Na tha v by PS Be Writ bate psmsiehtrit we f PAF TAO Ae, SF ht Ft ABA Py resierrs: Chath en AL CAT enere erry at RAL we 3-1: ate Phe Himile MORN Beh 6S g Weel riicaaeeiatne NpF eetleD- KC edasaRe Ta fich Hi Fad Melt heh 4 A hake Fake rete beKeS AF oUt eet eH af tbe Ss NaN MeN LaF oe PMN CLA A howl & rep barra MEF BETA teh MO TE ea et oe tt ASUS Vahey . in ae ato ake ey eA MTR RT ORF LAM Fate POEM ee ee ee 2 car ood et he a ae eer yas Pal hgh Oe Tp abet Bea Met FB BP ght P oPet APP adage ° VM ASEM IDES o> et bs hed ate ee tate ten se aie eae tie tee eS % et ae ASR Ate he = = ONT teh ty PORN cE es te ee : So a iehyere A ae Se a me ie tenet” a sors Aa ASS Ne ARN cate A oe Sette We ache ty he hee Be et aceite pele Cert.” setae te ae SS Sera emcee F od - c = be ~ ~ aes eee a ~ as Pp < = J ~ =f = « am - ~"? . - - —— A = 2 - J Boe Tae _ HARVARD UNIVERSITY Els EO LAS LIBRARY OF THE Museum of Comparative Zoology id is = =" ~ ge MUS. COMP. 4£U0L, LIBRARY ULI 14 1968 POSTILLA PEABODY MUSEUM YALE UNIVERSITY NUMBER 124. 9 SEPT. 1968 A NEW DEVONIAN FISH (CROSSOPTERYGII: RHIPIDISTIA) CONSIDERED IN RELATION TO THE ORIGIN OF THE AMPHIBIA KEITH STEWART THOMSON POSTILLA Published by the Peabody Museum of Natural History, Yale University Postilla includes results of original research on systematic, evolution- ary, morphological, and ecological biology, including paleontology. Syntheses and other theoretical papers based on research are also welcomed. Postilla is intended primarily for papers by the staff of the Peabody Museum or on research using material in this Museum. Editors: Jeanne E. Remington and Nancy A. Ahlstrom Postilla is published at frequent but irregular intervals. Manuscripts, orders for publications, and all correspondence concerning publications should be directed to: Publications Office Peabody Museum of Natural History New Haven, Conn., 06520, U.S.A. Lists of the publications of the Museum are available from the above office. These include Postilla, Bulletin, Discovery, special publications, and available back numbers of the discontinued journal, Bulletin of the Bingham Oceanographic Collection. All except Discovery are available in exchange for relevant publications of other scientific institutions anywhere in the world. A NEW DEVONIAN FISH (CROSSOPTERYGII: RHIPIDISTIA) CONSIDERED IN RELATION TO THE ORIGIN OF THE AMPHIBIA KEITH STEWART THOMSON Department of Biology and Peabody Museum of Natural History Yale University ABSTRACT Hyneria lindae, gen. et sp. nov., is a large rhizodontid rhipidis- tian fish from the Upper Devonian (Oswayo Formation) of Penn- sylvania. It shows a remarkable similarity to the intermediate stages that must have preceded the first true Amphibia. The similarity is due to the parallel evolution of rhizodontid Rhipidistia and the Amphibia from a common stock in the Middle or early Late Devonian. POSTILLA 124: 13 p. 9 SEPTEMBER 1968. 2, | POSTILLA INTRODUCTION The fossil fishes of the suborder Rhipidistia (Osteichthyes, Crossopterygii) have been the subject of intensive research for most of the present century because of their well-established posi- tion as the closest known ancestors of Amphibia. Research has been conducted according to a variety of approaches (taxonomic, mor- phological or functional), but one principal purpose has _re- mained — to test the hypothesis of a direct rhipidistian-tetrapod relationship by the study of resemblances and differences between the two groups. There are certain major obstacles to such a study, the principal ones being that only a relatively small number of rhipidistians are known and that many of the better preserved forms must, in fact, have been contemporaries of the first Amphib- ia. We must therefore conclude that the rhipidistian-amphibian transition occurred before the Late Devonian. Probably no Late Devonian Rhipidistia should be considered to belong to a lineage directly antecedent to the Amphibia. Such forms must be parallel radiations with the immediate amphibian precursors from a com- mon stock. The present paper records the existence of a large rhipidistian fish that shows a closer general resemblance to the Amphibia than any other known rhipidistian. This fish is from Upper Devonian (Oswayo Formation) deposits in Pennsylvania and_ therefore belongs in the list of rhipidistian fishes that are too young to be other than contemporaries of the first tetrapods. However, as will be considered later (under Discussion) the very close resemblance between this fish and the Ichthyostegalia offers some new, if indirect, evidence concerning the relationships of the known rhipidistian families to the first Amphibia. TAXONOMIC DIAGNOSIS AND DESCRIPTION Superfamily Rhizodontoidea Family Rhizodontidae Hyneria', gen. n. 1 From the village of Hyner, Clinton County, Pennsylvania, near which the specimen was collected. saw, GOUIVIE. ow we ie LIBRARY A NEW DEVONIAN RHIPIDISTIANL | 14 19693 TYPE SPECIES. Hyneria lindae, sp. n. HARVARD DIAGNOSIS. Large fish, estimated total length of Only! KRG WH Bpeci- men 250 cm. Proportions of trunk unknown. Skull with heavy dermal bones arranged in typical rhizodontoid pattern. Dermal elements lacking enamel layer, ornamented with a coarse network of anastomosing dentine ridges; isolated tubercles absent. Length of postparietal shield is contained approximately 2.8 times in length of parietal shield. Length of postparietal shield is contained approximately 4.5 times in length of lower jaw. At least one median postrostral element present, with two main lateral post- rostrals, all three elements being overlapped from behind by parietals. Pronounced spiracular notch in lateral margin of post- parietal shield. Large ventral ridge under each lateral portion of postparietal shield, passing anteromedially at a right angle to supratemporal-tubular suture. External parietal opening (not pre- served), if present, must be situated behind level of centers of ossification of parietals. Vomers triangular in shape but lacking extensive posterior flange. Lower jaw relatively elongate and shal- low. Principal gulars narrow and gently curved, lacking marked median angle. Length of principal gular is contained approximately 1.6 times in length of lower jaw. Teeth stout, rhizodontoid, conical with deeply furrowed base. Operculum subrectangular. Cleithrum in form of flat lamina of bone ornamented with anastomosing ridges running parallel to long axis of bone. Clavicle with stout ascending process. DESCRIPTION. The specimens are illustrated in Figures 3-13. These illustrations will serve in part for a description of the material. A unigue feature of Hyneria is the ornamentation of the dermal bones of the skull and the shoulder girdle, isolated denticles being almost completely absent from the surface pattern. On the skull elements, the dentine ridges form an extremely regular pattern of small symmetrical spaces enclosed by tuberculated ridges. On the shoulder girdle, the ridges tend to be more parallel and the en- closed lacunae are therefore elongated (in the direction of the axis of the cleithrum). The skull of Hyneria has been reconstructed (Figs. 1 and 2) on the assumption that the proportions of the various skull ele- ments (for example, the length of the parietal bone relative to the 4 POSTILLA Figure 1. Hyneria lindae gen. et sp. n. Reconstruction of skull in dorsal view, approx. 3/10. A NEW DEVONIAN RHIPIDISTIAN 5 Figure 2. Hyneria lindae gen. et sp. n. Reconstruction of skull in ventral view, approx. < 3/10. 6 POSTILLA whole parietal shield) are similar to those seen in other rhizo- dontid fishes such as Eusthenodon wangsjoi Jarvik (1952). Since the posterior portions of the parietal bone and the rostrum anterior of the median postrostral bone are missing, these parts are neces- sarily restored only tentatively. However, in no rhizodontoid does the center of ossification of the parietal lie at a level anterior to the midpoint of the bone (in most forms it lies in the posterior half of the bone). Furthermore, there is no indication of the parietal foramen in the preserved portion of the parietal shield (Figs. 11 and 12). A conservative restoration therefore places the center of ossification of each parietal at the midpoint of the length of the bone, as in the genus Eusthenodon. The relatively short rostrum is also a conservative restoration, based on the proportions in Eusthenodon. A further indicator of the cranial proportions is given by the relative length of the lower jaw. In all known Rhipi- distia, there is a more or less constant proportional relationship between the length of the lower jaw and the length of the post- parietal and parietal shields (Thomson, 1967a, fig. 10). Assum- ing that the present fish is typically rhipidistian we may make a conservative estimate of the cranial proportions from the known length of the postparietal shield and the approximate length of the lower jaw. The estimated proportions derived from the two meth- ods give the same ratio of the length of the parietal shield to that of the postparietal shield—approximately 2.8:1. If these estimates are correct they indicate that this fish has the longest parietal shield of any known rhipidistian. In this important feature, there- fore, Hyneria approaches the early tetrapod condition (e.g., Ichthyostega, in which the same ratio is 3.2:1) more closely than any other fish. The separate nature of the postparietal shield (Figs. 9 and 10) in Hyneria indicates that the skull in this form was divided by a typical crossopterygian intracranial joint (Thom- son, 1967a). The presence of the marked spiracular notch in the postparietal shield also indicates that Hyneria is a typical rhipi- distian, The functional significance of the ridge under the supra- temporal and tabular is not clear at present, but it may have served as an anchor-point preventing relative movement of the dermal skull roof and braincase, in addition to forming the medial margin of the groove for the spiracle. The shape of the vomer is significant. In all known rhizodontoids (but in no other Rhipidistia; Thomson, 1967b) the vomer has a A NEW DEVONIAN RHIPIDISTIAN 7 marked posterior flange passing along the side of the median tooth- bearing lamina of the parasphenoid, on the undersurface of the ethnosphenoid. The vomer of Hyneria has but a short posterior flange; this condition seems easily derivable from the more extreme condition seen, for example, in Eusthenopteron foordi (Jarvik, 1942, fig. 56). The mandible is narrow, elongate and slightly tapered anteriorly. No sign is seen of separate infradentary elements, although these are presumably present and obscured by the external surface orna- mentation (as is the case with the parietal shield, compare Figs. 11 and 12). The prearticular is large and a well-marked cavity for the coronoids was present. The tip of the mandible bears a large dentary tusk, as in Rhizodus. The gulars are elongate. The right principal gular shows a smooth medial margin where the left gular overlapped onto it. The right lateral gular series is preserved and its width may be accurately determined (Figs. 2, 5, and 6). However, this series of bones is badly fragmented in the available material and thus it has not been possible to determine the precise number of separate lateral gulars present. The sutures shown in the restora- tion (Fig. 2) are purely hypothetical, being based on the situa- tion in other rhizodontoid rhipidistians. The lateral gular series overlapped onto the lateral margin of the principal gular of each side, as indicated by the smooth overlap zone seen on the surface of each principal gular (Figs. 2, 5, and 6). The opercula of Hyneria are both slightly incomplete in the specimen (Figs. 7 and 8) but a fairly accurate restoration may be made. Each operculum is somewhat square in shape and con- siderably shorter along the anteroposterior axis than is the case in most Rhipidistia (anteroposterior length of operculum con- tained approximately 3.25 times in the length of the lower jaw). The opercula are superficially ornamented in the same style as the other cranial elements. No suboperculum has been found. The dermal shoulder girdle consists of a typical cleithrum and clavicle (Figs. 7 and 8). The cleithrum is a simple lamina, very similar to that of other Devonian rhizodontoids such as Eusthenop- teron. The clavicle (Figs. 3-6) shows a marked dorsal process not unlike that seen in the Carboniferous Rhizodus and much smaller than in other Devonian forms. No supracleithral elements have been found except for a right lateral extrascapula (Figs. 9 and 10) 8 POSTILLA which shows obvious overlap areas for a large anocleithrum and for the median extrascapula, the latter apparently also having been a relatively large element (Fig. 1). The only parts of the trunk that are known are isolated scales, one of which is associated with the cranial fragments. The scales are thicker relative to their diameter than most rhizodontoid scales and there is a prominent thickened rim along the margin of the posterior half of the scale (Fig. 13). Unfortunately, only the inner surface of the scale is shown on any of the available speci- mens (YPM 4938-4943). A curious feature of the scales is that the free margin is deeply notched. In each of the scales at hand, one of these notches is considerably more marked than the other (Fig. 13). Undoubtedly these notches in the free margin represent emarginations between the dentine ridges that make up the super- ficial ornamentation of the exposed portion of the outer surface of the scale. However, this notching is not known in other Rhipidistia. The inner surface of the scale is marked with a prominent central boss that seems to be continued posteriorly in the form of a thin ridge that cuts across the growth lines (Fig. 13). The boss itself is contained within the central area of the scale. It is possible that the scales showing this rather unusual structure came from a specialized region of the body, and that their structure is not typical. However, the same shape is seen in all available scales that have a complete margin. Hyneria lindae’, sp. n. SYNONYMY. “rhipidistian” Thomson, 1967a, p. 239 HOLOTYPE. MCZ* 9284: disarticulated skull in three blocks. PARATYPES. YPM?® 4938, 4939, 4940, 4941, 4942, 4943: isolated scales. OCCURRENCE. Uppermost Devonian |Oswayo Formation (Ebright, 1952) | of Clinton and McKean counties, Pennsylvania. From two ~Name derivation — for my wife. who has assisted me in collecting fossil fishes in many countries. ° Abbreviations used: MCZ — Museum of Comparative Zoology, Harvard University YPM — Peabody Museum, Yale University FIGURES 3-13 ABBREVIATIONS USED IN FIGURES AN anocleithrum CL cleithrum CV clavicle D dentary G principal gular ID infradentary IT intertemporal LE lateral extrascapular LG lateral gular LPR lateral postrostral MD mandible ME median extrascapular MPR median postrostral O operculum PA prearticular P parietal PP postparietal PQ palatoquadrate QJ quadratojugal SQ squamosal ST supratemporal T tabular V vomer VT vomerine tooth ‘pO X ‘xoidde aedfjojoy jo wonlog ‘u ‘ds ja ‘uad anpuy viuaudy *¢ omy *sdoRJINS QUOG [WISP po}USWeUIO soedIpUI SuIpeys AAvoy oy} ‘OT pue g ‘9 soins ul se ‘oinsy sty} uy ‘odAjyofoy Jo uonsod JO MOA ONeWUIeIseIq ‘U ‘ds ja ‘Uuas avpul] DidaUck ‘p 9INSIA ‘p'Q X ‘xoidde ‘gdAjojoy Jo uonsiog ‘u ‘ds 30 ‘uod appuy Diuaudy *¢ 91st 4 ‘gd{yojoy Jo uonsod appuy viuaudH “9 9Inslj JO MOIA oneuuedeiqd ‘U ‘ds yo ‘wos ‘adXjyojoy jo ‘yp’ X ‘xoidde uonlog ‘u ‘ds 3a ‘uas appuy viuaudy *L 9In3I4 ‘odAjojoy Jo uoniod JO MOIA oeMUEIseIq ‘u ‘ds 30 ‘uads anpuy vIUaUdT °g oMSIJ Ee, ‘v0 X -xordde ‘ds 39 ‘uod appuy visaudy “6 oIn314 ‘adAjojoy Jo uonszod JO MOIA oNeUUIeIseIG “u “ds jo ‘ued anpuy DIUaUKFT “QT omsty we te = — ———— Figure 11. Hyneria lindae gen. et sp. n. Portion of holotype, parietal shield in dorsal view, < 0.8. Figure 12. Hyneria lindae gen. et sp. n. Portion of holotype, parietal shield in ventral view, x 0.8. .n. Reconstruction of scale Q, n —_— oO ro] oO OD YS Ss) 3 iS roe is: op SX Sr oe ~ s m oO = is oa 48 ol ees =} Eas ie oa A NEW DEVONIAN RHIPIDISTIAN os Jocalities: 1) holotype and paratypes from a large road cut on the north side of U.S. highway 120, between the villages of North Bend and Hyner, Clinton County, Pennsylvania, and 2) fragments of a shoulder girdle which are cautiously referred to the same taxon from a locality on the east side of Route 46, 0.6 miles north of the McKean/Cameron County line, near Emporium, Pennsylvania. DIAGNOSIS AND DESCRIPTION. As for the genus. COMPARISONS Hyneria clearly belongs in the rhipidistian superfamily Rhizo- dontoidea, as is indicated by the structure of the scales, by the absence of enamel on the dermal bones and scales, and by the pat- tern of ornamentation on the dermal bones. At the present time, the Rhizodontoidea are classified into two groups, comprising on the one hand the genus Rhizodopsis (Family Rhizodopsidae) and on the other, the genera Tristicopterus, Eusthenopteron, Platy- cephalichthys, Eusthenodon, and Rhizodus (which probably includes Strepsodus). The genus Sauripterus is questionably included in this second group (see below) which normally forms the single family Rhizodontidae. Hyneria naturally falls with the rhizodontids. Of these, with the possible exception of Rhizodus (the skull of which is unknown), Hyneria is the most advanced form. In fact, it is remarkable that the Devonian genera in the family Rhizodontidae may be arranged in a temporal sequence which also corresponds to a direct sequence of structural modifica- tion. Within the group we may see a rapid achievement of a diphycercal tail (incompletely symmetrical in Tristicopterus), progressive elongation of the snout region of the skull (Fig. 14), and progressive modification of the dermal bone ornamentation from a pattern of more or less isolated denticles (Tristicopterus) to a pattern with the denticles arranged in short rows forming ridges (Eusthenopteron and Platycephalichthys), to the uniting of these ridges into a coarse incomplete network (Eusthenodon), to a complete regular network of ridges with no separate tubercles (Hyneria). The Carboniferous genus Rhizodus has a pattern of ornamentation in which some separate tubercles may be retained as in Eusthenopteron (more evident in R. ornatus than R. hib- berti). Hyneria clearly differs from each of the above-mentioned 10 POSTILLA Figure 14. Dorsal view of the skull in: A. Tristicopterus alatus, B. Eusthenopteron foordi, C. Eusthenodon wangsjoi, D. Hyneria lindae, E. Ichthyostega sp., F. Acanthostega gunneri, showing pro- gressive elongation of parietal shield. After Jarvik (1952) and original. A NEW DEVONIAN RHIPIDISTIAN Tel genera. However, since Rhizodus is rather poorly known it may be well to emphasize that in respect to those features of its struc- ture that are known, such as the shoulder girdle and operculum (Smith-Woodward, 1891), this genus seems quite separate from Hyneria. The possibility also exists that another very poorly- known rhipidistian, the genus Sauripteris from the Upper Devonian of Pennsylvania, is related to the material described above. How- ever, the shoulder girdle again seems rather different, apparently having a different structure of the clavicle. ENVIRONMENT OF DEPOSITION Study of the total fauna of the deposits from which Hyneria was collected will be deferred to a later date. We may note here that in addition to the large rhipidistian fish, very numerous small fragments of an antiarch similar to Bothriolepis (the presence of which helps to confirm the Upper Devonian correlation of the beds) and spines of “Xenacanthus’ are found. The antiarch remains are highly fragmentated, although very well preserved. Thin sections of the matrix in which the fossil materials are pre- served have been studied by Dr. D. C. Rhoads of Yale University, to whom I am grateful for the following observations. The deposits were laid down in quiet freshwater, probably in a flood plain between distributary channels. There is evidence of rework- ing of the sediment by burrowing organisms. Frequently the small bone fragments are seen to be displaced to a high angle with respect to the bedding by these burrowings. The red color is in the ferruginous cementing material. There were no great concen- trations of organic matter in the deposit other than the bone frag- ments. The environment was strongly oxidizing. From this I con- clude that the small bone fragments were probably carried into the region by flood waters (but from no great distance) and then accumulated temporarily in pools where larger fishes later became stranded. DISCUSSION As noted in the Introduction, there are many drawbacks to present discussion of rhipidistian-tetrapod relationships. If the Ichthyostegalia were not known, then the sequence of structural evolution seen within the rhizodontid Rhipidistia, culminating in 12 POSTILLA the genus Hyneria as described here, would seem a natural stage in the evolution of tetrapods. However, Hyneria cannot have evolved before the middle Late Devonian, while the advanced nature of the Late Devonian Ichthyostegalia shows that any struc- tural stages in their evolution corresponding to the rhizodontid sequence must have occurred much earlier. However, the fact of the close resemblance between the known Rhizodontidae and our analysis of what the ancestors of the first Amphibia must have been (see, for example, Thomson, 1966, 1967a, 1967b), in terms of both structure alone and the pattern of structural evolution, suggests a strong genetic relationship between the two groups. It also seems logical to assume that the rate of evolution in the immediately pre-tetrapod line(s) must have been far greater than in more “normal” Rhipidistia since a major series of adaptive changes was occurring. Possibly the best interpreta- tion of the out-of-phase evolution of rhizodontid Rhipidistia and ichthyostegal Amphibia is that they arose in the Middle Devonian from the same ancestral stock with a large number of features in common. Thereafter both lineages showed similar adaptations in terms of such features as the feeding mechanism (elongation of the snout), but with evolution proceeding much faster in the lineage leading to the Ichthyostegalia. ACKNOWLEDGMENTS [ am grateful to Professor A. S. Romer of Harvard University for the generous loan of material described in this paper and to Dr. D. C. Rhoads of Yale University for valuable interpretations of the lithology. The drawings were prepared by Miss Ward Whittington. The study has been supported financially by National Science Foundation grant GB-4814. A NEW DEVONIAN RHIPIDISTIAN 13 LITERATURE CITED Ebright, S. R. 1952. The Hyner and Ferry anticlines and adjacent areas Center, Clinton and Lycoming Counties, Pennsylvania. Penn. Geol. Surv. (4) Bull. M 35:1-32. Jarvik, E. 1942. On the structure of the snout of crossoterygians and lower gnathostomes in general. Zool. Bidrag. 21:237-675. 1952. On the fish-like tail in the ichthyostegid stegocephalians. Meddr. Grgnland 114:5-87. Smith-Woodward, A. 1891. Catalogue of fossil fishes in the British Mu- seum. Vol. 2. London. 567 p. Thomson, K. S. 1966. The evolution of the middle ear in the rhipidistian- amphibian transition. Am Zoologist 6:379-397. 1967a. Mechanisms of intracranial kinetics in fossil rhipi- distian fishes (Crossopterygil) and their relatives. Jour. Linn. Soc. London (Zoology) 46:223-253. 1967b. Notes on the relationships of the rhipidistian fishes and the ancestry of the tetrapods. Jour. Paleont. 41:660-674. REVIEW STYLE FORM ane ABSTRACT YVOMENCLATURE ILLUSTRATIONS FOOTNOTES TABLES REFERENCES 'THOR’S COPIES PROOF COPYRIGHT INFORMATION FOR AUTHORS The Publications Committee of the Peabody Museum of Natural History reviews and approves manuscripts for publication. Papers will be published in approximately the order in which they are accepted; delays may result if manuscript or illustrations are not in proper form. To facilitate review, the original and one carbon or xerox copy of the typescript and figures should be submitted. The author should keep a copy. Authors of biological papers should follow the Style Manual for Biological Journals, Second Edition (Amer. Inst. Biol. Sci.). Authors of paleontological manuscripts may choose to follow the Sugges- tions to Authors of the Reports of the U.S. Geological Survey, Fifth Edition (U.S. Govt. Printing Office). Maximum size is 80 printed pages including illustrations (= about 100 manuscript pages including illustrations). Manuscripts must be typewritten, with wide margins, on one side of good quality 814 x 11” paper. Double space everything. Do not underline any- thing except genera and species. The editors reserve the right to adjust style and form for conformity. Should be precise and short. Title should include pertinent key words which will facilitate computerized listings. Names of new taxa are not to be given in the title. The paper must begin with an abstract. Authors must submit com- pleted BioAbstract forms; these can be obtained from the Postilla editors in advance of submission of the manuscripts. Follow the International Codes of Zoological and Botanical Nomen- clature. Must be planned for reduction to 4 x 612” (to allow for running head and two-line caption). If illustration must go sideways on page, reduction should be to 3%4 x 634”. All illustrations should be called “Figures” and numbered in arabic, with letters for parts within one page. It is the author’s responsibility to see that illustra- tions are properly lettered and mounted. Captions should be typed double-spaced on a separate page. Should not be used, with rare exceptions. If unavoidable, type double-spaced on a separate page. Should be numbered in arabic. Each must be typed on a separate page. Horizontal rules should be drawn lightly in pencil; vertical rules must not be used. Tables are expensive to set and correct; cost may be lowered and errors prevented if author submits tables typed with electric typewriter for photographic reproduction. The style manuals mentioned above must be followed for form and for abbreviations of periodicals. Double space. Each author receives 50 free copies of his Postilla. Additional copies may be ordered at cost by author when he returns galley procf. All copies have covers. Author receives galley proof and manuscript for checking printer’s errors, but extensive revision cannot be made on the galley proof. Corrected galley proof and manuscript must be returned to editors within seven days. Any issue of Postilla will be copyrighted by Peabody Museum ot! Natural History only if its author specifically requests it. v WANN 3 2044 066 305 BOUND DEC 1972 fend Dre PAPEETE pe te Sorte: ars + Pane sone teria a " PSF FL 7 , ; sf ; : : r = ’ PRE SE Ne aera es EP C : - ate nme Spry mere a ee eet §epnatt ere of TE RRL ea pe F PERE A SER PIIIL a oa ot a ee eR ee id