares, near Puentes Grandes, Habana. 169. Smaragdus stigmaticus Poey = Gobionellus stigmaticus (Poey) Smaragdus stigmaticus Poey, 1860:281; 1861:380. Gobionellus stigmaticus, Poey, 1868:294; 1876:126. Gobius stigmaticus, Jordan & Evermann, 1898:2224. Holotype: M. C. Z. 13104. No. Poey 289. howell y rivero: types of poey fishes 215 Chonophorus Chonophorus Poey, 1860:274 {Chonophorus bucculentus Foey = Gobius banana Cuvier & Valenciennes) 170. Chonophorus bucculentus Poey = Chonophorus banana (Cuvier & Valenciennes) Chonophorus bucculentus Toey, 1860:275; 1861:381. Rhiuogobius bucculentus, Poey, 1868:394; 1876:125. Gobius banana, Cuvier & Valenciennes, 1837b :78. Aioaous taiasica, Jordan & Evermann, 1898:2236. Cotypes: M. C. Z. 13330, 13379-80, 3 specimens, 290 to 330 mm. No. Poey 441. 171. Chonophorus contractus Poey = Chonophorus banana (Cuvier & Valenciennes) Chonophorus contractus Poey, 1861 :424. Rhinogobius contractus, Poey, 1870:322; 1876:125. Gobius banana, Cuvier & Valenciennes, 18371) :78. Awaous taiasica, Jordan & Evermann, 1898:2236. Holotype: M. C. Z. 31220. No. Poey 471. MiCROGOBIUS Microgobius Poey, 1876:126 (Mierogobius signatus Poey) 172. Microgobius signatus Poey Microgobius signatus Poey, 1876:127, pi. 5, fig. 3; Jordan & Evermann, 1898:2246. Types: M. C. Z. 13127, 3 specimens, the largest of which is the holo- type, the other two paratypes. No. Poey 513. SiCYDIUM Sicydium Cuvier & Valenciennes, 1837b: 126 {Gobius plumieri Bloch) 173. Sicydium siragus Poey = Sicydium plumieri (Bloch) Sicydium siragus Foey, 1860:278; 1861:380; 1868:395; 1876:124. Gobius plumieri Bloch, 1786:125, pi. 178, fig. 3. Sicydium plumieri, Jordan & Evermann, 1898:2206. Cotypes: M. C. Z. 13328, 2 specimens 36 & 38 mm. long. No. Poey 574. Type locality: Santiago de Cuba. 216 bulletin: museum of comparative zoology GOBIOIDIDAE GOBIOIDES Gobioides Lacepede, 1800:580 {Gobioides broussonnetii Lacepede) 174. Gobioides barreto Poey = Gobioides broussonnetii Lacepede Gobioides barreto Poey, 1860:282; 1861:380; 1868:394; 1876:125. Gobioides broussonnetii, Lacepede, 1800:580; Poey, 1866a :335; Jordan & Evermann, 1898:2263. Holotype: M. C. Z. 13246. No. Poey 294. ECHENEIDAE ECHENEIS Echeneis Linnaeus, 1758:261 (Echeneis naucrates Linnaeus) 175. Echeneis verticalis Poey = Echeneis naucrates Linnaeus Echeneis verticalis Foey, 1860:253; 1861:376. Echeneis naucrates Linnaeus, 1758:261; Jordan & Evermann, 1898:2269. Leptecheneis naucrates, Poey, 1868:376; 1875b :90. Paratype: M. C. Z. 8709, the specimen he mentions as having 22 laminae in the disc. No. Poey 390. Rhombochirus Rhombochirus Gill, 1863a :88 (Echeneis osteochir Cuvier) 176. Echeneis tetrapturorum Poey = Rhombochirus osteochir (Cuvier) Echeneis tetrapturorum Poey, 1860:256, pi. 18, fig. 2; 1861:376. Rhombochirus tetrapturorum, Poey, 1868:377; 1876:89. Echeneis osteochir Cuvier, 1829:348. Rhombochirus osteochir, Jordan & Evermann, 1898: 2273. Holotype: M. C. Z. 8652. Paratypes: M. C. Z. 21805, 27228, 27229. No. Poey 130. HOWELL y RIVERO: TYPES OF POEY FISHES 217 OPISTHOGNATHIDAE Gnathypops Gnathypops Gill, 1862b :241 (Opisthognathus tnaxillosus Poey) 177. Opisthognathus macrops Poey = Gnathypops macrops (Poey) OpisiJwcimdhus macrops Poey, 1860:287; 1861:382. Gnathypops macrops, Poey, 1868:400; 1876:133; Jordan & Evermann, 1898:2284. Holotype: M. C. Z. 12514. No. Poey 485. LONCHOPISTHUS Lonchopisthus Gill, 1862b :241 {Opisthognathus micrognathus Poey) 178. Opisthognathus micrognathus Poey = Lonchopisthus micrognathus (Poey) Opisthognaihus micrognathus Poey, 1860:287; 1861: 382. Lonchopisthus micrognathus, Poey, 1868:400; 1876: 134; Jordan & Evermann, 1898:2287. Cotypes: M. C. Z. 12515, 12517, 2 specimens 90 & 125 mm. long. No. Poey 357. CLINIDAE Malacoctenus Malacoctenus Gill, 1860:103 {Clinus delalandi Cuvier & Valenciennes) 179. Myxodes macropus Poey = Malacoctenus macropus (Poey) Myxodes macropus Poey, 1868 :399 ; 1876 :131 . Malacoctenus macropus, Jordan & Evermann, 1898: 2357. Types: M. C. Z. 12511, 2 specimens, the larger of which is the holo- type. No. Poey 285. BLENNIIDAE Blennius Blennius Linnaeus, 1758:256 {Blennius galerita Linnaeus) 180. Blennius vinctus Poey 218 bulletin: museum of comparative zoology Blennius vindus Poey, 1867:243; 1868:397; 1876:129; Jordan & Evermann, 1898:2382. Holotype: M. C. Z. 12647. No. Poey 616. Entomacrodus Entomacrodus Gill, 1859c :168 {Entomacrodus nigricans Gill) 181. Salarias margaritaceus Poey = Entomacrodus margaritaceus (Poey) Salarias margaritaceus Poey, 1860:289; 1861:381; 1876:132. Entomacrodus margariiaceus, Poey, 1868:397; Jordan & Evermann, 1898:2398. Types: M. C. Z. 12513, 3 specimens, the largest of which is the holo- type. No. Poey 615. BROTULIDAE Stygicola Stygicola Gill, 1863b :252 (Lucifuga dentatus Poey) 182. Lucifuga dentatus Poey = Stygicola dentata (Poey) Lucifuga dentatus Poey, 1860:102, pi. 9, fig. 1, pi. 10, figs. 5-6, 9, pi. 11, figs. 6-8, 15, 17; 1865:113. Stygicola dentatus, Poey, 1868:401; 1876:137; Jordan & Evermann, 1898:2500. Holotype: M. C. Z. 32329. No. Poey 255. OPHIDIIDAE Otophidium Otophidium Gill, 1885:914 {Genypterus omostigma Jordan & Gilbert) 183. Ophidium graellsi Poey = Otophidium graellsi (Poey) Ophidium graellsi Foey, 1861:425, 1868:402; 1876:137. Ophidion graellsi, Jordan & Evermann, 1898:2488. Holotype: M. C. Z. 12440. No. Poey 480. HOWELL Y RIVERO: TYPES OF POEY FISHES 219 GOBIESOCIDAE SiCYASES Sicyases Miiller & Troschel, 1843:298 {Sicyases sanguineus) 184. Sicyases rubiginosus Poey Sicyases rubiginosus Poey, 1868:391; 1876:124. Gobiesox rubiginosus, Jordan & Evermann, 1898:2337. Holotype: M. C. Z. 12923. No. Poey 4. 185. Sicyases carneus Poey Sicyases carneus Poey, 1868:392; 1876:124. Gobiesox carneus, Jordan & Evermann, 1898:2337. Cotypes: M. C. Z. 12925, 8 specimens 16 to 24.5 mm.; no measure- ments were given in the original description. No. Poey 676. BALISTIDAE Xanthichthys Xanthichthys Kaup, in Richardson, 1856:313 (Balistes curassavicus Gmelin) 186. Balistes cicatricosus Poey = Xantichthys ringens (Linnaeus) Balistes cicatricosus Poey, 1861:327, 361; 1863:181; 1867:171. Xantichthys cicatricosus, Poey, 1868:435. Balistes ringens Linnaeus, 1758:329. Xantichthys ringens, Poey, 1876:164; Jordan & Ever- mann, 1898:1709. ' Holotype: M. C. Z. 11953. No. Poey 97. ANTENNARIDAE Antennarius Antennarius Lacepede, 1798:421 {Antennarius bivertex etc Commerson = Lophius commersonianus Lacepede) 187. Chironectes Tigris Poey = Antennarius Tigris (Poey) Chironectes tigris Poey, 1853:217, pi. 17, fig. 2; 1861 :382. 220 bulletin: museum of comparative zoology Aniennarius tigris, Poey, 1868:405; 1876:134; Jordan & Evermann, 1898:2723. Holotype: M. C. Z. 11611. Paratypes: M. C. Z. 11617, 11619, 11621, 3 specimens, 70 to 108 mm. No. Poey 207. 188. Antennarius corallinus Poey = Antennarius multiocellatus (Cuvier & Valenciennes) Antennarius corrallinus Poey, 1865:188; 1868:405; 1876:135. Chironectes imdtiocellatus, Cuvier & Valenkiennes, 1837a :422. Antennarius multiocellatus, Jordan & Evermann, 1898:2724. Holotype: M. C. Z. 11620. No. Poey 301. HOWELL Y RIVERO: TYPES OF POEY FISHES 221 BIBLIOGRAPHY Agassiz, Louis. 1829. In Spix, "Selecta genera et species piscium quos in itinere per Brasiliam annis 1817-1820 etc.," 1st. part. 1831. Idem, 2nd. part. Ahl, Jonas Nicholas. 1787. "De Muraena et Ophichtho." Artedi, Petri. 1793. "Sueci Descriptiones specierum piscium etc., Ichthyologiae, pars V. Bennett, F. D. 1840. "Narrative of Whaling Voyage around the Globe, etc.," 2. Bleeker, Pieter. 1852. "Diagnostiche beschrijving en van nieuve of weining bekende vis- schsoorten van Sumatra." Nat. Tijdschr. Neder-Indie, 3. 1854. "Bijdrage tot der Kennis der ichthyologische fauna van bet ieland Flores." Naturkundig. Tijdschr. Neder-Indie, 6. 1856. "Beschrijvingen van nieuve of wenig bekende visschsoorten van Menado en Makassar." Act. Soc. Sci. Indo-Neerl., 1. 1862a. "Sur quelques genres de la famille des Pleuronectoides." Versl. Akad. Amsterdam, 13. 1862b. "Notices Ichthyologiques." Comptes-rendus Acad. Royal Sci., Sec. Sci. exactes, Amsterdam, 14. 1877. "Memoire sur les Chromides marins ou Pomacentroldes de I'lnde archipelagique." Nat. Verb. HoU. Maatsch., Wetensch., 2. 1878. "Quatrieme memoire sur la faune ichthyologique de la NouveUe- Guinee." Arch. Neerl. Acad. Sci., 13. Bloch, Marc Elieser 1785. "Ichthyologie, ou Histoire naturelle, general et particuliere des poissons." 1786. "Ichthyologie, ou Histoire naturelle, general et particuliere des poissons." 1787. "Naturgeschichte der Auslandischen Fische." 3. 1790. Idem, 4. 1791. Idem, 5. 1792. Idem, 6. 1793. Idem, 8. 1795. Idem, 9. 1789. "Tu& Utlandskar Fiskar." Kongl. Vetenskaps Academiens Nya Handlingar, Stockh., 10. Bloch, Marc Elieser and Schneider, Johann Gottlob 1801. "Systema Ichthyologiae iconibus ex illustratum etc." 222 bulletin: museum of comparative zoology Bonaparte, Charles Lucien 1837. "Iconografia della Fauna Italica per le quattro classi degli animali vertebrati." Fasc. 91. Castelnau, Francois L. 1855. "Expedition dans les parties centrales de I'Amerique du Sud, etc. pendant les annees 1843 a 1847." Poissons. Cocco, Anastasio 1829. "Lettere al Signore Risso su alcuni pesci novelli." Giorn. Sci. Lett. Sicilia, 42. Cope, Edward Drinker 1870. "Contribution to the Ichthyology of the Lesser Antilles," Trans. Amer. Philos. See, 14. CuviER, George 1817. "Le Regne animal distribue etc.," Ed. 1, Poissons, 2 (The Latin forms supplied by Oken, 1817, in his Isis.) 1829. "Le Regne animal distribue etc.," Ed. 2, Poissons, 2. CuviER, George and Valenciennes, Achille 1828. "Histoire Naturelle des Poissons," 2. 1829a. Idem, 3. 1829b. Idem, 4. 1830a. Idem, 5. 1830b. Idem, 6. 1831a. Idem, 7. 1831b. Idem, 8. ■ 1833. Idem, 9. 1836. Idem, 11. 1837. Idem, 12. 1839a. Idem, 13. 1839b. Idem, 14. 1847. Idem, 20. Eigenmann, Carl H. and Eigenmann, Rosa Smith 1890. "Additions to the Fauna of San Diego." Proc. Calif. Acad. Sci., Ser. 2, 3. Forskal, Pehr 1775. "Descriptiones Animalium avium, amphiborium, piscium, etc." Fowler, Henry W. 1905. "New, rare, or little known Scombroids." Proc. Acad. Nat. Sci. Phila., 57. 1911. "Notes on Clupeoid Fishes." Proc. Acad. Nat. Sci. Phila., 63. Garman, Samuel 1896. "Cross Fertilization and Sexual rights and lefts among Verte- brates." Amer. Nat., 30. 1913. "The Plagiostomia." Mem. Mus. Comp. Zool., 36. HOWELL Y RIVERO : TYPES OF POEY FISHES 223 Gill, Theodore 1859a. "Description of Hyporhamphus, a new genus of fishes allied to Hemirhamphus Cuv." Proc. Acad. Nat. Sci. Phila., 11. 1859b. "Description of a third genus of Hemirhamphinae." Proc. Acad. Nat. Sci. Phila., 11. 1859c. "Description of a new genus of Salarianae from the West Indies." Proc. Acad. Nat. Sci. Phila., 11. 1860. "Monograph of the genus Labrosomus Sw." Proc. Acad. Nat. Sci. Phila., 12. 1861. "Catalogue of the fishes of the Eastern Coast of North America, from Greenland to Georgia." Supp. Proc. Acad. Nat. Sci. Phila., 13. 1862a. "Appendix to the Synopsis of the Subfamily of Percinae." Proc. Acad. Nat. Sci. Phila., 14. 1862b. "Remarks on the relations of the genera and other groups of Cuban fishes." Proc. Acad. Nat. Sci. Phila., 14. 1863a. "Catalogue of the fishes of Lower California collected by Mr. J. Xantus." Proc. Acad. Nat. Sci. Phila., 15. 1863b. "Descriptions of the genera of Gadoid and Brotuloid fishes of North America." Proc. Acad. Nat. Sci. Phila., 15. 1865. "On a new genus of Serraninae." Proc. Acad. Nat. Sci. Phila., 17. GiRARD, Charles 1858. "Notes upon various new genera and new species of fishes in the Museum of the Smithsonian Institution, etc." Proc. Acad. Nat. Sci. Phila.-, 10. 1859. "Ichthyological Notices." Proc. Acad. Nat. Sci. Phila., 11. Gmelin, Johann Frederick 1788. "Systema Naturae per regna tria naturae, etc.," Ed. 13. Gronow, Lorenz Theodor 1777. In Scopoli, "Introductio ad Historiam Naturalem." 1854. "Catalogue of fish collected and described by Laurence Theodore Gronow, now in the British Museum." Guichenot, a. 1853. In Ramon de la Sagra, "Historia Fisica, Politica y Natural de la Isla de Cuba," Pesces. GiJNTHER, Albert 1859. "Catalogue of the fishes in the British Museum," 1. 1862. Idem, 4. 1868. Idem, 7. HuBBS, Carl L. 1926. "Studies of the fishes of the Order Cyprinodontes." Misc. Pub. Univ. Mich. Mus., 16. 224 bulletin: museum of comparative zoology Jenyns, Leonard 1842. "The Zoology of the voyage of H. M. S. "Beagle," during the years 1832-1836." Fishes. Jordan, David Starr 1885. "A Catalogue of the fishes known to inhabit the waters of North America North of the Tropic of Cancer." Rept. U. S. Fish Comm., 13. 1886. "Revision of the Sciaenidae of Europe and America." Rept. U. S. Fish Comm., 14. Jordan, D. S. and Evermann, Barton W. 1895. "A Check List of the fishes and fish-like Vertebrates of North and Middle America." Rept. U. S. Fish Comm., 21. 1896. "The Fishes of North and Middle America." U. S. Nat. Mus. Bull. No. 47, 1. 1898. Idem, 2 and 3. 1927. "New Genera and species of North American fishes." Proc. Calif. Acad. Sci., 4th Ser., 16. Jordan, D. S., Evermann, B. W. and Clark, H. W. 1930. "Check List of the Fishes and Fish-like Vertebrates of North and Middle America." Rept. U. S. Fish Comm., Part 2. Jordan, D. S. and Hubbs, Carl L. 1919. "A Monographic Review of the Family of Atherinidae or Silver- sides." Stanf. Univ. Pub., Univ. Ser. Kaup, Johann Jacob 1856. "Catalogue of the Apodal fishes in the Collection of the British Museum." Lacepede, Bernard Germain 1798. "Histoire Naturelle des Poissons," 1. 1800. Idem, 2. 1802. Idem, 3. 1803a. Idem, 4. 1803b. Idem, 5. Lichtenstein, Martin Heinrich Carl 1821. "Die Werke von Marcgrave und Piso uber die Naturgeschichte Brasiliensis, erlautert aus den wider aufgefundenen Original- zeichnungen." Abhandl Berlin Acad. Wiss. Linnaeus, Carl 1758. "Systema Naturae sive regna tria naturae, etc.," Ed. 10. 1766. Idem, Ed. 12. Lowe, Richard Thomas 1838. "Piscium Maderensium species quaedam novae vel minus rite cognitae breviter descriptae, etc." Trans. Cambridge Phil. Soc, 6. HOWELL Y RIVERO: TYPES OF POEY FISHES 225 LtJTKEN, Christian Frederik 1851. "Nogle bemaerj kinder om naeseborenes stiling hds de i gruppe med Ophisurus staaende slaegter af aalefamilien," Videusk Meddel. Naturhist. Foren. Kjobenhavn. MtJLLER, Johannes and Troschel, Franz H. 1843. "Beitrage zur Kenntniss der natiirlichen Familien der Fische." Wiegmann's Archiv. Naturgesch. 1847. In Schombuigk, "The History of Barbados, etc." Myers, George S. 1927. "An Analysis of the Genera of Neotropical KilHfishes alhed to Rivulus." Ann. Mag. Nat. Hist., Ser. 9, 19. Norman, J. R. 1934. "Monograph of the Flatfishes (Heterosomata)." Oken, Lorenz 1817. "Les Congress Cuvier," in the Isis. Osbeck, Pehr 1765. "Iter Chinensis 1757; reprinted as "Reise nach Ostindien und China, etc." Deutsche Uebersetzung von J. G. Georgius Rostock. Parr, Albert Eide 1928. "Deep-sea fishes of the Order Iniomi from the waters around the Bahama and Bermuda Islands." Bull. Bingham Ocean. Coll., 3, Art. 3. Poey, Felipe 1851. "Memorias sobre la Historia Natural de la Isle de Cuba," 1, pp. 1^0, pis. 1-8. 1852. Idem, pp. 40-200, pis. 9-22. 1853. Idem, pp. 201-280, pis. 23-30. 1854. Idem, pp. 281^53, pis. 31-34. 1858. "Memorias sobre la Historia Natural de la Isla de Cuba," 2, pp. 1-96, pis. 1-9. 1860. Idem, pp. 97-336, pis. 10-12, 14. 1861. Idem, pp. 337^28, pis. 13, 15-19. 1863. "Descriptions des Poissons nouvelles ou peu connues." Proc Acad. Nat. Sci. Phila., 15, pp. 180-188. 1865. "Repertorio Fisico-Natural de la Isla de Cuba," 1, pp. 1-278. 1866a. Idem, pp. 279-420. 1866b. "Repertorio Fisico-Natural de la Isla de Cuba," 2, pp. 1-48. 1867. Idem, pp. 49-276. 1868. Idem, pp. 276-484. (The Synopsis Piscium Cubensium forms part of Vol. 2 of the Repertorio, pp. 279-484.) 1870a. "Review of the fish of Cuba belonging to the Genus Trisotropis, with an introductory note by J. Carson Brevoort." Ann. Lye. Nat. Hist. N. Y., 9, pp. 301-310. 226 bulletin: museum of comparative zoology 1870b. "New species of Cuban Fish." Ann. Lye. Nat. Hist. N. Y., 9, pp. 317-322. 1871. "Genres des Poissons de la Faune de Cuba appartenant a la Famille Percidae." Ann. Lye. Nat. Hist. N. Y., 10, pp. 27-79, pl.l. 1872. "Monographie des Poissons de Cuba compris dans la sous-famille des Sparini." Ann. Lye. Nat. Hist. N.Y., 10, pp. 170-184, pis. 6-7. 1875a. "Poissons de I'ile de Cuba." Ann. Lye. Nat. Hist. N. Y., 11, pp. 58-70, pis. 7-10. 1875b. "Enumeratio Piscium Cubensium." Ann. Soc. Esp. His. Nat., 4, pp. 75-162 (1-88), pis. 5-8. 1876. Idem, 5, pp. 131-218 (89-176), pp. 373-404 (177-208), pis. 7-10, 13-14. 1877. Idem, 6, pp. 139-154 (209-224). (This work has two numbers on each page, one corresponding to the vol., the other continuous for the work itself.) 1880. "Revisio Piscium Cubensium." Ann. Soc. Esp. Hist. Nat., 9, pp. 243-261 (1-19), pis. 7-10. (This work also has two numbers on each page.) Putnam, Frederick Ward 1863. "List of fishes sent by the Museum to different institutions in ex- change for other specimens, with annotations," Bull. Mus. Comp. Zool., 1. QuoY, Jean R. C. and Gaimard, Paul 1824. "Voyage autour du monde execute sur la corvette de S. M. L'Uranie etc." Zool., 1. Rafinesque, Samuel 1810a. "Caratteri di alcuni nuovi Generi e nuove specie di Animale e Piante della Sicilia." 1810b. "Indice d'lttiologia Siciliana." Ranzani, Camillo 1840. "De novis speciebus piscium; " Acad. Sci. Inst. Bonon. 3. 1842. "De novis speciebus piscium." Nov. Comm. Acad. Sci. Bonon., 5, Richardson, John 1844a. "The Zoology of the Voyage of H. M. S. Erebus and Terror during 1839-43." Fishes. 1844b. "Zoology of the voyage of H. M. S. Sulphur, etc." Ichthyology. 1856. "Ichthyology," in Encyclopaedia Britannica, Ed. 12. RiVERO, Luis Howell 1934. "Some new and rare Cuban eels." Mem. Soc. Cub. Hist. Nat., 8, No. 6. Shaw, George 1803. "General Zoology or systematic natural History, etc.," 4. HOWELL Y RIVERO: TYPES OF POEY FISHES 227 SwAiNSON, William 1839. "Natural History and classification of fishes, amphibians and reptiles, or monocardian animals," 2. Troschel, Franz H. 1865. In Miiller's "Reisen in den Vereinigten Staaten, Canada und Mexico," 3. Van Hasselt, J. K. 1824. "Notice anatomique sur quelques poissons." Bull. Sci. Nat. (Ferussac), 2. Walbaum, J. J. 1792. Petri Artedi, "Bibliotheca et philosophia ichthyologica . . . " Piscium, 3. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. LXXXII, No. 4 . CONTRIBUTION TO THE KNOWLEDGE OF THE GENUS SMIN TH U RIDES BORNER By J. W. FoLsoM and H. B. Mills With Nine Plates CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM September, 19.38 . . Zoology ''^, SEP 14 lyjo No. 4. — Contribution, to the Knowledge of the Genus Smintkurides Bonier By J. W. FoLSOM AND H. B. Mills INTRODUCTION Justus Watson Folsom (September 2, 1871 — September 24, 1936) The bronze grasshopper which served as a knocker on the door of Dr. Samuel Scudder's laboratory in Cambridge, Massachusetts, was agitated by the hand of one of the youngsters of the neighborhood. The former student and assistant of Louis Agassiz arose, opened the door, and invited the young enthusiast with his box of butterflies across the threshold. Thus began the entomological career of Dr. Justus Watson Folsom, and thus are linked the names of three of America's most brilliant and colorful zoologists, Agassiz, Scudder and Folsom. The entomological career which had its beginning that day in Cam- bridge ended in Vicksburg, Mississippi, on September 24, 1936, with Dr. Folsom's death. Between these two dates are packed endeavors, accomplishments and contributions which set a high standard among the entomologists of the world. Very early in his training Dr. Folsom's interests began to revolve about that still poorly understood suborder of insects, the Apterygota, and this interest continued unabated until his death. His entomologi- cal textbook, especially the third edition, was a departure from the usual pattern of texts and far ahead of its time. It was the first Amer- ican text to deal in any measure with the important fields of Insect Ecology and Physiology, and is a truly valuable inclusion in the ento- mologist's library. Dr. Folsom was born in Cambridge, Massachusetts, September 2, 1871. Upon the death of his parents the responsibility for his training and education was shouldered by Mrs. Josephine Seymour, the mother of one of his friends. He continued to make his home with her and she with him as long as he lived. After preliminary schooling he entered Harvard College, obtaining an S.B. in 1895 and a Sc.D. in 1899 under the direction of Dr. E. L. Mark. Then followed a year as Professor of Natural Sciences at Antioch College. In 1900 he received an appoint- ment to the Department of Entomology of the University of Illinois 232 bulletin: museum of comparative zoology remaining at that institution until 1923. In 1925 he entered the Bureau of Entomology of the United States Department of Agriculture, remaining in its service until the time of his death. Dr. Folsom was an extremely careful worker, and if the definition of a genius is "one who has an infinite capacity for taking pains" he must be so classed. While working on the collembolan genus Orches- ella in the summer of 1930 he spent several days mounting and exam- ining many hundreds of specimens of OrchcscUa hcxfasciata (Harvey), a very definite and clear-cut species and one with which he had long been familiar. "The microscopic examination of these insects is hard physical labor," he told the writer at one time, "But I am not satisfied until I have seen all that there is to see." He was constantly in search of better methods and new techniques for the examination and preser- vation of minute insects, and most unselfish with the information acquired. His care in giving credit to those who assisted him amounted to an obsession. He was extremely unselfish. When the writer first worked with him he pointed to a filing cabinet one day and said, "That file is full of new species which I have accumulated. Help yourself to it; work them up and describe them." That is a spirit not often demon- strated in the field of taxonomy. The following paper is based on notes and sketches upon which Dr. Folsom was working at the time of his death. Harlow B. Mills The genus Sminthuridcs was proposed by Borner (1900, p. 616) to include the species violaceus Renter, aquaticus Bourlet, malmgreni Tullberg, ■pcnicillifer Schiiifer, signatus Krausbauer, j^arvuhis Kraus- bauer, and assimilis Krausbauer, which previously had been included in the genus Sminthurus Latreille. The subgeneric description was as follows : "Tibiotarsal organ present, antennal segment IV of the male developed into a clasping organ as in many copepods. In the species known up to now the under claw of the third pair is differ- ent from the other two. The upper claw is more slender and usually longer on the first two pairs than on the third. The dorsal inner edge of the mucro differs from the outer edge ; the inner edge toothed. Mucronal bristle absent." This description was corrected later (1901, p. 6), the clasping organ of the male not limited to the fourth antennal segment and the mu- cronal bristle noted as present. FOLSOM AND MILLS: SMINTHURIDES BORNER 233 The same author (1901, p. 91) raised the group to generic standing, and later (1906, p. 181) indicated Smi/nthurus aquaticus Bourlet as the type. We follow Linnaniemi (1912, p. 247) in the use of the name Smin- thurides Borner in place of Prosminthurus Willem (1900, p. 55). Both of these generic names were proposed in 1900, with S. aauaticiis Bourl. as the t\'pe (indicated later by Borner). Borner's name appeared in December, 1900, and Willem's is undated except for the year. It is very possible that Prosminthurus has time priority, and it certainly has page priority over Sminthurides. The literature for the past thirty years has almost exclusively employed the latter name, following Borner's error (1901, p. 91) of postdating the appearance of Prosmin- thurus to 1901. Until this matter can be settled definitely, it seems best to conform to usage and use the name Sminthurides. Linnaniemi (1912, p. 247) observed but one pair of bothriotricha on the anogenital segment and suggested an emendation of the descrip- tion to that effect. In all of the species which we have examined thus far two pairs were seen. The posterior pair is, however, subdorsal and usually much shorter and more bristle-like. On either side of the furcal segment the bases of these sensory hairs form an oblique, nearly straight line in all of the subgenera but Denisiella, where the bases are rather close together and form the points of a triangle. There is a rather definite progression from Sminthurides s. str. through Stenacidia to Denisiella. The position of Sphaeridia, on the other hand, is somewhat anomolous. In the absence of a tibiotarsal organ and in the shape of the tenaculum it resembles Denisiella. The position of the bothriotricha of the furcal segment connect it with Sminthurides, while the simplification of the clasping antennae of the males isolate it from the other subgenera. Our present knowledge of the group leaves Sphaeridia's position debatable. BIOLOGY The species of this genus are predominantly water surface and humus inhabitants, although occasionally they have been taken from beneath bark and upon grass. Some species are beautifully adapted for life on water surfaces. xVquatic forms furnish food for small fishes. The biology of Sminthurides (Sminthurides) aquaticus Bourlet has been studied by several workers, Renter (1883), Levander (1894), 234 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Strebel (1927), and especially Falkenhan (1932). Although this species may not be typical of all of the others of the genus it is better known, and a summary of its biolog}' is given here. S. aquaticus feeds almost exclusively on soft dead and living plant material, unicellular algae, fungus spores, etc. The ventral tube is not an adhesive organ, probably not functioning as such in any Collembola, but functions exclusively for respiration. One of the conspicuous phases of copulation is the grasping of the female antennae by the male antennae, the male usually being carried about for several days completely off the water surface. Transmission of sperms takes place at the beginning of the copulation process. Females are capable of copulation before they are fully grown. Par- thenogenesis does not occur. Oviposition follows copulation ordinarily by fourteen to eighteen days, the time depending on the temperature and the copulation age of the female, and occurs in moist soil surrounding water surfaces. A female may lay as many as sixty-six eggs. Temperature largely controls the length of the egg stage, which varies from three and one- half days to five months. S. aquaticus can remain submerged for as long as four days. If it is not hindered from so doing, it usually succeeds in getting back to the water surface in from one to two days, where it shows no ill effects from the submersion. All of the eggs from one female develop either into males or females, and the number of male- and female-producing mothers is about equal. Only six per cent of the females observed by Falkenhan (1932) pro- duced broods of mixed sexes. Males moult three times and females between five and seven times, becoming sexually matiu-e after the third moult. Males usually live for four or five weeks and the females from two to three and one-half months. The sex ratio, which is originally about 1 :1 changes later in favor of the females due to their longer life. Males and females are found together throughout their active period, and from seven to ten generations occur each season. The winter is passed nor- mally in the egg stage, but in southern latitudes, or in exceptionally warm winters in northern regions, a few adult females may hibernate. The chief enemy of S. aquaticus in Europe is the mite Episcius sphagni (Halbert), which is frequent about the edges and on the sur- faces of small pools which the Collembolans inhabit. FOLSOM AND MILLS: SMINTHURIDES BORNER 235 Genus Sminthurides Borner 1900 Vesicles of ventral tube short, smooth, without lateral tubercles but sometimes with apical papillae. Segmentation of the body usually evident through intersegmental sutures or light lines. Corpus of ten- aculum with lateral clavate processes at the bases of the rami. Integu- ment granulate. Anal segment ankylosed with the genital segment, which is broadly united to the furcal segment. Body with five pairs of bothriotricha, three pairs on the furcal segment and two on the ano- genital division. Anal appendages of female wanting. Second and third segments of the antennae of the male modified into a clasping organ, the fourth segment always simple. Antennae of female not modified, the fourth segment simple, ringed or definitely subsegmented. Tibiotarsal organ of posterior legs present or absent; when present, composed of two tooth-like eversions and a heavy spine which may be lamellate, notched, or definitely bifid. Except for the subgenus Ben- isiclla, the first two pairs of ungues are longer, more slender, and usually definitely unidentate on the inner margin, differing from the posterior ungues which are shorter, broader, and usually weakly toothed or un- toothed; first two pairs of unguiculi usually short and narrow, the third pair much broader and definitely lamellate ; subapical unguicular filaments present, tenent hairs absent. Eyes normally eight on either side. Length seldom over 1 mm., usually much less. Key to the Subgenera of Sminthurides Borner 1. Tibiotarsal organ of 3rd pair of legs (figure 5) present, composed of 2 sacs and an enlarged spine 2 Such tibiotarsal organ absent 3 2. Mucronal edges with weak, or without true lamellae, dorsal inner edge toothed. Mucro slender, definitely narrowed in the apical third. Ant. IV simple (figure 89) Stenacidia Borner. P. 262 Mucronal edges more or less broadly lamellate, inner lamella toothed and ribbed (figure 12). Ant. IV simple, ringed, or subsegmented. . . . Sminthurides Borner. P. 235 3. Mucronal bristle absent (figure 106) Sphaeridia Linnaniemi. P. 268 Mucronal bristle present (figure 100) Denisiella n. subg. P. 264 Subgenus Sminthurides Borner s. str. 1843, Sminthurus Bourlet p. 58 {ad. partitn). 1900, Smithurides Borner p. 616 {ut subgenus). 1900, Prosminthurus Willem p. 55. 1901, Sminthurides Borner p. 91. 236 bulletin: museum of comparative zoology Tibiotarsal organ present. Muerones with 3 well developed lamellae, the inner always toothed; mueronal bristle present. Antennae usually longer than the head, the 4th segment simple, ringed, or definitely sub- segmented. Corpus of tenaculum usually subconical, exceeding the rami of the tenaculum, and with anterior and apical bristles. Vesicles of ventral tube simple or with as many as 6 apical tubercles. Anogen- ital segment broadly united with the rest of the body, the 5th and 6th segments sometimes separated dorsally by a constriction. Type: Smiiifhurides aquaticus (Bourlet). Key to the species of the subgenus Sminthurides Borner s. str. 1. Mucro apically bulbous or apparently so, with coarse teeth on inner lamella (figure 83) 2 Mucro not apically bulbous. Inner lamella with finer teeth (figure 32) . . 4 2. Ventral mueronal lamella ending in a sharp tooth before the apex (figure 83) lepus Mills 1934 Ventral mueronal lamella entire (figure 78) 3 3. Ungues stout, inner tooth beyond the middle. Mucro not hooked apically. Basal segment of fourth antennal segment related to apical about as 2:1. (figure 78) occultus Mills 1934 Ungues slender, the inner tooth situated before the middle. Mucro hooked apically. Basal segment related to the apical about as 2.2-2.5:1 (figure 76) macnamarai n. sp. 4. Mucro trough-shaped, both sides alike (figure 88) .... plicatus Schott 1891 Mucro not as above, the sides dissimilar 5 5. Filament of hind unguiculus of female branched (figure 45) 6 Filament of hind unguiculus of female simple (figure 57) 9 6. Filament 2- or 3-branched. Fourth antennal segment of female ringed (figure 45) bifidus Mills 1934 Filament of hind unguiculus 4- to 6-branched 7 7. Seta of organ of third tibiotarsus simple. Hind unguicular filament 4-branched (figure 40) appendicidatus Imms 1912 Seta of organ of third tibiotarsus wide, bifid. Unguicular filament 5- or 6-branched 8 8. Fourth antennal segment not subsegmented. Apex of bristle of tibiotarsus III organ not attaining the base of the unguiculus. Unguicular fila- ment of the first 2 pairs of legs simple, sharp (figure 49) penicillifer Schaffer 1896 Fourth antennal segment subsegmented with 7 (?) irregular divisions. Apex of bristle of tibiotarsus III organ passing the base of the un- guiculus. Unguicular filament of first 2 pairs of legs flat, lanceolate in its distal portion pauliani Denis 1936 FOLSOM AND MILLS: SMINTHURIDES BORNER 237 9. Antennae shorter than the head (figure 1) 10 Antennae equal to or longer than the head 11 10. Mucro with 10-12 inner teeth. Vesicles of ventral tube with several ter- minal papillae cruciatus Axelson 1905 Mucro with about 9 inner teeth, apically pointed. Vesicles ? Clothing not remarkably short hospes Borner 1907 11. Fore unguiculi very long, lanceolate, appressed, apex exceeding apex of unguis (figure 39) spegazzinii Borner 1907 Fore unguiculi not as above 12 12. Fourth antennal segment of female simple 13 Fourth antennal segment of female distinctly subsegmented 17 13. Organ of 3rd tibiotarsus of male with a bifid seta, 2 elongate swellings, and a lateral blunt club (figure 3, 4) stagnalis Womersley 1932 Organ of 3rd tibiotarsus usually with a simple seta. Accessory club absent in male 14 14. Mucro bearing a subapical fingerlike projection. Fourth antennal segment of female with a basal subglobose portion, separated from the apical part by a constriction, apical part gradually and rather evenly nar- rowing to apex (figure 6) globocerus n. sp Fourth antennal segment of female without a conspicuous basal globose region or finger-like projection of mucro 15 15. Mucro at least half as broad as long (figure 12). Seta of 3rd tibiotarsus organ usually extending far beyond the apex of the tibia, not lamel- late. Dorsal segmentation of body evident, .aquaticus Bourlet 1843 Mucro one-third as broad as long (figure 30) . Seta of 3rd til>iotarsal organ not greatly exceeding the apex of tibia, lamellate. Dorsal segmenta- tion of body not evident 16 16. Abdominal segments 5 and 6 demarcated dorsally. Mucro apically rounded (figures 26, 27). Ventral tube vesicles with 6 papillae. Seta of tibiotarsal organ broad, bifid ludovicianus n. sp. Abdominal segments 5 and 6 confluent. Mucrones apically pointed (figure 30). Vesicles of ventral tube simple. Seta of tilnotarsal organ tapering, normally simple lamellate malmgreni Tullberg 1876 17. Fourth antennal segment joints 7 and 8 weak ringlike subsegments, with narrow ringlike divisions between them (figure 41) melanolus Borner 1907 Fourth antennal segments definitely subsegmented usually with 4 or 5 subsegments 18 18. Subsegments of 4th antennal segment of female 7 or 8, none greatly longer than the rest (figure 70) annulicornis Axelson 1905 Antennae with 4 or 5 subsegments in the 4th segment (figure 73) 19 19. Basal subsegment subequal to the apical (figure 52) 20 Basal subsegment about twice the length of the apical (figure 79) 21 238 bulletin: museum of comparative zoology 20. Fourth antennal segment of female with 4 subsegments. Mucro strongly narrowed apically and rounded schotti Axelson 1903 Fourth antennal segment with 5 subsegments. Mucro not strongly nar- rowed apically, truncate parvulus Krausbauer 1898 21. Fourth antennal segment of female with 4 subsegments 22 Fourth antennal segment with 5 subsegments. . . .inequalis Borner 1903 22. Mucro strongly narrowed toward the apex. Length of females 0.5 mm. Body light, with dark blue stripe on the sides and a dorsal, broken colored area (figure 58) assimilis Krausbauer 1898 Mucro slightly narrowed distally. Length of females 0.35-0.45 mm. Color lighter or darker violet, pigment diffuse (figure 64) krausbaueri n. nom. Sminthurides cruciatus Axelson (Linnaniemi) Plate I, fig. 1 1905, Sminthurides cruciatus Axelson, p. 792. Color of body largely dark violet. Body with a median-dorsal dark stripe with transverse branches. Sides of abdomen mostly pale, or ventrolaterally dark. On the sides, especially posteriorly, are several roundish dark spots. Anogenital segment almost entirely violet. Antennae, legs, and furcula pale violet. Head pale, marked with violet. Oral region dark. Eyes 8 on either side. Antennae clearly shorter than the head (figure 1), as 6:7, the segments about as 7:8:13:23. Last antennal segment simple, unringed. First 2 pairs of ungues slender with an inner tooth at about the middle and small lateral teeth. Un- guiculi of these feet short, slender, with small lamellae, scarcely half the length of the unguis; filaments exceeding the apex of the unguis. Third pair of ungues shorter than the others, with small inner tooth and lateral teeth. Unguiculi of these feet 73 the unguis, with broad lamel- lae and filaments which exceed the ungues. Organ of 3rd tibiotarsus composed of 2 elongate swellings and a short, simple, basally broad seta which extends a little beyond the tibiotarsus. Ventral tube with several papillae at the end of the vesicles. Tenaculum with the corpus conical and bearing 2 strong anterior setae ; rami 3-toothed, each with a basal clavate appendage. Dens 2.3 times the mucro. Mucrones bent apically, with relatively narrow lamellae; about Ys as broad as long; inner lamella with 10-12 teeth, narrowing toward the apex and ending in a sharp tooth-like projection, outer lamella ribbed but without teeth, ventral lamella narrow but evident. Mucronal seta present. Clothing FOLSOM AND MILLS: SMINTHURIDES BORNER 239 exceptionally sparse and short. Integument finely granulate. Length up to 0.66 mm. Sweden, Finland. This species occurs on water surfaces with such species asS. malmgreni elegantus Reut. and S. aquaticus Bourl., sometimes among water plants. Sminthurides hospes Borner Plate I, fig. 2 1907, Sminthurides hospes Borner, p. 172. Weakly pigmented with violet, darker on the posterior dorsum. Dentes and mucrones pale, claws not plainly pigmented. Antennae violet, becoming darker distally. Eyes 8 on either side. Head only a little longer than the antennae, the segments of which are related about as l:P/5:2 ^5:4, or 1:1^2:2^4:4^4- Last antennal segment simple, not subsegmented. Body very highly arched. Ungues apparently without inner or lateral teeth, the first 2 pairs somewhat longer and more slender than the last (figure 2). Unguiculi on the first 2 pairs of feet with a concave inner lamella and subapical filament which attains the apex of the unguis (exceeds it in Borner 's figure 40). Ungues and unguiculi of 3rd feet almost as in aquaticus but narrower, the fila- ment exceeding the apex of the unguis. Tibiotarsal organ with the bristle bearing a short ventral accessory branch, scarcely reaching the base of the unguiculus. Anterior lobe of the tenaculum extending as far as the rami, apparently with only 1 seta. Dens to the mucro as 3 :1, furcula short. Dens dorsally and toward the apex in some degree clearly granulate. Mucrones more slender than in spegazzinii and mclanotus, recalling signatus Krausbauer. About 9 teeth on the inner lamella, pointed. Clothing relatively thick and long. Length up to 0.5 mm. La Plata, Argentina. Taken from water surface among water plants. Sminthurides stagnalis Womersley Plate I, figs. 3-4 1932, Sminthurides stagnalis Womersley, p. 16. Light brownish violet, with a median dark stripe. Eyes 6 (?) on either side. Antennae a little longer than the head, as 6 :5 in the male. Male with the usual well developed clasping organ. Ungues similar on 240 bulletin: museum of comparative zoology all feet, with no inner teeth. Unguieuli with moderately broad inner lamellae, and subapical filaments. Tibiotarsal organ with the seta short, broad, bifid. On the inner side of each posterior tibia of the male is a short strong spine above which is a longer thumb-like projection (figure 3). Mucro 0.4 the dentes, the inner lamella with strongly ribbed teeth (12 in Womersley's figure 3c), of the aquaticus type, with a simple pointed apex. In the male on the anogenital segment are 2 long pro- tuberances, each with 2 or 3 subapical spines (figure 4); just behind these is a short peglike projection. Clothing sparse, of long fine hairs. Length male 0.57 mm., female 0.69 mm. Collected on surface of stagnant pool, Denmark, West Australia. Sminthurides globocerus spec. nov. Plate I, figs. 5-7 Female. Ground color in light specimens pale yellow; pigment dark purple. Dorsum posteriorly purple, anteriorly pale yellow and usually crossed by 3 or 4 irregular, subparallel, purple bands which are broken on the midline; laterally purple. Head yellow dorsally, purple about the oral region. First antennal segment pale yellow with an apical mark; 2nd yellow with an apical ring; 3rd violet, pale basally; 4th entirely violet. Legs tinged with pale violet. Furcula unpigmented. Dark forms have the head and body blackish purple, sometimes almost slate black, with irregular lighter spots vaguely showing through. The venter is somewhat lighter especially at the insertion of the fur- cula, and the appendages darkly, diffusely purple. Eyes probably 8 on either side. Antennae definitely longer than the head, the segments about as 10:12:28:53 or 12:13:29:59. Fourth segment simple, a sub- basal constriction cuts off a globose basal region which does not, how- ever, represent a subsegment (figure 6). Beyond this constriction the segment gradually tapers to a rather acute apex. First 2 pairs of' ungues rather slender, with a weak inner tooth beyond the middle and 2 lateral teeth on each side. Unguieuli narrow, rather wider basally, with subapical filaments exceeding the ungues. Ungues of the hind legs (figure 8) without the inner tooth, broader. Unguieuli broadly lanceolate with definite, well developed lamellae and filaments which exceed the ungues. Seta of tibiotarsal organ heavy^ lamellate, bifid, reaching slightly beyond the apex of the tibiotarsus; the 2 basal sacs large, elongate, overhanging their insertions (figure 5). Dentes 2.2 times the mucrones, with a small inner thumblike projection. Mucro FOLSOM AND MILLS: SMINTHURIDES BORNER 241 with a prominent blunt projection dorsally at tlie apex. Inner dorsal lamella with 10-15 teeth, outer lamella usually with a small sub-basal tooth which is difficult to see, ventral lamella entire; mucro 3 or more times as long as broad; mucronal bristle present (figure 7). Dentes with numerous curved hairs dorsally and 3 longer suberect dorsal setae, the clothing recalling somewhat the condition in Deriisiella. Ventral bristle formula 1,1,1,1,2,2,2,2, the 2 single basal hairs minute. Body anteriorly with short moderately heavy hairs, posteriorly with longer, more numerous, backward-curving setae. Bothriotricha of the body 3 on either side, nearly in a straight line. Anogenital segment with 2 on either side. Tenaculum with the corpus subconical, bearing 1 apical and 2 anterior bristles. Rami 3-toothed, with basal clavate appendages. Fifth and 6th abdominal segments clearly separated dorsally. Length 0.43. Male. Coloration similar to that of the female, anterior dorsum sometimes without the dark crossbands, head mostly purple dorsally and orally. Clasping organ highly developed. Metanotum with a pair of subdorsal vesicles. Inner dorsal lamella of mucro with 10 or 11 teeth. Length 0.41 mm. North Carolina. Asheville, February 9, 1935. A. P. Jacot. Several specimens of both sexes from Andropogon sod. Sminthurides aquaticus (Bourlet) Plate I, figs. 9-13; Plate II, figs. 14-19 1843, Smynthurus aquaticus Bourlet p. 58. 1883, Sminthurus apicalis Reuter p. 20. 1896, Smynthurus amicus Folsom p. 446. 1900, Prosminthurus aquati- cus Willem p. 6. 1900, Sminthurus (Sminthurides) aquaticus Burner p. 616. 1901, Sminthurides aquaticus Borner p. 96. This widespread and variable species has been separated into the following forms : Var. AQUATICUS f.p. Yellow or brownish yellow; first 2 antennal segments pale brown; 3rd and 4th purple; legs pale brownish. This principal form of the species occurs in most parts of Europe but has not been recorded from North America. 242 bulletin: museum of comparative zoology var. viridulus (Renter) 1891, Sminthurus apicalis var. viridulus Reuter p. 231. 1893, Sminthurus aquaticus var. viridulus Schott p. 37. Sminthurides aquaticus var. viridula Borner p. 98. Greenish; often with a narrow dark median dorsal line. Antennae and legs tinged with purple. This European variety has not been recorded from North America. Linnaniemi (1912, p. 260) found it abundant in sphagnum moss. var. levanderi (Reuter) 1891, Sminthurus apicalis var. levanderi Reuter p. 232. 1893, Sminthurus aquaticus var. levanderi Schott p. 37. 1901, Sminthurides aquaticus var. levanderi Borner p. 98. Female. General color from rose pink to deep rose purple or violet; sides of the abdomen often olivaceous, with pale spots as in fig. 19. Sternum pale. Vertex yellow or whitish, with a wide median purple mark. Oral region and apices of legs blackish purple. Antennae mostly purple to violet; 1st and 2nd segments pale yellow or whitish. Legs pale purple with femora and tibiotarsi often violet. Manubrium and dentes dilute purple, dentes darker proximally and distally. Claws and mucrones pigmented. Antero-dorsum typically with strong seg- mented folds separated by pale intersegmental lines; the folds may, however, be weak or absent. Head with a dorsal ridge behind the vertex (figure 19). Fifth and 6th abdominal segments are not sep- arated. Eyes 8 on either side, 2 in each group being smaller than the others (figure 9). Antennae slightly longer than the head, as 1.2 :1 ; the 4th segment simple; the segments related as 10:14:25:51 (specimen from England) or 6:8:21:32 (specimen from Finland). First and 2nd ungues typically very slender but often comparatively stout, with an inner tooth (sometimes obscure) beyond the middle and 2 pairs of lateral teeth (figures 14, 15, 17, 18). First and 2nd unguiculi also slender with a subapical filament of variable length. Third ungues shorter than the 1st and 2nd, the inner tooth lacking (figures 13, 16). Third unguiculi subovate with a feebly subapical filament exceeding the ungues. Tibiotarsal organ with a strong bristle which extends far beyond the apex of the tibiotarsus (figure 11). Vesicles of ventral tube simple. Rami of tenaculum tridentate, with basal clavate appendages. Corpus with 2 long anterior setae and 2 smaller ones on the apex (figure 10). Dentes from 2.3 to 3 times as long as the mucrones. Mucrones convergent, spoonlike in general form, elliptical from above, FOLSOM AND MILLS: SMINTHURIDES BORNER 243 more than half as long as broad (as 32 :51) ; inner dorsal lamella with as many as 17 teeth; mucronal seta present (figure 12). Clothing of moderate length. Posterior dorsum with stiff setae. Often the setae of head, body, antennae, and legs situated each on a black spot. There are 5 pairs of bothriotricha as usual. The first (most anterior) is on the 2nd abdominal segment, the 2nd apparently on the 3rd, the 3rd probably on the 4th ; the 4th and 5th are apparently on the 5th abdom- inal segment. Integument minutely granulate. Length 1.0 mm. Male. In coloration like the female. Antennae modified for clasp- ing. Metanotum with a pair of subdorsal bladderlike eversible hyaline vesicles. Maximum length 0.5 mm. The foreign material of aquaticus which has been examined consists of numerous specimens from Germany (C. Schaft'er), Poland (J. Stach), Finland (W. Linnaniemi), England and Iceland (W. M. Davies). In all of this material the 1st 2 pairs of ungues are very slender, although varying somewhat in this respect. In the United States this typical form with slender ungues is present and widespread, though we record it as yet only from Massachusetts, North Carolina, Louisiana, and Utah. The prevailing form in North America is one with compara- tively stout ungues. Both forms may occur in the same locality. The unguicular filament varies greatly in relative length in European ma- terial. It may extend not quite so far as the inner tooth of the unguis or it may exceed the unguis, but is commonly short on the 1st pair. In North American material with the typical slender ungues the filament is short on the anterior feet ; it exceeds the unguis in the specimens with stouter ungues. The 2 vesicles of the tibiotarsal organ vary in size and form; sometimes they are slender. Both European and American examples have on all of the ungues 2 pairs of lateral teeth which have not been mentioned in previous de- scriptions. This species is common on the surface of the water of ponds and streams and on various aquatic plants. It is found also on adjacent damp humus, in which the eggs are laid. Occasionally it is encountered in moss. It is not limited to fresh water but occurs also on pools of salt water. Some of the individuals examined had desmids in the alimentary tract.