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SMITHSONIAN
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¥YOLUME: XLVIL
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The present series, entitled SMITHSONIAN MISCELLANEOUS COL- LECTIONS, is intended to include all the publications issued directly by the Smithsonian Institution in octavo form, excepting the ANn- NUAL Report to Congress; those in quarto constituting the SMITH- SONIAN CONTRIBUTIONS TO KNOWLEDGE. ‘The quarto series includes memoirs embracing the records of extended original investigations and researches, resulting in what are believed to be new truths and constituting positive additions to the sum of human knowledge. The octavo series is designed to contain reports on the present state of our knowledge of particular branches of science; instructions for collecting and digesting facts and materials for research; lists and synopses of species of the organic and inorganic world; reports of explorations; aids to bibliographical investigations, etc., generally prepared at the express request of the Institution, and at its expense.
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ARTICLES
The Structure of Wing Feathers. E. Mascua. (PLATES I-XVI.) Mawes 7s. steaplished WMay,.6; L905... ie 2' een ole pe ees I The Tarpon and Lady Fish* and their Relatives. THEODORE GILL. (PLatTEs xvui-xx1.) No. 1576. Published May 13, 1905.. 31 Diagnosis of a New Genus and Species of Fossil Sea Lion from the Miocene of Oregon. FREDERICK W. TruE. No. 1577. Pub-
Pete Wha 145, TOOR 202 2) nt Manet po eioe a rip oir sat Sere sie alates 47 Diatoms, the Jewels of the Plant World. AtBert MANN. (PLATES eati-xxy.y No. 1578. Published May 23, 1905.......... 50 Notes on the Nomenclature of Certain Genera of Birds. THarry C. OBERHOLSER. No. 1579. Published May 13, 1905...... 59 The Bibliography of Halley's Comet. E. F. McPixe. No. 1580. eee ME FOP EGUS vo eons vr ce dn vy aie eee «Uke et ors 69
The Ancestral Origin of the North American Unionide, or Fresh- Water Mussels. Cuartes A. WHITE. (PLATES XXVI—XXXI.)
Biehnente Eplished: | WME. TO, TOOK sc. cae aagers coer + eee ies eyahe 75 Brain Weight in Vertebrates. Aes Hrpiticxa.” No. 1582. Pub- Reem TO. MOG: Ait cs lecenc eo 3 et, ve atch wes soe ae Sitiale » 89 ores, “(PLATES XxXxII, xxx.) “No. 1583. (Smithson “Mor- emer ICUS ue, ole tiess feed «a ~ cjuas MRRIOR oy en Sart donne 113
The Diplomatic Service of the United States, with some Hints to- ward its’ Reform. ANDREW D. Wuite. No. 1586. Published Dire EUR IV Phe on «2 ao vd ska 0 320s what aie tie LOK ee ee 117
The History of the Whale Shark Rhinodon typicus Smith. Barton A. BEAN. (PLATES XXXIV-xxxvi.) No. 1587. Published July
2 DOES ene cla: Ue pea ae RA Mee SON! emir edi area 139 The Avian Genus Bleda Bonaparte and some of its Allies. Harry C1Oeernorser (No. 1588. . Published July 1; 1905.......«.' 149.
Scaphoceros tyrelli, an extinct Ruminant from the Klondike Gravels. WILFRED H. Oscoop. (PLATES xxxviI-xLi.) No. 1589. Pub- Rpm CHEE 030.5 oo, win wind wialar a’ don poareumeaniye sist gs ns “es Ones 073
A new Genus and several new Species of Landshells Collected im Central Mexico by Doctor Edward Palmer. Witt1aM HEALEY Parr, .) (PrAres \ xiiii—xiiv.): No. 1590... Published) July \ 1, TOS AS Vo ee Mel neiAb mart bees mitre ar 187
The Family of Cyprinids and the Carp as its Type. THEODORE GILL. (PLATES xLv-Lyi1u.) No. 1591. Published September SM ee eee al val sh aia Tas, wine: ae unl Soaynichy een cee eine ao = 195
The International Catalogue of Scientific Literature. Cyrus ADLER. Noetne2. Published September 8, 1905... 22... a a 219
Vv
vi SMITHSONIAN, MISCELLANEOUS COLLECTIONS [voL. 48
Instances of Hermaphroditism in Crayfishes. WILLIAM PERRY Hay.” No) 1593. Publisted September 10,2 1005 co. 0s cracane 222 PN ODES. 7) INGE GOA coi ester oases MT oso vty oa 229 The Species of Mosquitoes in the Genus Megarhinus. Harrison G. Dyar and Freperick A. Knas. No. 1657. Published Septem- IDES 7. POCO. os, ou cicese eres fora ted yn Saale nee pete eaep ely cia tx ees 241 A Contribution to the Knowledge of some South American Hymen- optera, chiefly from Paraguay. C.ScuHrotrKy. No. 1658. Pub- lished’ ebrtiaty. 4, 1G07 >. ....:'s <b Samos ciate erate ay Seueiena tome reriens 259 Description of a New Squirrel of the Sciwrus prevosti Group from Pulo Temaju, west coast of Borneo. Marcus Warp Lyon, Jr. No. 1659. Published February 4, 1907. .2.:-% iy. emmy 275 The Squirrels of the Sciurus vittatus Group in Sumatra. Marcus Warp Lyon, Jr. No. 1660. Published February 4, 1907.. 277 A Study in Butterfly Wing-Venation, with special regard to the radial vein of the front wing. Tuomas J. HEADLEE. (PLATES
LIX-Lx1.) No. 1661. Published February 4, 1907...... 284 Some Noteworthy Extra-European Cyprinids. THEODORE GILL. No: 1662... Published Pebsuary. 4:) 10070. c inet i ees 297 A Review of the American Volutide. Witt1AM HEALEy DALL. No. 1663.; Published) Pebruaty 4) 1007... naa cena ae 341 INO TES Ne TOBA Ns c's: alate eleetretenees otoos oy es Aa tse aie ects eee ae 374
The Breeding Habits of the Florida Alligator. ALBertT M. REESE. (PLatEs Lx1v-Lxv.) No. 1696. Published May 4, 1907.. 381 Life Histories of Toadfishes (Batrachoidids) Compared with those of Weevers (Trachinids) and Stargazers (Uranoscopids). THEO- DORE /GICL.-: No.» 1607. > Publisheds May. <4, - 1007-22 see 388 The Letter of Dr. Diego Alvarez Chanca, Dated 1494, Relating to the Second Voyage of Columbus to America (bemg the First Written Document on the Natural History, Ethnography, and Ethnology of America). ‘Translated, with notes, by A. M. FEr- NANDEZ DE YBARRA. (PLATE LxviI.) No. 1698. Published May Ais STA Ee ccs o eso 3 ata cee Ie ance pce eee 428 The Origin of the So-called Atlantic Animals and Plants of Western Norway. LEONHARD STEJNEGER. (PLATES LxviI-Lxx.) No. 1699. Published May 4, 4GG7: mec eine, oe ee coer 458 Manners and Customs of the Tagbanuas and other Tribes of the Island of Palawan, Philippines. MANuEL Huco VENTURELLO, Translated by Mrs. Edw. Y. Miller. No. 1700. Published Naas 1907... «see ay eee eee ete ols pean eter oe On ee 514 INGPESs | SINOL ZOD coi oeg ea ee cee oaks inact ele toe ase eee a 559 1 S101 ah eo RRR Th Rm fe Snake Cone ae: 563
PLATE
I-XVI. XVII-XXI. XXII-XXV. XXVI-XXXI. XXXII, XXXII. XXXIV. XXXV. XXXVI. XXXVII-XLII. Pi XLV. XLV-LVIII. LIX-LXIII. LXIV.
LXV.
LXYVI.
LXV, LX VIII. XTX.
Ex.
LIST. OF} PLATES
PAGE Structure) of - wine festhersic. 6 2%) oe 5s 30 We CAC POR epee AISI 8 Sie eso wo Bote 46 TPO MIS \ ReeUelaRts re A eee ore tte Tenor Ane Sake 58 Si aslsicbtee aa are eine wm clans Uegne Saab gilts Weer amg 88 Smithson Mortuary Ciapel Ppa: ie Saheim 114 Whale Shark (Rhinodon typicus)....... 139 Rhinodon typicus, Smith’s plate.......... 140 Teeth of Microstodus punctatus......... 146 SCOPMOCETAS, PYEREUR nt. oO te eimnl anid Sard scnion 186 Land shells’ from, Central Mexico. ....)...25 194 Bamily: Of Gyprinids, | ey erar. cots sss 218 Butterfly. wine-vyenstion. 22.4. 2096 PU OOM EMEStGN eee NTL Me aE 6 ats cag ae 386 PUI ACOr estan Ces SN. 2 ashes os 387 Ea Cosacmup or, Ameren. L500e <5 <4). a 430 skal: of \Cenmus) ahlamitens? yo i. foe ews 404 Morweotai “yonds herses! 3) Ceg5 sc). -s eae 472
Hypsographic map of Northern Europe. .
Remy H7 SAAN “| Ah
oF a o8
VOL. 48 1905
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VCL. Ill QUARTERLY ISSUE PART I
THE STRUCTURE OF WING-FEATHERS By DOCTOR E. MASCHA
[The following paper on the “ Structure of Wing-Feathers ”’ gives an account of an investigation conducted by Doctor E. Mascha under the direction of Professor R. von Lendenfeld, of the Imperial German University in Prague. The original German text of this paper is to be published in the Zeitschrift fiir wissenschaftliche Zoologie.
Doctor von Lendenfeld, who was a competitor in the Hodgkins competition of the Smithsonian Institution, has more recently been awarded a grant in aid of his research on the anatomy of the flight organs, the investigation being continued under his direction by Doctor Mascha, as here described. |
Il. INTRODUCTION
The object of this paper is in the first instance to give a detailed account of the morphology of the wing-feathers of birds as used in flight. As our knowledge of this subject is far from satisfactory, I have made a special study of the organs of flight, hoping that I might be able to supply needed and valuable information for those interested in the great problem of aerial navigation.
The remiges of Columba livia were first investigated. Although the structure of these feathers is in many respects simple and typical, several questions concerning the function of particular parts could be satisfactorily answered only by a comparative study of the wing- feathers of other birds. Through this extension of my investiga- tion I was enabled to find out which parts are constant and therefore probably essential, and which not being always present are infer- entially of secondary importance. Among the most noteworthy dis- coveries made through this comparative study are a knowledge of
I
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
the variability in the size and structure of the secondary fibers, the recognition of the importance of their ventral ridge, the variation of the hook fibers and the constancy of the curved fibers.
II. THe STRUCTURE OF THE REMIGES
1. The Structure of Feathers in General
According to the nomenclature adopted in this paper each feather consists of the following parts: (1) Quill or primary quill, (2) secondary quills, (3) tertiary fibers (a) hook fibers, (b) curved fibers.
The primary quill is the bearer of all parts of the feather. Its shorter proximal part is circular in section and hollow, its longer distal part more or less rectangular in section and filled with medul- lary air-cells. The primary quill is slightly curved downward and inward; on its sides the two feather vanes extend. They lie nearly in the same plane. The two vanes are nearly equal in the remiges attached to the ulna; distally the outer vanes diminish in breadth more rapidly than the inner. The outermost of the hand-remiges attached to the metacarpals and phalanges have a broad inner and “a comparatively narrow outer vane. The vanes are composed of the secondary quills which rise obliquely from the upper part of the primary quill, and of the tertiary fibers rising in a similar manner from the secondaries. The tertiaries are closely connected with each other ; together with the secondary quills they form the apparently continuous surface of the feather vane or feather plate.
2. Material and Methods Remiges of the following species were examined:
PasseErES: Fringilla spinus, Turdus vulgaris, Garrulus glandarius. Coracit®: Merops apiaster, Galbula viridis.
Bucerotes: Buceros monoceros.
Macrocuires: Cypselus apus, Micropus melba, Macropteryx mystaceus. CaprimMu_et: Podargus humeralis, Caprimulgus europeus.
Srrices: Strix fammea, Bubo maximus, Bubo mexicanus, Nyctea nivea. Psittact: Chrysotis @estiva, Sittace cerulea, Stringops habroptilus. CucuLi: Cuculus canorus.
Musopuaci: Turacus albocristatus.
CoLumBa&: Columba livia.
TUBINARES: Diomedea exulans.
ANSERES: Cygnus olor.
Accipirres: Aquila chrysetus.
*The systematic arrangement adopted is that of F. E. Beddard, ‘“ The structure and classification of Birds.” London, 1808.
MASCHA] STRUCTURE OF WING—FEATHERS 3
This list, although small, comprises birds of most types of flight, my aim being not to examine many different species but only rep- -resentatives of the different types. The remiges of Columba, Cypselus, and Diomedea were studied most carefully ; those of birds not capable of flight did not fall within the scope of my investiga- tions."
I have taken no notice of the numerous new terms concerning the arrangement of the feathers proposed by Alix (1864, p. 10), Wray (1887, pp. 344-345) and others, the older divisions into hand- remiges and arm-remiges being quite sufficient for my purpose.
It was to be foreseen that the treatment of such brittle and hard material as the horny substance of feathers would present serious technical difficulties. Former investigators have left no record of the way in which their examinations on developed feathers were conducted. It is true that Strong (1902, pp. 148-151) has described his mode of procedure rather fully, but he speaks merely of the treatment of “ feather germs.”
I treated the material to be examined in the following manner: I took one of the three outermost hand-remiges, usually the longest, and one of the arm-remiges of each species of bird to be examined, and placed a portion of the vane on the slide mounted in balsam.
Then one of the secondary quills was cut off and the tertiary fibers, attached to it, removed to the slide by means of a sharp scalpel. In this way large numbers of isolated tertiary fibers lying in different positions were obtained and mounted in balsam. Parts of white colorless feathers became so transparent in this medium that recourse to staining was resorted to. It should be noted that of the numerous stains tried, only two proved useful: picric acid, which quickly produced a yellow coloring, too faint however to be effective, and safranin, which stains a deep color and gives good results when properly applied. I used the safranin in a semi- alcoholic solution and left the objects in it from six to twelve hours. After being stained the feathers were dried and then further manipu- lated. The most difficult part of the work, but at the same time the one that gave the best results, was section cutting with the micro- tome, in which operation the fragility of the material proved very troublesome. The specimens were placed in chloroform to expel the air, and imbedded in paraffin or celloidin. In cutting the paraffin
*The wings of birds not capable of flight have no hook fibers on the ter- tiaries, as their wings do not need to form such an impenetrable surface as is requisite in birds of flight. In the ostrich the tertiary fibers are entirely smooth (branchless) ; in the cassowary and the apteryx they show small thorn- like protuberances which are also present in the penguin, but somewhat longer.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS [vor. 48
blocks I successfully adopted the method of covering the top of the block with melted paraffin after each section, and congealing it by means of a current of cold air.1 A very oblique position of the knife is essential to procure good sections. In the paraffin method I stained with safranin before imbedding. The sections were glued to the slide with Schallibaum’s collodion-clove oil and cleared with xylol. Although thin sections could be made by this method it had many drawbacks. The sections frequently split, and while remov- ing the paraffin with xylol numerous portions of them were lost. By experiment I found that the splitting was almost entirely ob- viated by the employment of celloidin, although it is true that the celloidin method also presented difficulties. It is hard to make as thin sections of celloidin blocks as of paraffin, and staining is much more difficult. If one stains (with safranin) before imbedding, the ether extracts even the strongest stain. If one stains the sections after cutting, the celloidin also takes the stain. The sections were at first made only on the two directions parallel to one of the two kinds of tertiary fibers. These proved to be very instructive for the exact study of the tertiaries but did not throw sufficient light upon the structure of the secondaries. To study these, sections were made vertical to the secondaries, which gave the desired information concerning them. 3. The Primary Quill
The morphology of the primary quill (fig. 6, Htk)? have already been fully discussed. Ahlborn (1896, p. 15-16) in particular has given a complete description of this part of the feather, illustrated by.a sketch, and has drawn attention to the importance of its struc- ture for flight. I therefore turn at once to the description of the parts composing the feather vanes, beginning with the considera- tion of the secondary quills.
4. The Secondary Quills The secondary quills are the bearers of the feather-vanes. Just as they themselves spring obliquely from both sides of the dorsal part of the primary quill, so they bear dorsally the tertiary hook and curved fibers. Clement (1876, p. 282) has drawn attention to this in a somewhat different sense. He calls the plate, composed of the two vanes of a feather, ve-villwm, and the vane formed by the tertiary
*The method employed was described by Professor von Lendenfeld in Zeitschrift fiir wissenschaftliche Mikroskopie und mikroskopische Techmk, v, 18, pp. 18-10.
2For figures see plates at end of the paper.
MASCHA | STRUCTURE OF WING—FEATHERS 5
fibers on each side of the secondary quills vexillum primitif. After him Strasser (1885, p. 197) noticed these structures, and introduced the expressions secondary quills and secondary vanes, to distinguish the former from the primary quill and the latter from the primary vanes of the whole feather.
The secondary quills arise dorsally from the sides of the primaries, and extend obliquely outward. The angle between them and the primary quills is about 50° in the arm-remiges and changes but little in the whole length of these feathers in both vanes. It is subject to greater variations in the hand-remiges. Here it is greatest at the base of the feather (about 50°), towards the end of the feather it becomes smaller, the outermost secondary quills rising from the primary quill at an angle of only 20-25°. In the central broadest part of the hand-remiges this angle is 30-40°. In the outer vane of the hand-remiges the angle is always smaller than in the correspond- ing part of the inner vane. At the distal end of the feather the secondary quills are curved and their tips turned in the direction of the continuation of the primary quill. The secondary quills are for the greater part of their length considerably compressed laterally, becoming band-like. They are highest at their origin from the pri- mary quill and decrease in height outwards, finally tapering to a fine point (fig. 12, a, b, c).
A few words must be said here concerning the histological struc- ture of the secondary quills. Like the primary quill they appear as horny tubes (figs. 1, 2, 3, 4, Hus) enclosing a medullary sub- stance or core composed of large polyedrical cells filled with air (figs. I, 2, 3,4 Mks). Klee (1886), p. 92-30) has pointed out that the cortex and the core are essentially different, but that both arise from similar elements, at first homogeneous. Davies (1889, pp. 588-589) has fully and clearly described this process of conversion of the intermediary cells into cortical and medullary substance. He says: “ The modification consists in a considerable increase in size of the central space which, in horned cells, contains the nucleus, combined with a change in the form of the cell.” The relative posi- tions of the cortex and the core are shown in the transverse sections of secondary quills represented in figs. 1-7, 29.
Two types of secondary quills can be distinguished. In the first, which is by far the most frequent, the core is irregular, composed of many layers of irregularly arranged core cells and the interior of these secondary quills presents a honeycomb-like structure. In the second type, met with in the Owls and Caprimulgi (figs. 2, 4, 7, 29), the core cells are arranged in a single vertical layer, irregularly
\
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
polyhedral and overlap each other like roof slates. Here and there a more irregular polyhedral cell is interposed between the others. The transverse sections show that these quills are not simple bands but curved more or less transversely, convex on one side and con- cave on the other. The manner in which the secondary quills, which are highest at their proximal end, become lower toward their distal end is correlated in many birds to that curvature in such a way that the curvature is greatest where the quills are highest and decreases with the height outward, finally disappearing almost entirely near the distal end. Strong (1902, pp. 158-159), draws attention to an extension of the lower margin of the ‘secondary quills which he terms “the ventral ridge,” and which he represents as a constant character. In early stages of the development of the feather this ridge is said to be very large, being much reduced later. This ven- tral ridge of the secondary quills, which also appears in the covert feathers of the wing (fig. 5), is functionally a very important char- acter of all remiges. It is small in Columba, Cypselus and others, larger in the Caprimulgi (figs. 2, 4, 7, Hnl), still further developed in the owls (fig. 29, Hnl) and in many water birds, and largest in Aquila and Diomedea. Hand in hand with the development of this ventral ridge goes the development of the curvature. The great development of the ventral ridge, combined with the strong curva- ture of the secondary quills in Aquila and Diomedea probably enables these birds to perform the sailing flight characteristic of them. It must be noted, however, that the ventral ridge is very well developed in owls and various water birds also, which are not such excellent sailors of the air as the albatross and the eagle. Ahlborn (18606, ». 20) describes a peculiarity of certain parts of the remiges in the duck, the swan and other birds. He says that portions of the lower sides of these feathers seem to be covered by a fine membrane, this appearance being due to the fact that here the secondary quills are not only connected with each other dorsally by the tertiary fibers but also ventrally by delicate membranous extensions of their ventral margins, which bridge over the spaces between them. These ex- tensions enclose a right angle with the band shaped secondary quills from which they arise and exactly fit on the ventral margins of the quill in front, to which they adhere on account of their intrinsic elasticity. This membranous extension of Ahlborn .is only the strongly developed ventral ridge of the secondary quills.
The decrease outwardly in the height of the secondary quills is uniform in some birds but not so in others. In the latter there is a sudden decrease in height in the secondaries of the broad inner
MASCHA | STRUCTURE OF WING—FEATHERS 7
vane about half way between the primary quill and the margin of the feather, or nearer to the former. Here also the ventral ridge suddenly becomes lower in the region where the height of the quills themselves abruptly becomes less. In the vanes with uniformly de- creasing secondary quills the central margins of these quills stand out free, so that one clearly sees the ribbing on the under side of the feathers of which Parseval (1889, p. 70) has spoken. It is quite different in the Striges, Accipitres and Diomedea, birds described by Ahlborn. Here the secondary qnills are very high and markedly concave at their point of origin and the ventral ridge is of considerable size in the basal part of the secondary quill; being nearly vertical to the quill itself it lies almost horizontally, and ex- tends to the convex rear side of the secondary quill in front of it (fig. 33). In the middle of the length of the secondary quill the ridge becomes so small and the angle between it and the quill to which it belongs so large, that the ventral connection of the second- ary quills formed by these ridges ceases. Outside this well marked line their ventral margins are free. If one looks at the ventral side of the feather it is noticed that in the proximal third of the vane the light is reflected from the ridges producing a silvery luster which ceases rather suddenly at a line parallel to the primary quill, an appearance which can be observed more or less clearly in the remiges of all these birds.
In the development of this ventral ridge two types can be distin- guished. One is represented by the remiges of Columba, the other by those of the Striges, Tubinares, and Accipitres. In the first the ridge is a stout, low ventral projection of the cortex of the secondary quills and appears wedge-shaped in transverse sections (figs. I, 3). In the second it is a band-shaped membrane thin down to its base, attached vertically to the high and markedly convex secondary quill (fig. 29). These types are connected by transitional forms. A peculiar development is observed in the secondary quills of the remiges of the Caprimulgi in which the ventral ridge is not sharp but blunt, of uniform thickness throughout and rounded at the margin. On the upper edge also the secondary quill bears a ridge, which is broad, low and rounded (figs. 3, 4, 29, Vd), and of no functional importance for flight.
The tertiary fibers are attached to the secondary quills in a com- plicated manner. In examining sections parallel to one of the two kinds of tertiary fibers, I could only indistinctly make out their junction, but the sections vertical to the secondary quills were much more instructive. I could find no description of this apparatus in
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
the whole literature on the subject. The works of Haecker (1890, pl. Iva, 1900, pl. x1v) contain some figures of transverse sections of secondary quills, which are not absolutely correct, but are essen- tially better and more true to nature than some others (e. g., Chad- bourne, 1897, pl. 1a). Unfortunately I was not able to see the treatise of Jeffries (1883) mentioned by Strong (1901, p. 160). Strong himself (1902, figs. 7, 8, 9, pl. 1) has published several good figures of transverse sections of secondary quills. Since, however, he made such sections of covert-feathers only, it is natural that the results I obtained deviate considerably from his. It seems that these formations are much more complicated in the remiges than in the covert feathers. According to my observations in the latter they have the following structure: Under the starting points of the tertiary fibers there arises from each side of the secondary quill a longitudinal ridge, not vertical but extending obliquely upward (figs. 3, 4, G), which consists entirely of horn-substance. Its outer surface is smooth and continuous with the side of the secondary quill. Its inner side, which is turned toward the upper part of the secondary quill, is grooved, the grooves being separated by parallel secondary ridges (figs. 3, 4, L), which extend obliquely upward and outward. These ridges are highest at their origin and decrease in height dis- tally ; in the transverse section they appear like the teeth of a comb, and become shorter the further they are away from the secondary quill, the last being a slight prominence only. The ridges themselves are broadest at the base and grow thinner above. Their number in the sections varies from two to four and they may be more numerous in the secondary quills of larger feathers. In the small covert- feathers this main longitudinal ridge occurs, as has been described by Strong, but here the secondary ridges rising from its inner face are absent. The most difficult question to answer is, how the outer edge of the main ridge is formed. Without having reached a per- fectly satisfactory conclusion, I think I may pronounce it as probable that it is not smooth but interrupted by incisions lying between the ends of the secondary ridges. If this were not so, in a succession of transverse sectiows of secondary quills the secondary ridges would always terminate at exactly the same level, which, as serial sections show, is not the case. The level and the appearance of the edge of the ridge vary in every section, and on this fact the above supposi- tion is based. In the grooves between the successive secondary ridges lie the ventrally thickened margins of the proximal parts of the tertiary fibers. In sections one sees the first between the secondary quill and the first secondary ridge, and the second be-
MASCHA] STRUCTURE OF WING—FEATHERS 9
tween this and the second secondary ridge (figs. 3, 4). If there are more than two secondary ridges in the section, the third and fourth are so small and the grooves lying between them so shallow, that the position of the fiber-sections relative to them becomes doubtful. In surface views one sees parallel lines at the side of the dorsal ridge of the secondary quill. These lie between the basal parts of the tertiary fibers and enclose an angle with the secondary quill con- siderably more acute than the angle between the tertiaries and the secondary quill. They cross the direction of the latter at angles of 20-25°. Ina line parallel to the secondary quill, which is the edge of the longitudinal ridge, these lines, which are the free margins of the secondary ridges described above, terminate suddenly (fig. 20,1)’.
If we combine the results of these observations we see that the basal parts of the tertiary fibers lie between the partitions like the bats of a stidiron, These bars arise from the inner face of a longitudinal ridge arising from the side of the secondary quill and extending obliquely upward. The bars decrease in height out- wardly and finally pass into the margin of the longitudinal ridge. The thickened ventral margins of the proximal portions of the ter- tiary fibers are enclosed between these bars; beyond the margin of the longitudinal ridge they are free. At the margin of the ridge a sharp bend occurs in the ventral margin of the tertiaries, its free outer portion enclosing a larger angle with the secondary quill than its proximal part. In the nearly symmetrical vanes of the distal remiges the secondary quills extend almost horizontally and, there- fore, the two vanes also are nearly horizontal. In the broad inner vane of the hand-remiges on the other hand, only the basal part of the secondary quills extends horizontally. Farther out they are at first bent a little downward but rise up toward their ends. The inner feather-vane therefore appears bent slightly in the form of an S, the hind margin of the feather being distinctly turned up. In the outer vane the secondary quills, which are here very short, are curved slightly downward. In the remiges of many birds of prey the inner vane suddenly becomes narrower in the upper third of its length. In this case the inner vane has a distinctly S-shaped curva- ture in its broad, proximal portion, the outer edge being much raised, while this curvature suddenly ceases where the broad, proxi- mal part passes into the narrow, distal part, the latter being like the front vane slightly bent downward. Since according to Ahlborn (1896, p. 18) the S-shaped curvature of the inner vane of the suc- ceeding remiges serves to insure a close overlapping of all parts
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS LvoL. 48
of the wing, it is undoubtedly of interest that in the wings of these birds this curvature is developed only so far as the feathers really overlap each other.
The secondary quills have the same structure in all remiges of a wing, but we observe a regular variation of their size in the different feathers of the same wing and in the different parts of each feather. An increase of the stiffness of the secondary quills is noticeable, as we proceed from the proximal to the distal part of the wing. Hand in hand with this goes an increase of the development of the ventral ridge, which is relatively small in the arm-remiges but attains enormous dimensions in the hand-remiges of Diomedea, the Accipitres, and Striges. At the same time the outer vane in the hand-remiges becomes constantly narrower, as compared to the inner, and at the same time stiffer and firmer. The secondary quills in the narrow vanes are’basally just as high as the sec- ondary quills of the corresponding broad vanes lying opposite them, and they do not terminate in fine threads like those of the broad vanes but are only slightly pointed at the end. The elasticity of the secondary quills contributes to keep the elements of the narrow outer vane in their proper position and order.
Besides these differences of size in the secondary quills of the different feathers of the wing, similar variations occur also in the different parts of each feather as mentioned above. The supposition is obvious that the secondary quills are thickest and highest at the base, and that they should become lower towards the end of the feather, where the primary quill also decreases in thickness. Such conditions are really found in the arm-remiges of all birds and in the outer hand-remiges of Diomedea. In the outer hand-remiges of numerous birds, such as the Columbinze, Coccygomorphe, Cypselide, Strigide, and Natatores, however, the secondary quills are moder- ately high at the base of the feather, increase gradually in height distally and attain their maximum height at or just beyond the middle of the feather. Measurements in corresponding portions of different hand-remiges gave the following results:
| Height of Secondar Juills. | Length of . y@
Sue | Whole Vane. | 3 cm. from the Middle of the | 3 cm. from the Bas Vane. | Tip. : AM ihe ss a es | ea z z Macropteryx mystaceus. | 18cm. | 300 kL | 389 uC 233 be Bubo maximus. 28 | 550 g1o 700 Cygnus olor. 26 | 489 | 1,223 678
The secondary quills are measured close to the primary quill near their point of origin, that is at the point of their greatest height.
MASCHA] STRUCTURE OF WING—FEATHERS II
A comparison of these measurements shows how great, especially in Cygnus olor, is the increase in height of the secondary quills from the base to the middle of the feather. The decrease in height towards the end of the feather is gradual, the secondary quills 3 cm. from the tip of the feather are sometimes considerably higher than at a similar distance from the base.
We know that the development of the parts corresponds to the demands made upon them and that in general an organ is most strongly constructed and most capable of resistance exactly at the point where this strength is most needed. The task of these feathers is to bear and to raise the bird’s body, by pressing on the air below, therefore the secondary quills are highest and as a consequence the feather vane the strongest and most capable of resistance where the greatest force is to be withstood. The wing being a concave surface, we can assume that its parts will be subjected to different pressures, and further, that where the pressure is greatest the feathers will be strongest. Apart from the variations of curvature we must also take into consideration the length of the feathers relative to the whole wing. In Diomedea the arm-remiges are short in comparison to the length of the wing and the secondary quills highest at the base. Where the remiges are relatively long, the secondary quills attain their maximum height in or just above the middle of the feather. The functional meaning of this correlation is however not quite easy to make out, but physiologists and flight-engineers should pay due attention to the fact that such morphological peculiarities are always correlated to the mechanical or other demands.
In addition to these general remarks on secondary quills I must mention the peculiar structure of the margin of the outer vane of the three outermost remiges in owls, which are toothed or rather comb-shaped (figs. 31, 34 7). The finer structure of this comb how- ever seems not to have been understood; its teeth are nothing else than the tips of the secondary quills of the outer vane, which in this instance, do not bend inward terminally and lie close together, but, after extending in a straight line for some distance, suddenly bend outward, so that their ends stand almost at right angles to the direc- tion of the primary quill. Of course the separate teeth of this comb, as parts of the secondary quills, are provided with tertiary fibers (fig. 34). It is to be noted that this formation, which is absent in only few species of owls, as for instance in Nyctea nivea, a day bird of prey, is to be found also in some of the Caprimulgi (e. g., some species of Podargus, fig. 30, Z). On the other hand, they seem to be entirely absent in the genus Caprimulgus itself, nor do they occur in Stringops habroptilus, although it also is a night bird.
“ SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
5. The Tertiary Fibers
Between the secondary quills lie two systems of fine fibers, which in their close connection form the greater part of the feather plate that acts on the air in flight. The tertiary fibers spring from the upper part of both sides of the secondary quills, and form two little fiber-vanes (fig. 22). In a work by Schroeder (1880, pp. 3-14) there is a critical discussion of all papers on this subject published up to 1880. Nitzsch (1840, pp. 5-15) was the first to give a fairly correct description of these structures, with a number of good draw- ings. In 1887 Wray made an attempt to construct an enlarged model of a feather and from it prepared some very schematic drawings (TE87, pl. scr):
Among works of later date are Ahlborn’s (1896, pp. 17-21) and Strong’s papers (1902, pp. 156-161), to which and to a small num- ber of others I shall have occasion to refer. What has been said with regard to the nomenclature of the secondary quills holds good for the tertiary fibers also. Uniformly termed “ barbules”’ in Eng- lish and French, they figure in German works promiscuously as “ Strahlen,” “ Nebenstrahlen,’ “ Faserchen,” “ Fiedern II. Ord- nung,” “ Fiederchen,’ etc. We must distinguish two essentially different kinds of tertiary fibers, one being on the whole straight and bearing on the ventral side several hooks, the hook-fibers (fig. 21, Hkf); the others being curved and without hooks, the curved fibers (fig. 21, Bgf). This division of the fibers into two kinds seems to be justified also by the fact that they start from different sides of the secondary quills.
If we cut sections through the feather vane, according to the method before described, in the direction of the hook-fibers, or in that of the curved fibers, in the first case, between the transverse sections of every two adjacent secondary quills we get a longitudinal section of a hook fiber and a series of transverse sections of curved fibers, and in the other case, a longitudinal section of a curved fiber and a series of transverse sections of hook-fibers (figs. 24,25). The series of transverse sections of different parallel fibers thus obtained are identical with a series of transverse sections of the same fiber. Accordingly, we can reconstruct from these sections any particular fiber we like.
A. Hook-Fibers
The structure of the hook-fibers is as follows. The proximal por- tion is band-like, transversely curved so as to become groove-like, the concavity of the groove being turned towards the secondary quill, from which the fibers arise (fig. 26). This band-like basal portion
MASCHA] STRUCTURE OF WING—FEATHERS 13
is continued in a distal part which is more rod or thread-like, and from which numerous projections arise ; the proximally ventral hooks characteristic of these fibers, and the distally ventral and dorsal spines lying in pairs opposite each other.
We will first consider the hook-fibers of the remiges of Columba livia as a typical example (figs. 2, 26). Here the proximal band- like portion forms about half of the whole fiber. It is transversely bent, its upper half being vertical, the lower turned obliquely toward the secondary quill and the front. As transverse sections clearly show, the fiber is here not only groove-shaped, but thickest at its upper margin, decreasing in thickness downward and passing finally into a thin ventral membrane. Only at the proximal end of the fiber the ventral membranous extension is absent. In studying trans- verse sections through the secondary quills (figs. 3, 4, 26) we see that in the first two the hook-fibers are thickest at the base and become thinner towards the top; in the third section the fiber is nearly equally thick at the top and the bottom and in the next sec- tion the reversed relation, holding good for the remainder of the hand-like part, sets in. At the upper margin a distinctly recogniz- able swelling (figs. 8, 26, WIt) is to be seen, which generally dis- appears distally, where the lower part of the fiber band passes into the thin ventral membrane mentioned above. The dorsal swelling is sometimes, particularly when pigment is present, plainly visible even in surface views. The histological structure of this part of the hook-fiber is as follows: If we look at a hook-fiber from the side we sometimes notice in its proximal half a row of oval spots usually with dark margins, their long axes extending obliquely upward and backward. These spots were first noted as depressions, later authors recognized them as dried-up nuclei (fig. 9, k). Schroeder (1880, p. 30) was the first to state that each fiber consists of a single row of cells. This supposition found confirmation in the subsequent investigations of Klee (1886) and Davies (1889).
The nuclei of the cells forming the fiber lie in its thinner portion and are clearly visible. Apart from the nuclei, one can frequently perceive the cell limits also, either as fine, dark lines, as I found them especially in the hook-fibers of Cypselus apus (fig. 16, Zgr) after staining, or as clear, pigmentless lines separating the upper por- tions of the cells in which the greatest amount of pigment is con- tained (figs. 8, 9, 10). Strong (1902, p. 156) has fully described these cell limits and has drawn particular attention to the fact that they first extend from the upper margin obliquely forward and downward and suddenly turn in the proximity of the nuclei, being
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
continued obliquely downward from here toward the base of the fiber. At the end of the proximal band-like portion of the fiber the lower halves of these cell limits change their direction and form a straight line with the upper halves, which in the distal part of this portion of the fiber attains a convexity towards the base (fig. 16, Zgr). The transverse curvature is not the same throughout the proximal portion of the fiber, being slightest at the base and be- coming greater distally. The upper and lower part of the band enclose an angle which is very obtuse proximally and becomes more acute distally; at the distal end of this portion of the fiber it is nearly a right angle (fig. 26). The series of transverse sections show that the difference in thickness between the upper, vertical and the outwardly lower, nearly horizontal part becomes greater distally. The upper vertical and the lower horizontal part of the fiber separate distally. <A little below the spot where these separa- tions occur the vertical upper portion passes. into the thread-like, distal part of the fiber. In ‘Columba livia the lower, horizontal mem- branous part terminates in several large lobes narrow at the base and broadened distally like leaves (fig. 8, wuLl,). The upper por- tions of these lobes stand vertical and enclose with the under ones an angle of nearly 90°. Columba livia is the only bird in which I have met with such high development of these lobes (fig. 26, VnL,). In others the lobes appear as digitate processes, one or two in num- ber, turned down distally as in Cypselus, Diomedea, Nyctea, Podar- gus (figs. 9, I1, 12, 13, VnL) and others. Each lobe springs from a different cell and is to be conceived as a simple cell-diverticulum.
Hook-shaped extensions hang down from the lower side of the distal thread-like portion of the hook-fibers which forms the con- tinuation of the upper thickened portion of the basal part. The bases of these hooks enfoliate the fiber. The hooks themselves are band-like, twisted in their upper part, and terminate in a strong backward turned spine (fig. 8, K). While the greater part of the band forming the hook is thin its end is thicker and forms a sort of swelling which bears the short terminal point, which turns back- ward to form the hook. In Columba livia and in numerous other birds 4 or 5 hooks occur on each hook-fiber (figs. 8, 10). I found fewer hooks in Cypselus (fig. 11), Micropus melba, Macropteryx mystaceus, where only 2 to 4 are present, and more in Diomedea (fig. 9) where 6 to 8 are to be found on each hook-fiber. It is usually relatively broad, rarely long and slender, as in Diomedea, the Striges, and Caprimulgi (figs. 9, 12, 13); those nearest the base of the fibers are the shortest and are directed vertically downward.
MASCHA] STRUCTURE OF WING—FEATHERS 15
Each succeeding hook is a little longer and directed more obliquely forward. While further differentiations of the hooks in general do not occur, in Turacus albocristus and in Cuculus canorus, on the anterior margin of the proximal hooks I found one to three small spines which gave them a peculiar appearance (figs. 14, 15). In the proximity of the hooks the transverse sections of the fiber itself changes. They are regularly oval where the last lobes of the ventral membrane arise. Farther out they become curved again, but the curvature is here far slighter than in the proximal portion of the fiber and the concavity turned toward the opposite side. The hooks are ventral projections of the successive cells forming the fiber. Beyond the last hook the fiber becomes rapidly thinner and gives off upward and downward, not as Nitzsch (1840, p. 14) erroneously supposed laterally, pairs of spine-shaped projections and terminates in a thin thread of varying length. The spines are the most variable parts of the whole hook-fiber. They are paired projections of the cells which form the distal portions of the fiber, each cell having two, a ventral and a dorsal one. The ventral spines are as a rule longer than the dorsal. The spines extend obliquely outward, the upper ones upward, the lower ones downward. They are broadest at their origin, become thinner distally and terminate in fine points. The proximal, ventral spines lying next to the hooks are blunt and fre- quently slightly curved in the form of a hook. They are transi- tions between the hook-shaped cell processes and the spines with straight-pointed ends. While, however, in the region of the hooks the fiber cells have no dorsal processes, in the distal, spined part of the fiber the cells have dorsal as well as ventral processes (spines). The spines originate from the distal, broader side of the cells, which here have the shape of flattened cones and are attached to each other in such a way that the narrow, proximal end of each is inserted into the broad, distal end of the next foregoing. The first two proximal, dorsal spines sometimes become very large and attain a lobose shape. This is especially well developed in the Cypselomorphze and in most water birds (figs. 17, 22, W,). These lobes lie horizontal, are directed towards the margin of the feather and extend as far as the next hook-fiber. In the hook-fibers of the proximal portions of the secondary quills, these lobes are smaller and more pointed than in the distal portions, where they become larger and relatively broader. Fatio (1886, p. 257) says of the spines of the hook-fibers of the feathers of water birds in general that “they are very long and numerous and by their irregular arrangement make the feather- ing bulky and so afford protection against the water.” In the
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
microphotographs of the surface of a wing feather of Cygnus olor one sees the strongly developed dorsal spines very plainly (fig. 23, W,). Surface views show that each hook-fiber is curved in a horizontal plane, the convexity being turned towards the secondary quill from-which it arises. This curvature is confined to the central part of the fiber. The proximal and distal parts of the fiber remain straight and lie in the same straight line (figs. 17, 18, 19, 22). The upper, thickened margin, which is vertical in the proximal part of the fiber, becomes oblique in the neighborhood of the hooks, in- clining above toward the secondary quill; distally it resumes its former vertical position. A similar twist, more or less pronounced, is observed in the hook fibers of all birds with the exception of the owls and Caprimulgi, where the upper margin of the fiber is vertical throughout its whole length.
The ventral part of the proximal band-like portion of the fiber, which terminates in the processes of the ventral membrane, is in Columba about as long as the distal portion; in smaller birds like Fringilla and also in the Psittaci, the proximal part is longer than the distal; in the Striges and Caprimulgi the distal part greatly ex- ceeds the proximal in length (figs. 12, 13), this being one of the differences between the remiges of the diurnal and nocturnal birds.
The most striking peculiarity of the remiges of the owls is the down covering the dorsal side particularly of the proximal part of the broad inner vane, which gives them a velvety appearance. This velvet-like down consists of the lengthened distal portions of the hook-fibers. Fatio (1886, p. 257) says that they are long, slen- der and covered with numerous lateral spines (barbicels), which on account of their length and irregular arrangement make the fiber appear like a down feather. The proximal band-shaped part of the hook-fibers of the remiges of owls (fig. 12) has the same struc- ture as in other birds. The terminal processes of the ventral mem- brane are usually one, less frequently two in number. They are small, narrow and show a slight hook-shaped curvature. In the region of the long and thin fourth and fifth hooks, the fiber is very narrow, and here the cell limits, usually indistinct, are sometimes conspicuous enough. The most remarkable feature of the remiges of owls is the extraordinary development of the distal, thread-shaped part of their hook-fibers. These terminal threads consist, according to the position of the fibers, of a row of 10 to 50 cells, each one bearing one or two peculiar, long and very thin spines (fig. 12, W). The dorsal spines are not raised but lie horizontal, so that they point inward toward the primary quill of the feather. The ventral spines
ft |
MASCHA] STRUCTURE OF WING—FEATHERS 17
do not extend downward but raise themselves and point toward the margin of the feather.
The spines of adjacent hook-fibers therefore cross each other at an angle of about go°, and thus form a system of rectangularly crossing threads similar to that formed by the hook and curved fibers. It is this peculiar structure of the distal portion of the hook- fibers which gives to the remiges of owls their great softness.
If one reflects that the distance between the secondary quills amounts only to 250 to 300m while the hook-fibers reach a length of 2 mm., it becomes clear that the hook-fibers attached to one secondary quill project a good distance beyond the next one, and often reach the second, or even the third or the fourth secondary quill. The many long terminal threads of the hook-fibers thus forming several overlapping layers, together with their numerous rectangu- larly crossing spines, form a dense, felt-like mass on the dorsal side of the feather. The constituent parts of this mass are prevented from becoming deranged through exterior mechanical influences by an extremely interesting arrangement. If one cuts through the proximal portion of the feather vane parallel to the direction of the hook-fibers and slightly magnifies the section thus exposed, it is seen that the proximal band-shaped portion of the fiber les almost horizontal and that the distal part rises abruptly up at an angle of aon, 40. (figs: 12, 32).
The hook-fibers of the Caprimulgi are very similar (fig. 13) to those of the owls, but the proximal band-shaped part of the hook- fibers of their remiges is relatively short and the terminal threads considerably lengthened. There are three to five rather narrow hooks, the distal ones being considerably longer than the proximal. The limits of the cells forming the fiber are often clearly recognizable in the hook region on account of their being somewhat raised and also on account of their color. As the hook-fibers of the remiges of the owls, so also those of the remiges of the Caprimulgi are dis- tinguished from the hook-fibers of other birds by the peculiar de- velopment of their thread-like, distal part. In the hook-fibers of the first hand-remiges of Podargus humeralis, the proximal, band- shaped part is 200 », the terminal thread 2.3 mm. long, that is, more than eleven times as much. While, however, the terminal threads of the hook-fibers of the Striges bear a great number of spines dor- sally and ventrally, such processes are entirely absent on the terminal part of the Caprimulgus hook-fibers. There are indeed immediately beyond the hooks one or two dorsal spines and four or five ventral the first remex. In the lower proximal third of this vane
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
ones, but such nearly spineless hook-fibers are not met with in the remiges of any other birds. All the other conditions described as present in owls, such as the teeth on the outer vane of the first hand- remex, the down on the upper surface, the upraising of the terminal threads and the overlapping of the layers of the latter, are also met with in the remiges of Caprimulgi (figs. 13, 30). Clark (1894, pp. 569-570) arrived at the result that these two groups agree in being aquincubital, a character which he considers of importance, and that the number of rows, the arrangement of the coverts and the relative position in the hand-remiges is the same in both. On the other hand, the number of hand-remiges in the, Striges is always greater than in the Caprimulgi. To this we can now add that the Striges and Caprimulgi possess peculiar feather structures common to both through which they deviate from all other birds.
The differences between them are the different development of the ventral ridge of the secondary quills, which in Striges is high and sharp while in Caprimulgi it is short and blunt, and the different structure of the terminal threads of the hook-fibers, which in the former possess numerous spines, while in the latter they are nearly spineless. Common to both are the length and upraising of the terminal threads of the hook-fibers, which form the down on the dorsal side of the feather, and the comb-like structure of the anterior margin of the first hand-remex.
The hook-fibers spring at an angle of 30 to 40° from the dorsal part of the inner side of the secondary quill. All the hook-fibers of a secondary quill are parallel to each other and equally far apart. It is to be noted that the distance between adjacent hook-fibers is nearly the same in all birds, varying only from 20 to 30p. In Cypselus this distance is 25 p, in Columba 22p, and in Diomedea 27 p.
The hook-fibers arising from one secondary quill extend as a rule about as far as the next secondary quill, excepting that in the Striges and Caprimulgi they extend, as we have seen, much further, al- though unusually long hook-fibers occur in some other birds also. In Diomedea, for instance, the spined terminal threads of the hook- fibers extend to the second secondary quill. The hook-fibers are not of equal length in different parts of the wing of the same bird, and are different even in the different parts of the same feather. These local variations in form and size, met with in all birds, are particu- larly striking in Nyctea nivea. In this owl the hook-fibers on the broad inner vane are longer than on the outer vane. Considerable variations occur also in the vanes of different feathers. I will illus- trate this by a few measurements made in the inner broad vane of
MASCHA| ° STRUCTURE OF WING-FEATHERS 19
the hook-fibers are longest. Here the proximal band-shaped part of the fibers measures on an average 250 in length, the terminal thread which is composed of about fifty cells, 1.77 mm. and the whole hook-fiber 2.02 mm. The terminal thread is about eight times as long as the proximal band-shaped part. Towards the middle of this vane the proximal part of the hook-fibers becomes longer, measuring 300 to 350m, and the terminal thread, here composed of only about 30 cells, shorter, measuring only 7op, so that the whole fiber is only 1.27 to 1.32 mm. long, and the terminal thread about three times as long as the proximal part. Near the tip of the feather we find hook-fibers, the proximal part of which measures 200 » and the terminal thread, here composed of only 8 to Io cells, 270m. Here the whole fiber is 470 long and the terminal thread only slightly longer than the band-shaped proximal part. The hook-fibers also, arising from the same secondary quill, are not all alike. The proximal band-shaped part remains fairly constant but the terminal thread varies considerably. Let us take a secondary quill of the middle of the above-mentioned feather vane: The proximal fiber portion on the whole length of the secondary quill is 300 to 350 p long.
The terminal thread is shortest and composed of fewest cells in the fibers arising proximally, nearest the main quill. Distally the terminal thread lengthens and the number of cells composing it in- creases until just beyond the middle of the secondary quill; further on toward the feather margin it again becomes a little shorter.
In the anterior narrow vane of the same feather the hook-fibers are considerably shorter than in the broad inner vane. Measure- ment in the middle portion of the feather gives an average length of 600 » for the hook-fibers of the narrow vane as against 1.28 mm. for the hook-fibers in the corresponding part of the broad vane. The proximal band-shaped part is 300 long in the narrow vane, that is to say, just as long as the terminal thread. There are four or five hooks on the fibers of both vanes.
In this place I may mention a few developments of hook-fibers characteristic of the feathers of owls. The cells forming the termi- nal thread bear one or two spines each, but whether one or two depends upon the position of the fiber. In the middle of the narrow outer vane of the distal hand-remiges the terminal threads of the hook-fibers are composed of about ten cells each, and provided with ventral spines only. In the middle of the broad inner vane the terminal threads of the hook-fibers arising from the proximal parts of the secondary quills are composed of cells which likewise bear
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
only ventral spines ; these ventral spines are, however, extraordinarily long. In the terminal threads of the hook-fibers arising toward the middle of the secondary quill, spines appear on the dorsal side also. Where dorsal spines are developed the ventral ones are shorter than elsewhere. Below the middle of the secondary quill the dorsal and ventral spines are equal and about half as long as the ventral spines of parts where no dorsals are developed. The down becomes grad- ually thicker outwards and is thickest and highest beyond the middle of the feather-vane, measuring transversely, just where the termi- nal threads of the hook-fibers and the spines attain their greatest length (fig. 32). In some birds the proximal, band-shaped part of the hook-fiber is remarkably strong when compared with the termi- nal thread. In Cypselus the proximal portion is actually longer than in Podargus, the hook-fibers of which have a much greater total length than those of Cypselus.
B. The Curved Fibers
Opposite the hook-fibers and a little lower down there rises from the side of each secondary quill a second system of fibers, the curved fibers (figs. 3, 4, Bgf). At first they extend obliquely outward at an angle of 35 to 40°. At about half their length they suddenly bend toward the secondary quill from which they arise in such a way that their distal portion comes to lie parallel to the latter (fig. 21). Although many anatomical characteristics indicate that the curved fibers and the hook-fibers are homologous, yet these two kinds of fibers are in many respects morphologically and func- tionally so different that a special terminology and a separate de- scription are necessary.
In the curved fibers as in the hook-fibers two parts, a broader, proximal, band-shaped portion and a slender terminal thread can be distinguished (figs. 9, 28).
The fiber increases in breadth from its point of origin up to the middle of its length and then narrows again. In sections vertical to the secondary quill one can see (figs. 3, 4, 27), that the proximal ends of the curved fibers are band-shaped with a thickened ventral and a thin dorsal margin. Where the fiber leaves the longitudinal ledge on which its basis rests, the two margins become equal in thickness. Further on, the dorsal margin remains stout while the ventral one becomes thin, membranous and sharp. So far the curved fibers resemble the hook-fibers, but a difference between them can already be noticed here. In the proximal part of the curved fibers the strip forming the upper margin is not merely thickened, as in
MASCHA] STRUCTURE OF WING-FEATHERS 21
the hook-fibers, but appears involuted (fig. 28, R). Klee (1886, p. 18) was the first to show that the dorsal margin of the curved fiber is not thickened but involuted to form a groove in which the hooks of the hook-fibers are inserted. As before-mentioned these fibers are transversely bent in the shape of a groove. The concavity lies towards the secondary quill from which the fiber arises. It is most pronounced above and flattens out below, the section being an evolvent (fig. 27, .). Perhaps one may consider the slight thicken- ing of the upper margin in the proximal part of the hook-fiber a formation corresponding to the involution of the curved fiber (fig. 26, Wit). The sections show that the fiber is thickest at the dorsal involuted margin, decreases in thickness downward and thins out to a fine membranous lamella below. This lower lamellate portion has not the same curvature as the upper part but displays toward its margin an inclination to bend in an opposite direction so that the transverse section becomes slightly S-shaped. The curved fibers like the hook fibers consist of a single row of cells lying one behind the other. A row of nuclei extending obliquely upward and out- ward is clearly visible (fig. 27, K), and one can also make out the cell limits.
The ventral margin of the proximal part of the fiber is continuous up to the bend, or only interrupted by shallow incisions so as to appear-somewhat wavy. The dorsal involuted margin retains its direction, the diminution of the fiber in width toward the end is caused only by the falling back of the ventral margin. In the region of the bend three or four tooth-like projections are observed at the upper, here less strongly involuted margin of the fiber. These teeth are turned backward and pointed. The middle ones are usually best developed (fig. 27, Zf), and correspond to a certain extent with the hooks on the hook-fibers. In cases where the hooks are broad, the teeth also are large, while in the case of birds whose hooks are long and thin, the teeth are often very small and hardly discernible. The first case is exemplified by the Psittaci, Columba and Anseres, the second by Diomedea and the Striges. It is remarkable that these tooth-like projections, which are not particularly difficult to see, have hitherto been mentioned in only a single description of feathers (Wray, 1887, p. 421, pl. x, fig. 2). They have either quite escaped the other observers, or have appeared to them not worthy of notice. I believe that their function is far from unim- portant.
In and beyond this region the ventral margin is divided by sev- eral deep incisions into a number of processes. These processes are
ty bo
SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
triangular, pointed distally and sometimes slightly curved hook-like at the end (figs. 27, 28, wnL,). Ahlborn (1896, p. 20) mentions these processes in his description of the curved fibers and calls them “the finest saw-toothlets.” Strong (1902, fig. 25, pl. 5) has rep- resented them in his figure of a curved fiber. These processes of the curved fibers seem to correspond to and be homologous with the hooks of the hook-fibers. They are most highly developed in Diomedea where they are extremely narrow and terminate in long, fine points. In this region, where on the dorsal margin the teeth, and on the ventral the processes appear, the curved fiber is already considerably narrowed and it continues to grow narrower distally till it finally terminates in a long, fine thread (fig. 28, F). The point of transition of the band-shaped proximal portion into this terminal thread coincides with the completion of the bend. The ventral processes take no part in this bend and retain the direction of the basal part of the fiber unchanged.
We will now consider again the series of transverse sections. We have seen that in the proximal parts the upper margin of the fibers is involuted spirally and that the curvature decreases toward the lower margin, the concave band gradually unrolling itself, as it were, toward the latter. Toward the middle of the length of the fiber there is a change. The portion of the transverse section lying midway between the upper and the lower margins, which before was considerably curved first flattens itself out and then forms an obtuse angle the reverse way, so that here the section attains the shape of a 3 (fig. 27, X). At the same time the whole fiber is spirally twisted so that the axis of the transverse section assumes first a vertical position and then inclines above toward the secondary quill.
The distal termination of the curved fibers is very similar to the long, thin terminal thread of such hook-fibers as those of the Capri- mulgi. While in most cases the thread in which the curved fiber terminates appears as a simple filament without any differentiation, sometimes slight thickenings appear in it at about equal distances, which apparently correspond to the nuclei of the cells arranged one behind the other, which compose this portion of the fiber (fig. 27, V-). The terminal threads of the successive curved fibers are parallel with each other and lie close together (figs. 19, 21).
The curved fibers spring from that side of the secondary quill which is turned toward the feather base. Their points of origin lie lower than those of the hook-fibers (figs. 3, 4).
In length the curved fibers surpass the hook-fibers, particularly in
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MASCHA | STRUCTURE OF WING—FEATHERS 23
the terminal threads of the former which are always longer than those of the latter in the same part of the feather. In Nyctea nivea, for instance, the proximal portion of the hook-fibers is on an aver- age only 300 to 350, those of the curved fibers, g00 long. The shape of the curved fibers is very constant. With the exception of a few isolated cases, such as Diomedea, in which the ventral mem- branous processes display a peculiar structure, they are entirely similar in all birds, differing only in respect to their size, which is proportional to that of the whole feather. The number of the curved fibers is somewhat less than that of the hook-fibers. The reason for this is that they are 30 to 40» apart, which is a little greater than the distance between the hook-fibers.
6. The Formation of the Feather-Plate
The two kinds of tertiary fibers described above together form the greater part of the plate represented by the feather. As Schroeder (1880, p. 3) tells us in the historical part of his work, Marcellus Malpighi, the first investigator who occupied himself with the study of the feather, could only speak of an interweaving (amplicatio) of its smaller elements. Subsequent investigators de- clared that these elements were too small for exact study. Nitzsch (1840, pp. 14, 15) was the first to try to solve the problem. He pointed out that the hooks of the upper “ rays” (hook-fibers) were designed to grip the lower fibers. This they do by inserting them- selves into small depressions in the sides of the latter. These “ de- pressions ’ of Nitzsch’s are however not depressions at all but the nuclei of the cells composing the curved fibers. Burmeister in a note pointed out this error (Nitzsch, 1840, p. 15), and stated that the hooks were too short to reach these “* depressions ”’ of the curved fibers. According to him the hooks are intended to grip the upper margin of these fibers, which he considers thickened and which he says they just reach and actually grasp. Schroeder (1880, p. 10), and Klee (1880, p. 18) discovered that the upper margin of the lower (curved) fibers is not thickened, as Burmeister had thought, but involuted to form a groove, which makes the impression of a thickened edge in surface views of the feather magnified with the microscope. This involuted margin is grasped by the hooks of the hook-fibers, which can glide backward and forward beneath it with- out relaxing their grip. Wray (1887, p. 422) and Ahlborn (1896, p. 20) also describe the formation of the feather-plate. According to them the hooks of the hook-fibers penetrate into the layer of underlying curved fibers and take hold of the membranous ventral
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
processes of the latter. The incorrectness of this statement can easily be demonstrated by the examination of a surface view by which it is to be seen that in their natural position the hooks lie some distance beyond the zone of these ventral processes and in reality grasp the more proximal, band-like portions of the curved fibers. They are moreover too short to reach as far down as the ventral margins of the curved fibers from which these processes protrude horizontally. Finally, one must consider that if the hooks actually gripped these processes they would not be capable of gliding backward and forward, but would be wedged in firmly between them. According to my own observations the hook-fibers overlie the curved fibers and cross them at an angle of nearly go° (figs. 17, 18, 19). The hooks hanging down cling by their lower margin to the in- voluted upper margin of the curved fibers. All the hooks of one hook-fiber hold different curved fibers (figs. 24, 25), so that as many curved fibers are held by one hook-fiber and as many hooks grip each curved fiber as there are hooks on each hook-fiber. The hooks can glide backward and forward in the smooth groove of the upper margin of the curved fiber, a fact which is of great importance in flight. During each downward wing stroke a pressure is exercised on the lower side of the feather-plate. As Parseval (1880, p. 70) asserts, the angles between the secondary and the primary quills grow more obtuse under the action of this pressure, and the distance between the outer part of the secondary quills increases. At the same time the feather changes its curvature and its surface in- creases. These changes are made possible by the gliding of the hooks which connect the two systems of tertiary fibers. If the sec- ondary quills move apart, the hooks glide from the middle of the ribbon-like portion of the curved fibers which they clasp during rest, towards their distal end (fig. 19. See the direction of the arrow). If the pressure were still increased the hooks would glide still further and slip off the curved fibers, thus losing their hold altogether if it were not for the peculiar arrangement for arresting them, which effectually prevents this. The bending of the curved fiber itself resists such a slipping off and the dentate protuberances of this part of their upper margin (fig. 28, sf) described above, make it still more difficult. These teeth being directed backward it becomes easy at the same time for the hooks to glide back into their proper position, should they have been carried beyond by some unusual force.
MASCHA ] STRUCTURE OF WING—FEATHERS 25
III. SumMaAryY OF RESULTS
1. In the core of the secondary quills the cells may be so arranged as to form several irregular, or one single regular layer. The first type is the more frequent. The second is met with chiefly in the Striges and Caprimulgi. |
2. All secondary quills have a ventral hornridge. It varies be- tween a low crest (Cypselus, Columba) and a large, curved, mem- branous plate (Diomedea, Striges, Cygnus).
3. The degree of concavity of the secondary quill is correlated to the size of the central hornridge.
4. At the origin of the tertiary fibers from the secondary quills very complicated structures are met with. Here a projecting longi- tudinal ledge occurs from the upper side of which oblique crests protrude, parallel and close together like the bars of a gridiron. Between these crests lie the proximal portions of the tertiary fibers.
5. The diminution in height of the secondary quills from the main quill outward may be gradual, constant and moderate, or in- terrupted by a step, as in Striges and Cygnus.
6. The height of the secondary quills is greatest either at the feather base (Aquila, Diomedea) or more frequently above the mid- dle of the length of the feather (Cygnus, Bubo, Macropteryx).
7. The teeth of the outer feather-vanes of the first three remiges of the owls are peculiarly differentiated tips of secondary quills.
8. The hook-fibers always spring from the inner side of the sec- ondary quill which is turned toward the main quill and the feather- tip. They are transversely curved bands composed of a single row of cells. These cells may possess projections which are, proximally, dorsal lobes and ventral hooks; distally, dorsal and ventral spines.
g. The number of hooks on each hook-fiber is 2 to 8; it is con- stant in the same species.
10. Peculiarly shaped hook-fibers are found in the Striges and Caprimulgi. In the former they are very long and provided with numerous large spines; in the latter likewise they are long, but have no distal spines.
11. The distance between the hook-fibers in the functional remiges of all classes of birds varies only between 20 and 30 p.
12. The curved fibers spring from the outer side of the secondary quill which is turned away from the primary quill and toward the feather basis. They are transversely curved bands, like the hook- fibers which they resemble in many points.
13. The dorsal, dentate projections at the bend of the curved fibers are an arresting apparatus.
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
14. The distance between the curved fibers in the functional remiges of all classes of birds varies only between 30 and 40 p.
15. The powerful development of the ventral hornridge of the secondary quills in Diomedea and Aquila is doubtless functionally related to the sailing flight of these birds.
16. The teeth on the outer vane of the first hand remiges in the Striges and Caprimulgi, and the down on the upper surface of the remiges of these birds formed by the lengthened, distal portions of the hook-fibers deaden the sound of the flight of these night birds.
IV. List or AuTHORS CONSULTED AND QUOTED
Ahlborn, F. 1896 Zur Mechanik des Vogelfluges. Hamburg, 1896. Beddard, F. E. 1898 The Structure and Classification of Birds. London, 1808. ° Clark, H. L. 1894 The Pterylography of certain American Goatsuckers and Owls. Proc. U. S. Nat. Mus., v, 17, pp. 551-572. Clement, C. 1876 Note sur la Structure microscopique des Plumes. Bull. Soc. Zool. France, I, 1876, pp. 282-286. Cuvier, G. 1809 Lecons d’Anatomie comparée. Recuillées et publiées par C. Dumeril, TTP. 603 (ck Mem: id) Wins). 13')330)). Davies, H. R. 1889 Die Entwickelung der Feder und ihre Beziehung zu anderen Integu- mentbildungen. Morph. Jahrb., Bd. 15, pp. 560-645, Taf. 23-26. de Meijere, J. C. H. : 1895 Ueber die Federn der V6gel, insbesondere tiber ihre Anordnung. Morph. Jahrb., Bd. 22, pp. 562-591. Fatio, V. 1886 Des diverses modifications dans les formes et la coloration des plumes. Mem. Soc. Phys. et Hist. Nat. Genéve, T. 18, Part 2, pp. 249-308. 3 pls. Gadow, H. 1882 On the color of Feathers as affected by their Structure. Proc. Zool. Soc., London, pp. 409-421, Pl. 27, 28. Haecker, V. 1890 Ueber die Farbe der Vogelfedern. Arch f. mikr. Anat., Bd. 35, Hit. I, pp. 68-87, Taf. 4. Haecker, V., und Meyer, G. 1901 Die blaue Farbe der Vogelfeder. Zool. Jahrb., Abth. Syst. Geog. und Biol. d. Tiere, Bd. 15, Hft. 2, pp. 267-294, Taf. 14. Heusinger, C. F. 1882 System der Histologie. Bd. 2, p. 207.
MASCHA | STRUCTURE OF WING-FEATHERS ay,
Holland, Th. 1868 Pterologische Untersuchungen. Cabanis, Journal f. Ornith., Jhg. 12, pp. 195-217. Klee, R. 1886 Bau und Entwickelung der Feder. MHaller’sche Zeitschrift f. Naturw., Bd eetonCNeeh., sd. 12), Eitt 1,° pp: 174—327. Mascha, E. tg0o2 Der Bau der Fltigelfeder. Verh. Ges. D. Naturf. u. Aerzte, 74, Vers. 2, Th. 1, H, pp. 159-162. Nitzsch, Ch. L. 1840 System der Pterylographie. Herausgegeben von H. Burmeister, Halle, 1840, 10 Taf. Parseval, A. von. 1889 Die Mechanik des Vogelfluges. Wiesbaden, 1880. Schroeder, R. 1880 Pterographische Untersuchungen. Diss. Inaug., Halle, 188o. Strasser, H. 1885 Ueber den Flug der Vogel. Jenaische Zeitschr. f. Naturw., Bd. 19 CNE EB). Gitt. 1,° pp. 174-927. Strong, R. M. 1902 The development of color in the Definitive Feather. Bull. Mus. Comp. Zool., v, 40, pp. 147-185. Pl. 9. Sundeval, C. J. 1843 Om Foglarnes vingar. K. V. Akad. Handlingar, 1843, Isis, v. Oken, Hit. 5, 1846. Wray, R. S. 1887 On the Structure of Barbs, Barbules and Barbicels of a typical pen- naceous Feather. Ibis, 1887, pp. 420-423, Pl. x11.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
EXPLANATION OF FIGURES.
1. Three transverse sections of a secondary quill of Columba livia. a at the base close to the primary quill; b in the middle; c at the tip on the outer edge of the broad feather vane. Magnified 1: Ioo.
' 2. Three transverse sections of a secondary quill of Caprimulgus europaeus. a at the base close to the primary quill; b in the middle; c at the tip near the outer margin of the broad feather vane. Magnified 1: 100.
3. Transverse section of a secondary quill of Columba livia from the outer margin of the proximal feather vane. Magnified 1: 600.
4. Transverse section of a secondary quill of Caprimulgus europaeus, from the middle of the proximal feather vane. Magnified 1: 600.
5. Section through the feather vane vertical to the secondary quills; through the wing covert of Garrulus glandarius. Magnified 1: 190.
6. Section through the narrow vane of the proximal part of the hand-remex of Columba livia, vertical to the secondary quills. Magnified 1: 25.
7. Section through the proximal part of the broad vane of a hand-remex of Caprimulgus europaeus, vertical to the secondary quills. Magnified 1: 8o.
8. Hook-fiber of Columba livia. Magnified 1: 220.
9. Hook-fiber of Diomedea exulans. Magnified 1: 125.
10. Hook-fiber of Columba livia. Magnified 1: 125.
11. Hook-fiber of Cypselus apus. Magnified 1: 125.
12. Hook-fiber of Nyctea nivea. Magnified 1:125.
13. Hook-fiber of Podargus humeralis. Magnified 1: Ioo.
14. Hook-bearing part of a hook-fiber of Turacus albocristatus, showing the spines of the proximal hooks. Magnified 1: 480.
15. Hook-bearing part of a hook-fiber of Cuculus canorus, showing the spines on the proximal hooks. Magnified 1: 480.
16. Proximal portion of a hook-fiber of Cypselus apus, showing the cell boundaries and the swelling on the upper margin. Magnified I: 400.
17, 18, 19. Surface views of the upper side of a hand-remex of Cypselus apus, focussed at three different levels. Magnified 1: 270.
17. Highest level showing the terminal threads of the hook-fibers. 18. Middle level showing the proximal portions of the hook-fibers. 19. Lowest level showing the curved fibers.
20. The points of origin of the hook-fiber in a hand-remex of Columba livia, seen from above. Magnified 1: 400.
2t. A secondary quill with hook and curved fibers of Columba livia. Mag- nified 1:60.
22. Hook-fibers of a secondary quill of Columba livia. Magnified 1: 110.
23. Surface view of the upper side of a hand-remex of Cygnus olor, showing the terminal parts of the hook-fibers. Magnified 1: 270.
24, 25. Schematic representation of the connection of the hook and curved fibers.
MASCHA] STRUCTURE OF WING—FEATHERS 29
24. Section parallel to the curved fibers. 25. Section parallel to the hook-fibers.
26. Section of a hand-remex of Columba livia, parallel to the curved fibers. Magnified 1: 225.
27. Section of a hand-remex of Columba livia, parallel to the hook-fibers. Magnified 1: 450.
28. Curved fibers of Columba livia. Magnified 1: 270.
29. Transverse section of a secondary quill near the primary quill of Bubo maximus. Magnified 1: 100.
30. Outer vane of the ninth hand-remex of Podargus humeralis, showing the dentate margin. Magnified 1:2.
31. Outer vane of the ninth hand-remex of Bubo maximus, showing the dentate margin. Magnified 1:2.
32. Section through the outer portion of the broad feather vane in a hand- remex parallel to the curved fibers of Bubo maximus, showing the down on the upper side. Magnified 1:12.
33. Section through the middle of a hand-remex of Bubo maximus, perpen- dicular to the secondary quills. Magnified 1:7.
34. The teeth of the margin of the outer feather vane of the tenth hand- remex of Bubo magellanicus. Magnified 1:70.
30
SMITHSONIAN MISCELLANEOUS COLLECTIONS
Key To LETTERING, WHICH APPLIES TO ALL FIGURES.
Curved fibers.
Terminal thread.
Ledge below the point of origin of the tertiary fibers. Hooks of the hook-fibers.
Hook-fibers.
Ventral hornridge of the secondary quill.
Horny substance.
Primary quill. «
Nuclei of the cells forming the tertiary fibers.
Crests on the ledges of the secondary quills. Medullary substance.
Groove in the upper margin of the curved fibexs. Dorsal thickening of the secondary quills.
Processes of the ventral membrane of the hook-fibers. Processes of the ventral membrane of the curved fibers. Spines of the hook-fibers.
Dorsal lobes of the hook-fibers.
Teeth on the outer edge of remiges.
[voL. 48
Dentate projections on the dorsal margin of the curved fibers.
Limits of the cells composing the tertiary fibers.
Siw? , oe
Fas als 5 aR RS ae Rat Mae eae a NEOUS COLLECTIONS mie VOL. 48 PL. I
x, Three transverse sections of a secondary quill ot Columba livia. a at the base close to the primary quill; din the middle; c at the tip on the outer edge of the broad feather vane. Magnified 1:100
2. Three transverse sections of a secondary quill of Caprimulgus ecuropeus. aat the base close to the primary quill; 4 in the middle; ¢ at the tip near the outer margin of the broad feather vane. Magnified 1:100
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3. Transverse section of a secondary quill of Columba livia from the outer margin of the proximal feather vane. Magnified 1:600 /
4. Transverse section of a secondary quill of Caprimulgus europaeus, from the middle ot the proximal feather vane. Magnified 1:600
VOL. 48, PL. Ill
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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VOL. 48, PL. IV
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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VCL. 48, PL. V
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 48, PL. VI
125
Magnified 1
12. Hook-fiber of Vyctea nivea.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 48, PL.VII
13
100
Magnified r
13. Hook-fiber of Podargus humeralis,
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. VIII
16
14. Hook-bearing part or a hook-fiber of Turacus albocristatus, showing the spines of the proxi- mal hooks. Magnified 1:480
15. Hook-bearing part of a hook-fiber of Cucudus canorus, showing the spines on the proximal hooks. Magnified 1:480
16. Proximal portion of a hook-fiber of Cyfse/us apus, showing the cell boundaries and the swell- ing on the upper margin. Magnified 1:400
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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, FL. IX
17, 18, 19. Surface views of the upper side of a hand remex of Cyfse/us afus,, focussed at three dif- ferent levels. Magnified 1:270
17 Highest level showing the terminal threads of the hook-fibers
18. Middle level showing the proximal portions of the hook-fibers
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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. X
19. Lowest level showing the curved fibers 20. The points of origin of the hook-fiber in a hand-remex of Columba livia, seen from above. Mag- nified 1:400
a®
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. XI
21. Asecondary quill with hook and curved fibers of Columba livia Magnified 1:60
22. Hook-fibers of a secondary quill of Columba livia. Magnified 1:110
23. Surface view of the upper side of a hand-remex of Cygnus olur, showing the terminal parts of the hook-fibers. Magnified 1:270
VOL. 48, PL. XII
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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VOL. 48, PL. XIII
ISCELLANEOUS COLLECTIONS
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VOL. 48, PL. XIV
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30. Outer vane of the ninth hand-remex of Podargus humeradis, 31. Outer vane of the ninth hand-remex of Budo maximus.
howing the dentate margin.
Magnified 1:2 showing the dentate margin.
Magnified 1:2
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, FL. XV
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32. Section through the outer portion of the broad feather vane in a hand-remex parallel to the curved
fibers of Bubo maximus, showing the down on the upper side. Magnified 1:12 33. Section through the middle of a hand-remex of Budo maximus, perpendicular to the secondary
quills. Magnified 1:7
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. XVI
34. The teeth of the margin of the outer feather vane ot the tenth hand-remex of Budo magellanicus.
Magnified 1:70
<a
PI BE Nuns
THES TARPON AND LADY—FISH - AND: -THEIR RELATIVES
By THEODORE GILL
Data respecting the habits of the tarpon and lady-fish are here brought together from widely distant sources and the little infor- mation at hand respecting the habits of their relative, the Ptero- thrissus of Japan, is added. The statements here collected, it is hoped, may serve as hints to observers respecting facts to be looked for in the biology of those fishes.
The morphological or rather chief osteological characteristics of the types here considered have been recently investigated and eluci- dated by Dr. W. G. Ridewood in a weli illustrated article ““On the Cranial Osteology of the Fishes of the Families Elopide and Albu- lide”? published in the Proceedings of the Zodlogical Society of London for 1904 (vol. II, pp. 35-81). The illustrations are so good and so much to the point that they are here reproduced. They show well the distinctive characterers of the genera as well as the common characteristics which evince the relationship of the Elopids and Albu- lids and distinguish them from the Clupeids to which the former have so strong but illusive likeness.
Tue TARPON AND ITS FAMILY
One of the most remarkable of the families of fishes is that of the Elopids, and of that family the tarpon of the Floridian waters is the most notable. Yet comparatively little is known of the habits of any of the species. Much—very much—has been written about the tarpon, but most of it has been of a personal or subjective nature and not about the fish itself. To elicit new facts and indicate desi- derata is the object of the present article. What is known may be briefly summarized.
I
The family of the Elopids (Elopidz) is composed of a few living fishes which have much superficial resemblance to the herring fam- ily; they have a compressed fusiform body, covered by smooth sil- very cycloid scales; the head is bony and scaleless; the mouth and jaws nearly like those of Clupeids and more or less oblique ; the dor- sal is submedian and the other fins are essentially like those of the
31
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
herrings. The distinctive characteristics are the very numerous (22-30) branchiostegal rays, an unpaired gular plate or intergular plate or bone (also called “ jugular plate’’) between the rami of the lower jaw, and the development of the parietal bones so that they connect along the middle of the skull and consequently superficially separate the frontal bones from the supraoccipital ; the supramaxil-
1 if d f] ! ' PE PE pV CUE OSITE: BIA
Fic. 1.—Elops saurus; skull from right side. (After Ridewood.)
ope
: g XN ! RY / (Ze Fic. 2—Megalops cyprinoides ; skull from right side.
Figs. 1 and 2.—Elopoid skulls showing trend of mouth, composition of supramaxil- laries (sm.), shape of circumorbital bones (cor.), etc. (After Ridewood.)
GILL] THE TARPON AND LADY—FISH 33
laries are very large, each composed of three pieces, and mostly outside of but adjoining the suborbitals (cor) ; the circumorbitals are peculiarly modified, there being a well developed preorbital, fol- lowed by a narrow suborbital, above the supramaxillary, and then by broad ones beginning above the hinder portion of the supra- maxillary and continued back of the orbits. The parasphenoid bone is narrow.
The family of the Elopids, like that of the Chirocentrids, is a decadent one—one of the past rather than of the present. It was represented by numerous genera and still more numerous species during the Cretaceous epoch. Some of those were of large size, even exceeding the recent tarpon in dimensions, and almost all of them became extinct by the end of that period. The family was far less conspicuous during the Tertiary epoch, but as early as the Lower Eocene the still existing generic types Elops and Megalops made their appearance. At least remains of fishes found in the London Clay have been referred to these genera by A. Smith Woodward. Their later tertiary history is unknown.
The living species are few in number—only four—and belong to two very distinct groups which are usually considered the only genera—Elops and Megalops. These are distinguishable by a num- ber of important characters.
II
i
; ! 7 ' } 1 / ! t
P&P Lop SY g
Fic. 3.—Elops saurus; hyopalatine arch, opercular bones, etc., with mandible ; left side, mesial aspect. j, jugular plate, dorsal view. (After Ridewood.)
Elops, the genus which has given name to the family, has small scales, a small head, lower jaw not projecting, pseudobranchiz
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
large, an oblong dorsal fin with the last ray not produced, and the anal fin small. Further, the head is slender and pike-like, the mouth not very oblique, the lower jaw especially slender and its articulation far behind, under the postorbitals, and the suspensorium angular, the hyomandibular being inclined backwards. It contains two spe- cies, the long known and wide-ranging Elops saurus, and the local- ized Elops lacerta of the Congo and western Africa.
The Elops saurus is common in the open sea along the coast of the southern United States and is best known as the ten-pounder, though it has received many other names.
The accepted name was current at least as early as the seven- teenth century for Dampier, in his “ Voyages to the Bay of Cam- peachy,” for 1676 records (p. 71) “ten-pounders” among the fishes (including tarpons, parricootas, etc.) he found in “ the lagunes, creeks and rivers.” ‘“ Ten-pounders,” he adds, “are shaped like mullets, but are so full of very small stiff bones, intermixt with the flesh that you can hardly eat them.”
The species needs no further attention for the present at least.
Ill
Fic. 4—Megalops cyprinoides; hyopalatine arch, opercular bones, and mandible of left side, mesial aspect. j, jugular plate, dorsal view. (After Ridewood.)
The genus Megalops has very large scales (30 or more along the lateral line), the head is comparatively large and the lower jaw projecting ; there are no pseudobranchiz ; the dorsal fin is inserted more or less behind the ventrals and its last ray is produced into an elongated “filament.” In contrast also with Elops the head is short
GILL] THE TARPON AND LADY—FISH 35
and oblong by reason of the very oblique mouth, the lower jaw is abbreviated, truncated in front, and with the articular fosse under the eyes, and the suspensoria are each continuous, the hyomandibulars being inclined forwards.
Two very distinct forms are known—so distinct indeed that they have been referred to different genera—the Megalops cyprinoides of the Indian Ocean and northern Australia and the Megalops atlanticus or celebrated tarpon of America.
The tarpon (Megalops atlanticus) has an elongated fusiform shape ; the forehead slightly incurved (rather than straight) to the snout; the chin projects and is obliquely truncated ; the dorsal (with ‘twelve rays) is on the posterior half of the body, nearly midway between the ventrals and anal; its free margin is very sloping and incurved and its long hind ray reaches nearly to the vertical of the anal; the anal (with twenty rays) is about twice as long as the dorsal and falciform; the caudal fin has a very wide V-shaped emargination. The scales are in about forty-two oblique rows. It reaches a length of about six feet—sometimes more.
IV
The oldest form of the name seems to have been Tarpon; such is the guise it has in Dampier’s “ Voyages to the Bay of Cam- peachy” in 1675, and in Roman’s “Concise Natural History of Florida” (1775). Dampier found that “the fish which they take near the shore with their nets are snooks, dog-fish and sometimes tarpon. The tarpon,” he says, “is a large scaly fish, shaped much like a salmon, but somewhat flatter. *Tis of a dull silver color, with scales as big as a half crown. A large tarpon will weight 25 or 30 pounds. ’Tis good, sweet, wholesome meat, and the flesh solid and firm. In its belly you shall find two large scallops of fat, weighing two or three pounds each. I never,” continues Dampier, “* knew any taken with hook or line; but are either with nets, or by striking them with harpoons, at which the Moskito-men are very .expert.” Such are the ideas of the fish gained by Dampier in its southern resorts. How different they are from those now prevalent in the United States will appear hereafter.
The name in most general use is tarpon and this may be con- sidered to be the literary and accepted phase. Tarpum was also an early form, but is now obsolete. Along the Texan coast Savanilla is still in general use, but is gradually being superseded by tarpon on account of the influence of anglers. The apt descriptive name Grande-écaille (pronounced grandykye and meaning large-scale) was
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
given by the French settlers of Louisiana. Other names of still more limited use are silver-fish (Pensacola), and jewfish (Georgia and parts of Florida). Jewfish it shares with many other fishes, and another fish of Florida, a gigantic Serranid, is better known by the term. Silver-king is a euphemistic designation. Caffum is a name current in the island of Barbados.
V
The tarpon may be briefly defined as a littoral fish of warm Ameri- can seas often entering into rivers and acclimated in some inland lakes.
The boating excursionist along some favored shore of Florida or Texas during the spring and summer months at least—perhaps dur- ing all but the winter months—may be startled by the sudden pro- jection from the water of a silver-like mass, which, after describing a low arch, will splash into the water again at a distance of maybe twenty feet from the starting point; that mass is the tarpon, or the “silver king.” Florida and Texas are the states in whose waters the fish is most freqently seen, because there most looked for, but its range extends far beyond those coasts in all directions. In summer wanderers visit the north as far as Massachusetts, where large individuals of the “ big-scale fish,” as they are there called, are “taken every year in traps at South Dartmouth” in the “ latter part of September’; southward they may be found in Brazil and sporadically in Argentina. Around all the islands of and in the Caribbean Sea and Gulf of Mexico schools may be met with. Further, immigrants have found their way into rivers that enter into the tropical seas and the Lake of Nicaragua has long been famous as the home of the species.
Being essentially a warm water fish, it is only in the warm months that the tarpon is to be found at its northern and southern limits. On the approach of cold weather it retires towards the tropics. Along the southern Floridian coasts some “ appear in February, in- creasing rapidly in numbers in March, April and May”; in Texas, “early in March.” At first they refuse the bait but “during the latter part of May and in June” bite freely. ‘ About the first of December ” they “ disappear entirely’ from the Texan waters. In the tropical seas they may be found always, and about Tampico, in Mexico, their “season is from November first to April, the time when the tarpon practically disappears from Florida and Texas.”
The tarpon is sensitive to sudden changes of temperature and especially to cold, and to such changes it is sometimes subject in
GILL] _ THE TARPON AND LADY—FISH 37.
its northern range. During a cold wave which invaded Florida towards the end of January (26-27) 1905, according to a letter of E. J. Brown in “ Forest and Stream,” “the tarpon especially were affected by the cold.” There were brought to “ Lemon City between forty and fifty tarpon which had been so benumbed by the cold as to be easily speared by parties who were searching for them. The largest fish was in length seven feet one and three-quarters inches, girth thirty-nine and three-quarters inches, weight one hundred and ninety-four pounds. Several others were nearly this size. . . . The tarpon were salted, to be sent to Key West market, where there is a ready demand for them.”
VI
That the tarpon is a most active fish may be inferred from its form which is especially adapted for swift and enduring action. Its life is spent in the enjoyment of its power and in pursuit of food ; a carnivorous fish, it preys “eagerly upon schools of young fry, or any small fish that it is able to receive into its mouth, and in pursuit of which it ascends fresh-water rivers quite a long dis- tance.” The schools of mullets contribute largely to the great fish’s supply. Such it attacks by darting upon them and generally seizing them tail foremost. Its frequent leaps into the air, like those of the salmon, seem to be mostly in sportive manifestation of its intense vitality and not for food or entirely from fear. C. F. Holder tells that one leaping tarpon “fell headlong” into a “boat, passing through the bottom ”’; that another leaped over man and’ boat; and that still another sprung up to the “ deck of a steamer” and “ fell headlong into a passenger’s lap.” Other wonderful tales are told of the activity of the tarpon. According to Holder (at second hand from another), a fish made an “ initial leap of twelve feet ” and fol- lowed this up “ with six leaps all equally high.’ The same observer believed that “ the ordinary height a tarpon leaps is from seven to eight feet.” While leaping, its gill-covers are frequently spread out and its blood-red gills visible. Withal it sometimes goes into very shallow water and seeks out a quiet nook in which it may rest, “perfectly stationary,” for quite a long time.
VII
The life history is very imperfectly known, but it does not appear to breed at any place along the continental coast of the United States, for none except large individuals have been recorded from those places most resorted to by anglers. For a very long time one of thirty pounds weight was the smallest obtained in Florida and
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
one of eleven pounds in Texas. It apparently demands a tempera- ture and conditions which the reef-forming coral animals require and sheltered brackish or fresh water for oviposition. In such locali- ties about Porto Rico, in February, 1899, Evermann and Marsh found not eggs, but very young, and there “it evidently breeds.” Thirteen fry, “2.25 to 3.25 inches ” long, were collected at Fajardo; at Hucares, “in the corner of a mangrove swamp” in “a small brackish pool of dark-colored water,” “entirely separated from the ocean by a narrow strip of land, four from 7.5 to 11.5 inches long were seined.”’ The smallest previously known was about nine inches long. All these are probably the young of the first year.
The very young or larvze will doubtless be found to be, like those of Elops and Albula, elongate ribbon-like animals of translucent and colorless texture, with a very small head and small fins. They are probably so transparent that their eyes alone are apparent in the water unless a very close examination is made. The youngest of the specimens (2.25 inches long) observed by Evermann and Marsh, were probably not long before developed from the larval condition. Such are the little fishes to be looked for as the very young of the great tarpon.
Most of the large tarpons caught along the coasts of Florida and the Southern states have attained full maturity; of such the length is about six feet, and the weight approximates one hundred pounds ; they are probably nearly or over three years old. Growth, however, is continued in some much beyond the average, one of three hundred and eighty three pounds, it is claimed, having been harpooned.
WILT
“The silver-king is the greatest of game fishes.” So declare Evermann and Marsh, and they echo the belief of many. Volumes and countless articles in periodicals have been devoted to detail of its excellencies. Its activity and gameness are proportioned to its size. The northern salmon affords tame sport compared with the “ silver-king.” Those of the average full-grown size (six feet long and one hundred pounds in weight) are caught in numbers with the rod and line; one weighing two hundred and twenty-three pounds closes for the time the record of feats with the rod, and it took the captor “three hours and a half before it was brought to gaff.”
The tarpon is now considered to have little or no edible value. It has, indeed, been declared by Schomburgk to be “considered a delicate eating”? in Barbados, and in the United States has been experimented with occasionally ; one (W. H. Burrall) who did so in
GILL] THE TARPON AND LADY—FISH 39
1874, declared (in Forest and Stream, 1, p. 324) that it was very palatable, but his taste was exceptional. It has been frequently tried since but rejected for the table. An effort was made on one or two occasions in Massachusetts when considerable numbers had been caught, “to find a market for them,” as at New Bedford, ‘ but the people did not like them, owing to the toughness. of the flesh.” Holder’s negro oarsman aptly replied to the suggestion that it was “the finest looking fish in the world,’ “ Yes, Sa, hit looks fine, so does hay. I'd rather eat hay dan tarpon, yes, Suh, I would.” It is truly, as Holder remarks, almost the only great game fish “ which is utterly scorned as a food fish.” Dampier’s opinion, expressed in 1675, and that of some Barbadians, has not been adopted by modern gourmands. It is “full of numerous small bones, which is a great inconvenience,’ says Schomburgk. In almost all cases where it has given anything like satisfaction the fish was of small size, and the truth may be that small ones are tender and savory but large ones coarse and tough, like overgrown individuals of other species. The results of unprejudiced judgment are still wanting.
It may be recalled here, however, that the Indian congener of the tarpon, the ox-eye (Megalops cyprinoides) is, according to Saville Kent as well as others, “highly esteemed for food,” and in the Malay archipelago, where it likewise abounds, it is cultivated in tanks after the same manner as the milk-fish, Chanos salmoneus.
Far from being sought by the fisherman for the market, the tarpon is detested by him. ‘‘ The Pensacola seine fishermen dread it while dragging their seines, for they have known of persons having been killed or severely wounded by its leaping against them from the seine in which it was enclosed. Even when it does not jump over the cork line of the seine, it is quite likely to break through the netting before landing.” Nevertheless even a dead tarpon yields some compensation for the trouble he gives. There is quite a demand for its great beautifully silvered scales, some of which may be as large as a lady’s palm. They find customers who are willing to pay as high as from five to twenty-five cents apiece and they are made up in various ways to attract the winter visitors to Florida.
IX
A species congeneric with the tarpon, but not very closely related, is the Megalops cyprinoides which, indeed, is the type of the genus. It is a less slender fish and the outline of the back and head is dif- ferent from that of the tarpon; further, the dorsal fin is not so far backward, that fin and the anal have more rays (dorsal, 19 to 21;
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
anal, 24 to 27), and the proportions of all the fins are more or less different. The size, also, is never so great as in the giant tarpons, for it rarely, if ever, attains to a length of more than five feet.
Like the tarpon, the Asiatic fish readily accommodates itself to fresh water. According to H. S. Thomas (1897), in India “ they acclimatize very readily to fresh water, and grow fast,’ and also breed, he was told, “in ponds.” The natives, too, “are fond of keeping them in ponds.”
They are more prone to associate in schools or shoals—that is, close together like herring—than the tarpon, especially when young. Thomas came “ across them coming up an estuary in a shoal, and it was like hauling in mackerel; and they run about the same size. There was a fish on as fast as ever you could get your line in the water. But the fun was very short-lived. It was in mid-stream, and they were all past the boat in a very little time.” Thomas took them “on a May fly and a Carnatic Carp fly.” In “30 minutes,” “on a light trout rod,” he “ took six of three-quarters of a pound each, lost four among weeds, and had one fly bitten off. Some of them sprang a foot in air, and all fought well.”
The fame of the tarpon has, in recent years, been reflected on its eastern relative and the lesser species has found advocates for its pursuit as a game fish. “ Enthusiastic anglers disposed to initi- ate” angling for it as for the American fish are referred by T. Saville Kent (1897) to the Badminton Magazine for 1895 for in- formation. “ There can be no doubt, in the writer’s opinion,” that the Australian fish, popularly known as the ox-eye herring, possesses “the most conspicuous potentialities for sport,” and “would yield equally exciting sport on the same lines.” Unlike its American relative, too, there might be the after satisfaction of seeing it on the table for, according to Kent, the ox-eye affords “ most excellent eating.” In India, it is raised to some extent for the table in tanks.
Tue Lapy-FisH
The Albulids are unique in the development of two transverse rows of valves in the bulbus anteriosus in advance of the heart, in which respect there is an approximation to the Ganoids. The form and physiognomy are peculiar but there is more superficial resem- blance to a whitefish (Coregonus) than to a herring ; the body is fusi- form but the dorsal arch much more curved forwards than the ven- tral; the scales are cycloid and brilliantly silvery ; the head bony and scaleless, the snout prominent forwards; and the supramaxillaries are moderate, composed of only two pieces, and partly retractile
GILL} THE TARPON AND LADY—FISH Al
under the large preorbitals; the mouth is small and overhung by the prominent snout; the circumorbitals strongly contrast with those of the Elopids, a large horizontal preorbital being developed along
Fic. 5—Albula conorhynchus ; skull from right side. Showing the peculiar snout (me.), circumorbitals (cor.), ete. (After Ridewood.)
the side of the snout and this is followed by a short and equally wide suborbital, itself succeeded by wide, angular and postorbital bones, having a well defined sensory canal. According to Ridewood “there are in all twelve bones of” the circumorbital series. “The most anterior ones are curious, basket-like bones, not much wider than the sensory canals which they carry. The canals in this region are particularly large.’ There is no gular plate. The parasphenoid
Fic. 6.—Albula conorhynchus ; hyopalatine arch, opercular bones, and mandible. left side, mesial aspect. Showing the peculiar dental pavement of the entoptery- goid bones, lower jaw, etc. (After Ridewood.)
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
bone is broad. Further the family is characterized by the villiform jaw and palatal teeth, pavement-like dentition of the hinder part of the mouth and wide parasphenoid bone.
is
The Albulids, like the Elopids, are an ancient family and repre- sentatives have existed from Cretaceous times to the present, al- though the affinities of the extinct forms have not been precisely determined. Remains of a fish that lived in the Lower and Middle Eocene and that was formerly called Pisodus owenw have been re- ferred even to the genus Albula. In the present seas there is only one genus, Albula, represented by a single species, Albula vulpes.
It is found in almost every tropical sea, but it is not confined te such for individuals not a few extend their wanderings quite far beyond the tropical zone, occasionally even roaming northward to Massachusetts.
It attains a length of from a foot and a half to three feet and a weight of about three to ten pounds, but the average is far below the maximum mentioned.
Tf
Notwithstanding its wide geographical distribution, it is in truth a shore fish and seeks its food close to the shore or on muddy or sandy flats where shellfish—especially small bivalve shellfish—most abound. When the flood tide begins and “up to full-tide” is the select time for feeding, and “ flats in water varying from a depth of eight to ten inches,” a choice place for hunting for food. As the fishes feed in such shallow water, their heads go down and their tails come cut of the water, and as they work in shorewards their dorsal fins cut the water, and the sunlight is reflected from their silvery sides.’ The actions of the fish thus seen have suggested to some the name “ grubber.”’
There is a beautiful correlation between the fish’s food and the structural means for assimilating it. The dentition as a whole is quite peculiar—unlike that of any other animal. The bony roof of the mouth is closed in by the juxtaposition of the parasphenoid and pterygoid bones and covered with roundish molar teeth and the floor of the mouth has opposed teeth so that the fish is well pro- vided with the means for crushing the shells which it takes; ex- ternally is provision for finding and rooting them up in the pro- jecting conic snout, which is so prominent as to have suggested one of its early names—Conorhynchus or cone-snout.
GILL] | THE TARPON AND LADY-FISH 43
iaet
Fic. 7.—Albula vulpes (Linneus). Transformation of the Ladyfish, from the translucent, loosely compacted larva to the smaller, firm-bodied young—slightly enlarged. Gulf of California. (After Gilbert and Jordan, Mss.)
A favorite region for the discharge of procreative duties is the Gulf of California. There the young may be found in immense quantities and they are “often thrown by the waves on the beach in great masses.” But so different are those young from the mother fish that they would not be recognized by the casual observer. They are “elongate, band-shaped, with very small head and loose
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS [VvoL. 48
transparent tissues.” In the water in fact their eyes alone are visible. Gilbert tells that “ from this condition they become gradu- ally shorter and more compact, shrinking from three or three and a half inches in length to two inches.” Then their form becomes much like that of maturity and from that stage they grow regularly till the proportions of ripe age are attained.
The various phases of this remarkable growth during which, with advancing days, the developing fish grows smaller and smaller, have all been found by Professor Gilbert and drawn under his direction.” Having at length shrunk to almost half the length of the longest esunculoid stage and acquired a roundness and com- pactness of body as well as shape of the adult, it starts anew in growth and continues till the size and other characteristics of the adult are attained. The history of the metamorphosis of the species is quite as remarkable as that of the butterfly. With diminishing length, with increased compactness, the myotomes or muscular folds grow closer together and less obvious, the dorsal fin and, to a less extent, the anal become better developed and advance towards the middle, and innumerable minor or, rather, less evident changes accompany such until the adult form in miniature is obtained.
av
One of its haunts is “the waters of Biscayne Bay and those extending some 60 miles further south,’ and by residents of that shore it “is not known to be found anywhere ” else. There probably, at least, it is angled for as much if not more than elsewhere and is quite generally regarded as the gamiest fish that swims. There near Miami, August Thomas (1903) verified to his own satisfac- tion the verdict of the neighborhood. He approached a school, as is generally done, in a boat with a guide.
“Your guide works the boat towards them carefully, for they are as timid as a deer, and once frightened are very difficult to ap- proach. When within 50 or 60 feet, which is as close as it is pos- sible to get without frightening the fish, you cast the bait to a spot in line with the direction the fish are working, and not nearer than
1A brief notice of Prof. Gilbert’s observations was communicated to Jordan and Evermann (1, 411), but Prof. Gilbert has informed the writer that his results are still unpublished. A figure of an esunculoid stage has been given by Boulenger (C. N. H., vu, 548) “after Gilbert,’ but otherwise no authority is mentioned.
2The writer is indebted to Professor Gilbert for a proof of the accompanying figure which is to be published in President Jordan’s great forthcoming work on fishes.
a
a
GILL] THE TARPON AND LADY—FISH 45
20 or 30 feet to them. The bait is one of the shell-fish upon which the fish feed, and it must be absolutely fresh.” This bait must be allowed to “lie immovable until the fish find it. The first indication is a slight nibble, for they are not vigorous biters, and they must be hooked, for they rarely hook themselves.”
At length one is hooked. Then commences the sport. ‘“ From three to five hundred feet of line is taken out on the first rush, and this is often repeated twice or even three times, making from one thousand to fifteen hundred feet of line in all that is taken out in this manner. When these bursts of speed are over it is fight, fight, fight, every inch of the way to the boat, the runs growing shorter as the fish fails. When at length he sees the boat the mighty strug- gle comes, but not having strength to make a dash, he circles about the boat at a distance of from 10 to 20 feet, often making the circuit half a dozen times—when he finally comes alongside, belly up, he is dead—died as he had lived—dead game—and may be lifted into the boat with safety by the guide.”
Fishes may be caught “from November to April, but it is at its best in December.”
There is much diversity of opinion respecting the culinary charac- teristics of the lady-fish. Thomas thought that “as a table fish they have few equals, either planked or broiled.” Goode, “ from personal observation,” testified that “its reputation is by no means a false one.” In the Bermudas, too, “where large schools are taken ” and where it is known as the Bone fish or Grubber, it is con- sidered “a most excellent food-fish.” Others, however, hold it in little esteem. Goode himself tells that along the southern coast of California where it is “ found in some numbers,” on account of “ its beautiful color it sells readily, but it is not especially esteemed as a table-fish.”’
But it is by all with common consent exalted as a game fish. The celebrated angler Henshall, in 1884, declared that, of all the fishes he had caught in the Indian river inlet, “a bone-fish of about 3 pounds ” gave more real sport than any of the others.” He found that it “fights in the water and in the air like the black-bass, but mostly in the air—a silver shuttle.”
THE GISU OR PTEROTHRISSUS OF JAPAN
A fish occurs in the deep sea off Japan, named gisu by the Japa- nese fishermen, which was considered to be the type of a peculiar family (Bathythrisside) related to the white fishes and other Salmonids by Giinther but which later ichthyologists (Boulenger
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
and Ridewood) have associated with the Albulids. It has- been scientifically named Pterothrissus gisu and Bathythrissa dorsalis and may still be considered as the representative of an independent family, very closely related to the Albulids, but named Pterothris- side. The Pterothrissids are essentially like the Albulids but the branchiostegal rays are in reduced number (about six), the vomer and palatines are toothless, and the peculiar dentigerous bone which covers the basibranchials of the A/bula is undeveloped. The dorsal fin is prolonged over most of the back and caudal region.
This family also is one of ancient origin, especially if Smith Woodward is correct in his statement that Pterothrissus “is not clearly distinguished from Jstieus.” Species of Istiews flourished in the seas whose deposits formed the upper Cretaceous beds of modern Westphalia and Mount Lebanon.
Only one species (Pterothrissus gisw) is known from existing seas; it is a deep-sea fish which occurs at depths of three hundred to four hundred fathoms off the Japanese archipelago. Nothing is
known of its habits.
ABBREVIATIONS EMPLOYED IN THE LETTERING OF FIGURES (AFTER RIDEWOOD. )
al, alisphenoid. max, maxilla.
an, angular. m, nasal.
bb, dentigerous plate covering the of, opisthotic. basibranchials. Opc, opercular.
bo, basioccipital. or, orbitosphenoid.
br, branchiostegal rays. p, parietal.
bs, basiphenoid. pb, pharyngobranchial.
cb, ceratobranchial. pl, palatine.
ch, ceratohyal. pm, premaxilla.
cor, circumorbital bones. pof, postfrontal.
ct, cartilage. pop, preopercular.
d, dentary. prf, prefrontal.
eb, epibranchial. pro, pro-otic.
ecar, ectosteal articular. ps, parasphenoid.
ecp, ectopterygoid. pt, post-temporal.
eo, exoccipital. ptf, posterior temporal fossa.
ep, epiotic. qd, quadrate.
f, frontal. sar, sesamoid articular.
gh, glossohyal. sm, surmaxilla.
hb, hypobranchial. soc, supraoccipital.
hh, hypohyal. sop, subopercular.
hm, hyomandibular. sp, spicular bone.
ah, interhyal. Sq, squamosal.
top, interopercular. st, supratemporal.
j, jugular or gular plate. stf, subtemporal fossa.
me, mesethmoid. sy, sympletic.
mpt, metapterygoid. v, vomer.
VOL. 48, PL. XVII
SMITHSONIAN MISCELLANEOUS COLLECTIONS
pounder (EJlops saurus).
Ten
1es).
IN O16
pr
The Oxeye (Megalops cj
Tarpon (Megalops atlanticus).
(Albula vulpes).
Lady fish
gist).
othrissus
ter:
£
(
Gisu
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. XVIII
eb* ct ‘cbs
Elops saurus : hyobranchial skeleton from dorsal view. (The epibrauchials and pharyngobranchials of the right side are not shown. )
Albula conorhynchus: hyobranchial skeleton from dorsal view. (The epibranchials and pharyngo- branchials of the right side are not shown.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. XIX
Elops saurus; cranium, A, dorsal view; B, back view; C, left side.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 43, PL. XX
Ss
,; B, back view; C, left side
Megalops cyprinoides: cranium, A, dorsal view
VOL. 48, PL. xx!
SMITHSONIAN MISCELLANEOUS COLLECTIONS
& |
a
A, dorsal view; 8, back view; C, left side.
Albula conorhynchus: cranium.
DIAGNOSIS. OF A NEW GENUS: AND .SPECIES OF FOSSIL SEA-LION FROM THE MIOCENE OF OREGON
By FREDERICK W. TRUE,
HEAD CurATor, DEPARTMENT OF BioLocy, U. S. Nationat MusEumM
At the suggestion of Mr. Wm. H. Dall, the National Museum purchased from Mr. B. H. Cammann of Empire City, Coos County, Oregon, in 1898, a portion of a large fossil skull from the soft Miocene sandstone of that locality. The specimen, as I am informed by Mr. Dall, was found by Mr. Cammann in the sandstone bluff on the east side of the lower part of Coos Bay, between Empire City and the “south slough,” in the formation to which Mr. J. S. Diller has given the name of the “ Empire Beds.”
Upon examination, the skull proves, as Mr. Cammann had sup- posed, to be that of a sea-lion. It represents a genus allied to Eumetopias, but much larger. The fragment consists-of the brain- case, or cranium proper, together with the pterygoids and the palatines as far forward as the posterior end of the hard palate. Both zygomatic processes of the squamosal are broken off near the root, and the right parietal bone has been lost, leaving a large opening through which the whole interior of the brain-case can be examined. The tympanic bullz are crushed and splintered off down to the level of the basioccipital and so mingled with the matrix that their form is lost. The surrounding foramina are also obliterated, and the base of the skull thus presents a broad, nearly flat surface, the appearance of which is, at first sight, very misleading. In other respects, however, the fragment is in an excellent state of preserva- tion, and presents characters which plainly indicate its affinities.
It has been deemed desirable to publish the following diagnosis and measurements in advance of a full description, with figures, which will appear later in one of the publications of the U. S. Geological
Survey. PONTOLEON new genus
Similar to Eumetopias, but with the ventral surface of the basi- occipital nearly plane, and the dorsal surface strongly concave. Postglenoid process of the squamosal strongly produced distally and directed somewhat posteriorly, so that the glenoid fossa is broader
47
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
distally than proximally. Dorsal surface of squamosal, between the wall of the cranium and the zygomatic process, concave antero- posteriorly throughout its whole extent. Hard palate abbreviated, the posterior margin concave.
PONTOLEON MAGNUS new species
Size much larger than that of the largest of existing eared-seals. Skull when complete probably about 50 cm. (or 20 inches) long.
DIMENSIONS OF THE TYPE-SKULL OF Pontoleon magnus AND OF TWO ADULT SKULLS OF Eumetopias jubata.
| Pontoleon Eumetopias jubata. Measurements. / tel Bering Id, St. Paul Id., | No. 3792. Kamchatka | | Alaska, | | No. 49720. | No. 49730. | cm. { cm. cm. Total length (basi-cranial)...........c.06 cssesneeesesseeeenes | 40.1 36.5 Total height posteriorly (from line of occipitomastoid processes to top of occipital crest in a straight line’... ig. E7cOn |, SeERe Greatest breadth between occipitomastoid processes... 24.81 De eee ge 5) Greatest breadth between outer margins of zygomatic | | PYOCESSES.OF SQUAMIOSAl,. isvoccedang-ed.cmrnetnatemeserineds 25.08: 1. 23.9 | 1s" sam Greatest breadth between outer margins of occipital EO IAA OER Se oss ancanGaapete nb SoHE ne caseuc, coueor danacassmane PLO. 9.0 8.6 Height of occiput from upper margin of foramen mag- num to top of Geipiial crést.....0.00i. vie soes one toetaends | ‘eny2 grat 4 8.1 Heishtiof foramen) macnn, .2..2..cssa.- seen yentons «es =ee | 5-7 Aiea 4.7 Breadth of foramen Maelo me gososdanesoabonuonrey cesciscccc 4.1 BLOmne| Bon Beneth of anioccipitalscond yle-2-.. ssrsnrencorscse «use ee em / 8.1 6.2 6.4 Breadthhof anvoccipital condyle. .-. << nssnedsecsaceneeasere Bai 3-4 ant Greatest transverse breadth of occipital crest.............. 15.4 15.8 13.8 Breadth between occipital condyles inferiorly.......2..... 172 2.0 Dey, Greatest breadth of occipitomastoid process antero- “pos: | | | [ardtoy elk PAS aeNS tach accscnnaparradeease: door scocpepp oc saceaker| i Na 5-On a 52 Distance from inferior margin of foramen magnum to. | | outer inferior angle of exoccipital.............02. ceeeeeee $26) 4) 8.2 7.8 Distance from outer inferior angle of exoccipital to postglenoid process of squamosal........ ...2.scseeeeeeee 10.2 7.8 7:2 Distance from inferior margin of foramen magnum to tip of hamular process of pterygoid...............0.se0e0 Ag © Es Ok ete Distance from tip of hamular process of pterygoid to | postevion ‘cmd of hat palates. asc <c: ans ccsse~soqued cerned 10.2 Nos | 5.1 Distance from surface of occipital condyles to end of | | astral pioksibets 5 th, Sete 2 ean ait a8 dha eer ee 24.1 1H {ly ze Greatest breadth between outer walls of ascending’ | j plates of palatines at their posterior end................ 5:5) [aries || 5.6 Greatest breadth of posterior nares..................--see00e Sake | 3.6 3.9 Length of glenoid fossa of squamosal (transverse)...... For 7.5 (on Length of glenoid fossa of squamosal (antero-posterior). Aa s-| BRO 352
* Actually 24.0 cm., but the left side is broken and 0.8 cm. has been added to agree with the right side.
* Actually 23.0 cm., but the right side is broken.
*The condyles are a little defective below.
— ee
a
TRUE] NEW SEA—LION FROM MIOCENE OF OREGON 49
Distance from the occipital condyles to the posterior end of the hard palate nearly equal to the mastoid breadth of the skull. Oc- cipitomastoid processes widely divergent, compressed laterally, nearly plane internally. External wall of the ascending plate of the palatines thickened, forming a strong rounded ridge. Posterior nares as broad as deep.
Type.—No. 3,792, U. S. N. M. (Vert. Pal.). Empire Beds (Miocene) of Empire City, Oregon. Collected by B. H. Cammann.
DIATOMS, THE JEWELS OF THE PLANT—-WORLD"*
By ALBERT MANN
To anyone familiar with the beautiful plants which form the sub- ject of this lecture it seems strange that so few people know of their existence, for they are abundant everywhere. The evident explana- tion of this, however, is the extreme minuteness of these organisms, most of which are wholly invisible to the naked eye. Among the 4,000 or more species there is one, Coscinodiscus rex, a perfect Goliath among his brethren, which is nearly as big as the head of an ordinary pin; but with this exception, the larger forms can better be compared to the point of the pin, while many are so extremely small that the highest powers of the microscope are needed to dis- play their form and the carvings with which they are ornamented.
The diatoms belong to the group of flowerless, aquatic plants known as the Algz. Where these are divided into six groups the Diatoms constitute one of the six; thus (1) Rhodophycez, the red alge; (2) Phzophycee, the brown alge; (3) Chlorophycez, the green alge; (4) Bacillariz, the diatoms; (5) Heterokonte, the yellow-green alge; (6) Cyanophycee (Myxophycez), the blue- green alge. Jam disposed, however, to classify them as a sub-order in the Order Conjugate belonging to the green alge, or Chlorophy- cee; and for the following reasons—(1) they have a unicellular thallus, (2) they have large, elaborate and symmetrically arranged chloroplasts, (3) they frequently produce resting-cells with thick walls, (4) they secrete gelatinous masses in which the individuals are embedded, (5) they display a double mode of multiplication, namely, that by fission, or division of one cell into two, and that by sexual reproduction by means of non-motile isogametes. The at- tempt to classify them with the brown alge is absurd.
The distribution of the diatoms is practically universal. They occupy all waters, torrid, temperate, and arctic; fresh, salt, and brackish; still and running. There is hardly a brook, pond, puddle, lake, river, or sea on earth that is destitute of these plants, unless the water be so contaminated with poisonous matter as to inhibit all life. The largest and most elegant forms belong to the tropics, but
1 Delivered at the U. S. National Museum, Washington, D. C., March 18, 1905, under the auspices of the Washington Biological Society.
50
MANN] DIATOMS, THE JEWELS OF THE PLANT-WORLD 51
the most amazing numbers of individuals are found in the arctic regions. Dr. Nansen found them in undiminished abundance at the northern limits of his polar journey.
Geologically the diatoms seem to have first appeared in the closing measures of the Middle Cretaceous ; in other words, they are, though low in organic complexity, comparatively recent in the scale of suc- cessive life-forms. The statement of Castracane, that they have been found in the Carboniferous Era, and the still more amazing claims that have been made of finding them in Devonian and even Silurian deposits are generally discredited. It should be remarked here that the precise period of entrance of these organisms should be quite clear, because of the prolific multiplication of individuals character- istic of these forms on the one hand and of the indestructible nature of their remains on the other.
Diatom structure can be best understood by looking first at the external sxeleton or casing, and then at the living substance within it. Each plant, a unicellular individual, secretes for itself an external case or box of clear and very dense silica, consisting of two valves, an upper and a lower, the one slipping over the other like the lid and bottom of an ordinary pasteboard box, Figs. 8 and 9, A, B. This case is not of uniform shape; but among the 4,000 or more species there can be found al- most every conceivable form, so long as the form displays symmetry on one or both of its axes. Thus there are round, square, triangular, stellate, oval, ovoid, crescent, sigmoid, cuneate, ba- cillar, etc., forms. It is evident that this great variety in the symmetrical con- tour of these structures adds considerably to their beauty and attractiveness. The variety is further enhanced by numerous outgrowths in the form of spines, horns or domes, so arranged as to preserve the symmetry of the valves from which they spring. The two valves themselves, which are with few exceptions identical in shape and markings, are carved and ornamented with an elegance and variety that is well-nigh incon- ceivable. Indeed it may safely be stated that there is hardly a kind of surface ornamentation known that has not been utilized in beauti- fying these structures. Polished beads of varying size arranged in radiating or concentric rows, shining bars, wavy ridges, delicate
[Eee ae}.
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
watch-case milling, hexagonal network, the interspaces of which are often further ornamented with secondary sculpture, intricate ara- besque designs, in short a diversity and delicacy of embellishment that makes these plants the most ornate of all living objects. The highest efficiency of the microscope is often taxed in revealing some of the minuter markings; and valves that were formerly thought to be quite smooth, and which, therefore, bear the inappropriate name, “hyaline” or “pellucida,” are now under better objectives found to be intricately carved with intersecting lines. It should be here stated that the few illustrations accompanying this description con- vey no adequate idea of the objects they represent; for in black and white it is quite impossible to reproduce the appearance of these structures of shining silica, often further beautified by pris- matic colors refracted from their various surfaces.
Outside of this silica casing there is a very thin and perfectly transparent organic pellicle, in vital connection with the living sub- stance within, erroneously called a gelatinous sheath. It is this internal living substance which determines the position of the organ- ism as a plant, and which presents some of the most interesting prob- lems connected with its life-history. It is made up of normal plant protoplasm (cytoplasm) with a single large centrally placed nucleus. Rarely there are two nuclei, found only in two or three species, where they are said to be constant. Generally there are two large vacuoles filled with cell-sap, and two or more chloroplasts. These latter are usually symmetrically arranged, in the elongated diatoms on either side of the median line and in the circular forms in evenly distributed granules or larger masses radially disposed. The green chlorophyl composing these bodies is disguised by an overlying brown or buff pigment called diatomin, which is so readily soluble in alcohol that when living diatoms are treated with that liquid the diatomin in- stantly disappears and the plants are seen to be bright green. The reserve food material stored up by the diatoms is not in the form of starch grains, but of globules of deep yellow, dense and highly re- fractive oil, either floating in the sap of the vacuoles or embedded in the cytoplasm.
Turning now to the physiology of the diatoms the question of their nourishment may be considered. This takes place as in other chloro- phyl-bearing plants, by the assimilation of inorganic substances in solution in the water about them through the agency of sunlight in conjunction with the chlorophyl masses; and in consequence of this fact it is plain that they are precluded from such waters as are not sufficiently lighted; as, for example, subterranean streams and the
MANN | DIATOMS, THE JEWELS OF THE PLANT-WORLD 53
deeper parts of the sea. No actual test has been made of the ocean depth at which diatoms can flourish, but the limit is probably some- thing below 100 fathoms. Specimens of diatoms are, it is true, ob- tained from all depths, even from the abysses of 6,000 fathoms or more; but in such cases they are invariably the dead and empty frustules of plants that have been transported there by surface cur- rents. There are a few diatoms partly or wholly destitute of chloro- phyl and which therefore live a saprophytic life. Such is Nitgschia putrida, a colorless form, and Bacillaria paradoxa another Nitzschia, which is only partly saprophytic and therefore not wholly colorless.
The food-product of assimilation can never be utilized by the individual plant to any great extent; for being encased in an in-
TG or 1
flexible silica box its chance of growth is restricted to a very slight increase in breadth by the slipping apart of the upper and lower valves or lids: in other words, each diatom is formed at its own maximum length. Most of the reserve food is therefore utilized in the multiplication of individuals. This takes place in two curious ways. The first and common method is the asexual one of fission, or the separation of a single plant into two along a median dividing line. Although this is the usual method of multiplication in unicellu- lar plants as well as in single cells of multicellular plants, the process in the diatoms is peculiar in taking place, not transversely, that is along the short axis, as is done in the bacteria, the other alge, etc., but longitudinally from end to end. This peculiarity is the origin of the name “diatom,” from déa toyéw, to cut through. An examination of Figs. 10 and 11 will make this process clear. It is easy to see that
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
smaller individuals must be formed at each repetition of this process, as the two new valves are always formed within the old ones. The diatom loses approximately one-sixtieth of its length by this method ; and if it were continued indefinitely the forms would necessarily dwindle to the vanishing point! This however is corrected by means of the second or sexual method of reproduction, a process that brings about two important results ; the diatom’s vitality is rejuvenated and its ancestral size is restored. This process, called conjugation, may take place in any one of three ways ; —
1. The nucleus of a single diatom divides karyokinetically; the cell contents swell, bursting apart the valves; the mass passes out into the water, becomes spherical, secretes a large quantity of jelly- like substance; the two daughter nuclei reunite; a large “auxo-
spore” is formed, and within this a single large diatom is built, like the parent frustule, but approximately double the size. See Figs. 12 and 13.
2. The second method is where two diatoms come into contact; the contents swell, as before; the two nuclei remain undivided, but fuse together and produce a single auxospore, within which a single diatom, double the former size, is again formed. See Fig. 14.
3. In the third method two parent diatoms join; the nucleus of each divides karyokinetically; the four daughter nuclei unite, the two from one plant with the two from the other, producing two auxospores and giving rise to two large frustules. See Fig. 15.
The first method is common among diatoms that are fixed, and especially those which grow in long filaments. The second is
MANN] DIATOMS, THE JEWELS OF THE PLANT-WORLD 55
rather uncommon, being confined to a few species. The third is perhaps the most frequent of all and is especially characteristic of the moving diatoms.
Conjugation takes from eight to twenty days for its completion; and as the diatoms lose by each act of fission about one-sixtieth in length, and as they divide every five or six days under normal condi- tions of nourishment, it would require not more than one act of con--
jugation yearly to balance the reduction. Frequently conjugation does take place only once in a year; and then it is, at least in our latitude, quite uniformly early in spring, often before the ice has entirely disappeared from the streams.
Two other methods of reproduction are claimed as taking place among the diatoms ; namely, by means of exceedingly minute spores ; and by means of daughter plants, two to sixteen in number, formed within the body of the parent plant. But as the former is wholly unsubstantiated, and the latter, described by G. Murray (in Proc. Roy. Soc. Edin., vol. 21) is, to say the least, most anomalous, no at- tempt will be here made to describe them.
There is one other physiological process of the diatoms which has up to the present time puzzled all its investigators, their motion. Many of the diatoms grow attached to some support, some of the round and oval forms lying flat on the fronds of other algz, while others are fixed at the ends of gelatinous stalks, singly or in clus- ters, or grow as zigzag chains or in rows like beads or in flat bands as long filaments. But large numbers are free; and these, especially
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
Niteschia and Navicula, display a liveliness of motion that is easy to watch but hard to understand. The majority of these forms are boat-shape, and their motion has a stately character quite different from the erratic movements of the lower animal organisms. But these tiny crafts move without oar or sail, paddle-wheel or propeller. They apparently present the anomaly of moving without any organ of locomotion! Many theories have been invented to explain this mystery; as fine protruded pseudopodia (Ehrenberg), osmotic cur- tents of water (Naegli and H. L. Smith), rows of cilia (J. D. Cox), a stream of protoplasm moving along the “raphe” a line on each valve, generally in the middle that is thought to be a narrow cleft (Muller). But none of these fit the case. No method of staining known has been able to show any protrusions. Whatever be the final explanation, it must explain not swimming but creeping; for these organisms are perfectly inert and helpless unless in contact with some fixed surface. This alone disposes of all theories requir- ing cilia, flagella, osmotic currents, etc. The power, too, must be considerable ; for diatoms will push aside in their course inert mat- ter many times their bulk. The theory must also apply te the ends
of the frustules; for the plant will often stand upon end and swing about most vigorously. It is not at all improbable that the so-called “gelatinous sheath,” which overlies uniformly the entire external surface and is connected with the living cell contents through numerous minute pores, is the seat of this motion, and by undulatory movements over its surface produces the phenomenon that is so evident and so puzzling. Any one who will watch the strange ac- cordeon-like extending and retracting movements of those well- named diatoms, Bacillaria paradoxa, especially when, fully extended, they touch each other only at the tips and yet form a series as rigid as a rod, will see that some explanation based on the external mem- brane fits the case better than any other. See Fig. 16, A and B.
A few words should be said upon the practical uses of the diatoms. The first of these is as polishing powders. Under the name of “ tri-
a ei
MANN] DIATOMS, THE JEWELS OF THE PLANT-WORLD 57
poli” they have been long in use to give brilliancy to metallic and other surfaces ; while, either mixed with soap or put up dry, they are sold under fancy names for the same purpose. One widely known brand of tooth-powder is composed entirely of diatom shells. As an absorbent of nitro-glycerine they have been extensively used in the manufacture of various dynamite compounds. They are also em- ployed as a substitute for asbestos in the composition of jackets for steam-pipes, as packing in refrigerators, etc. They have at least a semi-practical use in refuting the utilitarian theory of the origin of species; their rigidly exact yet infinitely diversified carving and ornamentation refusing absolutely to fit into such a conception. Diatoms have a most curious use among the more abject inhabitants of Lapland and Bohemia as a substitute for or an adulterant of food. Under the name of “berg mehl”’ diatomaceous earth is mixed with flour, fat, etc., and eaten. It is hardly supposable that this fossil earth contains any appreciable amount of nourishment. The philosophy of the practice is probably the fact that where a hungry man has a stomach capacity of two quarts and a food supply of only a pint, he can cajole himself and gain a sense of plethoric bounty by adding three pints of inert matter to his supply,—a sort of “square meal,” it is true, but a very hollow one! The diatoms do, however, form a considerable part of the world’s food supply, at least in an indirect way; for they are one of the principal sources of nourish- ment for mollusks, the clams, oysters, etc., whose stomachs always contain large quantities of these plants; as well as constituting a good part of the food of small fishes and of the animal organisms on which larger fish feed. Thus they are a sort of primary source of organic food, on the abundance of which many of our most valued food products depend.
It is well to mention here that the diatoms give promise of great practical value in determining the origin of sea-bottoms and the direction and extent of the sea-currents by which they are trans- ported. Their use to applied science in this respect is now being investigated.
There will be no difficulty for anyone interested in the examination of these plants in finding them, either as living or fossil forms. Wherever there is a brown coloration of the surface of the mud, sub- merged stones or twigs, not a red-yellow, which is due to iron, but a brown-yellow to almost black, there are diatoms in abundance. It is their characteristic color, when found in masses; and a little of the material placed under the microscope will reveal thousands of them. Or if fossil material is needed, diatomaceous earth can be
58 SMITHSONIAN: MISCELLANEOUS COLLECTIONS [voL. 48
found in almost every State in the Union and almost every land on earth. Immense beds exist, for example, at Nottingham, Md. ; Rich- mond, Va.; Keene, N. H.; Monterey, Santa Monica, Rodondo Beach, Cal.; near Spokane, Wash. ; etc.; while smaller deposits are frequent in many other localities. In foreign lands there are large deposits at Sendai, Japan; Ananino and Simbirsk, Russia; Alicate, Sicily; Bilin, Bohemia ; Luneberg, Germany; Mors, Jutland; Oamaru, New Zealand; Springfield, Barbados; etc.
The cleaning of diatoms of organic matter and the preparation of these and fossil forms as permanent microscopic mounts cannot be entered into here. The processes are easily learned from any good work on the microscope.
SMITHSONIAN INSTITUTION, Wasuincton, D. C., March, 1905.
VCL. 48, PL. XxiIl
SMITHSONIAN MISCELLANEOUS COLLECTIONS
DIATOMS
Pirate XXII FicurE 1. Coscinodiscus asteromphalus, E., X 185. Sendai, Japan. Photo- graph by A. A. Adee. 2. Lepidodiscus elegans, Witt., X 620. Simbirsk, Russia. Photo- graph by A. A. Adee.
FIGURE I.
PLATE XXIII Biddulphia Roperiana, Grev., variety mollis, Mann, X 400. Pacific Ocean, S. S. “ Albatross,” station 3608. Plagiogramma sceptrum, Mann, 375. Galapagos Islands.
Stephanopyxis ferox, Grev., X 400. California guano. From Moebius’ Plates of Diatoms.
Triceratium sp.? X 660. Bering Sea. Entogonia Davyana, Grev., X 300 (= Heibergia Barbadensis) Barbados. From Moebius’ Plates of Diatoms.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, Pi. XXIII
DIATOMS
+ pe cg. tke Pe » ewe = aad ie ton os | A ue y a 7 »
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 48, PL. XXIV - 48, .
ilies. eit
DIATOMS
PLaTE XXIV
‘Ficure 1. Secondary markings within the hexagonal network of Triceratium favus, Bright, X 1060. Oamaru, N. Z. Photograph by A. A. . Adee. 2. Amphora, n. s., X 565. Pacific Ocean. 3. Triceratium Campechianum Cl., 720. Florida. Photograph by A. A. Adee.
FIGURE I.
PLaTE XXV
Cestodiscus ovalis, Grev., X 335. Moron, Spain. From Moebius’ Plates of Diatoms.
Actinoptychus Waittianus O. Jan., X 200. Hayti. From
_ Diatomaceen, Jeremie in Hayti.
Brunia japonica, Temp., 100. Japan.
Aulacodiscus n. s.. X 1000. Pacific Ocean, S. S. “ Albatross,” Station 4029 H.
Navicula invenusta, Mann, X 375. Galapagos Islands.
Navicula bullata, Norm., 350. Western Australia. From Moebius’ Plates of Diatoms.
VOL. 48, PL. XXV
SMITHSONIAN MISCELLANEOUS COLLECTIONS
fi
ian
SS ere
‘ttre
‘Ls
4
.-
DIATOMS
NOTES ON THE NOMENCLATURE OF CERTAIN GENERA OF BIRDS
By HARRY C. OBERHOLSER
The following notes concern the status of some seventeen generic and a few specific terms that seem to require change. Most of these, though for several years held in abeyance by the writer, appear not yet to have been published by others; a few are revivals of former changes that lately have been ignored; and one or two have been mentioned as probably necessary by recent writers who failed to go farther. The alterations in specific names pertain only to species belonging to the genera treated.
The writer is under obligation to Dr. Charles W. Richmond for various courtesies in connection with the preparation of this paper, and wishes here to express his consequent appreciation.
BELLONA Mulsant and Verreaux
This name,’ employed by authors for a genus of West India hummingbirds, is, as already pointed out by Mr. J. H. Riley,? un- tenable, being preoccupied by Bellona Reichenbach,? a genus of ornithicnites. In seeking a name for the group, however, Mr. Riley rejects the once used Orthorhyncus Lacépéde* as a nomen nudum because “ no type was specified and the diagnosis is not diagnostic,” but revives it ta date from Froriep,* and by elimination fixes as its type Trochilus mosquitus Linneus. Then, since Orthorhyncus would thus take the place of the present Chrysolampis, Mr. Riley, still by process of elimination, transfers the name Chrysolampis to the group now known as Eulampis, and the term Eulampis to the unidentified “ Trochilus niger”? Wied. This arrangement leaves the preoccupied Bellona without a name, and it is accordingly christened Microlyssa.* These changes, however, can not stand, because Orthorhyncus is the proper name for Bellona, as may easily be shown; and they furthermore constitute a forcible illustration of the
* Bellona Mulsant and Verreaux, Classif. Troch., 1866, p. 75. * Auk, 1904, p. 485.
3 Riley, Auk, 1904, p. 485.
4 Natirl. Syst. Vogel, 1852, p. xxx.
= abl, Ots:, 1790, ‘p: 0:
6 Dumeril’s Analyt. Zool., 1806, p. 47.
59
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
instability and unsatisfactory nature of generic type determinations by elimination.
The genus Orthorhyncus was instituted by Lacépéde? for the “ Oiseaux mouches,” undoubtedly of Buffon, a group of twenty-four species, to one of which the name of course must be applied; so that Orthorhyncus stands on equal basis with the other names of Lacépéde proposed in the same place, which have been subsequently accepted without question. The type of Orthorhyncus was fixed as Trochilus cristatus Linneus by Gray in 1840;? and happily enough the same species also becomes the type if this be determined by elimination.
The species of this group should therefore stand as follows:
Orthorhyncus cristatus cristatus (Linnzus).
Orthorhyncus cristatus emigrans Lawrence.
Orthorhyncus ornatus Gould.
Orthorhyncus exilis (Gmelin).
DROMZAUS Vieillot
This name, spelled as above, does not occur in Vieillot’s “ Analyse,” and so far as we are aware was never used by this author. He does, however, in the main part of this work propose Dromiceius for the emus, type Casuarius novaechollandie LatHam ;° and in the supple- mentary list where he gives the derivations of his generic names, he inserts instead of Dromiceius the term Dromaius* which Ran- zani later emended to Dromeus? Since Dromiceius can scarcely be considered a typographical error for Dromaius, it follows that the former, standing first in the book, becomes the proper name for the genus.
The species are:
Dromiceius novehollandie (Latham).
Dromiceius ater (Vieillot).
Dromiceius irroratus (Bartlett).
Dromiceius patricius (De Vis) (fossil).
Dromiceius gracilipes (De Vis) (fossil).
Dromiceius queenslandie (De Vis) (fossil).
HYDRORNIS Milne-Edwards The fossil genus Hydrornis Milne-Edwards® is preoccupied by Hydrornis Blyth, used for a member of the Pittidee (Paludicola
1Tabl. Ois., 1799, Pp. 9.
2 List Gen. Birds, 1840, p. 14.
3 Analyse, 1816, p. 54.
4 Analyse, 1816, p. 70.
5 El. di. Zool., w1, pt. 1, 1821, p. 98.
6 Rech. Oiseaux Foss. France, 1, 1867, p. 362, Tb. 57, fig. 18-22.
OBERHOLSER] NOTES ON NOMENCLATURE OF BIRDS 61
nipalensis Hodgson).' It may be replaced by Dyspetornis, from
dvenetys, difficilis, and ¢pytc, avis. The type and only species,
Hydrornis natator Milne-Edwards, should therefore now be called: Dyspetornis natator (Milne-Edwards).
NENIA Boie
The name Nenia Boie® is untenable by reason of Nenia Stephens,‘ employed for a genus of Lepidoptera. The next available name is apparently Larosterna Blyth;* but the book in which this was pub- lished, though bearing on its title page the date 1849, contains in- ternal evidence to show that it did not appear until at least 1852. This gives priority to /nca Jardine,’ which has the same species, Sterna inca Lesson, as its type. The only species of this group, therefore, now becomes:
Inca inca (Lesson).
GNATHOSITTACA Cabanis An earlier name for Gnathosittaca Cabanis? which is based on Gnathosittaca heinet Cabanis (== Conurus icterotis Massena and Souancé) is found in Ognorhynchus Gray,* type Conurus icterotis Massena and Souancé. The sole species is: Ognorhynchus icterotis (Massena and Souancé).
DASYPTILUS Wagler The generic name commonly applied to Psittacus pecquetu Lesson is Dasyptilus Wagler ;° but this is, however, antedated by Psittrichas Lesson,'°used for the same bird. This species should therefore stand as: Psittrichas pecquetw (Lesson.)
NANODES Vigors and Horsfield The term Nanodes Vigors and Horsfield" for a group of Psittacidz
1 Journ. As. Soc. Bengal, xt, 1843, p. 960.
ILO (HE
3 Tsis, 1844, p. 1809.
‘fil. Brit. Ent., 11, 1820, p. 165.
5 Cat. Birds Mus. As. Soc., 1852, p. 293.
5 Contrib. Orn., 1850, p. 33.
7 Journ. f. Ornith., 1864, p. 414.
§ List Psitt. Br. Mus., 1850, p. 33.
8 Abhandl. Ak. Wissensch. Miinchen, 1, 1832, p. 502.
'Tllustr. Zool., 1831, pl. i; Ferussac’s Bull. des Sci. Nat., xxv, June, 1831,
P. 341. " Trans. Linn. Soc., xv, Feb., 1827, p. 274.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
is rendered untenable because of Nanodes Schonherr,' a genus of Coleoptera. Some time ago Forbes proposed to put Lathamus Lesson in place of Nanodes Vigors and Horsfield, dating the former from 1831,° and considering its type to be Lathamus rubrifrons Lesson (= Psittacus discolor Shaw); but the earlier use of La- thamus, also by Lesson, as a subgenus of Psittacus, for Psittacus, aurifrons Lesson,’ makes it a synonym of Bolborhynchus and thus unavailable for Nanodes. ‘The next and only other synonym of Nanodes, Euphema Wagler,’ becomes consequently its tenable title, since this is not invalidated by Euphemus Rafinesque,? a nomen nudum. The type and sole species ought therefore to be called:
Euphema discolor (Shaw).
DENDRORNIS Eyton
The name of the group of Dendrocolaptidz to which the generic term Dendrornis Eyton’ has been applied must apparently be changed. The type of Xiphorhynchus Swainson as usually cited® is Dendrocolaptes procurvus Temminck; but earlier in the same year Swainson had used this generic name in describing Xiphorhynchus flavigaster,? which is a member of the present genus Dendrornis. Although Swainson evidently intended to make Dendrocolaptes pro- curvus Temminck the type of Xiphorhynchus, he defeated his pur- pose by allowing the previous publication of Xiphorhynchus in com- bination with the name of a species of another group, such publica- tion being quite sufficient to fix the name of a genus. Since in this case the question is not complicated by the mention of any other species, Xiphorhynchus flavigaster Swainso * must be considered the type of Xiphorhynchus, and this generic term therefore transferred to displace Dendrornis.
The species are as follows:
Xiphorhynchus guttatus. (Lichtenstein).
Xiphorhynchus guttatoides (Lafresnaye).
1Curc. Disp. Meth., 1826, p. 322.
2 Proc. Zool. Soc. Lond., 1879, p. 166.
3 Traité d’Orn., 1831, p. 205.
4 Cent. Zool., 1830, p. 63, pl. 18.
5 Abhandl. Ak. Wissensch. Miinchen, 1, 1832, p. 492. 8 Anal. Nat., 1815, p. 144.
7 Jardine’s Contr. Ornith., 1852, p. 23.
§ Zool. Journ., 11, Aug—Nov., 1827, p. 354.
° Phil. Mag., 1, June, 1827, p. 440.
'0 Phil. Mag., 1, June, 1827; p. 440.
OBERHOLSER] NOTES ON NOMENCLATURE OF BIRDS 63
Xiphorhynchus
palliatus ‘(Des Murs).
Xiphorhynchus rostripallens rostripallens (Des Murs).
Xtphorhynchus Xtphorhynchus Xtphorhynchus
rostripallens sororius (Berlepsch and Hartert). eytomt (Sclater). d’orbignianus (Pucheran and Lafresnaye).
Xiphorhynchus flavigaster flavigaster Swainson.
Xiphorhynchus Xiphorhynchus
flavigaster eburneirostris (Eyton). flavigaster mentalis (Lawrence).
Xiphorhynchus flavigaster megarhynchus (Nelson).
Xiphorhynchus Xiphorhynchus Xiphorhynchus Xiphorhynchus Xiphorhynchus mann). Xiphorhynchus Xtphorhynchus Xiphorhynchus Xiphorhynchus Xiphorhynchus Xiphorhynchus Xtiphorhynchus
striatigularis (Richmond).
erythropygius (Sclater).
punctigulus (Ridgway).
triangularis triangularis (Lafresnaye). triangularis bogotensis (Berlepsch and_ Stolz-
lacrymosus lacrymosus (Lawrence). lacrymosus eximius (Hellmayr). nanus nanus (Lawrence).
nanus costiricensts (Ridgway). nanus confinis (Bangs).
susurrans (Jardine).
fraterculus (Ridgway).
Xiphorhynchus pardalotus (Vieillot).
Xiphorhynchus Xiphorhynchus Xiphorhynchus Xiphorhynchus Xtphorhynchus Xiphorhynchus Xtiphorhynchus Xiphorhynchus Xtphorhynchus Xiphorhynchus Xtphorhynchus Xiphorhynchus
polystictus (Salvin and Godman). ocellatus (Spix).
lineatocapillus (Berlepsch and Leverkiihn). insignis (Hellmayr).
elegans (Pelzeln).
weddell. (Lafresnaye).
kienerw (Des Murs).
spit (Lesson).
chunchotambo (Tschudi). multiguttatus (Lafresnaye). obsoletus obsoletus (Lichtenstein). obsoletus notatus (Eyton).
XIPHORHYNCHUS Swainson
As explained under the previous heading, the generic name Xiphorhynchus Swainson,’ since its type is clearly Xiphorhynchus flavigaster Swainson, belongs to Dendrornis. As the group now called Xiphorhynchus is thus left without a name, it may be known
* Phil. Mag., 1, June, 1827, p. 440.
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
as Xiphornis, from tyos, ensis, and épus, avis, and its type desig- nated as Dendrocolaptes procurvus Temminck. The species are: Xiphornis procurvus (Temminck). Xiphornis venezuelensis (Chapman). Xiphornis trochilirostris (Lichtenstein). Xiphornis thoracicus (Sclater). Xiphornis lafresnayanus (d’Orbigny). Xiphornis rufodorsalis (Chapman). Xiphornis falcularius (Vieillot). Xiphornis pusillus (Sclater). Xiphornis subprocurvus (Reichenbach). Xiphorms dorsoimmaculatus (Chapman). Xtiphornis pucheranii (Lafresnaye).
SHARPIA Bocage
The generic term Sharpia, bestowed by Bocage? on a group of Ploceidz, is preoccupied in coleoptera by Sharpia Tournier.2 It may be replaced by Notiospiza, from véteos, meridianus, and «zé£a, fringilla.
The type is Sharpia angolensis Bocage; and the two species will stand as:
Notiospiza angolensis (Bocage).
Notiospiza sanctithome (Hartlaub).
MALACOPTERON Eyton
Doctor Sharpe has already noted’ that Malacopteron Eyton is preoccupied in Coleoptera by Malacopterus Serville,’ and proposes to use Setaria Blyth® in its place. Unfortunately this also is debarred, by Setaria Oken' for a genus of Vermes. The genus Ophrydornis Bittikofer,* based on Setaria albogularis Blyth, is quite distinct from Malacopteron proper, and therefore can not be employed as a sub- stitute for the latter. Doctor Charles W. Richmond calls the writer’s attention to the fact that Dr. Sharpe has recently, in seeming inad- vertence, transferred this name Ophrydornis to the Malacocercus
1 Jorn. Sci. Math. Phys. e Nat. Lisboa, v1, 1878, p. 258.
2 C. R, But. Bele., XV 1873,-D: Cxmavas.
3 Bull. Brit. Orn. Club, xt1, 1902, p. 54.
4 Proc. Zool. Soc. Lond., 1839, p. 102.
5 Ann. Soc. Ent. France, 1, 1833, p. 565.
6 Journ. As. Soc. Bengal, xt, pt. 1, 1844, p. 385.
™Lehrb. d. Naturg., 1, 1815, p. xiii.
® Notes Leyd. Mus., xvi, 1895, p. IoT.
OBERHOLSER | NOTES ON NOMENCLATURE OF BIRDS 65
albogularis of Blyth, which is a Dumetia, and at the same time left Setaria albogularis Blyth, the type of Ophrydornis, in Malacopteron (Setaria) !* Since in view of these circumstances it becomes necessary to provide a new name for Malacopteron, it may be called Horizillas, from dp, limito, and ‘Adds, turdus, with Malacopteron magnum Eyton as the type.
The species to be referred to this group are:
Horizillas magna (Eyton).
Horizillas cinerea cinerea (Eyton).
Horizillas cinerea bungurensis (Hartert).
Horizillas rufifrons (Cabanis).
Horizillas palawanensis (Buttikofer).?
Horizillas pyrrhogenys (Temminck).
Horizillas affinis (Blyth).
Horizillas notata (Richmond).
Horizillas melanocephala (Davison).
Horizillas cinereicapilla (Salvadori).
HEDYMELA Sundevall
The generic term Hedymela Sundevall,*? recently employed by Dr. Sharpe for the pied flycatchers,* is long antedated by Ficedula Bris- son.° The type of both is the same—WMotacilla atricapilla Linnzeeus— and if Brissonian genera are accepted, as is now the all but universal practice, the latter name (Ficedula) must be used for this group. The Motacilla atricapilla of Linnzus,® moreover, must give place to Motacilla ficedula Linneus,’ a prior name for the same species. Also, the bird commonly known as Muscicapa collaris Bechstein*® must be called Ficedula albicollis (Temminck), because Muscicapa collaris Bechstein® is preoccupied by Muscicapa collaris Latham, a synonym of Platysteira cyanea, and Muscicapa albicollis Temminck"” is the next available name.
The species of this genus should consequently stand as follows:
1 Hand-List Gen. and Spec. Birds, 1v, 1903, pp. 27, 38, 39.
2This is Trichostoma rufifrons Tweeddale, nec Malacopteron rufifrons Cabanis, and is the Turdinus rufifrons of Sharpe, Hand-List Gen. and Spec. Birds, iV, 1903, p. 33.
3 Ofvers. Kongl. Vetensk. Ak. Forhandl. Stockholm, 1846 (1847), p. 225.
§ Hand-List Gen. and Spec. Birds, 11, 1901, p. 213.
5 Orn., II, 1760, p. 369.
O Syst WV ab, ed.) 20%, 1758," p: 187:
7 Syst. Nat., ed. 10, 1, 1758, p. 185.
8 Gem. Naturg. Deutschl., 1v, 1795, p. 495.
° Ind. Orn., 11, 1790, p. 471.
0 Man. d’Orn., 1815, p. 100.
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
Ficedula ficedula ficedula (Linnzus). Ficedula ficedula speculigera (Bonaparte). Ficedula semitorquata (Homeyer). Ficedula albicollis (Temminck).
CHENORHAMPHUS Oustalet
Chenorhamphus Oustalet,* based on Chenorhamphus cyano pectus Oustalet (— Todopsis grayi Wallace), is rendered untenable by Chenoramphus Gray? of which the type is Ardea oscitans Boddaert. Since it has no other name it may be called Conopotheras, from zwvwzoOnpas, muscicapa.
The type and sole species is:
Conopotheras grayi (Wallace).
HELMINTHOPHILA Ridgway
The name Helminthophila Ridgway,’ long in use for a genus of Mniotiltide in place of the preoccupied Helminthophaga Cabanis, * must itself be supplanted by Vermivora Swainson’ of much earlier date. Swainson evidently intended Vermivora as the generic name for Sylvia vermivora Wilson (== Helmitheros vermivorus Auct. recent.), and he so published it;’ but in another article previously appearing, he made use of this term’ in combination with Sylvia solitaria Wilson (— Certhia pinus Linneus), which species therefore becomes the type of the genus. Furthermore, V ermivora Swainson is not, as often considered, preoccupied by “ Vermivora” Meyer,® for this latter is merely a group name—“ Vermivore,” and not used in a generic sense at all.
The species of this genus should therefore stand as:
Vermivora chrysoptera (Linnzus).
Vermivora lawrenceii (Herrick).
Vermivora leucobronchialis (Brewster ).°
Vermivora pinus (Linneus).
1 Bull. Assoc. Scient. de France, xxt, 1878, No. 533, p. 248.
*Gen. Birds, 11, 1848, p. 562.
3 Bull. Nutt. Orn. Club, vit, 1882, p. 53.
4 Mus. Hein., 1, 1850, p. 20.
5 Phil. Mag., 1, June, 1827, p. 434.
6 Zool. Journ., ut, Apr.—July (published in July or later), 1827, p. 170.
7 Phil. Mag., 1, June, 1827, p. 434.
8 Besch. Vig. Liv- und Esthl., 1815, p. 118.
8 Probably a xanthochroic phase of V. chrysoptera, or a hybrid between /.
chrysoptera and V. pinus. 10 Almost certainly a leucochroic phase of V. pinus.
a
OBERHOLSER | NOTES ON NOMENCLATURE OF BIRDS 67
Vermivora bachmam (Audubon).
Vermivora peregrina (Wilson).
Vermivora celata celata (Say). ‘ Vermivora celata sordida (Townsend). Vermivora celata lutescens (Ridgway). Vermivora rubricapilla rubricapilla (Wilson). Vermivora rubricapilla gutturalis (Ridgway). Vermivora virgie (Baird).
Vermivora crissalis (Salvin and Godman). Vermivora lucie (Cooper).
TIARIS Swainson
Doctor Charles W. Richmond has already shown? that Tiaris Swain- son? belongs properly to Euetheia, but he failed to provide a name for the consequently nameless group of South American Fringillidze for which Tiaris has commonly been employed. This, therefore, may be called Charitospiza, from yéprc, gratia, and oxfa, fringilla. The type and only species, Fringilla ornata Wied,* needs a new specific designation on account of the earlier Fringilla ornata Vieil- lot, * and as it has no synonyms, may be known as:
Charitospiza eucosma Oberholser.
COTURNICULUS Bonaparte
An earlier name for Coturniculus Bonaparte® is found in Ammo- dramus Swainson,° the real type of which is Ammodramus bimacu- latus Swainson—not, as commonly considered, Fringilla caudacuta Wilson (= Oriolus caudacutus Gmelin)... This is a case precisely similar to those of Xiphorhynchus and Tiaris, since the first use of Ammodramus® is in the original description of Ammodramus bimacu- latus, the western continental form of Ammodramus savannarum (Gmelin), antedating by several months the publication of an article wherein Fringilla caudacuta Wilson is given as the type.”
The forms of this group will be therefore once more in possession of their former generic designation, and pass as:
1 Auk, XIX, 1902, p. 87. 2 Phil. Mag., 1, June, 1827, p. 438 (type Tiaris pusilla Swainson). 3 Reis. Brasil, 11, 1821, p. 191. 4 Nouv. Dict. d’ Hist. Nat., x1, 1817, p. 243 (Polynesia). 5 Geog. and Comp. List Bds. Eur. and N. A., 1838, p. 32. 5 Phil. Mag., 1, June, 1827, p. 435. 7Cf. Zool. Journ., m1, Aug.—Nov., 1827, p. 348. ' § Antea, pp. 62, 67. 9 Phil. Mag., 1, June, 1827, p. 435. 1 Zool. Journ., 11, Aug.—Nov., 1827, p. 348.
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
Ammodramus savannarum savannarum (Gmelin). Ammodramus savannarum passerinus (Wilson). Ammodramus savannarum obscurus Nelson. Ammodramus savannarum floridanus (Mearns). Ammodramus savannarum bimaculatus (Swainson).
AMMODRAMUS Swainson
Since the term Ammodramus Swainson" belongs to Coturniculus, as already shown,” another name is required for the group to which the former has been applied, and as there is none such available, it may be called Ammospiza, from 4p,u0s harena, and ozt£a, fringilla, with Oriolus caudacutus Gmelin as the type.
The species and subspecies are:
Ammospiza maritima maritima (Wilson).
Ammospiza maritima macgillivrau (Audubon).
Ammospiza maritima peninsule (Alien).
Ammospiza maritima fisheri (Chapman).
Ammospiza maritima sennetti (Allen).
Ammospiza nigrescens (Ridgway).
Ammospiza caudacuta caudacuta (Gmelin).
Ammospiza caudacuta nelsoni (Allen).
Ammospiza caudacuta subvirgata (Dwight).
Ammospiza leconteu (Audubon).
Ammospiza henslowii hensloww (Audubon).
Ammospiza henslowti occidentalis (Brewster).
* Phil. Mag., 1, June, 1827, p. 435. * Antea, p. 67.
Peele seOME TT TistPAST HISTORY AND roio RETURN
A SHORT BIBLIOGRAPHY WITH NOTES :
By EUGENE FAIRFIELD McPIKE
Memeer B. S. A., anp I. I. B., BRUSSELS
The formation of this collection (mentioned in The Observatory, 28: 141), has been facilitated by data courteously supplied by Herr Berthold Cohn, Strassburg, Germany; Dr. Chas. F. Forshaw, fh SL. Bradiord,. Eneland;- the. Rev. S.J. _Johnson,, M-A., F.R.A.S., Melplash Vicarage, Bridport, England; Professor Kurt Laves of the University of Chicago; the Hon. John S. Lawrence, Grand Rapids, Michigan, and others, to whom the compiler makes grateful acknowledgment. A supplemental note will appear in The Observatory, probably in June, 1905.
Part rt. Comets Past History
Bronte, Rev. Patrick. On Halley’s Comet, in 1835. Popular astronomy, 12: 571, Northfield, Minnesota, October, 1904.
A poem reprinted from The Bradfordian, no. 11, p. 176, Bradford, York- shire, England, August 1, 1861.
CHAMBERS, GEORGE F. 1841. A Handbook of descriptive and prac-
tical astronomy, 3 vols. Oxford, 1889, 1890.
Contains a cut (1:438) of a portion of the Bayeux Tapestry showing the comet of Halley, in 1066. “The comet of Halley appeared again in 1066, at the time when William the Conqueror invaded England. The chroniclers unanimously write: ‘The Normans, guided by a comet, invaded England.’ The Duchess-Queen Matilda, wife of William, has represented this comet and the amazement of her subjects on the tapestry (230 feet long) which may be seen at Bayeux.” See Popular Astronomy ... by Camille Flam- marion, translated by J. Ellard Gore, p. 479, New York, n. d.
FLAMMARION, CAMILLE. Popular astronomy: a general description of the heavens, by Camille Flammarion, translated by J. Ellard Gore, New York, n. d.
“Queen Victoria has in her crown a jewel the design of which was sug- gested by the tail of this comet, which had the greatest influence on the vic- tory at Hastings.” (P. 479.)
69
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
FLAMMARION, CAMILLE. Astronomie populaire: description gén- érale du ciel. 867 p. Paris, 1890. (See p. 609.)
Grecory, Davip. Astronomice, geometrice et physicze elementa. Oxford, 1702, fol. Translated into English; with additions. To which is added, Halley’s Synopsis of the astronomy of comets, revised and corrected by Edmund Stone, London, 1713, 1726, 2 vols. 8vo.
This book is mentioned in the ‘ Bibliotheca Britannica,’ by Robert Watt, vol. I, section 439v, Edinburgh, 1824.
Hatitey, Epmonp. 1656-1742. Astronomiz cometice synopsis, autore Edmundo Hallieo apud Oxoniensis geometriz professore Saviliano, & Reg. Soc. S. See Philosophical Transactions .. . vol. 24, for the years 1704 and 1705, pp. 1882-1899, London, 1706. Halley’s “most celebrated work ... was ‘Astronomiz cometice synop-
sis, communicated to the Royal Society in 1705, and separately published
in English at Oxford the same year. It was reprinted with his Tables in
1749, and translated into French by Le Monnier in 1743.” (See Dictionary
of National Biography, 24: 108, New York, 1890.)
Hatiey, EpmMonp. 1656-1742. Astronomiz cometicz synopsis, autore Edmundo Hallieo apud Oxoniensis geometriz professore Saviliano, & Reg. Soc. S. See the philosophical transactions of the Royal Society of London from their commencement, in 1665, to year 1800: abridged, with notes and biographic illustrations, by Charles Hutton . . .., London, 1809.
This abridgment gives (5: 201) only the caption of Halley’s memoir, with the note following: “Dr. Halley’s Astronomy of comets is here omitted, as it has been elsewhere published in a much fuller and more complete state, and that both in the English and Latin languages; as in the Miscellanea Curiosa, of Dr. Halley, in 3 volumes 8vo, in English; also translated and published in English, by G. T. Gent, in 8vo, 1757; and still more complete
and perfect in Dr. Halley’s Astronomical Tables, in 4to, both in Latin and English.”
HALiLEey, EpMonpD. 1656-1742. Astronomical tables with precepts both in English and in Latin, for computing the places of the sun, moon, planets and comets. 1 vol. 4to. London, 1752.
This rare work, of which a copy is in the Hon. John S. Lawrence’s private library, Grand Rapids, Michigan, was probably edited by James Bradley, suc- cessor of Halley as astronomer-royal. (Cf. Notes and Queries, 9th series, 11: 464.) In this book a revision of Halley’s Synopsis of comets is included. (See Philos. Trans., abridged, Hutton, 5: 201, note, London, 1809.)
MCPIKE] HALLEY’S COMET vi
HERSCHEL, Sir JoHN F. W. Results of astronomical observations made during the years 1834, 5, 6, 7, 8, at the Cape of Good Hope; i Loudon smut, Kider &°Co., 1847. See chapter V., pp. 393-400.
HERSCHEL, Sir J[OHN] F.W. Observations of the comet of Halley, after the perhelion passage in 1836, made at Feldhausen, Cape of Good Hope, by Sir J. F. W. Herschel. Memoirs of the Royal Astronomical Society; 10: 325-335, London, 1850.
Hrnp, J. R[ussELL]. On the past history of the comet of Halley. Monthly Notices of the Royal Astronomical Society, 10: no. 3 (January, 1850) 51-58, London.
Hinp, J. Russert. The comets: a descriptive treatise upon those bodies . . . London: John W. Parker and Son, 1852. See chap- ter iv., ‘ The comet of Halley,’ pp. 35-57.
Lynn, Witt1AM THyNNE. Remarkable comets; a brief survey of the most interesting facts in the history of cometary astronomy. Eighth edition, revised. 45 p. London: E. Stanford, 1900. See
pp. 11-13.
Mactear, [ ....] Observations of Halley’s comet, made at the royal observatory, Cape of Good Hope, in the years 1835-1836. Read April 14, 1837. Memoirs of the Royal Astronomical So- ciety, 10: 91-155, London, 1838.
McPixe, EuGeNne FarrFieLtp. Halley’s comet. Popular As- tronomy, 12: 685, Northfield, Minnesota, December, 1904. A note referring to Dr. Halley’s discovery of the identity of the comet of 1682 known as Halley’s comet.
SmytH, Captain W. H. Observations of Halley’s comet. Mem- oirs of the Royal Astronomical Society, 9: 229-246, London, 1836. Read June 10, 1836.
Wuiston, WittiAmM, Lectures on Sir Isaac Newton’s mathematic philosophy, and Dr. Halley’s account of comets. London, 1716,
8vo. This work is mentioned in the ‘ Bibliotheca Britannica,’ by Robert Watt. vol. i., section 961h, Edinburgh, 1824.
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
WintTHRoP, [JOHN]. 1714-1799. Two lectures on comets; Bos- ton, I811. Inside title-page, Boston, 1759. (Sce appendix, pp. 39-52.)
[.... ] A Famous comet. Quarterly Review, 188: 113-138, London, 1808.
This is a very readable account of Halley’s comet.
Part II. Comet’s 1910 RETURN
Astronomisches Gesellschaft. Vierteljahrsschrift. 1910 Return of Halley’s comet. Vierteljahrsschrift der Astronomischen Gesell- schaft. 39 Jahrgang, drittes Heft, pp. 149, 152, Leipzig, 1904. Contains the announcement of a prize of 1000 Mark, offered by the As-
tronomisches Gesellschaft, “for the best determination of the positions of
Halley’s comet for the year of its return.” See The Observatory, 28: no.
355 (March, 1905) 141.
Deminc, W. F. The meteoric shower of Halley's comet. The Journal of the British Astronomical Association, 12: 175-176, 288-289, London, 1902.
Relates to comet’s 1910 return.
Lynn, W[iLt1AM] T[HYNNE]. Halley’s comet. Notes and queries, tenth series, 1: 152, London, February 20, 1904. Note referring to Pontécoulant’s prediction of next perihelion passage “17 May, 1910”; also to the same astronomer’s investigation as to “the position of the comet at the preceding return.”
McPIKeE, EUGENE FarrFIELD. Halley’s comet. Notes and queries, tenth series, 1: 86, London, January 30, 1904. :
Note containing an intimation that ‘“malheureusement la tache entreprise
ne puisse pas étre accomplie” by the Russian Astronomical Society, in re- spect of its computing bureau’s proposed “calculation of the true path of Halley’s comet, with a view to predicting the exact date of the next return.”
PONTECOULANT, Count M. G. DE. 1766-1853. Notice sur la comeéte de Halley et ses apparitions successives de 1531 a IQIO. Comptes Rendus Hebdomadaires des Séances de l Academie des : Sciences, LVIII., 706-709, 766-769, 825-828, Paris, 1864.
Pontécoulant predicts the next perihelion passage for 1910, May 16.95, Paris meridian time.
Smart, D. Halley’s comet: 1910 return. The Journal of the Brit- ish Astronomical Association, 12: 134-136, London, 1902.
MCPIKE] HALLEY’S COMET 73
[....] The next return of Halley’s comet. Nature, 11: 286-287, London, February 11, 1875.
Note consisting of a short review of Pontécoulant’s prediction of the re- turn of Halley’s comet, 1910.
[2352.) Toro Return of Halley's comet. Nature, 49: 442, Lon- don, March 8, 1894.
“Prof. Glasenapp announces that the computing bureau established by the Russian Astronomical Society has undertaken the calculation of the true path of Halley’s comet, with a view to predicting the exact date of the next return.” It is said that a similar notice appeared in Astronomische Nach- richten, No. 3216 (1894).
Part III. BrsLtioGRAPHIES oF Dr. E. HALLEY.
British Museum. Lonpon. Catalogue of printed books. H. Hages, London: William Clowes and Sons, Limited, 1888. See cols. 273-276.
British Museum. Lonpon. Catalogue of printed books. Sup- plement. H-Henrivaux, London: William Clowes and Sons, Limited, 1903. See col. 102.
See, also, later ‘ accessions.’
British Museum. Lonpon. Catalogue of the printed maps, plans, and charts in the British Museum. London, 1885. See [vol. 1, A-K] col. 1733-1734.
C[LterKe] A[cnes] M. Halley, Edmund. Dictionary of National Biography, 24: 104-109, New York, 1890. Biographical sketch, accompanied by a bibliography. A few corrections in the text appear in the ‘Dict. Nat. Biog., 67: Errata.’
McPixe, Eucene FarrFieLp. Dr. Edmond Halley. Notes and queries, ninth series, 10: 361-362; 11: 85-86, 205-206, 3066, 463- 464; 12: 125-126, 185, 266-267, 464-465; tenth series, 2: 224. A bibliography with notes.
RupotpH, ALEXANDER J. AND McPIKE, EUGENE FAIRFIELD. A Bibliography of Dr. Edmond Halley by Alexander J. Rudolph,
_ with some notes and addenda by Eugene Fairfield McPike.
_. . Bulletin of Bibliography, Boston: Boston Book Co., June (?):
1905. To be published.
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
Watt, Rosert. Bibliotheca Britannica; or a general index to British and foreign literature, 4 vols. Edinburgh, 1824. See vol. I, authors, sections 4590—460f, and vol. 3, subjects (no pagina- tion) article Halley, Edmund.
THE ANCESTRAL ORIGIN OF THE NORTH AMERICAN UNIONID/, OR FRESH-WATER MUSSELS
By’ CHARLES A. WHITE
The subordinal group of fresh-water mollusks, the Naiades, includes two recognized families, the Unionide and the Mutelidz. These two families have many essential characteristics in common and, together, they are distinctly separate from all other molluscan families. They are not only peculiar as regards certain portions of their structure and life history but, with few exceptions, also as regards the restrictions of their inhabitation. That is, the Naiades form the great group of mollusks which are commonly known as fresh-water mussels, all of which are confined to a fresh- water habitat and all will die quickly if immersed in salt water, or if removed to the land. This article is written with special refer- ence to the family Unionide, to the geographical distribution of its living representatives, and to the character and succession in time of its fossil representatives in North America. Therefore the Mutelidz, which are far inferior in numbers and variety to the Unionidez, and are confined to Africa and South America, will not be further referred to except in a general way. The following elemen- tary statements concerning the structure and physiological functions of the Unionide are given for the purpose of emphasizing certain of the facts which are to be stated concerning the integral survival of the family through long geological ages, its present separateness from other molluscan families and the wide geographical distribution of its living members; and also to illustrate the characteristics by which the fossil shells of the family are recognized as such.
In a general way, the animal which produces, and is protected by, the shells of the fresh-water mussels is much like that of the common edible clam or, less closely, like the oyster. It is without a proper head, and also without some of the functional organs possessed by other animals; but it performs the function of locomotion, plowing slowly through mud and sand, by means of a muscular projection called the foot; that of respiration by gills, somewhat like those of fishes ; that of circulation by means of a rude pulsating organ which serves as a heart; that of digestion by a stomach; and that of repro- duction by minute eggs. The body, which consists of soft parts
75
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
only, is enveloped in a delicate membrane called the mantle. This organ, although so simple in structure, is a most important one, because, besides other uses to the animal, it forms the shell by secreting a milky substance which exudes, mostly from its free edges, and hardens layer upon layer, until the shells have reached their full thickness and, with the animal, their full size. It is in, or upon, the mantle also that true and valuable pearls are sometimes formed. Although these soft parts differ more or less in details of structure in different genera and species of the Unionide, it is the structure and texture of their shells which are generally used in recognizing their systematic relationships as well as largely in their classification; and it is of course those features and properties of the shell alone that are used in the classification of the fossil species.
In structure the shell of each individual consists of two convex valves which are generally equal in size, and of symmetrical shape. They are held together at their upper edges by a horny ligament, and are also drawn together by the two strong muscles of the animal within. Their joined edges under the ligament are usually provided with interlocking projections, the so-called hinge teeth, but some have no such teeth. The free, or lower, edges of the valves open a little way, and it is upon the forward portion of these edges that the shell rests when it is in its natural position.
The shell-substance consists of three distinct layers, each being of different quality and texture. First, a more or less thick inner pearly layer, which is usually iridescent and often of beautiful tints ; second, a very thin vertically prismatic layer outside of, and firmly adhering to the pearly one; and, third, outside of all, a thinner horny layer, called the epidermis. This shell-structure is the same for all the members of the Unionidz in all parts of the world, and it was the same for all members of the family that have existed in former geological ages, as is shown by their fossil remains.
Although the family characteristics of the Unionide, in whatever part of the world they are found, are clearly defined by structure and shell-texture, the species and genera in certain great regions are dis- tinctly different from those of other regions. The family is of world- wide distribution, representatives of it being found in the fresh waters of all the continents, in those of all the large islands, and in those of some of the smaller sea-girt islands. The number of known species of the family now living in the whole world is about one thousand. Of this number about six hundred species live in North American waters, and of the latter number the Mississippi River system alone contains about four hundred, or about four tenths of all the known species in the world.
WHITE] ANCESTRAL ORIGIN OF UNIONID fd
While the species and genera of the Unionide are different in different regions, the family is so distinct from all other mollus- can families except the Mutelide that naturalists, reasoning from present physical conditions and biological data only, have gen- erally assumed for it a common genetic origin in some one region, its subsequent differentiation, and its final distribution to other regions. Because of the fresh-water requirements of those mol- lusks and the separation by marine waters of the regions which they occupy, and also because there are many cases of intraconti- nental restriction of regional areas of distribution, the question, how the great distribution of the family could have occurred has been a most perplexing one. The marine waters of the earth cover the larger part of its surface and they are everywhere continuous and of essentially the same character. It is therefore easy to understand how any family of marine animals might gain a universal distribu- tion by successive migrations ; but the case is very different with the Unionidz. Since every member of that family dies quickly if placed in sea water or upon land they are confined to rivers and brooks, lakes and ponds and they cannot by their own act pass from one congenial habitat to another, either overland or through marine waters. Voluntary migration from one region to another being out of the question, some method of distribution by agential transportation has been generally advocated. Local dispersion by the shifting of drainage lines has been suggested; and even the inde- pendent origination of members of the family in each region has been assumed. It is desirable to give some account of the views which have been held concerning the geographical distribution of the Unionidz, both for completeness of statement and for the purpose of comparing those views with the hypothesis concerning the geo- logical origin of the North American species which I shall propose.
Those who have suggested the distribution of the family by trans- portation have attributed it to the agency of birds and fishes respec- tively. For all such cases it has been assumed that it was the eggs, or the newly hatched fry, technically called the glochidium, or the byssus-bearing fry, and not the adult mollusks, that have been trans- ported. Immediately after the fry have been hatched from the eggs, and while they are exceedingly minute, some, if not all of the species develop hooklets upon the temporary shell by which each can attach itself to other objects. Ata later stage the fry attaches itself to other objects by a slender, thread-like byssus.t It has been thought that these larval mollusks may become attached to, or
‘I make this statement from personal observation many years ago.
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
entangled upon, the feet of aquatic birds and carried by them in their flight from the fresh waters of one region to those of other regions and there set free. This suggestion seems to be plausible but no known migratory range of aquatic birds will connect any con- siderable part of the regions of the earth which the Unionide are known to inhabit. Besides this all the species, and even the genera, of those mollusks which live in certain of the widely separated regions are different from those of other regions, and those differ- ences are not accounted for in the supposition that the one region was stocked from the other. Moreover, very little interchange of species seems to have occurred under primeval conditions in cer- tain of the intracontinental regions which are constantly visited and revisited by aquatic birds. For example, the Mississippi and St. Lawrence river systems closely approach each other by some of their head waters, and yet each system originally contained a very differ- ent Unione fauna from that of the other, although the annual migra- tory range of millions of aquatic birds has for centuries traversed both regions. It is true that a small number of species are now known to inhabit both of those river systems, most of which prob- ably owe their double habitat to the agency of man. It is also likely that the traffic-canals which are now constructed or projected will increase the number of emigrants from each fauna.
The suggestion that fishes have been instrumental in the distri- bution of the Unionidee refers to the glochidia, or minute fry, before mentioned. It is well known that these minute larval mollusks attach themselves to fishes which live in the same waters, and that thev burrow into their skin, where they become hermetically encysted for further development. Migratory fishes coming from the sea into fresh waters to spawn may thus become infested if the spawning season of the mollusks and fishes should be approximately coinci- dent. If such fishes should return to the sea bearing in their skins the hermetically encysted parasites, and then enter another river with them, it has been thought that the young mollusks might escape into the new congenial waters and stock them with their kind. Unfortunately for this suggestion the encysted term for the young mollusk is only about seventy days, while the fishes would not nat- urally return from the sea to fresh waters before the spawning season of another year. Even then they would be much more likely to return to"the same river again than to enter any other. Long before that time the young mollusks would have dropped from their cysts and died in the salt water.
The suggestion that the dispersion of the Unionide has been
WHITE] ANCESTRAL ORIGIN OF UNIONIDE 79
effected by changes in the direction of drainage, caused by physical changes in the land surface, seems to be applicable to certain cases, but it is of course not of general application. For example, the Upper Mississippi and the Red River of the North have a closely similar Unione fauna, although the one empties into the Gulf of Mexico and the other into the Arctic Ocean. Their head waters are now far apart, and the land surface between them has only slight elevation. This suggestion is pertinent to the hypothesis which I shall present concerning the survival of the Unionide through suc- cessive geological periods.
Only one more of the suggestions that have been offered in expla- nation of the manner in which the present distribution of the Unionide has been accomplished will be noticed. Indeed, this one is of so improbable a character that it is presented only to show what extreme views have been held upon this subject. This sug- gestion is that the Unionide of every river system which they in- habit originated independently from somewhat similar molluscan forms that existed in marine waters near the river mouths, that those mollusks entered the rivers, acquired the characteristics of the fresh water family, and differentiated into new species-and genera. If one should consider this suggestion seriously it may be remem- bered that many of the rivers which contain closely similar Unione faunas flow into arctic and tropical seas respectively, and that the molluscan faunas of those seas are correspondingly different. Also that some of the rivers which contain species of the Unionide and other fresh water gill-bearing mollusks flow into inland seas, the character of whose waters is such that no molluscan life can exist in them. Such, for example, as the Jordan, flowing into the Dead Sea, and the Bear and Utah rivers flowing into Great Salt Lake. The primary origin of fresh water mollusca from certain marine forms that became land-locked in local waters which gradually freshened as the surrounding land was elevated above sea-level, is of course admitted. But that a distinct and well characterized family of fresh water mollusks could have originated from among incon- gruous marine faunas at a multitude of distinctly separated centers, and entered the.rivers by self migration, is not to be accepted as a rational proposition. .
The attempts that have been made to explain the manner in which the present distribution of the Unionidz has been accomplished are not only defective from a biological point of view, but none of them has had special reference to fossil Unione faunas. I shall presently show what I regard as good evidence that certain North American
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
fossil faunas are ancestrally related to the living fauna of the Mis- sissippi River; and by inference that other living faunas had a like ancestral origin.
Of late years students of the living North American Unionidze have recognized among the abundant species a considerable number of genera; properly basing their determinations largely upon the structure of the animal itself as well as upon that of the shell, and also to some extent upon group-differences that were formerly much overlooked. The earlier North American naturalists, however, clas- sifying those mollusks by means of the shells alone, usually recog- nized only three genera, namely, Unio, Anodonta and Margaritana or Alasmidonta, many of them having regarded the latter name as only a synonym of Margaritana. In studies of the fossil Unionide one is necessarily confined to the shells alone; and the fossil material which is available is usually insufficient for the recognition of such groups of species as are recognizable among living faunas. Because of these facts, and partly from a long established habit, I have retained that older classification in my studies of the fossil species. As the character of this article does not really require it, I do not now make any special reference to the improved classification.
Although the Mississippi fauna contains about four hundred spe- cies only about a dozen of them are referred to the genus Anodonta. Their preferred habitat is in still waters apart from the two other genera, and their shells are all of plain, simple type. The species that are referable to the genus Margaritana, including Alasmuidonta, in the same fauna are less in number than are those of Anodonta. They live in immediate association with Unio, and their shells have considerable diversity of form and surface features. It is therefore almost only among the teeming species of Unio that occurs the great variety of form and surface features by which the shells of these mollusks have given expression to what naturalists have long recognized as North American types of the Unionidz. I shall show that this term is properly so applied, not only because these Missis- sippi River types are different from those which are found among the living members of the family in other parts of the world, but because they evidently have been derived from ancient North Amer- ican ancestry. The illustrations upon the accompanying plates ex- press this prototypal character of the fossil species, so far as is practicable by such means, with the aid of the material that has hitherto been discovered.t It is only claimed that the expression
*The specimens from which these figures were drawn are all the property of the U. S. National Museum.
WHITE] ANCESTRAL ORIGIN OF UNIONID/E 81
given by these illustrations is of a general character, but one who is familiar with the living fauna of the Mississippi River will not fail to recognize a close similarity of some of its members to certain of the fossil species. Full artificial expression of the general relation- ship that exists between these fossil species and those which are now living in the Mississippi River system would require a large number of figures of the living, as well as of the fossil, species. As such a full illustration is, for obvious reasons, not now practicable, the reader is referred to the publications mentioned below’ or, better still, to the mollusks themselves in their native waters.
Before proceeding with special references to the figures upon the accompanying plates and to the fossil species which they represent, some explanation of relevant paleontological and geological facts in their relation to ancient physical geography is necessary. For the sake of brevity these explanatory remarks are mostly made in sen- tential, rather than in strictly consecutive, form.
Fossil shells of the Unionidz have been discovered in great num- bers and variety in many parts of the world and in formations of various geological periods. They are found imbedded in more or less hardened rocky strata that originally consisted of muddy or sandy sediment at the bottom of bodies of fresh water. Those lacustrine waters were finally shifted to other areas by oscillations of land surface or drained away by the deepening of the channels of outlet, but they left an unmistakable record of their fresh-water character in their fossiliferous sediments, which remained. So re- stricted are the living Unionide to fresh waters, and so distinctive are the shell characters and the shell texture of all the members of the family, that the geologist is as certain that the strata containing their fossil remains were deposited in fresh, and not in marine, waters as if he had then been there and analyzed them. Moreover, there are usually found with the fossil shells of the Unionide the shells of other mollusks which are similar to those of the associates of their living congeners.
The existence of a lake, or a body of fresh water, implies the coexistence of a surrounding land surface upon which flow drainage streams of inlet and outlet. The existence of a stratified deposit or formation containing remains of fresh water mollusks implies that the
* Observations on the Genus Unio. By Isaac Lea. Vols. xin quarto. Profusely illustrated by full-page plates, part of which are colored. Synopsis of the Naiades, or pearly fresh-water Mussels. By Charles Tor- rey Simpson. Proc. U. S. National Museum, vol. xxu, pp. 501-1044, and plate xvi. The literature of the Unionide is very extensive. That for North America is catalogued by Mr. Simpson in the forementioned work.
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
deposit was made in lacustrine waters, and that such a surrounding land surface as that just mentioned existed at the time the deposit was made. The study of North American geology has revealed the pres- ence, especially in the broad interior region of the continent, of many such lake deposits containing remains of the Unionide, the earliest one of which that will be referred to being of Triassic age. These deposits alternate with marine formations, showing that the conti- nent has risen by repeated oscillations of the land surface with rela- tion to sea-level, and not by one uniform upward movement extend- ing through successive geological ages. The aggregate gain of these oscillatory movements is the present elevation of the continent.
The first land that appeared above sea-level was drained of its surface waters by brooks; and as the land increased in extent the waters of the brooks increased in volume and became rivers. The unequal elevation of continental land occasionally caused broad de- pressions of its surface, which filled with drainage water and became lakes. Each lake, together with its outlet and inlets, became stocked with a fresh water fauna which was derived from some pre- existing fauna. Because all existing lakes and rivers contain mol- luscan life, and because all lacustrine deposits contain remains of such life, it is necessarily inferred that formerly existing lakes and rivers were stocked in like manner.
Lakes are parts of unfinished river systems. The deepening of the outlet portion of such a river system by its running water, aided by sedimentation in the still water, drains the lake and finishes the river system. For example, referring to existing rivers, all parts of the Mississippi system are finished except the slight expansion called Lake Pepin. The St. Lawrence system is very far from finished because of the great, and many smaller, lakes that still remain in both its principal and subordinate courses.
As a rule, abrupt land elevations, including mountain ranges, which have resulted from foldings and other displacements of the earth’s crust, have risen so slowly from previously plain regions, that the rivers which were already established there were not only not thereby obliterated, but usually they were not even materially deflected from their courses. By the corrasive action of its running water and the detritus which it carried by its flow, each stream abraded and carried away the earth-material, even including solid rock, as it slowly rose beneath its channel. Some of the now ex- isting rivers have thus made deep cafions with precipitous sides, through the rocky strata of elevated regions, and some have even cut their way through mountain ranges. The cafion sides represent
WHITE] ANCESTRAL ORIGIN OF UNIONID& 83
the rising of the land, not the lowering of the river. The cafion of Green river through the Uinta mountain range, and the Grand Cafion of the Colorado of the West through the Great Plateau, are cases of this kind. Still, some rivers have suffered vertical displace- ments in at least parts of their course. For example, the prolongation of the channel of some existing rivers of North America, is trace- able by soundings beneath sea-level, where they sank by subsidence of the continental border. If that border should be raised again such rivers and their faunas would come into their former posses- sions. At the beginning of the Tertiary period the Upper Mississippi and Ohio rivers emptied separately into the Gulf which then extended northward above the present confluence of the two rivers. That is, the whole of what is now the Lower Mississippi was then beneath sea-level. It has since been added to the upper portion of the great river system and stocked with its fauna.
These, and many other similar facts show that rivers, once estab- lished, although often modified in extent by land elevations and subsidences, and changed in direction by the opening of new lake outlets, have been among the most persistent features of the earth’s surface. The lakes which occupied portions of the course of ancient rivers have all been obliterated; and doubtless also in rare cases some rivers or small river systems, with their molluscan faunas, have been wholly destroyed. The facts which have been stated, however, warrant the assumption that, as a rule, some portions of those an- cient rivers have preserved a continuous flow of fresh water to the present time. I do not doubt that at least some portions of the present Mississippi River system represent a continuous fluvatile flow from a time at least as remote as the Cretaceous period. Rain waters have always fallen upon the land ever since its first elevation above the sea, and a constant flow of drainage streams has been necessary to remove it. It is only by a constant flow that genetic lines of fresh water denizens could have been preserved ; and I there- fore assume that the Unione fauna of the Mississippi River system has in this way been, at least in part, genetically derived from the fossil faunas some of whose remains are figured on the accompany- ing plates.
Some of the types of former fresh water denizens whose remains have been discovered are not found among living faunas, and it is therefore inferred that these were among the faunas of those rivers which failed entirely to preserve their continuity of flow through successive geological periods. For example, although Unio belliplicatus, which is represented by figures 4, 5 and 6, on plate
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
XXVIII, has all the structural and textural characteristics of the genus _ Unio, it is not only the earliest known species of that genus to possess well marked surface ornamentation, but its type of orna- mentation is different from that of any known living North Amer- ican species. Besides this, several species of the gasteropod mollusks which are associated with this Unio are also different in certain characteristics from any of their kind upon this continent, either fossil or living. Moreover the Bear River formation, in which this fossil fauna is found, is of small extent compared with the other North American fresh water formations. From all these facts I infer that the body of water in which the Bear River beds were deposited, together with its inlets and outlet, constituted a small separate river system with a distinctive fauna. Also that its case was an exception to the rule of the persistence of the rivers, and that this whole small river system with its fauna became destroyed by some geological disturbance of the land surface. The types of the Bear River fauna which were not thus destroyed, for example, the simple type of Unio nucalis, which existed before, and have existed ever since the Bear River epoch, were probably preserved in other bodies of fresh water by collateral lines from an original genetic source. These remarks upon ancient physical geography may be closed with the following summary statements, together with references to the figures upon the accompanying plates and to the species which they represent.
Fresh-water gill-bearing faunas have as certainly descended genetically through successive geological ages to the present time as have marine faunas. The genetic successors of each fauna have necessarily descended in a continuous fresh-water habitat. Such continuity of habitat has been produced and preserved by the sea- sonal rains which have always fallen upon the land and caused a constant drainage flow in its rivers and their branches. There has never been any intermission of such continuity because the fresh water supply has never failed, and because, as a rule, rivers have been among the most persistent of the earth’s surface features. While some rivers, or small river systems, have doubtless been from time to time destroyed by certain special movements of the earth’s crust and their peculiar faunas utterly exterminated, it is not prob- able that through all the great vicissitudes of continental devel- ment any greater proportion of fresh-water types have been thus destroyed than of marine types which have perished by volcanic eruptions, local elevation or depression of sea-bottom, changes of sea-currents, and other causes.
WHITE] ANCESTRAL ORIGIN OF UNIONID 85
Measured geologically, the life-time of species as such has been short, but genera, and the types which they embrace, have per- sisted through successive geological ages. The types of Unio which are represented on the accompanying plates have been thus preserved, while the species which successively bore them became extinct in the successive geological opochs. They so much resemble certain members of the living Mississippi river fauna as to warrant the assumption that the fossil faunas represent the living fauna ancestrally.
Many specimens of fossil shells of the Unionide have been dis- covered in the Triassic strata of New Mexico and Wyoming. All of them are very imperfect and of comparatively small size, but they unmistakably belong to the genus Unio. One of these Triassic specimens is represented by figure 1, plate xxvi. The specimens re- ferred to are the earliest of the certainly known examples of the Unionidz in North America, although certain shells found in De- vonian and Carboniferous rocks have been supposed to belong to that family. These Triassic shells are all of simple form, and none of them exhibits distinctive prototypal relationship to the living Mississippi River fauna. Their structure and shell texture, how- ever, clearly show that the genus Unio was fully established at that early period; and their wide distribution indicates that a large Unione fauna was then established.
In all, seven species of the Unionide have been discovered in the fresh water Jurassic strata of Colorado, Wyoming and South Dakota. All of them belong to the genus Umio, and five of the seven species are represented on plates xxvr and xxvii. They are all of simple, plain types, none of them exhibiting any special relationship to the Unione fauna of the Mississippi, unless it be U. stewardi. It is, however, not improbable that all these species, as well as those found in Triassic strata, are ancestrally related to the simpler forms of the Mississippi fauna.
While there evidently was a large representation of the Unionidze in the Triassic and Jurassic periods, it was in the closing period of Mesozoic time, the Cretaceous, that the family received an extra- ordinary development. This fact is shown by the discovery at numerous places within a large geographical area, and in several successive formations, of a large number and great variety of fossil species of Unio, and of the addition among them of a few species of Anodonta and Margaritana. The increased diversity of the Unionide in this period is also shown in the exhibition by many of the species of Unio of those peculiarities which I have designated
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS [voL. 48
as North American prototypal characteristics. These discoveries of Cretaceous species have been made in the states of Colorado, Utah, Wyoming, South Dakota and Montana; and in the Canadian territories of Alberta, Assiniboia and Saskatchewan. In vertical range these discoveries extend from the base to the top of the Creta- ceous series of formations as it exists in the great region just indi- cated. The formations, or groups of strata, are, beginning with the lowest, the Dakota, Colorado, including the Bear River beds, Pierre, including the Fox-Hills, Judith River and Belly River beds, and the Laramie. The Dakota group has furnished comparatively few molluscan fossils, and the most that need now be said of it is that it is not of marine origin. The Colorado and Pierre formations con- sist mainly of unquestionably marine strata, with which the fresh water groups alternate. The Laramie is the uppermost formation of the Cretaceous series and the character of its molluscan fauna gives evidence that it was deposited in a body of water that was in part fresh and in part brackish. This formation aso contains plant remains which have been referred to the Tertiary; and dinosaurian remains which are regarded as of Cretaceous age. I now provision- ally refer the formation to the latter age, although its molluscan fauna might with propriety be referred to the Tertiary. It is in the Laramie strata that the greatest number of species of Unio have been found that bear the prototypal features which have been frequently referred to. Most of these species were found in a few fossiliferous layers of limited extent, each of which was probably deposited near the mouth of an inlet and not in the stiller waters of the lake. The formation from which each of the species repre- sented upon the accompanying plates were obtained is noted upon the page of explanations which accompanies each of the plates. Besides the species which are referred to in the foregoing para- graphs and figured on the accompanying plates, Professor R. P. Whitfield has published descriptions and figures of six new species
of Unio which were discovered in strata of the Laramie Group of
Montana, and which he has named as follows: Unio e@sopiformis, U. verrucosiformis, U. retusoides, U. browni, U. percorrugata, and U. postbiplicata. All these fossil species present prototypal char- acteristics of the living Mississippi Unione fauna in a marked degree. Three of them are so closely like three living species re- spectively that Professor Whitfield has given names to the fossil
*“" Notice of Six New Species of Unios from the Laramie Group,” by R. P. Whitfield, Bull. Am. Museum of Nat. Hist., vol. x1x, pp. 483-487, plates XX XVIII-XL.
a
WHITE] ANCESTRAL ORIGIN OF UNIONID/E 87
forms which are only modifications of the names of the living forms which they so closely resemble. One cannot doubt that further dis- coveries will yield additional evidence of the prototypal relationship of the fossil and living Unione faunas of this continent.
Following the Laramie in the order of time and of geological sequence, are the Eocene, Miocene and Pliocene Tertiary formations, all three of which, in the great interior region of North America, consist of fresh water lacustrine deposits. From the fact that the Laramie Group has been found to contain so many prototypal exam- ples of the North American Unionide one might naturally expect to find among the Tertiary molluscan faunas numerous species of Unio that would, by similar prototypal features connect the Laramie forms more or less directly with the living Mississippi River fauna. Such, unfortunately, is not the fact, for only a few species of the Unionide have been found in any of those Tertiary deposits, and they are all of simple type and plain surface. If only such plain forms of Unio really existed in those Tertiary waters between the Laramie period and the present time, my assumption of the ances- trally prototypal character of the Cretaceous Uniones would be unsupported. Without any exception known to me, however, the