^0 HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Bulletin' of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 1. NEW AND OLD SILURIAN TRILOBITES FROM SOUTH- EASTERN WISCONSIN, WITH NOTES ON THE GENERA OF THE ILLAENIDAE. By Percy E. Raymond. With Four Plates. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM. January, 1916. No. 1. — New and old Silurian Trilobites from Southeastern Wisconsin, with Notes on the Genera of the Illaenidae. By Percy E. Raymond. In the F. H. Day collection, received in January 1881, as a gift of Mr. Alexander Agassiz, the Museum of Comparative Zoology secured one of the three great collections of the Silurian fossils of southeastern Wisconsin. It is particularly rich in Illaenidae, but contains also such rare forms as Harpes telleri, Trochurv^ nasutus, and Dicrano- peltis telleri, to be found elsewhere only in Mr. Teller's magnificent collection. The trilobite fauna of the quarries around Milwaukee and Racine differs considerably from that found in the vicinity of Chicago, so that some of the species described by Weller from the latter area are either absent from the M. C. Z. collection or represented by speci- mens from other sources than the Day collection. The Phacopidae have been omitted from the present study, and only such species are mentioned as are represented by specimens which add something to what has already been published. ILLAENIDAE Hawle and Corda. The Illaenidae form a remarkably homogeneous group, and in spite of the great number of species which have been described only three generic or subgeneric names (Illaenus, Bumastus, and Thaleops) are in common use. Holm recognized only Illaenus and Bumastus and other writers have been even more conservative, referring all the species to Illaenus. The only serious attempt to subdivide the genus is that made by Salter, who recognized eight subgenera (including Illaenus), but none of Salter's names has ever come into general use, although some of them could be adopted advantageously. In all, seventeen subgenera or genera have been proposed for inclusion in this family, but only seven of these seem to be valid. The names, in chronological order, are: — Cryptonymus Eichwald 1825, Illaenus Dalman 1826, Deucalion Shtsheglov 1827, Bumastus Murchison 1839, Archegonus and Dysplanus Burmeister 1843, Thaleops Conrad 1843, Alceste Hawle and Corda 1847, Rhodope Angelin 1854, Actino- 4 bulletin: museum of comparative zoology. lobus Eichwald 1860, Panderia Volborth 1863, lUaenurus Hall 1863, Illaenopsis Salter 1S66, Ectillaenus, Hydrolaenus, Octillaenus Salter, 1867, and Illaenoides Weller 1907. Certain other genera, such as Symphysurus and Nileus have often been placed with lUaenus, but they have more generally been recognized as belonging to the Asa- phidae. The fundamental difference between the lUaenidae and the Asaphidae is, of course, the presence in the former family of an epistoma, and its absence in the latter. The absence of the grooves on the pleura of the thorax in the Illaenidae and their very general presence in the Asaphidae affords quite a safe criterion for judging of the dorsal surface. The genera enumerated above may be taken up in order. Cryptonymus Eichwald, 1825. Observations geognostico-zoologicae per Ingriam marisque Baltici Provincias nee non de Trilobites. Casani, 1825, p. 44. In this paper, Eichwald, evidently not understanding Brongniart's genus Asaphus, describes the new genus Cryptonymus, and under it, eight species. The first four are species of Asaphus and the last four are species of Illaenidae. Asaphus had been described in 1822, but Illaenus was not published till 1826, so that, if the four asaphids were eliminated, it would really leave an illaenid as the type of Cryptony- mus. The first of the illaenids described, Cryptonymus roscnhergi, is a Bumastus, the second, Cr. wahlenbcrgi, third, Cr. rudolphii, and the last, Cr. parkinsonii, are all species of Illaenus. To choose among these a type for Cryptonymus would be to upset one of two well- established names without any possible gain. Eichwald himself ac- cepted Illaenus as the name of these species, and in 1840 transferred his name Cryptonymus to Trilobites pimcfatus Wahlenberg, a pro- ceeding which he afterward stoutly defended (1855). If a new generic name is at any time necessary for any of the eight species described by Eichwald, Cryptonymus is still available, but till such a con- tingency arises, it seems best not to revive the name. Illaenus Dalman, 1826. Type, Asaphus crassicauda Wahlenberg. Om Palaeaderna eller de sa kallade Trilobiterna. Kgl. Vet. akad. Handl., 1826, p. 248. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 5 Dalman divided Asaphus into four sections, the third of which he designated as Illaenus, using the name in a subgeneric sense. lUaenus he divided into two divisions. Divisio 1, Cornigeri, contained the single species Asaphus {Illaenus) centrotus, which Burmeister made the type of Dysplanus in 1843. Divisio 2, Mutici, contained Asaphus {Illaenus) crassicauda and A. {Illaenus) laticauda. The first of these has by general consent been made the type of Illaenus. Holm has, in several papers, redescribed the typical species, so that its charac- teristics are well known. Beside the characters of the family, the species shows a short and wide, strongl}^ convex and curved cephalon and pygidium, both without concave borders, large prominent eyes which are situated far back, deep but short glabellar furrows, short but rather wide free cheeks without spines on the genal angles. The thorax has a narrow axial lobe and ten segments. The pygidium has a short but prominent axial lobe, and is wider than long. As Clarke has pointed out, Illaenus americanus Billings is an American species which is very similar to /. crassicauda, and it seems that only such species as conform to the kind of structure exhibited by the type should be admitted to the restricted genus Illaenus. Deucalion Shtsheglov, 1827. Sur les Trilobites en general et en particulier sur ceux de Zarskoe- Selo. Journ. fur neue Endeckungen in der Phys. Chem. Natur. und Technologic St. Petersburg, 1827, no. 1, 2, p. 234, pi. 7, f. 9 a-c. I have not seen this paper, but judge from what Holm says that Deucalion is a svnonvm of Illaenus. The genus was founded on a new species, D. hrongniarti, which Holm was unable to recognize. BuMASTUs Murchison, 1839. Type, BuMASTUS barriensis Murchison (Partim). Silurian system, 1839, p. 656 (non figs.). The particular features of this genus upon which Murchison himself laid most stress were the absence of dorsal furrows, and the presence of ten segments in the thorax. The general usage, however, has been that of referring all illaenids having the axial lobe of the thorax very 6 bulletin: museum of comparative zoology. broad, its width equaling or exceeding one half the total width of the body, to the genus Bumastus. This practice will probably prevail, for we now know that other illaenids beside Bumastus have ten seg- ments in the thorax, and there is no illaenid known from which dorsal furrows are absolutely absent. In view of the somewhat numerous subdivisions of the illaenids, it may be well to reexamine the type- species, Bumastus barriensis Murchison, in a little detail. Salter has explained that the specimens figured by Murchison really do not belong to this species, so that Ave are obliged to use Salter's figures of the "Barr Trilobite." Fortunately the M. C. Z. collection contains two plaster casts of the original specimen figured by Jukes in 1829 and later by Salter. The casts are rather carelessly made, but are in general in fair agreement with Salter's figures. From these sources may be derived the statement that the typical species of Bumastus is a large Silui'ian illaenid with smooth, subequal cephalon and pygi- dium, rounded, spineless genal angles, large eyes, situated near the posterior margin of the cephalon, a very wide axial lobe, shallow dorsal furrows, ten segments in the thorax, and no trace of an axial lobe on the pygidium. The dorsal furrows on the cephalon are short, extend- ing but little ahead of the eyes. The cephalon does not appear to have any rim or concave depression, but the pygidium shows a slight concavity, so that the profile of that member does not present a smooth convex curve, but the curvature is reversed near the posterior end of the pygidium. Both cephalon and pygidium are wider than long. Archegonus Burmeister, 1843. Die organisation der trilobiten, 1843, p. 120, 121, pi. 5, f. 3. The type of this genus is Calyviene ? aequalis H. von Meyer, as this was the only species cited by Burmeister in the first edition of his Organisation der trilobiten. As the type is evidently not an illaenid, but one of the Proetidae, the genus automatically disappears from the family. Dysplanus Burmeister, 1843. Die organisation der trilobiten, 1843, p. 120. Type, Illaenus centrotus Dalman. The type of this genus is an illaenid with rather long, parabolic head and abdomen-shields, spines at the genal angles, small eyes far RAYMOND: NEW AND OLD SILURIAN TRILOBITES. / back, narrow axial lobe, and nine segments in the thorax. The profiles of both shields are rather flat but uniformly convex curves. (See especially Holm, Bihang Kogl. Vet. akad. Handl., 1883, 7, pi. 4, f. l-l2). Holm was not able to see any value in this genus, pointing out that the only real characteristic brought forward by Burmeister and by Angelin was the presence of genal spines, and the species with genal spines are so highly variable among themselves as to suggest that this character in itself does not denote any real relationship. It seems, however, that a certain group of illaenids can properly be de- noted by this term, and the genus will be referred to later. Thaleops Conrad, 1843. Proc. Acad. nat. sci. Phil, 1843, 1, p. 331. Type, Thaleops ovata Conrad. The presence in this species of very high eyes on long peduncles, long narrow genal spines, deep dorsal furrows sharply delineating a prominent glabella, a narrow axial lobe, ten thoracic segments, and a small short pygidium, mark an unusually well-defined genus which seems to be confined to North America, and probably to the Ordovi- cian, though one Silurian species has been referred to the genus. Alceste Hawle and Corda, 1847. Prodr. monog. Bohm. tril., 1847, p. 66, pi. 4, f. 31.. Type, Alceste latissima Hawle and Corda (which is the same as lUaenus hisingeri Barrande, according to Barrande). This genus is not valid, for it was based upon an immature speci- men showing only four segments, and was very incorrectly described and figured. Without Barrande's explanation, no one would be able to identify the Alceste laiissima with any Bohemian trilobite. Rhodope Angelin, 1854. Pal. Scandinavia, 1854, pt. 1, Trilobita, p. 38, pi. 22, f. 17. Type, Rhodope lineata Angelin. The name Rhodope was used for a gastropod by von Siebold in 1848 (Anatomic, p. 296), and Volborth replaced the name by Panderia. 8 bulletin: museum of comparative zoology. AcTiNOLOBUS Eichwald, 1860. Type, Illaenus atavus Eichwald, 1857. Lethaea Rossica, 1860, 1, p. 1488. The type is an illaenid which seems sufficiently peculiar to deserve a distinct generic name. The cephalon is short and the pygidium long, and both cephalon and pygidium have a-concave border; the cephalon a narrow lip, and the pygidium as wide a border as the average Isotelus. The eyes are rather large, far back and far apart, free cheeks small, genal angles rounded. The dorsal furrows of the cephalon are short, the axial lobe of the thorax is narrow; ten segments are present; and the axial lobe of the pygidium is short and triangular. Actinolobus ataviis is a Russian Ordovician species (Cla), and another species with a wide border on the pygidium is the one from the Silurian described by Schmidt as Illaenus masckei (From F, Estland). Panderia Volborth, 1863. Type, Panderia triquetra Volborth. Mem. Acad. imp. sci. St. Petersburg, 1863, 6, no. 2, p. 31. Although proposing this name primarily to replace the preoccupied Rhodope of Angelin, Volborth made his own new species the type, and the genus must rest upon it. Holm does not actually use Panderia, but he seems to have considered it a fit receptacle for the group of small trilobites with only eight thoracic segments, and gives (1883, p. 161) a new definition according to his interpretation of the genus. The presence of only eight segments in the thorax does not appeal very strongly to the present WTiter as a generic characteristic. Panderia triquetra does, however, present some rather unusual charac- teristics in its very short, strongly convex cephalon with extremely large eyes, the high, well-defined glabella, and the short pygidium with long, prominent axial lobe. Species of this type are not at all common, and may be referred to Illaenus without doing violence to the definition of that genus. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. \f Illaenurus Hall, 1863. Type, Illaenurus quadratus Hall. 16th Rept. N. Y. state cab. nat. hist., 1863, p. 176, pi. 7. Although Hall believed this species to be closely allied to Illaenus, as indicated by the name, it seems more probable that it belongs to the Asaphidae and is allied to Symphysurus. Illaenopsis Salter, 1866. Type, Illaenopsis thomsoi^i Salter. Mem. Geol. surv. Gt. Britain, 1866, 3, p. 256. As has been repeatedly pointed out, the grooved pleura of the thorax of this trilobite exclude it from the Illaenidae, and place it near Symphysurus in the Asaphidae. OcTiLLAENUS Salter, 1867. Type, Illaenus hisingeri Barrande. Monog. Brit. Silurian trilobites, 1867, pt. 4, p. 182. This genus was erected by Salter to contain the type, a species in which the pleura of the first thoracic segment are produced into spines. There are eight segments in the thorax, the axial lobe is narrow, the glabella well defined, eyes of medium size and far back, free cheeks with sharp genal spines Pygidium about as long as wide, without defined axial lobe. This species could probably be placed with Dysplanus, but I would follow Salter in the recognition of the remark- able development of spines on the first thoracic segment, it being a unique example of such a characteristic among the smooth trilobites. Ectillaenus Salter, 1867. Type, Illaenus perovalis Murchison. Monog. Brit. Silurian trilobites, 1867, pt. 4, p. 182. Holm has pointed out that in proposing this genus Salter confused the true /. perovalis of Murchison and a new species afterward de- 10 bulletin: museum of comparative zoology. scribed by Hicks as /. hughesi, and not understanding clearly the characteristics of either species, produced a name of no particular value. The name should be dropped, unless it can be shown to be of more value than now appears to be probable. Hydrolaenus Salter, 1867. Type, Illaenus conifrons Billings. Monog. Brit. Silurian trilobites, 1867, pt. 4, p. 182. The type-species is a Thaleops, and Hydrolaenus is therefore a synonym of that genus. Illaenoides Weller, 1907. Type, Illaenoides trilobus Weller. Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 226. The type of this genus is remarkable chiefly for its small eyes which are situated halfway to the front of the head. The glabellar furrows are narrow and shallow, the facial suture cuts the cheeks very close to the genal angles, making the free cheeks of unusual shape. The genal angles are rounded. The axial lobe of the thorax is wider than in typical Illaenus, but less wide than in Bumastus. The pygidium is long with a narrow concave border, but no trace of an axial lobe. Type and only known species from the Niagaran at Bridgeport (Chi- cago), Illinois. Su7nmary. — It appears that of the genera proposed, Illaenus, Bumastus, Thaleops, Actinolobus, and Illaenoides have unquestion- able value. Dysplanus and Octillaenus are more or less valuable but need further study and redefinition. Cryptonymus, Deucalion, Panderia, and Ectillaenixs are names which cannot be used at present, but might possibly be revived. Archegonus, Illaenurus, and Illae- nopsis belong to other families. Alceste was never properly defined. Rhodope was preoccupied, and Hydrolaenus is a synonym. It is in- teresting to note the lapse of forty years between Salter's new names and the next generic name applied to a member of this group. It is evident that the type-genus Illaenus contains the great majority raymomd: new and old Silurian trilobites. 11 of the species in this family, the other genera having, as a usual thing, only from one to two or three species each. Bumastus comes next to lUaenus in the number of species, and has its greatest development in America where there are at least six species in the Middle Ordovician and about fifteen in the Middle Silurian, as contrasted with three or four species in the Silurian of Great Britain, about the same number in Scandinavia and Russia, and two in Bohemia. Considering the great abundance of the illaenids, we have sur- prisingly little information as to their ancestry or relationships. Following the usual theory, which seems to be borne out by the facts in most cases, one would expect these smooth forms to be the descend- ants of more normal trilobites with glabellar furrows and with ribs on the pygidium. But among all the illaenids there does not seem to be one which shows any trace of ribs on the pygidium, while only a few show indications of glabellar furrows. And such indications of furrows as exist are merely spots or slight depressions on the smooth glabella. The nearest relatives of the Illaenidae are undoubtedly the Goldiidae (Bronteidae) not the Asaphidae, with which family they have usually been classed. The presence in both the Illaenidae and the Goldiidae of an epistoma, similar hypostomas, forward expanding glabella, large eyes which are placed far back, unfurrowed pleura in the thorax, and short axial lobe on the pygidium, indicate a very close relationship, some of these characteristics being apparently too funda- mental to admit of explanation on the ground of parallelism. The Goldiidae, in spite of their specialization, are more like the typical trilobite than the Illaenidae, and it would be natural to place them in the ancestral position. The geological range at once negatives this attempt, for the Goldiidae did not appear until the Middle Ordovi- cian, are very rare in the Ordovician and reached their greatest de- velopment in the Silurian and Devonian. The illaenids, on the other hand, appeared in the basal Ordovician, possibly even in the Cambrian, reached their greatest development in the Middle Silurian and did not survive that period. That the Goldiidae should have been de- rived from the Illaenidae, however, seems highly improbable, for the phylogeny of the former family pursues a normal course, the oldest members of the family being most highly segmented, and the usual ''smoothing out" process producing such (relatively) lUaenus-like species as Goldius dormitzeri, G. campanifer, and G. hrongniarti in the Devonian of Bohemia. If these species, without glabellar furrows and with highly convex almost ribless pygidia occurred in the Ordo- vician, and forms like Goldius hmatvs (Billings) in the Devonian, we 12 bulletin: museum of comparative zoology. should probably conclude at once that the Goldiidae were derived from the Illaenidae. The oldest illaenids which are well known are those which Barrande described from the base of the Ordovician in Bohemia. Of these, Illaenus advena is a quite typical Illaenus, while the other species are peculiar. Illaenus bohemicus, the type of which is in the M. (". Z., was founded on a badlj^ preserved single specimen, which does not seem to be an Illaenus. The pygidium sKows a long, distinct, and ringed axial lobe and if this member were found alone, it would at once be assigned to the Asaphidae. The nine segments of the thorax are, however, without pleural furrows. The cephalon is too poorly preserved to indicate any characters of value. Long wide genal spines are present, the glabella appears to have nearly parallel sides imtil the vicinity of the anterior end is reached, when it expands abruptly. Rather faint glabellar furrows seem to be present. With only this single specimen, it does not seem possible to assign the species to any genus or even family, and it certainly throws no light on the origin of the Illaenidae. Another species from Di, described by Barrande, is the common Illaenus katzeri. This species differs from other illaenids in its eyes. Barrande supposed it to be blind, but Holub has recently shown (Bull, international Acad. sci. Boheme, 1908. German abstract, p. 7, pi. 7) that it has small eyes, situated forward. Both shields are rather flattened, not so strongly incurved as in the typical genus Illaenus, and the pygidium is long and parabolic in outline, thus suggesting Dysplanus. There is nothing, however, to indicate that /. katzeri is primitive, but it seems rather, like the Silurian Illaenoides trilobus, to be a degenerate form. If the genus is divided at all, this species cannot be considered as congeneric with Illaenus crassicauda, and I would suggest that the name Wossekia be applied to it, since Wossek, Bohemia, is the locality from which practically all the specimens have been obtained. Illaenus ])uer seems to be, as Brogger was the first to point out, a Symphysurus. Illaenus calvus has a peculiar marginal rim. The species is known from a single fragment. Illaenus aratus has a narrow glabella, and the eyes are very far apart or absent. It is primitive for an illaenid, but gives no suggestion as to the origin of the group. A imthologic specimen of Illaenus. — Since this paper was written, I have been enabled, through a grant from the Shaler Memorial fund at Harvard. College, to visit northern Europe, and some time was spent in studying the collections in London, Berlin, St. Petersburg, Stockholm, and Christiania. One of the most interesting trilobites seen was the specimen of Illaenus revelensis Holm, which was figured RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 13 Ly Holm in his description of the Russian Ordovician Illaenidae (Mem. Acad. imp. sci. St. Petersburg, 1886, ser. 7, 33, pi. 2, f. 5a). This is the only example I have seen among trilobites of a malforma- tion due to moulting. As partially shown in Holm's figure, there is an impressed line between the facial suture and the dorsal furrow on the left side, which follows exactly the course of a facial suture, even extending across the front of the cranidium, a point not shown in Holm's figure. Across the posterior part of the cephalon, close to the edge, is a furrow marking the posterior edge of the shell at the time of the previous moult. The eye on the left side is smaller than its opposite, and the palpebral lobe is malformed. The cephalon is decidedly unsymmetrical, the left free cheek being drawn backward. All these pathological features seem to be due to the partial retention of the shell at the next previous moult. Classification of the Illaenidae. From the first, all classifications of the Illaenidae have rested mainly upon the number of segments in the thorax, and secondarily upon the width of the axial lobe of the thorax. Thus, Dalman, the describer of Illaenus, separated the species with nine segments from those with ten. Holm, the principal writer upon the genuS; while recognizing only the genus Illaenus and the subgenus Bumastus, divided Illaenus into three groups, those with ten, nine, and eight segments in the thorax. A study of the American illaenids does not favor a classifica- tion of this sort, for it has been repeatedly shown that the nvunber of thoracic segments in species of both Illaenus and Bumastus is variable, even within the limits of a single species. Likewise, the presence or absence of genal spines is not a characteristic justifying, in itself, the . erection of a genus of Illaenidae, for, as has been several times pointed out, species are found with all sorts and conditions of spines, and if all species having this characteristic were to be referred to a single genus, there would be hardly another characteristic common to the assem- blage. The length and convexity of the cephalon and pygidium, the size and position of the eyes, the width of the axial lobe of the thorax, and the shape of the glabella seem of the most importance, but I would also take into consideration the sort of genal spines which may be present. It is still too early to make any natural classification, and the genera here recognized are based primarily upon the more con- spicuous peculiarities of the type-species of each. 14 bulletin: museum of comparative zoology. ILLAENIDAE Hawle and Corda. Opisthoparia with large, convex, nearly smooth, cephalic and abdominal shields. Epistoma large, hypostoma convex, ovoid. Thorax of eight to ten segments, with unfurrowed pleura. Pygidium without ribs, axial lobe short or absent. Ordovician and Silurian. ILLAENINAE, subf. nov. Illaenidae with narrow axial lobe, cephalon and pygidium without concave border. Illaenus Dalman. Cephalon and pygidium very convex, wider than long, abruptly deflected; eyes large and far back, axial lobe of pygidium high. Genal spines, when present, rounded in section. Type, Illaenus crassicauda (Wahlenburg). Ordovician and Silurian. Europe, North and South America, India, and Australia. Thaleops Conrad. Cephalon and pygidium similar to Illaenus, but with eyes on long stalks and elongate genal spines always present. Type, Thaleops ovata Conrad. Ordovician and possibly Silurian. North America. Dysplanus Burmeister. Cephalon and pygidium long and flattened, parabolic in outline, genal spines present, usually flattened. Type, Illaenus centrotus Dalman. Ordovician. Northern Europe. Wossekia, gen. nov. Cephalon and pygidium as in Dysplanus, genal spines absent, eyes small and far forward. Type, Illaenus katzeri Barrande. Basal Ordovician. Bohemia. Octillaenus Salter. Similar to Dysplanus but with rounded genal spines, and pleura of first segment elongated into spines. Type, Illaenus hisingeri Barrande. Ordovician. Bohemia. BUMASTINAE, subf. nov. Illaenidae with (usually) concave border on one or both shields , axial lobe generally wide, though sometimes narrow. • Bumastus Murchison. Axial lobe of thorax equal to more than half the total width; eight to ten segments. Typically, a concave border is present on the pygidium, frequently on both cephalon and pygidium, rarely (in small Middle Ordovician species) without con- RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 15 cave border on either shield. Eyes very large. Type, Bumastus harriensis Murchison. Ordovician and Silurian. North America. Silurian. Europe. Actinolobus Eichwald. Axial lobe narrow, cephalon short, pygi- diura long, with very wide concave border. Eyes large. Type, Illaenus atavus Eichwald. Ordovician, Russia. Silurian. Russia and United States. lUaenoides Weller Axial lobe with a width between that of Actino- lobus and Bumastus, eyes very small and far forward, narrow con- cave border on pygidium. Type, lUaenoides trilobus Weller. Silurian. United States. The above classification is designed to separate the species with long more or less flattened shields from the more typical illaenids with short and abruptly deflected cephalon and pygidium. The first two genera are modifications of the central Illaenus type, the other three of the more flattened Dysplanus group. In defining the subfamily Bumastinae as I have, all the forms with a more or less Isotelus-like pygidium are removed from the typical Illaenus group. Among the small species constituting the earliest of the Bumastinae one finds species like B. glohosus Billings, B. bellevillensis Raymond and Narra- way, and a few others, which lack a concave border. On the other hand, so large a Middle Ordovician Bumastus, as B. indeterviinatvs (Walcott), the type of which is figured (Plate 2) for the first time, has a distinctly concave border. It is very possibly true that the small species mentioned above should be given a distinct name and placed in the lUaeninae, but it still seems somewhat early to take so radical a step. Description of species. We owe to Professor Weller a complete and careful description of the Illaenidae of the Chicago area, and, as he had access to collections made at Racine and near Milwaukee, his description in large measure covers the Wisconsin area also. In the large collections which I have been able to examine, I have, however, found a few specimens more perfect than those previously described, and also a few new species. When first studying the excellent figures given by Weller, one is struck by the apparent triviality of the specific characteristics employed in the discrimination of the species, but with a large collection, it is found that the characteristics are remarkably constant. The study of these illaenids is unusually interesting, in fact, for it seems to be 16 bulletin: museum of comparative zoology. one of the few cases where characteristics arbitrarily chosen may serve to define natural groups. The bumastids particularly, which are simple trilobites with few variable characters, seem to be susceptible to this sort of treatment. The chief variable characters are amount of convexity, ratio of length to breadth, length of dorsal furrows, pres- ence or absence of "lip" on cephalon and concave border on pygidium, and size of eyes. Almost every possible combination of these few characteristics seems to be present among the species, and each com- bination is usually exhibited by a large number of specimens. That these variations are not the attributes of one very plastic species but of a number of distinct species is shown by the geographical distribu- tion as well as by the numbers in each group. -For example, Bumastus insignis seems to be confined to a small area in the immediate vicinit;\' of Chicago, and B. ioxus to Joliet, Illinois and Racine, Wisconsin, while B. cunimdus is found in vast numbers near Milwaukee, but is rare in the Chicago area. Others of the species are equally local in distribution. Bumastus cuniculus (Hall). Illaenus cuniculus Hall, 20th Rept. N. Y. state cab. nat. hist., 1868, p. 377, pi. 22, f. 12; 1870, rev. ed., p. 421, pi. 22, f. 12. Weller, Bull. Chicago acad. nat. hist., 1907, no. 4, pt. 2, p. 219, pi. 19, f. 1-6. This is by far the most common species at Wauwatosa, Wise. There are several nearly complete specimens in the M. C. Z., most of them enrolled. They show the axial lobe of the thorax to be extremely wide, and the dorsal furrows very shallow. There are ten segments. The diagnostic specific characteristics are: — elongate, moderately con- vex cephalon with very narrow rim which is prominent at the front, but disappears before reaching the genal angles, and eyes of medium size. Dorsal furrows faint, hardly visible at all in front of the scar- like spots just inside the eyes. Pygidium elongate, moderately convex, with practically no depressed border. A flattening of the convexity near the border can be seen if the pygidium is viewed in profile. Measurements: — A cephalon of average size is 45 mm. long, 55 mm. broad; the eye 9 mm. long, or one fifth the total length. A large cephalon is 56 mm. long. The pygidium of an enrolled specimen whose cephalon is 45 mm. long is 50 mm. long and 52 mm. wide. The axial lobe of this specimen is 42 mm. wide and the total width of the thorax is 53 mm. Formation and locality: — Very common in the Niagaran at Wauwa- tosa, near Milwaukee, Wisconsin. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 17 BuMASTUS NiAGARENSis (Whitfield). Plate 1, fig. 3. Illaenus niagarensis Whitfield, Ann. rept. Geol. surv. Wisconsin for 1879, 1880, p. 68. Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 219, pi. 19, f. 7-11. Illaenus madisonianus Whitfield. Geol. Wise, 1882, 4, p. 307, pi. 20, figs. 8-9. Foerste, Bull. sci. lab. Den univ., 1885, 1, p. 106 pi. 14, f. la-b, 2a-b; 1887, 2, p. 93, pi. 8, f. 8, 9, 10, 10a; Geol. surv. Ohio, 1893, 7, p. 526, pi. 26, f. 1, 2, varieties, pi. 27, f. 7-10. Van Ingen, School of mines quar- / terly, Columbia univ., 1901, 23, p. 35. This seems to be one of the few illaenids of the Chicago-Wisconsin Niagaran area which has hitherto been known from entire specimens. Whitfield figured an entire one, Weller had a nearly complete speci- men, and there are five in the M. C. Z. (Day collection). One of these is figured as it shows some characters not shown in either Whit- field's or Weller's figure. Whitfield's figure, if it really represents this species, is inaccurate in respect to the glabellar furrows, which are really much longer than is indicated by his figure or description. From the general proportions of the body, and the position of the eyes, it would seem that his figure really does represent this species. The specimen here figured is a little longer and narrower than those previously figured, and has a longer and more pointed pygidium. It will be noted that the thoi'ax is exceedingly short, though ten segments are present. The dorsal furrows do not show in the figured specimen, but they do on another specimen in the M. C. Z. collection, and are also indicated in Whitfield's figure. The specific characteristics are : — cephalon rather convex, with long dorsal furrows, no lip or concave border on the cephalon. Eyes of medium size, situated nearly their own length from the posterior margin, thorax short, pygidium long, rather pointed behind, with narrow concave border. Measurements: — The specimen (Plate 1, fig. 3) is 58 mm. long; cephalon 22 mm. long, 31 mm. wide; thorax 13.5 mm. long; pygidium 29 mm. long, 29 mm. wide; the eyes are damaged. On a very good enrolled specimen preserving the test, the cephalon is 24 mm. long, the eye 5 mm. long, and 5 mm. from the posterior margin. On a larger specimen the eye is 1 mm. more than its own length from the posterior margin. 18 bulletin: museum of comparative zoology. Formation and locality: — This species is quite common in the Niag- aran at Wauwatosa, though not nearly so common as B. cunicuhis. BUMASTUS DAYI, Sp. nOV. Plate 1, fig. 8-10. The Day collection contained two fine specimens labeled sp. nov., and as they have proven to be such I have named them in honor of the collector. The specific characteristics are: — cephalon and pygi- dium short and convex, eyes large and far back, no lip or concavity at the front of the glabella, though there is one at the sides of the free cheeks, dorsal furrows of the cephalon long, reaching the pits in front of the eyes. Thorax of ten segments, axial lobe very wide, furrows shallow. Pygidium short and evenly convex, with only a trace of a concave margin. It will be at once noted that this species is much like B. niagarensis, but has large eyes far back, and the pygidium is shorter and with less depressed margin. It differs from B. cuniculus in having long in- stead of short dorsal furrows on the cephalon, and in lacking the rim at the front. Of the species found near Chicago, B. chicagoensis (Weller) is very similar to this but that species has a much shorter and more abruptly deflected cephalon. Measurements: — The type is 46 mm. long; cephalon 19 mm. long, 26 mm. wide; eye 7 mm. long and 2 mm. from posterior margin; thorax 14 mm. in length; pygidium 23 mm. long, 25 mm. wide. A large cephalon is 34 mm. long, 46 mm. wide; eye 11 mm. long, 4 mm. from posterior margin. Formation and locality: — Quite common in the Niagaran at Wauwa- tosa, -Wise. BuMASTUS DECIPIENS, Sp. nOV. Plate 1, fig. 1, 2. Exceedingly like B. dayi is a species of which the collection contains only five or six specimens, two of which are approximately entire. The specimens are of about the same size as the smaller ones of B. dayi, and the principal difference between the two is that B. decipiens RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 19 has short glabellar furrows and no lip on the free cheek, while B. dayi has long furrows and a lip on the free cheek. The eye of B. decijnens seems also to be a little longer and further back than in B. dayi. The pits in front of the eyes are exceedingly faint. Buviastus transversalis (Weller) likewise has a short head, large eyes, and short dorsal furrows, but is much wider than B. decipiens. B. armatus (Hall) is much like B. decipiens but has spines at the genal angles. Measurements: — The holotype (Plate 1, fig. 2) an imperfect cranidium, is 19 mm. long; the eye is 7.5 mm. long, and 2 mm. from the posterior margin. The figured paratype (Plate 1, fig. 1) is about 51 mm. long; cephalon 21 mm. long; thorax 15 mm. long. A second small, unfigured paratype has the cephalon 12 mm. long, 16 mm. wide; eye 5 mm. long; 1 mm. from posterior margin; thorax 8 mm. long; pygidium 12 mm. long, and about 16 mm. wide. Formation and locality: — All the specimens are from the Niagaran at Wauwatosa, Wise. BUMASTUS TENUIS, Sp. nOV. Plate 1, fig. 6, 7 and ? 11. This species is like B. decipiens, rare, and the collection contains five specimens showing the cranidium only. It is characterized by its slight convexity and the very wide concave lip at the front. The dorsal furrows are long, reaching to the pit in front of the eyes. These pits are themselves very far forward and close to the margin. The eyes are large and close to the posterior margin. This species of course suggests B. cuniculus, but the lip is much wider, the dorsal furrows longer, and the pits in which they end very close to the margin. It has a flatter cephalon and wider lip than any known Bumastus. A pygidium (Plate 1, fig. 11) which does not seem to belong with any other species found at Wauwatosa is tentatively referred to B. tenuis. It is too short and not convex enough for B. niagarensis, and has too wide a concave border for B. cuniculus, B. dayi, or B. decipiens. Measurements: — The type cranidium is 33 mm. long and 34 mm. wide at the palpebral lobes. The eye was approximately 8 mm. long and 2 mm. from the posterior margin. Formation and locality: — All the specimens are from the Niagaran at Wauwatosa, Wise. 20 bulletin: museum of comparative zoology. BUMASTUS INSIGNIS (Hall). Illaenus insignis Hall, 18th Rept. N. Y. state cab. nat. hist., 1865, p. 27, f. 5, 6 adv. sheets; 20th Rept. N. Y. state cab. nat. hist., 1868, p. 331, f. 5, 6, pi. 22, f. 13, 14; 1870, rev. ed., p. 419; f. 10. 11, pi. 22, f. 13, 14. WeUer, Bull. Chicago acad. nat. sci., 1907, no. 4, pt. 2, p. 215, pi. 17, f. 1-5. Not of Salter, Whitfield, Foerste, and Kindle. (See WeUer, op. cit.). It is interesting to note that though Hall cites Waukesha and Milwaukee, Wise, first in his list of localities for this species, the species is not known at all from Wisconsin, and the specimens which Hall figured were undoubtedly from near Chicago. Whitfield's Illaenus insignis was B. cuniculus (Hall). The English specimens figured by Salter as belonging to this species, show the same long furrows and cephalon with a lip, but the front of the head is not so pointed. The eyes of the English form are likewise less- elongate and further back than in specimens from near Chicago. It seems probable that the English form deserves a distinct name, and B. pomiata, the name used by Salter on the plate of his publication, is still available. BuMASTUs loxus Hall. Illaenus (Bumastus) barriensis ? Hall, Geol. surv. Wise, 1862, 1, p. 433, (no description); 18th Rept. N. Y. state cab. nat. hist., 1865, p. 28 adv. sheets; 20th Rept. N. Y. state cab. nat. hist., 1868, p. 332. Illaenus ioxus Hall^ 20th Rept. N. Y. state cab. nat. hist., 1868, p. 387, fig., pi. 22, f. 4-11, pi. 23, f. 1. Whitfield, Geol. Wise, 1882, 4, p. 304, pi. 21, f. 11, 12. Foerste, Proc. Boston soc. nat. hist., 1890, 24, p. 268, pi. 5, f. 20. Van Ingen, School of mines quarterly, Columbia univ., 23, 1901, p. 35 (no description). Kindle, 28th Ann. rept. Dept. geol. and' nat. res. Indiana, 1904, p. 480.. pi. 22, f. 7, pi. 23, f. 3. Weller, BuU. Chicago acad. sci., 1907, no. 4, pt. 2, p 222, pi. 18, f. 1-3. Illaenus (Bumastus) ioxus Hall, 20th Rept. N. Y. state cab. nat. hist., 1870, rev. ed., p. 420, f. 12, pi. 22, f. 4-10. ? 11th Rept. Dept. geol. and nat. hist. Indiana, 1882, p. 335, pi. 38, f. 14, non 13; Trans. Albany inst., 1883, 10, p. 76. The status of the name of this species is somewhat like that of Cheirurus niagarensis. In the Pal. N. Y., 2, Hall identified the rather common large Bumastus of the Rochester shale as B. barriensis Murchison, and in his earlier references to the Wisconsin specimen he RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 21 used the same name. When he came to figure a Wisconsin specimen, however, he proposed a new specific name for it, and the name was so obviously suggested by the Wisconsin specimens that I agree with Weller that those specimens should be considered the types of B. ioxus. Professor Weller has suggested that the specimens from New York may belong to another species. I have investigated the point as fully as the material at my command would permit, and have not so far been able to find any really good characteristics on which to base a separation. The best specimens from New York are usually small, and considerably flattened. The study of better material will prob- ably reveal characteristics not now evident, and I have therefore omitted from the synonymy the references to the New York specimens. Hall figured a pygidium which he assigned to the species, undoubt- edly correctly, but up to the present the thorax has been unknown. The M. C. Z. (Day collection), contains a large specimen, whose label states that it is the "only perfect specimen found at Racine." It is not exactly a perfect specimen, though it retains cephalon, parts of ten thoracic segments and the pygidium. The axial lobe of the thorax ,is somewhat less wide than one would have expected from the large size of the animal, but, being 62% of the total width, is about the general average among the bumastids. The pygidium does not show an actual concave border, but there is a very decided flattening of the curve of the profile at the back. As the specimen is preserved, the pygidium is somewhat unnaturally drawn in, so that the actual length is not shown. On the other hand, the last thoracic segment is displaced from the others and there is a considerable space between the thorax and cephalon at the anterior end, and between the thorax and pygidium behind. The length of this specimen, therefore, gives only a rough approximation of the correct length. Incidentally it should be noted that the cephalon of this species has a large median tubercle near the posterior margin. It is shown in Hall's figure, but omitted from Weller's. Measurements: — Length, about 180 mm.; cephalon 75 mm. long, 110 mm. wide; thorax about 75 mm. long, about 100 mm. wide at middle, axial lobe 62 mm. wide; pygidium 65 mm. long, 102 mm. wide. A well-preserved pygidium is 68 mm. long and 100 mm. wide. Formntion and locality: — Hall mentions Waukesha and Wauwa- tosa as localities for this species, but in very extensive collections from these places no specimens of this species are present, while we have a number of specimens from the Racine dolomite at Racine, Wise. 22 bulletin: museum of comparative zoology. BuMASTUS graftonensis Meek and Worthen. Illaenus (Bumastus) graftonensis Meek and Worthen, Proc. Acad. nat. sci. PhU., 1870, p. 54; Geol. surv. Illinois, 1875, 6, p. 508, pi. 25, f. 4. Buniastus sp. ind. Meek and Worthen, Geol. surv. Illinois, 1875, 6, pi. 24. f. 3. Illaenus graftonensis Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 223, pi. 16, f. 4-6. Only the cephalon of this species has previously been known, but the Day collection contains two complete, though somewhat flattened specimens from Waukesha, Wisconsin, where this species seems to be fairly common. The cephalon is too well known to need further description, except to note that as in B. ioxus, there is a prominent median tubercle be- tween the eyes and near the posterior margin. The thorax has ten segments, a broad axial lobe, rather well-defined dorsal furrows. The pleura of the thoracic segments are more promi- nent and ridged, and not so flat as in most species of Bumastus. The pygidium is short and moderately convex, with a flattening around the margin, but not a real concave border. The pygidium resembles that of B. ioxus, but is shorter and wider, the average ratio of length to width in B. ioxus being .64 and in the two pygidia of B. graftonensis which we have, .54. Formation and locality: — Nine specimens (M. C. Z. coll.) are from the Niagaran at Waukesha, Wise, a locality from which this species has not previously been reported. Bumastus indeterminatus (Walcott). Plate 2. Illaenus indeterminatus Walcott, 31st Ann. rept. N. Y. state mus. nat. hist., 1877, p. 19 adv. sheets; 31st Ann. rept. N. Y. state mus. nat. hist., 1879, p. 70. Illaenus cf. /. indeterminatus Clarke, Pal. Minn., 1897, 3, pt. 2, p. 716, f. 24. Bumastus indeterminatus Raymond and Narraway, Ann. Carnegie mus., 1908, 4, p 253, pi. 62, f. 8, 9. The M. C. Z. contains the type of this species, and it is figured (Plate 2) for the first time. The specimen consists of a good cephalon, part of one free cheek, a very much dislocated thorax, of which only se\en segments can be definitely made out, and a very fine pygidium. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 23 The pygidium is considerably longer and more convex than the cepha- lon, and the axial lobe is evidently very wide and without very strong dorsal furrows. The dorsal furrows of the cephalon fade out just before reaching the pits in front of the eyes, but this is probably an individual variation. These pits are not directly in front of the eyes as in most species of Bumastus, but considerably inside the projection of a line drawn through the length of the eye. The pygidium is very convex, with concave slopes to the lateral and posterior margins. Formation and locality: — The type, M. C. Z. no. 650, is from Russia (Newport), Herkimer Co., New York. The horizon is the Leray- Black River (Ordovician). ACTINOLOBUS AMERICANUS, Sp. UOV. Plate 1, fig. 4, 5. The M. C. Z. contains a single pygidium which, on account of its great length and wide flat border, is referred to Actinolobus, a genus previously unrepresented in this country. The specimen is damaged on the right side and has been restored in plaster. There is also a break on the left side not shown in the figure, but the outline is believed to be accurate. The specimen is 38 mm. long and 20 mm. wide at the front. At the widest part, the border has a width of 10 mm. and is somewhat concave. The central part of the pygidium is strongly convex. The anterior margin indicates that the axial lobe of the thorax was wide, though not so wide as in the species of Bumastus found with it. Formation and locality: — From the Racine dolomite at Racine, Wisconsin. LICHADIDAE Hawle and Corda. Trochurus nasutus (Weller). Plate 3, fig. 1, 2. Dicranopeltis nasuta Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 240, pi. 22, f. 5-7. A figure of the specimen of this species in the Day collection is introduced in order to show the wav in which the frontal lobe of the 24 bulletin: museum of comparative zoology. glabella tapers into a spine, this feature not being correctly shown by Weller's figures. I had at first believed that this specimen repre- sented a distinct species, but fortunately was able to see the type in Mr. Teller's collection. That the species belongs to Trochurus in- stead of Dicranopeltis is shown by the dorsal furrows, which curve inward instead of outward, at the posterior ends. Formation and locality: — The specimen is from the Niagaran at Wauwatosa, Wisconsin. ODONTOPLEURIDAE Burmeister. Ceratocephala goniata Warder. Plate 3, fig. 3-5. Ceratocephala goniata Warder, Amer. journ. sci., 1838, ser. 1, 34, p. 378, fig. Clarke, 44th Rept. N. Y. state mus. nat. hist., 1892, p. 91-100, pi. 1, f. 1. Kindle, 2Sth Ann. rept Dept. geol. and nat. res. Indiana, 1904, p. 480, pi. 24, f. 13. Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 255, pi. 23, f. 1-2. Raymond, Bull. Victoria mem. mus., 1913, 1, p. 38. Acidaspis danai HaU, Geol. surv. Wise, 1862, 1, p. 432 (no description); 18th. Rept. N. Y. state cab. nat. hist., 1865, p. 28, adv. sheets; 20th Rept. N. Y., state cab. nat. hist., 1868, p. 333, pi. 21, f. 8-9; 1870, rev. ed., p. 423, pi. 21, f. 8, 9. Acidaspis ida WincheU and Marcy, Mem. Boston soc. nat. hist., 1865, 1, p. 106, pi. 3, f. 13. The cephalon of this species is, thanks to Professor Weller, now well known, but hitherto the thorax and pygidium have not been noticed. The Day collection in the M. C. Z. contains parts of two pygidia and a fragment showing a portion of five segments of the thorax. The pygidium is like that of Acidaspis portlocki Barrande, with a large median spine, on each side of which are two smaller spines, then a large spine, and finally an outer small spine. Each spine gives off small thorns on each side. The thorax is similar to that of C. verneuili Barrande. However, as only the central portion has been seen this part of the body would not be expected to show specific characteristics. The cephalon seems to be more nearly allied to C. vesiculosa than to C. verneuili, and a cephalon in the M. C. Z. gives indication that marginal spines were present on the free cheeks. The pygidium found with the cranidia at Wauwatosa are unlike those ascribed to C. vesiculosa by RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 25 Barrande, and, on the other hand, thou.^h the pygidia in the Day collection are much like that of A. portlocki, their cephalons are very unlike those assigned to A. portlocki by Barrande. Formation and locality: — The specimens figured are from the Niagaran at W,ajiwatosa, Wise. ENCRINURIDAE Angelin. Encrinurus reflexus, sp. nov. Plate 3, fig. 7, 8. The Day collection contains several pygidia of an Encrinurus which is larger and differs in various ways from any species of this genus heretofore described from Silurian strata of this country. Cephalon and thorax unknown. Pygidium large for the genus, triangular, pleura bent abruptly downward, the posterior end somewhat turned up. Axial lobe very long, tapering backward to a point. It is marked by about thirty rings which are prominent and sharp toward the front, but very faint at the posterior end. With the exception of two at the anterior end the rings do not cross the axial lobe, but leave a narrow smooth space along the median line. Along this smooth lane are disposed eight pustules, approximately evenly spaced. On the pleural lobes are eight pairs of broad flat ribs which curve backward, making a rather abrupt turn near their outer ends. They do not reach quite to the margin, and end in blunt free spines which project from the sides a little above the margin. At the posterior end the last two ribs from each side converge alongside the axial lobe, and, with a small median spine, project beyond the end of the axial lobe. The ribs have small pustules scattered somewhat irregularly over them, and not aligned in longitudinal rows. Nearly every rib has a pustule at its inner end •and one near the middle. Some of the ribs have only these two, but the longer ones near the front have another. Measurements: — The more complete of the cotypes is 28 mm. long, 26 mm. broad at the front. The axial lobe is 9 mm. wide at the front and 26 mm. long. Comparison with other species : — Nine species of Encrinurus have previously been described from the Silurian of America, but most of them are of much smaller size than the present species and only two 26 bulletin: museum of comparative zoology. species, E. deltoideus Shumard and E. nereus Hall, have as many as eight pairs of ribs. Most of the species have seven pairs, one, E. americanus Vogdes, has six, and E. elegantulus Billings has only five. The species may be taken up in alphabetical order. E. americanus Vogdes (Description new Crustacea from Clinton of Georgia, 18G6, p. 1), has only six pairs of ribs on the pleura, and no pustules, thus ruling it out at once. E. deltoideus Shumard (Geol. Missouri, 1855, p. 198, pi. B, f. 10), is similar in many ways to E. reflexus, having the rings very numerous, twenty-four in nvimber, and interrupted by a smooth lane. There are, however, no pustules except very indistinct granules, on this lane, and the ribs, of which there are eight pairs, are likewise smooth. E. egani S. A. Miller (Journ. Cine. soc. nat. hist., 1880, 2, p. 254, pi. 15, f. 1, lb), has a long terminal spine, only seven pairs of ribs, which are narrow with wide interspaces, and while the axial lobe of the pygidium is on the same plan as in E. reflexus, it has fewer rings and tubercles. E. elegajitulus Billings (Cat. Silurian fossils Anticosti, 1866, p. 62), has only five pairs of ribs, the median lane on the axial lobe is without tubercles, but the first eight rings cross it. E. indianaensis Kindle (28th Ann. rept. Dept. geol. and nat. res., Indiana, 1904, p. 482, pi. 24, f. 14, 15), is a very peculiar species, not at all of the same type as the one under discussion. It has fifteen rings on the axial lobe and ten pairs of ribs. Each rib crosses the axial lobe and has three to five tubercles. E. nereus Hall (20th Rept. N. Y. state cab. nat. hist., 1868, p. 375, pi. 21, f. 15) has no flattened lane along the top of the axial lobe, and no nodes on the rings or ribs. E. ornatus Hall and Whitfield (Pal. Ohio, 1875, 2, p. 154, pi. 6, f . 16) is c^uite similar to the present species, but has only seven pairs of ribs, and the ribs themselves are narrower and the spaces between them wider. There are also only twenty rings and five nodes on the axial lobe. E. thresheri Foerste (Bull. Sci. lab. Denison univ. 1887, 2, p. 101, pi. 8, f. 26) is a small species which is similar to the last and to E. reflexus. There are, however, only seven pairs of ribs, which are themselves exceedingly narrow; there are also only eighteen rings on the axial lobe, and six pustules on the smooth lane. E. tuber culifrons Weller (Bull. Chicago acad. sci.. 1907, no. 4, pt. 2, p. 259, pi. 24, f. 12, 13) is a small form with a short wide pygidium which is without nodes and the rings of which cross the axial lobe without interruption. Formation and locality: — From the Niagaran at Wauwatosa, Wise. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 27 CALYMENIDAE Milne Edwards. Calymene niagarensis Hall. Calymene niagarensis Hall, Geol. N. Y., 1843, pt. 4, p. 102, f. 3 on p. 101; tab. org. rem. 10, f. 3. Calymene hlumenbachi var. niagarensis Hall, Pal. N. Y., 1852, pt. 2, p. 307, pi. 67, f. 11, 12. Calymenes are difficult fossils to differentiate satisfactorily, but it is possible to draw a little closer limits to some of the species than has been done in the past. The Silurian species do not present so difficult a problem as do those in the Ordovician, specific characteristics being apparently more fixed and constant in later times. Before venturing to separate two new species, it is best to direct attention for a moment to the well-known (in name) Silurian form. The name was applied originally by Hall to specimens from the Rochester shale at Lockport, N. Y. The figures and description show the original specimens to have been of the Calymene hlumenbachi type, that is with a narrow lip in front of the glabella, three pairs of glabellar lobes, anrl pygidium with an impressed line on each rib, distinctly bifurcating the outer portion. The ribs also reach practically to the margin. It seems that the species Calymene niagarensis should be restricted to such trilobites as show these important characteristics of the types and these may be seen in most of the Calymenes in the Rochester shale. Another Calymene found at the same horizon, C. vogdesi Foerste, has the same bifurcated ribs on the pygidium, but a much longer snout-like lip in front of the glabella. It is also a much larger form, one of the largest of the Calymenes. In the Ordovician the common Calymene senaria of the Trenton has the same type of bifurcated rib, while the later C. meeki Foerste, so abundant in the Eden and IMaysville at Cincinnati, shows only a trace of an impressed line on the ribs, and often the line is absent entirel\'. Calymene breviceps, sp. nov. Plate 3, fig. 11. ■Calymene niagarensis Hall, 28th Rept. N. Y. state mus. nat. hist., doc. ed., 1877, pi. 32, f. 8-15; mus. ed., 1879, pi. 32, f. 8-15; 11th Rept. Dept. geol. and nat. hist. Indiana, 1882, p. 331, pi. 34, f. 8-15. Hall and Clarke, Pal. N. Y., 1888, 7, pi. 1, f. 10-14. 28 bulletin: museum of comparative zoology. The Calymene abundant at Waldron, Indiana, has always been identified with C. niagarensis, but differs from that species in at least two marked details. The first and most obvious characteristic is that there is no lip, nor any furrow between the glabella and the rim, so that the glabella reaches upon, and in some cases, overhangs the rim, a feature usual in the Cheiruridae but extremely uncommon among the Calymenidae. This gives the cephalon the high, short appearance which suggested the name breviceps. On the pygidium the ribs reach nearly to the margin but become faint on approaching it. Ordinarily the ribs do not bear any median impressed line though traces of one may be seen on some specimens. This species is in many ways much like C. celebra, the next species described. Formation and locality: — This species is so far known only from the (Silurian) Waldron shale at Waldron, Indiana, where it is very com- mon. Calymene celebra, sp. nov. Plate 3, fig. 9, 10. Calymene hlumenhachii var. niagarensis Hall, Geol. surv. Wise, 1862, 1, p. 432. Calymene niagarensis Hall, 18th Rept. N. Y. state cab. nat. hist., 1865, p. 30, adv. sheets; 20th Rept. N. Y. state cab. nat. hist., 1868, p. 334; 1870, rev. ed., p. 425 Weller, Bull. Chicago acad. sci., no. 4, pt. 2, p. 261, pi. 23, f. 9-10. One of the most abundant of the trilobites of the Chicago area and of southeastern Wisconsin is a Calymene which is constantly identified as C. niagarensis. It is quite commonly found entire, but always so far as I have seen in the condition of a cast of the interior. Moulds of the exterior are common, but seldom complete. The cephalon is like that of C. niagarensis, with a short lip and narrow furrow in front of the glabella. The dorsal furrows are always very deep and sharp, but this is due to the state of preservation. The glabella tapers rather abruptly toward the front. The basal lobes are large, rounded, almost isolated; the second lobes small and rounded, the intermediate "extra lobes" not very prominent. The third lobes are very small and the fourth ones just barely indicated. The frontal lobe is short and rather square at the front. The eyes are close to the glabella and opposite the furrows between the second and third pairs of lobes. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 29 The pygidium is the most characteristic portion of the animal. The axial lobe is narrow, well defined, and has rings. The pleural lobes show four pairs of narrow ribs, without impressed line, which reach only halfway to the margin. The fourth of the four pairs are very faint and short. Each pleural lobe is thus divided into a small triangular ribbed portion near the axial lobe and a much longer smooth portion below. This pygidium presents the greatest possible contrast to C. niagarensis, in which the ribs are more conspicuous near the margin than near the axial lobe. The peculiarities of the pygidium have doubtless been noticed before, and probably have been explained as due to the state of preservation, the specimens all being internal casts. Internal casts of either cephala or pygidia of trilobites are practically always less and not more smooth than the exteriors, how- ever, and cleaned interiors of C. senaria, C. breviceps, and C. meeJci, all show that Calymene follows the general rule. Calymene celebra shows a halfway stage to what is achieved in C. clintoni Vanuxem, namely, a pygidium with smooth pleural lobes. The latter species is too far removed from the Calymenes with typical ribbed pygidia to be in- cluded in the same genus. Formation and locality: — - Calymene celebra is common in the Niagaran of the Chicago district in northern Illinois, in the same portion of the Silurian in southeastern Wisconsin, and also near Madison, Indiana, and Eaton, Ohio. LiocALYMENE, gen. nov. Calymeninae (as distinguished from the Homalonotinae) with distinct glabella, three pairs of glabellar lobes, narrow thorax, pygidium with ringed axial and smooth pleural lobes. Type, Hemicrypturus clintoni Vanuxem. Liocalymene cJintoni, in perfect preservation, appears to be an exceedingly rare fossil. The single specimen in the M. C. Z. is in about the same condition as that figured by Hall (Pal. N. Y., 2, p. 298, pi. A 66, f. 5a), and is from the Clinton shale at Clinton, Herkimer Co., N. Y. 30 bulletin: museum of comparative zoology. CHEIRURIDAE Salter. Cheirurinae Raymond. Cheirurus niagarensis (Hall). Plate 4, fig. 4, 5, 6, 9. Ceraurus insignis Hall, Pal. N. Y., 1852, 2, p. 303, pi. 67, f. 9, 10. Ceraurus niagarensis Hall, 20th Rept. N. Y. state cab.' nat. hist., 1868, p. 376, ? Whiteaves, Geol. surv. Canada. Pal. foss., 1884, 3, pt. 1, p. 42; 1895, 3, pt. 2, p. 107. ? Van Ingen, School of mines quarterly, Columbia univ , ■ 1901, 23, p. 35 (no description). Kindle, 28th Ann. rept. Dept. geol. and nat. res. Indiana, p. 483, pi. 23, f. 1, 2, pi. 24, f. 8. Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 263, pi. 24, f. 20, non 21. The name Ceraurus niagarensis appears for the first time on p. 376 of the first edition (1868) of the 20th Ann. Rept. of the New York State Cabinet of Natural History. Earlier in the same paper, (p. 335), Hall referred certain trilobites from the Silurian at Wauwatosa and other localities in Wisconsin to Ceraurus insignis (Beyrich). On p. 376 he states that he has reexamined the specimens and considers them different from C. insignis. The name Ceraurus niagarensis is used as a heading, but is not designated either as a new name or a new species. On p. 427 of the Revised edition, published in 1870, the remarks are reprinted, but the letters N.S. follow the name. In neither case is there any description of the species given, but the plates contain representations of an imperfect cranidium and a broken hypostoma. The next use of the name by Hall was in 1879, in the 28th Rept. of the N. Y. State Museum, p. 189. He here describes the pygidium at some length from specimens obtained at Waldron, Indiana, and remarks, at the end of his description: "From the above it will be seen that the separation first made in the revised edition of the 20th Rept. St. Cab., was necessary, and that it constitutes a distinct spe- cies." The single figure given represents a pygidium. After studying collections from a number of localities it becomes evident that the American forms now referred to Ceraurus niagarensis include two or three species, and it is therefore, necessary to determine the type for C. niagarensis. From the absence of description accom- RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 31 panying the first use of the name niagarensis it would appear that Hall did not apply the name to the Wisconsin specimens alone, but meant to assign this name to all American forms previously ascribed to Cheirurus insignis or Ch. hivincronatus . This idea is strengthened by the fact that he states that the New York and Wisconsin specimens show the same characteristics. He also refers to Roemer's Ceraurus bimucronatus from Tennessee. This idea is still further strengthened by the remark quoted above from his description of the Waldron fauna. If this is the case, then we should return to Hall's first descrip- tion of a Ceraurus insigiiis in America to get at his idea of the species. If we take the other view, that the specimens from Wauwatosa, which seem to have been the first ones to cause Hall to doubt the correctness of his reference of all the Silurian cheirurids to the Bohemian species, are the real types of C niagarensis, we are confronted by the fact that he did not describe his specimens, and, moreover, he was evidently in doubt about them, as evidenced by his pleasure at finding distinguish- ing features in the pygidia from Waldron. This latter description was the first real description published after the name niagarensis was proposed, and it might well be argued that the last described of the group should be the type. It seems simpler, however, to accept what was Hall's evident intent, and believe that in proposing the name Jiiagarensis he was merely proposing a new name for the specimens he had previously described as Ceraurus insignis. The first description of Ceraurus insignis Beyrich, by Hall occurs in vol. 2 of the New York State Paleontology, 1852. On page 300 there is mentioned, without description, a glabella from the Clinton which is figured on plate 66A. On page 306 of the same volume Hall describes two cranidia from the Rochester shale at Rochester, N. Y. Both specimens are figured. As these are the first American speci- mens which are both figured and described, I propose to designate as the type of Cheirurus niagarensis (Hall) the one represented in fig. 10, pi. 67, of the above volume. This specimen is in the American Museum Natural History, No. 1827. The specimen so designated is a typical Cheirurus, with the glabella expanding rather rapidly forward, the frontal lobe occupying less than half the length of the glabella, and the first two pairs of glabellar furrows nearly straight, and following a direction approximately parallel to the posterior margin of the cephalon. Their inner ends are separated by a smooth space equal in width to about half the glabella. The eyes are near the dorsal furrows, and about opposite the second glabellar furrows. 32 bulletin: museum of comparative zoology. This type of cranidium is quite often seen in collections from the Rochester shales, but one also sees another type, the one which Hall figured from the Clinton. The cranidium is similar to the one just described, but the glabellar furrows, instead of being short and straight, are long, curve backward, and their inner ends almost meet. This type of head deserves to be recognized as distinct from the other; it is the type of head figured by Hall from Wisconsin and though both types are very common there, this is by far the more abundant. To this same type, though possibly not to the same species, belongs the cephalon of Ceraurus himucronatus figured by Roemer. No entire specimen of Cheirurus has, so far as I know, been found in America, and it is therefore difficult to decide what pygidium shall be associated with each type of cephalon. It would appear that no Cheirurus pygidium had been figured from New York. The M. C. Z. possesses a single small pygidium of a Cheirurus from the Rochester shale at Rochester, N. Y. It is of the familiar Cheirurus insigiiis type, with three pairs of long slender spines, and a short median spine. It is very dift'erent from the pygidium from the Waldron shale ascribed to Ceraurus niagarensis by Hall, for that specimen was described as having broad flat spines, each spine with a depressed line on the sur- face. Pygidia found at Wauwatosa are like the one from Rochester and it seems probable that this type of pygidium is to be referred to Cheirurus niagarensis. The following description of Cheirurus niagarensis is based on three glabellas (M. C. Z. €25) and a pygidium (M. C. Z. §24) from the Rochester shale at Rochester, N. Y., and a cranidium with three segments attached and an associated hypostoma, from Wauwatosa, Wise. (M. C. Z. 626). A large cranidium with a part of the thorax (M. C. Z. 627) from Wauwatosa was also consulted. Cheirurus niagarensis (Hall) restricted. A Cheirurus of medium size. Cranidium semicircular in outline, gently convex, the glabella forming the highest and most prominent part, but not standing much above the cheeks. The glabella reaches the front of the cranidium, expands toward the front, and is widest at the middle of the frontal lobe. Dorsal furrows narrow and sharp, but not very deep. Glabellar furrows short, sharp, the first two pairs extending only a short distance onto the glabella. Their direction is RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 33 approximately vertical to the axis of the animal, but they are usually not absolutely straight, but bend a little forward in mid length. The posterior furrows run diagonally inward and connect with the neck furrow, as in the genus generally. The eyes are close to the glabella and opposite the second pair of glabellar furrows. Free cheeks small, pitted, with a smooth, convex rim. Fixed cheeks pitted. Glabella granulose. The associated hypostonia is roughly tetragonal, the surface with sharp, scattered pustules, and the posterior margin nearly straight, without spines at the angles. The furrow around the body portion is wide and deep. Of the thorax only three segments are known. The axial lobe is narrow, the inner part of the pleural lobe is crossed by a narrow diagonal furrow which separates two triangular nodes, and there is a prominent node at the fulcrum. Beyond the fulcrum the pleuron projects as a blade-like spine. The pygidium is short, with three pairs of slender spines which are oval in section and unfurrowed. The last pair extend further back than the ones ahead of them. A median spine is present, but very short. The axial lobe is narrow, cone-shaped, with the point backward, bearing three rings and a node. The pleural lobes are narrow, and .show a single short divided rib on either side at the anterior end. This species is very much like Cheirurus insignis Beyrich. The glabella seems to be a little shorter and wider in the American form, and the Bohemian species has the eyes further from the glabella and has eyelines. Of the latter, however, the specimens in the M. C. Z. show a trace. The hypostoma of the Bohemian form is similar to that of the American species, but the posterior margin is somewhat wider and more flattened. The pygidia are practically the same, though the median spine is a little stronger in Ch. insignis. It will be seen from the above description, that if we restrict Cheiru- rus niagarensis to those forms which Hall first identified with Cheirurus insignis, we eliminate both the forms which caused him to change his mind about the identification, and propose the new name niagarensis. This would seem to vitiate the argument above, but it must be re- membered that Hall did not recognize that he was dealing with more than one species, and he did not apply the new name to any definite specimens. In fact, it would seem that he did not become fully con- vinced that a new name was needed till he studied the pygidium from Waldron, and if the name niagarensis is not to be interpreted as has been done here, it would be almost impossible to decide whether the Wisconsin or the Waldron specimens should be selected as the types. 34 bulletin: museum of comparative zoology. In spite of the similarity of Cheirurus niagarensis, as above defined, to Cheirurus insignis, I believe that a separate specific name should be maintained, especially as the discovery of further material may show unsuspected differences between the two. Measurements: — A cranidium from Rochester (M. C. Z. 625). Length 20 mm., width 35 mm. ; width of glabella at neck-ring 12 mm., at front 16 mm.; length of frontal lobe 10 mm. A cranidium from Wauwatosa, (M. C. Z. 626). Length 9 mm., width 18 mm.; width glabella at neck-ring 5 mm., width at front 8 mm. ; length of frontal lobe, 3 mm. Formation and locality: — Rochester shale at Rochester and Lock- port, N. Y., Niagaran in Indiana, Wisconsin and Illinois. Cheirurus welleri, sp. nov. Plate 3, fig. 6; Plate 4, fig. 7, 8, 10. Ceraurus insignis Hall, Pal. N. Y., 1852, pt. 2, p. 300, pi. 66A, f. 4. Geol. surv. Wise, 1862, 1, p. 433; 18th Rept. N. Y. state cab. nat. hist., 1865, p. 31, adv. sheets; 20th Rept. N. Y. state cab. nat. hist., 1868, p. 335. ? Ceraurus bimucronatus Roemer, Silurian faun. west. Tenn, 1860, p. 80, pi. 5, f. 19. Ceraurus niagarensis HaU, 20th Rept. N. Y. state cab. nat. hist., 1868, p. 376, pi. 21, f. 10-11; 1870, rev. ed., p. 427, pi. 21, f. 10, 11 ; 11th Rept. Dept. Geol. and nat. hist. Indiana, pi. 33, f. 10, non pi. 34, f. 16. WeUer, Bull. Chicago sci., 1907, no. 4, pt. 2, pi. 24, f. 21 (non 20). The most abundant American Silurian cheirurid in the collections of the M. C. Z. is one whose glabellar furrows nearly cross the glabella, and is therefore the nearest approach to a Crotalocephalus so far found in America. This type has long been known, but constantly confused with Ch. niagarensis or Ch. insignis. As has already been mentioned, Hall figured one in 1852, and Roemer a similar one in 1860. The two species sometimes occur together, as at Wauwatosa, but Ch. welleri is easily recognized by its long glabellar furrows. A second distinction is that the posterior margin of the hypostoma is rounded in Ch. welleri and straight in Ch. niagarensis. Further, the hypostoma of the latter species is tuberculated and of the former smooth. A large Cheirurus with approximately semicircular cephalon. The glabella is long, expands gradually forward. The frontal lobe occu- RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 35 pies less than half the length, and the furrows are long, curve back- ward, the last pair meeting, as usual, and the inner ends of the other pairs being very close together. The free cheeks are small, the eyes opposite the ends of the second pair of glabellar furrows and near the dorsal furrows. The fixed cheeks are wide, with coarse pits, and the genal spines are short and slender. The glabella seems to be devoid of granulation. Of the thorax, only five segments have been seen. It seems to be in all respects like that of Ch. niagarensis. No pygidium can defi- nitely be assigned to this species. All the specimens found at Wauwa- tosa seem to agree with the pygidium from Rochester which has been assigned to Ch. niagarensis. Weller has, however, figured a cheirurid pygidium found near Chicago, in which the central spines of the pygidium are shorter than the others. This pygidium is in this respect unlike the one here assigned to Ch. niagarensis and may belong to Ch. welleri. In cephalon and thorax, Ch. welleri is closely allied to Ch. quenstedti Barrande, of Bohemia. This species has the inner spines so short that the pygidium appears to possess only two pairs of spines. It may be that Ch. welleri has a pygidium intermediate in form be- tween Ch. insignis or Ch. niagarensis, and Ch. quenstedti. Measurements: — A large cranidium, one of the cotypes (M. C. Z. 14) is 33 mm. long and 62 mm. wide; the glabella is 19 mm. wide at the neck-ring and 25 mm. wide at the frontal lobe; the frontal lobe is 15 mm. long. This has about the same length as Ch. dilatatus, but the disparity of the other dimensions should be noted. The figures for Ch. welleri are always given first. Length, 33, 35, width, 62, 52; width glabella at neck, 19, 20, at frontal lobe, 25, 31. The longer specimen is the narrower, and has a wider glabella, thus showing a great reduction of the cheeks. The cranidium of a smaller cotype (M. C. Z. 630) is 14.5 mm. long, 25 mm. wide, the glabella is 7 mm. wide at the neck-ring, 11 mm. wide at the frontal lobe, and the frontal lobe is 7 mm. long. Large specimens of this species seem to have been abundant, and the largest glabella in the collection is 45 mm. long, and, so far as I know, the largest American cheirurid. Restored with the proportions of Ch. insignis, this trilobite would have a length of 145 mm. or nearly 6 inches. The Bohemian Eccoptochile claviger (Beyrich) equals this size. Formation and locality: — The types are from the Niagaran at Wauwatosa, Wise. The species occurs also in the Clinton of New York, the Waldron of Indiana, Silurian of Tennessee, and Guelph of Ontario. It is probably the most cosmopolitan species of Cheirurus. 36 bulletin: museum of comparative zoology. ChEIRURUS DILATATUS, Sp. nov. Plate 4, fig. 1, 3. Sphaerexochus romingeri ? Hall, 28th Pi,ept. N. Y. state mus. nat. hist., 1877, doc. ed., pi. 32, f. 16. Ceraurus (Cheirurus) niagarensis Hall, 28th Rept. N. Y. state mus. nat. hist., 1879, mus. ed., p. 189, pi. 32, f. 16; 11th Ann. rept. Dept. geol. and nat. hist. Indiana, 1882, p. 335, pi. 34, f. 16, non pi. 33, f. 10. In the discussion of Cheirurus niagarensis (p. 30) frequent mention has been made of the pygidium from Waldron which Hall figured. This pygidium differs radically from the pygidia which have been referred to Ch. niagarensis and Ch. welleri, in having broad, short spines, each marked by a depressed line. This appearance of the spines is probably due to crushing, in so far as the depressed line is concerned, but the spines are decidedly shorter and broader than those of the species previously described. Cheirurids seem to be rare at Waldron for a search through an ex- tensive collection from that locality in the M. C. Z. has revealed only one good cranidium, one poor one, and a pygidium. The best speci- men is a fairly well-preserved cranidium, having much the general appearance of Ch. niagarensis, only larger. On a closer examination of the proportions, however, it is seen that this form is longer and narrower than the typical specimens of Ch. niagarensis, and the glabella makes up a larger proportion of the cephalon. In a specimen of Ch. niagarensis from Rochester (M. C. Z. 625) the length is .57 of the width and in the specimen from Waldron (M. C. Z. 628) it is .67. In the first species the width of the frontal lobe of the glabella is less than half the width of the cephalon (.46). In the latter specimen it is somewhat more (.60). These appear, in figures, relatively small differences, but when the areas involved are compared it is at once seen that the glabella of the Waldron specimen is much larger than that of the specimens of Ch. niagarensis. It is of course uncertain whether the pygidium described by Hall is to be associated with the cranidium here discussed. It may belong to the same species, and they are provisionally associated. The cranidium is, however, made the holotype of the species, and the pygidium a paratype. A large Cheirurus. The cranidium is dominated by the glabella, whose frontal lobe is more than one half as wide as the total width RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 37 of the head. The glabella expands rapidly forward, the frontal lobe occupies less than one half the length, and the glabellar furrows are like those of Cheirurus niagarensis. The free cheeks are small, the eyes opposite the ends of the second pair of furrows, and a little further from the glabella than in Ch. niagarensis. The surface of the cheeks shows numerous pits, while the glabella is granulose. Genal spine rather long and straight. Only one kind of Cheirurus pygidium has been found at Waldron. The spines are short and subequal in length, broad, and flat. The ribs on the pleural lobes are distinctly separated by furrows, and the first rib on each side has a short sharp diagonal furrow. The median un- paired spine at the back is about one half as long as the ones adjacent to it. Measurements: — The type is 35 mm. long, 52 mm. wide; the glabella is 31 mm. wide at the front and 20 mm. wide at the neck-ring; the frontal lobe is 17 mm. long. Forviaiion and locality: — Found only in the Waldron shale at Waldron, Indiana. Cheirurus patens, sp. nov. Plate 4, fig. 2. This species, like the previous one, is notable for the small size of its cheeks. The glabella is of the insignis type, with short furrows, but is rather more convex and prominent than in that species. It expands rapidly toward the front, and the frontal lobe is a little longer than in any other species of the genus, forming a trifle more than one half the length of the glabella of the type. The outline of the frontal lobe is almost exactly semicircular, and thus differs from most of the other specimens seen, the ordinary outline being that of half an ellipse. Since the first pair of glabellar furrows are a little further back than on most species, the fixed cheeks which terminate here are shorter than usual, and the eye is a little further back. The cheeks are full of pits, and the glabella granulose. The pos- terior glabellar lobes are not strictly triangular but pentagonal, a statement which is true of most species of Cheirurus, but more obvious than usual in this species. Measurements: — Length about 31 mm., width, about 53 mm.; width glabella at frontal lobe 26 mm., length frontal lobe 17 mm. Forviaiion and locality: — The type is a single imperfect cranidium 38 bulletin: museum of comparative zoology. from the "Niagara" at Cicero, Illinois (M. C. Z. 629). A second specimen, tentatively referred to this species, is in the Museum of the Geological Survey of Canada, and is from the Guelph of Hespeler, Ontario. Cheirurus tarquinius Billings. Cheirurus tarquinius Billings, Proc. Portland soc. nat. hist., 1863, 1, p. 121, pi. 3, f. 22. This is a little-known species of the Ch. insignis group. It has a short, wide cephalon, narrow triangular basal lobes on the glabella, a short frontal lobe, and the first two pairs of furrows turn backward, are quite straight, and are intermediate in length between those of Ch. niagarensis and Ck. welleri. The species is especially characterized by the forward position of the eyes, which are opposite the second glabellar lobes, and the consequent small free cheeks and long fixed cheeks. The genal angles appear to be spineless. The type is No. 3081 in the Museum of the Geological Survey of Canada. Associated with it is a pygidium from the same locality. It is of the insignis type, with three pairs of spines, but the median spine is shorter and more rounded than in either Ch. insignis or Ch. niagarensis. A poorly preserved hypostoma in the same collection has the posterior end more rounded than that of Ch. niagarensis, and thus more like that of Ch. welleri. Measurements: — The type (G. S. C. 3081) is 19 mm. long, 35 mm. wide; and the glabella is 12 mm. wide at the neck-ring and 17 mm. wide at the frontal lobe. Formation and locality: — Middle Silurian at Port Daniel, Bay, Chaleur, P. Q., Canada. Also reported by Billings from Masardis, Maine. Cheirurus hydei (Weller). Ceraurus hydei Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 264, pi. 24, f. 22. This species is of more than ordinary interest, from its resemblance to a Ceraurus. The cephalon and thorax are those of a typical Cheirurus, but the pygidium is that of Ceraurus. This at once raises the question as to whether this is a Cheirurus which has developed a Ceraurus-like pygidium, or whether it is a Ceraurus whose cephalon and thorax have developed in a manner paralleling that of Cheirurus. baymond: new and oll silurian trilobites. 39 There is still a third possibility, namely, that Ceraurtis hydei is the young of Cheirurus niagarensis, with which it occurs. On this third point, Weller states that the fixed cheeks of C. hydei lack the pitted surface characterizing C. niagarensis, and that C. hydei has a border all around the cephalon, while C. niagarensis lacks it in front of the glabella. These facts seem to be borne out by the type which is now before me, and Professor Weller might have added that the glabella expand more rapidly in the young of C. niagarensis than in C. hydei, and has deeper glabellar furrows. The eyes too, of the young of C. niagarensis are much further back than those of C. hydei. Against these differences we may, however, place the fact that the thorax is alike in the two species, and more similar to the thorax of Cheirurus insignis Bey rich than to any of the Ordovician species of Ceraurus. In both Cheirurus niagarensis and Ceraurus hydei, the part of each pleural lobe between the dorsal furrow and the fulcral line is very much reduced, the diagonal furrow is very short, and the two small nodes which it separates are narrow, and one directly in front of the other, a point not brought out in Weller's somewhat generalized figure. On the fulcral line there is a row of nodes, and just inside this row is a longitudinal furrow parallel to the dorsal furrows. Beyond the fulcral line, the pleura are free, not contiguous as shown in Weller's figure. These same characteristics are shown in two specimens of Cheirurus niagarensis from Wauwatosa, Wisconsin. On the whole, it does not seem very probable that C. hydei is the young of Cheirurus niagarensis, especially as there is another species, Ceraurus nuperu^ (Billings) which has a Cheirurus-like cephalon and Ceraurus-like pygidium. The choice seems to lie between calling it a Ceraurus or a Cheirurus. Theoretically, it would seem that the Ceraurus pygidium was more specialized, and, therefore, less apt to be dupHcated than the Cheirurus head. Most of the other Cheiruri- dae, except Ceraurus, have all the spines of the pygidium approxi- mately equal. In Ceraurus -pleurexantherrvm there is a tendency in some specimens to have the basal lobes of the glabella triangular instead of square, and in Ceraurus viisneri the glabella occupies a large part of the cephalon, and the cephalon is long. Further, Ceraurus reaches the climax of size and abundance in the Trenton, the late species being smaller, and the specimens rarer. As to the thorax, I have shown that this portion of the test changes in parallel directions in many lines of the Asaphidae, and the same might well happen in the Cheirurudae. On the other hand, one would not expect a decadent race to show new characters similar to those of a race which is at its best. 40 bulletin: museum of comparative zoology. Though no other cheirurid exactly dupHcates the Ceraurus pygi- diuni, there are numerous cases among the trilobites referred to the genus Cheirurus in which there is a reduction of the inner pairs of spines of the pygidium. Thus, Weller has figured a Cheirurus pygi- dium from Lemont, near Chicago, in which spines of the inner pair are shorter than the others. In Cheirurus qucnsiedti there are only two pairs of spines, the inner pair being reduced to mere rudiments. In Cheirurus hawlei there is a still further reduction, so that there is only one pair of long spines, thus producing a pygidium which is a parallel to that of Ceraurus, though differing considerably from it in detail. The typical number of segments in a Cheirurus pygidium seems to be five, a protopygidium and four pairs of coalesced segments which originally had free spines. Among the species referred to Cheirurus by Barrande may be seen Ch. minutus Barrande with four pairs of spines, Ch. bifurcatus with four pairs, the central pair partly united, Ch. insignis and many others with three pairs and a central spine. In England, Ch. biniucronatus with three pairs without the central spine. In Bohemia again, Ch. quenstedti Barrande with two pairs of spines and two rudiments, Ch. hawlei Barrande with one pair spines and four rudiments, and in America Ch. hydci Weller and Ch. nuperus Billings with one pair of spines and three rudiments. Differ- ences in the cephalon show that this is not a progressive (or regressive) series, but apparently a number of cases of parallel development by the loss of the posterior inner pairs of spines. In view of this general tendency among the cheirurids to a reduction of the spines of the pj^gidium, it seems that more weight should be given to the cephalon then to the pygidium in determining relation- ships, and Ch. hydei and Ch. nuperus are therefore referred to Cheiru- rus. It may be proper, when the family has been more fully studied, to erect a new genus for these peculiar species. The M. C. Z. has recently acquired a fairly complete specimen (M. C. Z. 631) of this species, of which only two other specimens are now known. The specimen is from an unknown locality near Chicago, 111. This speci- men shows that the eye is very far forward, opposite the first pair of glabellar furrows. Both the genal and pygidial spines are longer than had been supposed, and as pointed out above, the pleura of the thorax, beyond the line of nodes denoting the fulcra, are free blade-like spines. Measuremejits: — The specimen figured by Weller is 24 mm. long, 14.5 mm. wide at the genal angles, and the glabella is 5 mm. wide at the back. Formation and locality: — Known onl\' from the Niagaran near Chicago, Illinois. RAYMOND: NEW AND OLD SILURIAN TRILOBITES. 41 I Cheirurus nuperus Billings. Cheirurus nuperus Billings, Cat. Silurian fossils Anticosti, 1866, p. 60, f. 20. This species was described from an isolated glabella and pygidium from Div. 3 at East Point, Anticosti. Schuchert and Twenhofel have listed it with a query from the upper part (D9) of their Gun River formation, where it is associated with Bilobites hilohus and Triplecia ortoni, in strata of Clinton age. Like Ch. hydei, this species shows the Cheirurus type of basal lobes on the glabella and the pygidium shows three pairs of spines. The outer pair or great spines are large, flat and not so long or so much curved as in most of the species of the genus. C. hydei has the great spines much more slender and further apart than in C. nuperiis. The type of this species is lost, and no further specimens have been described. Sphaerexochus romingeri Hall. S-phaerexochus romingeri Hall, 20th Rept. N. Y. state cab. nat. hist., 1868, p. 375, pi. 21, f. 4-7. (See Weller, Bull. Chicago acad. sci., 1907, no. 4, pt. 2, p. 209, for earher and later references to this species). This is an exceedingly common species in the Niagaran in the Chicago and Wisconsin areas but the pygidium is rare and usually incorrectly figured. Hall started the misrepresentation figuring the pygidium as having three spines on each side and a rounded projection at the back. As a matter of fact, the margin of the pygidium is entire, and the spines figured by Hall are the ribs on the pleural lobes. Weller produced practically a similar figure, and one of Kindle's is about the same, but the other, being of a mould, is more correct. Other describers of the species have refrained from figuring the pygidium. PLATE 1. Raymond. — New and Old Silurian Trilobites. PLATE 1. All figures natural size. 1. Bumastus decipiens Raymond. A specimen lacking the free cheeks and with the posterior portion of the pygidium incomplete. Niagaran at Wauwatosa, Wise. M. C. Z. 641. 2. The same species. Side view of the holot>'pe, showing the large eye and the absence of lip on the free cheek. Wauwatosa, Wise. M. C. Z. 642. 3. Bumastus niagarensis (Whitfield). An entire specimen. Wauwatosa, Wise. M. C. Z 643. 4,5. A ctinolobus americanus R&ynioud. Two views of the holotype. From the Racine dolomite (Niagaran), Racine, Wise. M. C. Z. 644. 6, 7. Bumastus tenuis Raymond. Two cranidia, to show the flattened form and the wide concave lip. Wauwatosa, Wise. M. C.Z. 645, 646. 8; 9. Bumastus dayi Raymond. Dorsal and side views of the holotype. Wauwatosa, Wise. M. C. Z. 647. 10. The same species. A large cephalon. Wauwatosa, Wise. M. C.Z. 648. 11. Bumastus tenuis ? Raymond. A pygidium referred with doubt to this species. Wauwatosa, Wise. M. C. Z. 649. BULL. MUS. COMP. 200L. Raymond.— Trilobites, Plate 1 3 4 W 8 II. E. f!. FISCHER, DEL. PLATE 2. Raymond. — New and Old Silurian Trilobites. PLATE 2. \ Bumastus indeterminatus (Walcott) The holotype. From the Leray- Black River at Newport, Herkimer Co., N. Y. Natural size. M. C. Z. 650. BULL. MUS. COMP. ZOOL. Raymond.— Trilobites, Plate 2 E. N. FiSCHER, DEL. Batmond. — New and Old Silurian Trilobites. PLATE 3. All figures natural size. 1,2. Trochurus nasutus (WeUer) Dorsal and side views of a cranidium from the Niagaran at Wauwatosa, Wise. M. C. Z. 632. 3. Ceratocephala goniata Warder. Mould of the exterior of a pygidium. From the Racine dolomite, Racine, Wise. M. C. Z. 633. 4. The same species. A fragment of a thorax. Racine, Wise. M.C. Z. 634. 5. The same species. A natural cast of the central part of a pygidium. Racine, Wise. M. C. Z. 635. 6. Cheirurus welleri Raymond. A cephalon and parts of five thoracic segments. Wauwatosa, Wise. M. C. Z. 630. 7, 8. Encrinurus reflexus Raymond. Dorsal view of one, and profile view of the other of the cotypes. From the Niagaran at Wauwatosa, Wise. M. C. Z. 636, 637. 9, 10. Calymene celebra Raymond. The cephalon of one entire specimen and the pygidium of another, the two specimens being the cotypes of the species. From the Niagaran at Grafton, Illinois. M. C. Z. 638, 639. 11. Calymene breviceps Raymond. An entire specimen from the Waldron shale (Silurian), at Waldron, Indiana. Holotype. M. C. Z. 640. BULL. MUS. COMP. ZOOL. Raymond.— Trilobites, Plate 3 3 n>\jrx /?>^^:::^'Cn 70 E. N. FISCHER, DEL PLATE 4. Batmond. — New and Old SUurian Trilobites. PLATE 4. 1. Cheirurus dilitatus Raymond. The holotype. From the Waldron shale, Waldron, Indiana. Natural size. M. C. Z. 628. 2. Cheirurus patens Raymond. The holotype. From the Niagaran at Cicero, Ilhnois. Natural size. M. C. Z. 629. 3. Cheirurus dilatatus Raymond. A pygidium from the Waldron shale. X 2. M. C. Z. 1273. 4. Cheirurus niagarensis Hall. Cephalon and part of thorax from Wauwa- tosa, Wise. Natural size. M. C. Z. 627. 5. The same species. A pygidium from the Rochester shale at Rochester, N. Y. X 3.4. M. C. Z. 624. 6. The same species. An hypostoma from Wauwatosa, Wise. Natural size. M. C. Z. 1278. 7. Cheirurus welleri Raymond. A cotype. Wauwatosa, Wise. Natural size. M. C. Z. 14. 8. The same species. The other cotype. The same specimen is shown in Plate 3, fig. 6. Wauwatosa, Wise. X f. M. C. Z. 630. 9. Cheirurus niagarensis Hall. A pygidium from Wauwatosa, Wise. X 2. M. C. Z. 1277. 10. Cheirurus welleri Raymond. An hypostoma. Wauwatosa, Wise. The specimen is attached to a glabella. X 1.45. M. C. Z. 1269. BULL. MUS. COMP. ZOOL. Raymond — Trilobites, Plate 4 W^xM GEORGE NELSON. PHOTO. JAN 26 iBlu Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 2. THE AUSTRALIAN ANTS OF THE GENUS ONYCHOMYRMEX. By William Morton Wheeler. With Two Plates. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM. January, 1916. No. 2. — The Australian Ants of the Genus Onychomyrmex. CONTRIBUTIONS FROM THE ENTOMOLOGICAL LABORATORY OF THE BUSSEY INSTITUTION, HARVARD UNIVERSITY. NO. 104. ' / By William Morton Wheeler. Twenty years ago Emery described a singular ponerine ant taken by Podenzana on Mt. Bellenden-Ker, in Queensland, as the type of a new genus under the name of Onychomyrmex hedleyi, in honor of Mr. C. Hedley, a distinguished Australian naturalist. The worker, which was the only phase seen by Emery, exhibited an unusual combination of characters, especially in the shape of the mandibles, clypeus, petiole, and middle and hind tarsi, the terminal joints, pulvilli, and claws of which were conspicuously enlarged. He regarded the affinities of the genus as obscure. "Its mandibles and petiole," he says, "recall the species of Amblyopone and related genera, but the frontal carinae, approximated and dilated in front, resemble those of Ponera and Leptogenys. The tibiae without spurs are not found in any other Ponerinae. The tarsi, with their enormous claws and pul- villi, have no analogue, to my knowledge, except in the Dorylinae {Aenictus, Anomma), but the insertions of the antennae and the structure of the thorax lead me to think that these resemblances do not indicate a true relationship." Ashmead (Can. ent., 1905, 37, p. 382) regarded the genus Onychomyrmex as constituting a distinct tribe of Ponerinae (Onychomyrmicini). In his recent revision of the subfamily in the "Genera Insectorum" (1911), Emery adopts Ash- mead's name as that of a sixth and last subtribe in the tribe Ponerini. While collecting in the rich tropical "scrub" in the neighborhood of Kuranda, Queensland during the autumn of 1914, I succeeded in finding not only 0. hedleyi, which had not been recorded for nearly twenty years, but also two additional species of the genus. On returning to Boston I learned that Dr. E. Mjoberg had anticipated me in finding 0. hedleyi and one of the other species, while he was collecting for the Swedish scientific expedition to Australia during 1910-1913 and that Forel had just described the latter as 0. mjobergi. The third species is described in the following pages as 0. doddi, in 46 BT lletin: museum of comparative zoology. honor of Mr. F. P. Dodd, the well-known observer and collector of Queensland insects. I was so fortunate as to discover the females of mjobergi and doddi and the larva of the former species. The female Onychom^Tmex is of such an unusual type that it seems advisable to revise the genus in such a way that some of my Australian friends may be able to recognize all the known species at a glance and to make additions to our knowledge concerning their habits. Unfortunately the male Onychomyrmex is still unknown and will have to be found before the precise status of the genus in the sub- family Ponerinae can be ascertained. Forel does, indeed, describe a male ponerine taken by Mjoberg as that of 0. hcdicyi, but he says that he does this "with a very great interrogation point." He has, in fact, no evidence that the specimen is an Onychomyrmex, except the very inconclusive fact that it was taken in the same locality (Malanda, Queensland) as the worker of hcdleyi. I deem it advisable, therefore, to assume that the male is unknown till it is actually taken in nests with the workers. Such observations as I was able to make on the habits of the three species of Onychomyrmex are recorded below in connection with the taxonomic descriptions. So far as at present known all the species of the genus are confined to Queensland, and all live in red rotten logs in the tropical rain-forest (" scrub ").^ Onychomyrmex Emery. Emery, Arm. Soc. ent. Belg., 1895, 39, p. 349; Genera Insectorum, 1911, fasc. 118, p. 96; Forel, Arkiv. f. zool., 1915, 9, p. 2. Worker. Small, slender, monomorphic. Mandibles rather long, narrow at the base, broadest in the middle, with long, curved, acute tips, their inner borders armed with a number of unequal teeth, some of which, near the middle of the series, are directed backward. Both the maxillary and labial palpi very short, 2-jointed. Clypeus very short, abrupt, with rounded, entire anterior border beset with a regular row of minute teeth. Frontal carinae small, prominent, closely approximated, enlarged and dilated anteriorly, separated by a very narrow groove. Frontal groove lacking. Eyes very small, consisting of about 6 or 8 ommatidia, situated behind the middle of the head. 1 Emery believed that the Anommi erratica of Frederick Smith from New Guinea might be an Onychomyrmex, but the description mentions none of the distinctive characters of this genus and was, perhaps, drawn from an Aenictiis. wheeler: AUSTRALIAN ANTS. 47 Antennae 12-jointed, funiculus filiform, not clavate or conspicuously enlarged at the tip. Thorax slender, with very distinct promesonotal and mesoepinotal sutures; mesonotum small, discoidal, with distinct sutures on all sides. Petiole with a short peduncle in front and a large, prominent compressed ventral projection, the node rounded, scarcely narrowed behind where it articulates hy means of its whole posterior surface with the postpetiole. Postpetiole large, convex below, separated by a pronounced constriction from the gaster, which is rather short. Sting very long and well-developed. Legs long; middle and hind tibiae without spurs; terminal joints of the middle and hind tarsi conspicuously elongated and incrassated, with very large, strongly curved, simple claws and large pulvilli. Female. Apterous and ergatomorphic. Head broadened in front and more depressed at the anterior corners than in the worker. Eyes very small; ocelli absent. Mandibles more falcate, not abruptly curved at the tips, with only a few short, blunt teeth. Mesonotum somewhat longer than in the worker. Petiole differing from that of the worker in being much broader, with a very short and narrow peduncle and lacking the ventral projection. Constriction between the postpetiole and gaster much less distinct than in the worker. Gaster much larger, elongate elliptical, sting somewhat smaller. In other respects like the worker. Larva. Slender, smooth and nontuberculate, with twelve very distinct postcephalic segments, the constrictions between which are everywhere deep and conspicuous, even at the posterior end of the body. Head short, rounded, with well-de^'eloped, slender, acute, falcate mandibles, destitute of teeth. Clypeus rather long, project- ing. Antennae very small. Maxillary sensillae long and prominent. Head sparsely, remainder of body more densely and uniformly covered with short, straight, stiff hairs or bristles. Genotype: Onychomyrmcv hedleyi Emery. The discovery of the ergatoid female of Onychomyrmex only adds to our perplexity in regard to the precise taxonomic position of the genus. Similar females are known to occur in a few other ponerine genera, notably in Acanthostichus, Paranomopone, which I recently described from .Queensland, and Leptogenys (subgenus Lobopelta), but all of these, together with Onychomyrmex, belong to very differ- ent sections of the subfamily, and the resemblances between them seem to be due to "convergence" and not to morphological relation- ship, or common phylogenetic development. The thorax of the female has simply assumed the structure of that of the worker, while the 48 bulletin: museum of comparative zoology. gaster is greatly expanded to accommodate the voluminous ovaries. On closer examination it is found that in each of the four genera men- tioned the female differs from the congeneric worker in certain peculiar characters. This will best be seen by a comparison of the worker and female Lobopelta with the corresponding phases of OnychomjTmex. Many years ago I called attention to the fact that the female Lobo- "pelia elongata Buckley of Texas has no winged female, but that each colony contains a single egg-producing individual, which agrees in all respects with the worker, except in the larger size of the abdomen and the somewhat more compressed petiolar scale. While at Kuranda I succeeded in finding two females of another species (Leptogenys (Lobopelta) fallax Mayr subsp. fortis Forel), a small-eyed form which lives, like the species of Onychomyrmex, in red rotten logs in the primeval rain-forest. One of these females was the mother of a flourishing colony of perhaps 300 workers, the other was isolated in a small cavity in a large log and was, therefore, about to start a colony. I have figured one of the specimens (Plate 2, fig. 8, 9), with the worker (fig. 6, 7) to show the difference between them (in this case greater than those obtaining between the female and worker of Lobopelta elongata) and between the corresponding phases of Onychomyrmex mjobergi and doddi (Plate 1, fig. 3-6; Plate 2, fig. 3-5). It will be seen that in the Lobopelta female the petiole is very much more compressed and more curved forward than in the worker, the thorax more convex and furnished with a small scutellar sclerite and that the head is more orbicular and less rectangular and has distinctly larger eyes and a single ocellus. In the female Onychomyrmex the eyes are not larger than in the worker, there are no traces of ocelli, the head is dilated anteriorly, with rather straight, posteriorly converging sides, and with very different mandibles, while the petiole exhibits a peculiar modification as compared with that of Lobopelta, being greatly swollen behind and much contracted in front. The female Acantho- stichus differs from the worker, according to Emery, in its rounded head, larger eyes, the presence of three ocellar pits, more widely separated frontal carinae, broader thorax, much larger abdomen, the absence of prickles on the sides of the pygidium, and a different pubescence on the abdomen. The only external differences between the female and worker Paranomopone are the presence of a median ocellus in the former and a larger postpetiole and gaster. These comparisons all point to the conclusion that in each of the four genera ergatomorphic females have been developed independently by simpli- fication, or atrophy from the primitively winged type of female during wheeler: AUSTRALIAN ANTS. 49 the long phylogenetic history of the ponerine subfamily. It is also pi'obable that the very similar " diehthadiif orm " females of the ants belonging to the subfamily Dorylinae have had a like independent origin and development. The larva of Onj^ehomyrmex (Plate 1, fig. 7; Plate 2, fig. 1, 2), in the very distinct segmentation of the body and in the structure of the head, seems to be of a rather primitive type and resembles the larvae of the Dorylinae (Eciton) and lower Ponerinae (Acanthostichus, Cerapachys), but the larvae of ants have not been sufficiently studied to enable us to draw satisfactory conclusions concerning the phylo- genetic relationships of the various genera. A study of the worker Onychomyrmex certainly reveals a number of highly specialized characters. Such are particularly the shape of the mandibles, the vestigial condition of the palpi, the small size of the eyes, and the enlargement of the terminal joint, claws, and pulvilli of the middle and hind tarsi. The degenerate visual organs show that these ants belong to the hypogaeic series and that they pass their lives concealed in the logs which gradually decompose in the moist shade of the dense tropical jungle. The powerful, toothed mandibles, long sting and great hooked claws indicate that their possessors do not feed habitually on small feeble insects like termites, but on much larger creatures such as the larvae of passalids and scarabaeids and possibly on adult myriopods and scorpions. This I found to be the case in a colony of 0. mjobcrgi, for when the log containing it was broken open, many of the workers were detected in the act of biting and stinging to death a huge lamellicorn beetle larva more than two inches in length, which they had just found in a cavity in the wood. It is not improba- ble that the colonies move from place to place in search of their prey, like the colonies of the subterranean Dorylinae (Eciton coecuvi and Dorylus), which the}' very closely resemble in behavior, color, sculp- ture, and pilosity. The species of Onychomyrmex are far from common even in Queens- land, and the few colonies I secured were the reward of many hours of search and of the destruction of many old logs in places where I was frequently attacked by land-leeches and saw quite a number of the deadly black snakes (Pseudechis porphyriacus) . Perhaps it would be possible for the collector to attract colonies by placing large beetle or cossid larvae in holes in the rotten logs usually found along the paths through the " scrub." 50 bulletin: museum of comparative zoology. Onychomyrmex hedleyi Emery. Plate 1, fig. 1, 2. Emery, Ann. Soe. ent. Belg., 39, 1895, p. 350, f. 2. ^ ; Gen. Insect., 1911, fasc. 118, p. 97, pi. 3, f. 9, 9b; Forel, Arkiv. f. zool., 1915, 9, p. 3, pi. 1, f. 3 ^ cf (?). Worker. Length 3.5-4 mm. Head about I5 times as long as broad, subreetangular, a little broader in front than behind, with straight sides and posterior border and rounded posterior corners. Clypeus with the anterior border slightly flattened, arcuately rounded in the middle, sinuate at the sides, its edge beset with about 20 minute, regular teeth. Eyes with about 6-8 minute ommatidia, situated | the length of the head from the anterior margin. Mandibles with long, abruptly incurved apical tooth and seven basal teeth of different sizes, the two in the middle of the series largest and directed backward. Antennae slender, scapes fully | as long as the head, first and last funicular joints twice as long as broad, remaining joints about I2 times as long as broad. Thoracic sutures all strongly impressed; pronotum convex above, especially in front, with convex sides, a little longer than broad; mesonotum nearly twice as broad as long ; epinotum longer and nar- rower than the pronotum, longer than broad, Avith feebly convex sides and separated in dorsal view from the pronotum by a pronounced impression on each side. In profile the thorax is distinctly impressed at the mesonotum, the base of the epinotum is nearly twice as long as the straight declivity into which it passes through an obtuse angle. Node of petiole in profile with rather straight anterior slope and convex summit, slightly concave at the posterior border; from above the node is as long as broad, rounded in front, with straight posterior border; ventral projection long and blunt, compressed and somewhat translucent. Postpetiole as long as broad, first gastric segment a little longer than the postpetiole. Legs slender. Very smooth and shining; mandibles, clypeus, and cheeks sub- opaque, the mandibles finely striated, the cl.A'peus and cheeks finely rugulose-punctate. Bodj- with fine, sparse, piligerous punctures, which are most numerous on the head, especially on its sides. Hairs delicate, pale yellowish, short, suberect, covering not only the whole body, legs, and antennal scapes but also the funiculi; wheeler: austil\liax ants. 51 somewhat longer and sparser on the thorax, abdomen, and legs than on the head and antennae. Black; thoracic sutures, sides and terminal segments of abdomen, clypeus, cheeks, and anterior portion of gula reddish castaneous, mandibles, except their teeth, antennae, and legs paler, brownish red, middle portions of femora and tibiae more or less infuscated. Queensland: Mt. Bellenden-Ker, type locality (Podenzana); Ma- landa (E. ]Mj6berg) ; Kuranda (Wheeler and F. P. Dodd). I took two small companies of this ant, unaccompanied by larvae or females, Oct. 24 and 28, e\ddently on foraging expeditions in the heart of rotten logs. One of the companies comprised a dozen, the other about two dozen workers. Later Mr. Dodd sent me eight workers which he had taken in the same locality. The ants moved rather slowly and were easily captured. OXYCHOMYRMEX MJOBERGI Forel. Plate 1, fig. 3-7; Plate 2, fig. 1, 2. Forel, Arkiv. f. Zool., 191.5, 9, p. .3, pi. 1, f. 7; text fig. 1, S . JVorker. Length 3. .5-4 mm. Head subrectangular, not more than \ longer than broad, scarcely broader in front than behind, with feebl\' and evenly convex sides, feebly concave posterior border and rounded posterior corners. Clypeus with broadly arcuate anterior border, sinuate on each side, minutely and evenly denticulate. Eyes scarcely smaller than in hedleyi, situated about § the distance from the anterior to the posterior border of the head. ^Mandibles similar to those of hedleyi. Antennae shorter, scapes only § as long as the head, first and last funicular joints nearly twice as long as broad, remaining joints not longer than broad, the more basal joints a little broader than long. Thorax differing from that of hedleyi in being stouter and in haAing the dorsal outline nearly straight in profile, the pronotum being convex only at the ex- treme anterior end and the mesonotum less impressed. Thoracic sutures very distinct but less impressed than in hedleyi. Mesonotum fully three times as broad as long. Petiole with very short peduncle, anterior surface of node more concave, its upper surface seen from aboAe distinctly broader than long, with very convex sides. Con- striction between the postpetiole and gaster somewhat deeper than in hedleyi, legs stouter. 52 bulletin: museum of compakative zoology. Smooth and shining; mandibles shining, not striate but sparsely punctate, like the remainder of the body. Punctures on the head coarser than in hedleyi, and more abundant, especially on the cheeks and sides of the front. Clypeus subopaque, rugulose-punctate. Hairs similar to those of hedleyi but coarser and of rather uneven length, pale yellow. Rich ferruginous red, clypeus darker; tarsal claws, sutures of thorax and gaster, articulations of antennal funiculi dark brown, mandibular teeth black; legs and anal segments of gaster paler and more yellowish. Female. Length nearly 5.5 mm. Head a little longer than broad and nearly as broad in front as long, with prominent, depressed anterior corners, the sides converging posteriorly, with two transverse impressions, one half-way between the anterior corner and the eye and one at the eye. Eyes as small as in the worker, but more elongate. Mandibles with less abruptly incurved tips than in the worker and with only two indistinct teeth. Thorax more robust than in the worker, the pro- and epinotum with more convex sides and the pronotum more convex above, so that the megonotum is more impressed in profile. From above the mesonotum is scarcely twice as broad as long. Petiole much larger than in the worker, with very short, slender peduncle, without ventral projec- tion; node large, very convex in front, from above more than twice as broad as long, broader than the epinotum and nearly half as broad as the postpetiole. Gaster very much larger than in the worker, more than twice as long as broad, suboblong, flattened dorsoventrally. Sculpture as in the worker, but the piligerous punctures, especially on the head, much coarser, almost foveolate and somewhat elongated on the sides of the front. Cheeks and sides of epinotum subopaque, finely rugulose-punctate. Hairs coarser and longer, especially on the body, than in the worker. Color more brownish ferruginous; mandibles, antennae, and legs more yellowish ; pleurae, sides of petiole, and sutures of gaster brown- ish yellow. Queensland: Herberton (type locality), Atherton and Cedar Creek (E. Mj5berg); Kuranda (Wheeler). October 24, I found two fine colonies of this species in rotten logs. One comprised at least 400 workers, a single queen, with the abdomen greatly distended with eggs, and a large number of nearly mature larvae but no pupae. The other colony was somewhat less populous but also contained many larvae. The ants moved rather slowly in long files through the cracks in the wood, evidently endeavoring to- wheeler: AUSTRALIAN ANTS. 53 keep in close touch with one another by means of their antennae, after the manner of the DoryHnae. They stung severely for such small insects. The worker of 0. mjobergi is readily distinguished from that of hedleyi by its paler color, shorter head, antennal scapes and funicular joints, the straight dorsal profile of the thorax, broader epinotum and petiole, deeper constriction between the postpetiole and gaster, and smooth, shining, and sparsely punctate mandibles. Onychomyrmex doddi, sp. nov. Plate 2, fig. 3-5. Worker. Length: 2-2.5 mm. Head subrectangular, about \ longer than broad, scarcely broader in front than behind, with nearly straight lateral and posterior borders and rounded posterior corners. Clypeus with broadly arcuate, finely denticulate anterior border, sinuate on the sides. Eyes very similar to those of the preceding species, situated about | the distance from the anterior to the posterior border of the head. Mandibles with the long terminal tooth less abruptly bent inward, remaining teeth rather small. Antennal scapes | as long as the head; first and terminal funicular joints fully twice as long as broad, remaining joints scarcely longer than broad. Thorax rather stout, shaped much as in mjobergi, with straight, horizontal dorsal outline, the pronotum longer than broad, rising rather abruptly from the neck, but posteriorly flattened above, its sides only feebly convex. Mesonotum somewhat more than twice as broad as long. Thoracic sutures very distinct. Epino- tum in profile with the base feebly and evenly convex and longer than the declivity which is sloping and distinctly concave. Petiole in profile with a short basal peduncle and prominent, compressed, somewhat translucent ventral projection; the node with subequal anterior and dorsal surfaces, both feebly convex; seen from above as long as broad, subrectangular, with rounded sides and straight, subequal anterior and posterior borders. Postpetiole as long as broad, very convex below and separated by a pronounced constriction from the gaster. Legs as in mjobergi. Smooth and shining, covered with small piligerous punctures, which are most abundant on the head and especially on the cheeks. Mandi- bles, clypeus, and cheeks opaque, the mandibles finely and sharply 54 bulletin: museum of comparative zoology. striate and sparsely punctate, the clypeus densely transversely rugu- lose. Pilosity pale yellow, much as in hedleyi but shorter. Color also like that of hedleyi, deep castaneous, nearly black; mandibles, except the teeth, clypeus, and frontal carinae deep brown- ish red; antennae, legs, and tip of gaster yellowish brown; coxae and middle portions of femora and tibiae darker. Female. Length nearly 4 mm. Resembling the female of mjbhergi in form, but the head is pro- portionally broader behind and without lateral impressions; differing from the worker in the shape of the head, which is broadened in front, the feebly dentate, less curved mandibles and the stouter thorax and larger petiole, postpetiole, and gaster. The sides and dorsal surface of the pro- and epinotum are more convex than in the worker and the promesonotal and mesoepinotal sutures are more impressed so that the dorsal outline is much less straight and continuous. Mesonotum not more than twice as broad as long. Petiole like that of the female mjohcrgi, the peduncle very small, the node very large, convex and rounded in front and on the sides, with straight posterior border; seen from above it is only a little more than 1^ times as broad as long, scarcely broader than the epinotum and more than half as broad as the postpetiole. The latter is separated by a very slight constriction from the gaster, w^hich is large and shaped much as in the female mjohcrgi. • Sculpture and color as in the worker, hairs considerably longer and coarser, especially on the postpetiole and gaster. Queensland: Kuranda (Wheeler). I found onh- one colony of this ant (November 1), consisting of a female and nearly 50 workers, but without larvae, in a small log in a damp, shady spot in the dense "scrub." The worker is readily distinguished from both hedleyi and mjobergi by its smaller size and less abruptly curved mandibles; from mjobergi by its color, longer head, striated mandibles and finer pilosity; from hedleyi by the straight dorsal outline of the thorax and less convex pronotum, shorter petiole, scapes, and funicular joints. i PLATE 1. Wheeler. — Aiistralian Ants. \ PLATE 1. 1. Onychomyrmex hedleyi Emery. Worker, lateral view. 2. Head of same; dorsal view. 3. O. mjobergi Forel. Worker, lateral view. 4. Head of same, dorsal view. 5. 0. mjobergi Forel. Female, lateral view. 6. Head of same, dorsal view. 7. Adult larva of 0. mjobergi, lateral view. BULL. MUS. COMP. ZOOL. Wheeler. — Omychomyrmex, Plate 1 W. M. WHEELER, Del. PLATE 2. Wheeler. — Australian Ants. PLATE 2. 1. 0. mjobergi Forel. Head of larva, lateral view. la. Mandible of same. 2. Head of same, dorsal view. 3. 0. doddi Wheeler. Worker, dorsal view. 4. Thorax and abdomen of same, lateral view. 5. O. doddi Wheeler. Female, dorsal view. 6. Leptogenys {Lohopelta) fallax Mayr subsp. fortis Forel. Worker, lateral view. 7. Head of same, dorsal view. 8. L. (L.) fallax Mayr subsp. fortis Forel. Female, lateral view. 9. Head of same, dorsal view. BULL. MUS. COMP. ZOOL. Wheeler. — Onychomyrmex, Plate 2 W. M. WHEELER, Del. FFB 12 1916 Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 3. THE SPERMATOGENESIS OF PHRYNOTETTIX MAGNUS, WITH SPECIAL REFERENCE TO SYNAPSIS AND THE INDIVIDUALITY OF THE CHROMOSOMES. By D. H. Wenrich. With Ten Plates. CAMBRIDGE, MA8S., U. S. A.: PRINTED FOR THE MUSEUM. February, 1916. No. 3. — The Spermatogenesis of Phrynotettix magnus, with special Reference to Synapsis and the Individuality of the Chromosomes. By D. H. Wenrich. contributions prom the zoological laboratory of the museum op comparative zoology at harvard COLLEGE. No. 266. TABLE OF CONTENTS. I. Introduction A. Outline of the problems B. Material and acknowledgments II. Observations A. Outline of spermatogenesis: nomenclature a. Introductory ..... b. Outline of successive stages 1. 2. 3. 4. 5. Spermatogonia Primary spermatocytes Secondary spermatocytes Spermatids Spermatozoa c. Additional features B. Synapsis a. The postspireme stages 1. Chromosome-pair A 2. Chromosome-pair B 3. Chromosome-pair C h. The conjugation of chromosomes . 1. The formation of leptotene threads 2. The zygotene stages 3. The pachytene stages C. The individuahty of the chromosomes . a. The selected chromosome pairs 1. Chromosome-pair A 2. Chromosome-pair B 3. Chromosome-pair C b. The accessory chromosome c. The spermatogonia! divisions d. The somatic nuclei D. Summary of observations PAGE. 68 58 60 60 60 60 61 61 61 63 64 64 64 65 65 66 69 70 72 72 74 75 76 76 76 79 84 86 87 91 92 58 bulletin: museum of comparative zoology. III. Discussion ....... A. Synapsis and the maturation divisions . a. Results from Orthoptera b. Recent work on synapsis B. Individuality ..... a Constancy of metaphase chromosomes 1. Constancy in number 2. Constancy in size and shape . b. Persistent organization of chromosomes 1. The selected chromosomes 2. The heterochromosomes . 3. Plasmosomes and nucleoli . 4. Persistence of chromosomes between mitoses C. Chromosomes and heredity . a. Mendelism and maturation b. Some experimental evidence D. Summary of conclusions IV. Bibliography .... V Explanation of plates Page. 95 95 95 98 105 105 105 106 110 110 111 112 114 118 118 122 125 126 135 I. INTRODUCTION. A. Outline of the Problems. Two of the most important subjects which have claimed the atten- tion of cytologists for many years are the two named in the subtitle of this paper. Every species of animals and plants is thought to have a definite number of chromosomes, which is characteristic for the species. In the process of maturation this typical, or diploid, number becomes reduced so that each functional gamete contains only half that number, the haploid number. It is generally believed that the process of reduction is initiated by a pairing of the chromosomes in the prophase of the first maturation division. It is also generally admitted that of the two chromosomes which united to form a single pair, one has been derived from the maternal, and the other from the paternal ancestor, and that these become separated again at one of the two maturation divisions. But there has been a considerable amount of disagreement as to how the pairing of the chromosomes takes place, and also differences of opinion as to which of the two maturation divi- sions results in their separation. As to the process by which the pairing of chromosomes is accom- WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 59 plished, the two opposing views that have been most wudely discussed are: — (1) that homologous chromosomes unite side-by-side (para- synapsis), or (2) that they unite end-to-end (telosynapsis). The adoption of either view, however, involves the very important assump- tion that there is a continuity between the chromosomes that appear in the earlier divisions and those that conjugate. Doubt has been expressed by some writers as to the existence of any such continuity, or individuality, of the chromosomes, and the question is regarded as one that is still unsettled. Many geneticists, on the other hand, are readily inclined to correlate the behavior of the chromosomes in maturation with the behavior of Mendelian factors in heredity. And in some cases an organization of the individual chromosomes has been assumed of such a nature that a definite portion or region of a chromo- some is concerned with the transmission of a particular factor. Such assumptions call for an analysis of the individual chromosomes to determine their inner constitution or architecture. The author of the present study has sought to throw light on all these problems. That as to how synapsis takes place was the first considered; it was taken up from the standpoint of the origin and constitution of the chromosomes of the first spermatocytes. Early in the work it was found that the only method by which conclusive results could be obtained was that of following the history of individual chromosomes. Owing to the favorableness of the material, at least three chromosome-pairs were found that possessed individual peculiari- ties by which they could be recognized through all stages from the growth-period to their division in the first spermatocyte mitosis. Pursuit of this method naturally led to a consideration of the problem of the individuality of the chromosomes, and it was found to be possi- ble to recognize one pair of chromosomes at all stages from sperma- tognia to spermatids. A further study of chromosome-individuality led to the interesting discovery that each chromosome has a definite organization, or architecture, which appears at the same stages in all the animals studied. In the following description, I have not followed the usual method of adhering to the chronological seqvience of events, but have adopted the order in which the problems presented themselves. I believe I have been able through a study of this material to demonstrate that in Phrynotettix (1) parasynapsis occurs, (2) usually the first matura- tion is equational, (3) each chromosome preserves its individuality throughout the spermatogenic cell-generations, and (4) at least cer- tain chromosomes, and probably all, have a recognizably constant organization. 60 bulletin: museum of comparative zoology. B. Material and Acknowledgments. Phrynotettix magnus belongs to the subfamily Oedipodinae of the orthopteran family Acrididae. The specimens that furnished the basis for this investigation were collected in 1907 near the Santa Rita Mountains of southern Arizona, by a collecting party from the Uni- versity of Kansas. The testes were dissected out and fixed in Flem- ming's stronger solution. Sections were cut 6-12 micra thick and stained either by Heidenhain's iron-haematoxylin, or by Flemming's tricolor, method. Material from thirteen animals was available and consisted partly of the slides used by Miss Pinney as the basis of her paper of 1908, partly of other slides prepared in Dr. McClung's labora- tory, and lastly of material sectioned and stained by the writer. The work was begun in 1911 at the University of Kansas under the direction of Prof. C. E. McClung, to whom I am indebted for the material used and for advice and kindly interest throughout. The greater part of the work was done at Harvard University during the years 1912-1915 under the direction of Prof. E. L. Mark, to whom I owe my warmest thanks for valuable criticism and suggestions and for sympathetic interest at all times. I am also indebted to Miss Eleanor Carothers, formerly a fellow student, for some collaboration, especially with reference to the so-called "plasmosomes." II. OBSERVATIONS. A. Outline of spermatogenesis: Nomenclature. a. Introductory. There is some confusion in the literature on maturation in regard to the use of the terms applied to the various steps and processes in the history of germ-cells undergoing development into gametes. This is due in part to differences in the details of the processes in the various forms investigated, and in part to different interpretations of similar stages by different authors. It therefore seems necessary, or at least expedient, to explain the terms that one wishes to use in description. A brief outline of acridian spermatogenesis follows, in connection with which the nomenclature employed will be explained. In addition, WENEICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 61 enough of the pecuUarities of Phrynotettix will be described to render clear any new terms made necessary by them. Wilcox ('94) and Davis ('08) have both given detailed descriptions of the structure of the acridian testis and have given figures or dia- grams to show the topography of the follicles of which the testes are composed. It will therefore be unnecessary to reproduce such figures and descriptions here. b. Outline of successive Stages. 1. Spermatogonia. — The spermatogonia of Phrynotettix (Plates 1, 2, fig. 1-20) behave in a manner typical for the Acrididae as de- scribed especially by Sutton ('00), and by Davis ('08). As Pinney ('08) has shown, there are 23 chromosomes, of which 22 can be arranged in pairs, leaving an odd one, the accessory chromosome (McClung, '99), or monosome (Montgomery, '06). The paired chro- mosomes may be referred to as the autosomes (Montgomery, '06). All the divisions of the spermatogonia are mitotic and are usually considered as equivalent to somatic mitoses. A detailed account of these divisions is given on pages 87-90. 2. Primary spermatocytes. — The daughter cells produced at the final spermatogonial division, as is well known, are characterized, among other things, by the growth-period and by the formation of the reduced, or haploid, number of chromosomes. For distinguishing the different stages in the prophases, Davis ('08) employed a non- descriptive method, designating successive stages by the successive letters of the alphabet. Here it seems advisable to use largely the terminology introduced by Winniwarter and by Gregoire. The telophase of the last spermatogonial division embraces a series of processes similar to those in the telophases of the earlier spermato- gonial divisions. Following the telophase, a series of changes takes place which results eventually in the formation of fine single threads. This fine-thread stage may be called the leptotene stage (Winniwarter, '00). Between the telophase and the leptotene stages occur changes which are of the utmost importance in any attempt to solve the prob- lems of synapsis and the individuality of the chromosomes. These stages may be called the jjreleptotene stages (Gregoire, '07). There may be distinguished an earlier (Plate 2, fig. 23, 24) and a later (Plate 3, fig. 25-27) preleptotbue stage. When the leptotene threads are first formed, they seem to be greatly {)2 bulletin: museum of comparative zoology. tangled and lack definite arrangement. This condition, which may be called early leptotene (fig. 28), is followed by a later leptotene (fig. 29), in w^hich the threads become oriented with one end attached at one side (the polar side) of the nucleus. Soon there appear among the single threads others which are double and twice the width of the single ones. The proportionate number of double threads gradually increases until all the threads appear double. The stage during w^hich the doubling takes place (fig. 30, 31) is the zygotene stage of Gregoire ('07). When all the threads have become double the pachytene stage (Winniwarter, '00) has been reached (Plate 3, fig. 32-34). This term continues to be applicable throughout the relatively long growth- period, and until the spireme ^ breaks up into the haploid number of segments, which become tetrads. The number of pachytene threads seems to be much less than that of the leptotene threads. The stages characterized by the appearance of separate segments of the spireme may be designated by the term diplotene of Winniwarter ('00). This term is used for the sake of consistency with the others employed, although the conditions in Phrynotettix differ somewhat from those described by Winniwarter for mammals. He describes the longitudinal split as disappearing in the pachytene stages, on account of the threads becoming twisted, and reappearing in the diplo- tene stage. In Phrynotettix the longitudinal split remains visible and little or no twisting occurs. Soon after becoming independent, a second longitudinal split occurs in the spireme segments at right angles to the first, thus forming typical tetrads, each composed of four chromatids (McCIung, '00). The first longitudinal split, which persists from the pachytene stage, may be called the priviary split, and the one at right angles to it may be called the secondary split. From the time of their formation until the succeeding metaphase, the tetrads undergo a gradual shortening and thickening. During this period they pass through the well- known figures, X's, K's, 8's, rings and crosses (Plate 3, fig. 38). The stage during which these changes occur is frequently referred to as the diakinesis stage (Hacker, '95*), but it may be simpler to call it the postspircine stage (Gregoire, '07), or the tetrad stage."^ The postspireme stages end with the establishment of the tetrad- ' The term spireme will be used to embrace the stages included under the names leptotene, zygotene, and pachytene without, however, implying anything as to the existence of a con- tinuous thread. 2 I have avoided the use of the term, prophase, in connection with the postspireme stages because it might properly be applied to the whole series of stages from the preleptotene to the metaphase. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 63 chromosomes upon the mitotic spindle of the first spermatocyte divi- sion. The number thus appearing on the spindle is twelve (Plate 4, fig. 39). One of them is the accessory chromosome, which is a dyad and passes to one pole undivided (Plate 4, fig. 41, X). The eleven tetrads represent the other twenty-two spermatogonial chromosomes arranged in pairs. One daughter cell of each spermatocyte receives eleven dyads and the other receives twelve, the additional one in the latter case being the accessory chromosome. In the anaphase all the chromosomes appear as V's, thus showing their dyad constitution (fig. 42 and 43). Before this, in the meta- phase, the separate chromatids are not discei-nible, but early in the anaphase they separate from each other at the end opposite that which is attached to the spindle-fiber, in this way giving rise to the V-shaped figures. The V-shaped arrangement persists until the metaphase of the succeeding division is reached. 3. Secondary spermatocytes. — The secondary spermatocytes pre- sent only a short resting stage. For this stage, between the formation of the secondary spermatocytes and their division, we may employ the term interkinesis (intercinese) proposed by Gregoire ('05). The extent of diffusion reached by the dyads in interkinesis is much greater than that usually described for this stage, as will be seen from figures 46 and 47 (Plate 4). The djads reappear however, in the same orientation and relative positions that they had before dift'usion. In the metaphase the dyads show the same double structure that they did in the anaphase of the immediately preceding division (Plate 5, fig. 50-52). The two monads composing each dyad are separated from each other in the metaphase, and in the anaphase are carried to the poles of the spindle (fig. 54). Half of the secondary spermato- cytes show in the plates of the metaphase eleven chromosomes and the other half twelve chromosomes, as was to have been expected owing to the non-division of the monosome in the division of the primary spermatocytes. Figure 50 (Plate 5) shows eleven and figure 51 shows twelve chromosomes. The term reductio^ial will be used to designate that one of the two maturation divisions which results in the separation of the chromo- somes that conjugated in synapsis. Correspondingly the term equa- tional will be applied to the division in which the halves of whole chromosomes are separated. Employing the terminology of Korschelt und Heider ('03), we may use the terms prereduction when the first maturation division is reductional, and postreduction when the second division is reductional. 64 bulletin: museum of comparative zoology. 4. Spermatids. — The spermatids, daughter cells of the secondary spermatocytes, undergo gradual transformation, without further divi- sion, into the mature spermatozoa. Their chromosomes undergo dis- solution, having, however, first formed a network not unlike that found in the telophase of ordinary mitoses. 5. Spermatozoa. — This term is used, as usual, to designate the functional male gametes — the end products of all the preceding processes. c. Additional Features. In Phrynotettix, as pointed out by Pinney ('08), there appear in many of the stages of spermatogenesis condensed and deeply staining granules at the ends of the chromosomes. These granules are recog- nizable in the stages where the greater part of the chromatin is ex- tended or diffuse, so that their density, contrasted with that of the rest of the chromatin, brings them into view. Figures 8, 10, 12 (Plate 1) and 14-20 (Plate 2) show them for the spermatogonia; figures 28-38 (Plate 3) for the primary spermatocytes; figures 45-48 (Plate 4) for the inter kinesis stage, and figure 55 (Plate 5), for the spermatids. These granules appear at that end of each chromo- some — including the accessory — to which the spindle-fiber attaches. They were named accordingly by Miss Pinney polar granules. In the case of certain clu'omosomes, as noted by her, similar granules also occur at the end of the chromosome opposite that to which the spindle- fiber attaches. The chromosomes are thus seen to exhibit polarity and it will therefore be convenient to designate the two ends by differ- ent terms. In the. absence of better terms, I shall call the end to which the spindle-fiber attaches the proximal or synaptic end, and the opposite one the distal end. At various stages there is a tendency for some of the polar granules to fuse together, as noted by Pinney '08, forming what I shall call coviposite granules. These are to be seen in the telophase and pro- phase of the spermatogonia (Plate 1, fig. 12; and Plate 2, fig. 14, 16), in the spireme stages of the primary spermatocytes (Plate 3, fig. 33-36), and even in the connective-tissue nuclei (Plate 9, fig. 108-110). They are particularly noticeable in the pachytene stages, for during that period quite large masses of chromatin may form by the coales- cence of a nimiber of these polar granules. The number of granules making up a composite granule is variable, but may usually be de- termined by the number of spireme threads attached to it. These WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 65 sometimes radiate out from the composite granule like the spokes of a wheel from its hub. Such a stage corresponds to the bouquet stage of Eisen ('00). At the end of the pachytene stage the granules composing the composites separate out again, apparently without having changed their identity (Plate 3, fig. 35-37). The tendency of the polar granules to remain on one side of the nucleus may be interpreted as evidence of a somewhat persistent polarity of the nucleus as a whole. It will therefore be convenient to speak of that region of the nucleus where the majority of the polar granules are congregated as the ■proximal pole, and the opposite side as the distal pole, of the nucleus. In my description of the leptotene and zygotene stages it will have been noticed that no mention is made of the contraction, or synezesis, stage (McClung '05). Such a phenomenon has not appeared in my material and, as has been claimed by McClung ('00, '05), Davis ('08) and others, is probably not normal in the Orthoptera. I shall use the term synapsis in the same sense in which it was originally used by Moore (*95), that is, to indicate the process of coupling or conjugation of the chromosomes of the last spermatogonia to form those of the jSrst spermatocyte. Following Wilson ('09), I shall use parasynapsis to denote side-by-side conjugation, and telo- synapsis to denote end-to-end conjugation. For the purpose of determining more accurately the history of the changes undergone by the chromatin through the successive stages outlined above, three individual autosome-pairs have been selected for detailed study. To distinguish them from the other autosomes, I shall call them the selected chromosomes. B. Synapsis. a. The Postspireme Stages. Of the various methods by which the diploid series of chromosomes could unite in pairs to form the haploid, or reduced, series, the two which have been more frequently defended are: — (1) that by which the members of each pair unite end-to-end (telosynapsis), and (2) that by which they unite side-by-side (parasynapsis). Evidence in favor of both methods has been gained from observations on orthop- teran material. The writer, without prejudice in favor of either view, undertook to discover which of these processes occurs in Phrynotettix. 66 bulletin: museum of comparative zoology. Efforts were first directed to a study of the postspireme stages in the hope of discovering how the segments of the pachytene spireme became the tetrads exhibiting the shapes of V's, X's, 8's, crosses and rings. Such a variety of shapes and forms presented themselves at any one of the tetrad stages, however, that it was impossible to decide which were the more primiti^'e and which the derived forms. Figure 38, a-g (Plate 3); for example, shows some of the different shapes of tetrads seen in a single stage and, indeed, in the same cyst. The only method that seemed to offer a means of securing decisive evidence on the problem was that of following the history of individual chromo- some-pairs through a large number of stages. For this purpose it was necessary to find pairs which possessed individual characteristics by which they could be recognized in all the stages concerned. Fortu- nately, at least three pairs were found which fulfilled these require- ments. For convenience in description they have been designated 'M," "5," and "C." 1. Chromosome-pair A. — This element was first distinguished in the pachytene stage, where it is a very deeply stained spireme segment. Examples of it are shown in figures 56 and 57, (Plate 5). Its differen- tial staining property is so marked and constant that it can be recog- nized by this character alone up to the later postspireme stages. But there is an additional means of identification. Like most of the pachy- tene threads, this one normally makes a loop the two ends of which approach to, or attach at, the proximal pole of the nucleus (Plate 5, fig. 56). One or both ends may become free from entanglements, but more frequently only one. In the latter case the free end, or if both ends are free, one of them, is nearly always terminated by two knobs, of which one is usually larger and less deeply stained than the other (Plate 5, fig. 57.(7; P'ate 10, fig. 113). These knobs, I believe, may be identified as the polar granules described by Miss Pinney ('08). But in this instance, as shown by numerous observations, the more promi- nent granules occur at the distal end of the chromosome instead of the proximal end, where they are found on the majority of the other chromosomes. That the expanded condition of one of the granules furnishes a means of identification, will be apparent from an examina- tion of figure 62 (Plate 6). As an exceptional occurrence these two terminal granules may be equal in size, neither one being expanded. In order to test the relative frequency of these two conditions, some counts were made and tabulated for both the spireme and postspireme stages, as follows: — WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 67 Stage Total Expanded Not expanded % Expanded Spireme HI 94 7 93.06 Postspireme 162 146 16 90.12 Both stages 273 240 23 91.26 "f^ It will be seen from this table that approximately 90% (examples counted at random) have one of the granules in the expanded condi- tion. In the postspireme stages this peculiarity appears less like an expanded single granule than as a group of closely associated small granules, typically three in number. This condition will be discussed more fully in another place (p. 112). In both the spireme and the postspireme stages the modified polar granule furnishes a ready means of identification of chromosome-pair A, especially when its staining qualities, already described, are taken into consideration. The con- stant relative size of A in the tetrad stages is also a help in identifica- tion. Figure 62 (Plate 6) indicates clearly the processes by which the spireme loop becomes first transformed into a typical tetrad, and then condensed to a metaphase chromosome. From the zygotene stage onward, there is a gradual shortening of the spireme loops or segments. The later stages of this process are to be seen in figure 62. Through- out the pachytene stage the spireme loops exhibit a median longi- tudinal cleft, usually referred to as the longitudinal split. I shall call this the primary longitudinal split. Occasionally paired granules, or chromomeres, appear to be fused together, but as a general rule, the split is continuous throughout the length of the loop. In my opinion this so-called longitudinal split is really the space between two spireme (leptotene) threads which have conjugated side-by-side. Fm-ther evi- dence for this belief will be presented later. Figure 62, c, indicates the first step in the process of forming the four chromatids of the tetrad. A second longitudinal split, at right angles to the first or primary split, begins at the proximal end (upper end in the figures) of the free spireme segment (fig. 62, c) and gradually proceeds toward the distal (lower) end (fig. 62, c-c). It will be seen from these figures that as the separation produced by the secondary split proceeds distally, the separated chromatids at the same time reunite along the plane of the primary split. The separation due to the secondary split gradually increases until the diverging pairs of chromatids extend in opposite directions, thus forming a rod-like element the two ends of which correspond to the proximal pole of the 68 bulletin: museum of comparative zoology. original spireme segment, and its middle point to the distal pole. The rod-shaped tetrad becomes oriented in the spindle of the first matura- tion division with its long axis parallel to the spindle-axis, and at metaphase separates in the middle. In other words, the plane of the secondary split becomes the plane of the first maturation division, which is therefore equational. If now we may assume that the longi- tudinally split spireme segment has represented a pair of chromatin- threads which had conjugated side-by-side throughout their length, the plane of the primary split must be the plane of the reductional division, which becomes effective in the second spermatocyte mitosis. The tetrad A also forms rings, as shown in figure 62, j, k, I (Plate 6). I have not been able to trace these rings into the metaphase to de- termine their orientation on the spindle, and furthermore I am quite uncertain whether the ring shape persists as far as the metaphase. Most of the metaphase figures show one tetrad in the form of a rod with its axis parallel to the spindle-axis, and with a constriction in the middle, as shown in figure 62, i and figure 79, A (Plate 7). Sometimes two or more rod-shaped tetrads are to be seen in the same spindle and with the same orientation. However, one of them is alwavs in a more advanced stage of division than the others, and I have been in- clined to identify this precocious one with tetrad A. Figure 62, c-t, indicates that such a conclusion is justified. Since the straight-rod condition is so characteristic of the metaphase, it may be that the rings also become transformed into straight rods by the time the metaphase is reached. The rings seem to have been formed either by a failure of the proxi- mal ends to separate during the formation of the secondary longi- tudinal split, or by a secondary union of these ends, i. e. after the split had begun. For example, if a tetrad in the condition of figure 62, c, has the secondary split completed without the separation of the proxi- mal ends, a ring would result. So also would a ring be formed by a secondary union of the two proximal ends of a stage such as is seen in figure 62, d or e. In either event the region within the ring would represent the space formed as a result of the secondary longitudinal split. If the chromatids should now begin to separate at the proxi- mal end along the plane of the primary split, as seems to be indicated in figure 62, k and /, and if this process should be continued until a metaphase chromosome such as that shown in i is produced, there is every reason to believe that it would result in a separation of the original conjugants of the pair, and therefore constitute a reductional division. On the other hand, it is possible that the separation along WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 6^ the plane of the primary split is never completed, but that the chroma- tids again become separated at the proximal end, assuming the forms shown at /to i, figure 62, and that the first division is therefore always equational. However, the possibility of an occasional reductional division as a result of the ring-formation must be taken into considera- tion. 2. Chromosome-pair B. Figure 63 (Plate 6) presents a series of stages for B corresponding to those in figure 62 for A. This series of stages of B supports the conclusions reached from a study of A in regard to: — (1) a probable parallel, association in the pachytene stages of pairs of threads, each representing individual chromosomes; (2) the formation of the tetrad by, first, a separation along the plane of conjugation (?". e., the primary longitudinal split) and, secondly, by a splitting of each of the original conjugants (the secondary longi- tudinal split) ; and (3), as a result, an equational division of the tetrada at the first division. This chromosome-pair (B) is characterized by the presence of large and well-marked polar granules at both ends and by a similar large granule not far from the middle, though always somewhat nearer the distal end. Leaving aside the formation of rings, the chief difference in behavior between A and B is that jn the former the plane along which the greatest separation takes place before metaphase is that of the secondary longitudinal split, while in the latter the greatest separa- tion takes place along the plane of the primary split. This results in A becoming extended in the direction of the spindle-axis, as already described, while B becomes extended at right angles to this axis. In the latter case the separation along the plane of the primary split does not become complete at the expense of the separation along the plane of the secondary split, but the latter separation persists for a short distance, giving rise to a cross with unequal arms (fig. 63, g, h). The short arms terminate in the proximal or synaptic ends of the chroma- tids, while the longer arms terminate in the distal ends. However, these differences in behavior between A and B are not fundamental, since the final result, an equational division, is the same in both cases. But they are indications of the individual peculiarities of these elements. It should also be pointed out that such differences could easily be misinterpreted, if only parts of the histories of the pairs were known. It is important to note that the drawings of the series shown in figure 63 were all taken from sections of a single testis. In searching- for the same element in other individuals, I was surprised to find the 70 bulletin: museum of comparative zoology. condition shown in figure 64, a-h. In this series are found the same differentiating characters that have ah-eady been described for B, except that one member of the large pair of granules at the distal end is lacking. In other words, we have to do here with a pair, composed of unequal elements, which differs from its homologue in another individual, composed of equal elements, by the absence of a definite part of one of the components. Examination of all the thirteen individuals demonstrated that eleven of them possessed this second or unequal type, while only two showed the equal type. If there could have been any doubt about the sequence of events in the transformation of a spireme segment into a tetrad and the sub-' sequent equational division in the case of chromosome-pair A or the equal type of B, the beha^•ior of this unequal type of B, as shown in figure 64, must certainly make the subject clear. In this instance, on account of the difference between the two members, it is possible to identify them in such a way that there can be no question as to the two planes of longitudinal splitting. The figures have in all cases been made with great care with the aid of a camera lucida and are faithful reproductions of the conditions seen under the microscope so far as they can be represented by the method of reproduction used. In the early stages of the transformation of the spireme segments into tetrads, the separate chromatids are not distinguishable through- out the whole length of the segment. This is due in part to a closer association of the chromati<^ls anfl in part to the fact that one of the longitudinal splits becomes more pronounced at one end and the other split at the other end of the tetrad. Somewhere between the ends, therefore, there is a crossing or apparent chiasma. At the point of the crossing the chromatids at first appear to be fused together (figs. 63, d and 64, d). Very soon, however, the confusion disappears, the chromatids become distinct, and their relationships easily discerni- ble, as shown in figures 68, r, and 64, r. In botli these cases the wide separation at the proximal end has been along the plane of the second- ary longitudinal split, and that at the distal end along the plane of the primary split. The resulting crossing, or apparent chiasma, is a perfectly normal and natural result of these processes and indicates nothing in the wa^^ of a breaking or recombining of the parts of chromatids. 3. Chromosome-pair C. — Figure 65 (Plate 6) shows one form of the third of the three selected chromosome-pairs. In this case the two components are very unequal in size, one of them possessing a very large, condensed mass, or granule, of chromatin at its distal end, WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 71 while the other has none. It will be noticed (fig. 65, b, c) that the details of the two components are quite homologous up to the large distal granule, and that the point of attachment of this large granule seems to correspond to the distal end of the smaller component. These considerations would lead us to suppose that here, too, as in the un- equal type of B, the difference between the members of the pair may be due to the loss by one of them of a definite part possessed by the other. In this case, however, no such equal pair has been found as occurs in B when both members possess the part in question. The side-by-side association of the members of this pair is as evident as it was for A and B and the relations of the two longitudinal splits are the same. In regard to the mode of distribution of tetrad parts in the first maturation division, however, we meet in this case a curious exception to the general rule. This pair divides equationally, as shown in figure 65, h-j; but it sometimes divides reductionally as shown at k-m, same figure. From casual inspection it appeared that the divi- sion occurred as frequently in the one manner as in the other. But in order to test the relative frequencies of the two methods, 928 cases chosen at random were counted and it was found that of this number 472, or 50.8%, were in process of reductional division, while 456, or 49.2%, were dividing equationally. It would seem from these counts that the method by which the tetrad divides is a matter of mere chance. This is the more apparent when we take into consideration the fact, brought out by extended observations, that the two methods occurred side-by-side in the same cysts. It may be that the shape or position of the tetrad when it is first brought under the influence of the mitotic spindle determines the mode of division. The fact that this unequal pair divided in the first division reduc- tionally a part of the time made it possible to study the distribution of the two conjugants with reference to the accessory chromosome, which goes to one pole undivided. It was soon found that either member of the pair could accompany the accessory into the second- ary spermatocytes. Consequently counts were made to determine whether the two kinds of distribution occurred with anything like equal frequency. Out of 421 cases counted at random 216, or 51.3%, were found to show the larger member going to the same pole as the accessory (Plate 10, fig. 121, C), while in 205, or 48.7%, of the cases the srnaller member was going with the accessory to the same pole (fig. 120, C) . These results seem to furnish a good example of chance dis- tribution of clu-omosomes at maturation. The behavior of these three selected clu-omosome-pairs, as described 72 bulletin: museum of comparative zoology. above in detail, seems to me to establish very definitely that the association of paired eliromosomes in the pachytene stages is one in which the members lie side-by-side throughout their entire length, and therefore exliibit parasynapsis. I should further add that while I have not singled out any other members of the complex for individual study, a careful analysis of the other spireme segments and the deriva- tive tetrads indicates that the condition of parasynapsis is realized for the entire series. I was thus able to analyze the stages of the complex as a whole after following the history of the selected indi- vidual pairs, whereas previously I was unable to reach a definite conclusion. As to the method of division in the first spermatocytes, the evidence presented indicates that B, always, and A, in most cases, divide equa- tionally, while C divides either reductionally or equationally and with equal frequency by each method. My study of the other tetrads leads me to think that, as a general rule, they divide equationally in the first division. Where the first division is equational the second is regarded as reductional, and we therefore have postreduction. The general rule has its exceptions, however, as already noted in the case of C and possibly sometimes in the case of A. b. The Conjugation of Chromosomes. 1. The formation of leptotcne threads. — The evidence for para- synapsis derived from a study of the postspireme stages, as presented in the preceding paragraphs, has not embraced the actual process of conjugation; and it therefore remains to be demonstrated that a side-by-side conjugation does take place. But it is even more im- portant to show that the conjugants are actually chromosomes, the morphological descendants of the telophase chromosomes of the final spermatogonial division. Figure 21 (Plate 2) shows a side view and figure 22 a transverse (optical) section through the chromosomes of cells nearing the end of the telophase of the last spermatogonial divi- sion. The side view shows the chromosomes already partly diffused, but each one occupies a definite territory, so that there is no question as to their persistent individuality, except for the coalescence of some of the polar granules. But, as I shall point out later, the polar gran- ules do not necessarily lose their identity when they unite into the compound masses. The optical section, figure 22, shows even more plainly the persistent individuality of the chromosomes up to this WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 73 point, for there still can be seen the remnants of the vesicular walls which surrounded each chromosome in the earlier telophase. There are 21 of these chromatin-masses, or "blocs" (Janssens, '01), shown in this optical section, and that is sufficiently close to the total number, 23, to indicate that all the chromosomes are still independent, except for the union at the polar ends, as already mentioned. In figure 21 it will be noticed that the diffusing chromatin is dis- posed roughly in the form of spirals. Figures 23 to 29 indicate what becomes of these spirals in the " blocs" of chromatin. I am not quite sure of the exact succession of stages here, but believe they are about as shown in the successive figures. It is possible that figures 23 and 24 — which are side view and optical section, respectively, of the same stage — are no earlier than the stages shown in figure 25 (Plate 3). However that may be, the evidence seems to indicate that each of the blocs at stages such as those shown in figures 21 and 22 gives rise to a single fine thread, at first much coiled but later much elongated. The side view shown in figure 23 is at a stage the casual examination of which might lead one to suppose that the chromatin was in a hope- less tangle without any definite arrangement whatever. But careful focussing and patient study revealed what I have tried to show in figure 23, viz., that the chromatin is still disposed, for the most part, in separate blocs, but that a very much coiled and convoluted thread is forming within each one of these territories. Some have unraveled to a considerable extent, and have become extended in various direc- tions through the nuclear sap. But each seems to be a continuous thread, despite some tendency for the ragged edges at times to be connected with adjacent threads. In the optical section of this stage (fig. 24) it will be seen that the blocs have remained in place and separate from each other for the most part, though some anastomosis of the linin fibers has taken place at the periphery of the blocs. On the other hand, there are still some remnants of the previously existing vesicular walls, as shown in the left side of the figure. When one focusses up and down on such a cell, it is possible to follow in some cases the tliread which is differentiating out of the net-like structure of each bloc, but in optical section the reticulum is more apparent than the continuous thread. The section shows nineteen blocs, which number is not far from the somatic number of chromosomes (23). Figures 23 and 24 represent what I have called the preleptotene stage. At the stage shown in figure 25 (Plate 3), which I believe to be slightly more advanced than the one in figures 23 and 24, the amount of anastomosis between adjacent chromatic elements seems consid- 74 bulletin: museum of comparative zoology. erably greater than in the stage last described. The anastomosis is to be seen more particularly at the sides of figure 25. Through the middle of this figure the individual spiral threads seem to be more easily distinguishable, and I am inclined to believe that the two which stain more deeply than the others are the members of the A pair of chromosomes. The stages including and following this reticular stage are hard to represent in a drawing of the kind employed, owing to the difficulty of portraying in their natural relations the parts seen at different planes of focus. Careful study has always convinced me, however, that the uncoiling and elongating threads are single, con- tinuous, and not united into an indiscriminate network. I have selected in figm-es 26 and 27 views favorable for di'awing where some of the threads, at least, are definitely separate and continuous across the diameter of the nucleus. At the stage represented in figure 28 (Plate 3) the unwinding of the coiled threads has been completed, but the threads have as yet no definite orientation. At the somewhat later leptotene stage shown in figure 29 the threads are finer and less homogenous than in the earlier stage, the substance of the thread seeming to have become more distinctly differentiated into a linin fiber and cliromatic granules, the latter scattered at irregular intervals along the fiber. Moreover, in this later stage the threads appear to be definitely oriented, with one end attached at the proximal pole of the nucleus. The threads then take a course thi'ough the center of the nucleus or near its periphery, extending wholly or partly across and then turning back with a wide curve. 2. The zygotene stages. — In figure 30 (Plate 3) some of the threads are double, others are single, and it would be difficult to decide from a casual examination of this stage alone whether or not the double threads had arisen by a splitting of the single ones. In the case of one or two of the double threads, however, as may be seen at the left side of the nucleus, the double condition does not continue throughout the whole length, but towards the distal end of the nucleus the thread is seen to branch into two single threads. I interpret tliis branching thread as one in which the parallel conjugation has not yet been completed. Another instance of the same kind may be seen in figure 31, which represents a stage somewhat more advanced than that of figure 30. These appearances lead me to believe that conjugation begins at the side of the nucleus corresponding to the proximal ends of the leptotene threads, and proceeds gradually toward their distal ends. It is further evident from these figures that conjugation is WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 75 not a simultaneous process for all the chromosome-pairs, but that it is a gradual process, some conjugating earlier than others. Just how the members of the different pairs are enabled to select their mates is a very puzzling question, but probably the stretching and orientation of the threads as shown in figure 29 might facilitate this process. That some of the pairs conjugate quite early, is shown in figure 30, where it may be seen that in selected pair B conjugation is complete. In figure 27, which is of a very much earlier stage, there are to be seen two of the still hazily defined threads lying side-by-side. They are similar enough in their constitution to be regarded as the two members of a pair, and it would not be surprising if conjugation should begin at a stage as early as this. As an additional detail it should be pointed out that the bead-like granules which are strung along the threads of the leptotene and zygotene stages are not always of exactly equal size in the two con- jugating elements. Tn figure 58 (Plate 5) the example of chromo- some-pair B well illustrates the disparity in size between the two members of some of the pairs of granules. This condition may well answer the criticisms of those who hold that the accuracy with which the granules are paired could be accounted for only on the assump- tion that they arose by a splitting of single granules into equal parts. I am able to show in this case that the members of each pair of granules are not always of equal size. 3. The pachytene stages. — ■ Figures 32 and 33 (Plate 3) are of early pachytene stages. It sometimes happens that even at such stages there may remain one or two pairs of threads that are not fully conju- gated, though I have not added a drawing of such a condition. In the case of some of the pachytene threads of figure 32, complete loops have been formed, both ends being attached at the polar region. The formation of such loops is not necessarily the rule, however, as has been indicated already in connection with the spireme loops of the selected chromosome-pairs (fig. 56-61, Plate 5). In figure 33 a scat- tered arrangement of the polar granules is to be seen, though they have coalesced to form several composite granules. Figure 34, of a later pachytene, exhibits one of the large composite granules. Figures 35-37 indicate how the composite granules break up into their com- ponent polar granules. A comparison of the examples of chromo- some-pair B in figures 30 and 35 will indicate the extent of the process of gradual shortening which takes place during the pachytene stages. It will be noticed that the line of separation between the threads which have conjugated (i. e., the primary longitudinal split) remains visible throughout the pachytene stages. 76 bulletin: museum of comparative zoology. C. The Individuality of the Chromosomes. a. The selected Chromosome Pairs. The method adopted in the study of the subject of synapsis — that of following the history of individual chromosome-pairs — has natu- rally led to a consideration of the subject of the individuality of the chromosomes, that is, their persistence as morphological entities tlirough all the stages of nuclear activity. I have already attempted to demonstrate that each of the chromosomes of the last sperma- togonial division gives rise to a single leptotene thread and that these single threads conjugate two-by-two in the zygotene stage. It will be more convincing, however, if we can follow some particular chromo- some-pair through these difficult stages. 1. Chromosome-pair A. — As the chief characteristic by which the chromosome-pair A could be recognized in the pachytene and later stages of the first spermatocytes, I have already described its great density and staining capacity. If there is a persistence of individual chromosomes from the spermatogonia to the spermatocytes, we should expect to find in the former a pair of chromosomes exhibiting th( same peculiarities that the pair did in the later generation. Such a pair can, indeed, be found in the telophases not only of the last spermato- gonia! division but of the earlier spermatogonia as well. Figures 66 and 67 (Plate 6) show such pairs of chromosomes more deeply stained than their fellows. Figure 66 shows one of the earlier generations of spermatogonia, as is indicated by the vesicular condition of the ac- cessory chromosome, and figure 67 represents a telophase of the last spermatogonial division, as is shown by the condensed accessory at this stage. It is difficult to follow all the changes that these chromosomes undergo in their transformation into pachytene tlu'eads, but I believe that most of the stages are represented in the series of drawings, figures 67-78. Figure 67 corresponds to a stage midway between those shown in figures 21 and 23 (Plate 2). Figure 68 is of a stage corresponding very closely to that in figure 25 (Plate 3). In figures 68 and 25, two bands or "blocs" of chromatin can be seen which are more deeply stained than the other chromatin-blocs. The accessory chromosome is distinguishable by its characteristic density and its position at the periphery of the nucleus. The polar granules are also distinguishable. The chromatin in these darker blocs {A in both WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 77 figures 67 and 68) shows a more or less well-defined spiral condition. This spiral is better shown in figure 69, where it is more unravelled. Very soon after the process of uncoiling gives rise to the leptotene threads, stages in conjugation may be seen. Figure 70 shows an early leptotene stage with two threads which stain more deeply than the others, having conjugated as far as they can be traced in this particular section. I think we may identify these denser threads as the members of the chromosome-pair A. The two sides of figure 70 are drawn differently. The left side is diagrammatic and is intended to represent the apparent entanglement of the leptotene threads. On the right side an attempt has been made to follow individual threads. Careful study makes it evident that the tlireads, instead of anastomosing, as they appear to do when one makes only a superficial examination of them, are really continuous and distinct for certain distances. The difficulty in following in- dividual threads is due to the fact that after the early leptotene stage the chromatin collects into chromomeres, which are strung along a linin fiber, so fine and stainless in some places that it is scarcely trace- able. When two such fibers cross each other in close proximity it is sometimes almost impossible to trace the independent courses of the two in the region of the apparent intersection. There is less difficulty, however, in tracing the threads of ^. At the stage shown in figure 71 — which corresponds with that in figure 29 (Plate 3) the threads are very fine and well oriented. In this nucleus there can be seen a loop of heavier threads (A), which have the ap- pearance of being two, loosely wrapped around each other. The ac- cessory, as shown at X, also forms a he^vy spireme loop at this stage, but it is so much heavier than the one described that there can be no confusion between the two. The deeply staining loop of interlaced threads I interpret to be the spireme of the chromosome-pair A. In figure 73 is shown an yl-spireme which has not completed its conjuga- tion. It will be noticed that of the other threads in this nucleus some are double and some are single; and, furthermore, that the double ones are twice the width of the single ones. In figure 74 the spiremes of the pair A have completely conjugated, though the general appearance of the cell indicates that the stage is no further advanced than that shown in figure 73. Figures 75-78 (Plate 7) show the pair A in various stages of conjugation at stages closely corresponding to those shown in figures 73 and 74. I have already traced the pair A from the pachytene stage to the metaphase of the first spermatocyte division, so that it now remains to 78 bulletin: museum of comparative zoology. examine only the stages following that division. Figure 80 (Plate 7) shows a telophase of the first spermatocyte division as seen when looljing from the equator toward the centrosome. There are eleven dyads here, and since the whole number could easily be counted, the accessory is not present. One of these dyads is more deeply stained than the others, and, judging from its size relations, I think we may identify this dyad as one from chromosome-pair A. This conclusion receives still stronger support from figures 81 et seq. Figure 81 is of a stage slightly later than the one in figure 80, and here we can see the dyad A in addition to the accessory dyad, which is less deeply stained than the others and is surrounded by a well-defined clear space, as indicated by the dotted line. In figure 82 is drawn a telophase in which the dyad A is shown in both the daughter cells. From these figures (80-82) it is apparent that this element cannot be confused with the accessory at these stages. In figures 83 and 84, however, it is less easy to distinguish between them. But a long and careful study has convinced me that the accessory, having early passed through a stage of greatest diffusion, soon becomes condensed, while the other dyads are undergoing dissolution. Dyad A, on the other hand, at first remains more condensed than the others and then gradually be- comes diffused like them. Figure 83 shows an early interkinesis stage in which the large accessory dyad (X) is more condensed than that shown in figure 81, but where dyad A is still more dense. In figure 84, which is of a stage not much further advanced, the accessory is seen to be the most condensed dyad (X), whereas A has gone far toward its stage of diffusion corresponding to that of the other chromosomes. That the accessory remains condensed throughout interkinesis is further shown in figures 46 and 48 (Plate 4). Figure 48 further shows that in the prophase of the second spermatocyte the A dyad condenses earlier than any other dyads except that of the accessory. It was impossible to trace the A dyad into the metaphase of the secondary spermatocyte, but in the telophase it may again be recog- nized by its characteristic deeper staining and by its size relations. In figure 55 (Plate 5), which is a polar view of such a telophase, three deeply staining chromatic masses are shown. The larger one (X) is probably the accessory, the next in size, the monad of A, and the smallest, a monad of B (p. 79). Figure 85 (Plate 7) shows a somewhat later telophase, in which diffusion has progressed a little beyond that seen in figure 55. About the same relative staining qualities and relative sizes are seen as in figure 55. The accessory appears in only half of the secondary spermatocytes and spermatids, however, and WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 79 figure 86 is of a pair of spermatids in which the accessory does not occur. The diffusion process has here proceeded beyond that shown in figure 85, but the two more deeply staining masses, representing the monads of A and B, can readily be distinguished. We may on the strength of this evidence say that the chromosome- pair A can be traced from the spermatogonia to the spermatids, thus demonstrating a case of morphological identity through all these generations and stages. 2. Chromosome-pair B. — If morphological continuity is the gen- eral rule, and if the peculiarities of the chromosome-pairs B and C are distinctive enough, we should be able to trace the latter as we have traced A. In many stages, however, these smaller pairs are not so easily recognizable as was the pair A, but it has been possible to ob- tain good evidence for individuality even through them. I have called attention to a dyad in interkinesis, and a monad in the spermatids, which seem to satisfy requirements for identification as the element B. In figures 80 and 83 (Plate 7), for example, is seen a dyad smaller than A, which stains almost as deeply as the latter. An element with similar properties is to be seen in figures 55 (Plate 5), 85 and 86 (Plate 7). This element {B in the figures) has such size relations when compared with A and the smallest element (as seen in figure 80) as we should expect in B; when we consider, further, that in the postspireme stages B stained more deeply than the majority of the other tetrads, the staining qualities exhibited in these later stages should also furnish a means of identification. When we look at the spermatogonia! telophases of the same indi- vidual from which figure 63 was taken, that is, one in which the components of pair B are equal, we can readily find a pair of chromo- somes that possesses the chief characteristic by which B was recog- nized in the postspireme stages, namely, the presence of a prominent polar granule at each end and a third not far from the middle, though nearer the distal end. Examples of such spermatogonia! telopliases are sliown in figures 87-96 (Plate 8). A further consideration of these stages is given on page 83. The study of cliromosome-pair B in the growth-period has furnished some of the most interesting data on the subject of chromosome individuality that I have secured. An analysis of this pair in its extended condition in the pachytene stages of the first spermatocyte was made for one of the specimens (no. 772) and then comparisons drawn between the conditions in this and those in all the other animals in the series studied. 80 bulletin: museum of comparative zoology. Figure 58 (Plate 5) shows the element during the zygotene stages — as indicated by the incompletely conjugated pair of threads near the middle of the figure — in a condition of complete parallel association for the two conjugants, but a condition in which the members of the pairs of granules, the chromomeres, are distinct. A close examination of this spireme of B discloses a series of cliromomeres in addition to, and smaller than, the three already mentioned as characterizing the element. For convenience in description the more prominent granules or chromomeres will be given separate designations. The five granules which I wish to mention more particularly will be numbered in order from the proximal (no. 1) to the distal end (no. 5, figure 58). I shall also call attention to the two paii's of small granules between numbers 3 and 4 and to the two pairs of still smaller ones between 2 and 3. I should not omit to direct attention to the series of granules between numbers 1 and 2 and between numbers 4 and 5, but detailed considera- tion' of those already mentioned will probably suffice for the purpose in view. I was at first impressed by the constancy in relative size and posi- tion with which some of these granules recurred in different examples of B and at different stages in a single individual (no. 772). It then occurred to me to compare the same element at about the same stage for all the thirteen animals from which material was available. Figure 97 (Plate 8) is the result, each of the separate draAvings having been taken from a different animal. The constancy with which the minute details of size and arrangement of the parts of this pair were repeated in all of the individuals was surprising. Not only are the five more prominent chromomeres repeated in approximately the same relative sizes and positions, — as shown in figure 97, where corresponding granules are connected by dotted lines, — but there is likewise a strik- ing correspondence in the more minute details. For example, the segment between the granules numbered 3 and 4 always contains two pairs of granules of about the same relative size, though they vary somewhat in relative position. On the other hand, the segment be- tween 2 and 3 is characterized by the entire absence of any prominent granules. In some cases, however, as in/, i, and k, figure 97, granules can be made out in this segment, and when this is possible there are always two pairs of very small ones in the same relative positions. It is true that there are some variations in the appearances of the segments between granules numbered 1 and 2 and between 4 and 5, as well as differences in the actual size of the numbered granules. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 81 These variations may be due to one or more of several causes: — (1) DiflFerenees arise on account of slightly different reactions to the fixatives and stains. (2) There is a tendency for adjacent granules to fuse, thus causing apparent variations in number and relative size. (3) There is a slight difference in appearance at different stages. (4) The different positions assumed by the element with reference to the optical axis of the microscope may account for some variation in appearance. (5) Some individual variation from animal to animal might be expected. It will be noticed that the distal granule (no. 5) is single in all the individuals except those represented at b and c, where it is double. This is in accordance with the statement previously made (page 70) that chromosome-pair B is unequal in eleven and equal in only two of the thirteen animals studied. It will also be noticed that the granules at the proximal end (no. 1) frequently become associated with other polar granules in a composite granule (a, h, i, j, I, m, fig. 97), and that with one exception (/) the distal end is free. The formation of composite granules is a characteristic feature of this material, as already noted on page 64. One of the granules of the proximal pair (no. 1) of individual /, figure 97, is seen to be enlarged and less deeply stained than its mate. Another example may be seen at k. I believe this to be an example of a modification similar to that described in connection with the distal granules of the pair A (p. 66). In B, this condition appears with much less frequency, for in a count of 84 cases taken at random from one individual only 14 (16|%) had one of the granules in the expanded condition. This modification may persist into the tetrad stages, as was the case with A. No case was found in which both granules were expanded. In order to test the variability of the details of constitution of the element 5 in a single animal, a study was undertaken with this object in view. Sixteen drawings (fig. 98, a-p) were made of examples taken at random from a single slide. Comparison shows about the same degree of constancy in the composition of the elements here as in the set from different animals. Some of the variations may be pointed out. For example, the relative lengths of the segments 1-4 and 4-5 in example a, figure 98, are somewhat different from those in example h. I think we may assume that the spireme threads possess some elasticity and that the variation in arrangement, association, and posi- tion of the several segments of the spireme may frequently bring about stresses which may stretch some of the threads or parts of threads to a 82 bulletin: museum of comparative zoology. greater or less degree. The tendency for adjacent granules to fuse probably accounts for some of the variations to be noticed. If one will compare in order the examples I, m, a, and c, (fig. 98) the different steps in the fusion of granule no. 4 with the smaller, yet prominent, granule close to it will be seen. As the threads shorten during the later pachytene and postspireme stages, this coalescence of adjacent granules becomes more noticeable and the individual granules all finally lose their visible identity in the compact metaphase chromo- somes. It will be observed that the members of a pair of granules may also appear to be fused together into a single ma|SS. An example of this is seen in figure 98, n, granules 4 and 5. This fusion must be very temporary in character, since it is not the general rule, and since the granules separate again in the postspireme stages, as shown in figure 63 (Plate 6) ; yet so close an association of these granules apparently offers opportunity for the exchange of chemical substances between them. In the case of the proximal granules (no. 1), the members may not only fuse with each other but, as previously noted, characteristi- cally unite with the polar granules of other chromosome-pairs to form the composite granules. The association is fully as close as in that of any single pair, for frequently all traces of the outline of individual granules is entirely lost, as, for example, in figure 34 (Plate 3). Al- though the individual granviles separate out again in the postspireme stages, if we admit that there is an exchange of chemical substances between members of a single pair of granules, I think we must also assume it for the polar granules of the different chromosome-pairs. At j (fig. 98, Plate 8) may be seen another example of an expanded polar granule, such as has already been mentioned. The possible significance of this peculiarity will be discussed on page 112. It will be instructive to compare the members of particular pairs of granules. Figure 58 (Plate 5), as already mentioned, represents a zygotene stage. The paired chromatic threads near the middle have jiist begun to conjugate, while in the case of chromosome-pair B, in the left half of the figure, the two conjugating threads have only recently come to lie side-by-side, for the members of the different pairs of granules are yet distinct. This condition fortunately gives us an opportunity to compare the relative sizes of the members of each pair. On examination it will be seen that the members of the pair numbered 4 are not equal in size. This is also true for the pair numbered 3. In the case of number 4, the disparity in size between the two granules is considerable, and it is interesting to observe that this difference in WENKICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 83 size can frequently be noticed throughout the pachytene stages. Examples of this may be seen at b, e, i, I, and o in figure 98 (Plate 8), which are drawn from the same individual as figure 58. Similar conditions are also to be found in other individuals, as will be seen in figure 97,. a, c, e, j, and /. This pair of chromosomes can be recognized in the spermatogonia by the presence, in the telophase, of the three most prominent granules, those I have numbered 1, 4, and 5 in the pachytene stages. Examples of such telophases are represented in figures 87-96. In two cases, where the cliromosomes had become considerably elongated in the general diffusion process of the telophase, I was able to make out granules 2 and 3 also with their characteristic relative positions and sizes. These are shown in figures 95 and 96. Where both chromo- somes of the pair are recognizable in the same nucleus, there seems to be in every case a difference in size between the two middle granules (no. 4). This difference is probably directly related to the difference noted in the zygotene stage (fig. 58) and the pachytene stages (figs. 97 and 98). Thus, aside from finding a striking degree of correspondence in the minute organization of the chromosome-pair B for all the individuals studied (in the pachytene stages), it has also been possible to trace the pair through all the stages from the spermatogonia to the spermatid, except in the preleptotene and leptotene stages. Figures 30 (Plate 3) and 58 (Plate 5) show that conjugation is completed at a relatively early stage in the zygotene. This precocious conjugation is possibly facilitated by the relatively small size of this pair. The failure to recognize the pair in the leptotene and immediately preceding stages is probably due to the fact that it has not so great a differential stain- ing capacity as has pair A, and to the lack of sufficiently long con- tinued study with this object in view. A further peculiarity of chromosome-pair B may be seen upon an examination of figures 99 and 100 (Plate 9). There it will be seen that one end of the tetrad has a peculiar roughened or brush-like appearance, to which McCIung ('14) has already called attention. It will be noticed in the same drawings that the accessory chromosome also presents a similar appearance. Furthermore, a like condition is to be seen at the longer end of C, as shown in figure 100, and at the end of some of the other autosomes, as seen in figure ^9. The rough- ened contour of the accessory in both metaphase and anaphase of the first spermatocyte division was noted for Phrynotettix by Miss Pinney ('08), and has been described for other species of Orthoptera, 84 bulletin: museum of comparative zoology. for example by Davis ('08) for Dissosteira and Stenobothrus, and McClung ('14) for various Acrididae. But no one, so far as I am aware, has described such a condition for any of the autosomes. Figure 99 is from a sUde that had been treated with Heidenhain's iron-haematoxyhn stain, but the destaining process had been carried farther than in most of the other sUdes. Figure 100 is from another individual, the slides of which had been stained by Flemming's tri- color method, but had not been excessively differentiated. It will be noted that the autosomes in this figure do not exhibit the roughened synaptic ends that are seen in figure 99. It seems probable, therefore, that differences in the staining process may have much to do with the appearance or non-appearance of the roughened condition. In heavily stained slides even the accessory, as well as the tetrads B and C, may appear with a smooth contour. In this connection, I may call attention to these several points: — (1) Tetrad B is unequal in both the cases figured and the roughened end corresponds to the large distal granule on the larger conjugant (see fig. 64, Plate 6). (2) Tetrad C is likewise unequal and the roughened end also corresponds with the large distal granule at the end of the larger of the two com- ponents (see fig. 65). (3) The polar granules usually occur at the proximal end, i. e. the end to which the spindle-fibers attach, and therefore the roughened tips of the autosomes in figure 99 probably correspond to the polar granules of these elements. (4) The accessory chromosome and the polar granules have the common property of remaining condensed while the rest of the chromatin is diffuse, as well as the common property exliibited in these two figures (99 and 100, Plate 9). The suggestion therefore offers itself that there may be some common physical or chemical properties underlying the corre- spondence in behavior between the accessory and the polar granules. 3. Chromosovw-pair C. — The drawings of chromosome-pair B in figure 64 (Plate 6) and those of C in figure 65 were made from sections cut from the same testis. An examination of the spermatogonial telophases of tliis individual revealed the larger members of each of these pairs very well defined, as indicated in figures 101-105 (Plate 9). No attempt was made to recognize the smaller members of these pairs, because they lacked characteristics, other than size, distinctive enough to make recognition certain. With the larger members of these pairs, however, the distinguishing features are so pronounced that I think there can be no doubt about the identification. I did not attempt to follow these elements through the preleptotene and leptotene stages, but I have no doubt that careful enough study WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 85 would enable one to trace them, as was done in the case of chromo- some-pair A. It is a matter of no small importance, I believe, that each of the "selected" chromosome-pairs has been recognizable by means of one or both its members, in the spermatogonia as well as in the spermatocytes. On the other hand, when I came to search through the postspireme stages of the other individuals for tetrad C, I was able to find the condition shown in figure 65 in only two instances; but a careful study of these stages in the rejnaining animals of the series revealed, in place of the large unequal type shown in figure 65, two other types, which are shown in figure 107, c~m. Figure 106 presents an example of tetrad B from each of the thirteen animals from which material was available for study, and figure 107 a similar series of tetrad C. The corresponding letters, a, h, c, etc., in the two series represent the same animal. We may therefore, speak of the different animals as a, b, c, etc. Cliromosome-pairs B and C are the smallest in the whole com- plex and it will be seen from these two series of drawings that, except in a and b, the pair C is the smaller of the two. In a and b, C is slightly larger than B, as was determined by numerous comparisons in the metaphases of the first spermatocytes. The difference in quantity of chromatin in these two cases is quite small, however, and differences in shape and 'behavior were largely depended on for identification. For convenience in description, we may designate the three types of chromosome-pair C as Ci, C2, and C3. By Ci will be indicated the type, previously described, which is represented in figure 65, and at a and b in figure 107. The type shown in figure 107, c-h, may be designated Co, and that shown in figure 107 at i-vi, as C3. Thus it will be seen that (with a possible exception yet to be discussed) of the thirteen animals studied, two exhibited the type Ci, six the type C2, and the remainder, five, the type C3. If now we compare types Ci and C2, it will be apparent at once that both members of the pair C2 resemble the smaller member of Ci. The homology is striking if one notices the polar granules and the pair of granules close to them, both of which appear in about the same relative size and position in all the examples of both types (except h). It is therefore not difficult to believe that type Co does actually repre- sent a pair of chromosomes homologous to the smaller conjugant in type Ci. Turning to type C3, as shown in figure 107, i-m, it will be observed that this is quite different from either Ci or C2. It represents an unequal pair but the larger member is very different from the larger 86 bulletin: museum of comparative zoology. one in type Ci. Furthermore the prominent chromomere near the polar granules does not seem to be present, except possibly at vi (fig. 107). On the other hand the smaller eonjugant resembles those in 0% in size and otherwise except for the prominent granule already men- tioned. We might therefore be led to suppose that the smaller com- ponent in Cs is homologous to the smaller one in Ci and the two small ones in Ci. But if the example at h (fig. 107) be regarded, it will be seen that this is a small pair lacking any prominent cliromomere near the polar granules, and might therefore be thought to be homologous with the smaller eonjugant in type Cz, if it be considered different from those in type C2. However, even if the somewhat questionable posi- tion of example h, be disregarded as to homologies, it still must be admitted that we have at least three different types of chromosomes appearing in these examples of tetrad C. I may again point out that there is no chance of making a mistake as to the identity of these elements, for the chromosome-pairs B and C are the smallest pairs in the complex, and the different types of C are mutually exclusive, that is, no two of them are ever found in the same animal. I might further add that all the drawings were carefully outlined with a camera lucida and the details filled in so as to represent as accurately as possible the actual conditions as seen in the microscope. The matter of the possible recombination and redistribution of these different types is discussed on page 121. h. The Accessory Chromosome. The accessory chromosome has not been made an object of special study here. Since it has been so thoroughly and so frequently de- scribed for orthopteran material, it will suffice to give only a brief account of it in this connection. In the first place, it should be stated that the accessory can be recognized as a distinct chromatic individual at practically every stage from the primary spermatogonia to the spermatid. The fact that it forms a large and faintly staining vesicle or "sac" in all the spermagonia except the last, probably accounts for the occasional statement that it can be first recognized in the telo- phase of the last spermatogonia, where it appears as a condensed mass of chromatin, or a chromatin nucleolus. Two points deserve to be emphasized: — (1) The accessory, more than the other chromosomes, maintains an exclusive individuality in nearly all stages. However, it sometimes does become associated WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 87 with other chromosomes, especially in the growth-period. Here its polar granule may unite with those of the other chromosomes to form a composite granule. (2) Its behavior, while unique in many respects, differs from that of the autosomes in the degree and the chronology, rather than in the kind, of its changes. The autosomes form vesicles in the telophase of the spermatogonia, as Sutton ('00) long ago pointed out, just as does the accessory, but they are not quite so large or persistent as with the latter. In the growth-period the accessory forms a looped spireme, just as the autosomes do (see fig. 71 and 72, Plate 6), but its thread is much more dense and heavily stained than the others. Although it fails to find a mate in synapsis, its behavior is very like that of the autosomes and its spireme loop may occupy the entire circumference of the nucleus. The process of shortening and thickening, which all the chromosomes undergo, occurs very early in the case of the accessory and it passes through most of the growth- period as a rather compact mass of chromatin. In the postspireme stages, at the time when the chromatids separate from each other by the formation of the secondary longitudinal split, the accessory forms a more or less bent or twisted rod, which often shows a longitudinal split. This split must be homologous to the secondary split seen in the autosomes, which divides longitudinally each of the chromosomes united in synapsis. In the anaphase of the first spermatocyte division its halves separate at the distal end, so that it forms a dyad similar in all respects to those of the autosomes, except for its more roughened condition. In fact, the accessory dyad cannot always be distinguished from the others in the late anaphase. In the metaphase of the second- ary spermatocytes it divides along with the autosomes and usually is indistinguishable from them. Its behavior may therefore be more nearly parallel to that of the whole series of chromosomes than we are sometimes led to suppose. c. The Spermatogonial Divisions. Let us now consider the subject of persistent chromosomal organiza- tion from the standpoint of the spermatogonial divisions. Figures 1-20 (Plates 1 and 2) are intended to represent the most important stages included in the cycle of changes from one cell division to the next. In this description no reference will be made to the selected chromosomes, but the general behavior of the chromatin material will be considered. We shall also leave out of account the mechanics 88 bulletin: museum of comparative zoology. of the division process and concern ourselves chiefly with the fate of the chromosomes after their division and separation has been accom- phshed. In my account of the accessory chromosome, I have already men- tioned the formation of sacs or vesicles in the telophases of the sperma- togonia. In an early telophase, such as is shown in figure 5 (Plate 1), the chromosomes are clumped together in a rather compact mass at the pole of the spindle. But the distal tips of the larger chromosomes may be seen projecting in various directions. Following the clumped condition, stages occur during which the chromosomes begin to ex- pand and to separate from one another. iVt the same time there is developed about each chromosome a hyaline area, at first small in extent, but gradually enlarging as the chromosomes continue to ex- pand. These conditions are shown in figures 6-9. Figure 6 is a side view and figure 7 a transverse (optical) section of the same stage. Figures 8 and 9 are likewise side view and optical section, respectively, of a later stage. At this later stage it will be seen that a membrane has been formed at the boundary between the hyaline area and the cytoplasm. We are therefore probably dealing with sacs or vesicles similar to those described by Sutton ('00) for Brachystola. What is the origin of these sacs? Does the hyaline region as it first appears represent material from the cytoplasm, or from the chro- mosomes, or is it an artifact resulting from the contraction of the chromatin under the influence of the fixative? That it is not an arti- fact, will be apparent, I believe, from the following considerations: — (1) The chromosomes themselves, at the stages shown in figures 6 and 7, are larger than in the earlier stages represented in figures 3 and 4. (2) The chromosomes continue to expand and the vesicles expand still more rapidly, as will be seen from the later stages (fig. 8 and 9). (3) The hyaline region as seen in figures 6 and 7 appears more highly refractive than the cytoplasm which would not be the case if it were a space produced by shrinkage of the chromatin. A comparison of the conditions shown in figures G and 7 with those shown in figures 10 and 12 will, I believe, show that the expansion of the vesicles has been at the expense of the cytoplasm. The relative volume of the space within the vesicle as compared with the volume of the cytoplasm, is much less in these earlier stages (fig. 6 and 7) than in the later stages (fig. 10 and 11). Further, it will be seen that the expansion of the vesicles is accompanied by: — (1) an increase in the size of the cell-body, (2) a diffusion of the chromatin into a kind of reticulum within the space of each sac, (3) the breaking down of the WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 89 vesicular membranes between adjacent vesicles within the group, especially at the polar end, (4) the formation of an irregular nuclear membrane from the outer walls of the vesicles, (5) the apparent anastomosing of the edges of the networks arising from the diffusion of the chromosomes in adjacent vesicles. The walls bounding the original vesicles are still to be seen in figures 8-13, and this is particu- larly true of the accessory chromosome, the vesicle of which persists till a late prophase. What can we now say as to the continuity of the individual chromo- somes? Let us first follow the changes undergone by the accessory chromosome. Figures 1 and 2 are of metaphases, in which all the chromosomes except the accessory are compact and smooth in outline. This is roughened in outline and seems to have already begun the process of expansion which characterizes its behavior immediately after division. In figure 4 a hyaline area of considerable extent has already been formed about the accessory, and close examination reveals also a narrow hyaline area just beginning to develop around each of the autosomes. By the time the stage shown in figure 6 is reached, the substance of the accessory has become distributed through the entire space of the vesicle which accompanied the formation of the hyaline area. In its distribution within the vesicle, the chromatic substance is more concentrated on the periphery of the sac, than through the central space. The vesicle continues to expand along with the expansion of the nuclear material as a whole, until the stage of greatest diffusion of the autosomes has been reached (fig. 12 and 13). At the stage shown in figures 14a and 14b (Plate 2) the chromatin has begun to concentrate towards the axes of the sacs, but this process seems to be less advanced in the accessory {X, fig. 14b) than in the autosomes. These are the earliest of the prophases. In the later prophases, as shown in figures 17 and 20, the accessory becomes concentrated as a coiled thread running down through the middle of the vesicular space. The wall of the vesicle persists longer than does that of the nucleus as a whole or that of the other autosomes (fig. 20). There can be no question, it seems to me, that the accessory main- tains a persistent individuality through all these stages. If now the changes undergone by the autosomes be followed, we shall find for them also evidences of persistent individuality. I think no one would deny a persistent individuality up to the stages shown in figures 8 and 9 (Plate 1) . In these figures the chromatin has become reticular, but the masses representing individual cliromosomes are still quite distinct and surrounded for the most part by the persisting 90 bulletin: museum of comparative, zoology. walls of the vesicles. The method of formation for these vesicles parallels very closely that described for the accessory, the chief difference being that in the case of the accessory the process is much more rapid. In figures 10 and 11 we find the chromatin much diffused and occupying most of the space within the original vesicles. The vesicular walls are no longer visible, however, except on the periphery as an undulating nuclear membrane, and around the accessory. In spite of this fact, the chromatic masses or blocs, each of which has arisen from a single chromosome, are still recognizable as distinct from one another. This is especially well shown in the optical section drawn in figure 11. There are only eighteen masses shown in this section, but the apparent reduction in number need cause no appre- hension as to the fate of the other members of the complex. It fre- quently happens that the chromosomal vesicles do not all lie parallel to each other, so as to be represented in a single transverse section, and some may even assume a position at right angles to the axis of the majority. Such a case is shown in the upper left-hand corner of figure 10. If, now, we examine the stages shown in figures 12 and 13, which are of the period of greatest diffusion that I have been able to find, we may still see, both in optical section (fig. 13) and in side view (fig. 12), the positions of the individual chromosomes represented by a more condensed band or core. In the case of the optical section, nearly the complete number of chromosomes, as represented by these denser masses or cores, can be counted. It is true that there seems to be an anastomosing system of fibrils connecting the adjacent masses, but this need not mean that there has been a loss of chromosome-iden- tity in a common nuclear mass. An early prophase is represented by figures 14a and 14b (Plate 2), which show the two sections of a single cell. We see at this stage the beginning of the process of chromatin concentration which results, finally, in the formation of the condensed chromosomes ready for the next division. The chromatic material of each chromosome first con- centrates near the middle of the region that it occupied in the nucleus in the diffuse condition. There is thus formed a loosely reticulated core (fig. 14a and 14b), out of which there develops a spirally coiled thread, as shown in figures 15a and 15b. The two stages represented in figures 14 and 15 are very close together in time, for they occurred side-by- side in the same cyst. These coiled threads are at first rather small in diameter, but they rapidly thicken and shorten, as indicated in figures 16-20 (Plate 2). During the process of shortening and thick- ening the outlines of the vesicular walls become more distinct. This is WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 91 especially true of the distal pole, as shown in figure 18. It would seem, therefore, that the vesicular membranes first became formed, then largely disappeared, and later reappeared in part. I am inclined to believe that they actually persist to a greater extent than is apparent. There cannot be any doubt, however, that the vesicles do coalesce at the polar end of the nucleus, for there the individual polar granules frequently fuse to form composite granules, such as may be seen in figure 12 (Plate 1) and figures 14, 15, 16, and 19 (Plate 2). The first indication of the longitudinal split which forecasts the next mitosis was discernible at a stage such as is shown in figure 17. From this stage on to the metaphase, however, the split was clearly visible. I believe that the evidence here presented furnishes very good grounds for believing that the chromosomes do not lose their individ- uality in passing through the so-called 'rest-stage' between the two successive cell-di\'isions. d. The somatic Nuclei. Only slight attention has been given to somatic cells in connection with the subject of the individuality of chromosomes, but some points were noted which it seems worth while to record. The connective- tissue nuclei within the follicle always divide by the indirect or mi- totic method. The details are similar to those just described for the spermatogonia, except that individual chromosome-vesicles, even for the accessory, are less conspicuous — in fact, in my limited study of these cells I have not recognized the accessory chromosome with certainty. The only evidence of amitosis is a lobulated condition of the resting nuclei ; that condition is a very characteristic one, but has no more significance as to amitosis than the lobulated appearance of the spermatogonia! nuclei. In the diffused chromatin-stages — telo- phase, rest-stage, and early prophases — the polar granules appear, coalesce more or less to form composite granules, and separate out again just as they do in the spermatogonia. Furthermore, it is possi- ble to find chromosomes in the telophases that exhibit all the chief characteristics of the "selected" chromosomes. For example, in Plate 9, B, figures 108, 109 and 110, are to be seen diffusing chromo- somes with the characteristic features of one of the larger members of chromosome-pair B. It would seem from this evidence that the same morphological constitution of individual chromosomes persisted even in these somatic cells. 92 bulletin: museum of comparative zoology. Going outside the follicle, it is of interest to note what appears in the nuclei of the follicular investment. This investment is a thin membrane inclosing the follicle, forming the outer of the two laj^ers composing the follicular wall. In this membrane the nuclei are very much flattened, so that the chromosomes lie nearly all in one plane. Figures 111 and 112 indicate the chromatic conditions in two such nuclei. It is, I believe, a significant fact that the chromatic masses to be found in these nuclei are in number approximately equal to the unreduced number of chromosomes found in the spermatogonia. Exceptions, it is true, occur; adjacent cliromosome-masses may be- come intunately associated, or one individual mass may become divided into partially separated masses. These nuclei are fully differentiated and are destined never to undergo another cell-division. They must gradually lose their functions and will finally " die in their tracks." The different conditions of the chromatin in the different nuclei suggests that the process of senescent degeneration may have already set in. The important fact still remains, that the individual chromosomes have a tendency to remain distinct from each other, even in these highly differentiated nuclei in a period not only of 'rest' but perhaps of senescence. D. Summary of Observations. 1 . The general topographical relations of the different generations of male sexual cells in the testes of Phrynotettix magnus are typical for the Acrididae. 2. For purposes of accurately following the history of the changes undergone by the chromosomes from the pachytene stages of the first spermatocyte to the time of mitosis, three individual chromosome- pairs were selected, each of which possessed characteristics by which it could be recognized in all the stages concerned. These three pairs were designated, for convenience, "A," " B," and "C." A study of these three chromosome-pairs showed : — (a) that there is a longi- tudinal split in the pachytene stages, which persists into the tetrad and later stages (this is called the primary longitudinal split); (b) that a tetrad is formed out of a spireme segment by (1) a separation along the primary split, and (2) the appearance of a secondary longi- tudinal split along the middle of each of the two parts separated by the primary split. 3. Tetrad " ^ " opens out along the plane of the secondary split. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 93 the proximal ends separating and moving about 90 degrees apart, so that a rod-shaped element is formed the middle of which represents the distal end of the original segment. The rod, thus extended, be- comes oriented with its long axis parallel to that of the spindle and it separates in the middle, thus bringing about an equational division. Tetrad "A" also forms rings, but these were not traced into the meta- phase, and their later behavior is not known. 4. Tetrad " B" occurs in one or the other of two forms: either (1) as an equal pair (in two of the thirteen animals), or (2) as an unequal pair (in the other eleven animals) . The unequal pair differs from the equal in the absence of a large terminal granule at the distal end of one of its members. Both types show the same behavior, opening out at both ends of the segment so that a cross is formed. The separation along the plane of the primary split is the greater and occurs at the distal end; but the cross becomes so oriented on the spindle that the short arms (i. e. the proximal end of the original segment) are attached to the spindle-fibers. Separation in metaphase is therefore along the plane of the secondary split, thus constituting an equational division. 5. Tetrad "C" occurs in three forms, designated Ci, C2 and C3. Ci is composed of very unequal elements, the larger of which possesses a relatively very large terminal knob or granule that is not present on the other. C2 is a pair with equal members each of which appear to be homologous to the smaller member of Ci. C3 is a pair of unequal elements neither member of which appears to be exactly homologous to the components of Ci and C2. The smaller member resembles those of Co and may be homologous to them. The larger member is midway in size between the two members of Ci. Co and C3 divide equationally in the first maturation mitosis, but Ci divides half the time equationally and half the time reductionally in this first division. When dividing reductionally the two unequal dyads follow the law of chance in their distribution with reference to the accessory chromo- some, which passes to one pole undivided. 6. Study of the early growth-stages of the first spermatocyte shows that each of the chromosomes of the telophase of the last sperma- togonial division forms a long spirally coiled thread, which uncoils and stretches out to form the leptotene threads of the primary sperma- tocyte. The leptotene threads conjugate side-by-side (parasynapsis) to form the double threads of the pachytene stage. 7. It was possible to recognize the chromosome-pair A in the spermatogonia as two separate chromosomes (telophases) and to 94 bulletin: museum of comparative zoology. trace the pair through all the stages from the spermatogonia to the spermatids, thus constituting a demonstration of a case of continuous identity, or individuality, through these stages. It was also possible to recognize chromosome-pairs B and C in the spermatogonial telo- phases as well as in the second spermatocytes and spermatids. 8. In the earlier pachytene stages, chromosome-pair B was found to have a definite arrangement of granules or chromomeres, and it was shown that the relative sizes and positions of these chromomeres re- mained constant for similar stages, not only in different cells of a single individual, but also in all the thirteen animals. 9. The spermatogonial divisions showed that each chromosome forms a sac or vesicle in the earlier telophases, and that it expands and becomes diffused within these vesicles; that, although the vesicles appear to coalesce, there is always a remnant of each chromosome visible in the center of the region occupied by the vesicle, and that in the prophase the chromatin concentrates about this remnant or core and there forms a spirally coiled thread, which develops into a prophase chromosome. 10. Study of somatic cells showed:— (1) that chromosome B could be recognized in the connective-tissue cells within the follicle, and (2) that cells of the follicular envelope, which are probably in a state of senescence, still preserved the normal number (23) of chromatic masses. 11. The polar granules are constant features of the organization of the individual chromosomes, as was shown by Pinney ('08); but in some cases (chromosome-pairs A and B) they may become modified to give rise to expansions which resemble the "vesicles" described by Carothers ('13), as well as the "plasmosomes" of most authors. The polar granules tend to unite into composite granules at all of the diffuse stages of chromatic evolution. 12. The accessory chromosome behaves in the manner that is typical for the Acrididae. It forms a large separate sac or vesicle in the earlier spermatogonial generations and a peripheral compact mass in the telophase of the last spermatogonial division. During the leptotene and zygotene stages it may unravel into a long loop, which in some cases is equal in length to a great circle of the nucleus. In the pachytene stages it reassumes a compact form, but may b 5 attached by its polar granule to the polar granules of other cln-omosomes and thus become attached to a composite granule. It passes to one pole undivided in the first maturation division but divides in the second. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 95 III. DISCUSSION. A. Synapsis and the Maturation Divisions. It is very difficult to separate the subjects of synapsis and the matura- tion divisions from the subject of chromosome-individuaHty. Yet for the sake of clearness it seems best to make such an artificial separa- tion. It might also be possible to separate from each other the sub- jects of synapsis and maturation divisions, but since the two are so intimately related, it seems better to discuss them at the same time. Anything like a complete review of the literature on the subjects of synapsis and reduction divisions will not be attempted here, in view of the extensive general reviews in the monographs of Gregoire ('05, '10) and Vejdovsky ('11-12), and the reviews relating particularly to orthopteran spermatogenesis by Davis ('08) and McClung ('14). a. Results from Orthoptera. McClung ('14) has so recently reviewed the literature on Orthop- tera dealing with this subject that it will suffice here to summarize briefly the results. The different views may be classified as follows : — I. Synapsis not considered. a. Both maturation divisions reductional. 1. Wilcox ('94, '96, '97, '01), Caloptenus. b. Both maturations equational. 1. De Sinety ('01), various Orthoptera. 2. Granata ('10), Pamphagus. c. First division transverse. 1. Vom Rath ('92, '95), Gryllotalpa. 2. Farmer and Moore ('05), Periplaneta. 3. Jordan (*08), Aplopus. II. Synapsis described or assumed. A. Telosynapsis described or assumed. a. First maturation division reductional. 1. Montgomery ('05), Syrbula. 2. Stevens ('05), Blatta. 3. Wassilieff ('07), Blatta. 4. Zweiger ('06), Forficula. 96 bulletin: museum of comparative zoology. 5. Davis ('08), Acrididae and Locustidae. 6. Buchner ('09), Gryllus, Oedipoda. 7. Stevens ('10b), Forfieula. 8. Brunelli ('09, '10), Gryllus, Tryxalis. b. Second maturation division reductional. 1. Sutton ('02, '03), Brachystola. 2. Baumgartner ('04), Gryllus. 3. McClung ('05, '08a, '14), various Orthoptera. 4. Stevens ('05), Stenopalmatus. 5. Nowlin ('08), Melanoplus. 6. Finney ('08), Phrynotettix. 7. Robertson ('08) Syrbula. 8. Carothers ('13), Acrididae. B. Parasynapsis assumed or described. a. First maturation division reductional. 1. Gerard ('09), Stenobothrus. 2. Morse ('09), Blattidae. 3. Stevens ('12a), Ceuthophilus. 4. Robertson ('15), Tettigidae. b. Both divisions equational. 1. Vejdovsky ('11-12), Locustidae. c. Division neither reductional nor equational. 1. Otte -('07), Locusta. This classification ^ is interesting from two points of view. In the first place, it indicates the diverse results that have been obtained by the various investigators working on a limited group within which one might reasonably expect to find a high degree of uniformity in chromo- somal behavior. In the second place, the results that I have obtained do not come under any of the classes in the above outline. As stated on previous pages, I have shown (1) that the spermatogonial chromo- somes develop into the fine leptotene threads, which conjugate by paras;>aiapsis without the conjugants losing their identity, that is, the line of conjugation is visible tlu-oughout the growth-period as the 'primary longitudinal split'; (2) that a second longitudinal split at right angles to the first occurs in the early postspireme stages; and (3) that the tetrads become so oriented on the first maturation spindle that the resulting division is equational. Each of the dyads of the second spermatocj^tes consists of parts of the two original conjugants, ' I have omitted reference to some papers which were non-committed on the points under discussion. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 97 and these conjugant-halves become separated in the second matura- tion mitosis, the result being, therefore, a reductional division. I am thus able to support the careful studies of McClung and his students as to the orientation of the tetrads in the first maturation spindle, where the spindle-fibers become attached at the so-called 'synaptic' or proximal ends, and therefore bring about an equational division. I can likewise support the findings of those investigators who describe parasynapsis. If we accept the view that one of the longitudinal splits is in reality the line of separation between parallel conjugants, we can also accept the observations of De Sinety ('01) as to the existence of two longitudinal divisions. McClung and those of his students who have worked on orthopteran material have derived their results from studies confined largely to spermatogonia and the postspireme stages. There is nothing in any of their figures, however, which would be incompatible with parasynap- sis. And the figures by Sutton ('02, fig. 5a, 5b, 6 and 7) of early postspireme stages in Brachystola are much more satisfactorily inter- preted from the standpoint of a preexisting parasynapsis than from the standpoint of telosynapsis. I may also state that I have recently examined some Brachystola material and am well satisfied that the conditions there are quite comparable to those prevailing in Phry- notettix. McClung in his latest paper ('14) accepts the possibility of parasynapsis, and Robertson, who in 1908 argued foi- telosynapsis in Syrbula in no uncertain terms, has recently found parasynapsis in the Tettigidae (Robertson, '15). A glance at the outline of the results of orthopteran studies given above reveals the fact that parasynapsis has relatively few adherents. I believe the failure to recognize this important stage has been due (1) to the general unfavorableness of these synapsis, or lepto-zygotene, stages foi" the elucidation of the conditions and a consequent failure properly to interpret them, or (2) to attention having been largely confined to the postspireme stages. That a study of the latter stages could allow of quite diverse interpretations, I am keenly aware, for it was not till I undertook to follow the history of individual chromo- somes that I was able to arrive at any satisfactory conclusion as to the sequence of events. I am confident that the use of the same method on other material will reveal conditions similar to those that I have described for Phrynotettix. Where the chromosomes differ among themselves as to shape, as they do in Stenobothrus, another source of confusion is encountered, for very few authors have recognized the fact that cliromosomes of 98 bulletin: museum of comparative zoology. different shape may behave differently in their orientation on the maturation spindles. McClung has recently gone over this matter in a very painstaking way, and I can agree with his conclusion that, in general, the chromosomes with the spindle-fiber attachment termi- nal, that is, rod-shaped chromosomes, are oriented in the first matura- tion spindle so as to produce an equational division, while those wliich have the spindle-fiber attachment non-terminal, that is, at the apex of V-shaped cln-omosomes, become oriented so as to bring about a reduction at the first division. This general rule is of course violated when the pairs of rods are of unequal length, which usually (Baum- gartner, '11; Payne, '12; Carothers, '13; Robertson, '15), but not always (Ci, described in this paper), divide reductionally in the first division. Davis ('08) sought to establish the behavior of the V- shaped chromosomes of Stenobothrus as the type for the Orthoptera in general. He correctly described the behavior of these chromosomes in the maturations, but fell into error by attempting to make the rod- shaped chromosomes conform to the same type of behavior. He also failed to recognize parasynapsis. I have recently made a study of the conditions in Stenobotlirus and may say that I found parasynapsis for both forms of cln-omosomes, and that the V-shaped chromosomes divide reductionally in the first maturation mitosis, as Davis de- scribed, but that the rod-shaped chromosomes divide equationally in the first division, as I found that they did in Phrynotettix. > b. Recent Work on Synapsis. That parasynapsis has a wide occurrence, is evident from a glance at the cytological literature, especially within recent years. Gregoire in his two admirable monographs ('05, '10) has reviewed most of the previous literature bearing on the subject of the behavior of the chromosomes in maturation, and has endeavored to find a common type of behavior for both plants and animals. He says ('10, p. 384) : "Dans un bon nombre d'objets animaux et vegetaux, les cineses de maturation s'accomplissent suivant le type d'une prereduction hetero- homeoty pique preparee par une pseudo-reduction prophasique par parasyndese ou zygotenie." In this " heterohomeotypique " scheme, however, Gregoire has failed to distinguish the difference in behavior between the chromosomes with terminal and those with non-terminal spindle-fiber attachment. Since the publication of Gregoire's later monograph, a considerable number of investigators have reported the existence of parasynapsis. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 99 De Saedeleer ('13) finds in Ascaris all the typical stages of the growth-period: — leptotene, zygotene, pachytene, and diplotene; he consequently believes that parasynapsis occurs. Among the Crustacea parasynapsis has been found by McClendon and by Kornhauser for Copepoda and by Fasten for Canibarus. McClendon ('10) found parasynapsis in both the oogenesis and the spermatogenesis of Pandarus sinuatus, but could not decide which of the maturation divisions were reductional. Kornhauser ('15) gives a very full account of a careful study of the process of parasynapsis in Hersilia apodiformis, thus confirming the earlier results, as to the existence of parasynapsis in Copepoda, of Lerat ('05), Matschek ('09), and McClendon ('10). In this paper he clears up the uncertainty in regard to this group brought about by the unique theories held by Hacker ('92) and his followers. Kornhauser demonstrates very clearly that the so-called ' Querkerbe,' which Hacker and his followers interpreted as the point of end-to-end union, is nothing more than the synaptic point of the chromosomes which have a median or non- terminal spindle-fiber attachment. The Copepoda are thus brought into line with the majority of other forms. Fasten ('14) finds para- synapsis in Cambarus, and although he is dealing with a very large number of clu-omosomes (the diploid number is about 200), his figures of the leptotene and zygotene stages are quite convincing. With respect to work on insect material, I have already mentioned that on Orthoptera. The results of Stevens are unusual in that she has described telosynapsis for Blatta ('05), Stenopalmatus ('05), and Forficula ('10b), while in Ceuthophilus ('12a) she found parasynapsis. In the last mentioned article she says (p. 227) " I should not be sur- prised if the range of variation should prove to extend from (a) cases where there is nothing that could be called conjugation, but merely such a pairing without contact even, as will secure segregation of homologous maternal and paternal chromosomes to different daughter cells, through (b) an intermediate condition of telosynapsis and less intimate paras;yTiapsis, to (c) cases where homologous chromosomes are so completely fused in parasynapsis that it is impossible to tell whether the resulting cliromosomes which are segregated in mitosis are identical with those that went into synapsis or not." It may be that more intensive studies will reveal greater uniformity of behavior than Stevens advocated. Payne ('14), in a brief description of tetrad formation in Forficula sp., reaches only tentative explanations and conclusions. He finds a variable number of chromosomes in the two maturation divisions and suggests that this might be accounted for b;^- supposing that some of 100 bulletin: museum of comparative zoology. the spermatogonial chromosomes had failed to pair. He describes two methods of ring-formation. The correctness of his conclusions as to the succession of stages in some of his series might be questioned on the ground that they are not different stages of the same chromo- some. The series shown in his figures 2 to 11 probably represents a normal method of ring-formation, viz., by the opening out of a para- synaptic spireme segment along one of the longitudinal splits, with the ends remaining in contact. The series in figures 12 to 16 might also easily be derived from a parasynaptic segment. It is extremely questionable whether the figures in the series 18-20 are arranged by the author in their natural sequence; the reverse order is more likely to be the correct one. His figures 19 to 27 (Plate 2) doubtless repre- sent different shapes of the same chromosome-pair, the so-called "middle granule" serving to identify the element. I would suggest, however, that the stage that he represents in figure 19 may have resulted from an opening out of a parasynaptic segment in the same way that I have described for chromosome-pair A, in which case the "middle granules" would be polar granules instead of "middle" ones. In view of the rather far-reaching conclusions that Robertson ('15) has drawn from his work on the Tettigidae, I would call attention to some differences, as well as similarities, between his work and mine. In the first place, he describes parasynapsis in the early stages of the growth-period, as I have done, but in the postspireme stages he as- sumes that the conjugants separate along the plane of conjugation (primary longitudinal split), the separation beginning at the proximal end. He shows in the metaphase of the first spermatocyte most of the chromosomes as elongated rods with appearance and orientation similar to that seen for my tetrad A. It will be remembered that in the latter case the separation from the proximal end of the spireme segment toward the distal end is not along the plane of the primary split, but along the plane of the secondary split. I believe that Robertson may have overlooked a similar behavior in the chromo- somes of his material. Curiously enough the unequal elements that he describes are very similar to the unequal type of chromosome B and of chromosome C, in PhrynotettLx. I have shown that the behavior of the cliromatids in B and C is similar to that in A. And that in the cases in which C divides reductionally in the first division the spindle-fiber attachment is necessarily shifted to the distal ends. I believe that such a condition probably occurs in the unequal tetrads described by Robertson, and if so, theoretical explanations, as to how the elements came to be related to each other in the way they are. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 101 would be unnecessary. Robertson says he believes that the unequal tetrad in Tettigidea -parvipennis has arisen by a loss at the distal end. And he finds in other individuals an homologous pair each member of which is equivalent to the larger member of the unequal pair. This is just the condition that is presented by B in my material. And, furthermore, by analysis of the pachytene and postspireme stages, I am able to say just what part has been lost. Another striking analogy between my observations and those of Robertson occurs in connection with the larger unequal pair that he has found in Acridium granulatus. He finds in some individuals a small equal pair and in two individuals an homologous unequal pair, the smaller component of which corresponds to either of the two elements of the equal pair. This, again, is precisely the relationship between types Ci and C2 in Phrynotettix. Here, too, we both failed to find the other possible combination, namely, that of a pair of the larger conjugants. These striking similarities lead me to think that the elements described by Robertson may be explained in the same way that I have explained them in Phrynotettix. If such be the case, then the various assumptions as to doubling and "sesquiva- lent" chromosomes will be unnecessary. I believe, further, that without a doubt the unequal tetrads described by Robertson do divide reductionallv in the first maturation division, but that the spindle-fiber attaches at the distal and not at the proximal end; and, furthermore, that there is a very good chance that the other chromo- somes may behave as does chromosome-pair A in Plirynotettix, and therefore divide equationally, as it does. Of recent works on Hemiptera, the most interesting from the stand- point of synapsis are the papers by Montgomery, Wilson, and Korn- hauser. Montgomery ('11) advocated telosynapsis for many years, but in this late paper, in which he described the spermatogenesis of Euschistus, he concludes that pairing is by a process of parasynapsis. I believe it to be a highly significant fact that Montgomery, at the end of his very active career as a cytologist, and with his wide ex- perience back of him, should reverse his former position on the subject of synapsis and should find parasynapsis in this insect, which he had studied and reported on at an earlier date ('01). He says (p. 743): " In the growth period through the pachytene stage there is no longi- tudinal splitting, for what I had previously ('01) interpreted as such, I now find to be the line of conjugation .... Frequently at certain points along a geminus the chromatin granules appear accurately paired. But this does not appear until a rather advanced stage of the 102 bulletin: museum of comparative zoology. strepsinema and is by no means regular" .... Further on (p. 753) he says : — " During the past year I have also convinced myself of the occurrence of parasyndesis in Plethodon, such as Janssens had de- scribed for this object and the Schreiners for Salamandra." Wilson ('12), in his critical study of the subject, first states the questions that he believes must be answered in connection with synap- sis and then gives his reasons for believing in the wide occurrence of parasynapsis. He regards the following questions as still awaiting a satisfactory answer: — "1. Is synapsis a fact? Do the chromatic elements actually conjugate or otherwise become associated two-by- two? 2. Admitting the fact of synapsis, are the conjugating ele- ments chromosomes, and are they individually identical with those of the last diploid or premeiotic division? 3. Do they conjugate side-by-side (parasynapsis, parasyndesis), or end-to-end (telosynapsis, telosyndesis), or in both ways? 4. Does synapsis lead to a partial or complete fusion of the conjugating elements to form 'zygosomes' or 'mixochromosomes,' or are they subsequently disjoined by a reduction division?" Wilson finds his own material (hemipteran) not altogether favorable for a solution of the problems enumerated, but has been able to study the preparations of Tomopteris, supplied by the Schreiners, and of Batrachoseps supplied by Janssens. He studied also some orthop- teran material, including Phrynotettix, secured from McClung. After a study of Tomopteris and Batrachoseps he says (p. 384): "Through the study of Batrachoseps and Tomopteris I have finally been convinced — for the first time, I must confess, as far as the auto- somes are concerned — (1) that synapsis, or the conjugation of chromosomes two-by-two, is a fact, and (2) that in these animals (perhaps also in the Orthoptera) the conjugation is a side-by-side union, or parasynapsis." And again (p. 399), "The few observations I have been able to make on McClung's preparations of Achurum, Phrynotettix, and Mermiria. . . .lead me to the impression that a side-by-side union of leptotene threads takes place here also." Browne ('13), from a comparative study of the spermatogenesis of three species of Notonecta, regards the evidence, though not abso- lutely conclusive, as indicating a conjugation by parasynapsis. Kornhauser ('14), as a result of a very careful study of the spermato- genesis of two species of Enchenopa, finds conclusive evidence of a parasynaptic union at the beginning of the growth-period. The evidence for parasynapsis in the Hemiptera is thus seen to be very strong. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 103 Of recent work on the Diptera, I may mention that of Stevens and of Taylor on Culex, and that of Metz on Drosophila. Stevens ('10a) finds parasynapsis in Culex, not merely in the growth-period of spermatogenesis, but among other kinds of cells. She says ('10a, p. 208) : — " That parasynapsis occurs immediately after the last oogonial mitosis is certain and it is equally certain that the chromo- somes are similarly paired in earlier generations of the oogonia," Again (p. 209) : — " In Culex it is quite certain that parasynapsis occurs in each cell generation of the germ cells in the telophase," and (p. 212), "It is interesting to find in Culex a clear case of parasynap- sis in oogonia, oocytes, spermatogonia, and spermatocyte prophases and then to see the same chromosome pairs appearing in the first maturation metakinesis as though united end-to-end." (It is prob- able that she has overlooked stages in the postspireme showing the changes undergone by a parasynaptic spireme segment in its trans- formation into a metaphase tetrad). Miss Stevens found six to be the somatic number of chromosomes in Culex, the reduced number being three. The side-by-side pairing of the chromosomes in nearly all generations of cells studied, gave the appearance of a reduced number in many situations where it would not have been suspected. Taylor ('14) states that she found in Culex pipiens only three chromo- somes in all the stages that she studied, i. e. in both the somatic and germ-cells of both sexes. This is a very surprising result, but an explanation may perhaps be found in the conditions observed by Miss Stevens, namely, the tendency for the chromosomes in all kinds of cells to pair between mitoses. A poor fixation might easily prevent one from recognizing the double nature of a closely adhering pair of chromosomes. Besides, Miss Taylor found some cells with six chromosomes, and shows figures of some others with more than three. It would seem more reasonable, then, to regard the prevalence of the reduced number found by Miss Taylor as the result of the constantly recurring tendency of the chromosomes to unite side-by-side between successive mitoses, and possibly to poor fixation. The results of Metz (*14) on Drosophila are interesting in this connection, for he reports conditions in these flies similar to those found in the mosquito. In this he confirms the earlier results of Stevens ('08). He finds (p. 55) that: — "The chromosomes not only exhibit a close association in pairs at nearly all times, but that before every cell division the members of each pair become so intimately united that they may be said actually to conjugate. Each pair, with the possible exception of the sex chromosomes, goes through what 104 bulletin: museum of comparative zoology. amounts to a synapsis in every cell di\'ision, so that in many cases the figures closely resemble the haploid groups. Apparently this takes place especially in early prophase, but a second conjugation may occur dm'ing metaphases, just a short time before division. In the second, or metaphase, conjugation, at least, it is worthy of note that the union is unquestionably a side-by-side or parasynaptic one." Thus we find parasynapsis in a greatly exaggerated form in these examples from the Diptera. Of recent studies on Vertebrata, we may note those of Snook and Long on an amphibian, of Jordan on an opossum, and of Wodsedalek on the pig. Snook and Long ('14) find the same kind of evidence for para- synapsis that has been presented for Batrachoseps by Janssens ('05), for Salamandra by the Schreiners ('07), and by Wilson ('12) as quoted at p. 102. This evidence, together with that announced by Mont- gomery ('11) for Plethodon, forms a series of observations which renders very probable a general occurrence of parasynapsis among amphibians. Jordan ('11) describes in the spermatogenesis of an opossum what he considers evidence for telos;vTiapsis. His figures, however, are far from convincing on this point, since they could as readily be inter- preted in favor of parasynapsis as telosjaiapsis. Wodsedalek ('13), in his studies of the spermatogenesis of the pig, is unable to find conclusive evidence on the subject of synapsis. He says (p. 13), however, that in the synezesis stage, "The thin tlireads become arranged in a very much tangled mass of loops, which later appear in about half the original number and twice as thick." Inas- much as these phenomena accompany every case of demonstrated parasjTiapsis, the evidence seems to favor the occurrence of this mode of conjugation in this case. In conclusion, I think it must be admitted that there is abundant evidence for a widespread occurrence of parasynapsis, especially as shown by the most recent investigations. While a majority of the authors who have worked on orthopteran material have reported telosynapsis, I believe there is some chance that many of them were mistaken, or that a more careful analysis of the critical stages would have given a different result. Whether we accept the hypothesis of Stevens, that all degrees of s\Tiapsis occur, or the idea of Gregoire, that parasynapsis is an almost universal phenomenon, we must at all events admit that the most careful of the recent investigations indi- cate that the latter condition is widespread throughout the animal kingdom. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 105 As to which of the two maturation divisions is equational and which is reductional, no absohite rule can be laid down. The evidence, however, points to the probability that generally chromosomes with terminal spindle-fiber attachment are not separated from each other until the second division, while those that have a non-terminal at- tachment are separated in the first, and that consequently in the former the reduction occurs at the second division, in the latter at the first division. B. Individuality. The theory of individuality was early championed by Van Beneden ('83), Rabl ('85), and Boveri ('88). In more recent years the theory has been supported by many wi'iters, who have accepted as substan- tial evidence in its favor the constancy in the number, size, and shape of the chromosomes reappearing in the mitotic spindle of any one species of animal or plant. On the other hand, some eminent zoologists have attacked the theory on the ground that the individual chromosomes cannot be traced through the so-called "rest" period between mitoses. It will, therefore, be convenient to discuss the two topics: — (a) constancy in metaphase chromosomes, and (b) persistent organization of chromosomes. a. Constajici/ of Metaphase Chromosomes. 1. Constancy in number. — The constancy in number of chromo- somes for any species is among the most commonplace of cytological observations. It will therefore be unnecessary to make any exten- sive references to the literature. Some exceptions to the general rule occur, however, and should receive attention. Supernumerary chromosomes have been reported from time to time, and have been studied especially by Stevens and Wilson. Wilson ('09) found in Metapodius variations in chromosome-number from 21 to 26, though the number for each individual animal was constant. The number of chromosomes was dependent neither on sex, nor locality of habitat, nor w^as it correlated with constant differences of size or of visible structures in the adults. But the variation affects only particular classes of chromosomes (the small idiochromosomes) and all exhibit the same behavior. Furthermore, Wilson found a few cases of mitoses in which both members of a pair of small chromosome were going to 106 bulletin: museum of comparative zoology. the same pole. Using this as a basis, he was able to find satisfactory explanation of the variation in number, and one which served to sup- port the theory of the individuality of the chromosomes. Stevens ('12b) found in Diabrotica supernumerary chromosomes varying in number from 1 to 5, and believed that they had their origin in trans- verse and longitudinal divisions of the X-chromosome, which normally divides only longitudinally. These anomalies can therefore be explained on the basis of some unusual method of distribution of the chromosomes in mitosis; the fact that such extra cliromosomes persist in all the cells of the animal in which they are found is a striking bit of evidence in favor of the belief that they maintain their individuality. Delia Valle ('09, '12) has attacked the theory of individuality, declaring that the chromosomes are temporary and variable structures, which form in the prophase and dissolve in the telophase. He thinks their number is the quotient of the quantity of chromatin divided by the average size of the clu-omosomes. The quantity of clu-omatin is said to vary with conditions of nutrition, and the number of cln-omo- somes with variations in external conditions. He made counts of chromosomes from cells of the peritoneum of salamander larvae and obtained numbers varying from 19 to 27. Montgomery ('10) points to the following grounds for doubting the accuracy of Delia Valle's conclusions : — " 1. The chromosomes counted are long, sinuous ribbons, that overlap and interlace, the most difficult kind to count with accuracy. 2. He included in the counts some cells in prophases, where one cannot be certain that all the clu'omosomes have fully separated. 3. The total number of the chromosomes is so large, about 24, that the chance of error in enumeration is great. It is but fair to conclude that while his technique was excellent, his choice of material was bad, consequently a degree of scepticism might well be maintained toward his results." Delia Valle in his latest paper ('12) argues that the chromosomes are variable structures, because he has been able to find transition stages between mitotic and amitotic methods of cell-division in the erythrocytes of young salamanders. It is a well-known fact that amitosis frequently accompanies degenera- tion, and the figures of Delia Valle present strong indications of being those of degenerating cells. It is precisely in degenerating cells that one would look for inconstancy in the behavior of the chromosomes. 2. Constancy in size and shape. — It will be convenient to consider the subjects of size and shape together. As to shape, we may dis- tinguish spheres, rods, and V-shaped elements. Spheres are invariably small and may be regarded as short rods. It will be convenient to A\ enrich: spermatogenesis of PHRYNOTETTIX MAGNUS. 107 include under the term "V-shaped" all the chromosomes which have a non-terminal spindle-fiber attachment. They may be regarded as rods which have become bent at the point of the spindle-fiber attach- ment. Broadly, therefore, we may look upon all cliromosomes as rod-shaped, but it will make description easier to distinguish the types just mentioned. In attempting to show that chromosomes have a constant size and shape for each species, as well as a constant number, it will be well to call attention to the fact, so clearly stated by McClung ('14), that the point of the spindle-fiber attachment is, as a general rule, constant and therefore one of the indications of a persistent organization for each individual chromosome. Some groups of animals exhibit a high degree of uniformity in the shape and size of the chromosomes in any species, as for example, among the Crustacea and the Amphibia, while others show a great variety of forms (Orthoptera, mammals). In the groups with diverse shapes and sizes of chromosomes, the striking fact was pointed out by Montgomery ('01) that there are two of each different size. Mont- gomery reached the logical conclusion that of the two equivalent series existing in «ach cell, one had been derived from the maternal and the other from the paternal ancestor. That the same series of sizes and shapes reappears in each cell- generation, is recorded by nearly every observer whose material is favorable enough to admit of such comparisons. The work of Mc- Clung ('00, '02, '04, '08b, '14), Sutton ('02, '03), Baumgartner ('04), Nowlin ('08), Pinney ('08), Robertson ('08) on orthopteran material has done much to establish this fundamental feature of individuality. A very interesting series of observations on this point is that of Meek ('12a, '12b) on Stenobothrus. He describes the results of a series of careful measurements of cliromosome-dimensions in diiTerent genera- " tions of cells, and as a result of these observations becomes convinced of the existence of persistent individuality. I may quote some of his conclusions ('12a, p. 24, ff.): — " (1) In all metaphases the relative positions of the chromosomes in the equatorial plate appear to be arbitrary. (2) The rods composing all ordinary chromosomes are cylindrical with rounded ends, and of an unifonn and constant diame- ter, viz., 0.83 micra. In each species eight lengths have been found, and these constitute members of a series in arithmetical progi'ession, of which the difference between consecutive terms is equal to the radius of the rod. The heterotropic chromosome does not belong to this general series, for, although equal in length to the longest rod, its 108 bulletin: museum of comparative zoology. diameter varies at different points and exceeds 0.83 micra. (3) The rods are indivisible units, and, since each spermatogonial and second spermatocyte chromosome is composed of two, and each primary spermatocyte chromosome of four, their morphological identity is metrically proved. (4) The eight rod-lengths are not the same in any two species; the longest and 5 short chromosomes occur in all, but identity is always established by the two remaining chromosome- rods. (5) The complexes of a species and its variety appear to be identical; differences if existing, are too small to be recognized. (6) The somatic chromosomes are identical with those of the germ cells. (7) The total volume of ordinary chromosomes is the same in sperma- togonial and primary spermatocyte metaphases, whereas only half this amount appears in that of the secondary spermatocyte." For some zoologists, the fact that for any species the chromosomes reappear in the different cell-generations possessing the same relations as to number, shape, and size is merely an expression of the activities of the cell and signifies nothing as to a persistent individuality of the chromosomes. Fortunately, we are not dependent on this kind of evidence alone, the results of studies of the chromosomes of hybrids, for example, offering still stronger evidence of individuality, as shown by the work of Moenkliaus on hybrids between Fundulus and Men- idia, and that on echinoderm hybrids by Baltzer and by Tenent. Moenkhaus ('04) could recognize the two sets of chromosomes arising from the pronuclei of the diverse parents by characteristic differences in size. For the first two or three cleavages the two groups tended to remain distinct, but in later cleavages the chromosomes of the two kinds become more and more intermingled, though they are still recognizable by their characteristic sizes. The value of this evidence is obvious, and on this point Moenkhaus says (p. 53) : — "As long as the two kinds remain grouped, as during the first two divisions, this fact has little added significance (i. c, that two groups of distinctly different kinds of chromosomes arise), since within each group it would be perfectly possible for the component chromosomes to ex- change chromatin granules during the resting period. If, however, as occurs in the later cleavages, the two kinds of chromosomes become mingled, the chromatin granules of both kinds must lie mingled to- gether within the resting nucleus. If from such a nucleus the two kinds of chromosomes again emerge, it amounts almost to a demon- stration that the chromatin substance of a given chromosome forms a unit and that unit persists." Baltzer ('09) was able to recognize in hybrids between Echinus and Strongylocentrotus certain chromo- WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 109 somes which were distinctive of the species from which they were derived. Tennent ('08) found in hybrids between Moira and Arbacia a mixture of two kinds of chromosomes, each variety of which could be distinguished. It is, indeed, difficult to understand how these distinctive chromosomes could recur with such definite characteristics in hybrid embryos, if there is no persistent identity for them. Variations from the general rule of chromosomal constancy have been recorded from time to time, for example, in the shapes of tetrad chromosomes. In many species there is a tendency for each of the forms of the tetrads to be reproduced in the first spermatocyte meta- phase. This is particularly true where the chromosomes are all of a similar shape and size. But even in such cases, there is a variation in the exact contour presented by different chromosomes. It has been made apparent by many investigators, especially by McClung and his students, that the shape of a metaphase tetrad is dependent upon the extent and character of the movement on each other of the constituent cliromatids. The work of these authors also shows that homologous chromosomes tend to assume about the same shape in all the cells at corresponding stages of mitosis, but that this condition of similarity has its exceptions. Baumgartner ('04) called attention to the constancy in the number of rings formed among the tetrads of Gryllus, and others have noted similar conditions. However, such a criterion for individuality is not always a safe guide, as was pointed out by Foot and Strobell ('05). Commenting on Baumgartner's paper, they say, in regard to chromosomes in Allolobophora foctida: — " We find no constant form differences of the chromosomes, the simplest form of the bivalent chromosomes is two rods«attached end to end, and these present a variety of shapes, rings, figures 8, crosses, etc., without any regularity or constancy. The free ends of the bivalent chromo- somes show a tendency to unite into a ring and in some cases nearly all the eleven chromosomes are rings, and sometimes not a single ring is formed" (footnote, p. 222). A glance at figures 39 and 40 (Plate 4) of this paper will also show a variability in shape of the eleven bivalent chromosomes. In my account of tetrad A, I have shown that this element may or may not form rings, so that this character could not be used as a criterion for identification in the earlier postspireme stages. But in spite of these exceptions, there does exist in many cases a strong tendency for a chromosome to assume the same shape at similar stages in all the cells of an animal, and the exceptions have no significance in relation to the question of a variation in the funda- mental organization of the tetrads. 110 bulletin: museum of comparative zoology. In the case of unequal tetrads, however, variation in shape does have some meaning with reference to chromosomal organization. In the specimens of Phrynotettix which I have studied, the shape of tetrad B in two individuals is fundamentally different from that in the other eleven, because, in the latter, a definite part of one member of the pair is lacking. Similarly, in the case of Ci and C2, the difference concerns a definite part of the members of the pairs. But the im- portant thing to be kept in mind is that the organization of each of these tetrads is constant for any individual animal, and such differ- ences as exist between individuals can be readily accounted for. b. Persistent Organization of Chromosomes. 1. The selected chromosomes. — One of the most important con- clusions arrived at in the present study relates to the constancy in the finer organization of the chromosomes, both from stage to stage in the same individual, and from one individual to another. This is shown in two ways : — first, by the existence in chromosome-pair B of an architecture that is constant both for any one individual in the various stages in which any architectural condition could be recognized, and likewise for all the individuals studied; secondly, by that of a particular pair of chromosomes (A) recognizable through all the stages from spermatogonia to spermatids, the recognition being made possible by the fact that the chromosomes in question possess properties which are characteristic and constant for all stages. Both of these selected ^chromosomes, A and B, tend to stain more deeply than the other autosomes, but this tendency is much more marked in A than in B. If chromosomes possessing similar peculiari- ties be found in related species, may they not be regarded as homolo- gous to the selected chromosomes A and B of Phrynotettix? I think such homologies could be established. Miss Carothers ('13) shows that the small unequal tetrad in Brachystola is usually associated with the accessory chromosome, and is more intensely stained than the other autosomes. Might it not be possible to analyse this un- equal element in Brachystola and determine its relation to the unequal tetrads of Phrynotettix? Since these two genera are closely related, I believe this would be possible. Furthermore, the other unequal tetrads described by Miss Carothers for Arphia and Dissosteira were among the small chromosomes and, on account of the similarity in behavior, might be found homologous to 5 or C of Phrynotettix. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. Ill Miss Nowlin ('08) describes for Melanoplus bivittatus a precocious tetrad (no. 11), which always appears in the metaphase as a rod extended parallel to the spindle-axis. Such is also the behavior and form of chromosome A. Furthermore, I have examined slides of Melanoplus material and find that it also has a spireme loop that stains more deeply than the others. May not this precocious tetrad of Melanoplus be related to chromosome A of Phrynotettix? Early in the course of my investigation I had the opportunity of looking over some of Dr. McClung's collection of slides of acridian material, and, though a thorough study was not made, I could easily recognize in the pachytene stage of a number of species a spireme loop which stained more deeply than the others. Such loops were found, for example, in species of Aeoloplus, Amphitornis, Arphia, Brachy- stola, Hadrotettix, Hesperotettix, Hippiscus, Melanoplus, Phaetaliotes, and Stenobothrus. One characteristic of such tlireads, which, how- ever, is not so marked in Phrynotettix, is a tendency to become asso- ciated with the accessory chromosome. This is particularly true of Melanoplus and Stenobothrus, the forms in which Davis ('08) was led by this close association to describe a "double monosome." There can be no question, I think, that these "double monosomes" were merely the accessory chromosome plus one of these deeply stained spireme segments. In view of these facts, the suggestion offers itself, that similarity in the properties and behavior of certain chromosomes in different species may be correlated with their taxonomic relation- ships. Such correlation was, indeed, seen and discussed some time ago by McClung ('08a). Meek ('12) has already made a comparative study of the sizes of the chromosomes in several species of Steno- bothrus, and has reached the conclusion that the five smaller pairs of chromosomes are of the same size in all species, but that the large (V-shaped) pairs differ from one species to another. It still remains to be seen whether or not the chromosomes of different species can be compared on the basis of their details of organization and behavior, as well as size. 2. The heterochromosomes. — I believe most observers agree that the heterochromosomes maintain their individuality tlii'ough the growth-stages of the male germ-cell cycle. On another page (p. 87) I have called attention to the similarity in behavior between the autosomes and the accessory, this has also been noted by many others, so that there is no very good ground for setting up a claim to funda- mental distinction between the two kinds. It seems to me, therefore, that if we admit a persistent individuality for the heterochromosomes. 112 bulletin: museum of comparative zoology. we must at the same time admit a high degree of individuahty for the autosomes. 3. PJasmosovies and nucleoli. — One of the most puzzHng problems that cytologists have to deal with is the behavior and function of the so-called ' plasmosomes ' or 'nucleoli.' They apparently exhibit such a variety of reactions to methods of technique, and exhibit such varying relationships to other structures in the cell, that it is almost hopeless even to attempt to classify them. That they play some important role in the physiology of the cell, there is not the slightest doubt, but what that role is, or what relation they bear to the question of chromosome-individuality, are problems that are far from a solu- tion at the present time. In my description of the tetrads A and B, I called attention to a peculiar modification of one of the terminal granules of each. I emphasized the fact that, in the case of tetrad A, this modified granule furnished a means of identification for this element. Just what the nature of this modification is, I cannot state definitely, but in the pachytene stage it has the appearance of an expansion of a previously condensed granule, and I have so treated it in my description. The similar condition in B appears to arise in the same way, but in this case there seems to be a more definite boundary to the modified gran- ule, which thus resembles the plasmosomes, or "vesicles," described by Carothers ('13). The expanded granule of A is usually not homo- geneous, some areas within it appearing more dense than others. This condition probably foreshadows that seen in the postspireme stages, where it appears more like a collection of small granules, typically tlu-ee in number. Miss Carothers described the 'vesicles' that she found as being attached to spireme threads, and in some cases to specific threads. Furthermore, she found that the occurrence of the vesicles extended to several species, and, in some species, through several generations of cells. I am indebted to Dr. McClung and to Miss Carothers for the privilege of looking over some of the material studied by the latter, as well as for the opportunity of studying slides of other species ; I can confirm Miss Carothers's observations, and can add that these so-called ' vesicles ' are present in nearly every species of grasshopper that I have studied with this object in view. I believe that the modified granules in Phrynotettix can be homol- ogised with the 'plasmosomes' of other species. I would especially call attention to the fact that these structures are always attached to chromosomes, and that, in Phrynotettix, at least, they always involve a certain part of the chromosome to which they are attached. I WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 113 believe we may therefore regard them as being related to the organiza- tion of the chromosomes, just as much as the polar granules are. A glance at the literature will show how constantly these structures are found; but no one, except Miss Carothers, so far as I am aware, has suggested that they are always attached to some definite region of a chromosome. It would seem to be worth while for some one to make a study of these structures from this point of view. The changes in staining capacity which the plasmosomes undergo at different stages raises an interesting question as to what may be their relation to the chromatin of the thread with which they are associated. Do they take up chromatin from the chromatin-thread, thus increasing their own stainability, and give it back again as they lose their power to hold the stain? Do they elaborate chromatin from raw materials in the surrounding cell substance and give it up to the chromatin-thread? Is their chromatic substance different from other chromatin? Or, do they have some other way of becoming for a time chromatic and later non-chromatic? May it not be possible to answer some of these questions by carefully resolving into its chromomeres the chromatin-thread with which they are associated, and comparing the constitution at different stages? I believe this could be done on favorable material. In Phrynotettix, these struc- tures are definitely related to polar granules. Are the polar granules to be classed in the same category as the plasmosomes? Is it possible for a polar granule to become transformed into a plasmosome, and then back into a polar granule again? The last question seems to be an- swered in the affirmative by the conditions in Phrynotettix. In the case of B, for example, one of the proximal granules becomes "ex- panded" in only about 16% of the cases counted. In becoming expanded it has become like a plasmosome. When it is not expanded, it remains a polar granule. Is it any wonder, then, that the plasmo- somes have been called 'variable' and 'uncertain' elements of the nucleus? Plasmosomes are associated with heterochromosomes, as well as with autosomes. Davis ('08), for example, noticed one on the mono- some of Stenobothrus, and I have confirmed the observation from slides of my own. Morse ('09) found in cockroaches a plasmosome constantly associated with the "chromatin nucleolus" (accessory), and in addition another body in the cytoplasm, which he called a plasmosome. A similar cytoplasmic body, which stains like chroma- tin, is found in a number of Acrididae. Dederer ('07) found a plasmo- some associated with the pair of idiochromosomes in Philosamia, and 114 bulletin: museum of comparative zoology. Blackman ('05) found a plasmosome attached to the accessory chro- mosome of Scolopendra. A long Hst might be added to show that plasmosomes have been found associated with particular chromosomes. Many attempts, not altogether successful, have been made to explain the baffling relations to the other cell-structures of such bodies as have been called plasmosomes, nucleoli, chromoplasts, karyospheres, etc., but any future attempts to elucidate these relations must, I believe, be accompanied by a recognition of the relations that these structures bear to the organization of individual chromosomes. 4. Persistence of chromosomes between mitoses. It still remains to discuss what may be the nature of the "organization" of the chromo- somes in the stages through which the nuclear substance passes from one metakinesis to the next. I shall consider briefly (1) the origin of the nucleus from the chromosomes, and (2) theories of continuity. (1) Origin of the nucleus. In my description of the spermato- gonial divisions of Phr\Tiotettix (p. 87-91), I pointed out that each chi-omosome becomes surrounded, as early as the anaphase, by a hyaline region, that this region expands in the telophase; that the chromatin of each chromosome becomes diffused to a certain extent within its own region; that a membrane becomes formed at the boundary between the hyaline region and the cytoplasm, producing the chromosomic "vesicle"; and that the nuclear membrane consists of the outer walls of the vesicles at the periphery of the nuclear group. I drew the conclusion that the hyaline region was formed at the ex- pense of the cytoplasm and that the material of each chromosome tended to remain within the space of its own vesicle, a core of chroma- tin being particularly noticeable in the center of this region, and that the prophase chromosome subsequently formed was developed out of the substance of one, and only one, of the previously existing telophase chromosomes. Sutton ('00) was the first to describe the vesicles of the spermatogonia of a grasshopper. Since then, Otte ('07) has seen similar structures in Locusta, and Davis ('08) in several Acrididae; Pinney ('08) has described them for Plirynotettix. Sutton stated that in the earlier stages of nuclear formation, each chromosome produced a separate vesicle, just as I have found for Plirynotettix, but that in later stages, the proximal ends fused together, giving a common nuclear cavity, from which the distal ends of the vesicles, particularly the longer ones, projected out like the fingers of a glove. Sutton interpreted these conditions as lending strong support to the theory of individuality. Otte believed that the individual vesicles remain distinct throughout the whole of the interkinetic phases, and WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 115 Pinney reached a similar conclusion. Davis, on the other hand, could recognize only an irregular outline for the nucleus, and did not identify the vesicle of even the monosome with certainty. Gerard ('09) saw the hyaline regions about the telophase chromosomes of Stenobothrus, and stated that the nuclear membrane was formed in connection with them. Since similar conditions have been reported by so many observers, it would seem that these vesicular structures are the result of normal processes and not, as claimed by Vejdovsky ('11-12), artifacts. If we turn to accounts other than those on orthopteran spermato- genesis, we find that the formation of chromosornic vesicles out of individual chromosomes in the telophase is by no means of rare occurrence. Van Beneden ('83) noted in his work on Ascaris that each of the two chromosomes of the female pronucleus often formed a separate 'half -nucleus.' Hacker ('95b) observed that the chromo- somes of the early cleavages of Cyclops brevicornis formed two groups of "Blaschen," one group from the maternal and another from the paternal pronuclei. Conklin ('02) calls attention to the occurrence of such chromosomic vesicles, and gives the history of the nuclear changes during the cycle of division in Crepidula as follows (p. 45) : — "(1) The cliromosomes, consisting of chromatin enclosed in a linin sheath, divide and move to the poles of the spindle, where they par- tially surround the spheres. (2) Here they become vesicular, the interior of the vesicle becoming achromatic, though frequently con- taining a nucleolus-like body, while the wall remains chromatic. (3) These vesicles continue to enlarge and then unite into the "resting nucleus." The nuclear membrane is composed of the outermost walls of the vesicles, while the inner walls stretch through the nucleus as achromatic partitions. The chromosomal vesicles for the egg and sperm nuclei remain distinct longer than those from the same nucleus .... Such vesicles are found generally, if not universally, in the early division of ova, though they are not usually found in other mitoses." Small wood ('05) describes similar chromosomic vesicles in the eggs of nudibranchs. He found that during the "rest-pause" between the first and second maturation divisions the chromosomes frequently have distinct vesicles. There may be a single vesicle for all the chromosomes, or a single vesicle for each chromosome; all conditions between these two extremes occur. Medes ('05) in her work on Scutigera found in the second spermatocytes (p. 174) that: — "There is no immediate formation of a nuclear membrane, but each separate chromosome, as it disintegrates, becomes enclosed in a membrane of its 116 bulletin: museum of comparative zoology. own, thus forming a structure similar to a nucleus but containing only a single chromosome." Kornhauser ('15, p. 408) says concerning the spermatogonia of Hersilia: — "The telophase chromosomes become gradually fainter in outline, and a clear area in the cytoplasm begins to form about them. It is, I believe, the boundary between this clear area and the more reticular cytoplasm which forms the new nuclear membrane." Thus it will be seen that it is quite usual for telophase chromosomes to form individual nuclei, which later fuse to form the whole nucleus, and with Smalhvood ('05) we may accept this tendency as an argument for chromosomal individuality. (2) Theories of continuity. Among those who support a theory of continuity, there is not always agreement as to what structures are carried from one cell-generation to the next. It is generally agreed that the chromosomes are composed of at least two substances; the chromatin and the ground substance (linin, plastin). Hacker ('04) formulated the " Successionshypothese," stating that the persisting structures of the chromosomes consisted of the "Grundsubstanz," or achromatic part. Bonnevie ('08a) and others, on the contrary, regard the chromatic substance as the persistent portion and the achromatin as the temporary part of the clu'omosome. Vejdovsky ('07, '11-12) has evolved a most elaborate theory touch- ing this problem. In his monograph of 1907, he based his conclusions on a study of the ovogenesis and maturation of some annelids. He concludes that the nucleus is derived from the chromosomes and from them alone. He divides the interkinetic stages into two periods; the one during which the nucleus is formed out of the chromosomes he calls " katachromasis," and the one during which the chromosomes are formed out of the nucleus he calls "anachromasis." In his later monograph ('11-12) he analyses these processes still further and attempts to describe in detail the events in the two periods. His conclusions may be briefly stated as follows : — A chromosome is com- posed of two substances, one a less deeply staining substratum, on the surface of which is the other, the more deeply stainable chromatin. In the early stages of katachromasis, the chromatin differentiates into a spiral thread, or "chromonema," which is coiled about the surface of the substratum. The substratum then dissolves, forming the nuclear sap, or " enchylema." The chromonema further differentiates into a finely coiled chromatic portion, inside of which is a linin core. In this condition, he recognizes the anlage of the chromosome of the succeeding generation. The linin substance of the chromonema is to become the substratum of the future chromosome, and the finely WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 117 coiled chromatic portion will become its chromonema. In successive generations, therefore, there is a changing composition of the chromo- somes. During each katachromasis, the ground substance of the chromosome dissolves, leaving the chromonema, which becomes differentiated into the two kinds of substance found in the chromo- some of the next generation. This is an ingenious theory, to say the least, and carries with it some measure of support for the theory of individuality, inasmuch as each new chromosome is formed out of the substance of a preceding one. I have found nothing in my studies to support any one of these theories to the exclusion of the others. It is rather surprising, how- ever, that Vejdovsky found no indication of the chromosomic vesicles in the spermatogonia of the Orthoptera that he studied and, that he regards those seen by others as artifacts. I find little evidence of a chromonema in the telophase of the spermatogonia, and what evidence there is would indicate that the chromatin becomes distributed on the inner surface of the vesicular walls, not on the outer surface of an achromatic core. In the telophase of the last spermatogonia, I find a spiral thread forming, but it develops out of the chromatin at the middle of the area occupied by a vesicle. But whether we accept any one of these theories, or reject all of them, there still remain the strongest grounds for believing, as they all indicate, that there is some underlying organization which is in some way perpetuated for each individual chromosome. I am inclined to the belief that this organization involves both chromatic and achromatic substance. In plant material evidence which indicates a continuity of the chromosomes has not been wanting. Gregoire ('07, '10) believes that the results of his own investigations and those of others on plants furnish strong support for the individuality theory. Stout ('12) has recently added evidence for this belief in his work on Carex aquatilis. He says ('12, p. 36): — "The chromosomes are present in all resting nuclei as visible units of a definite number. These individual chromor somes can be traced as such through all stages of both somatic and germ-cell divisions, with the exception of the various stages of synapsis (synizesis)." Lee ('13) also finds continuity of the chromosomes in plants through the " rest-stage." He beheves that the chromosomes of even the metaphase become vacuolated, that this vacuolization increases in the telophase, where, later, a spiral thread is formed out of each cliromosome. This spiral thread becomes the prophase chromosome of the succeeding division. He introduces the term "spirophase" to designate the so-called "rest-stage." 118 bulletin: museum of comparative zoology. There* seems to be a great amount of disagreement as to just what constitutes individuahty, but I beheve that we may class as instances of individuahty all cases where it can be shown that the substance of any telophase chromosome gives rise to one and only one prophase chromosome. In that event, any one of the three theories mentioned above would support the theory of individuality. I believe that I have demonstrated individuality for chromosome-pair A, and have shown good evidence for it among the other chromosomes of Phryno- tettix. Besides, it seems to me much more logical to regard the con- stant reappearance of the same architectural conditions of a given chromosome as a result of continuity of that architecture in some form or other through all the cell-divisions, than to assume that the organization is entirely destroyed and reestablished between successive mitoses. C. Chromosomes and Heredity. Any discussion of the relation of the chromosomes to heredity must deal to a considerable extent with theory and speculation. Yet there are many facts which tend to the belief that the chromosomes are, after all, directly concerned with the transmission of hereditary qualities. A few facts and some theory will be considered in the following paragraphs under the two heads: — (a) Mendelism and maturation, and (b) some experimental evidence. a. Mendelism and Maturation. Wilhelm Roux was apparently the first to formulate, in the early eighties, a theory in which an attempt was made to localize the carriers of hereditary qualities in the chromosomes; this was later elaborated by others, especially by Weismann, who postulated a reduction division which has since been identified with one or the other of the maturation divisions. Montgomery ('01) pointed out that the chromosomes of the diploid series occur in pairs, the members of each pair being of the same shape and size. There are thus two similar series of chromosomes. He concluded that one series was derived from the maternal, the other from the paternal ancestor. He concluded further that the members of each pair unite to form the bivalent chromosomes of the first spermatocytes. Boveri ('02) de- cided from the results of his experiments on dispermic echinoderm WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 119 eggs, that the chromosomes were quahtatively different. Sutton ('03) in the following year, explained how the behavior of the chromo- somes in matm*ation could be correlated with the behavior of Men- delian characters. He showed: — (1) that the union of clu-omosomes of diverse origin into pairs and their subsequent separation in one of the maturation divisions would insure to every gamete one of every kind of chromosome in the series: (2) that if the law of chance were operative in the orientation of the pairs on the maturation spindles, every possible combination of male and female chromosome could result; and (3) that such a recombination according to the law of chance would account for the transmission of Mendelian characters, if the chromosomes retained their individuality and really were the carriers of the qualities. This work of Sutton has been generally accepted as proving the correlation assumed, but it remained for Carothers ('13) to demon- strate that the law of chance actually does operate in the distribution of the chromosomes in the maturation spindles. In the case of the unequal tetrads described by her, it was shown that either the large or small member of the pair may go to the same pole as the accessory chromosome, which, as usual, was found to go to one pole undivided. Moreover, it was found that the ratio between the two results of distribution was approximately one to one. Robertson ('15) has very recently published some of his work on the Tettigidae, where he has found the same rule to hold for the unequal pairs that were present in his material. The behavior of tetrad Ci in Phrynotettix agrees with that described by Carothers and Robertson. These cases establish the fact that there really is a distribution of chi'omosomes in the maturation divisions according to the law of chance. A further consideration of the cases of unequal tetrads in Orthop- tera will show in how far the theoretical possibilities as to chance distribution have been realized. Baumgartner ('11) in reporting his results on Gryllotalpa borealis before the American Society of Zoolo- gists, stated that he found in the first maturation mitosis an unequal pair of chromosomes, of which the larger dyad always went to the same pole as the accessory. Payne ('12) found the same conditions in this species of Gryllotalpa. He regards the large member as possibly associated with the accessory to form a sex-group, similar to the groups in Conorhinus and Fitchia (Payne, '09), or in Thyanta (Wilson, '10), with the exception that in Gryllotalpa the grouping occurs in the first spermatocyte metaphase instead of the second. Payne suggests that the chromosomes instead of following a haphazard 120 bulletin: museum of comparative zoology. method of distribution in the maturation divisions, may always move the same way, i. e., all the chromosomes brought into the egg may pass into the female-producing sperm. It is extremely doubtful if the last suggestion will prove applicable as a general rule, but the conditions in Gryllotalpa are interesting exceptions to what has been found in the Acrididae and Tettigidae.^ Hartmann ('13) describes small chromosomes as dividing unequally in some male germ-cells of Schistocerca. In one first-spermatocyte cell he found two such chromosomes (tetrads) dividing unequally, and he found some cases of unequal division in the secondary sperma- tocytes. These observations, if correct, would lead one to suspect that he might" have been dealing with a condition similar to that in Phrynotettix, except that in the first division, either both the small chromosomes divided sometimes reductionally and sometimes equa- tionally, or, while one of them followed this method, the other always divided reductionally. Bringing together the results of Baumgartner and Payne for Gryllo- talpa, those of Carothers for Acrididae, Robertson for Tettigidae, and my own for Plirynotettix, we may arrange a graded series of condi- tions beginning with (1) tetrad B, in Phrynotettix, which is unequal, but divides equationally in the first division; passing (2) to Ci, which divides with equal frequency either reductionally or equationally in the first division, and when dividing reductionally shows chance distribution with reference to the accessory; thence (3) to the unequal types found by Carothers and Robertson, which always divide re- ductionally in the first division but show chance distribution, and finally (4) to Gryllotalpa, where division is always unequal in the first spermatocytes, but the larger dyad always accompanies the accessory. Whether this series offers any possible explanation as to the origin of these unequal elements, and their different kinds of behavior, is prob- lematical. Robertson's work deserves further consideration, because he has found two of the tkree possible combinations which would be expected out of a random recombination of two unequal elements which con- jugate. In the case of Tettigidea, he found the unequal tetrad in ' i^oslscrij)!. — Unfortunately Jif had [overlooked the results reported for Gryllotalpa vul- garis by Voinov ('14), who found in the first spermatocyte metaphase an unequal pair of dyads, which separate so that sometimes the larger dyad and sometimes the smaller one goes to the same pole as the accessory chromosome. These results are in accord with those mentioned above for the Acrididae and the Tettigidae and it may be surmised that similar conditions perhaps obtain for Gryllotalpa borealis but have so far been overlooked. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 121 only two individuals, all the others showing a pair both members of which were equal to the larger member of the unequal pair. The third possibility, an equal pair, homologous to the smaller of the dyads, was not found. This case is analogous to that of B in Phiynotettix.^ In Acridium, Robertson found two individuals, one a male, the other a female, possessing an unequal pair of chromosomes, whereas all the other individuals studied showed the homologous pair to be equal, both members being equivalent in size to the smaller of the two members of the unequal pair. This case is analogous to those of tetrads Ci and C2 in Phrynotettix, where, also, only two combinations, the same two, out of a possible three have been found. Robertson calls attention to the obvious possibility of a loss of chromatin from the unequal pair in Tettigidea, and suggests that the loss of Mendelian factors could be accounted for in this way. He also suggests that the loss of the distal ends of both the chromosomes, resulting in a pair of small dyads each equivalent to the smaller mem- ber of the unequal pair, might result in lethal conditions, or might mean the loss of factors necessary for development. In the case of the unequal pair in Acridium, he assumes that there has been an addi- tion to one member of the smaller pair. If this element is similar to Ci of Plu*ynotettix, as it seems to be, then the simpler explanation would be that a part had been lost, just as in the case of the one in Tettigidea. It is curious that in both Ci and the unequal pair in Acridium, the same combination, i. e., a pair both members of which would be equal to the larger member of the unequal pair, is lacking. I am inclined to believe, if sufficient material were available, that the remaining possible combinations would be found. The matter could, at least, be tested by experiment. It is the hope of the writer to conduct breeding experiments with this object in view. One further point remains to be considered in relation to chromo- some-pair C. I have described these tetrads in detail elsewhere (p. 85), but a reference to figure 107 (Plate 9) will recall that there are three types, which I have designated as Ci, C2, and C3. If similar types exist in the female, — Robertson ('15) found an unequal pair in a female of Acridium, — and random mating be assumed for the ani- mals possessing the three different types, then one ought to obtain in * Since writing this I have had an opportunity to examine slides frcn some new Phrynotettix material collected during the summer of 1915 by Miss Garothers of the University of Pennsyl- vania. In some of the individuals of the new material I have found the expected third type of chromosome-pair B composed of two elements both equivalent to the shorter member of the unequal type. 122 bulletin: museum of comparative zoology. the offspring all possible combinations of the three kinds of chromo- somes. Or, if mating involving any two of the types could be made, there should result all possible combinations between them. On account of the small number of animals available for my study, no conclusions as to whether these conditions are realized in nature could be drawn. The presence of the three types in these few animals, however, strongly suggests the possibility of realization, especially since two of the three possible combinations are realized for the two kinds of chromosome in type Ci. The presence of a third type also suggests that there may exist in this case the mechanism for the transmission of triple allelomorphs. b. Some experimental Evidence. The most extensive breeding experiments the results of which tend to show that the chromosomes are concerned in the transmission of hereditary characters are those on Drosophila by Prof. T. H. Morgan and his students. In the course of this work they have dealt with over a hundred unit-characters which show Mendelian inheritance, either in a typical or modified form. In Drosophila, there are four pairs of chromosomes, of which one pair is very small, and one is a pair of heterochromosomes, or " sex-chromosomes." In their behavior in inheritance, the hundred and more characters fall into four groups, each group tending to behave as a unit, just as it would be expected to do in case it were carried by a single pair of chromosomes. Of these groups of characters, one is very small, the others much larger, the largest one being the group of " sex-linked " characters. Naturally the small group of characters has been correlated with the small pair of chromosomes and the group of sex-linked characters with the sex- chromosomes. But there have been exceptions in the case of many pairs of allelo- morphs, especially those that are sex-linked, i. e., cases where factors belonging to a certain group have gone into a mating together, but have not always reappeared together, as they would be expected to do if they were all carried by a single chromosome and that cliromosome maintained its individuality. These phenomena have been explained by the so-called "cross-over" hypothesis. In this connection Morgan ('11) developed what has been termed the "linear arrangement" hypothesis, which was further elaborated by Sturtevant ('13). These authors assume that the factors, or "genes," which represent the WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 123 characters, are distributed in a linear series along the length of the chromosomes. Then, invoking the aid of the " chiasmatype " theory of Jannsens ('09), they attempt to explain the "cross-overs" by as- suming, first, that when two chromosomes conjugate side by side, they may become twisted around each other, and, secondly, that the later separation is along a plane, which cuts across the threads once for every complete twist. Considering the matter in relation to the tetrad stages, it might be imagined that the two tlireads cross each other, and that at the point of crossing, a weakness of the strands causes them to break and then recombine, forming threads each of which is composed of a part of both the original conjugants. Judging from his figures, Janssens founded his theory on conditions similar to those shown in my figure 38, a-d (Plate 3). I am quite sure that the evidence in Phrynotettix does not support the idea that the chromatids break and recombine in any of the postspireme stages. On the contrary, I believe that the chromatids maintain as strict an individuality as I have claimed for the chromosomes themselves. And since these tetrad figures are repeated in so many animals, and even in plants, there would seem to be ground for supposing the be- havior to be similar in all. On the basis of this hypothesis, however, Morgan and his pupils have been able to explain the anomalous behavior of the genes which they call the "cross-over" in a very satisfactory way; furthermore, they have been able to use it in connection with the linear-arrangement hypothesis to predict the behavior of any given character, with refer- ence to any other character in the group to which it belongs, provided its behavior in relation to one or two of the characters of the group is known. But there is one point yet to be noted. I have based my criticism of the chiasmatype theory on the conditions as found in spermatogenesis. One of the peculiar facts found in the work on Drosophila is that there is no "crossing-over" in the male. But why should such a phenomenon occur in the female and not in the male? Is it not possible that in the "great growth" period of the oocyte, — where the tetrads become so much more expanded and diffused than in the male, even seeming to disappear entirely in some cases, — the tetrads might suffer some such changes as those suggested by the experimental results? There is also to be considered the often re- peated condition of parasynapsis in Drosophila, as shown by Metz (*14), which might offer greater opportunities for such "cross-overs" to occur than are found in other animals. Whatever else may be said of the results of the experiments on 124 bulletin: museum of comparative zoology. Drosophila, it must be admitted that they go very far towards estab- lishing a direct relationship between the elii'omosomes and the trans- mission of Mendelian characters. Perhaps the most convincing evi- dence of this kind is that obtained b}^ Bridges. He has found that in certain strains involving sex-linked inheritance, some exceptional females appeared which were like their mothers in every respect, and showed no transmission of sex-linked characters from the father, although such transmission would be expected, since the male sex formula is XY and that of the female is XX. Furthermore, he found that such exceptionally produced females inherit directly from their mother the power of producing like exceptions (about 5%). The explanation advanced by Bridges ('14) was that "the sex-linked genes were borne by the X-chromosomes and that 10% of the eggs of the exceptional females retained both of the Z^-chromosomes, or conversely lost both to the polar body." This phenomenon was called "non- disjunction." Breeding experiments showed that an X-chromosome gene could not be the cause of the phenomenon, and the prediction was accordingly made that half the daughters of a non-disjunctional female would be found to contain in addition to the two X-chromo- somes a supernumerary chromosome which would be a Y. Cytologi- cal investigations have shown that approximately one-half of the daughters of a non-disjunctional female do, in fact, contain a super- numerary F-chromosome, while the remaining half contain only the two X'-chromosomes. I may add that thi'ough the kindness of Dr. Bridges, I have been able to examine some of his slides and convince myself of the presence of the extra chromosome. This brilliant piece of work makes it very hard to disagree with Bridges's conclusion ('14, p. 109) that, "there can be no doubt that the complete parallelism between the unique behavior of the chromosomes and the behavior of the sex-linked genes and sex in this case means that the sex-linked genes are located in and borne by the sex-chromosomes." Returning now to a consideration of the linear-arrangement hypo- thesis, it must be admitted that the theory has attractive possibilities, and up to the present time has stood the test of experimental breeding in Drosophila. It may not be out of place, therefore, to call attention in this connection to the constancy of the granular, or chromomeral organization of the chromosomes of Phrynotettix, particularly in chromosome-pair B. May not this constancy of architecture of the chromosomes have a meaning correlated with that assumed in the linear-arrangement hypothesis? This possibility seems to me to be worthy of further investigation. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 125 D. Summary of Conclusions. It is believed that the present study of the spermatogenesis of Phrynotettix magnus has demonstrated : — 1. That conjugation of the chromosome-pairs is by parasynapsis. 2. That the majority of the bivalent chromosomes divide equa- tionally in the first maturation division. 3. That the chromosomes retain their individuality through the spermatogenic cell-generations. 4. That the so-called ' plasmosomes ' take their origin from some definite region (granule) of particular chromosomes, but that they may be variable in occurrence and in extent of development. 5. That in the maturation divisions {e. g. chromosome-pair Ci) the law of chance is followed in the distribution of the chromosomes. 6. That each chromosome possesses a definite organization, which is expressed in the constancy of the relative sizes and positions of its chromomeres (as seen, e. g., in chromosome-pair B). 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Studies in spermatogenesis with especial reference to the " acces- sory chromosome." Carnegie instn. Publ. 36, 320 pp., 7 pis. '08. A study of the germ cells of certain Diptera, with reference to the heterochromosomes and the phenomena of synapsis. Journ. exp. zool., 5, p. 359-374, 4 pis. '10a. The chromosomes in the germ-cells of Culex. Journ. exp. zool., 8, p. 207-225. '10^. An unequal pair of heterochromosomes in Forficula. Journ. exp zool, 8, p. 227-241, 3 pis. '12a. Supernumerary chromosomes, and synapsis in Ceuthophilus (sp.?). Biol, bull., 22, p. 219-230, 35 text fig. '12^. Further observations on supernumerary chromosomes, and sex ratios in Diabrotica soror. Biol, bull., 22, p. 231-238, 13 text fig. Stout, A. B. '12. The individuaUty of the chromosomes and their serial arrangement in Carex aquatilis. Arch. f. zellf., 9, p. 114-140, taf. 11, 12. Sturtevant, A. H. '13. The linear arrangement of six sex-Unked factors in Drosophila, as shown by their mode of association. Journ. exp. zool., 14, p. 43-59. Sutton, W. S. '00. Spermatogonial divisions of Brachystola magna. Kans. univ. quart., 9, p. 135-160, pi. 32-35. 132 bulletin: museum of comparative zoology. Sutton, W. S '02. On the morphology of the chromosome group in Brachystola magna. Biol, bull., 4, p. 24-39, 11 text fig. '03. The chromosomes in heredity. Biol, bull., 4, p. 231-251. Taylor, Monica. '14. The chromosome complex of Culex pipiens. Quart, journ. micr. sci., 60, p. 377-398, pi. 27, 28, 3 text fig. Tennent, D. H. '08. The chromosomes in cross-fertihzed echinoid eggs. Biol, bull., 15, p. 127-134, 1 pi., 2 text fig. Vejdovsky, F. '07. Neue untersuchungen uber die reifung und befruchtung. Bohm. gesell. wiss., Prag., 103 pp., 9 taf., 5 text fig. '11-12. Zum problem der vererbungstrager. Bohm. gesell. wiss., Prag., 184 pp., 12 taf., 16 text fig. Voinov, D. '14. Sur un nouveau mecanisme determinant le dimorphisme des 616- ments sexuels; chromosome a polarity variable. Compt. rend, stances Soc. biologie (S6ances de la reunion biologique de Bucarest du 19 Fevrier, 1914), 76, p. 509-511. Wassilieff, A. '07. Die spermatogenese von Blatta germanica. Arch. f. mikr. anat., 70, p. 1-42, taf. 1-3. Weismann, A. '87. Ueber die zahl der richtungskorper und tiber ihre bedeutimg fur die vererbung. Jena, viii + 75 pp., 3 text fig. Wilcox, E. V. '94. Spermatogenesis of Caloptenus femur-rubrum and Cicada tibicen. Bull. M. C. Z., 27, p. 1-32, 5 pis. '96. Further studies on Caloptenus femur-rubrum. Bull. M. C. Z., 29, p. 191-203, 3 pis. '97. Chromatic tetrads. Anat. anz., 14, p. 194-198. '01. Longitudinal and transverse divisions of chromosomes. Anat. anz., 19, p. 332-335. Wilson, E. B. '09. Studies on chromosomes. IV. The " accessory " chromosome in Syromastes and Pyrrochoris with a comparative review of the types of sexual differences of the chromosome groups. Journ. exp. zool , 6, p. 69-99, 2 pi., 2 text fig. '10. Studies on chromosomes, VI. A new type of chromosome combina- tion in Metapodius. Journ. exp. zool., 9, p. 53-78, 5 text fig. '11. Studies on chromosomes. VII. A review of the chromosomes of Nezara; with some more general conclusions. Journ. morph., 22, p. 71-110, 1 pi., 9 text fig. WENRICH: spermatogenesis of PHRYNOTETTIX MAGNUS. 133 Wilson, E. B. '12. Studies on chromosomes. VIII. Observations on the matm-ation phenomena in certain Hemiptera and other forms, with considerations on synapsis and reduction. Journ. exp. zool., 13, p. 345-431, 9 pb. Winiwarter, H. von. '00. Recherches sur I'ovogfoese et I'organog^nese de I'ovaire des mam- miferes (lapin et homme). Arch, biol., 17, p. 33-199, pi. 3-8. Wodsedalek, J. E. '13. Spermatogenesis of the pig with special reference to the accessory chromosomes. Biol, bull., 25, p. 8-45, 6 pis. Zweiger, H. '06. Die spermatogenese von Forficula auricularia L. Jena, zeit., 42, p. 143-172, taf. 11-14. EXPLANATION OF PLATES. The drawings in plates 1 to 9 were in all cases made with the aid of a camera lucida, using a Zeiss No. 12 compens. ocular and a Spencer homogeneous immersion 1.8 mm. objective, N. A. 1.30. Drawings no. 63-65 were made at a magnification of 2,600 diameters, all the others at a magnification of 3,000 diameters. The drawings have been reduced one third in reproduction. Plate 10 consists of photomicrographs made at a magnification of 1,150 diameters. Symbols used: A designates selected chromosome-pair "A." D (( « « u li r> >> "C". X " accessory chromosome. G " composite granule. g ** polar granule. PLATE 1. Wenrich.— Spermatogenesis of Phrynotettix magnus. PLATE 1. All figures are of the early spermatogonia. Figs. 1, 2. — In metaphase. Figs. 3, 4. — In anaphase. In figure 4 the cytoplasm has been drawn in order to show the hyaline region around the chromosomes. Fig. 5. — Early telophase. Figs. 6, 7. — Side view and optical section (polar view), respectively, of an early telophase in which the chromosomes have begun to expand. The cytoplasm has been represented in order to show the hyaline region around the chromosomes. Figs. 8, 9. — Side view and optical section, respectively, of later telophase. Vesicles persistent. Figs. 10, 11. — Side view and optical section, respectively, of late telophase. Vesicles mostly disappeared. Figs. 12, 13. — Side view and optical section, respectively, of stage showing the greatest diffusion of chromatin that was found. BULL. MUS. COMP. ZOOL. Wenrich- Spermatogenesis Phrynotettix, Plate 1 Wenrich. — Spermatogenesis of Phrynotettix magnus. PLATE 2. Figs. 14-20. — Prophases of early spermatogonia. Figs. 14a, 14b. — Two successive sections of the same cell in earUest prophase. Figs. 15a, 15b. — Two sections of a single cell in early prophase. Formation of fine spiral thread. Fig. 16. — Polar view of later prophase. Fine spiral threads, all distinct. Composite granules prominent. Fig. 17. — Side view of later prophase. Accessory in vesicle. Longitudinal epUt beginning to appear in threads. Fig. 18. — Section from distal pole of nucleus. Vesicular walls persisting. Fig. 19. — Later prophase. Composite granules breaking up into polar granules. Fig. 20. — Later prophase. Accessory still in vesicle. Figs. 21, 22. — Side view and optical section, respectively, of telophase of last spermatogonial division. Figs. 23, 24. — Oblique side view and optical section, respectively, of early preleptotene stage. BULL. MUS. COMP. ZOOL. vVenrich- Spermatogenesis Phrynotettix, Plate 2 PLATE 3. Wenrich. — Spermatogenesis of Phrynotettix magaus. PLATE 3. Figs. 25-27. — Later preleptotene stage. Fig. 28. — Early? leptotene stage. Fig. 29.— Later leptotene stage. Fig. 30. — Early zygotene stage. «• Fig. 3L — Later zygotene stage. Figs. 32, 33. — Early pachytene stages. Fig. 34. — Later pachytene stage. Figs. 35-37. — Stages of breaking up of composite granule of pachytene stage into its component polar granules. Fig. 37.— Diplotene stage. Fig. 38, a-g. — Different types of tetrads from a single cyst of postspireme, or tetrad, stage. BULL. MUS. COMP. ZOOL. Wenrich.- Sp£Rmatogenesis Phrynotettix, Plate 3 Wenrich. — Spermatogenesis of PhrynotelUx magnus. PLATE 3. Figs. 25-27. — Later preleptotene stage. Fig. 28. — Earlji leptotene stage. Fig. 29. — Later leptotene stage. Fig. 30. — Early zygotene stage. Fig. 3L — Later zygotene stage. Figs. 32, 33.— Early pachytene stages. Fig. 34.— Later pachytene stage. Figs. 35-37. — Stages of breaking up of composite granule of pachytene stage into its component polar granules. Fig. 37. — Diplotene stage. Fig. 38, a-g. — Different types of tetrads from a single cyst of postspireme, or tetrad, stage. BULL. MUS. COiMP. ZOOL. WtNRiCH.- Spermatogenesis Phrynotettix, Plate 3 PLATE 4. Wenrich. — Spermatogenesis of Phrynotettix magnus. PLATE 4. Figs. 39-44. — Stages in division of first spermatocytes. Fig. 39. — Polar view of metaphase, showing twelve chromosomes. Fig. 40. — Polar view of metaphase (eleven chromosomes). Fig. 41. — Side view of anaphase, showing dyads. Fig. 42. — Polar view of anaphase, showing twelve dyads. Fig. 43. — Polar view of anaphase, showing eleven dyads. Fig. 44. — Telophase. Figs. 45-49. — Interkinesis stages. Fig. 45. — Dyads undergoing diffusion. Fig. 46. — Dyads (except the accessory) still more diffused. Fig. 47. — Stage of greatest diffusion. OutUne of dyads uncertain. Fig. 48. — A prophase of the second spermatocyte division. Outline of dyads reappearing in the middle of the area in which they diffused. Fig. 49. — Late prophase of second spermatocyte. Dyads fully reformed. BULL. MUS. COMP. ZOOL. Wenrich- Spermatogenesis Phrynotettix, Plate 4 PLATE 5. Wenmch. — Spermatogenesis of Phrynotettix magnus. PLATE 5. Figs. 50-55. — Stages in division of the secondary spermatocytes. Figs. 50, 51. — Polar views of metaphase, showing eleven and twelve chromo- somes, respectively. Fig. 52. — Side view of metaphase. Fig. 53. — Side view of early anaphase. Fig. 54. — Side view of late anaphase. Fig. 55. — Polar view of telophase. Figs. 56-61. — Pachytene stages of first spermatocyte. Figs. 56, 57 showing spireme loop of selected chromosome-pair "A," Figs. 58, 59 showing spireme loops of selected chromosome-pair "B," and Figs. 60, 61 showing spireme of selected chromosome-pair "C." Figure 61 shows in addition the "B" spireme. BULL. MUS. COMP. ZOOL. WENRICH- SPERMATOGENESIS PHRYNOTETTiX, PLATE 5 PLATE 6. Wenbich. — Spermatogenesis of Phrynotettix magnus. PLATE 6. Fig. 62. — Stages in the transformation of chromosome-pair "A," from pachytene spireme to metaphase of the first spermatocyte. Fig. 63. — Similar stages for the chromosome-pair "5," for the equal-type. Fig. 64. — Same stages for the pair "5," unequal type. Fig. 65. — Similar transformation stages for chromosome-pair "C" Fig. 66. — Telophase of an early spermatogonimn, showing members of chro- mosome-pair "A." Fig. 67. — Telophase of one of the last spermatogonia, showing members of chromosome-pair "A." Figs. 68, 69. — Preleptotene stages, showing persistence of chromosome-pair "A." Fig. 70. — Early leptotene. Conjugation of pair "A." Fig. 71. — Later leptotene. Conjugation of pair "A." Spireme of the ac- cessory. Fig. 72. — Spireme of the accessory. Fig. 73. — Zygotene stage, showing conjugation of pair "A." Fig, 74. — Zygotene stage, showing conjugation of pair "A" completed. BULL. MUS. COMP. ZOOL. WeNRicH.- Spermatogenesis Phrynotettix, Plate 6 '." €• f*' ^. / F " .* ' ' ■■ 64 «_ m^ ^ ««<^J 6 6 'A 70 < PLATE 7. Wenhich. — Spermatogenesis of Phrynotettix magnus.' PLATE 7. Figs. 75-78. — Zygotene stages showing incomplete conjugation of chromo- some-pair "A." Fig. 79. — Side view of first spermatocyte metaphase, showing tetrad "A," as an extended rod with a constriction in the middle. Fig. 80. — Polar view of a telophase of the first spermatocyte, showing dyads "A" and "B." Fig. 81. Side view of a telophase of the first spermatocyte, showing dyads of "A" and the accessory. Fig. 82. — Side view of first spermatocyte telophase, showing the "A" dyads in both daughter cells. Fig. 83. — Interkinesis stage, showing the accessory dyad and the "A" dyad (more condensed). Fig. 84. — Interkinesis stage, showing the accessory dyad and the "A" dyad (less condensed). Fig. 85. — Telophase of the second spermatocj^te, showing the "A," "B," and accessory monads. Fig. 86. — Telophase of the secondary spermatocyte, showing the "A" and "B" monads. BU LL. MUS. COMP. ZOOL. WeNRICH.- SPERMATOGENESIS Phrynotettix, Plate 7 PLATE 8. W ENRICH. — Spermatogenesis of Phrynotettix maenns PLATE 8. Figs. 87-96. — Telophases of spermatogonia, showing members of chromo- some-pair "5" (all from one animal, no. 772). Fig. 97, a-m. — One example of chromosome-pair "B," in the pachytene stage from each of the 13 animals studied. Fig. 98, a-p. — Examples of chromosome-pair "B" from the pachytene stage of a single animal (no. 772). BULL. MUS. COMP. ZOOL. Wenrich.- SPERMATOGENESIS Phrynotettix, Plate 8 I ^ rn^iJ PLATE 9. Wenrich. — Spermatogenesis of Phrynotettix maguus. PLATE 9. Fig. 99. — Side view of first spermatocyte metaphase, showing roughened condition of the accessory, of chromosome-pair "5," and of the polar tips of autosomes. Fig. 100. — Side view of first spermatocyte metaphase, showing roughened condition of chromosome-pairs "B" and " C," and of the accessory. Figs. 101-105.— Telophases of spermatogonia, showing the larger members of the chromosome-pairs " B" (unequal type) and "C" (type Ci). Fig. 106. — An example of tetrad "B" from each of the 13 animals studied. Fig. 107. — An example of tetrad "C" from each of the 13 animals studied. (The corresponding letters in figures 106 and 107 refer to the same animal) . Figs. 108-110. — Telophases of connective-tissue nuclei, showing polar gran- ules and chromosome "B." Figs. Ill, 112. — Nuclei from the foUicular envelope. BU LL. MUS. COMP. ZOOL. Wenrich- Spermatogenesis Phrynotettix, Pu A *'« ffll «i ^^ /- J e 106 f 9 h 107 5.-^ W ENRICH.' — SpermatxjeeneHis of Phrynotettii maRDUs. PLATE 10. Photomicrographs . Fig. 113. — Spireme of "A" in the pachytene stage. Fig. 114.— Tetrad of "A." Fig. 115. — Pachytene spireme loop of "B." Fig. 116.— Tetrad of "B." Fig. 117.— Tetrad of "C" (type Ci). Fig. 118. — Both tetrads, "B" and "C," dividing equationally in the same spindle. Fig. 119. — Tetrad "B" dividing equationally and tetrad "C" dividing re- ductionally in the same spindle. Fig. 120.— Unequal division of "C." Smaller dyad going to same pole as the accessory. Fig. 121. — Unequal division of "C." Larger dyad going to same pole as the accessory. BULL. MUS. COMP. ZOOL. vVenrich- Spermatogenesis Phrynotettix, Plate 10 113 *> H 114 ^-Ji 115 ^~ B 116 117 /■ X ♦ / '< *■ I! 118 X X \A I 119 y 4tH V 120 121 D. H. W. PHOTO. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 4. A REVISION OF THE LIZARDS OF THE GENUS CYCLURA. By Thomas Barboub and G. K. Noble. With Fifteen Plates. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM. February, 1916. No. 4. — A Revision of the Lizards of the Genus Cyclura. By Thomas Barbour and G. K. Noble. INTRODUCTION. Some years ago while working upon West Indian reptiles the senior author became interested in Cyclura. Every opportunity has been grasped which offered the slightest probability of securin'g specimens, so that now the Museum of Comparative Zoology con- tains more species of that genus than any other museum. That the series is by no means large, will appear at once. The preparing of this revision would have been difficult but for the friendly interest of Mr. H. W. Fowler of the Academy of Natural Sciences of Phila- delphia; the unique type of our C. nuchalis is in the Museum of the Academy. We take great pleasure in dedicating C. stejnegeri from Mona Island to Dr. Stejneger, through whose kindness a paratype from the small series in the U. S. N. M. has been retained for the M. C. Z. From the Carnegie Museum in Pittsburgh we have speci- mens of C. rileyi and of C. macleayi, from the Isle of Pines, presented in return for the identification, by the senior author, of the Carnegie Museum series of West Indian reptiles. These he was allowed to study through the kindness of Prof. L. E. Griffin. A number of Rhinoceros Iguanas have been received from time to time at the New York Zoological Park, have died and probably most of them have found a resting place in the American Museum in New York. These cannot now be found; one of the examples, however, now mounted in the Museum of Comparative Zoology, a gift of the N. Y. Zoolog- ical Society, was said to be from Navassa Island and seems to repre- sent the species confined to that island. In general, zoological park specimens, while very valuable for anatomical study, are often with- out locality, although this is sometimes supplied from the fertile imagination of an animal dealer. The fine series of examples of C. carinata in the New York Zoological Park, was, however, a conspicu- ous exception, since they were known to have come from Turks Island. Unfortunately this entire, valuable series seems to have been lost sight of, and a careful search at the American Museum of Natural History failed to reveal a single one. 140 bulletin: museum of comparative zoology. GENERAL CONSIDERATIONS. The members of the genus Cyelura form a small compact group of species confined to the Greater Antillean district of the West Indian region. Related to the Rock Iguanas (Ctenosaura) of the Central American mainland they are nevertheless well set off from the latter by the possession of the peculiar corneous combs or pectinations on the hind toes. Except for this character common to all the West Indian forms, some of these would appear more closely related to some race of Ctenosaura than to another of the island species. On the whole, it does not seem advisable to recognize the genus Metapoceras for the so-called Rhinoceros Iguanas of Navassa, Haiti, and Mona, since they are ob\aously but slightly advanced modifications of such a type as the Jamaican Iguana, which is a true, and probably an- cestral, Cyelura in every respect. The species in the Cayman Islands is nearly related to the Cuban, and the number of forms known from the Bahamas represent two groups of species, one showing affinities with the Cuban, Cyelura macleayi. In the Bahamas, haeo- lopha of Andros Island seems most like macleayi, with its neighbor, inornata, hardly less similar; while nuehalis of Fortune, rileyi of Watlings, and carinata from Turks Island form another well differ- entiated group of races. The latter species has head-scales of a simple and scarcely modified, one might, at first sight, say obviously primitive nature. We imagine, however, that this condition has been reached secondarily, the transition back through some of the other species being clearly traceable. So that while the scales of the head of earinata are of a very simple and undifferentiated character, it is nevertheless extremely improbable, especially in view of its habi- tat, that the species can be considered ancestral or anything more than a reversion to the probably, or possibly, primitive condition for Cyelura, and Ctenosaura, or their progenitors. It does not seem wise to lay much stress upon the distribution of the species of Cyelura as a basis for any zoographic deduction or sm-mise. We know but little of the habits of the species, the whole group is fast disappearing and will soon be wholly extinct, and even now we are able to character- ize but eleven species, probably a comparatively small part of those in existence even two hundred years ago. Early writers often mention Iguanas in the West Indies, and of these some referred to the genus Iguana and some to Cyelura ; among the latter was Catesby. This authority writing upon the Natural BARBOUR AND NOBLE: THE GENUS CYCLURA. 141 History of Carolina and the Bahamas, states in 1743, that Iguanas or Guanas were abundant upon many islands throughout the Bahamas, so common in fact that schooners were cargoed with them and that they were carried to Carolina for food. The name Guana is even now used among the "Conchs" of the Bahamas, who still speak a peculiar archaic English. A vague idea of how wide-spread these great lizards were in early colonial times may be gained from the Bahaman place- names. Thus, there is a Great Guana Cay, off the Abaco coast not far from Green Turtle Cay, a settlement which once had some impor- tance. This islet was visited by the senior author in 1904 but no Guanas were found, and none had persisted to within the memory of the elder folk living in the tiny hamlet. There is also a Guana Cay near Little Harbor about half way up the chain of the Berry Islands, and not far from one of the Bahaman Whale Cays, for this also is a common place-name. Then we find another Great Guana Cay in the Exuma chain of Cays. In all of these islands Guanas are now un- known. On Bitter Guana Cay, however, but a few hundred yards from the Great Guana Cay in Exuma, Mr. C. J. Maynard tells us that up to 1915 a few Cycluras were still to be found. He believes that these represent an undescribed race. As to the status of the other Bahaman species : — baeolopha is still not uncommon, since its habitat, Andros Island, is very large and contains much unsettled and indeed even unexplored territory. Of nuchalis from Fortune Island we know nothing. Stejneger's species, rileyi, is confined to two tiny islets in the saline lagoon of Watling's Island; here Riley obtained the types in 1903 and W. W. Worthington procured a few specimens in March, 1909. Our new species, inornata, is, or was, found upon a little island called U Cay, in Allen's Harbor, north of Highborn Cay and situated in the Ai'chipelago between Exuma and New Providence. Here in 1892 Maynard found the Iguanas not uncommon. He revisited the islet in 1915, was storm bound there and hence had ample opportun- ity to cover it very completely. He found but two Iguanas still living upon U Cay. Both of these he shot ; one, our type, he secured, the other escaped, wounded. Thus the species inornata, which once doubtless existed on several islands about Allen's Harbor, is now beyond doubt extinct. Since these creatures are excellent for food, they are constantly hunted by the native negroes, often with dogs trained for the purpose. These negroes during the course of their sponging and turtling voyages cover the entire Bahaman Archipelago, visiting even the most remote, inaccessible, and infertile cays. There is a constant search for animal food, which unfortunately is by no 142 bulletin: museum of comparative zoology. means abundant or easily obtained by the poor inhabitants of one of Great Britain's most deUghtful but poverty-stricken colonies. Iguanas are often brought from Andros to market in Nassau, upon New Providence Island. One of the authors has seen the creatures for sale there upon several occasions. We have concluded therefore that a magnificent adult male baeolopha in the Academy of Natural Sciences of Philadelphia (Reptile Coll. 8120) probably represents such a specimen, although it is said to have been collected by a Mr. Wilson on New Providence in 1861. Our belief is that the Iguanas disappeared from New Providence long before this date. For the benefit of the herpetologists we should also record that Salt Cay, near Hog Island, opposite the town of Nassau, has been stocked with Iguanas brought from Andros Island. Mr. Chamberlain, the owner of Salt Cay is reported to have stated that they have thriven and appear to have become well established in their new home. Of some of the other species of the genus we know even less than of these we have referred to. Cyclura collei has almost certainly com- pletely disappeared upon the mainland of Jamaica and it was only by the greatest stroke of good fortune that Mr. Arthur Perrin of Cam- bridge who kindly volunteered to make a special excursion for the purpose, was able to secure the specimen, which we describe, from Goat Island not far from Old Harbor off southern Jamaica. Dr. A. G. Mayer tells us that an Iguana was secured a few years ago on one of the cays near Montego Bay, and that he believes a few still exist there. Of the species on Navassa, Haiti, and Mona we know practically nothing. Mr. W. M. Mann who spent some months in Haiti, and who made an excellent collection of reptiles there in 1913 was unable to learn anything of Iguanas and secured none. Mr. Halter from the American Museum, visited Santo Domingo in 1915 and could learn nothing of existing Iguanas. Of the Cyclura on Mona Island we know only that Stejneger quotes Bowdish as say- ing that he got his specimens in 1901, among the rocks. The journeys which the senior author has made on a number of occasions permit us to speak with more authority regarding Cuba. Gundlach in 1880 wrote " Esta especie vive en varios cayos y en las costas de la isla de Cuba y de la isla de Pinos; pero es hoy una especie rara, aunque antiguamente fuese comun y llevada a los merca- dos, siendo su came estimada como excel ente manjar." He goes on to say that it usually lives in burrows in the sand dunes or in sandy places about the coasts where it is easily dug out. Now the Cuban Iguana has with increasing civilization become still more rare and restricted BARBOUR AND NOBLE: THE GENUS CYCLURA. 143 in range. C. T. Ramsden, the accomplished naturahst of Guanta- namo has found a few specimens on the coast cHffs of the extreme eastern end of the island. Wirt Robinson sent one to the M. C. Z. from Santiago in December, 1903. The only very young Cyclura we have seen was one loaned by Ramsden, and obtained by Oskar Tollin during a trip to Belig, near Cabo Cruz, in the summer of 1914. The species is still abundant on the Cayos near Manzanillo and those off the south coast near Santa Cruz del Sur. It is fairly abundant on the Isle of Pines. In 1915 while Prof, de la Torre and his assistant Sr. V. J. Rodriguez were collecting with W. S. Brooks and T. Barbour in the region of Guane, we learned that Iguanas were still not uncommon in the limestone mountains which encircle the glorious valley of Luis Lazo. Here we got two fine adult specimens, one for the Museo Poey in Havana and one for the M. C. Z. Prof, de la Torre says that Iguanas are also fairly common on the Pan de Guajaibon and he writes us that he has recently seen one near Baracoa. They persist as well on many of the small and remote cayos of the north coast. Curiously enough in spite of what Gundlach says — no one appears to eat Iguanas in Cuba at the present time. Ramsden has also observed this and writes me that he has been told that Cubans believe the Iguana to be very poisonous. When hung up by the tail, they say, a baba or burujo, as they call it, black drivel or vomit, runs from its mouth. This is supposed to be deadly. The black vomit due to blood in the stomach, which marks the final stage of a fatal case of yellow fever, is also called in Cuba burujo, a name also used for the grounds of coffee, and it may be that some imagined similarity between these burujos, coupled with forgetfulness which increased as Iguanas grew rare, has now spread the idea that Iguanas are unfit for food, when once, in Cuba as elsewhere, they were eagerly sought after. It will be noticed that some old, long standing specific names have been dropped. The reason for this is as follows: — the name nuhila was based upon a young specimen without locality. The descrip- tion given by Gray is worthless. The type is mentioned in Boulenger's Catalogue and hence doubtless is still in existence. When it is examined it will probably be possible to determine whether the name supercedes one used here or whether it represents another distinct species. It is impossible to determine this. Cope attached the name nuhila to a specimen which he said was from Cat Island and was U. S. N. M. 14576; but Stejneger tells us that this number is borne by a specimen of Leiocephalus and that there is now no Cat Island Iguana in the U. S. N. M. and no evidence that there ever was one. Cuvier 144 bulletin: museum of comparative zoology. first used cyclura as a specific name for what he called L'Iguane de la Caroline. After what Catesby said we imagine that he had a Bahaman specimen which had been carried to Carolina and had probably been sent to Paris from there. Which Bahaman species he had it is impos- sible to decide from his meagre descriptions. So unless the t;y^e is still in existence and sufficiently well preserved, which is improbable, it will not be possible to more than surmise that Cuvier probably had a specimen of C. baeolopha. Stejneger and Barbour have both used the name Cyclura cyclura Cuv. for the Cuban Iguana and this might be considered as restricting the name. It is probably better to drop the name altogether until someone studies this old type, and also the type of nubila in the British Museum. , KEY TO THE SPECIES. Median frontal shield enlarged and tubercular. fei Nasals broadly in contact with the rostral stejnegeri. b^ Nasals separated from the rostral by small scales, or granules. c^ Posterior prefrontals separated from the frontal shield by two rows of scales nigerrima. >%: .»>:, '.-^V GEORGE NELSON. PHOTO. PLATE^e. Barbour and Noble. — The Genus Gyclura. PLATE 6. Cyclura baeolopha Cope. Fig. 1.— Side view of head of adult female. M. C. Z. 5960. Fig. 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.— Cyclura, Plate 6 GEORGE NELSON, PlIQTO. PLATE 7. Barbour and Nobi«. — The Genus Cyclura. PLATE 7. Cyclura rileyi Stejneger. Fig. 1.— Side view of head of adult male. M. C. Z. 10918. Watling's Island, Bahamas. Fig- 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.— Cyclura, Plate 7 :^^^,. GEORGE NELSON, PHOTO. PLATE 8. Harbour and Noble. — The Genua Cyclura. PLATE 8. Cyclura nuchalis Barbour & Noble. Fig. 1. — Side view of head. Type, Collection Acad. nat. sci. Phil., 11985, Fortune Island, Bahamas. Fig. 2. — Upper view of head of the same specimen. Cyclura carinata Harlan. Fig. 3.— Side view of head of adult male. M. C. Z. 1252. Turk's Island, Southern Bahamas. Fig. 4. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.— Cyclura, Plate 8 GEORGE NELSON, PHOTO. PLATE 9. Barbour and Noble. — The Genus Cychira. PLATE 9. Cyclura collei Gray. Fig. 1. — Side view of head of adult male. M. C. Z. 9397. Goat Island, near Old Harbour, Jamaica. Fig. 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.— Cyclura, Plate 9 GEORGE NELSON, PHOTO. PLATE 10. Barbour and Noble. — The Genus Cyclura. PLATE 10. Cyclura cornuta (Bonnaterre) . Fig. 1. — Side view of head of half grown specimen. M. C. Z. 3597A. Jere- mie, Haiti. Fig. 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour a>jd Noble— Cyclura, Plate 10 ^-^ --'*^' GEORGE NELSON, PHOTO. PLATE 11. Barbour and I\oble. — The Genus Gyclura PLATE 11. Cydura nigerrima Cope. Fig. 1.— Side view of head of adult male. Cotype? M. C. Z. 4717. Na- vassa Island. Fig. 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble— Cyclura, Plate 11 V' \ GEORGE NELSON. PHOTO Babboub and Noble. — The Genus (lycliira. PLATE 12. Cydura stejnegeri Barbour & Noble. Fig. 1. — Side view of head of adult male. Paratype M. C. Z. 11145. Mona Island. Fig. 2. — Upper view of head of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.-Cyclura, Plate 12 -*>: GEORGE NELSON. PHOTO PLATE 13. Barboub and Noble. — The Genus Gyclura. PLATE 13. Cyclura baeolopha Cope. Fig. 1.— Segment of taH. M. C. Z. 5960. Cf. Plate 6, fig. 1. Fig. 2. — Foot of the same specimen. Cyclura carinata Harlan. Fig. 3.— Segment of tail. M. C. Z. 1252. C/. Plate 8, fig. 3. Fig. 4. — Foot of the same specimen. Cyclura viacleayi Gray. Fig. 5.— Segment of taU. M. C. Z. 11050. Cf. Plate 2, fig. 1. Fig. 6. — Foot of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.— Cyclura, Plate 13 > KSS mfi J'v 4Uttr E. N. FISCHER DEL. PLATE 14. Barbour and Nobla. — The Genus Gyclura. PLATE 14. Cyclura inornata Barbour & Noble. Fig. 1.— Upper view of head. Type M. C. Z. 11062. U Cay in Allen's Harbor, near Highborn Cay, Bahamas. Fig. 2. — Side view of head of the same specimen. Fig. 3. — Segment of tail of the same specimen. Fig. 4. — - Foot of the same specimen. BULL. MUS. COMP. 200L. Barbour and Noble —Cyclura, Plate 14 E. N. FISCHER DEL. PLATE 15. Barbour and Noble. — The Genus Cyclura. PLATE 15. Cyclura nigerrima Cope. Fig. 1.— Segment of tail. M. C. Z. 4717. Navassa Island. Fig. 2. — Foot of the same specimen. Cyclura rileyi Stejneger. Fig. 3.— Segment of tail. M. C. Z. 1091S. Watlings Island. Fig. 4. — Foot of the same specimen. Cyclura collet Gray. Fig. 5.— Segment of tail. M. C. Z. 9397. Goat Island, near Old Harbour Jamaica. Fig. 6. — Foot of the same specimen. BULL. MUS. COMP. ZOOL. Barbour and Noble.-Cyclura, Plate 15 .^ "T ^ /Xw-^Iil^^*^^ 2 J?^^ E. N. FISCHER DEL. fLD ^ i-i. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 5. THE ANTS OF THE PHILLIPS EXPEDITION TO PALESTINE DURING 1914. By W. M. Wheeler and W. M. Mann. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM, February, 1916. No. 5. — The Ants of the Phillips Expedition to Palestine during 1914- CONTRIBUTIONS FROM THE ENTOMOLOGICAL LABORATORY OF THE BUSSEY INSTITUTION, HARVARD UNIVERSITY, NO. 107. By W. M. Wheeler and W. M. Mann. The junior author, while -accompanying Dr. John C. Phillips on a recent zoological expedition to Palestine and the adjacent countries for the Museum of Comparative Zoology, succeeded in amassing quite a collection of ants. As many collections of these insects have been made from time to time in Egypt and Asia Minor and have been care- fully described in numerous papers by Ern. Andre, Emery, Forel, Mayr, and Ruzsky, it seemed improbable that another collection would contain anything new. After the specimens were mounted and examined, however, we were surprised to find among them a new and peculiar species of Deromyrma and a few undescribed varieties and subspecies of well-known Mediterranean species. We decided, therefore, to publish a list of all the forms collected, together with such field-notes as seemed interesting. FORMICIDiE. 1. Ponera eduardi Forel. ^ 9 (ergatoid).— Baniyas, Syria. 2. Sima bifovcolata Mayr var. syriaca, var. nov. Worker. Agreeing very closely with the typical form described from Delagoa Bay and Zanzibar, except in the following characters: — the tibiee have no suberect hairs, the mandibles have only three instead of four or five teeth, and the petiolar node is semicircular in profile. The tip of the gaster is not brown. The peculiar paired granular pits on the occiput seem to be quite as distinctly developed as in the type. Several workers found running on plants at Wady Gazelle, Sinai Peninsula. The typical form has been recorded by Mayr only as far north as the White Nile where it was taken by Tragardh. 3. Aphaenogaster splendida ^oger. y — Rasheya, Syria; in damp, shady places. 168 bulletin: museum of comparative zoology. 4. Aphaenogaster {Deromyrma) -phillipsi, sp. nov. (Fig. 1). Worker. Length 6-6.8 mm. Very slender. Head twice as long as broad, with the rather small but convex eyes well in front of the middle of its sides; cheeks sub- parallel, distinctly convex, postocular borders of head straight and gradually converging to the occipital border, which has a strongly reflexed margin and is only about half as broad as the distance between the eyes. In profile the upper surface of the head is convex anteriorly, the gular surface flat. Mandibles with slightly concave external borders, three stout apical teeth and several basal denticles. Clypeus flat and subcarinate, the middle of its anterior border broadly and Fig. 1. Aphaenogaster {Deromyrma) phillipsi, sp. nov. Body of ijporker in profile; head of sEtme from above. sinuately excised. Frontal area impressed, distinct, with a median cannula. Frontal carinae approximated and parallel behind. An- tennae very slender, the scapes surpassing the occipitaF margin by about § their length; funiculus without a club, its joints very slender, the second, third, and terminal longest. Thorax slender, pro- and mesonotum subequal, the former in profile feebly convex, the latter sloping, slightly concave in the middle and with a small but distinct convexity in front just behind the promesonotal suture. Mesoepinotal constriction abrupt, short and moderately deep. Epinotum in profile about 1| times as long as high, its base convex in profile and twice as WHEELER AND MANN: ANTS OF PALESTINE. 169 long as the concave declivity, armed with two small, rather acute teeth, which are directed upward and slightly backward and outward. Petiole more than twice as long as broad, in profile with a rather low, rounded node. Postpetiole about 1| times as long as broad, less than half again as broad as the petiole, in profile with a similar but some- what larger node. Gaster elongate elliptical, narrowed in front. Legs very long and slender; spurs of the posterior tibiae short but distinct. Gaster very smooth and shining, remainder of body more feebly shining. Mandibles subopaque, finely and densely striate; head, thorax, petiole, and postpetiole shagreened, the meso- and epinotum opaque, rugulose-punctate; the anterior portion of the head above, including the antennal foveae and excluding the clypeus, longitudi- nally rugose, becoming reticulately rugulose and punctate posteriorly ; pronotum and upper surfaces of petiolar and postpetiolar nodes smoother and more shining. Epinotal declivity shining, feebly and transversely rugose. Hairs yellow, very short, blunt, sparse on the body, entirely lacking on the legs; very short, but distinct and oblique on the antennal scapes, especially towards their tips. Pubescence absent. Head, thorax, petiole, postpetiole, and antennae pale ferruginous; legs brownish yellow; gaster clear, pale yellow, with the posterior I of the first segment dark brown. Described from eleven workers from Petra, Palestine. These were found in the early morning eating portions of the bait with which small mammal traps had been baited. This species differs considerably from either of the two previously described Palaearctic species of Deromyrma, cecconii Emery from Crete and rhaphidiiceps Mayr from Turkestan. The Cretan species is smaller (5.7 mm.), has the body black, the tibiae with oblique hairs, the petiolar node is angular in profile, the epinotal teeth are longer, the head is shorter and of a different shape behind and the antennal funiculi have an indistinct, 4-jointed clava. In coloration and the shape of the head phillipsi resembles rhaphidiiceps, but the latter is smaller, the occipital margin of the head has no reflected margin, the clypeal border is entire, the first funicular joint is longer than the second and the tibiae are hairy. 5. Messor rufotestaceus Foerster. U . — Wady Gharandel, Sinai Peninsula and Petra, Palestine; in the former locality living in crater nests, in the latter under stones and more abundant. 170 bulletin: museum of comparative zoology. 6. Mcssor barbanis Linne sviljsp. strudor Latr. var. oricnialis Emery. y . — Rashe\'a, Animik, and Baruk, Syria; Ain Gleidat and Wady Hisa/ Palestine. 7. Mcssor harharus Linne subsp. semirufus Ern. Andre. ^ . — t ' Ammik, Hasbeiya, Zahleh, and Baniyas, Syria; Wady Hisa, ^ Palestine. One of the commonest ants in Syria, in crater nests. 8. Mcssor harharus Linne subsp. semirufus Ern. Andre var. concolor Emery. ^ . — Shiba, Syria; Fuweila, Arabia; Wady Feran, Sinai Peninsula; Wady Hisa, Palestine. 9. Messor harharus Linne subsp. mcridionalis Ern. Andre. ^ . — Petra and Wady Mojeb, Palestine; Shiba and Wady El Katana, Syria. 10. Messor harharus Linne subsp. acgyptiacus Emery. ^ 9 cf • — W'ady Feran, Wady Gazelle and Mt. Sinai, Sinai Peninsula; Cairo (Mann) and Fayum (Wm. Granger), Egypt. 1 1 . Phcidolc paUidula Nyl. Ql y . — Petra and Wady Kerak, Palestine ; Ammik, Syria; Wady Feran and Wady Gazelle, Sinai Peninsula. 12. Phcidolc incgacephala Fabr. 2i y . — Baruk, Syria. 13. Phcidolc sinaitica Mayr. % y . — Cairo, Egypt. 14. Crematogastcr scutcllaris Oliv. subsp. schmitti Mayr var. ionia Forel. y . — Rasheya, Syria; Petra, Palestine. We refer these specimens to Forel's variety on account of the distinct infuscation of the head and thorax. 15. Crematogaster auherti Emery subsp. jehovae Forel. ^ . — W^ady Kerak and Ain Gleidat, Palestine; Shiba, Syria. 16. Crematogastcr auherti Emery subsp. antaris Emery. ^ . — Mt. Sinai, Sinai Peninsula; Fuweila, Arabia. 17. Crematogastcr incrmis Mayr. y. — - Mt. Sinai, Wady Gazelle, and W'ady Feran, Sinai Peninsula; Wady Mojeb, Palestine. This is the commonest species of the genus in the Sinaitic Penin- sula. IS. Crematogaster lortcti Forel. y .— Ain Gleidat, Palestine. Many colonies, nesting under stones in moist localities. 19. Monomorium vcnustum F. Smith subsp. niloticum Emery, y . — Wady Gazelle, Sinai Peninsula. 20. Monomorium solomonis Linne. y . — Fuweila, Arabia; Wady Gharandel, Sinai Peninsula; Wady Mojeb, Palestine. 21. Monomorium solomonis Linne subsp. suhopacum F. Smith var. Phoenicia ^jinery. y 9 cf. — Akaba, Arabia; Petra, Palestine. 22. Monomorium hicolor Emery subsp. nitidiventre Emery, y 9 . — Cairo, Egypt. WHEELER AND MANN: ANTS OF PALESTINE. 171 The female (dealated) measures 3.8-4 mm. and resembles the worker in color and sculpture, except that the base of the gaster above is yellowish red like the head, thorax, and pedicel and more opaque. Head longer than broad, with straight, subparallel sides, distinctly excised posterior border and rather angular posterior corners. Thorax elongate elliptical, fully 2| times as long as broad ; in profile the dorsal surface of the mesonotum, praescutellum and scutellum form a straight line; epinotum with a pronounced median longitudinal impression. Postpetiole not broader than the petiole, distinctly broader than long. 23. Monoviorium abeUlei Ern. Andre. ^ . — Wady Feran and Wady Gazelle, Sinai Peninsula; Petra, Palestine. 24. Monomorium {Holcomyrmex) dentigerum Roger. ^ . — Petra, Palestine. 25. Monomorium {Holcomyrmex) dentigerum Roger var. haal, var. nov. Worker. Differing from the typical form in its decidedly darker color, the body and antennal scapes being castaneous or blackish brown; the discs of the mandibles, the clypeus, front, cheeks, mesono- tum, pleurae, and lower portions of the petiole and postpetiole deep red; the legs and tips of antennal scapes yellowish brown. Numerous specimens from Shiba, Syria and Wady Kerak, Palestine. 26. Leptothorax tuberum Fabr. var. luteus Forel. Three workers from Ain Gleidat, Palestine agree closely with the description of this form except that they have the posterior half of the first gastric segment and the whole of the remaining segments pale brown instead of yellow, like the remainder of the body. We deem it inadvisable to describe this form as a new variety on the basis of so little material. 27. Tetramorium striativentre Mayr. ^ . — Wady El Katana, Syria; Wady Mojeb, Palestine. 28. Tetramorium caespitum Linne. ^ . — Petra, Palestine. 29. Tetramorium caespitum Linne var. forte Forel. ^ . — Baruk, Syria. 30. Tetramorium caespitum Linne var. schmitti Forel. S . — Baruk and Ammik, Syria; Wady Mojeb, Palestine; Mt. Sinai, Sinai Peninsula. 172 bulletin: museum of compakative zoology. 31. Tetramormm caespitum -Linne subsp. punicum F. Smith var. lucidulum 'Emery. ^. — Petra, Palestine; Ammik and Baniyas, Syria; Mt. Sinai, Sinai Peninsula. The colonies at Petra were very populous and were nesting under stones. 32. Tetramorium caespitum Linne subsp. judas, subsp. nov. Worker. Length 2.3-3.5 mm. Allied to the subspecies semilaeve Ern. Andre but the whole body, except the mandibles and clypeus, shining and the sculpture very feeble. Head much as in semilaeve, with the sides, posterior corners and a streak between the front and the posterior corners smooth and shining, the rugae on the front delicate and numerous, continued nearly or quite to the occiput. Pro- and mesonotum smooth and shining, with only traces of rugae at the sides, epinotum subopaque and rugose; pleurae more or less rugose as are also the sides of the petiole and postpetiole, the summits of the nodes of the latter shining, nearly smooth, or merely indistinctly punctate-rugulose. Gaster smooth and shining throughout. Color dark brown; mandibles, cly- peus, cheeks, antennae, and legs testaceous. Nine specimens from Wady Mojeb, Palestine. This form seems to be near the var. splendens Ruzsky of the subsp. semilaeve, but we infer that the thorax is more strongly sculptured than the head in this form. 33. Bothriomyrmex meridionalis Roger var. syria Forel. ^ . — Ain Gleidat, Palestine; Rasheya and Wady El Katana, Syria; Wady Gazelle, Sinai Peninsula. 34. Tapinoma erraticum Latr. ^. — Petra, Palestine; Baruk, Syria. 35. Tapinoma erraticum Latr. subsp. nigerrimum Nyl. y . — Petra, Palestine; Fuweila, Arabia. 37. Acantholepis frauenfeldi Mayr. ^ 9 . — Wady Hisa, Palestine. 38. Acantholepis frauenfeldi M.a,yT Ya.r. bipartita F. Smith. ^ 9. — Rasheya and Wady El Katana, Syria. 39. Acantholepis carbonaria Emery. ^ . — Wady Gazelle, Sinai Peninsula. 40. Acantholepis capensis Mayr var. canescens Emery. ^ . — Two workers from Fuweila, Arabia agree very closely with a single specimen of this ant from Erythraea in the senior author's col- lection. This variety is also recorded from Kaka on the White Nile and from Bogosland and Somaliland. WHEELEfl vVND . MANN : ANTS OF PALESTINE. 173 41. Plagiolepis pygmaea Latr. ^ 9 . — Ain Gleidat and Petra, Palestine; Ammik, Syria; Wady Feran, Sinai Peninsula. 42. Prenolepis {Nylanderia) jacgerskjoeldi Mayr. y . — Cairo, Egypt; Wady Kerak, Palestine. 43. Formica rufibarhis Fabr. var. clarorufibarbis Ruzsky. ^ . — Baruk, Syria. The specimens have the base of the gaster red as in the var. clara Forel, but the top of the head is infuscated as in the typical rufibarhis. 44. Cataglyphis bombycina Roger. "^ g . — Lake Fayum, Egypt (Wm. Granger). 45. Cat(iglyphis bombycina Roger var. sinaitica, var. nov. Soldier and Worker. Differing from the typical form in the much darker coloration, the occiput, thorax, petiole, gaster, and femora being deep castaneous brown or even blackish, the knees, tibiae, head, antennal scapes, and first funicular joint and in some specimens also the thoracic dorsum, paler brown; the mandibles in the worker yellowish red. Antennal funiculi beyond the first joint and the teeth of the mandibles black. The hairs on the tibiae are distinctly longer than in the typical form and the silver pubescence, especially on the gaster, is even denser. A single soldier and numerous workers from Wady Gazelle, Sinai Peninsula. 46. Cataglyphis albicans Roger subsp. livida Ern. Andre. S 9 . — Petra, Palestine; Wady Gharandel, Sinai. 47. Cataglyphis viatica Fabr. subsp. bicolor Ern. Andre. ^ . — Baruk and Ammik, Syria; Wady Mojeb, Palestine. 48. Cataglyphis viatica Fabr. subsp. bicolor Ern. Andre var. nigra Ern. Andre, y .— Cairo, Egypt; Lake Fayum, Egypt (Wm. Granger)., 49. Cataglyphis viatica Fabr. subsp. bicolor Emery var. orientalis Forel. y .— Wady Mojeb and Wady Hisa, Palestine; El Katana, Syria; Wady Feran and Wady Gharandel, Sinai Peninsula. 50. Cataglyphis cursor Fonsc. subsp. aenescens Nyl. S . — Shiba, Syria. 51. Camponotus (Myrmoturba) macidatus Fabr. subsp. thoracicus Fabr. var. oasium Forel. y. — Ammik, Syria; Wady Feran, Sinai Peninsula. 52. Camponotus {Myrmoturba) maculatus Fabr. subsp. thoracicus Fabr. var. cypriacus Forel. ^ 9 . — Wady Gharandel, Sinai Peninsula. 174 bulletin: museum of comparative zoology. 53. Camponotus {Myrmoturha) maculatus Fabr. subsp. thoracicus Fabr. var. xerxes Forel. y . — Zahleh, Syria. 54. Camponotus {Myrmoturha) viaculatm Fabr. subsp. thoracicus Fabr. var. sanctoides Forel. ^ — Wady Feran and Mt. Sinai, Sinai Peninsula. 55. Camponotus (Myrmoturha) maculatus Fabr. subsp. thoracicus Fabr. var. mortis Forel. y . — Wady Feran, Sinai Peninsula. 56. Camponotus (Myrmoturha) maculatus Fabr. subsp. sanctus Forel. S. — Shiba, Bakeyas, El Katana, and Rasheya, Syria; Petra, Palestine. 57. Camponotus (Myrmoturha) maculatus Fabr. subsp. turkestanicus Ern. Andre. ^ . — Wady Hisa, Palestine. 58. Camponotus (Myrmoturha) maculatus Fabr. subsp. haldaccii Emery. ^ . — Baniyas, Syria. 59. Camponotus (Myrmoturha) maculatus Fabr. subsp. aethiops Fabr. var. concavus Forel. g . — Shiba, Syria; Mt. Herraon, Palestine. 60. Camponotus (Orthonotomyrmex) lateralis Oliv. var. atricolor Nyl. ^ . — Rasheya and Ammik, Syria. 61. Camponotus (Orthonotomyrmex) interjectus Wsi\T. ^. — -Wady Kerak, Palestine. 62. Polyrhachis (Myrmhopla) simplex Mayr. y 9 cf . — Wady Kerak, Palestine; Wady Feran, Sinai Peninsula. This species was very abundant in both of these localities, always in damp places and always associated with a certain tree. In Wady Feran several nests were seen on plants. These were made of portions of leaves and twigs fastened together with films of silk. In Wady El Katana, near the Dead Sea, the only nest found was beneath a stone at the base of a tree. It contained many larvae, some of which were lying on the ground, and others on a sheet of silk. On nearly every tree in the vicinity of this nest there were many Membracidae which were constantly attended by the Polyrhachis workers and in most cases even sheltered in sheds' constructed by the ants. The workers were observed while carrying the larvae up the trees and using them to spin the silk of the sheds. MAR 23 ]m Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. Vol. LX. No. 6. RESULTS OF THE YALE PERUVIAN EXPEDITION OF IQU. THE ARACHNIDA. By Ralph V. Ch.\mberlin. With Twentt-pive Plates. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM. March, 1916. No. 6. — Results of the Yale Peruvian Expedition of 1911. — The , Arachnida. By Ralph V. Chamberlin. The arachnids, upon a study of which this paper is based, were collected for the most part by Prof. H. W. Foote while a member of the Yale Peruvian Expedition of 1911. The collection has proved to be extraordinarily interesting, presenting a very large proportion of previously undescribed forms among which are eighty -two new species and twelve new genera. A few specimens were secured at Panama; but aside from these, the material all comes from localities in a little known section of southeastern Peru at elevations ranging from 3,000 to 11,500 feet above sea level. Comparatively few of the species seem to be identical with forms recorded from more northerly locali- ties of Peru by Taczanowski in his Les Araneides du Peru (Bull. Soc. nat. Mosc. 1878, 53, p. 278-374; Horae Soc. entom. Ross., 1878, 14. p. 140-175; 1879, 15, p. 102-136). Some of the localities mentioned are not on published maps. This collection of arachnids was turned over to the author for study through the kindness of Professor Foote, who has also consented that the entire lot remain the property of the Museum of Comparative Zoology. Prof. A. Petrunkevitch of Yale had planned to report upon the collection but the press of other duties prevented. He had made notes and drawings of several species which were kindly sent to me. The principal Peruvian localities from which the arachnids of the collection come with elevations and months during which specimens were secured are as follows : — Arequipa, 7,600 feet June Cuzco, 11,500 feet July Huadquina, 5,000 feet July Ollantaytanibo, 9,000 feet July Torontoy, 8,000 feet July San Miguel, 6,000 feet July Urubamba, 9,500 feet July Lucma, 7,000 feet August Paltaybamba, 5,000 feet August 178 bulletin: museum of comparative zoology. Santa Ana, 3,000 feet August Tineoehaca, 7,000 feet August Vilcabamba, 10,000 feet August Conservidayo River, —— August Sorontoy, 7,000 feet September The specimens from Panama were collected in June. SCORPIONIDA. BUTHIDAE. TiTYUs FOOTEi, sp. nov. Plate 1, fig. 1-4. Carapace and dorsum of preabdomen from testaceous to dark reddish with blackish markings near eyes and along the anterior border of the carapace, and especially across the caudal borders of the abdo- men, the dark bands widest at middle. Legs and pedipalps yellowish. Cauda proximally from yellow to testaceous of reddish cast becoming decidedly darker distad, the last one or two segments usually black or nearly so ; sting, in individuals in full color, more or less chestnut. Carapace and all tergites of preabdomen strongly granular. Pre- abdominal sternites each roughened with minute granules. All seg- ments of Cauda granular, the fifth most strongly so. All tergites of preabdomen with median dorsal keel well developed, that of the fifth incomplete behind, and with two well-developed dorsal keels each side of it; sternite of fourth and fifth segments with five keels of which the median is very weak. First segment of cauda with ten keels, a complete median lateral ridge being present between the usual upper and lower lateral ridges ; second segment with keels the same except- ing that the median lateral is developed only caudad; third and fourth segments w^ith the eight ridges, two median dorsal and median ventral and the two upper and two lower laterals; fifth segment with but five distinct keels, the odd one being a complete median ventral; all keels granular. Basal portion of sting finely granular. A well-developed acute spine beneath, (Plate 1, fig. 1). Ca^MBERLIN: THE ARACHNIDA. 179 Legs and pedipalps with strongly developed longitudinal granular ridges. Hand of pedipalp broader than tibia, its ridges strongly developed, these continuing upon the immovable finger where they become smooth. Finger with fifteen or sixteen oblique lines of granules, (Plate 1, fig. 2) ; not fully twice as long as proximal hand (cir. 7:4). Comb with 20-22 teeth. Basal middle lamella enlarged, Plate 1, fig. 4. Length of types 38 to 52 mm. Cauda of male proportionately longer and more slender than in the female. (Fourth segment of cauda in female 2.5 mm. wide X 5.0 mm. long, in male 2.5 mm. wide X 5.8 mm. long). Localities. — Huadquina, 5,000 feet, July 26. (Type, M. C. Z., 121; paratypes, M. C. Z. 122, five specimens.) Ollantaytambo 9,000 feet, July 20. (M. C. Z. 123, one young specimen). Named for Prof. H. W. Foote of Yale University. This species is apparently nearest to T. bahiensis (Perty). In coloration it differs in having the tibia of the pedipalp light and uni- form, not conspicuously darkened in the way characteristic of bahien- sis, and in having the fifth segment of the cauda conspicuously dark. The hand of the pedipalp is conspicuously more slender and its ridges are much more strongly developed and those along the immovable finger continue without break to the proximal end of the hand. The basal middle lamella of the comb is relatively much larger, being more inflated, and with its inner edge semicircular instead of straight. The spine on under side of sting is more slender and acute with at most only a single and relatively small granule on its upper edge. BOTHRIURIDAE. Brachistosternus andinus, sp. nov. Plate 1, fig. 5-8. General color yellow, the carapace and tergites of the preabdomen darkened with brown and blackish markings; carapace free from the darker color in a triangular area in front of eyes and in a narrower area behind them ; tergites of preabdomen lighter along caudal border and in a narrow median longitudinal line; postabdomen above pale 180 bulletin: mitsefm of comparative zoology. yellow, below darker, being mottled with brown which is most abun- dant on the fifth segment and is partially absent from the first. Carapace and first six segments of the abdomen smooth and shin- ing; seventh abdominal segment finely granular, (Plate 1, fig. o). First segment of postabdomen equal in length and breadth, the second a little longer than wide. Dorsal and upper lateral keels of first segment of cauda finely granular, the granules on those of other segments fewer and weaker or essentially absent. Ventral median keel of Cauda absent from first four segments, weakly developed on the fifth; lower lateral keels of fifth segment conspicuous, strongly granular, the lower lateral keels of the other segments obsolete. Ventral surface of the fifth segment conspicuously granular excepting at anterior end, that of the fourth segment with scattered granules, the venter elsewhere smooth. Marginal keels of femur of pedipalp smooth; tibia along posterior ventral edge with trichobothria eight in number, these arranged in two rows of four each; hand of chela thicker than the tibia, smooth, with a row of 8-10 trichobothria under the outer ridge. The lateral granules on the mesal surface of the movable finger remaining apart and distinct from the main row over entire length, eight in number in each row, (Plate 1, fig. 6). The distal lamella of comb abruptly much narrower than the proxi- mal ones so that the anterior edge appears indented at its l)eginning as in B. ehrenhcrqi. Teeth of each comb 28 in number, (Plate 1, fig. 8). Length 34 mm. Locality— Ollantaytambo 9,000 feet, July 20. (T\pe, M. C. Z. 124, one gravid female). SOLPUGIDA SOLPUGIDAE. MuMMUCiA VARiEGATA (Gervais). Caleodes variegata Gervais, Gay Hist. Chile. ZooL, 1849, 4, p. 15, t. 1, f. 2. Mummuda variegata Simon, Ann. Ent. soc. France, 1879, ser. 5, 9, p. 151, t. 3, f. 29, .30. Loco%.— Ollantaytambo, 9,000 feet, July 20. (M. C. Z. 125, one female). CHAMBERLIX: THE ARACHNIDA. 181 PHALANGIDA. GOXOLEPTIDAE. GONOLEPTES ENOPLUS/ Sp. nOV. Plate 2, fig. 7-8; Plate 3, fig. 1-5. Main portion of carapace very dilute ferruginous, its caudal and caudolateral borders dusk\' but integument along edge of these parts white; area in front of cervical furrow deeply mottled with black; tergite of abdomen blackish with caudal border lighter. Legs black, but the distal articles of tarsus III conspicuously light, whitish, and the metatarsus and especially the tarsus of leg IV also somewhat lighter. Coxa IV dilute ferruginous black across distal end both below and above excepting the white line along the edge, its process also black excepting the paler tips. Coxae of legs I to III densely mottled with black excepting niesal ends which are paler. Sternites of abdomen blackish excepting the paler caudal borders and the white integument adjoining the caudal edges. Body very wide toward caudal end of cephidothorax from where very stronglj' and abruptly narrowed cephalad to opposite bases of thirfl legs, then more gently narrowing to anterior end; also con- spicuousl}^ narrowing caudad to end of abdomen; the greatest width of carapace ecfual to body length, (Plate 2, fig. 7). Carapace with only four complete transverse sulci, but the first of these with two branches on each side; first sulcus lying in a deeper cer\ical furrow, as usual angularly bent caudad at middle; second sulcus moderately angularly bent forward at middle; third sulcus with ends bent abruptly caudad and then running sul)parallel with body axis; fourth sulcus angularly bent forward at middle; all sulcj connected by a median longitudinal sulcus which is deepest between the first two. Eye-tubercle sharply set off and elevated, wider than long, the two cones near its caudal border not especially high. An- terior border of carapace conspicuously elevated as usual, the elevated rim much widest at middle. Lower frontal margin extended lietween the notches for mandibles and laterad of each in an acute spine-like process. Upper frontal margin at each lateral corner bearing several low teeth. Lateral border of carapace behind conspicuously elevated ' efowXoi, armed. 182 bulletin: museum of comparative zoology. as usual, the ridge largely composed of a continuous row of tubercles which, large behind, decrease cephalad and disappear caudad of level of first sulcus, the row of tubercles continued also across caudal border. A row of two or three tubercles in a second low ridge occurs just mesad of the outer rim a little caudad of widest part of carapace. The cones, immediately in front of last sulcus and near middle line, are rather low, distally rounded. Area between last two sulci with numerous larger and smaller tubercles or granules, the area between the second and third w4th fewer and mostly very small granules and the area between the first and second with still fewer scattered and inconspicuous granules, (Plate 2, fig. 7). First three tergites of abdomen with a continuous transverse row of conspicuous tubercles. Anal scutum without processes; rather ob- scurely granular. First three pairs of coxae proximally parallel, the first two bending moderately cephalad from near middle of length, the third transverse for whole length ; second coxae scarcely one fifth longer than the third ; first coxae along midventral surface with a row of distinct granules, the second with a few almost obsolete granules and the third smooth. Fourth coxae very strongly enlarged and directed ectocaudad as usual; terminating on dorsoectal side in a stout process which has a shorter prong on the ventral side beyond middle of length; on mesal side of distal end with a shorter stout process which is toothed on its ectal side. Spiracles moderate, distinct. First article of mandibles strongly narrowed toward base; distal half with a strongly and abruptly elevated hump on dorsal surface. Distal article with fewer hairs in front and distad, (Plate 2, fig. 8). Pedipalps when extended shorter than carapace; not crossed. Trochanter roundly elevated and smooth above; ventrally with a single conical tubercle bearing distally a bristle. Femur a little complanate beneath and bearing few low granules from which hairs arise; dorsally also with scattered low granule-like elevations at bases of some of hairs. Patella smooth. Tibia not compressed; along mesoventral edge with two similar but stouter spines with proximal portions thicker as usual. Tarsus along mesoventral edge with two subequal spines of usual type and along ectoventral edge with two similar large spines and between these a much smaller spine and a similar one between the distal large spine and the base of the claw. Claw much shorter than tarsus, conspicuouslv curved, (Plate 3, fig. 1,2). chamberlin: the arachnida. 183 Femora of first two pairs of legs finely and conspicuously granular; femora of third legs more coarsely granular, the granules on ventral sur- face toward distal end largest; patella with a few granules; tibia with finer granules above but ventrall}' with larger distinct tubercles or teeth. Fourth leg with metatarsi abruptly more slender than tibia; trochanter IV on dorsoectal surface toward base with a stout, distally truncate process and on opposite side with two larger, distally acumi- nate processes; femur conspicuously curving dorsomesad, with nu- merous stout processes of differing lengths of which one from the dorsal surface, one opposite on ventral surface and one toward distal end from mesoventral surface are much the largest (Plate 3, fig. 4, 5) ; patella with numerous small seriate tubercles of uniform size; tibia above and laterally with setigerous tubercles like those of patella but ventrally with a median-longitudinal series of 7-9 long processes which decrease in length distad; metatarsus with a series of low setigerous tubercles. Tarsus I with five joints of which three are in the distal division; II with eight joints of which three are in the distal division; III with five joints of which the most distal is greatly enlarged and bears a setigerous process on its anterior end projecting between the claws, the ventral surface of articles densely clothed with short fine hairs, (Plate 3, fig. 3) ; IV with six joints. Length 8 mm. ; greatest width of carapace also 8 mm. Length of leg 111 mm.; of leg II 20 mm.; of leg III 16 mm.; of leg IV cir. 21+ mm. Locality — San Miguel, 6,000 feet, July 24. (Type, M. C. Z. 126, one male). GONOLEPTES HUADQUINAE, sp. nOV. Plate 4, fig. 3-8. General color above brownish grey, the head region darker and the anal scutum blackish; a series of lighter circular spots across dorsum of carapace and abdomen, one over and about each tubercle. Coxae of first three pairs of legs beneath black, the coxae of the fourth pair subtestaceous. Sternite of abdomen blackish with lighter transverse borders and spots. Trochanters of legs above and below brownish yellow; other joints of legs blackish but femora IV testaceous with fewer dusky mottlings; femora I to III paler at distal end with an obscure narrow light annulus near middle ; tibiae all with an obscure 184 bulletin: museum of comparative zoology. median pale annulus; metatarsi also with faint median annulus and metatarsi III and IV with a more distinct light ring at distal end; tarsi III and IV with distal article pale and also with a siihmedian pale annulus. Carapace broadest at level of third sulcus from where the body narrows conspicuously caudad, the comparati\ely long aljdomcn being more abruptly narrowed or constricted at third segment with the anal scutum narrowly rounded; cephalad the cephalothorax is abruptl\' narrowed near \eve\ of third legs as usual and a little in- dented on each side caudad of anterior end. Carapace with the usual four sulci of which the first is bent l)uck angularly at middle and the second is more moderately angulate in the opposite direction; the third sulcus is also weakly angulate and the fourth more strongly and acutely so between bases of the cones; a distinct median longitudinal sulcus between the first and the second sulci. Eye-tubercle distinctly limited, moderate in height, much wider than long; paired interocular processes close together, distinct, conically acimiinate, pale in color. Tubercles along lateral sub- marginal elevated rims smaller and more obscure, these, as usual, be- coming larger but of only moderate size caudad, the tubercles widely separated, those across caudal border similar. A pair of wideh' sepa- rated tubercles between eye-tuliercle and first sulcus. Area between sulci I and II with a few low tubercles; the second area with an irregular transverse row of more distinct setigerous tubercles and the tubercles of the last area more nvunerous as usual. Conical processes of carapace moderate in size, acutely acuminate with an acute curved branch on subdorsal side, (Plate 4, fig. 6, 7). Anterior border of head elevated as usual, being limited caudad by a transverse furrow as in other species. Frontal margin not dentate. First three tergites of abdomen each with a transverse row of tuber- cles which decrease in size laterad and which are well separated. Anal tergite with a transverse row of four tubercles. Sternites of abdomen with corresponding rows of smaller tubercles. Coxae I distally bent rather strongly forwards, the second more moderately so, the third straight and a little shorter than the second; coxae I to III inclusive each with a row of distinct tubercles along the midventral line. Coxae IV of the usual general form; tuberculate; the tubercles moderate, numerous but not dense; at distal end on mesal side with a proximally stout but not long process which distad is abruptly narrowed and terminates in an acicular point; on ectal side of distal end a shorter conical process. chamberlin: the arachnida. 185 Spiracles conspicuously exposed. First article of mandible strongly constricted as usual with the article abruptly expanded distad of the constriction and with the dorsal hump conspicuous. Second article long, moderately curved ventrad at middle, the dorsal (anterior) face being somewhat depressed or excavated and bearing a moderate number of short stiff hairs. Fingers of chela crossing at tips, the outer one stouter, longer and more strongly bent at tip, (Plate 4, fig. 3). Pedipalps in type about as long as body, but in a second specimen in which the abdomen is strongly retracted they are considerably longer. Coxa strongly elevated above, the ventral process long and subcylin- drical, white in color. Trochanter constricted proximally; at distal end on ventral surface a large conical tubercle. Femur subcylindrical, a few inconspicuous tubercles along ventral surface. Patella unarmed. Tibia along mesoventral line with two long curved spinous processes and two very small ones between the two large and in front of the distal one respectively; along ecto ventral line with two long less curved processes of which the anterior is the larger and near the base of the latter and distad of it a very much reduced spine. Tarsus in mesoventral line with two long processes and along ectoventral line with two large processes and a third smaller process on ectal side be- tween the others and in a second specimen with a small spine in front of others on ectal as well as mesal side. Claw much longer than the tarsus, (Plate 4, fig. 4, 5). Legs I to III with femora, patellae, tibiae, and metatarsi tubercu- late, more strongly developed on femora than on more distal joints and more conspicuously ventrally than dorsally. Fourth legs with tubercles of femora largely replaced by more conspicuous conical processes; tubercles of other joints also stronger than on other legs Tarsus I of five segments, of which three are in the second division; II with eight, also with three in the second division; III with four (the most proximal with two false sutures thus giving appearance of seven articles), these densely clothed ventrally with fine short hairs as usual, none of the articles modified specialh'. Leg IV, (Plate 4, fig. 8). Length 5.6 mm.; greatest width of carapace 4 mm. Length of leg I cir. 8 mm.; of leg II, 14 mm.; of leg III, 11 mm.; of leg IV, 14 mm. Locality.— Huadquina, 5,000 feet, July 24. (Type, M. C. Z. 127, one female). San Miguel, 6,000 feet, JuJy. (Paratype, M. C. Z. 128, one female). 186 Bl'LLETIX: MUSEUM OF COMPARATIVE ZOOLOGY. GONOLEPTES SCOTIUS/ sp. IIOV. Plate 3, fig. 6-8; Plate 4, fig. 1-2. Body throughout black of brownish cast, lines of white along cara- pace caudally and laterally and bordering tergites and sternites of abdomen. Caudal edges of coxae IV also bordered with white. Carapace under lens seen to be not densely mottled with small lighter spots in head region and over border behind. The legs similarly mottled but light spots more numerous, the trochanters of all legs and the tarsi of third and fourth legs and in some also of first and second lighter in color. Palpi paler than legs, the spots being larger and in part confluent. Body relatively narrower than in enoplus, the greatest width of carapace being much less than the total length of body and about equal to length of carapace. The general form of body shown in Plate 3, fig. 6. Carapace with four transverse sulci of which the first is deeper and is angulate at middle in the usual way, bifurcate on each side toward end; second sulcus angularly bent forwards at middle, the third toward each end bending back caudoectad; a strongly impressed longitudinal median sulcus between first and second transverse sulci which is more weakly continued between second and third and third and fourth. A transverse sulcus also indicated in line with caudal edge of eye-tubercle. Eye-tubercle sharply defined, moderate in height, wider than long, bearing a few small but distinct tubercles with paired ones present in place of the cones of enoplus rounded or boss-like, these two smaller than eyes but larger than other tubercles. Median portion of head just in front of eye-tubercle elevated and the anterior border each side of this part elevated as usual. Lower frontal margin excavated for mandibles as usual ; upper frontal margin without distinct teeth at each rounded lateral corner but with rounded boss at end of ridge there. From cervical region caudad on each lateral border of carapace the usual series of tubercles ; a tubercle-free area opposite the third sulcus and behind this about three noncontigu- ous larger tubercles and then along caudal margin a series of smaller tubercles which are widely separated and irregularly spaced. The conical processes at posterior end of carapace long, acuminate, close ' ffKoTios, dusky. chamberlin: the arachnida. 187 together. Dorsal area of carapace between first and fourth trans- verse sulci with scattered small granules. The three anterior tergites of abdomen each with a transverse row of well-separated conical tubercles which decrease in size from the middle ones laterad. Anal tergite caudally rounded, with two rounded ele- vations near base. First two pairs of coxae curving a little forwards, the third straight throughout its length; second coxae a little longer than the third; first coxae with the usual row of setigerous tubercles which are very small; the second and third coxae smooth or nearly so. Fourth coxae greatly enlarged in the usual way; ending distally on the ecto- dorsal side in a large curved process which is unbranched, and on opposite side in a shorter one which presents a low rounded process on ectal side below tip. Coxae all with numerous small granules each bearing a hair. Spiracles distinctly exposed. First joint of mandibles immediately distad of its constriction with an abruptly elevated rounded hump. Second joint with a few hairs in front toward distal end. Trochanter of pedipalp with a rounded hump above and a single setigerous conical tubercle below. Femur subcylindrical ; along ven- tral median line between base and middle three large conical tubercles. Patella unarmed. Tibia along meso ventral line with two spinous pro- cesses of which the anterior is much more slender and along the ecto- ventral line with two spinous processes of which the anterior is the larger, and in front of the latter a third much smaller spine. Tarsus along ectoventral line with five spines of which the first from caudal end and the third and fourth are larger; three spines in meso ventral line; claw large, as long or somewhat longer than the article (Plate 4, fig. 2). Femora of first three pairs of legs granulotubercular \entrally, the tubercles of the third pair largest; tibiae of these legs also tubercular, the tubercles small, those of the ventral surface and especially those of the third pair largest; metatarsi more obscurely granular. Tro- chanter of leg IV granular; on inner side with a rounded process and on outer side with a much longer acuminate process which curves back caudad above base ; femur tubercular and distad with a number of larger processes of which two much exceeding the others in size arise from the ventral surface toward the distal end and distally curve mesad; patella uniformly seriately tubercular; tibia with numerous tubercles which are larger, subconical, on ventral surface; metatarsus over proximal half with numerous tubercles, the distal, 188 bulletin: Must:u.M of comparative zoology. clavately enlarging lialf nearly smooth, (Plate 4, fig. 1). Tarsus I with five segments; II with six articles of which three are in the second division; III with six which are clothed ventrally with numerous fine hairs and none of which is specially enlarged, but the metatarsus is greatly enlarged toward its distal end which is abruptly narrowed and resembles_a tarsal article in form and pubescence, while from the ventral surface toward proximal end arises a very large, abruptly ciu'ved branch or process, (Plate 3, fig. 8). Length of type 7.8 mm. ; greatest width of carapace 6.1 mm. Length of leg I cir. 9 mm.; of leg II, 16 mm.; of leg III, 12-1- mm.; of leg IV cir. 17 mm. ioco///(/.— Lucma, 7,000 feet, August 7. (Type, M. C. Z. 129, one male; paratypes, 130, two males). Pachylus orinus,^ sp. nov. Plate 5, fig. 1-3. Body strongly narrowed cephalad; conspicuously constricted both laterally and dorsally along first furrow just back of eye-region and nearly in line with coxae of third legs; widest a little back of con- striction where the sides are convexlv rounded and from w^here the body narrows cavidad to end of abdomen with no constriction be- tween cephalothorax and abdomen; abdomen caudally semicircularly rounded. Carapace with five transverse sulci of which the most anterior lies in the previously mentioned constriction and at its middle is bent caudad at a distinct angle; the second one forms a slight angle at middle with apex cephalad, while the other sulci are straight; no connecting longitudinal median sulcus excepting a very weak one between the first and second. Eye-tubercle sharply set off, about equal in length and breadth; the cone between eyes high and acute; eye-tubercle separated by a transverse furrow or depression from the conspicuovisly elevated frontal border. Lower frontal margin deeply notched or excavated for insertion of mandibles and between the two notches extending ventrad in a spiniform process, otherwise unarmed ; imarmed above. Carapace with lateral borders strongly elevated, the marginal ridge over its middle, ectally curving portion conspicu- 1 ' opeicot, a mountaineer. CHAMBERLIN : THE ARACHNIDA. 189 ously granular, the end portions smooth; caudal border with a trans- verse row of tubercles; the area between fifth and fourth sulci with a transverse row of fewer and less developed tubercles, the two areas next cephalad with still fewer and more obscure tubercles similarly in a transverse line, (Plate 5, fig. 1). The first two abdominal segments each bearing above a transverse row of numerous, well-developed granules. Dorsal anal scutum bearing five conspicuous spinous processes of which the median is much longest, and in front of the most anterior of these on each side a series of much smaller teeth which decrease progressively cephalad; the second spine from the median one on each side is elevated dorsally along its whole length in a keel-like form, the elevated portion pro- jecting cephalad over base in a rounded, tubercle-like process. First three pairs of coxae proximally parallel, those of the first two pairs distally curved considerably cephalad, the third ones straight throughout. Second coxae but little more than one fifth longer than the third (15:11). The coxae of the second and third pairs along anteroventral surface with a low, sharply elevated edge which is obscurely tuberculate; first coxae with no elevated edge but wnth a row of very small tubercles. Fourth coxae very strongly enlarged as usual and directed caudad; each one terminating at distal end on ectal side above in a stout acute spine and on opposite mesal side of end in a similar but smaller spine; elsewhere smooth, not tuberculate or dentate. Spiracles distinct, rather large. ]Mandil)les stout; first article strongly narrowed proximally, the expanded distal portion with a strongly elevated, long, keel-like hump above which is smooth ; second article in front sparsely clothed with hairs and with fewer hairs behind toward distal end, (Plate 5, fig. 2). Pedipalps rather short, when extended scarcely as long as carapace ; not crossed. Trochanter short, . subcylindrical, with a low hump above on which are one or two slight tubercles; ventrally with a few hairs springing from slightly tubercular bases. Femur cylindrical, not compressed, moderately convexly elevated from end to end above; wholly without spines; a few short hairs from slightly tuber- cular bases below. Patella unarmed. Tibia not compressed; along mesoventral edge with two rather slender, acutely acuminate spines and between these a minute third one; along ecto ventral edge with three long spines and between the caudal and middle one of these a fourth minute one; the middle one of the larger spines much largest 190 bulletin: museum of comparative zoology. and a little surpassing the distal end of tarsus when joints are flexed, the other two spines proximally abruptly thicker than distad, anterior spine cognate to base of large spine. Tarsus on mesal side ventrally with a series of four slender, distally bristle-like spines; on ectal side with a series of about twelve pale slender spines which are very short excepting two which are much larger and of about same size as the three of mesal side ; tarsal claw of about same length as tarsus, (Plate 5, fig. 3)- . First three pairs of legs with femora, patellae, and tibiae especially on ventral surface finely but rather sparsely tubercular, a hair arising from each tubercle, distal joints wholly smooth. Fourth legs stout, with tarsi abruptly much more slender than the metatarsi; tro- chanter with two robust granules on ectal side and at distal end on mesal side with a stout thorn; femur granular above, along ventroectal edge with a series of mostly stout conical spines which, beginning at about one fourth the length from the proximal end as low tubercles increase regularly in length distad; along meso ventral surface a series of fewer and lower conical tubercles and on the mesal surface with two or three irregular series of conical tubercles and spines ; patella strongly tubercular above and laterally, and with tubercles replaced by stout conical spines of which three or four are comparable in length to the larger ones of femur; tibia above and laterally strongly tubercular, below with tubercles replaced by stouter conical tubercular elevations and longer spines like those of the proximal joints ; metatarsus above and laterally densely granular, below with stouter seriate cones or teeth, much smaller and more uniform than the spines of the proximal joints. Tarsus I with five joints, three of which are in the distal division; tarsus II with eight joints; III and IV with six joints. General color dilute ferruginous, the head region weakly dusky; caudal border of carapace and the first two tergites of abdomen black; patellae and distal ends of femora and tibiae of legs dusky or black, the fourth legs a darker, more strictly ferruginous cast than the others. Abdomen darker beneath than coxae of legs. Length to base of median caudal spine 10.2 mm. ; to tip of caudal spine 12.3 mm. Greatest width of carapace 6 mm. Length of leg I, (exclusive of coxa) cir. 15 mm.; of leg II, 22 mm.; of leg III, 7 mm.; of leg IV, 30 mm. Lom/%.— Huadquina, 5,000 feet, July. (Type, M. C. Z. 131). San Miguel, 6,000 feet, July. (Paratype, M. C. Z. 132, one specimen). chamberlin: the arachnida. 191 COSMETIDAE. Paravanones peruvianus, sp. nov. Pl'ate 2, fig. 1-6. General background of body a dilute ferruginous. Carapace dusky from a close network of black lines, the color less dusky in a deltoid area back of eye-tubercle with apex caudad and embracing two short black stripes, one caudad of each eye, in a narrow mid- longitudinal line, and along the transverse sutures and in a broader area along each side of the carapace, tubercles pale. Ventral surface of cephalothorax and coxae paler, a clearer light ferruginous; under lens seen to be covered with a less dense network of dark lines. Ab- dominal segments both above and below black or nearly so, a line back of each segment white. Legs also with an inconspicuous net- work of fine dark lines; femora, patellae, and tibiae dark at distal ends and tarsus of leg I also blackish. Carapace widest at level of third sulcus, convexly rounded, abruptly narrowed a little in front of caudal end and at level of caudal edge of third legs in front of which the sides converge but moderately, the anterior corners more oblique. Carapace crossed by five transverse furrows which are wide, not suture like, and of which the first is bent back angularly at the middle. Eye-tubercle much wider than long, concavely depressed from ends toward middle; a series of four tubercles in a curved line on each elevated end subparallel with edge of ocellus and several weaker tubercles immediately mesad of these; the median portion of eye- tubercle smooth. Anterior margin not elevated or only obscurely so on each side, the median portion between eye-tubercle and front margin broadly elevated. At each anterolateral corner a prominent process, (Plate 2, fig. 3), the border near level of anterior edge of second coxae with a process projecting caudoventrad and meeting a process from second coxae. Edges of carapace in its widest part with a number of rather low conical tubercles, the tubercles across caudal border more numerous and with two much larger subcylindrical processes near middle; the more anterior lateral margins essentially smooth. On areas between furrows a limited number of tubercles which are larger and somewhat more numerous in caudal than in anterior region, and in each area a pair of tubercles which are higher I 192 bulletin: museum of comparative zoology. and more cylindric than the others; surface of carapace in general granulate, (Plate 2, fig. 3). A row of conical tubercles across each abdominal segment. Anal scutum proximally with smaller granules and tubercles, distally with a number of larger tubercles, about five of which appear as dentations on the caudal margin, one at middle and two or three on each side of it. First coxae bent forwards at .ends, the second slightly so. Third coxae but little shorter than the second. Coxa I in both male and female on caudal side above middle with two conical processes and at distal end on caudo ventral corner with a longer process which is bent dorsad; along its antero ventral surface a row of tubercles the most distal of which appears at the anteroventral corner as a freely pro- jecting larger conical process; from anterior surface at distal end a long cylindrical process which bends dorsad close to surface and over- laps the end of a process springing from anterodorsal surface. A low broad process from distal end of coxa II near anterodorsal corner; a row of a few tubercles along anteroventral line and a similar one on third coxae. Coxa IV at distal end on mesal side with a low rounded process, (Plate 2, fig. 4-6). Spiracles clearly exposed. First joint of mandible strongly elevated above distad of the con- striction; with a transverse row of cavidally projecting teeth or conical tubercles across upper border of caudal surface of the dorsal hump; dorsal surface of hump with fine teeth or granules; second article rather short and broad with a few hairs at distal end below^ (caudad) and more in front, (Plate 2, fig. 1). Pedipalps short, closely flexed ventrad and caudad, not crossed. Coxa subcylindric. Trochanter narrow at base, expanding distad, with a broad conical projection beneath. Femur strongly flattened from side to side, being abruptly very high immediately distad of the narrow base; a row of teeth along the middorsal line and a series of longer subcylindric, distall}^ rounded processes along ventral line. Patella of usual form; bearing small setigerous tubercles above. Tibia strongly flattened from side to side, especially on ventral half, clavately increasing in height from base distad; with a number of small tubercles on dorsal surface proximad. Tarsus subcylindric, with a low thin keel from ventral surface at proximal end, (Plate 2, fig.^2). Femora, patellae, and tibiae of legs I to III finely tubercular or granular, the metatarsi more abruptly so proximally and especially chamberlin: the arachnida. 193 on legs III. Leg IV in female similar to leg III; in male with a series of much stronger teeth along dorsal or mesodorsal line and on ventral surface at distal end with a series of five conspicuous long processes of which the second is longest and is bent at distal end, the others being straight and decreasing in length distad; the patella with a stout spine above at proximal end. Tarsus I with five segments; II with thirteen segments; III with six segments, of which the last three are abruptly more slender; IV with seven segments of which three are in the second division; tarsi III and IV clothed with fine hair on ventral surface. Length of male type 6 mm.; greatest width of carapace, 4.5 mm.; length of leg I cir. 1 mm.; of leg II, 2Lo mm.; of leg III, 13 mm.; of leg IV, 8 mm. Length of female paratype 6 mm.; greatest width of carapace 4.5 mm.; length of leg I, 10.5 mm.; of leg II, 20 mm.; of leg III, 14 mm.; of leg IV, 18 mm. Localities. — Santa Ana, 3,000 feet, August 4. (Type, M. C. Z. 133, male; parat^'pe, M. C. Z. 34, female). San Miguel, 6,000 feet, July. (M. C. Z. 135, one female). PHALANGIIDAE. LlOBUNUM MONTICOLA, Sp. nOV. Plate 5, fig. 9; Plate 6, fig. 1-3. Body above brown, a paler longitudinal median stripe which narrows to a point at caudal end of abdomen, this stripe on anterior portion of abdomen embracing a deep black, sharply defined median stripe. Eye-tubercle black. Ventral surface of body yellowish. Trochanters all deep black and coxae dusky distad; patellae also solid black and tibiae black at distal ends. Body blunth^ rounded behind, the abdomen broadest across caudal end; carapace narrowed markedly cephalad. Entire surface of body above and below densely granular, the granules conically acutely pointed. Abdomen clearly set off from cephalo thorax by a suture; the two segments at end distinct. Eye-tubercle high, its anterior and posterior faces subvertical or a little constricted at base; a curving line of acute spinous points 194 bulletin: museum of comparative zoology. mesad of each ocellus and curving caudoectad and ventrad on the caudal surface of tubercle, (Plate 6, fig. 1). First joint of mandible nearly smooth, a little roughened over dorsal surface; hairs above few and straight, more on mesal surface. Second joint also smooth except for a light dorsal roughening; with a few longer hairs at distal end above and laterally with a more numerous patch on mesal side distad, (Plate 6, fig. 2, 3). Pedipalps moderately long. Femur ventrally along ectal and similarly along mesal side with a row of acute teeth ; also with a patch of teeth above and laterally at distal end. Dorsoectal and dorso- mesal surfaces of patella with fine teeth which are largest and most numerous on the dorsomesal process, a narrow dorsal stripe free from teeth or nearly so. Tibia with teeth ventrally and especially laterally and dorsolaterally, a middorsal stripe free from teeth. The long slender tarsus free irom teeth but with numerous short fine and a few coarser short hairs. Legs with numerous fine teeth which become smaller and less con- spicuous in going distad. Length of body 3.8 mm. Length of palp cir. 4.5 mm.; of leg I, 32.5 mm.; of leg II, cir. 80 mm.; of leg III, 41 mm.; of leg IV, 5L5 mm. Localities.— Paltaybamba, 5,000 feet, August 27. (Type, M. C. Z. 136). Santa Ana, 3,000 feet, August 3. (Paratype, M. C. Z. 137, one specimen). LioPAGUS,^ gen. nov. Eye-tubercle weakly longitudinally furrowed; wholly smooth, bearing no teeth or spines or other processes. Body essentially smooth. Scutum unarmed. First joint of mandible at base with the usual ventral process. Patella of pedipalp with a moderate inner apophysis. Claw dis- tinctly pectinate. A one pointed process on dorsal margin of distal end of all coxae. Legs thin; moderately long. Femur II with two false sutures; III with none and IV with one. Genotype. — Liopagus simplex, sp. nov. Related to Prionostemma Pocock in the presence of false sutures only on second and fourth femora. From that genus it is most e'asily distinguished in having on femur II only two sutures or joints instead of the three uniformly present in Prionostemma. I Xcios. smooth, Trdyos, a peak. chamberlin: the arachnida. 195 LiOPAGUS SIMPLEX, sp. nov. Plate 5, fig. 4-8. Body above brownish grey, darker caudad; similar below with a darker brown or dusky stripe across each abdominal segment. Femora toward distal end each with a conspicuous deep black annu- lus. Legs brown; patellae and the distal ends of tibiae darker. Body nearly parallel sided from level a little back of eye-tubercle to one a little caudad of ends of coxae IV, then abruptly narrowing to caudal end and semicircularly rounded in front. Obscurely granular in a line across border of segments. Eye-tubercle as wide as high. Viewed from behind appearing conspicuously constricted at base, the front face subvertical, not so convex as the caudal. Tubercle wholly smooth; slightly depressed longitudinally along middle line, (Plate 5, fig. 4, 5). All joints of legs free from tubercles but with scattered very fine teeth on femora and a large one at distal end above on femora and patellae. Coxae with a single slender process from distal end above. First joint of mandible smooth, with the process below as usual ; a few short hairs above and a series of them along a meso ventral line. Second joint small; with a few short hairs above, glabrous beneath, (Plate 5, fig. 6). Femur of pedipalp between two and three times as long as the patella without its apophysis; with a series of short stout hairs be- neath and smaller, less erect ones on dorsal surface but with no teeth. Patella with apophysis moderately slender, cylindric, more densely clothed with hair. Tibia also with more numerous short stiff hairs; a slight rounded process at distomesal corner, this clothed with hairs as elsewhere on joint. Tarsus with numerous fine short appressed hairs and fewer shorter, stiff erect hairs. Claw pectinate, (Plate 5, fig. 7, 8). Length dr. 3.5 mm. Leg I missing. Length of leg II cir. 42 mm. ; of leg III, 2L5 mm. ; leg IV broken off at tip. Locality.— Huadquina, 5,000 feet, July 30. (Type, M. C. Z. 138). 196 bulletin: museum of comparative zoology. ARANEIDA. AVICULARIIDAE. Hemirrhagus peruvianus, sp. nov. Plate 6, fig. 4-10; Plate 7, fig. 1-2. Integument of carapace, chelicerae, femora of legs and palpi, sternum and labium dark chocolate-brown, the other parts of legs and palpi lighter brown, in part of dilute reddish tinge; sternum and femora of legs beneath sometimes lighter than above. Integument of abdomen brown, paler beneath than al:)ove; above with a small light area toward anterior end. Carapace covered with a woolly coat of russet-brown hair of a more or less coppery lustre, and some longer grey ones at caudal edge especially. The legs are thickly clothed with short brownish hairs with deep black ones more sparsely inter- mixed; the numerous longer bristles are dark proximally, becoming grey, grey or white distad, these forming rather indistinct longitudi- nal streaks on the femora and patellae. Abdomen densely clothed with brown hair of a distinctly coppery lustre, that of anterior surface black and forming a distinct black area; longer bristles of dorsal surface grey distally; sometimes dark hairs form a larger element on the ventral surface and this sm-face then appears dark grey or blackish instead of copper-brown as more usual. Fringes of endites and cheli- cerae orange proximally, lighter distad. Eye-tubercle sharply limited; of moderate height. Eye-area less than twice as wide as long in both male and female (67 : 37) ; rows subequal or the posterior one slightly longer than the anterior (up to ratio 67:65). Anterior row of eyes in dorsal view a little procurved; in anterior view rather strongly procurved. Anterior median eyes with diameter not much more than half that of the laterals {cir. 6:11), a little more than their radius apart, but decidedly less than their radius from the laterals. Posterior row of eyes conspicuously' re- curved. Posterior laterals smaller than the anterior laterals from which separated by less than their radius. Posterior medians three fourths the diameter of the laterals ; mesal side more curved than the ectal, more or less angulate anteroectally, (Plate 6, fig. 4). Head low, only slightly rising in front of the thoracic groove; chamberlin: the arachnida. 197 highest some distance eaudad of eyes. Thoracic fovea some distance back of middle of carapace; straight, transverse. Sternum subequal in length and breadth, being sometimes slightly longer than wide (male) and sometimes slightly wider than long (female). Moderately convex. Sigillae submarginal. Labium a little wider than long. Spinules in a transverse band of about four rows, irregularly and closely arranged. All tarsi densely scopulate; the scopulae of the anterior tarsi divided by a narrow setose line, those of the posterior tarsi by a broad setose band which, however, is narrow^er than the joint. Anterior metatarsi scopulate mostly more than half way to base, the posterior metatarsi scopulate at distal ends only. Hair on anterior surface of coxa I both above and below suture moderately long, in part prone, with in addition a number of finer and shorter straight hairs which tend to be somewhat clubbed at tip; no spinescent bristles. Paired claws bearing from three to four moderate teeth, commonly three or two being fully developed with one or two appearing as mere points, (Plate 7, fig. 1, 2). In addition to the scales of the ordinary type occurring on dorsal surface of tarsi, (Plate 6, fig. 5) occasionally one of the second type, (Plate 6, fig. 6), is to be seen. Metatarsus I ventrally with an apical and a subbasal spine, a long one also on anterior surface. Tibia of male with three spines along ventrocaudal line and a fourth a little more dorsad; or with spines as many as seven to none, there being on caudoventral surface three l)asal, two submedian and one apical, and on anteroventral a pair toward apex and one lateral surface (male from Huadquina); the number may differ on the right and left legs of same specimen. Patella armed with a spine on caudal surface (male) . Metatarsus IV with three pairs of ventral spines and two on anterior and also on posterior surface (female) or with five or six on each lateral surface (male). Inferior spur of tibia I of male longer than the superior, moderately curved and bearing a large stout black spine inserted on its dorso- caudal surface. The superior spur bearing a similar stout spine on mesal surface, (Plate 6, fig. 7). Tibia of male palpus thicker proximally than patella or femur, narrowing distad ; a narrow ridge on mesal side from middle distad, elevated at distal end into a low tubercle. Tarsus short, bilobate as usual. Palpal organ with spine in lateral view appearing at right angles to the main axis of bulb, narrowing distad, with apex slender and acute, below tip with a short, stout, subtriangular spur. In 198 bulletin: museum of comparative zoology. anterior view the tip of process is seen to curve rather abruptly ectad, the spine occurring at the bend on the convex side, (Plate 6, fig. 10). Male (Tincochaca) . Length, 15 mm. Length of cephalothorax, 7.3 mm.; width, 6.0 mm. Leg I Leg II Leg III Leg IV fem. 6.2 mm. 6.0 5.1 7.2 tib.+pat. 9.0mm. 7.0 6.2 8.5 met. 4.0 mm. 3.6 4.6 7.0 tar. 3.5 mm. 3.0 3.0 4.0 total 24.7 mm. 19.6 18.9 26.7 Female {Tincochaca). Length, 17 mm. Length of cephalothorax, 8.5 mm.; width, 7.0 mm. Leg I Leg II Leg III Leg IV fem 5.8 mm. 5.1 4.6 6.0 tib. + pat. 7.8 mm. 6.2 5.7 8.0 met. 3 . 3 mm. 2.7 3.3 5.4 tar. 3 . 2 mm. 2.6 3.2 3.2 total 20.1 mm. 16.6 16.8 24.6 Localities. — Huadquina, 5,000 feet, July. (Type, M. C. Z. 139, male; para type, immature female 324). Tincochaca, 7,000 feet, August. (M. C. Z. 140, male and female). Santa Ana, 3,000 feet, August. (M. C. Z. 141, one female). Urubamba, 9,500 feet, July 18. (M. C. Z. 325). Hemirrhagus major, sp. nov. Plate 7, fig. 3-8. Integument of carapace, sternum, legs, and palpi dark brown or chocolate to chestnut; that of labium and endites reddish brown or chestnut. Hairs of carapace somewhat wavy, moderately thick, light brown of more or less dark golden or bronze lustre. Hair of chelicerae dense, light brown or bronze lustre. Sternum clothed with a dense coat of short brown hair with black ones sparsely inter- mixed; bristles dark proximally, rufous distally, mostly of moderate length. Legs clothed with a coat of short brown hair with darker dusky or blackish hairs intermixed and especially abundant on dorsal surface of femora, the brown hair in part of shining brown lustre; a narrow stripe of grey or white hairs across ends of joints above ; two longitudinal stripes on patellae above formed by hair free and bristle chamberlin: the arachnida. 19^ free areas over which the lighter hairs unmixed with dark extends from each side. Bristles of legs shorter proximad, becoming longer and more numerous especially on tibiae ventrally and on metatarsus above as well as laterally and ventrally; bristles dark proximally, becoming light rufous distally, rather coarse. Integument of abdomen above dark, somewhat dusky brown, ventrally lighter brown, with several small obscure white spots on each side; bristles of dorsum subdense, forming a light brick-red area. Eye-tubercle black, well defined, moderate in height, highest between posterior median eyes, more strongly convex anteriorly than posteriorly. Eye-area trapeziform, the posterior row being distinctly longer than the anterior, the difference typically distinctly more marked than in peruvianus (up to 85: 78); area from a little less than half as long as greatest width to a little more thus also differing from peruvianus. Anterior row of eyes from moderately procurved to nearly straight, much more procurved in anterior view than in dorsal. Anterior median eyes with diameter two thirds that of laterals ; about two thirds their diameter apart, less than their radius from the laterals. Posterior lateral eyes about three fourths the diameter of the anterior laterals, less than their radius from the latter. Posterior medians pyriform in outline, being narrowed to a point caudad and widely rounded cephalad, (Plate 7, fig. 3). Head moderately elevated, in outline slightly convex, highest a little caudad of eye-tubercle. Fovea straight or vaguely procurved, short and deep. Sternum longer than wide. Posterior sigillae about their length from margins. Median ones submarginal ; anterior marginal. Mesal margin of furrow or chelicera bearing a row of fourteen teeth. Labium wider than long, truncate distad. Spinules in a narrow band (about four rows deep) across distal end, the spinules not dense. Spinules on proximal end of endite rather numerous. Anterior surface of coxa I with longer in part semiprone hairs and some of the short, slender, distally obtuse or clubbed hairs such as are present in peruvianus but in addition below the suture with numerous dark, strongly chitinized spiniform bristles, these more numerous distally, some rather more slender ones also occurring above suture. Tarsal claws four or five (anterior and posterior of leg I respectively) , proportionately shorter, more slender and more uniform than in peruvianus. On the posterior claws, which are longer and propor- 200 bulletin: museum of comparative zoology. tionately more slender, one or two extra points may occur in addition to the five ordinary teeth, (Plate 7, fig. 5-7). Scopulae of anterior tarsi divided by a narrow setose line, the posterior ones by a broader one, but this much narrower than the joint. Anterior metatarsi scopulate well toward base; posterior with scopular hairs only distally and there sparse or absent. Meta- tarsus I (female) ventrally with a pair of small apical spines and a single one proximad of middle. Tibia I (female) also ventrally with a pair of spines at distal end and a single submedian one ; in the male with seven spines on ventral and ventrocaudal surface and two on anterior surface. Metatarsus IV with fifteen to eighteen spines irregularly arranged; tibia IV with eight to ten. In the male the metatarsus of leg I is abruptly much more slender than the tibia and is strongly bowed dorsad. The spurs of the tibia are elevated on a conspicuous common basal process standing at right angles to the article; the inferior process is not much longer than the superior but it is proximally stouter being narrowed distad, moderately curved toward end, bearing on dorsal surface a dark, stout, acute process attached near apex and corresponding to the larger one of penivianus; superior process directed more cephalad, more uniform in diameter, curved a little mesad toward tip, the spine rather slender, closely applied to surface excepting at tip which is divergent, (Plate 7, fig.^4). In the male palpus the tibia is thicker than the patella or femiu", narrowing distad, a narrow ridge-like thickening mesoventral edge from middle distad and elevated at distal end into a low tubercle, ventrally with a dense growth of long bristles. Tarsus short, bilobate as usual, the mesal lobe at distomesal corner adjacent to bulb extended into a blackish, densely chitinized tubercle. Spine of bulb propor- tionately much longer than in pcruvianus, basal part with axis corre- sponding with long axis of bulb, curving semicircularly first ventrad and then forwards as shown in the figure, a slight tooth a little distad of middle, in anterior view the process bends first somewhat ectad. Male (Type, Cuzco Valley). Length, 29 mm. Length of cephalo- thorax, 14 mm.; width, 12.3 mm. fem. tib. + pat. met. tar. total Leg I 10.8 mm. 14 mm. 8.1 mm. 5.4 mm. 38.3 mm Leg II 10 12.2 7.8 5 35.0 Leg III 9 11.2 8.3 4.5 33.0 Leg IV 11.2 14.2 11.2 5.S 42.4 Tibia I, 7.1 mm. Til.ia IV, S.9 mm. chamberlin: the arachnida. 201 Female (Urubamba). Length, 37 mm. Length of cephalothorax, 15.8 mm.; width, 13.1 mm. (In a female from Cuzco Valley the cephalothorax is proportionately broader, the ratio of length and breadth being 16:14.3). ' fern. tib.+pat. met. tar. total Leg I 11.8 mm. 1.5.3 mm. 7.5 mm. 5.6 mm. 40.2 mm. Leg II 11.0 13.0 7.1 5.3 36.4 I^glll 10.0 11.8 8.8 5.5 36.1 Leg IV 12.5 15.5 13.0 6.4 47.4 Localities. — Cuzco Valley. (Type, M. C. Z. 142, one male; para- type, M. C. Z. 143, one female) E. D. Flint coll. Urubamba, 9,500 feet, July. (M. C. Z. 144, one female). Hemirrhagus sp. Locality. — Urubamba, 9,500 feet, July. (M. C. Z. 326, several immature females). Eurypelma aymara,^ sp. nov. Integument of carapace when dry black or nearly so, when wet appearing of slight chestnut tinge. Sternum chestnut, darker cepha- lad. Labium nearly black. Endites chestnut. Legs proximall\' blackish like the carapace, becoming slightly more chestnut distad. Carapace clothed with a coat of sandy grey and light brown hair of a golden lustre. Hair of femora dusky brown of rufous tinge, that of more distal joints with a larger proportion of grey intermixed; bristles numerous and long, of rufous or rust color proximally, paler, greyish yellow distad. Hair of venter of abdomen brown and black intermixed, chiefly the latter. Dorsum with a thick coat of long rufous bristles. Pars cephalica moderately high, highest a little caudad of eye- tubercle. Eye-tubercle rather high, sharply limited ; highest along the median longitudinal line which descends a little from between the posterior median eyes cephalad. Eye-area considerably less than twice as wide as long (25:16). Anterior row of eyes slightly shorter than ' Aymara, a tribe of the indigenes of Peru. 202 bulletin: museum of comparative zoology. the second (25 : 24) ; in dorsal view considerably procurved, a line tangent to the anterior edges passing through the anterior third of the lateral eyes. Anterior median eyes much smaller than the laterals, their diameters being about as 2 to 3; medians three fourths their diameter apart and near the same distance from the laterals. Pos- terior lateral eyes of nearly same size as the anterior laterals from which they are separated by less than their radius. Posterior median eyes much smaller than the laterals; oblong, with sides straight and ends more rounded. A line tangent to caudal edges of the two posterior eyes of each side intersects the anterior median eye of opposite side. Metatarsus I and II scopulate very nearly to base; metatarsus III scopulate over distal third; metatarsus IV scopulate only at distal end. Female. Length 38 mm. Length of cephalothorax, 21.2 mm.; width, 18.1 mm. Length of pars cephalica, 13.8 mm. fem. tib.+pat. met. tar. total Leg I 15 mm. 20.9 mm. 10 mm. 6.5 mm. 52.4mm. • Leg II 13.8 18.1 9.9 6.1 47.9 Leg III 12.3 16 11 6.1 45.4 Leg IV 15.7 20 15.6 6.8 58.1 Locality — Feru: Aymas, Dr. W. H. Jones. (Type, M. C. Z. 145, one female). This seems to be the only true Eurypelma thus far recorded from Peru. DiPLURA MONTICOLENS,^ sp. nOV. Plate 7, fig. 9-10. Cephalothorax dusky brown, the margins of carapace and the eye- area darkest. Edges of sternum darker than median portion. Cheli- cera palpi and legs testaceous. Integument of abdomen dark above, light beneath; a series of three pairs of obscure light stripes extend- ing from near dorsal median line obliquely ventrocaudad across sides. Spinnerets testaceous, in part dusky, especially at ends. Hair of carapace light brown, those of sternum sparse, darker. Bristles of abdomen dark rufous brown, the shorter hairs mostly lighter. Eye-area twice as wdde as long or very nearly so (67:34). Eye- tubercle highest beneath anterior median eyes, behind which it descends almost immediately and also slopes laterally from this ' iVIons, mountain, incolens, inhabiting. chamberlin: the arachnida. 203 point, the eyes being set obliquely to the horizontal plane. Anterior row of eyes in dorsal view a little procurved, in anterior view strongly recurved. Anterior median eyes inclusive of rim with diameter more than three fourths as great as that of the anterior laterals, less than their radius from each other and closer to the laterals; exclusive of rim the median eyes are less than their diameter apart and are near their radius from laterals. Anterior laterals very oblique, the anterior, narrow end lying in front of the ectal edge of the median eye. Pos- terior lateral eyes clearly smaller than the anterior laterals (diameters nearly as 3:4). Posterior median eyes smaller than laterals, closer to them than to the anterior medians. A line tangent to the caudal edges of posterior eyes on each side is tangent to or passes near caudal edge of anterior median eye of opposite side, (Plate 7, fig. 9, 10). Cephalothorax broad and low anteriorly (the width across eye area to greatest width about as 47:76). Thoracic fovea small, oval, transverse, situated considerably behind middle of length of carapace. Labium much wider than long, with anterior margin mesally in- curved. On anterior edge with two spinules present, a scar in t>T)e seeming to indicate the normal presence of a third one. Endites at proximal corner with a small group of few (8-10) spinules. Sternum as wide as long. The sigillae marginal. Tarsi of legs slender, all conspicuously curved, those of the anterior pairs most strongly so. Paired claws each with a double row of mostly eight teeth. Inferior claw smooth, slender. Metatarsus I with three pairs of ventral spines; femur I with a spine at distal end on ante- rior side and above and another smaller one on the posterior side; other femora with corresponding spines, the posterior one increasing in size in going caudad. Posterior metatarsi much more strongly spined. None of the tarsi spined. Posterior spiimerets as long as abdomen; slender; articles sub- equal, the first thickest, the distal one most slender, narrowing distad. Anterior spinnerets short, distally acuminate. Female. Length 8.4 mm. Length of cephalothorax 3.4 mm.; width, 2.8 mm. fern. tib.+pat. met. tar. total Leg I 3.4mm. 4.5 mm. 2.9 mm. 2.2 mm. 13.0mm. Leg II 3.1 4.1 2.8 2.2 12.2 Leg III 2.9 3.8 2.8 2.0 11.5 Leg IV 4.0 5.0 4.0 2.5 15.2 Loca%. — Huadquina, 5,000 feet, July. (Type, M. C. Z. 146, female). 204 bulletin: museum of comparative zoology. Brachythele keithi, sp. nov. Plate 7, fig. 11. Integument of cephalothorax and legs light chestnut-brown; that of the chelicerae dark chestnut; integument of abdomen and spin- nerets clear brown. Pubescence of carapace brown of golden lustre ; that of abdomen dense, of similar golden brown color or of more coppery lustre. Eye-tubercle strongly convexly elevated between the posterior median eyes. Eye-area twice as wide as long (45:22); anterior row a little shorter than the posterior (45:42). x\nterior row a little procurved. Anterior median eyes a little more than their radius apart, closer to the lateral eyes; diameter but little more than two thirds the longer diameter of the lateral eyes (ratio near 23:31). Posterior lateral eyes slightly longer than the anterior (33: 31). Pos- terior median eyes elongate, narrowed caudad, length to width about as 3: 5 — smaller than lateral eyes (lengths as 25: 33), nearer to these than to anterior medians. A line tangent to caudal edges of posterior eyes on each side in type passing through anterior third of opposite anterior median eye, (Plate 7, fig. 11). Labium clearly wider than long; distal margin mesally incurved. Spinules none. Spinules on proximal corner of endites short and stout, constricted near or a little above base, distally expanded and rounded. Thoracic fovea recurved. Sternum longer than w4de; gently convex. The sigillae at \e\e\ between second and third legs most distinct, marginal. Sternum with numerous fine tubercles from each of which arises a hair. Paired claws with teeth in two rows; teeth of each row mostly seven or eight in number. Unpaired claw smooth. Tarsal scopulae dense, extending to base; none divided by a setose line or band. Spines long and moderately stout, black. Posterior spinnerets more than two thirds as long as the abdomen ventrally ; the three joints subequal in length, decreasing in diameter distad, the second being more slender than the first and the third than the second. Anterior spinnerets twice or more their diameter apart at base. Female. Length, 30 mm. Length of cephalothorax, 14 mm.; width, 12 mm. chamberlin: the arachnida. 205 fern. tib. + pat. met. tar. total Leg I 12 mm. 15 mm. 9.5 mm. 8 mm. 44.5 mm. Leg II missing Leg III 10 12.5 10 7.5 40.0 Leg IV missing Length of spinnerets 12.2 mm. Locality. — Huadquina, 5,000 feet, wet season. (Type M. C. Z. 147, one female). Named for Minor C. Keith of New York. Brachythele incursus, sp. nov. Plate 7, fig. 12. Integument of cephalothorax and legs brown, that of carapace and femora of legs above deeper in color; chelicerae above blackish brown. Integument of abdomen above dusky brown; below light brown. Spinnerets blackish. Hair of carapace and abdomen gold- en brown. Hair of sternum blackish. Fringe of chelicerae rufous. Eye-area less than twice as wide as long (57:30). Anterior row of eyes of same length as posterior. Anterior row straight. Anterior median eyes slightly more than their radius apart (6:5.5), nearer to the laterals; diameter but little more than half that of lateral eye (11:20). Posterior lateral eyes clearly smaller than the anterior laterals (ratio of diameters 3:4), lateral eyes nearly contiguous as usual. Posterior median eyes elliptic, about three fourths as wide as long, four fifths as long as the laterals. A line tangent to caudal edges of two posterior eyes of each side passes clearly caudad of anterior median eye of opposite side, (Plate 7, fig. 12). Labium wider than long in the ratio 9.5:6.8; distal margin a little incurved. A row of low stout, rounded spinules across apical portion. Endites at proximal corner with the patch of spinules as usual. Thoracic fovea a little recurved. Sternum longer than wide. Hairs not dense, black, longer about margins. Paired claws of legs each with two rows of from six to eight teeth. Single claw smooth. In the stage of growth represented by the type the anterior tarsi are not densely scopulate and the metatarsi are scopulate only distally; the posterior tarsi are scopulate only 206 bulletin: museum of comparative zoology. distally, ordinary bristles covering the proximal portion while the metatarsi are not at all scopulate. The tarsi are not armed. Patel- lae, tibiae, and metatarsi armed with stout black spines as usual. Spinnerets short, about one half the length of the abdomen on ven- tral side, much less slender relatively than in keithi: each joint atten- uated distad. Black in color. Female {not fully mature). Length, 13 mm. Length of cephalo- thorax, 6.2, mm.; width, 5 mm. fern. tib.+pat. met. tar. total Leg I 4.3 mm. 5.6 mm. 2.8 mm. 2.0 mm. 14.7 mm Leg II 4.0 4.7 2.3 1.8 12.8 Leg III 3.2 4.1 2.2 1.8 11.3 Leg IV 4.3 5.5 3.2 2.0 15.0 Loca/i/?/.— Huadquina, 5,000 feet, July. (Type, M. C. Z. 148, one young female). The female described above is apparently not fully grown; but the differences it presents especially in eye sizes and relations added to the differences in proportions of joints of legs, the presence of spinules on the labium and especially the pronounced differences in form and relative length of spinnerets render it impossible to regard it as identi- cal with the preceding species which comes from the same locality. ULOBORIDAE. Orinomus,^ gen. nov. Posterior eyes small, subequal, the series strongly recurved, nar- rower than the cephalothorax, the medians farther apart than from the laterals. Anterior eyes in a procurved series; medians farther from laterals than from each other. Area of median eyes much wider than long and much narrower in front than behind. First leg much the longest, the others in order IV, II, III. Meta- tarsus I clearly shorter than tibia + patella I. Calamistrum not reaching distal end of metatarsus IV. . Cribellum narrow; entire. Abdomen subglobose; bigibous above, not produced or acuminate behind; spinnerets not quite terminal. ' opeii-o/ios, mountain ranging. chamberlin: the arachnida. 207 Genotype. — Orinonius lamprus, sp. nov. Apparently closest to Uloboms, a genus occurring widely in this and adjoining regions as well as elsewhere. Orinomus lamprus/ sp. nov. Plate 8, fig. \-4. Carapace nearly black, a narrow median longitudinal line yellow- ish, the clypeus also paler. Carapace clothed with grey hair. Ster- num black; a pale median mark. Coxae dusky beneath. First legs with femora, patellae, and tibiae black, with a few small light spots; metatarsi and tarsi yellow. Other legs yellow, with dusky annul i. Abdomen somewhat yellowish white, finely spotted with black, a dark median longitudinal line over entire length and a deeper, more distinct one on each side from the hump caudad; venter with a geminate black stripe along the middle. Posterior row of eyes conspicuously recurved, a line tangent to caudal edges of median eyes passing through the anterior third of laterals; median eyes three times their diameter apart, half as far from the laterals which are nearly of same size. Posterior lateral eyes as far from the anterior laterals as from the posterior medians (once and a half their diameter), the anterior laterals with diameter two thirds as large. Anterior median eyes only a little more than their diameter apart but more than once and a half their diameter from the laterals; row conspicuously procurved. Area of median eyes wider than long nearly in ratio 5:3; twice as wide behind as in front. Cly- peus slanting ventrocephalad ; about 6qual in w^idth to diameter of median eye. Labium wider than long; subtriangular, the sides being straight and meeting at an angle, (Plate 8, fig. 4). Sternum caudally produced as a short tongue between the separated posterior coxae, the tongue rounded at end. More than twice as wide as long (20:9), (Plate 8, fig. 2). Female. Length 3 mm. fem. tib. + pat. met. tar. total Leg I 1.6 ram. 1.6 mm. 1.2 mm. .6 mm. 5.0 mm. Leg 11 1.0 1.1 1 .3 3.4 Locality.— Tiale). -Urubamba, 9,500 feet, July. * \aiivp6s, distinct. (Type, M. C. Z. 149, one 208 bulletin: museum of comparative zoology. DICTYNIDAE. Amaurobius, sp. a. An immature male and female from Ollantaytambo (9,000 feet, July. M. C. Z. 150). The abdomen is marked with a distinct median longitudinal black stripe of narrowly hastate form and with indented edges and with a bright white spot each side of middle of its length. Carapace with head dark brown, elsewhere dusky. Sternum dark, dusky brown. Two immature males apparently of same species from Urubamba (9,500 feet, July. M. C. Z. 151). They are darker throughout, the sternum being nearly black and the abdomen so dark that the dorsal median black line is almost obliterated ; but the two white spots are conspicuous. Amaurobius sp. b. One immature female with lateral eyes nearer together than usual in the genus. Carapace yellow with each eye enclosed in black. Sternum light brown, a little dusky. Legs yellow, tibiae very ob- scurely ringed. Abdomen grey of olive cast; above conspicuously marked with a series of dark chevron lines from middle caudad, all of which are broken at middle. ioca%.— Urubamba, 9,500 feet, July. (M. C. Z. 152). Amaurobius platei Cambridge. Journ. Linn. soc. London, 1898, 27, p. 18, pi. 2, f. 3. One specimen probably this species was secured at Ollantaytambo, 9,000 feet in July. (M. C. Z. 166). Aymarella,^ gen. nov. Pars cephalica relatively broad, convex, and high. Anterior row of eyes substraight or slightly procurved, the medians much smaller than the laterals, farther from the laterals than from each other. Posterior row of eyes recurved ; subequal, the medians ' Diminutive of Aymara. chamberlin: the arachnida, 209 farther from the laterals than from each other. Area of median eyes distinctly narrower in front than behind. Lateral eyes close together, not more than their radius apart. Clypeus narrower than the anterior lateral eyes. Chelicera with lower margin short, but little oblique, armed with two (or three) small teeth; upper margin with three teeth of which the median one is largest. Labium longer than wide, narrowed distad, obtuse. Length of legs in order I-IV, II, III. Robust. Moderately spined. Metatarsi much longer than the tarsi. Cribellum bipartite. Calamistrum uniseriate. Genotype. — Aymarella viunda, sp. nov. This genus would seem to be closely related to Calleva, a genus based by Simon upon an Argentine species; but it differs clearly in having the lateral eyes much closer together, in having the legs spined, and in having the metatarsi much longer than the tarsi. Callevopsis, a Chilean genus, differs, e. g., in having the eyes of the anterior row equidistant and the lower margin of the chelicera armed with four teeth. Aymarella munda,^ sp. nov, Plate 9, fig. 1-5. Carapace brown to fuscorufous with darker lines radiating from thoracic furrow. Sternum fuscous to nearly black; clothed with numerous black hairs. Legs testaceous, typically of rufous tinge distad on anterior pairs; femora darker, dusky to nearly black. Palpi fuscous. Chelicerae nearly black. Labium and endites very dark, pale across tips. Abdomen olive-grey; without distinct mark- ings but in some with a median hastate mark very vaguely outlined in dark and followed behind by several faint light colored chevron lines, the lines very thin, and also a small light dot each side of middle median mark. Posterior row of eyes decidedly longer than the anterior (3.4:2.9), a little recurved; median eyes smaller than laterals (7:9), L7 their diameter apart, a third farther from the laterals. Anterior row of eyes viewed from above with line of apices a little recurved; viewed from in front the line of centres is a little procurved; median eyes ' mundus, fine, neat. 210 bulletin: museum of comparative zoology. with diameter about three fifths that of the laterals ; laterals on well- marked tubercles. Area of median eyes narrower in front than behind in ratio 10:13. Lateral eyes usually their radius apart, the distance being somewhat variable. Pars cephalica large, conspicuously elevated, highest immediately behind eye-area, smooth and shining, (Plate 9, fig. 1). Sternum longer than wide in about ratio 5:4. Caudal process short, narrow, (Plate 9, fig. 2). Labium grooved across base as usual; narrowed conspicuously from above groove distad; distally obtuse, (Plate 9, fig. 3). Tarsus of palpus armed with a claw and a number of spines ; other joints unarmed. Femur I armed with a single spine on anterior surface toward distal end; other femora unarmed. Patellae unarmed. Tibia III armed on anterior surface in subapical position with one spine; other tibiae unarmed. Metatarsi I and II armed ventrally with three pairs of spines; IV also with spines which are not distinctly arranged in pairs but are more distributed along surface of joint. Superior margin of furrow of chelicera armed with three teeth of which the median is largest; lower margin nearly transverse and with two rather small teeth. Abdomen broadly subelliptic in outline. Cribellum bipartite, (Plate 9, fig. 5). Feviale {San Miguel). Length 8 mm. Length of cephalothorax 3.2 mm.; width, 2.2 mm. fern. tib. + pat. met. tar. total Leg I 2.5 mm. 3 mm. 2 mm. 1.2 mm. 8.7 mm. Leg II 2.25 2.7 1.9 1 7.85 Leg III 2.1 2.2 1.5 1 6.8 Leg IV 2.5 3.1 2 1.1 8.7 Localities.— San Miguel, 6,000 feet, July. (Type, M. C. Z. 153 female; paratypes, M. C. Z. 154, females). Torontoy, 8,000feet, July. (M. C. Z. 155, several females). Lucma, 7,000 feet, August (M. C. Z. 156). Urubamba, 9,500 feet, July. (M. C. Z. 157, one female). Huadquina, 5,000 feet, July. (M. C. Z. 158, one female). Ollan- taytambo, 9,000 feet, July. (M. C. Z. 159, one female). AuxiMUS sp. A single immature male of a species of this genus was secured at Lucma in August. (M. C. Z. 160). chamberlin: the arachnida, 211 AuxiMUS PRODUCTUS, sp. nov. Plate 8, fig. 5-7. Carapace dusky brown or fuscous. Caelicerae similarly dusky over a more reddish or chestnut background. Legs testaceous; the tibiae, tarsi, and metatarsi of legs I and II dusky, almost black or tibia with an apical and a broader subbasal dark annulus indistinctly set off; tibiae of legs III and IV with apical and subbasal dusky annu- lus distinct and metatarsus of these legs with an apical and a broader, less distinctly limited subbasal dark band. Patellae dusky. Femora without rings or distinct marks, usually dusky beneath proximally. Tibia and tarsus of palpus dusky or almost black like the distal articles of anterior legs. Sternum dusky brown. Labium and endites darker than sternum, pale across tips. Abdomen above with background of greyish yellow of slight olivaceous tinge; at base above with a black median sagittate stripe the acute apex of which is at middle; this mark is followed caudad by a series of four, successively smaller and smaller chevron-shaped black lines the angle of the first of which is at the point of the sagittate mark; chevron marks with ends on each side united by a longitudinal black line. Dorsum elsewhere with small black, in part angular, dots, these more numerous on the sides where they are confluent with enclosed areas dusky and entire surface thus appearing blackish. A broad median band along venter immacu- late greyish yellow of slight olivaceous cast like background of dorsum. Posterior row of eyes considerably procurved, a line through middles of median eyes being nearly tangent to caudal edges of laterals; median eyes clearly less than twice their diameter apart (11:7) and nearly the same distance from the laterals; smaller than the laterals (7:9). Anterior row of eyes with line of apices as seen from above very slightly procurved; viewed from in front the row is nearly straight, the centres of the laterals being slightly lower; median eyes with diameter half as great as that of laterals, four fifths their diameter apart, nearly same distance or slightly farther from laterals. Ante- rior laterals equal to posterior laterals from which they are separated by a distance not more than equalling half the radius. Labium about two thirds as broad at base as long; narrowed distad; apically truncate. Sternum longer than wide, the length inclusive of caudal process being to width as 9 : 6.5 and the length only to level of anterior proximal corner of coxa IV being as 7:6.5. 212 bulletin: museum of comparative zoology. Abdomen in outline broadly subovate. Lower margin of furrow of chelicera with six teeth of which the two farthest from claw are much smaller and the first is smaller than the succeeding three. All femora dorsally with a submedian spine and at distal end with one toward anterior side and there may be a smaller one in corre- sponding position on posterior side. Tibia I with a submedian pair of spines on ventral side and a single one on anterior side between middle and proximal end. Tibia II with a single submedian ventral spine; and tibiae III and IV unarmed. Metatarsus I with three pairs of ventral spines. Femur of palpus with a spine in submedian position above; tibia with three long, almost bristle-like, spines, two toward mesal side above and one ectal in position; tarsus with numerous spines, the proximal ones long and slender, the distal ones shorter and stouter. Female. Length, 7.5 mm. Length of cephalothorax, 3 mm.; width, 2.5 mm. fern. tib.+pat. met. tar. total Leg I 2.4 mm. .3.25 mm. 2.1mm. 1.1mm. 8.85 ram. Leg 11 2.2 2.9 2 1.1 8.2 Leg III 2.2 2.2 1.6 1 7.0 Leg IV 2.4 2,9 2 1.25 8.55 Locality.— Tincochaca, 7,000 feet, August. (Type, M. C. Z. 161, one female). While a number of species of Auximus have been described from Ecuador, only one was previously known from Peru, A. funestus (Keys.), a species differing rather widely from the present one in hav- ing its posterior row of eyes straight, in the sparser spining of legs, in the different form of epigynum as well as in color. DiCTYNA HESPERIA,^ sp. nOV. Plate 8, fig. 8. Thoracic part of carapace fuscous, yellow along edges; head testa- ceous. Legs and palpi yellow, the tarsi in part dusky. Sternum yellow, faintly dusky. Labium dusky yellow. Abdomen opaque yellowish white, venter covered with a broad, more or less dusky band, a more blackish area about the spinnerets; above there is a ' flffo\os, ambiguous. 296 bulletin: museum of comparative zoology. Tooth of lower margin of chelicera (female) short, stoutly conical, acute; teeth of upper margin two, small (or an obsolete third tooth may be present). Ocular quadrangle much wider than long, conspicuously wider behind than in front, with the minute second eyes in front of the middle as usual. Eyes of the first row as in the preceding species. Length of female 5 mm.; length of cephalothorax 2 mm.; width, 1.4 mm. Localities. — Huadquina, 5,000 feet, July. (Type, M. C. Z. 302, female; para types, M. C. Z. 303, six females). Paltaybamba, 5,000 feet, August. (M. C. Z. 304, three females). Dendryphantes sp. a. A female from Huadquina, 5,000 feet, July. (M. C. Z. 309) cannot be satisfactorily determined. Dendryphantes sp. b. An immature female of doubtful species from San Miguel, 6,000 feet, July. (M. C. Z. 310). Wala sp. An immature male and female of uncertain species from San Miguel, 6,000 feet, July. (M. C. Z. 305). Wala noda,^ sp. nov. ^ Plate 25, fig. 2. Carapace with integument brownish black, rubbed in part but apparently clothed in life with greyish scales which on the head are more brownish. Legs brown. Sternum brownish black. Abdomen grey-brown beneath. x\bove clothed with light grey scales except for two pairs of elongate subtriangular dark areas of which the apices are directed forwards and the edges are curved; a narrow chevron-mark between the two pairs of dark spots. 1 puio^, edentate. chamberlin: the arachnida. 297 Lower margin of furrow of chelicera unarmed or with but a minute pale rudiment of a tooth. Tibia I armed beneath with three pairs of spines; metatarsus I with two longer pairs; tibia II beneath with an apical pair and a single spine toward middle in the usual way. Ocular quadrangle much wider than long (13:9), fully as wide in front as behind. Eyes of second row slightly in front of the middle. Length of female, 6 mm. ; cephalothorax, in length, 2.8 mm. ; width, L8 mm. Localiiy.— Torontoy, 8,000 feet, July 22. (T^-pe, M. C. Z. 306, female; paratype, M. C. Z. 307, one female). One female from the Conservidayo River in bad condition is this species or very close to it. EvoPHRYS CRUX Taczanowski. Bull. Soc. imp. nat. Moscow, 1878, p. 284. Described originally from Amable Maria, Peru. Locality — Tincochaca, 7,000 feet, August. (M. C. Z. 311, female). EvoPHRYS PERUVIANA Taczanowski. Bull. Soc. imp. nat. Moscow, 1878, p. 280. Previously known from Amable Maria and Pumamarca, Peru. Locality. — Conservidayo River, August. (M. C. Z. 312, four females) . EvoPHRYS sp. a. Locality — Santa. Ana, 3,000 feet, August. (M. C. Z. 313, one immature female). EvoPHRYS sp. b. Zoca%.— Santa Ana, 3,000 feet, August. (M. C. Z. 314, one immature female). EvoPHRYS sp. c. Locality.— Lucma, 6,000-7,000 feet, August. (M. C. Z. 315, one immature male in coloration much like E. crux Tacz.). 298 bulletin: museum of comparative zoology. EvoPHRYS sima/ sp. nov. Plate 25, fig. 1. Carapace with a broad median longitudinal light band anteriorly as broad as the ocular quadrangle but narrowing caudad to a point at the caudal edge, this band reddish on head and becoming yellow caudad; sides blackish brown with a narrow supramarginal pale stripe on each side. Sternum yellow. Last three pairs of legs yellow; first pair of legs with femora reddish brown, more yellow distally, the more distal joints light brown, the metatarsus clothed with a brush of long spatulate hairs on ventral surface, similar but more sparse ones on ventral surface of femur and patella. Palpi reddish brown, the femur with numerous white hairs above and with black scopulate hairs below which also occur on other joints. Dorsum of abdomen black with a narrow median longitudinal yellow stripe over the entire length, the edges of this stripe dentate along caudal half of length; sides yellow finely dotted with black and a more solid stripe below at the anterior end ; venter yellow with a few dots at the sides. Spinnerets dusky. Tibia I with the usual three pairs of ventral spines; one small spine toward the distal end on lower anterior surface. Metatarsus I with two pairs of spines as usual. Ocular quadrangle a little wider in front than behind and rather more than one fourth wider than long. Eyes of second row caudad of middle. Eyes of first row nearly contiguous, medians twice the diameter of the laterals. Length of male, 4 mm. Length of cephalothorax, 2.2 mm.; width, L6 mm. Locality.— San Miguel, 6,000 feet, July. (M. C. Z. 316, one male). Phiale panamae, sp. nov. Plate 25, fig. 4. Carapace black with a reddish yellow median longitudinal stripe crossed by a black band at level of the third eye row and narrowing to the posterior margin; also a reddish supramarginal stripe on each side. Sternum yellow, dusky at margins. Legs and palpi dusky brown, the posterior pairs somewhat paler. Spinnerets and venter ' Gosiute sima, one. chamberlin: the arachnida. 299 of abdomen dusky ; dorsum black, the anterior face and anterolateral corners and sides whitish, three pale spots in a triangle near middle and several fainter, smaller ones farther caudad. Ocular quadrangle slightly wider in front than behind; eyes of second row minute, behind middle. First row of eyes decidedly recurved ; eyes contiguous or nearly so, the median eyes with diameter two and a half times that of the laterals. Femora I and II with three moderately long spines at the distal end on anterior part of dorsal surface and with two spines in the middorsal line farther proximad. Tibia I with the usual three pairs of rather short ventral spines and two on anterior surface ; metatarsus I with two pairs of ventral spines and one on anterior surface at the distal end. Tibia of palpus with apophysis at distal end from dorsoectal corner subconical, distally curved. Length of male, 4 mm.; length of cephalothorax, 2.1 mm.; width, 1.3 mm. Locality. — Panama, June. (M. C. Z. 317, one male). Phiale huadquinae, sp. nov. Plate 25, fig. 3. ' Carapace with integument solid shining black, a band of white hair along each side. Sternum and mouthparts also black, the endites pale across tips. Last two pairs of coxae yellowish beneath, the anterior pairs darker; femora, patellae, and tibiae black. Metatar- sus black distally, testaceous proximally; tarsi testaceous. Integu- ment of abdomen black clothed with hair of golden brown lustre; hair of venter grey. Lower margin of furrow of chelicera with one stout tooth; upper margin with two teeth united at base. Ocular quadrangle wider in front than behind (46:43). Eyes of second row minute, at middle of length of quadrangle or scarcely in front. Anterior row of eyes strongly recurved; median eyes a little more than twice the diameter of the lateral, about one seventh their diameter apart and from the laterals. Tibia I with three pairs of spines below and three single ones on anterior surface; metatarsus with two pairs beneath. Length of male, 7 mm.; length of cephalothorax, 3.1 mm.; width, 2.1 mm. Locality. — Huadquina, 5,000 feet, July. (M. C. Z. 318, one male). PLATE 1. •Chamberlin. — The Arachnida. PLATE 1. Tityus foot&i Chamberlin. Fig. 1. Lateral view of sting and last two segments of abdomen. Fig. 2. Finger, inner surface, showing arrangement of granules. Fig. 3. Tarsus IV, anterior view. Fig. 4. Comb. Brachistostermis andinus Chamberlin. Fig. 5. Carapace, dorsal view. Fig. 6. Finger, inner view, showing granules. Fig. 7. Tarsus IV. Fig. 8. Comb. BULL. MUS. COMP. ZOOl. Chamberlin— Peruvian Arachnioa. Plate 1 R V. C.DEL. PLATE 2. Chamberijn.— The Arachnida. PLATE 2. Paravanones peruviantis Chamberlin. Right mandible, ectal view. Right pedipalp, ectal view. Anterior portion of carapace from above and a httle to the left. Femur and adjoiaing parts of leg IV, the proximal portion in dorsal view, the distal portion in more dorsoectal view. Trochanter IV (left), ventral view. Showing process of coxa IV, dorsal view. Gonoleptes enoplus Chamberlin. Fig. 7. Dorsal view. Fig. 8. Mandible, ectal view. Fig. 1. Fig. 2. Fig. 3. Fig. 4 Fig. 5. Fig. 6. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnioa. Plate 2 R V. C.DEL. PLATE 3. Chamberun. — The Arachnida. PLATE 3. Gonoleptes enoplns Chamberlin. Fig. 1. Coxa and trochanter of left pedipalp, ectal view. Fig. 2. Left pedipalp, excepting proximal joints, ectal view. Fig. 3. Leg III) distal portion, anterior view. Fig. 4. Right leg IV, dorsoectal view. Fig. 5. Coxal spur of leg IV, lateral view. Goiwleptes scolius Chamberlin. Fig. 6. Dorsal view. Fig. 7. Mandible, ectal view. Fig. 8. Left leg III, anterior view. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida, Plate 3 H. V. C..DE1.. ^ PLATE 4. Ghamberlin. — The Arachnida. PLATE 4. Gonoleptes scotiiLs Chamberlin. Fig. 1. Left leg IV, subdorsal view. Fig. 2. Right pedipalp, ectal view. Gonoleptes huadquinae Chamberlin. Fig. 3. Mandible, ectal view. Fig. 4. Right pedipalp, ectal view (Tjrpe). Fig. 5. Right pedipalp, ectal view (Paratype). Fig. 6. Caudal processes of carapace, anterodorsal view (Paratype). Fig. 7. Left process, lateral view. Fig. 8. Left leg IV, dorsoectal view. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida. Plate 4 R V. C.,DEL. PLATEjS. Cbamberlin. — The Arachnida. PLATE 5. Pachylus orinus Chamberlin. Fig. 1. Dorsal view. Fig. 2. Right mandible, ectal view. Fig. 3. Right pedipalp, ectal view. Liopagus simplex Chamberlin. Fig. 4. Eye-tubercle, caudal view. Fig. 5. Eye-tubercle, lateral view. Fig. 6. Right mandible, ectal view. Fig. 7. Right pedipalp, mesal view. Fig. 8. Part of left pedipalp, dorsal view. Ldobunum monticola Chamberlin. Fig. 9. Left pedipalp. ectal view. BULL. MUS. COMP. ZOOL. Chamberlin,— Peruvian Arachnida, Plate 5 R. V. C.,DEL. PLATE 6. Chamberum. — The Arachnida PLATE 6. lAobunum monticola Chamberlin. Fig. 1. Lateral view of eye-tubercle. Fig. 2. Left mandible, mesal view. Fig. 3. Left mandible, ectal view. Hemirrhagus peruvianvs Chamberlin. Eyes, dorsal view. Tarsal scale of usual type. Tarsal scale of a second type. Tibial spurs, right leg I of male. Claw of leg IV. Lock of tarsometatarsal joint (tarsus above). Palpal organ of male, left ectal view. Fig. 4. Fig, 5. Fig. 6. Fig. 7. Fig. 8. Fig. 9. Fig. 10. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida. Plate 6 9 9 R. V. C..DFL. PLATE 7. Chamberlin. — Tho Arachnida. PLATE 7. Hemirrhagus peruvianns Chamberlin. Fig. 1. Anterior claw of leg I. Fig. 2. Posterior claw of leg I. Hemirrhagus viajor Chamberlin. Fig. 3. Eyes (Type, Urubamba). Fig. 4. Tibial spurs, left leg I of male, mesal view. Fig. 5. Caudal claw, leg I. Fig. 6. Claw of leg III, female. Fig. 7. Claw of leg III, male. Fig. 8. Right palpal organ of male, ectal view (T3T)e). Diplura monticolens Chamberlin. Fig. 9. Eyes, dorsal view. Fig. 10. Eyes from in front and a Uttle above. Brachythele keithi Chamberhn. Fig. 11. Eyes, dorsal view. Brachythele incursiis Chamberhn. Fig. 12. Eyes, dorsal view (more enlarged than fig. 11). BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida. Plate 7 R. V. CDEL. i PLATE 8. Ch.\mbeklin. — The Aracbnida. PLATE 8. Orinomus lamprus Chamberlin. Fig. 1. Lateral view of body, Fig. 2. Sternum. Fig. 3. Epigynum. Fig. 4. Labium. AuximiLS productus Chamberlin. Fig. 5. Sternum. Fig. 6. Cribellum. Fig. 7. Epigyn\un, caudoventral view. Dictyna hesperia Chamberlin. Fig. 8. Epigynum. SULL. MUS. COMP. 200L. Chamberlin— Peruvian Arachnida, Plate 8 6 8 R V. C.DEL. PLATE 9. Chamberlin. — The ArachnMa. PLATE 9. Aymarella munda Chamberlin. Fig. 1. Lateral view of cephalothorax and mandibles Fig. 2. Sternum. Fig. 3. Labium. Fig. 4. Epigynum. Fig. 5. Cribellum. Thomisoides terrosus Nicolet. Fig. 6. Eyes, dorsal view. Fig. 7. Mandible, ectal view, showing stridulating plate. Fig. 8. Labium. Fig. 9. Distal portion of mandible, ventral view. Fig. 10. Posterior claw of leg L BULL. MUS. COMP. ZOOL. Chamberlin.— Peruvian Arachnida. Plate 9 ^.r^- R- V. C..DEL. Obamberun. — The Aracbnida. PLATE 10. Thomisoides terrosus Nicolet. Fig. 1. Dorsal view. Fig. 2. Base of spine from leg. Fig. 3. Distal end of bristle from leg. (Figs. 1 to 3 drawn by Prof. A. Petrunkevitch). Ariadna hotchkissi Chamberlin. Fig. 4. Eyes, dorsal view. Fig. 5. Labium. Nops beUiUa Chamberlin. Fig. 6. Sternum. Fig. 7. Eyes and anterior part of carapace. Fig. 8. Labium and left endite. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida, Plate 10 R V. C.GEl.. PLATE 11. CHAMBEnLirv. — The Arachnida. PLATE 11. Nops bellnla Chamberlin. Fig. 1. Dorsal view of abdomen. Fig. 2. Distal portion of right leg I, caudal view. Fig. 3. Mesal and anterior claw of leg I, caudal view. Drassodes araucanius Chamberlin. Fig. 4. Eyes, dorsal view. Fig. 5. Sternum. Fig. 6. Labium. Fig. 7. Epigynum. Fig. S. Hair of scopula, highly magnified. BULL. MUS. COMP. ZOOL. Ghamberlin— Peruvian Arachnida^ Plate 1 1 R V. C.DEL. PLATE 12. Cbamberlin. — The Arachoida. PLATE 12. Drassodes araucanius Chamberlin. Fig. 1. Epigj-num, possibly lacking one moult of maturity. Fig. 2. Tarsal claw. Apodrassus andiniis Chamberlin. Fig. 3. Sternum. Fig. 4. Hair of fascicula of claw. Fig. 5. Eyes, dorsal view. Fig. 6. Labium. Fig. 7. Claw of leg I. Fig. 8. Spermathecae. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida, Plate 12 "•^"wiltrli^iii- iivr->.-.Mt> 6 8 R V. C. DCL. PLATE 13. Chamberlin.- — The Arachnida. Fig. 1 Fig. 2. Fig. 3. Fig. 4. Fig. 5. Fig. 6. Fig. 7. Fig. 8 Fig. 9. PLATE 13. Hypsorinus binghamae Chamberlin. Abdomen, lateral view. Eyes, dorsal view. Sternum, labium, etc. Endite, subventral view. Tip of same, sublateral view. Patella, tibia and tarsus of palpus of adult female. Unpaired claw of leg I. Epigynum. Serrated bristle or accessory claw of tarsus I. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida. Plate 13 R. V. C.DEL. PLATE 14. Cbamberlin. — The Arachoida. PLATE 14. Hypsorinus binghamae Chamberlin. Fig. 1. Feather hair from near claw of leg I. Fig. 2. Bristle from end of tarsus I above claws. Fig. 3. End of chelicera of male from above. Fig. 4. Same of female from above. Fig. 5. Claw of leg I, female. Fig. 6. Palpus of male, ectal view. Fig. 7. Sfame, from above. Litoporus aberrans Chamberlin. Fig. 8. Sternum and labium. Fig. 9. Abdomen, lateral view. BULL. MUS. COMP. ZOO! Chamberlin— Peruvian Arachnida. Plate 14 i^ V. C .01:.. PLATE 15. Cb AM BERLIN. — The Arachnida. PLATE 15. Litoporus aberrans Chamhei-lin. Fig. 1. Eyes from above. Fig. 2. Right palpus of male, ectal view. Fig. 3. Palpus of male, anterior view. Argyrodes lucmae Chamberlin. Fig. 4. Sternum. Fig. 5. Abdomen, lateral view. Fig. 6. Right palpus of male, dorsal view. Theridion leguiai Chambeilin. Fig. 7. Sternum. Fig. 8. Eyes from above. Fig. 9. Middle and anterior claw, caudal view. Fig. 10. Epigynum. k. BULL. MUS. COMP. 200L. Chamberlin.— Peruvian Arachnida, Plate 15 O Or.1 O GO R. V. C.-DEL. PLATE 16. Chambeblin. — Tlie Arachnida. PLATE 16. Theridion tosum Chamberlin. Fig. 1. Labium. Fig. 2. Sternum. Fig. 3. Eyes from above. Fig. 4. Epigynxim. Garricola sanctiis Chamberlin. Fig. 5. Lateral view of body. Fig. 6. Sternum. Fig. 7. Epigyniun. Enoplognatha peruviana Chamberlin. Fig. 8. Sternum, Fig. 9. Paired claw of leg I. Fig. 10. Unpaired claw. Fig. 11. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin— Peruviaini Arachnida, Plate 16 ^:' ■■■■:■■■■ ■ •■ 1!!5'5»^ '^^-. R. V. C, DEL. PLATE 17. Chamberlin. — The Arachnida PLATE 17. Enoplognatha peruviana Chamberliii. Fig. 1. Eyes dorsal view (anterior row above). Fig. 2. Labium. Enoplognatha dubia Chamberlin. Fig. 3. Epigynum. Erigone niwina Chamberlin. Fig. 4. Epigynum. Erigone taibo Chamberlin. Fig. 5. Epigynum. Oedothoraz melacra Chamberlin. Fig. 6. Palpus. Fig. 7. Epigynum. Oedothorax orinus Chamberlin. Fig. 8. Epigynum. Tutibo debUipes Chamberlin. Fig, 9. Epigynum. Fig. 10. Palpus, distoventral view. BULL. MUS. COMP. ZOOL. Chamberlin— Peruv'an Arachnida, Plate 17 9 9 R- V. C. PEL. PLATE 18. Chambeblin. — The Arachnida. PLATE 18. Tetragnatha scopus Chamberlin. Fig. 1. Chelicera from above. Tetragnatha tincochacae Chamberlin. Fig. 2. Chelicera of male from above. Fig. 3. Palpus of male, mesoventral view. Tetragnatha quechua Chamberlin, Fig. 4. CheUcera from above. Meta explorans Chamberlin. Fig. 5. Epigyiimn. Acacesia -peruviana Chamberlin. Fig. 6. E}pig>Tium. Eustala monticola Chamberlin. Fig. 7. Epigynum. Aranea duocypha Chamberlin. Fig. 8. Abdomen, dorsal view. Fig. 9. Sternum. Fig. 10. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin.-Peruvian Arachnida, Plate 18 5 R V. C. DEL, PLATE 19. Crambbhi.in. — The Aracliuida PLATE 19. Aranea quechuana Chamberlin. Fig. 1. Palpal organ of male, meso ventral view. Aranea tigana Chamberlin. Fig. 2. Palpal organ of male, mesoventral view. Aranea orina Chamberlin. Fig. 3. Palpal organ of male, mesoventral view. Aranea calotypa Chamberlin. Fig. 4. Palpal organ of male, mesoventral view. Aranea plesia Chamberlin. Fig. 5. Epigynum. Aranea compsa Chamberlin. Fig. 6. Epigynum. Aranea sexta Chamberlin. Fig. 7. Dorsal view of abdomen. Gea panamensis Chamberlin. Fig. 8. Left palpal organ of male, mesoventral view. Scoloderus hyhus Chamberlin. Fig. 9. Epigynum. Aranea santa Chamberlin. Fig. 10. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian arachnida. Plate 19 R, V. C.DEL. PLATE 20. CuAMBERLiN. — The Arachnidu. PLATE 20. Anamxia atopa Chamberlin. Fig. 1 . Eyes from above and a little to the left. Fig. 2. Abdomen, lateral view. Fig. 3. Right leg II; anterior view of tibia, etc. Micrathena cola Chamberlin. Fig. 5. Dorsal view of abdomen. Gelanor innominatum Chamberlin. Fig. 6. Epigynum. Eusparassus shefleli Chamberlin. Fig. 7. Sternum. Fig. 8. Labium. BULL. MUS. CO MP. ZOOl Chamberlin.—PcR'jvian Arachnida. Plate 20 R V. C..DEL. PLATE 21. Chambe:ri.in. — The Arachnida. PLATE 21. Eusparassus shefteli Chamberlin. Fig. 1. Epigynum. Horiocte7ms lycosoides Chamborlin. Fig. 2. Labium. Fig. 3. Epigynum. Fig. 4. Distal portion of chelicera, ventral view. Quechuella Imnpra Chamberlin. Fig. 5. Labium. Fig. 6. Sternum. Fig. 7. Dorsal view of abdomen. Fig, 8. Eyes from in front and a little above. Trachelopachys bicolor Chamberlin. Fig. 9. Labium, Fig. 10. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin.— Peruvian Arachmida, Plate 21 4 0 ^ ^ 8 OgoO 6 9 R V. C.,DEL. PLATE 22. Chamberun. — The Arachnida. PLATE 22. Castaneira quechua Chamber lin. Fig. 1. Epigynnm. Anyphaena apora Chamberlin. Fig. 2. Labium. Fig. 3. Epigynum. Anyphaena andina Chamberlin. Fig. 4. Epigynum. Anyphaena poicila Chamberlin. Fig. 5. Dorsal view of abdomen. Gayenna monticola Chamberlin. Fig. 6. Epigynum. Tunabo peruvianus Chamberlin. Fig. 7. Labium. Fig. 8. Sternum. Fig. 9. Distal portion of chelicera, ventral view. BULL. MUS. COMP. ZOOL. Chamberlin.-Peruvian Arachnida. Plate 22 6 _^j^es2rEs!3EgsSEi!st_ 2 m 6 R. V. C..DEL. PLATE 23. Chamberlin. — The Arachnida. PLATE 23. Trechalea monticola Chamberlin. Fig. 1. Paired and single claw. Porrima harknessi Chamberlin. Fig. 2. Labium. Fig. 3. Male palpal organ, ventral view. Fig. 4. EpigjTium. Fig. 5. Lorum of pedicel. Fig. 6. Claws. Arctosa altamontis Chamberlin. Fig. 7. Epigynum. Fig. 8. Male palpal organ, ventral view. Fig. 9. Labium. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida. Plate 23 2 8 9 R. V. C.DEL. PLATE 24. Chambeiii.in. — The Arachnids. PLATE 24. Lycosa orinus Chamberlin, Fig. 1. Epigynum. Lycosa andina Chamberlin. Fig. 2. Epigynum. Fig. 3. Palpal organ, ventral view. Lycosa liopus Chamberlin. Fig. 4. Epigynum. Lycosa algina Chamberlin. Fig. 5. Labium. Fig. 6. Palpal organ. Orinocosa aymara Chamberlin. Fig. 7. Labium. Fig. 8. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin.-Peruvian Arachnida, Plate 24 H V. C.DEL. PLATE 25. Chamberun. — The Aracbnida. PLATE 26. Evophrys sima Chamberlin. P'ig. 1. Palpal organ, ventral view. Wala noda Chamberlin. Fig. 2. Epigynum. Phiale huadquinae Chamberlin. Fig. 3. Palpal organ. Phiale panamae Chamberlin. Fig. 4. Palpal organ of male. Dendryphantes andinus Chamberlin. Fig. 5. Palpal organ of male. Fig. 6. Distal end of chelicera of male, caudal view. Dendryphantes cuius Chamberlin. Fig. 7. Palpal organ of male, subventral view. Fig. 8. Distal end of chelicera of male. Dendryphantes amphiholus Chamberlin. • Fig. 9. Epigynum. BULL. MUS. COMP. ZOOL. Chamberlin— Peruvian Arachnida, Plate 25 R V. C.DEL. ^^ ibio Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE. * Vol. LX. No. 7. A COLLECTION OF BIRDS FROM THE CAYMAN ISLANDS. By Outram Bangs. CAMBRIDGE, MASS., U. S. A.: PRINTED FOR THE MUSEUM. March, 1916. No. 7. — A Collection of Birds from the Cayman Islands. By Outram Bangs. In the year 1911 the well-known collector, W. W. Brown, Jr., spent the spring and early summer, April to July, in the Caymans. He visited all three islands and made a practically complete collection of the resident, breeding land birds of the group. This beautiful lot of skins, in Brown's inimitable make, fortunately remained intact and was secured for the Museum of Comparative Zoology. Below I give an annotated list of the collection which represents only the resident ornis of the islands, being happily free from migrants. In the Ibis for 1911, (ser. 9, 5, p. 137-161), Mr. P. R. Lowe published a list of the birds of the Cayman Islands. Lowe's account of the islands and his description of them is so good and complete as to leave nothing more to be said. He also brought up to date all bird collecting that had been done there. I must, however, give my opinion upon the sources whence the bird life peculiar to the Caymans has been derived. Lowe justly states that on account of the very recent origin of the islands no genus and no very peculiar forms occur there alone. In the main, this is true, but I think for the moment he had forgotten Mimo- cichla ravida. This bird bears no close relationship to any other existing species of the genus. We must, however, bear in mind that Jamaica at present, alone among the Greater Antilles, is without a species of Mimocichla. It is highly probable that a form similar to M. ravida once occurred there and that the Cayman bird, now itself on the verge of extinction, was derived from that form. Coereba sharpei is a species of uncertain origin. The genus Coereba has been in the near past, and perhaps is still, so very plastic that the relationships of the various forms are hard to trace. Dendroica viteUina (which also occurs in Swan Island) and D. crawfordi, quite clearly indicate an instance, rather rare among birds, of a migratory species establishing itself upon islands that lie on the line of passage and becoming differentiated there; for clearly the nearest relation of these two wood-warblers is the migratory North American Dendroica discolor. Three other peculiar forms were, I believe, received directly from 304 bulletin: museum of comparative zoology. the near by mainland. These are the Elaenia which is much more Uke E. martinica siibpagana, than it is any of the gray Lesser Antillean forms; the Vircosyha caymancnsis which is very closely related to V. magister of the coast of British Honduras and unlike any West Indian form; and the Vireo, which so far as I can see is identical with V. crassirostris the bird inhabiting the Bahamas (which are of simi- lar formation). This species is so much like V. ochraccus of the opposite coasts of Central America and so unlike any of the species peculiar to the Greater Antilles, — Cuba, Jamaica, or Porto Rico, that there seems no question of its origin. The remainder of the Cayman birds have come from either Jamaica or Cuba, in some case being still identical with the parent stock, in others having differentiated into what may be called island species or subspecies according to the degree of change. From Jamaica the Caymans have received Leptoiila coUaris and Icterus bairdi. From Cuba the islands have derived the two forms of Amazona peculiar to them, CoJaptcs guruJIachi, Ccnturus caj/mancusis, Mimocichia cori/i, two forms of Holoquiscalus, Spindalis salvini, and Mdopyrrha faylori; probably also Tohnarckus caymanensis, although this species might be descended from either T. caudifasciatus of Cuba or T. janiaiccnsis of Jamaica. From the above which discusses every bird peculiar to the Caymans it will be seen that I am unable to recognize several forms which ha\e been described as species or subspecies peculiar to the islands, and these I comment upon at length in the following list. At the time of Brown's \isitto Little Cayman and Cayman Brae — June and July — the Boobies and Man-o'-War-Birds were not breed- ing and all he saw during his stay were occasional birds off shore. Besides these and the list of species following he saw and positively identified only two species, namely Hydranassa tricolor ruficoUis (Gosse) and Nyctanassa violacea (Linn,e). Brown took nests and eggs of a number of the species; these are preserved in the Museum of Comparative Zoology, and I believe some of them have not before been collected. I have marked Avith an asterisk each species of which he secured the nest and eggs; and with a dagger the species of which the eggs only were taken. As this paper was going to press, an article on the birds of Grand Cayman appeared in the Ibis, January 1916, p. 17-35, by T. M. Savage English. Mr. English apparently collected no specimens, but based all his identifications on living birds observed afield. During his three years' residence in the island he was able to add twelve species bangs: birds from the cayman islands. 305 to Lowe's list. A few of these are merely migrants. Four others I had already included in the following account of Brown's collection, and I let my words stand as first written. It would be of interest to know the bird that Mr. English found breeding in Grand Cayman and called Chordeiles virginicmus, distin- guishing it from Chordeiles minor by its larger size. On geographic grounds it certainly could not have been Chordeiles virginianus vir- ginicmus (Gmelin). Ardeidae. *BUTORIDES VIRESCENS BRUNESCENS (Lemb.). Two specimens, an adult cf and an immature 9 , Grand Cayman, May. These are similar to specimens from Jamaica and Cuba. I have already (Auk, Oct. 1915, 32, p. 481-484) given my reasons for using the name brunescens for the Green Heron of the Greater Antilles. Whether or not that form can be maintained as distinct from B. v. vutculatus (Bodd.) of Martiniciue remains, I think, to be proved. Oberholser has probablv' subdivided the Green Heron too much, and he had but three specimens from Martinique when he wrote his Revision of the subspecies of the Green Heron (Proc. U. S. N. M., 1912, 42, p. 529-577). Rallidae. *Gallinula chloropus CACHiNNANS Bangs. Brown found the Gallinule breeding in abundance in the many marshy ponds in Grand Cayman in April and May, and took several sets of eggs, but made up no skins. Laridae. t Sterna antillarum (Lesson). One adult cf , Little Cayman, July 26. Browfi found the Least Tern breeding in abundance. 306 bulletin: museum of comparative zoology. Charadriidae. Charadrius wilsonius Ord. Brown noted Wilson's Plover on several occasions on the beaches, where he thought it was breeding. He took no specimens. I include it in this list on Brown's identification in spite of the lack of specimens, in order to correct a rather curious error in Lowe's list where under Aegialitis semipalmata Lowe says, "This bird is resident in Jamaica and breeds there. Whether it does so in the Caymans I am unaware. I have included it among the residents." No mention is made of Wilson's Plover, and it seems certain that Lowe in some way confused that species with the Arctic-breeding Semipalmated Plover. t Hypsibates mexicanus (Miill.). The Black-necked Stilt was breeding in numbers in the mangroves in Grand Cayman in May. Brown took several sets of eggs, but did not shoot any birds. Columbidae. CoLUMBA leucocephala Linue. Three specimens, two males and a female, all adult, Grand Cayman and Cayman Brae, May and June. t Zenaida zenaida zenaida (Bp.). Zenaida spadicea Cory, Auk, 1886, 3, p. 498, Grand Cayman. Zenaida richardsoni Cory, Auk, 1887, 4, p. 7, Little Cayman. Thirteen specimens, both sexes, all adult. Grand Cayman, Little Cayman, and Cayman Brae, May, June, and July. I can find no difference in specimens from the various islands of the Cayman group, and after a most careful comparison, with adequate material, am unable to distinguish in any way Cayman specimens, which appear to me to be quite like examples from the Bahamas, Cuba, and Jamaica, in the same condition of plumage. Judging from Cory's name and description I fancy he took the Grand Cayman Pea Dove to be darker than true zenaida. This may have been because he compared Grand Cayman specimens killed when in worn summer plumage, with skins from elsewhere in autumn or winter dress, there being quite a change with season in bangs: birds from the cayman islands. 307 Z. zenaida. This is wholly due, I think, to wear and the loss of the bloom or sheen characteristic of the fresh plumage. * Chaemepelia passerina insularis (Ridg.). Eleven specimens, all adult males. Grand Cayman, Little Cayman, and Cayman Brae. April, May, and June. There is no difference in skins from the three islands. On comparing this series with an enormous number of Cuban birds, I could find no differences at all, in color, color of the bill, size, or anything else. I therefore sent the series to W. E. C. Todd, as the latest authority on this group of birds, and asked him for an opinion that I might quote in print. He replied that he was now fully prepared to say that the Cayman and Cuban forms are identical. Chaemepelia passerina afiavida (Palmer and Riley) therefore becomes a synonym of P. p. insularis (Ridg.). The Jamaican form, though very close still appears to be recognizable. • t Leptotila jamaicensis collaris (Cory). Five specimens, both sexes, all adult. Grand Cayman, May and June. Brown considers this Dove to be the rarest of all the peculiar Grand Cayman birds. The five specimens he took were the result of days spent hunting especially for it in its favorite haunts. I can detect no differences whatever in color in comparing these Grand Cayman skins with our twelve specimens of true L. jamaicensis (Linne) from Jamaica. The Cayman examples appear to have less white at the tips of the three outer rectrices. This may be partly, perhaps wholly, due to their tail feathers being more worn down at the ends. The Cayman specimens also average slightly smaller than Jamaican ones, as the following measurements show, but the differ- ence is so trifling that a larger amount of material might actually tiu-n it the other way, and I doubt much if the form can be maintained. Grand Cayman. Exposed No. Sex Wing Tail Tarsus Culmen 68334 & 148 103 29 16 68335 cf 146 90 31 16 68338 d^ 153 105 31.5 15.5 68336 9 142 95 28 15.5 68337 9 144 102 27.5 16 308 > bulletin: a lUSEUM ( OF COMPARAT] Jamaica. [VE ZOOLOG Y. Exposed No. Sex Wing Tail Tarsus Cvlmen 37732 d^ 159 111 31.5 16.5 37733 cT 161 112 31 17 37734 d" 157 109 29 16 54089 cf 154 109 32 16 3695 d" 153 108 30 16.5 3696 d' 158 108 33 17 37735 9 153 107 29 16 37736 9 151 105 29.5 16.5 37737 9 153 100 29 16 41841 9 159 106 32 16.5 54088 9 151 99 28.5 16 71554 9 151 96 28.5 16 PSITTACIDAE, t x\mazona leucocephala caymanensis (Cory). Ten specimens, both sexes, all adult, Grand Cayman, May and June. This well-marked form is peculiar to Grand Cayman. I agree with Todd (Annals Carnegie mus., 1911, 7, p. 418) that its relationship to true leucocephala of Cuba, which is close, is best expressed by the use of trinomials. Brown noted that the skin of the orbital region, ^•aried from white to flesh-color. He took a set of four eggs together with the female parent on May 12. Amazona leucocephala hesterna, subsp. nov. Ten specimens, both sexes, all adult. Little Cayman and Cayman Brae, June and July. Type, adult d^, M. C. Z. 68313, Cayman Brae, July 15, 1911, W. W. Brown, Jr. Characters. Similar to true A. leucocephala (Linne) of Cuba, but smaller. Paler green, lime-green to mignonette-green (in the Cuban bird about Kronberg's green); under tail coverts and under surface of tail (be;s-ond the red base) paler and more yellowish; red belly patch always large, more sharply contrasted and brighter red without bangs: birds from the cayman islands. 309 lavender shimmer — bright hych-anger-red — (dark vinaceous, almost always, more or less touched with lavender in true leucocephala); outer surface of closed wing paler and duller blue, more greenish, — much more as in A. l. caymanensis; shorter upper tail coverts, some- times also longer upper tail coverts and lower rump feathers, more or less extensively edged and tipped with red (Cuban examples of true A. leucocephala seldom show such red markings, and when they do only to a very slight extent). Brown noted the iris as brown and the orbital skin as white to gra^-ish white, the tarsus as yellow. Measurements. Caymax Brag. No. Sex 68312 cT 68313 . d" 68315 d" 68316 c? 68309 9 68310 9 68311 9 68314 9 68317 9 68308 9 Remarks. Brown found this Parrot to be not uncommon though of rather local distribution in Cayman Brae. He also took one speci- men in Little Cavman on Julv 25. CUCULIDAE. Crotophaga ani Linrie. Three specimens, both sexes, all adult, Grand Cayman and Little CajTnan, May and July. * CoccYZUS minor nesiotes (Cab. and Heine). Eleven specimens, both sexes, all | adult. Grand Cayman, Little Cayman, and Cayman Brae, May, June, and July. Culmen Wing Tail Tarsus from Cere 132 110 21 27 134 109 23 27 133 113 21 27 129 103 22 25 132 109 22 26 128 111 23 25 126 102 21 24 129 107 21 23 134 106 21 25 Little Caymax. 131 ill 22 25 310 bulletin: museum of comparative zoology. These specimens agree with Jamaican skins in size and proportions and are a Uttle larger than the Bahaman form C. m. maynardi Ridg. In the color of the under parts this series shows a wide range of indi- vidual variation. The darkest ones are exactly like the paler speci- mens from Jamaica and the palest ones like the darker examples of maynardi. Thus as a whole the series averages a little paler below than the average of a long series of nesiotes from Jamaica. All, how- ever, were taken later in the season, than any skin we have from Jamaica and are without doubt somewhat faded out. Hybrididae. Hybris perlata furcata (Temm.). One adult 9 , Cayman Brae, June 27. This is a very pale and gray individual, rather different from ordinary specimens from Jamaica or Cuba. In a long series from Jamaica, however, we have one skin that agrees with it exactly. Caprimulgidae. Chordeiles virginianus minor (Cab.). One adult cf , Little Cayman, July 17. Brown made no note on the abundance of the Little Nighthawk in the Caymans, which I believe has not before been recorded from the islands. It is possible that this individual may have been a migrant from Cuba, where the bird breeds abundantl^r', but apparently does not winter. PiCIDAE. t COLAPTES CHRYSOCAULOSUS GUNDLACHI Cory. Thirteen specimens, both sexes, all adult, Grand Cayman, April, May, and June. This series shows in a marked degree the two characters pointed out by Ridgway in his Birds of North and Middle America — smaller size and smaller and narrower black malar patch in the males — which distinguish it from the Cuban form. It is confined to Grand Cavman. bangs: birds from the cayman islands. 311 t Centurus caymanensis Cory. Nine specimens, both sexes, all adult, Grand Cayman, April, May, and June. This strongly characterized island species, is one of the commoner birds of Grand Cayman to which island it is confined. Tyrannidae. Tyrannus dominicensis dominicensis (Gmel.). Four specimens, both sexes, all adult, Grand Cayman and Cayman Brae, April and June. TOLMARCHUS CAYMANENSIS (NicoU). Twenty specimens, both sexes, young and adult, Grand Cayman, Little Cayman, and Cayman Brae, April, May, June, and July. This is a well-marked form peculiar to the Caymans, where, I be- lieve, it does not differ either in color or size in the three islands of the group. In fresh spring plumage (April specimens from Grand Cay- man) the back is distinctly olivaceous, as compared with the gray back in the Cuban form, T. caudifasciatiLS (D'Orbigny), in similar plumage. In birds killed by May 25 and from then on through the summer, the color of the back, by fading and wear, has changed to a dirty grayish, quite the same as in Cuban skins in the same condition of feather. In this plumage the Cayman bird can only be recognized by its much duller, browner head, less contrasted with the gray of the back — the head of the Cuban bird in worn plumage being very black and sharply contrasted against the color of the back. The Cayman bird also has a longer and more slender bill, this character being well marked as an average one, but unfortunately failing in the case of certain individuals. All the adults from Little Cayman and Cayman Brae, except one, are in the worn and faded midsummer plumage just referred to; the one exception is M. C. Z. 68248 Cayman Brae, June 29, which, though taken on a date earlier than some others that had not changed, has almost completed the postnuptial moult and has again an olivaceous back. The color of the back in this skin is quite the same as in the April specimens from Grand Cayman, while the more faded Grand Cayman individuals killed May 25 are like the 312 bulletin: museum of comparative zoology. ones from Little Cayman and Cayman Brae taken in June and July. Brown took fully fledged young from July 10 to July 28. The wing in the adults in this series ranges, in Grand Cayman skins, from 103- 107; in skins from Cayman Brae, 97-103; in the only adult from Little Cayman (a cf ) it is 108. The Cayman Brae specimens have the tips of the primaries a little more worn down than the Grand CajTTian ones. Myiarchus sagrae sagrae (Gundlach). Myiarchus denigratus Cory, Auk, Oct. 1886, 3, p. 500, 502, Grand Cayman. Ten specimens, both sexes, all adult. Grand Cayman, April and May. Apparently this bird is found in Grand Cayman only of the Caymans. The specimens in the present series are indistinguishable in any way from Cuban skins. * Elaenia martinica caymanensis Berlepsch. Elaenia martinica complexa Berlepsch, Proc. 4th International ornith. congress, 1905, p. 395, Cayman Brae. Twenty-six specimens, both sexes, all adult, Grand Cayman, Little Cayman, and Cayman Brae, April, May, June, and July. Specimens in exactly similar plumage from the three islands of the Cayman group are absolutely alike, and no subdivision can be made. I am sure Berlepsch was deceived by the artificial discoloration of Maynard's Cayman Brae skins, upon examples of which he based his E. vi. complexa. Two such skins are now before me and I do not wonder at such a mistake being made. The Cayman Elaenia fades and bleaches out late in summer, losing all its colors. Two skins collected in Grand Cayman in August, 188(3, by W. B. Richardson, have lost all traces of the colors and markings of the form when in fresh plumage. The April speci- mens in the present series from Grand Cayman are in beautiful fresh unfaded plumage. Some of the late July skins from Cayman Brae have nearly completed the postnuptial moult and are indistinguish- able from these. Others taken at the same time had not commenced to moult, and are nearly as faded out as the August examples just referred to. The Cayman bird appears an excellent form, but I cannot agree with /^ bangs: birds from the cayman islands. 313 some other ornithologists that its nearest relation is E. m. riisii Sol. of St. Thomas. Nor do I think it very closely related to any of the dis- tinctly gray Lesser Antillean forms. It seems obviously much more like E. m. subpagana Scl. and Salv. of the near by mainland, with which it exactly agrees in size and markings and in color except in being j)aler throughout. In good plumage the belly is uniformly yellow, the chest dull yellowish gray, the throat grayish white, and the upper parts olive. All these colors, however, are much paler than in the continental bird. I think that this bird was derived not through any of the Lesser Antillean forms, which on zoogeographical grounds would seem out of reason, but like Vireosyha caymanensis and probably Vireo crassi- rostris direct from the form occupying the adjacent mainland. MiMIDAE. ^ MiMUS POLYGLOTTOS ORPHEUS (Linne). One adult cf, Grand Cayman, May 14. The Jamaican Mockingbird is abundant in Grand Cayman, but apparently is wanting in the two smaller islands. TURDIDAE. MiMocicHLA RAViDA Cory. Thirteen specimens, both sexes, all adult. Grand Cayman, April, May, and June. The Grand Cayman Thrush belongs in a group of the genus Mimo- cichla by itself, and of all the birds peculiar to the Caj'mans is the only one that is very distinct, having no representative elsewhere. In view of the recent origin of the ornis of the Caymans, it is probable that there was somewhere, possibly in Jamaica, where no member of the genus now occurs, a related form which has disappeared. The Thrush is now extremely rare and local in Grand Cayman. Brown covered the whole island and found it only in two remote patches of woodland. Each of these tracts of rather heavier forest than is usual in the island now-a-days was inliabited by a few pairs of thrushes, which Brown believes to be the entire population of the island. In each of these woods Brown was careful to leave birds enough to perpetuate the species, if it is not gradually becoming extinct from some natural cause, as seems to be the case. 314 bulletin: museum of comparative zoology. Brown noted the colors of the soft parts to be as follows: — "Iris, brown; tarsus, bill, and bare skin of orbital region, coral red." MiMOCiCHLA RUBRiPES CORYI Sharpe. Twenty-three specimens, both sexes, all adult, Cayman Brae, June and July. Unlike its cousin of the larger island, the Cayman Brae Thrush is an extremely abundant bird. It is a very well-marked form, with a large light-colored bill. ViREONIDAE. ViREOSYLVA MAGISTER CAYTVLAJMENSIS (Cory). Twelve specimens, both sexes, all adult. Grand Cayijian, April and May. Brown found this Vireo in the mangroves in Grand Cayman, where it was not uncommon. It has been recorded from both Little Cayman and Cayman Brae by Cory, but Brown did not find it in either of the smaller islands, where its place seemed to be wholly taken by V. calidris barbatula. The Grand Cayman Vireo is very closely related to true V. magister LawT. of the coast of British Honduras, from which it differs only by its paler coloration. ViREOSYLVA CALIDRIS BARBATULA (Cab.). Fifteen specimens, both sexes, all adult. Little Cayman and Cayman Brae, June and July. ^ The Black-whiskered Vireo was very common in the two smaller islands of the group. The skins show no differences when compared with Cuban examples. ViREO CRASSiROSTRis CRASSiROSTRis (Bryant). Vireo alleni Cory, Auk, Oct. 1886, 3, p. 500-501, Grand Cayman. Seventeen specimens, both sexes, all adult, Grand Cayman, Little Cayman, and Cayman Brae, April, May, June, and July. This series critically compared with our sixty-fovu- skins from the Bahamas proves beyond a doubt that the much discussed V. alleni / bangs: birds from the cayman islands. 315 is absolutely identical with the Bahama bird. All the Cayman examples are in the yellow phase of plumage. They correspond exactly with yellow specimens from the Bahamas from Inagua to New Providence, the type locality of V. crassirostris. The three characters that Ridgway in his Birds of North and Middle America thought might distinguish V. alleni, all prove illusive. The browner back in the specimens he examined was due entirely to discoloration from the now famous chemical preservative used by Maynard; the outermost primary is not smaller; and the pale wing-bands are not broader. Todd (Annals Carnegie mus., 1911, 7, p. 428^30) has discussed at length the color-phases of V. crassirostris, and I wholly agree with him that the gray and the yellow (the so-called Vireo crassirostris fiavescens Ridg.) specimens, represent nothing but extremes of color-variation in one and the same subspecies. Examples from the different islands of the Caymans are all quite alike. Mniotiltidae. Dendroica petechia petechia (Linne). Dendroica auricapilla Ridg., Proc. U. S. N. M., Aug. 1888, 10, p. 572, Grand Cayman. Thirteen specimens, both sexes, adults and two young. Grand Cay- man, Little Cayman, and Cayman Brae, April, May, and July. This series together with four skins from the Cajnnans already in the M. C. Z. I have compared most carefully with a fine set of Jamaican speciniens, with the result that I find no way in which to separate them. Ridgway in his Birds of North and Middle America recognizes auricapilla as differing from petechia on the grounds of "decidedly shorter wing and larger bill and feet." His own measure- ments, however, which followed, show very trifling differences. My measurements of eight adult males from the Caymans, the wing is : — 62-65, (63.81); exposed culmen, 10-11.5 (10.62). In eight adult males from Jamaica, the wing is: — 62-67 (64.5); exposed culmen, 10-11 (10.68). I can see no differences at all in the feet. There are no differences in specimens from the three islands of the Caymans. Dendroica petechia petechia can be separated from D. p. gundlachi Baird of Cuba by slightly paler colors and more extensively ochraceous €rown. 316 bulletin: museum of comparative zoology. Dendroica vitellina vitellina Cory. Ten specimens, both sexes, all adult. Grand Cayman, April and May. This fine island form confined to Grand Cayman, was in Brown's experience a very uncommon bird and he told me that it was with difficulty that he got even the ten noted above. Dendroica vitellina crawfordi Nicoll. Thirty-seven specimens, both sexes, adults and young, Little Cay- man and Cayman Brae, June and July. This is a well-marked subspecies whose characters were accurately noted by Nicoll, (Bull. B. O. C, 1904, 14, p. 95) who also figured it (Ibis, 1904, ser. 8, 4, pi. 11, f. 1). It is an abundant bird in the two smaller islands, and is quite the same in both. Coerebidae. * COEREBA SHARPEI (Cory). Twenty-eight specimens, both sexes, all adult, Grand Cayman, Little Cayman, and Cayman Brae, April, May, June, and July. Brown's specimens from Grand Cayman are unfortunately not comparable with his series from Little Cayman and Cayman Brae, and I am unable to say whether the differences shown by birds from the two smaller islands, when compared with examples from Grand Cayman, are seasonal or not. I am inclined, however, to regard these differences as only seasonal. The Grand Cayman birds, all taken in April and May, were in worn and somewhat faded breeding plumage, while those from Little Cayman and Cayman Brae, taken in late June and July, had completed or were just completing the post- nuptial moult, and were therefore all in what might be called fresh autumnal plumage. The upper parts in the Grand Cayman specimens are dull brownish black; the yellow of the under parts is pale and dull. The upper parts in the Little Cayman and Cayman Brae skins are grayish black with a slight olivaceous cast; the yellow of the under parts is richer and rather more orange. Brown noted that the " skin at corners of mouth, red" in the Grand Cayman bird; " skin at corners of mouth, flesh-color" in Little Cayman and Cayman Brae specimens. This possibly also has to do with the breeding season. bangs: birds from the cayman islands. 817 icteridae. * holoquiscalus caymanensis caymanensis (cory). Foiu" specimens, three males and a female, all adult. Grand Cayman, May. Brown had to spend so much time while in Grand Cayman search- ing for the rare species, that he rather neglected the Grackle and some of the other very common birds. This is a very well-marked insular subspecies peculiar to Grand Cayman. HOLOQTJISCALUS CAYMANENSIS CARIBAETJS Todd. Fourteen specimens, both sexes, adults and one young. Little Cayman and Cayman Brae, June and July. The Grackle of the two smaller islands which differs from true H. caymanensis of Grand Cayman in its much larger size and stronger bill, has always been referred to H. gundlachii (Cassin) of eastern Cuba. I had in the present paper corrected this old error, and had named the form as new, arriving at the same conclusions as Todd, except that he did not know the bird of Cayman Brae and Little Cayman, which is identical with that inhabiting the Isle of Pines and western Cuba. Todd's paper. The Birds of the Isle of Pines, Annals of the Carnegie museum, 10, nos. 1-2, (dated Jan. 1916, but received by M. C. Z. Mar. 1, 1916), containing a description of the form, came just in time to allow me to change the name while reading proof. Icterus bairdi Cory. Seventeen specimens, both sexes, adults, and five immature (one year old?) birds still carrying a partly or wholly greenish yellow tail. Grand Cayman, April, May, and June. A nest made of palm fibres and attached to a hemp palm leaf about sixty feet from the ground was found 28 May; the nest contained three young birds. This splendid island species confined to Grand Cayman differs from /. leiicopteryx (Wagler) of Jamaica, from which it obviously was de- rived, in being bright golden yellow only slightly tinged with olive on the head, and just a trifle darker on the back than it is below. It is also a little smaller and has a slightly slenderer and more delicate bill. 318 bulletin: museum of comparative zoology. Baird's Oriole has always been extremely rare in collections, in fact besides our series there exist only Cory's original specimens and two in the Tring Museum that were collected by Taylor when he visited the island in 1896 for the Hon. Walter Rothschild. The species seems to be on the verge of extinction. Why this is I can offer no suggestion. Certainly I. leucopteryx is common enough in Jamaica and adapts itself to all the changes man makes there. Brown found this Oriole scattered here and there at wide intervals in the island and told me he thought it was one of the rarest birds he had ever himted for. Tanagridae. * Spindalis salvini Cory. Fifty-five specimens, both sexes (only five females) all adult. Grand Cayman, April and May. This is a fine, large species peculiar to Grand Cayman. Its nearest relative is clearly S. pretrei (Lesson) of Cuba. Its bill though of course larger than in the Cuban species, the bird itself being much larger, is very like it, and quite different from the heavy coarse bill of